On the Origin of Species, 6th Edition

Darwin, Charles

Published: 1872 Categorie(s): Non-Fiction, Science Source: Feedbooks

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

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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

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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."

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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 wellknown paper ("Edinburgh Philosophical Journal", vol. XIV, page 283) on the Spongilla, clearly declares his belief that species are descended from

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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

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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

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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

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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

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writes in 1854 ("Etudes sur Geograph. The "Philosophy of Creation" has been treated in a masterly manner by the Rev. page 250). so at certain periods the germs of existing species may have been chemically affected by circumambient molecules of a particular nature. "a natural in contradistinction to a miraculous process. twenty-seven have written on special branches of natural history or geology. as Sir John Herschel expresses it. Burdach. in Pander and Dalton's work on Fossil Sloths. Lecoq. Geoffroy Saint-Hilaire et Goethe. page 357). supposed to have been caused by some miasma have arisen and spread over the world. des Naturhist.. a similar belief." 11 . M. and thus have given rise to new forms." or. He infers that many species have kept true for long periods.celebrated botanist and palaeontologist Unger published. I may add. tom. Geolog. Dr.). 1853. 1855. in his "Essays on the Unity of Worlds". been maintained by Oken in his mystical "Natur-Philosophie". Similar views have. whereas a few have become modified. 2nd Ser. Poiret and Fries. Baden Powell. tom. in 1852. Nothing can be more striking than the manner in which he shows that the introduction of new species is "a regular. i.". Bot. not a casual phenomenon. that of the thirty-four authors named in this Historical Sketch. Count Keyserling ("Bulletin de la Soc. likewise. nous conduisent directement aux idees emises par deux hommes justement celebres. etc. Vincent. In this same year. but are to be regarded as their descendants through continued reproduction. expressed." A well-known French botanist. Rheinlands". Lecoq's large work make it a little doubtful how far he extends his views on the modification of species. or at least disbelieve in separate acts of creation. in 1821. The distinction of species he explains by the destruction of intermediate graduated forms. Schaaffhausen published an excellent pamphlet ("Verhand. his belief that species undergo development and modification. "Thus living plants and animals are not separated from the extinct by new creations. Dalton.) In 1853 a celebrated geologist. who believe in the modification of species. in which he maintains the development of organic forms on the earth. it seems that Bory St. suggested that as new diseases. have all admitted that new species are continually being produced. x. Vereins der Preuss. From other references in Godron's work "Sur l'Espece". "On voit que nos recherches sur la fixite ou la variation de l'espece." Some other passages scattered through M. as is well known.

and sadly damaged by some of its supporters. 12 . and supports this doctrine by many original observations. 1859. read July 1. Von Baer.The third volume of the "Journal of the Linnean Society" contains papers. Rudolph Wagner. If. 1859. 1861. 1860. In June. is yet the only one to which physiology lends any countenance. The first edition of this work was published on November 24. and the second edition on January 7. 1859. toward whom all zoologists feel so profound a respect. if we suppose that each species of animal and plant. In the first part of this great work he admits the truth of the descent and modification of species. he remarks. 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. though unproven. Referring to such cases. "Zoologisch-Anthropologische Untersuchungen". chiefly grounded on the laws of geographical distribution. Professor Huxley gave a lecture before the Royal Institution on the "Persistent Types of Animal Life". "It is difficult to comprehend the meaning of such facts as these. on the other hand. or each great type of organisation. Wallace and myself. in which. the theory of Natural Selection is promulgated by Mr. a hypothesis which. Dr. 1858. Hooker published his "Introduction to the Australian Flora"." In December. expressed about the year 1859 (see Prof. as stated in the introductory remarks to this volume. that forms now perfectly distinct have descended from a single parent-form. was formed and placed upon the surface of the globe at long intervals by a distinct act of creative power. by Mr. Wallace with admirable force and clearness. 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. 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. 51) his conviction. s.

these I enlarged in 1844 into a sketch of the conclusions. Lyell and Dr. Wallace's excellent memoir. No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts. with a few facts in illustration. For I am well aware that scarcely a single point is discussed in this volume on which facts cannot be 13 . who is now studying the natural history of the Malay Archipelago. seemed to throw some light on the origin of species—that mystery of mysteries. but which. No doubt errors may have crept in. in 1837. as will be seen in the latter chapters of this volume. After five years' work I allowed myself to speculate on the subject. and as my health is far from strong. as Mr. My work is now (1859) nearly finished. which then seemed to me probable: from that period to the present day I have steadily pursued the same object. I have been urged to publish this abstract. which I now publish. On my return home. Hooker. These facts. I can here give only the general conclusions at which I have arrived. This abstract.Introduction When on board H.S. and it is published in the third volume of the Journal of that Society. and drew up some short notes. Wallace. it occurred to me. who sent it to the Linnean Society. 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. I have more especially been induced to do this. as naturalist. I hope. some brief extracts from my manuscripts. Beagle. and I hope in a future work to do this. as it has been called by one of our greatest philosophers. I hope that I may be excused for entering on these personal details. Sir C. on which my conclusions have been grounded. and I must trust to the reader reposing some confidence in my accuracy. though I hope I have always been cautious in trusting to good authorities alone. has arrived at almost exactly the same general conclusions that I have on the origin of species. in most cases will suffice. must necessarily be imperfect. but as it will take me many more years to complete it.M. with references. with Mr. and in the geological relations of the present to the past inhabitants of that continent. with a request that I would forward it to Sir Charles Lyell. I cannot here give references and authorities for my several statements. who both knew of my work—the latter having read my sketch of 1844—honoured me by thinking it advisable to publish. I was much struck with certain facts in the distribution of the organic beings inhabiting South America. In 1858 he sent me a memoir on this subject. as I give them to show that I have not been hasty in coming to a decision.

It is. some of them personally unknown to me. in this and in all other perplexing cases I have invariably found that our knowledge.. might come to the conclusion that species had not been independently created. let this opportunity pass without expressing my deep obligations to Dr. would be unsatisfactory.adduced. of variation under domestication. who. In considering the origin of species. reflecting on the mutual affinities of organic beings. beak. or of habit. but had descended. but it is preposterous to attribute to mere external conditions. as the only possible cause of variation. which has seeds that must be transported by certain birds. from other species. of the highest importance to gain a clear insight into the means of modification and coadaptation. has aided me in every possible way by his large stores of knowledge and his excellent judgment. on their embryological relations. imperfect though it be. their geographical distribution. 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. Nevertheless. which draws its nourishment from certain trees. In one limited sense. as we shall hereafter see. food. 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. inhabiting this world have been modified. such as climate. so admirably adapted to catch insects under the bark of trees. even if well founded. for the last fifteen years. often apparently leading to conclusions directly opposite to those at which I have arrived. geological succession. tail. and tongue. and which has flowers with separate sexes absolutely requiring the agency of certain insects to bring pollen from one flower to the other. Naturalists continually refer to external conditions. afforded 14 . by the effects of external conditions. or of the volition of the plant itself. such a conclusion. etc. until it could be shown how the innumerable species. this may be true. for instance. and this is here impossible. with its feet. it is equally preposterous to account for the structure of this parasite. with its relations to several distinct organic beings. the structure. Hooker. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question. and other such facts. of the woodpecker. so as to acquire that perfection of structure and coadaptation which justly excites our admiration. like varieties. Nor have I been disappointed. therefore. however. In the case of the mistletoe. it is quite conceivable that a naturalist. I cannot.

secondly the subject of instinct. first. hybridism. and as. their geographical distribution throughout space. we shall see how great is the power of man in accumulating by his selection successive slight variations. the difficulties of transitions. From the strong principle of inheritance. and. both when mature and in an embryonic 15 . which inevitably follows from the high geometrical ratio of their increase. I will then pass on to the variability of species in a state of nature. will be considered. be enabled to discuss what circumstances are most favourable to variation. in the twelfth and thirteenth. if it vary however slightly in any manner profitable to itself. thirdly. I shall devote the first chapter of this abstract to variation under domestication. or the infertility of species and the fertility of varieties when intercrossed. consequently. In the next chapter the struggle for existence among all organic beings throughout the world. but I shall. however. or the mental powers of animals. unfortunately. their classification or mutual affinities. the most apparent and gravest difficulties in accepting the theory will be given: namely. although they have been very commonly neglected by naturalists.the best and safest clue. any selected variety will tend to propagate its new and modified form. in the fourteenth. and thus be NATURALLY SELECTED. I may venture to express my conviction of the high value of such studies. under the complex and sometimes varying conditions of life. it follows that any being. 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. there is a frequently recurring struggle for existence. applied to the whole animal and vegetable kingdoms. In the next chapter I shall discuss the complex and little known laws of variation. what is equally or more important. In the next chapter I shall consider the geological succession of organic beings throughout time. and fourthly. As many more individuals of each species are born than can possibly survive. and leads to what I have called divergence of character. We shall. be compelled to treat this subject far too briefly. This fundamental subject of natural selection will be treated at some length in the fourth chapter. the imperfection of the geological record. and we shall then see how natural selection almost inevitably causes much extinction of the less improved forms of life. From these considerations. In the five succeeding chapters. This is the doctrine of Malthus. 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. will have a better chance of surviving.

and why another allied species has a narrow range and is rare? Yet these relations are of the highest importance. I am fully convinced that species are not immutable. 16 . that each species has been independently created—is erroneous. that the view which most naturalists until recently entertained. 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. as I believe. but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species. Furthermore. in the same manner as the acknowledged varieties of any one species are the descendants of that species. the future success and modification of every inhabitant of this world. for they determine the present welfare and. I can entertain no doubt. means of modification. if he make due allowance for our profound ignorance in regard to the mutual relations of the many beings which live around us.condition. after the most deliberate study and dispassionate judgment of which I am capable. and which I formerly entertained—namely. Although much remains obscure. In the last chapter I shall give a brief recapitulation of the whole work. and will long remain obscure. Who can explain why one species ranges widely and is very numerous. I am convinced that natural selection has been the most important. and a few concluding remarks. No one ought to feel surprise at much remaining as yet unexplained in regard to the origin of species and varieties. but not the exclusive.

Chapter 1 Variation Under Domestication 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. their Effects — Methodical and Unconscious Selection — Unknown Origin of our Domestic Productions — Circumstances favourable to Man's power of Selection. anciently followed. their Differences and Origin — Principles of Selection. 17 .

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. also. when the organisation has once begun to vary. be little doubt about many slight changes. on the other hand. As far as I am able to judge.1. it generally continues varying for many generations. The former seems to be much the more important. There is. The effects on the offspring are either definite or in definite. the nature of the organism and the nature of the conditions. dissimilar variations arise under conditions which appear to be nearly uniform. dissimilar conditions. and. after long attending to the subject. those to which the parent species had been exposed under nature. that this variability may be partly connected with excess of food. There can. and as I have incidently shown in my work on "Variation under Domestication. and somewhat different from. and that. some probability in the view propounded by Andrew Knight. Our oldest cultivated plants. With respect to the direct action." there are two factors: namely. such as wheat. It is extremely difficult to come to any conclusion in regard to the extent of the changes which have been thus definitely induced. however. And if we reflect on the vast diversity of the plants and animals which have been cultivated. It seems clear that organic beings must be exposed during several generations to new conditions to cause any great amount of variation. that they generally differ more from each other than do the individuals of any one species or variety in a state of nature. No case is on record of a variable organism ceasing to vary under cultivation. 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. colour from the nature of the food. for nearly similar variations sometimes arise under. still yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification. and which have varied during all ages under the most different climates and treatment. 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. Each of the endless variations which we see in 18 . as far as we can judge. one of the first points which strikes us is. as Professor Weismann has lately insisted. thickness of the skin and hair from climate. we must bear in mind that in every case. Causes of Variability When we compare the individuals of the same variety or sub-variety of our older cultivated plants and animals. etc. such as size from the amount of food.

partly from the fact of this system being extremely sensitive to any change in the conditions. we may infer that variability is thus induced. but monstrosities cannot be separated by any distinct line from slighter variations. and that which may be observed with plants and animals when reared under new or unnatural conditions. 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. and yet rarely 19 . With respect to what I have called the in direct action of changed conditions. even when the male and female unite. Many facts clearly show how eminently susceptible the reproductive system is to very slight changes in the surrounding conditions. shows us what singular modifications might result in the case of plants from a chemical change in the nature of the sap. which appear among many individuals living together.the plumage of our fowls must have had some efficient cause. through the reproductive system of being affected. and has probably played a more important part in the formation of our domestic races. causing coughs or colds. according to their state of body or constitution. At long intervals of time. all probably would be modified in the same manner. and few things more difficult than to get it to breed freely under confinement. but erroneously attributed to vitiated instincts. and which cannot be accounted for by inheritance from either parent or from some more remote ancestor. whether extremely slight or strongly marked. Many cultivated plants display the utmost vigour. All such changes of structure. or inflammation of various organs. out of millions of individuals reared in the same country and fed on nearly the same food. may be considered as the in definite effects of the conditions of life on each individual organism. How many animals there are which will not breed. namely. Nothing is more easy than to tame an animal. and partly from the similarity. We see in definite variability in the endless slight peculiarities which distinguish the individuals of the same species. between the variability which follows from the crossing of distinct species. in nearly the same manner as the chill effects different men in an in definite manner. and in seedlings from the same seed-capsule. though kept in an almost free state in their native country! This is generally. In definite variability is a much more common result of changed conditions than definite variability. Even stronglymarked differences occasionally appear in the young of the same litter. rheumatism. deviations of structure so strongly pronounced as to deserve to be called monstrosities arise. as Kolreuter and others have remarked. and if the same cause were to act uniformly during a long series of generations on many individuals.

on the one hand. and as buds on distinct trees. though taken young from a state of nature perfectly tamed.or never seed! In some few cases it has been discovered that a very trifling change. but this is certainly an error. in the same condition as in the most sterile hybrids. and vary very slightly—perhaps hardly more than in a state of nature. and healthy (of which I could give numerous instances). acting irregularly. and when. I cannot here give the details which I have collected and elsewhere published on this curious subject. we see domesticated animals and plants. As a single bud out of many thousands produced year after year on the same tree under uniform conditions. such as a little more or less water at some particular period of growth. buds on peach. carnivorous birds. we see individuals. even from the tropics. but to show how singular the laws are which determine the reproduction of animals under confinement. with the rarest exception. breeding freely under confinement. etc. growing under different conditions. breed in this country pretty freely under confinement. though often weak and sickly. we need not be surprised at this system. When. yet having their reproductive system so seriously affected by unperceived causes as to fail to act. I may add that as some organisms breed freely under the most unnatural conditions—for instance. and sometimes by seed. rabbits and ferrets kept in hutches—showing that their reproductive organs are not easily affected. but are far from rare under culture. has been known suddenly to assume a new character. will determine whether or not a plant will produce seeds. Some naturalists have maintained that all variations are connected with the act of sexual reproduction. as they may be named. when it does act under confinement. that is. These bud variations. which seldom produce young. offsets.trees producing nectarines. long-lived.. whereas. 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. and producing offspring somewhat unlike their parents. Many exotic plants have pollen utterly worthless. 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 20 . with the exception of the plantigrades or bear family. hardly ever lay fertile eggs. have sometimes yielded nearly the same variety—for instance. so will some animals and plants withstand domestication or cultivation. They occur rarely under nature. for I have given in another work a long list of "sporting plants. can be propagated by grafts. on the other hand. I may mention that carnivorous animals." as they are called by gardeners.

has in determining the nature of the flames. by which a mass of combustible matter is ignited. perhaps of not more importance than the nature of the spark.the organism in determining each particular form of variation. 21 .

while dark-coloured individuals escape: Professor Wyman has recently communicated to me a good illustration of this fact. which coloured their bones pink.2. as they alone have a good chance of living. and many instances are given in Isidore Geoffroy St. in comparison with these organs in other countries. Colour and constitutional peculiarities go together. Some instances of correlation are quite whimsical. Virginia squatters) added. thus cats which are entirely white and have blue eyes are generally deaf. some few of which can be dimly seen. and this change may be safely attributed to the domestic duck flying much less. on asking some farmers in Virginia how it was that all their pigs were black. than do the same bones in the wild duck. and the view which has been suggested that the drooping is due to disuse of the muscles of the ear. Breeders believe that long limbs are almost always accompanied by an elongated head. they informed him that the pigs ate the paint-root (Lachnanthes). 22 . and one of the "crackers" (i. it appears that white sheep and pigs are injured by certain plants. from the animals being seldom much alarmed. but it has been lately stated by Mr. Important changes in the embryo or larva will probably entail changes in the mature animal. I will here only allude to what may be called correlated variation. Hilaire's great work on this subject. seems probable. Correlated Variation. With animals the increased use or disuse of parts has had a more marked influence. thus I find in the domestic duck that the bones of the wing weigh less and the bones of the leg more. Not one of our domestic animals can be named which has not in some country drooping ears." Hairless dogs have imperfect teeth. In monstrosities. From facts collected by Heusinger. long-haired and coarse-haired animals are apt to have. in proportion to the whole skeleton. as is asserted. the correlations between quite distinct parts are very curious. Many laws regulate variation.e. "we select the black members of a litter for raising. The great and inherited development of the udders in cows and goats in countries where they are habitually milked. is probably another instance of the effects of use. and will hereafter be briefly discussed. and walking more. of which many remarkable cases could be given among animals and plants. Inheritance Changed habits produce an inherited effect as in the period of the flowering of plants when transported from one climate to another. Tait that this is confined to the males. Effects Of Habit And Of The Use Or Disuse Of Parts. and which caused the hoofs of all but the black varieties to drop off. than its wild parents.

Every one must have heard of cases of albinism. The whole organisation seems to have become plastic. appears in the parent—say. are endless. apparently exposed to the same conditions. If strange and rare deviations of structure are truly inherited. and non-inheritance as the anomaly. Dr. or but dimly understood laws of variation are infinitely complex and diversified.. No breeder doubts how strong is the tendency to inheritance. and departs in a slight degree from that of the parental type. Any variation which is not inherited is unimportant for us. the mere doctrine of chances almost compels us to attribute its reappearance to inheritance. as on the hyacinth. that like produces like is his fundamental belief: doubts have been thrown on this principle only by theoretical writers. once among several million individuals—and it reappears in the child. due to some extraordinary combination of circumstances. why the child often reverts in certain characteristics to its grandfather or grandmother or more remote ancestor. why a peculiarity is often transmitted from one sex to both sexes. pigeons with feathered feet have skin between their outer toes. pigeons with short beaks have small feet. to look at the inheritance of every character whatever as the rule. no one can say why the same peculiarity in different individuals of the same species. unknown. Hence if man goes on selecting. less strange and commoner deviations may be freely admitted to be inheritable. The laws governing inheritance are for the most part unknown. But the number and diversity of inheritable deviations of structure. etc. we cannot tell whether it may not be due to the same cause having acted on both. more 23 . etc. The results of the various. 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. and we see it in the father and child. is the fullest and the best on this subject. even the dahlia. Perhaps the correct way of viewing the whole subject would be. but when among individuals. is sometimes inherited and sometimes not so. potato. prickly skin. he will almost certainly modify unintentionally other parts of the structure. or in different species. or to one sex alone. When any deviation of structure often appears. and thus augmenting. hairy bodies. appearing in several members of the same family. owing to the mysterious laws of correlation. any very rare deviation.. both those of slight and those of considerable physiological importance. and those with long beaks large feet. any peculiarity. It is well worth while carefully to study the several treatises on some of our old cultivated plants.long or many horns. in two large volumes. Prosper Lucas' treatise.

however. In many cases we do not know what the aboriginal stock was. Having alluded to the subject of reversion. Hence it has been argued that no deductions can be drawn from domestic races to species in a state of nature.commonly but not exclusively to the like sex. during many generations. 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. is clearly due to the male element. it seems to me not improbable that if we could succeed in naturalising. to the males alone. that our domestic varieties. in very poor soil—in which case. that only a single variety should be turned loose in its new home. In many cases this could not be otherwise. that peculiarities appearing in the males of our domestic breeds are often transmitted. I have in vain endeavoured to discover on what decisive facts the above statement has so often and so boldly been made. peculiarities in the silk-worm are known to appear at the corresponding caterpillar or cocoon stage. I believe this rule to be of the highest importance in explaining the laws of embryology. It is a fact of some importance to us. and so could not tell whether or not nearly perfect reversion had ensued. for instance. some effect would have to be attributed to the DEFINITE action of the poor soil —that they would. either exclusively or in a much greater degree. to a large extent. yet that it does tend to appear in the offspring at the same period at which it first appeared in the parent. at whatever period of life a peculiarity first appears. It would be necessary. I may here refer to a statement often made by naturalists—namely. or were to cultivate. Nevertheless. it tends to reappear in the offspring at a corresponding age. These remarks are of course confined to the first APPEARANCE of the peculiarity. in nearly the same manner as the increased length of the horns in the offspring from a shorthorned cow by a long-horned bull. gradually but invariably revert in character to their aboriginal stocks. A much more important rule. when run wild. though appearing late in life. thus the inherited peculiarities in the horns of cattle could appear only in the offspring when nearly mature. which I think may be trusted. the several races. and that. though sometimes earlier. or 24 . when there is no apparent reason why a peculiarity should appear at any particular age. is that. But hereditary diseases and some other facts make me believe that the rule has a wider extension. and not to the primary cause which may have acted on the ovules or on the male element. of the cabbage. in order to prevent the effects of intercrossing. as our varieties certainly do occasionally revert in some of their characters to ancestral forms.

would be opposed to all experience. 25 . and esculent vegetables. by blending together. If it could be shown that our domestic varieties manifested a strong tendency to reversion—that is. Whether or not the experiment would succeed is not of great importance for our line of argument. revert to the wild aboriginal stock. for by the experiment itself the conditions of life are changed. in such case. and poultry of various breeds. for an unlimited number of generations.even wholly. so that free intercrossing might check. any slight deviations in their structure. long and short-horned cattle. I grant that we could deduce nothing from domestic varieties in regard to species. 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. while kept under the same conditions and while kept in a considerable body. to lose their acquired characters.

It can be shown that this statement is not correct. bloodhound. although differing from each other and from other species of the same genus. spaniel and bull-dog. as we shall presently see. Character Of Domestic Varieties. and compare them with closely allied species. would be interesting. but naturalists differ much in determining what characters are of generic value. from not knowing whether they are descended from one or several parent species. which we all know propagate their kind truly. that. It has often been stated that domestic races do not differ from each other in characters of generic value. for the domestic races of many animals and plants have been ranked by some competent judges as the descendants of aboriginally distinct species. all such valuations being at present empirical. both when compared one with another. that the whole amount of difference between the several breeds of the dog has 26 . Domestic races often have a somewhat monstrous character. This must be admitted as true. and by other competent judges as mere varieties. and more especially when compared with the species under nature to which they are nearest allied. Difficulty Of Distinguishing Between Varieties And Species. as already remarked. it will be seen that we have no right to expect often to find a generic amount of difference in our domesticated races. 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. they often differ in an extreme degree in some one part. were the offspring of any single species. we are soon involved in doubt. this source of doubt would not so perpetually recur. of the many foxes—inhabiting the different quarters of the world. If any well marked distinction existed between a domestic race and a species. With these exceptions (and with that of the perfect fertility of varieties when crossed—a subject hereafter to be discussed). if. it could be shown that the greyhound. if it could be cleared up. In attempting to estimate the amount of structural difference between allied domestic races. for instance. terrier. then such facts would have great weight in making us doubt about the immutability of the many closely allied natural species—for instance. I do not believe. by which I mean. we generally perceive in each domestic race. in several trifling respects. but the differences in most cases are less in degree.3. Origin Of Domestic Varieties From One Or More Species When we look to the hereditary varieties or races of our domestic animals and plants. When it is explained how genera originate under nature. less uniformity of character than in true species. This point.

or are even identical with. and belonging to equally diverse classes and countries. during which the domesticated animals. those still existing. or the small power of endurance of warmth by the reindeer. and we know that at the 27 . that we find in the most ancient times. The lake-inhabitants of Switzerland cultivated several kinds of wheat and barley. as Heer has remarked.habitations of Switzerland. when he first tamed an animal. I believe that a small part of the difference is due to their being descended from distinct species. The argument mainly relied on by those who believe in the multiple origin of our domestic animals is. equal in number to our domesticated productions. but how could a savage possibly know. and likewise to withstand diverse climates. were taken from a state of nature. and they possessed several domesticated animals. it is not possible to come to any definite conclusion. on the monuments of Egypt. might have varied and given rise to distinct races. I do not dispute that these capacities have added largely to the value of most of our domesticated productions. whether it would vary in succeeding generations. and shows that animals were domesticated at a much earlier period than has hitherto been supposed. kept by different tribes in different districts. all geologists believe that barbarian men existed at an enormously remote period. the pea. and this again implies a long continued previous period of less advanced civilisation. All this clearly shows. But this only throws far backward the history of civilisation. Since the discovery of flint tools in the superficial formations of many parts of the world. and whether it would endure other climates? Has the little variability of the ass and goose.been produced under domestication. and that some of these ancient breeds closely resemble. and in the lake. whether they are descended from one or several wild species. there is presumptive or even strong evidence that all are descended from a single wild stock. In the case of most of our anciently domesticated animals and plants. or of cold by the common camel. and could be made to breed for an equal number of generations under domestication. In the case of strongly marked races of some other domesticated species. they would on an average vary as largely as the parent species of our existing domesticated productions have varied. prevented their domestication? I cannot doubt that if other animals and plants. the poppy for oil and flax. It has often been assumed that man has chosen for domestication animals and plants having an extraordinary inherent tendency to vary. that they had at this early age progressed considerably in civilisation. much diversity in the breeds. They also carried on commerce with other nations.

and that their blood. I am doubtfully inclined to believe. The origin of most of our domestic animals will probably forever remain vague. Even in the case of the breeds of the domestic dog throughout the 28 . In regard to sheep and goats I can form no decided opinion. come to the conclusion that several wild species of Canidae have been tamed. Spain.. Blyth. as well as that of the specific distinction between the humped and common cattle. some breeds of which differ much from each other. From facts communicated to me by Mr. and of others who have studied this bird in India. They believe that every race which breeds true. may. after a laborious collection of all known facts. and France but few distinct from those of Germany. The doctrine of the origin of our several domestic races from several aboriginal stocks. and several goats. voice. that all the races belong to the same species. etc. This conclusion. But I may here state that. Having kept nearly all the English breeds of the fowl alive. and examined their skeletons.present day there is hardly a tribe so barbarous as not to have domesticated at least the dog. In regard to ducks and rabbits.. it is almost certain that they are descended from a different aboriginal stock from our European cattle. constitution and structure of the humped Indian cattle. in Europe alone. Gallus bankiva. we must admit that many domestic breeds must have originated in Europe. the evidence is clear that they are all descended from the common duck and wild rabbit. and several even within Great Britain. but that each of these kingdoms possesses several peculiar breeds of cattle. and some competent judges believe that these latter have had two or three wild progenitors. whether or not these deserve to be called species. sheep. Blyth. etc. has had its wild prototype. has been carried to an absurd extreme by some authors. and so with Hungary. looking to the domestic dogs of the whole world. flows in the veins of our domestic breeds. for whence otherwise could they have been derived? So it is in India. from reasons which I cannot here give. having bred and crossed them. in some cases mingled together. on the habits. in opposition to several authors. as many sheep. let the distinctive characters be ever so slight. it appears to me almost certain that all are the descendants of the wild Indian fowl. and this is the conclusion of Mr. With respect to horses. 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. At this rate there must have existed at least a score of species of wild cattle. be looked upon as established by the admirable researches of Professor Rutimeyer. I have. indeed.

which I admit are descended from several wild species. but to obtain a race intermediate between two quite distinct races would be very difficult. pug-dog. The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) quite uniform in character. 29 . and if we account for our several domestic races by this process. in the wild state. and every thing seems simple enough. Moreover. the bloodhound. but when these mongrels are crossed one with another for several generations. we must admit the former existence of the most extreme forms. for who will believe that animals closely resembling the Italian greyhound. as the Italian greyhound. Sebright expressly experimented with this object and failed. the bull-dog. 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.. bull-dog. or Blenheim spaniel. bloodhound. hardly two of them are alike. etc. Sir J. the possibility of making distinct races by crossing has been greatly exaggerated.—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.world. and then the difficulty of the task becomes manifest. etc. it cannot be doubted that there has been an immense amount of inherited variation. but by crossing we can only get forms in some degree intermediate between their parents.

The short-faced tumbler has a beak in outline almost like that of a finch. has a very short and broad one. The turbit has a short and conical beak. very large external orifices to the nostrils. and it has. is also remarkable from the wonderful development of the carunculated skin about the head. after deliberation. The barb is allied to the carrier. 30 . Elliot from India. The diversity of the breeds is something astonishing. instead of a long beak. I have. and by the Hon. as their names express. and have been most kindly favoured with skins from several quarters of the world. may well excite astonishment and even laughter. The Jacobin has the feathers so much reversed along the back of the neck that they form a hood. Murray from Persia. more especially by the Hon. utter a very different coo from the other breeds. W. Breeds Of The Domestic Pigeon. proportionally to its size. Several other less distinct breeds might be specified. some of the sub-breeds of runts have very long necks. slightly. the upper part of the oesophagus. I have associated with several eminent fanciers. others very long wings and tails. with long. wings. which it glories in inflating. and it has the habit of continually expanding. and have been permitted to join two of the London Pigeon Clubs. The pouter has a much elongated body. and this is accompanied by greatly elongated eyelids. and legs. The fantail has thirty or even forty tail-feathers. Many treatises in different languages have been published on pigeons. instead of twelve or fourteen. and the common tumbler has the singular inherited habit of flying at a great height in a compact flock. I have kept every breed which I could purchase or obtain. and tumbling in the air head over heels. with a line of reversed feathers down the breast. Their Differences And Origin Believing that it is always best to study some special group. elongated wing and tail feathers. as being of considerable antiquity. more especially the male bird. and some of them are very important. The runt is a bird of great size. The trumpeter and laugher. C. The carrier. massive beak and large feet. and its enormously developed crop. but. entailing corresponding differences in their skulls. and a wide gape of mouth.4. taken up domestic pigeons. and see the wonderful difference in their beaks. Compare the English carrier and the short-faced tumbler. 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. others singularly short tails.

varies in a highly remarkable manner. the males and females have come to differ in a slight degree from each other. the number of scutellae on the toes. namely. they must have descended from at least seven or eight aboriginal stocks. the relative length of the wing and tail to each other and to the body. and in some breeds the voice and disposition. more especially as in each of these breeds several trulyinherited sub-breeds. The shape. together with their relative breadth and the presence of processes. if shown to an ornithologist. as does the number of the ribs. the development and abortion of the oil-gland. of the tongue (not always in strict correlation with the length of beak).In the skeletons of the several breeds. The manner of flight. the relative length of the leg and foot. the development of skin between the toes. which. for it is impossible to make the present domestic breeds by the crossing of any lesser 31 . in certain breeds. I am fully convinced that the common opinion of naturalists is correct. and he were told that they were wild birds. including under this term several geographical races or sub-species. are all points of structure which are variable. pouter. differs enormously. the runt. or species. The period at which the perfect plumage is acquired varies. Moreover. Lastly. could be shown him. as well as the breadth and length of the ramus of the lower jaw. and have not proceeded from the rock-pigeon. I do not believe that any ornithologist would in this case place the English carrier. The caudal and sacral vertebrae vary in number. Great as are the differences between the breeds of the pigeon. the number of the primary wing and caudal feathers. I will here briefly give them. The shape and size of the eggs vary. in length and breadth and curvature. of the orifice of the nostrils. as he would call them. The size and shape of the apertures in the sternum are highly variable. which differ from each other in the most trifling respects. As several of the reasons which have led me to this belief are in some degree applicable in other cases. the barb. The proportional width of the gape of mouth. and fantail in the same genus. as does the state of the down with which the nestling birds are clothed when hatched. the size of the crop and of the upper part of the oesophagus. would certainly be ranked by him as well-defined species. the short-faced tumbler. that all are descended from the rock-pigeon (Columba livia). so is the degree of divergence and relative size of the two arms of the furcula. the proportional length of the eyelids. the development of the bones of the face. Altogether at least a score of pigeons might be chosen. differ remarkably. If the several breeds are not varieties.

therefore. and. and further. that these very species have since all become extinct or unknown. or barb. 32 . and the common rockpigeon. for reversed feathers like those of the Jacobin. though agreeing generally with the wild rock-pigeon in constitution. voice. or that of the short-faced tumbler. they did not breed or willingly perch on trees. which has the same habits with the domestic breeds. for tail-feathers like those of the fantail. not only that half-civilized man succeeded in thoroughly domesticating several species. So many strange contingencies are improbable in the highest degree. that the above-specified breeds. and applicable in several other cases. livia. as to be quite prolific under confinement. Hence it must be assumed. or they must have become extinct in the wild state. or on the shores of the Mediterranean. and this. only two or three other species of rock-pigeons are known. habits and remarkable characters. for instance. has become feral in several places. yet are certainly highly abnormal in other parts. but that he intentionally or by chance picked out extraordinarily abnormal species.number: how. habits. Hence the supposed extermination of so many species having similar habits with the rock-pigeon seems a very rash assumption. Again. 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 rockpigeons. though the dovecot-pigeon. But besides C. all recent experience shows that it is difficult to get wild animals to breed freely under domestication. yet on the hypothesis of the multiple origin of our pigeons. are unlikely to be exterminated. and yet be unknown to ornithologists. seems improbable. we may look in vain through the whole great family of Columbidae for a beak like that of the English carrier. some of them must have been carried back again into their native country. the several above-named domesticated breeds have been transported to all parts of the world. which is the rock-pigeon in a very slightly altered state. has not been exterminated even on several of the smaller British islets. But birds breeding on precipices. with its geographical sub-species. is. for a crop like that of the pouter. but not one has become wild or feral. Hence the supposed aboriginal stocks must either still exist in the countries where they were originally domesticated. colouring. and in most parts of their structure. and good flyers. considering their size. and these have not any of the characters of the domestic breeds. Moreover. that is. it must be assumed that at least seven or eight species were so thoroughly domesticated in ancient times by half-civilized man. An argument of great weight.

the mongrels were dusky and mottled. although no other existing species is thus coloured and marked. and the mongrels were black. or at most within a score.Some facts in regard to the colouring of pigeons well deserve consideration. brown and mottled. To give one instance out of several which I have observed: I crossed some white fantails. The rock-pigeon is of a slaty-blue. and they produced a bird of as beautiful a blue colour. so that in each separate breed there might be a tendency to revert to the very same colours and markings. been crossed by the rock-pigeon: I say within a dozen or twenty generations. the wings chequered with black. as in each succeeding generation there will be less of the foreign blood. 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. intermedia of Strickland. The tail has a terminal dark bar. These several marks do not occur together in any other species of the whole family. and barred and white-edged tail-feathers. with the white loins. I also crossed a barb with a spot. for no instance is known of crossed descendants reverting to an ancestor of foreign blood. Moreover. even to the white edging of the outer tail-feathers. besides the two black bars. have. that all the several imagined aboriginal stocks were coloured and marked like the rock-pigeon. has this part bluish. and which notoriously breeds very true. that each breed. and there is a tendency in the breed to revert to a character which was lost during 33 . Some semi-domestic breeds. as any wild rock-pigeon! We can understand these facts. secondly. of generations. taking thoroughly well-bred birds. Now. we must make one of the two following highly improbable suppositions. and some truly wild breeds. in every one of the domestic breeds. with the outer feathers externally edged at the base with white. Or. sometimes concur perfectly developed. when birds belonging to two or more distinct breeds are crossed. which is a white bird with a red tail and red spot on the forehead. has within a dozen. even the purest. C. double black wing-bar. but when there has been no cross. The wings have two black bars. none of which are blue or have any of the above-specified marks. but the Indian sub-species. all the above marks. with white loins. if all the domestic breeds are descended from the rock-pigeon. But if we deny this. removed by a greater number of generations. first. Either. which breed very true. the mongrel offspring are very apt suddenly to acquire these characters. 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. I then crossed one of the mongrel barb-fantails with a mongrel barb-spot. on the well-known principle of reversion to ancestral characters.

livia has been found capable of domestication in Europe and in India. that although an English carrier or a short-faced tumbler differs immensely in certain characters from the rock-pigeon. this tendency. From these several reasons. Lastly. namely. the hybrids or mongrels from between all the breeds of the pigeon are perfectly fertile. Secondly. more especially those brought from distant countries. between them and the rock-pigeon. an almost perfect series. In favour of this view. From the history of the dog. this conclusion is probably quite correct. and fantails now are. and their not having become anywhere feral—these species presenting certain very abnormal characters. and that it agrees in habits and in a great number of points of structure with all the domestic breeds. that the wild C. pouters. I may add. but not with all the breeds. the mongrel offspring being perfectly fertile—from these several reasons. Now. firstly. may be transmitted undiminished for an indefinite number of generations. and the number of tail-feathers in the fantail. and of some other domestic animals. 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. 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. But to extend it so far as to suppose that species. seems to me rash in the extreme. Some authors believe that longcontinued domestication eliminates this strong tendency to sterility in species. hardly any cases have been ascertained with certainty of hybrids from two quite distinct species of animals being perfectly fertile. on the most distinct breeds. both when kept pure and when crossed—and lastly. those characters which are mainly distinctive of each breed are in each eminently variable. as I can state from my own observations.pigeon or Columba livia with its geographical sub-species. These two distinct cases of reversion are often confounded together by those who have written on inheritance. so we can in some other cases. and the 34 . as compared with all other Columbidae. purposely made. for all that we can see to the contrary. Thirdly. taken together. if applied to species closely related to each other. the shortness of that of the tumbler. inter se. we may safely conclude that all our domestic breeds are descended from the rock. for instance. should yield offspring perfectly fertile.some former generation. we can make. the wattle and length of beak of the carrier. tumblers. aboriginally as distinct as carriers.

as I have asked. will likewise be obvious when we treat of selection. because when I first kept pigeons and watched the several kinds. They have been domesticated for thousands of years in several quarters of the world. never less than 20. I have never met a pigeon. and he will laugh you to scorn. or poultry. a celebrated raiser of Hereford cattle. for instance a Ribston-pippin or 35 . or both from a common parent. immense prices were given for pigeons. as any naturalist could in coming to a similar conclusion in regard to the many species of finches. I felt fully as much difficulty in believing that since they had been domesticated they had all proceeded from a common parent. namely. about 3000 B. but Mr. It is also a most favourable circumstance for the production of distinct breeds. shows how utterly he disbelieves that the several sorts. that nearly all the breeders of the various domestic animals and the cultivators of plants.stock." Pigeons were much valued by Akber Khan in India. yet quite insufficient. about the year 1600.C. are firmly convinced that the several breeds to which each has attended. see how it is that the several breeds so often have a somewhat monstrous character. also. that they can reckon up their pedigree and race. well knowing how truly they breed. In the time of the Romans.explanation of this fact will be obvious when we treat of selection. as we hear from Pliny. The paramount importance of these considerations in explaining the immense amount of variation which pigeons have undergone. or duck. in nature. which method was never practised before. Fourthly. as was pointed out to me by Professor Lepsius." About this same period the Dutch were as eager about pigeons as were the old Romans. with whom I have conversed. Ask. Van Mons. has improved them astonishingly. continues the courtly historian. or other groups of birds. or rabbit fancier." and. by crossing the breeds. in his treatise on pears and apples. or whose treatises I have read. "His Majesty. pigeons have been watched and tended with the utmost care. We shall then. who was not fully convinced that each main breed was descended from a distinct species. whether his cattle might not have descended from Long-horns. One circumstance has struck me much. I have discussed the probable origin of domestic pigeons at some. the earliest known record of pigeons is in the fifth Aegyptian dynasty. are descended from so many aboriginally distinct species.. Birch informs me that pigeons are given in a bill of fare in the previous dynasty.000 pigeons were taken with the court. that male and female pigeons can be easily mated for life. "nay. "The monarchs of Iran and Turan sent him some very rare birds. length. and loved by many people. and thus different breeds can be kept together in the same aviary. they are come to this pass.

yet admit that many of our domestic races are descended from the same parents—may they not learn a lesson of caution. and though they well know that each race varies slightly. is simple: from long-continued study they are strongly impressed with the differences between the several races. when they deride the idea of species in a state of nature being lineal descendants of other species? 36 . May not those naturalists who. Innumerable other examples could be given. could ever have proceeded from the seeds of the same tree. yet they ignore all general arguments. The explanation. knowing far less of the laws of inheritance than does the breeder. and refuse to sum up in their minds slight differences accumulated during many successive generations. for they win their prizes by selecting such slight differences. and knowing no more than he does of the intermediate links in the long lines of descent.Codlin-apple. I think.

Breeders habitually speak of an animal's organisation as 37 . many botanists. and this is known to have been the case with the ancon sheep. we must. Some variations useful to him have probably arisen suddenly. when we compare the host of agricultural. It is certain that several of our eminent breeders have. man adds them up in certain directions useful to him. Principles Of Selection Anciently Followed. But when we compare the dray-horse and race-horse. and with the bantam so small and elegant. orchard. but he would be a bold man who would account by such agencies for the differences between a dray and racehorse. Some effect may be attributed to the direct and definite action of the external conditions of life. look further than to mere variability. and flower-garden races of plants. And Their Effects Let us now briefly consider the steps by which domestic races have been produced. which can not be rivalled by any mechanical contrivance. the various breeds of sheep fitted either for cultivated land or mountain pasture. believe that the fuller's teasel. with its hooks. The great power of this principle of selection is not hypothetical. even within a single lifetime. or so beautiful in his eyes. with the wool of one breed good for one purpose. for instance. with other breeds so little quarrelsome. each good for man in different ways. is only a variety of the wild Dipsacus. In order fully to realise what they have done it is almost necessary to read several of the many treatises devoted to this subject. indeed. The key is man's power of accumulative selection: nature gives successive variations. One of the most remarkable features in our domesticated races is that we see in them adaptation. a carrier and tumbler pigeon. when we compare the game-cock. and to inspect the animals. with "everlasting layers" which never desire to sit. most useful to man at different seasons and for different purposes. in many cases. or by one step. modified to a large extent their breeds of cattle and sheep. not indeed to the animal's or plant's own good. a greyhound and bloodhound. We can not suppose that all the breeds were suddenly produced as perfect and as useful as we now see them. either from one or from several allied species. when we compare the many breeds of dogs. I think. we know that this has not been their history. culinary.5. so pertinacious in battle. and that of another breed for another purpose. In this sense he may be said to have made for himself useful breeds. So it has probably been with the turnspit dog. and some to habit. but to man's use or fancy. and this amount of change may have suddenly arisen in a seedling. the dromedary and camel.

Youatt. What English breeders have actually effected is proved by the enormous prices given for animals with a good pedigree. by means of which he may summon into life whatever form and mould he pleases. The improvement is by no means generally due to crossing different breeds." Lord Somerville. but to change it altogether. to give a very trifling instance. thus. if he wants any of these qualities. and may make great improvements. says: "It would seem as if they had chalked out upon a wall a form perfect in itself. he will assuredly fail. speaking of what breeders have done for sheep. like a picture by a connoisseur. The same principles are followed by horticulturists. that men follow it as a trade: the sheep are placed on a table and are studied. which they can model almost as they please. not only to modify the character of his flock. If selection consisted merely in separating some very distinct variety and breeding from it. of differences absolutely inappreciable by an uneducated eye—differences which I for one have vainly attempted to appreciate. and these have been exported to almost every quarter of the world. If I had space I could quote numerous passages to this effect from highly competent authorities. and then had given it existence. he will succeed. If gifted with these qualities. during successive generations. the principle would be so obvious as hardly to be worth notice. but the variations are here often more abrupt. No one supposes that our choicest productions have been produced by a single variation from the aboriginal stock. Few would readily believe in the natural capacity and years of practice requisite to become even a skilful pigeon-fancier. the steadily 38 . and the sheep are each time marked and classed. And when a cross has been made. Not one man in a thousand has accuracy of eye and judgment sufficient to become an eminent breeder." In Saxony the importance of the principle of selection in regard to merino sheep is so fully recognised.something plastic. and who was himself a very good judge of animals. but its importance consists in the great effect produced by the accumulation in one direction. this is done three times at intervals of months. We have proofs that this is not so in several cases in which exact records have been kept. and he studies his subject for years. except sometimes among closely allied sub-breeds. speaks of the principle of selection as "that which enables the agriculturist. all the best breeders are strongly opposed to this practice. the closest selection is far more indispensable even than in ordinary cases. so that the very best may ultimately be selected for breeding. who was probably better acquainted with the works of agriculturalists than almost any other individual. and devotes his lifetime to it with indomitable perseverance. It is the magician's wand.

in comparison with the flowers of the same varieties. will produce races differing from each other chiefly in these characters. and yet the flowers present very slight differences. and how alike the leaves. how unlike the flowers of the heartsease are. and this may be compared to the "roguing" of plants by nurserymen. in which the full importance of the principle is acknowledged. In rude and barbarous periods of English history choice animals were often imported. and hairiness. and laws were passed to prevent their exportation: the destruction of horses under a certain size was ordered. and the result has been. or whatever part is valued. pods. in a corresponding degree. When a race of plants is once pretty well established.increasing size of the common gooseberry may be quoted. In regard to plants. the flowers. in comparison with the leaves and flowers of the same set of varieties. the diversity of leaves. shape. when the flowers of the present day are compared with drawings made only twenty or thirty years ago. this is hardly ever—I speak after careful observation—perhaps never. how much the fruit of the different kinds of gooseberries differ in size. it has certainly been more attended to of late years. the seedraisers do not pick out the best plants. either in the leaves. and pull up the "rogues. as a general rule. in fact. We see an astonishing improvement in many florists' flowers. but. I could give several references to works of high antiquity. the importance of which should never be overlooked. and how extremely alike the flowers. the case. it cannot be doubted that the continued selection of slight variations. See how different the leaves of the cabbage are. With animals this kind of selection is. It may be objected that the principle of selection has been reduced to methodical practice for scarcely more than three-quarters of a century. by comparing the diversity of flowers in the different varieties of the same species in the flower-garden. But it is very far from true that the principle is a modern discovery. likewise followed. The principle of selection I find distinctly given in an ancient Chinese 39 . and the diversity of fruit of the same species in the orchard. rapid and important. there is another means of observing the accumulated effects of selection—namely. for hardly any one is so careless as to breed from his worst animals. or tubers. in the kitchen-garden. or the fruit. colour. and many treatises have been published on the subject." as they call the plants that deviate from the proper standard. will ensure some differences. It is not that the varieties which differ largely in some one point do not differ at all in other points. but merely go over their seedbeds. The law of correlated variation.

and is now attended to by the lowest savages. The savages in South Africa match their draught cattle by colour. had attention not been paid to breeding. have been a strange fact. as is attested by passages in Pliny. From passages in Genesis. Some of these facts do not show actual selection. and they formerly did so. It would.encyclopaedia. for the inheritance of good and bad qualities is so obvious. it is clear that the colour of domestic animals was at that early period attended to. as do some of the Esquimaux their teams of dogs. indeed. but they show that the breeding of domestic animals was carefully attended to in ancient times. 40 . to improve the breed. Savages now sometimes cross their dogs with wild canine animals. Livingstone states that good domestic breeds are highly valued by the negroes in the interior of Africa who have not associated with Europeans. Explicit rules are laid down by some of the Roman classical writers.

but what concerns us is. Unconscious Selection At the present time. etc. Borrow has not seen. that the change has been effected unconsciously and gradually. is more important. 41 . Nevertheless we may infer that this process. did greatly modify. In some cases. to make a new strain or sub-breed. in the same way as Bakewell. the forms and qualities of their cattle. eminent breeders try by methodical selection. which may be called unconscious. and which results from every one trying to possess and breed from the best individual animals. by this very same process. a man who intends keeping pointers naturally tries to get as good dogs as he can. Mr. By comparing the accounts given in various old treatises of the former and present state of carrier and tumbler pigeons in Britain. which may serve for comparison. English racehorses have come to surpass in fleetness and size the parent Arabs. Some highly competent authorities are convinced that the setter is directly derived from the spaniel. By a similar process of selection. and has probably been slowly altered from it. Collins. for our purpose. Lord Spencer and others have shown how the cattle of England have increased in weight and in early maturity. It is known that the English pointer has been greatly changed within the last century. individuals of the same breed exist in less civilised districts. and by careful training. Thus. as I am informed by him. any native dog in Spain like our pointer. are favoured in the weights which they carry. even during their lifetimes. only carried on more methodically. 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. or but little changed. been chiefly effected by crosses with the foxhound. and Persia. continued during centuries.. we can trace the stages through which they have insensibly passed. so that the latter. but he has no wish or expectation of permanently altering the breed. where the breed has been less improved. it is believed. But. with a distinct object in view. would improve and modify any breed.6. and in this case the change has. India. however. though the old Spanish pointer certainly came from Spain. superior to anything of the kind in the country. unchanged. and come to differ so greatly from the rock-pigeon. There is reason to believe that King Charles' spaniel has been unconsciously modified to a large extent since the time of that monarch. and afterwards breeds from his own best dogs. by the regulations for the Goodwood Races. a form of selection. compared with the stock formerly kept in this country. and yet so effectually that.

and so 42 . and. No one would ever expect to get a firstrate heartsease or dahlia from the seed of a wild plant. the production of the distinct strains. The two flocks of Leicester sheep kept by Mr. though he might succeed from a poor seedling growing wild. for any special purpose. by their killing and devouring their old women. No one would expect to raise a first-rate melting pear from the seed of a wild pear. has been followed almost unconsciously. when compared with the older varieties or with their parent-stocks. sowing its seeds. We see the value set on animals even by the barbarians of Tierra del Fuego. as of less value than their dogs. Burgess. rose. 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. It has consisted in always cultivating the best known variety. Bakewell for upwards of fifty years. if it had come from a garden-stock. pelargonium. may plainly be recognised in the increased size and beauty which we now see in the varieties of the heartsease. In plants the same gradual process of improvement through the occasional preservation of the best individuals. as far as the final result is concerned. when a slightly better variety chanced to appear. in times of dearth. whether or not sufficiently distinct to be ranked at their first appearance as distinct varieties. or even wished. appears. as Mr. though cultivated in classical times. and other plants. in so far that the breeders could never have expected. would be carefully preserved during famines and other accidents. yet any one animal particularly useful to them. to produce the result which ensued—namely. to which savages are so liable. 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. from Pliny's description." If there exist savages so barbarous as never to think of the inherited character of the offspring of their domestic animals. to have been a fruit of very inferior quality. dahlia. and. and whether or not two or more species or races have become blended together by crossing. There is not a suspicion existing in the mind of any one at all acquainted with the subject that the owner of either of them has deviated in any one instance from the pure blood of Mr. Youatt remarks.Youatt gives an excellent illustration of the effects of a course of selection which may be considered as unconscious. Bakewell's flock. so that in this case there would be a kind of unconscious selection going on. Buckley and Mr. The pear. selecting it. and such choice animals would thus generally leave more offspring than the inferior ones. but the art has been simple. "Have been purely bred from the original stock of Mr.

onwards. and likewise their differences being so great in external characters." as will hereafter be more fully explained. It is not that these countries. or a 43 . individuals of the same species. This. have more of the character of true species than the varieties kept in civilised countries. On the view here given of the important part which selection by man has played. how it is that our domestic races show adaptation in their structure or in their habits to man's wants or fancies. excepting on variations which are first given to him in some slight degree by nature. so rich in species. though we owe our excellent fruit in some small degree to their having naturally chosen and preserved the best varieties they could anywhere find. having slightly different constitutions or structure. has afforded us a single plant worth culture. and indeed he rarely cares for what is internal. do not by a strange chance possess the aboriginal stocks of any useful plants. and relatively so slight in internal parts or organs. who cultivated the best pears which they could procure. further understand the frequently abnormal character of our domestic races. And in two countries very differently circumstanced. We can. and thus by a process of "natural selection. or only with much difficulty. as has been remarked by some authors. two sub-breeds might be formed. we can understand how it is that neither Australia. any deviation of structure excepting such as is externally visible. the wild parentstocks of the plants which have been longest cultivated in our flower and kitchen gardens. explains. it should not be overlooked that they almost always have to struggle for their own food. the well-known fact. that in a number of cases we cannot recognise. 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. never thought what splendid fruit we should eat. partly explains why the varieties kept by savages. Man can hardly select. would often succeed better in the one country than in the other. In regard to the domestic animals kept by uncivilised man. it becomes at once obvious. But the gardeners of the classical period. I think. nor any other region inhabited by quite uncivilised man. perhaps. at least during certain seasons. and therefore do not know. 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. 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. A large amount of change. He can never act by selection. thus slowly and unconsciously accumulated. as I believe. the Cape of Good Hope.

like a dialect of a language. selection. 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. or takes more care than usual in matching his best animals. that we know hardly anything about the origin or history of any of our domestic breeds. be judged of by the value which is now set on them. they will spread more widely. after several breeds have fairly been established. as it is not one of the points of the breed. that most fleeting of characters. A man preserves and breeds from an individual with some slight deviation of structure. in fact. which differ only in colour. It is known that with pigeons many slight variations now occasionally appear. But. in most cases. In semi-civilised countries. I have no doubt. 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. The common goose has not given rise to any marked varieties. But they will as yet hardly have a distinct name. As soon as the points of value are once 44 . in which as many as seventeen tailfeathers have been counted. and the improved animals slowly spread in the immediate neighbourhood. the spreading of a new subbreed will be a slow process. the more likely it would be to catch his attention. namely. and the more abnormal or unusual any character was when it first appeared. like the present Java fantail. The man who first selected a pigeon with a slightly larger tail. and from being only slightly valued. and will then probably first receive a provincial name. their history will have been disregarded. But to use such an expression as trying to make a fantail is. with little free communication. but these are rejected as faults or deviations from the standard of perfection in each breed. in one's own possession. have lately been exhibited as distinct at our poultry-shows. utterly incorrect. and will be recognised as something distinct and valuable. and it is in human nature to value any novelty. however slight. Perhaps the parent bird of all fantails had only fourteen tailfeathers somewhat expanded.pouter till he saw a pigeon with a crop of somewhat unusual size. or like individuals of other and distinct breeds. These views appear to explain what has sometimes been noticed. never dreamed what the descendants of that pigeon would become through long-continued. partly unconscious and partly methodical. and thus improves them. When further improved by the same slow and gradual process. hence the Toulouse and the common breed. can hardly be said to have a distinct origin. a breed. Nor must the value which would formerly have been set on any slight differences in the individuals of the same species.

45 . and insensible changes.acknowledged. the principle. as I have called it. according to the state of civilisation of the inhabitants—slowly to add to the characteristic features of the breed. whatever they may be. But the chance will be infinitely small of any record having been preserved of such slow. of unconscious selection will always tend—perhaps more at one period than at another. varying. as the breed rises or falls in fashion—perhaps more in one district than in another.

and raised seedlings from them. with extreme care. to man's power of selection. When the individuals are scanty all will be allowed to breed. I have seen it gravely remarked. to allow of the accumulation of a large amount of modification in almost any desired direction. from keeping large stocks of the same plant. and this is a great convenience to the fancier. the chance of their appearance will be much increased by a large number of individuals being kept. with respect to the sheep of part of Yorkshire.7. and again picked out the best seedlings and bred from them. are generally far more successful than amateurs in raising new and valuable varieties. and this circumstance must have largely favoured the formation of new breeds. and are mostly IN SMALL LOTS. they never can be improved. Wandering savages or the inhabitants of open plains rarely possess more than one breed of the same species. as gardeners picked out individual plants with slightly larger. earlier. But probably the most important element is that the animal or plant should be so highly valued by man. favourable or the reverse. in a country which is already stocked with other races. As soon. 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. Unless such attention be paid nothing can be effected. not that mere individual differences are not amply sufficient. nurserymen. however. Hence number is of the highest importance for success. that it was most fortunate that the strawberry began to vary just when gardeners began to attend to this plant. With animals. but the slight varieties had been neglected. that the closest attention is paid to even the slightest deviations in its qualities or structure. facility in preventing crosses is an important element in the formation of new races—at least. as freely giving the materials for selection to work on. Pigeons can be mated for life. On this principle Marshall formerly remarked. "As they generally belong to poor people. A high degree of variability is obviously favourable." On the other hand. A large number of individuals of an animal or plant can be reared only where the conditions for its propagation are favourable. for thus many races may be improved and kept true. 46 . In this respect enclosure of the land plays a part. But as variations manifestly useful or pleasing to man appear only occasionally. whatever their quality may be. and this will effectually prevent selection. though mingled in the same aviary. Circumstances Favourable To Man's Power Of Selection I will now say a few words on the circumstances. or better fruit. No doubt the strawberry had always varied since it was cultivated.

So with plants. from the difficulty in pairing them. and so it is with colour. as when killed they serve for food. For instance. in donkeys. cats. It would be equally rash to assert that characters now increased to their utmost limit. the donkey. With respect to fleetness. in geese. the seeds of the different 47 . could not. etc. goose. seems to have a singularly inflexible organisation.Pigeons. I may add. we rarely see a distinct breed long kept up. and this implies variation. more than do the distinct species of the same genera. as I have elsewhere described. as this will be determined by the friction to be overcome. as Mr. peacock. and the power of contraction in the muscular fibres. from only a few being kept by poor people. than any two natural species belonging to the same genus. Wallace has remarked with much truth. such breeds as we do sometimes see are almost always imported from some other country. can be propagated in great numbers and at a very quick rate. in peacocks. though it has varied to a slight extent. but the goose. and more especially from no pleasure having been felt in the display of distinct breeds.. and. which man has attended to and selected. and a dray-horse is comparably stronger. which depends on many bodily characters. a limit will be at last reached. from not being very easily reared and a large stock not kept. But what concerns us is that the domestic varieties of the same species differ from each other in almost every character. food and feathers. the weight of the body to be carried. from being valuable only for two purposes. It would be somewhat rash to assert that the limit has been attained in any one case. for almost all our animals and plants have been greatly improved in many ways within a recent period. Eclipse was far fleeter. although so much valued by women and children. and inferior birds may be freely rejected. and can never afterward be exceeded. Isidore Geoffroy St. and probably with the length of hair. On the other hand. may be attributed in main part to selection not having been brought into play: in cats. there must be a limit to the fleetness of any terrestrial animal. for recently in certain parts of Spain and of the United States this animal has been surprisingly modified and improved by careful selection. after remaining fixed for many centuries. No doubt. again vary under new conditions of life. Some authors have maintained that the amount of variation in our domestic productions is soon reached. Although I do not doubt that some domestic animals vary less than others. and little attention paid to their breeding. can not be easily matched. Hilaire has proved this in regard to size. from their nocturnal rambling habits. yet the rarity or absence of distinct breeds of the cat. under the conditions to which it is exposed when domesticated.

under all circumstances. but more efficiently. of which correlated growth is probably the most important. Over all these causes of change. In some cases the intercrossing of aboriginally distinct species appears to have played an important part in the origin of our breeds. the importance of crossing is immense. 48 . and still more strongly with the melon. When several breeds have once been formed in any country. buds. The same remark holds good in regard to the fruit of the several varieties of the plum. or unconsciously and slowly. Some. With plants which are temporarily propagated by cuttings. and indirectly by affecting the reproductive system. perhaps a great. Variability is governed by many unknown laws. has. may be attributed to the definite action of the conditions of life. largely aided in the formation of new sub-breeds. with the aid of selection. both by acting directly on the organisation. To sum up on the origin of our domestic races of animals and plants. and the sterility of hybrids. The greater or less force of inheritance and reversion determine whether variations shall endure. their occasional intercrossing. but plants not propagated by seed are of little importance to us. It is not probable that variability is an inherent and necessary contingent. but the importance of crossing has been much exaggerated. etc. as well as in many other analogous cases.. The final result is thus rendered infinitely complex. whether applied methodically and quickly. for their endurance is only temporary. the accumulative action of selection.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. for the cultivator may here disregard the extreme variability both of hybrids and of mongrels. but how much we do not know. effect may be attributed to the increased use or disuse of parts. seems to have been the predominant power. no doubt. both in regard to animals and to those plants which are propagated by seed. Something. Changed conditions of life are of the highest importance in causing variability.

and common species. related to each other. 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. much diffused. 49 . and in having restricted ranges.Chapter 2 Variation Under Nature Variability — Individual differences — Doubtful species — Wide ranging. but unequally.

but they graduate into varieties. It may be doubted whether sudden and considerable deviations of structure. are ever permanently propagated in a state of nature. If monstrous forms of this kind ever do appear in a state of nature and are capable of reproduction (which is not always the case). but these I shall reserve for a future work. Generally the term includes the unknown element of a distinct act of creation. more especially with plants. Thus pigs have occasionally been born with a sort of proboscis. and "variations" in this sense are supposed not to be inherited. as they occur rarely and singly. though it can rarely be proved. but I have as yet failed to find. Some authors use the term "variation" in a technical sense. Before applying the principles arrived at in the last chapter to organic beings in a state of nature. 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. Under domestication monstrosities sometimes occur which resemble normal structures in widely different animals. or dwarfed plants on Alpine summits. Nor shall I here discuss the various definitions which have been given of the term species. yet every naturalist knows vaguely what he means when he speaks of a species. their preservation would depend on unusually favourable circumstances. By a monstrosity I presume is meant some considerable deviation of structure. and if any wild species of the same genus had naturally possessed a proboscis. after diligent search. it might have been argued that this had appeared as a monstrosity. or the thicker fur of an animal from far northwards. No one definition has satisfied all naturalists. generally injurious.1. We have also what are called monstrosities. but who can say that the dwarfed condition of shells in the brackish waters of the Baltic. a long catalogue of dry facts ought to be given. 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. To treat this subject properly. as that a complex machine should have been invented by man in a perfect state. and these alone bear on the question. cases of monstrosities resembling normal structures in nearly allied forms. as implying a modification directly due to the physical conditions of life. They 50 . such as we occasionally see in our domestic productions. or not useful to the species. The term "variety" is almost equally difficult to define.

would. and thus their abnormal character would almost inevitably be lost. 51 . during the first and succeeding generations cross with the ordinary form. also. But I shall have to return in a future chapter to the preservation and perpetuation of single or occasional variations.

may be called individual differences. There is one point connected with individual differences which is extremely perplexing: I refer to those genera which have been called "protean" or "polymorphic. but under any other point of view many instances assuredly can be given. under this point of view. These individual differences are of the highest importance for us. yet Sir J. as I have collected. it might have been thought that changes of this nature could have been effected only by slow degrees. from being observed in the individuals of the same species inhabiting the same confined locality. No one supposes that all the individuals of the same species are cast in the same actual mould. and compare them in many specimens of the same species. sometimes vary in the individuals of the same species. 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. and that there are not many men who will laboriously examine internal and important organs. Individual Differences The many slight differences which appear in the offspring from the same parents. which he could collect on good authority. during a course of years. whether viewed under a physiological or classificatory point of view. and. This philosophical naturalist. by a long catalogue of facts. Lubbock has shown a degree of variability in these main nerves in Coccus. for these same authors practically rank those parts as important (as some few naturalists have honestly confessed) which do not vary. no instance will ever be found of an important part varying. I may add. Authors sometimes argue in a circle when they state that important organs never vary. in the same manner as man accumulates in any given direction individual differences in his domesticated productions. which may almost be compared to the irregular branching of the stem of a tree. and they thus afford materials for natural selection to act on and accumulate. has also shown that the muscles in the larvae of certain insects are far from uniform. but I could show. that parts which must be called important." in which species present an inordinate 52 . as must be familiar to every one. or which it may be presumed have thus arisen.2. These individual differences generally affect what naturalists consider unimportant parts. It should be remembered that systematists are far from being pleased at finding variability in important characters. I am convinced that the most experienced naturalist would be surprised at the number of the cases of variability. even in important parts of structure. for they are often inherited.

several genera of insects. describes a certain butterfly which presents in the same island a great range of varieties connected by intermediate links. hardly two naturalists agree whether to rank them as species or as varieties. and of Brachiopod shells. the castes are connected together by finely graduated varieties. and Hieracium among plants. both with animals and plants. I am inclined to suspect that we see. cases of dimorphism and trimorphism. polymorphic in other countries. independently of variation. Genera which are polymorphic in one country seem to be. and the other has antennae much more abundantly furnished with smelling-hairs. in the two or three castes of sterile females or workers among insects.amount of variation. Individuals of the same species often present. the several worker-castes are generally quite distinct. There are. are not now connected by intermediate gradations. but in some cases. regularly appear under two or even three conspicuously distinct forms. and the extreme links of the chain closely resemble the two forms of an allied dimorphic species inhabiting another part of the Malay Archipelago. great differences of structure. Wallace. Although in most of these cases. at least in some of these polymorphic genera. with a few exceptions. Thus also with ants. the male of a Tanais regularly occurs under two distinct forms. the two or three forms. 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. one of these has strong and differently shaped pincers. at former periods of time. it is possible that they were once thus connected. These facts are very perplexing. and likewise. Thus. Mr. as in the two sexes of various animals. Fritz Muller has described analogous but more extraordinary cases with the males of certain Brazilian Crustaceans: thus. in the Malayan Archipelago. With respect to many of these forms. and which consequently have not been seized on and rendered definite by natural selection. not connected by intermediate varieties. has shown that the females of certain species of butterflies. for instance. In most polymorphic genera some of the species have fixed and definite characters. for they seem to show that this kind of variability is independent of the conditions of life. variations which are of no service or disservice to the species. So it is. who has lately called attention to the subject. Rosa. as hereafter to be explained. We may instance Rubus. Wallace. both with animals and plants. and in the immature and larval states of many of the lower animals. as is known to every one. judging from Brachiopod shells. as we shall hereafter see. also. and that an hermaphrodite plant 53 . Mr. with some dimorphic plants. as I have myself observed.

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. bearing three different kinds of females and three or even six different kinds of males.capsule three distinct hermaphrodite forms.should produce from the same seed. 54 .

Practically. but sometimes the one first described as the species. even when they are closely connected by intermediate links. We must. or are so closely linked to them by intermediate gradations. one form is ranked as a variety of another. which are generally considered as varieties. but which have all been ranked by botanists as species. in many cases. as far as we know. and the other as the variety. nor will the commonly assumed hybrid nature of the intermediate forms always remove the difficulty. Mr. for as long. drawn up by different botanists. however. or of the United States. when a naturalist can unite by means of intermediate links any two forms. But cases of great difficulty. for few well-marked and wellknown varieties can be named which have not been ranked as species by at least some competent judges. Watson. whereas Mr. H. That varieties of this doubtful nature are far from uncommon cannot be disputed. sometimes arise in deciding whether or not to rank one form as a variety of another. Hence. of France. he treats the one as a variety of the other. Mr.C. that naturalists do not like to rank them as distinct species. Babington gives 251 species. are in several respects the most important for us. and here a wide door for the entry of doubt and conjecture is opened. which I will not here enumerate. Compare the several floras of Great Britain. and by another as mere varieties. the opinion of naturalists having sound judgment and wide experience seems the only guide to follow. and in making this list he has omitted many trifling varieties. as have good and true species.3. and see what a surprising number of forms have been ranked by one botanist as good species. but because analogy leads the observer to suppose either that they do now somewhere exist. however. Bentham 55 . We have every reason to believe that many of these doubtful and closely allied forms have permanently retained their characters for a long time. Doubtful Species The forms which possess in some considerable degree the character of species. Under genera. in determining whether a form should be ranked as a species or a variety. decide by a majority of naturalists. but which are so closely similar to other forms. In very many cases. including the most polymorphic forms. has marked for me 182 British plants. but which nevertheless have been ranked by some botanists as species. to whom I lie under deep obligation for assistance of all kinds. ranking the most common. or may formerly have existed. and he has entirely omitted several highly polymorphic genera. not because the intermediate links have actually been found.

I was much struck how entirely vague and arbitrary is the distinction between species and varieties. but are common in separated areas. they are almost universally ranked by naturalists as true species." Lastly. but as they do not differ from each other by strongly marked and important characters. ranked by one zoologist as a species and by another as a variety. namely. but as they are distinguished from each other by a greater amount of difference than that between the local forms and sub-species. Many years ago. The geographical races or sub-species are local forms completely fixed and isolated. now generally regarded as varieties. inhabiting the islands of the great Malayan Archipelago. one with another. Several experienced ornithologists consider our British red grouse as only a strongly marked race of a Norwegian 56 . have been ranked by one eminent naturalist as undoubted species. and as true representative species. when comparing. which differ very slightly from each other. "There is no possible test but individual opinion to determine which of them shall be considered as species and which as varieties. in several valuable papers on the various animals. and seeing others compare.gives only 112—a difference of 139 doubtful forms! Among animals which unite for each birth. representative species fill the same place in the natural economy of each island as do the local forms and sub-species. the differences are seen to be so slight and graduated that it is impossible to define or describe them. no certain criterion can possibly be given by which variable forms. Wallace. The local forms are moderately constant and distinct in each separate island. especially on the Lepidoptera. but when all from the several islands are compared together. as geographical races or sub-species. but which have been ranked as species by some zoologists. the birds from the closely neighbouring islands of the Galapagos Archipelago. as variable forms. though at the same time the extreme forms are sufficiently distinct. as they are often called. as local forms. local forms. Even Ireland has a few animals. doubtful forms. shows that they may be classed under four heads. How many of the birds and insects in North America and Europe. and which are highly locomotive. On the islets of the little Madeira group there are many insects which are characterized as varieties in Mr. or. The first or variable forms vary much within the limits of the same island. and by another as varieties. sub species and representative species can be recognised. Wollaston's admirable work. Nevertheless. and with those from the American mainland. but which would certainly be ranked as distinct species by many entomologists. can rarely be found within the same country. geographical races! Mr.

or in the nature of their secretions. Walsh ranks the forms which it may be supposed would freely intercross. In several cases. have been observed thus to differ in a slight degree. When the differences are rather more strongly marked.species. implying and assuming a separate act of 57 . however. or between the several islets of these small archipelagos. B. The term species thus comes to be a mere useless abstraction. The naturalist thus loses his best guide in determining whether to rank doubtful forms as varieties or species. Most vegetable-feeding insects live on one kind of plant or on one group of plants. even if he can do so for himself. which of these Phytophagic forms ought to be called species and which varieties. Walsh to present in their larval or mature state. When. insects found living on different plants. but then these same naturalists rank the slightest difference as of specific value. the forms are ranked by all entomologists as good species. Mr. on the other hand. as species. Some few naturalists maintain that animals never present varieties. a distinguished entomologist of the United States. has described what he calls Phytophagic varieties and Phytophagic species. or Madeira. an animal or plant ranges over the same continent. and presents different forms in the different areas. some feed indiscriminately on many kinds. and when both sexes and all ages are affected. which inhabit distinct continents or islands. will suffice if that between America and Europe is ample. both males and females. as varieties. or in two geological formations. have been observed by Mr. As the differences depend on the insects having long fed on distinct plants. A wide distance between the homes of two doubtful forms leads many naturalists to rank them as distinct species. though constant differences in colour. but what distance. or the Canaries. and these are then degraded to the rank of varieties. But no observer can determine for another. they believe that two distinct species are hidden under the same dress. or in both states. and when the same identical form is met with in two distant countries. and those which appear to have lost this power. or inhabits many islands in the same archipelago. In some instances the males alone. in other instances. be sufficient? Mr. but do not in consequence vary. This likewise necessarily occurs with closely allied organisms. whereas the greater number rank it as an undoubted species peculiar to Great Britain. there is always a good chance that intermediate forms will be discovered which will link together the extreme states. slight. Walsh. size. it has been well asked.D. it cannot be expected that intermediate links connecting the several forms should now be found. will that between Europe and the Azores.

resemble species so completely in character that they have been thus ranked by other highly competent judges. yet a German author makes more than a dozen species out of forms. 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. and estimates numerically the relative frequency of the variations. will often be ranked by some authors as species. After this discussion. Look at the common oak. etc. as Asa Gray has remarked in commenting on this memoir. or could have worked on them with more zeal and sagacity. in many cases. who repeat that the greater part of our species are clearly limited. I may here allude to a remarkable memoir lately published by A. He first gives in detail all the many points of structure which vary in the several species. It is certain that many forms. is vainly to beat the air. But to discuss whether they ought to be called species or varieties. which are almost universally considered by other botanists to be varieties. will no doubt bring naturalists to agree how to rank doubtful forms. have been brought to bear in the attempt to determine their rank. Such characters are not of course of specific value.. considered by highly competent judges to be varieties. he emphatically remarks: "They are mistaken. how closely it has been studied. de Candolle. analogical variation. This seemed to be true. but they are. and that the doubtful species are in a feeble minority. Many of the cases of strongly marked varieties or doubtful species well deserve consideration. moreover. on the oaks of the whole world. from geographical distribution. varieties of it will almost universally be found recorded. 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. These varieties. He specifies above a dozen characters which may be found varying even on the same branch. and never found connected by intermediate states. Yet it must be confessed that it is in the best known countries that we find the greatest number of them. sometimes according to age or development.creation. hybridism. so long as a genus was 58 . sometimes without any assignable reason. the result of so much labour. I have been struck with the fact that if any animal or plant in a state of nature be highly useful to man. such as generally enter into specific definitions. for several interesting lines of argument. or from any cause closely attracts his attention. before any definition of these terms has been generally accepted. but space does not here permit me to discuss them. No one ever had more ample materials for the discrimination of the species. Close investigation.

It should be added that De Candolle no longer believes that species are immutable creations. at least. with the amount of difference in the forms which he is continually studying. which will be enumerated in his Prodromus as belonging to the oak family. But if his observations be widely extended he will in the end generally be able to make up his own mind. De Candolle admits that out of the 300 species. and this shows. were provisional. for he knows nothing of the amount and kind of variation to which the group is subject. When he comes to study allied forms brought from countries not now continuous. geographical botany and zoology. as Asa Gray again remarks. that is to say. for he will encounter a greater number of closely-allied forms. Just as we come to know them better. and he has little general knowledge of analogical variation in other groups and in other countries by which to correct his first impressions. just like the pigeon or poultry fancier before alluded to. for he will become impressed. sessiliflora and pubescens. and the truth of this admission will often be disputed by other naturalists.species. Thus Quercus robur has twenty-eight varieties. pedunculata. The forms which connect these three sub-species are comparatively rare. how very generally there is some variation. are not known strictly to fulfil the definition above given of a true species. at least two-thirds are provisional species. 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. in which case he cannot 59 . As he extends the range of his observations he will meet with more cases of difficulty. namely Q. "and the most accordant with the known facts in palaeontology. intermediate forms flow in. that is. all of which. excepting six. but he will succeed in this at the expense of admitting much variation. and its species were founded upon a few specimens. of anatomical structure and classification. but concludes that the derivative theory is the most natural one. and." 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. are clustered round three sub. His general tendency will be to make many species. and doubts as to specific limits augment. 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. Finally.imperfectly known." He also adds that it is the best known species which present the greatest number of spontaneous varieties and sub-varieties.

The passage from one stage of difference to another may. or both might co-exist. The term variety. but do not quite arrive at. as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. he will be compelled to trust almost entirely to analogy. to a set of individuals closely resembling each other. and the species as the variety. the passage from one stage of difference to another may be safely attributed to the cumulative action of natural selection. But we shall hereafter return to this subject. between sub-species and well-marked varieties. for convenience sake. and at the latter. which is given to less distinct and more fluctuating forms. If a variety were to flourish so as to exceed in numbers the parent species. again.hope to find intermediate links. be the simple result of the nature of the organism and of the different physical conditions to which it has long been exposed. as has been shown to be the case by Mr. It need not be supposed that all varieties or incipient species attain the rank of species. is also applied arbitrarily. or they may endure as varieties for very long periods. 60 . in many cases. as leading to sub-species. and to the effects of the increased use or disuse of parts. the forms which in the opinion of some naturalists come very near to. but whether this belief is justifiable must be judged by the weight of the various facts and considerations to be given throughout this work. but with respect to the more important and adaptive characters. or. Certainly no clear line of demarcation has as yet been drawn between species and sub-species—that is. or it might come to supplant and exterminate the parent species. and a series impresses the mind with the idea of an actual passage. Hence I look at individual differences. and that it does not essentially differ from the term variety. They may become extinct. And I look at varieties which are in any degree more distinct and permanent. hereafter to be explained. though of small interest to the systematist. again. Wollaston with the varieties of certain fossil land-shells in Madeira. in comparison with mere individual differences. as of the highest importance for us. and then to species. A well-marked variety may therefore be called an incipient species. and his difficulties will rise to a climax. and both rank as independent species. and with plants by Gaston de Saporta. it would then rank as the species. the rank of species. for the sake of convenience. or between lesser varieties and individual differences. as steps toward more strongly marked and permanent varieties. These differences blend into each other by an insensible series. From these remarks it will be seen that I look at the term species as one arbitrarily given.

as varieties. it should be understood that reference is made only to the forms which come into competition with each 61 . Hooker. hereafter to be discussed. or. might have been anticipated. incipient species. and to a certain extent from commonness). for. Watson.C. Alphonse de Candolle and others have shown that plants which have very wide ranges generally present varieties. Wide-Ranging. as did subsequently Dr. by tabulating all the varieties in several well-worked floras. and as they come into competition (which. And Common Species Vary Most Guided by theoretical considerations. oftenest give rise to varieties sufficiently well-marked to have been recorded in botanical works. is rather perplexing. and the species which are most widely diffused within their own country (and this is a different consideration from wide range. and are the most numerous in individuals—which oftenest produce well-marked varieties. H. soon convinced me that there were many difficulties. that is abound most in individuals. are the most diffused in their own country. to whom I am much indebted for valuable advice and assistance on this subject. as they are exposed to diverse physical conditions.4. however. though in some slight degree modified. necessarily have to struggle with the other inhabitants of the country. he thinks that the following statements are fairly well established. perhaps. the species which are already dominant will be the most likely to yield offspring. But my tables further show that. but Mr. And this. and the tables of the proportional numbers of the varying species. in any limited country. The whole subject. treated as it necessarily here is with much brevity. still inherit those advantages that enabled their parents to become dominant over their compatriots. as we shall hereafter see. which. I shall reserve for a future work the discussion of these difficulties. and examined the tables." "divergence of character. even in stronger terms. Hooker permits me to add that after having carefully read my manuscript." and other questions. In these remarks on predominence. as they may be called. the dominant species—those which range widely. the species which are the most common. and allusions cannot be avoided to the "struggle for existence. or. At first this seemed a simple task. Much Diffused. is a far more important circumstance) with different sets of organic beings. Dr. Hence it is the most flourishing. as I consider them. and this might have been expected. I thought that some interesting results might be obtained in regard to the nature and relations of the species which vary most. in order to become in any degree permanent.

With respect to the number of individuals or commonness of species.other. the comparison of course relates only to the members of the same group. and more especially to the members of the same genus or class having nearly similar habits of life. and more widely diffused. 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. 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. But if the conferva or parasitic fungus exceeds its allies in the above respects. it will then be dominant within its own class. which live under nearly the same conditions. 62 .

those including many species) being placed on one side.e. But so many causes tend to obscure this result. all those in the larger genera (i. than the species of the smaller genera. consequently. as a general rule. we might have expected to have found in the larger genera. into two nearly equal masses. but this seems to be connected with the nature of the stations inhabited by them. Where many species of a genus have been formed through variation. Where many large trees grow. and 63 . species of the same genus) have been formed. for the mere fact of many species of the same genus inhabiting any country. shows that there is something in the organic or inorganic conditions of that country favourable to the genus. there is no apparent reason why more varieties should occur in a group having many species. be divided into two equal masses. The cause of lowly-organised plants ranging widely will be discussed in our chapter on Geographical Distribution. and has little or no relation to the size of the genera to which the species belong. Fresh water and salt-loving plants generally have very wide ranges and are much diffused. and all those in the smaller genera on the other side.5..e. if we look at each species as a special act of creation. To test the truth of this anticipation I have arranged the plants of twelve countries. that I am surprised that my tables show even a small majority on the side of the larger genera. Again. or those including many species. I will here allude to only two causes of obscurity. and hence we might expect that the circumstances would generally still be favourable to variation.. and. This might have been anticipated. On the other hand. the species of the larger genera on one side. and the coleopterous insects of two districts. plants low in the scale of organisation are generally much more widely diffused than plants higher in the scale. I was led to anticipate that the species of the larger genera in each country would oftener present varieties. to be now forming. and here again there is no close relation to the size of the genera. Species Of The Larger Genera In Each Country Vary More Frequently Than The Species Of The Smaller Genera If the plants inhabiting a country as described in any Flora. for wherever many closely related species (i. a larger proportional number of dominant species. than in one having few. circumstances have been favourable for variation. From looking at species as only strongly marked and well-defined varieties. the former will be found to include a somewhat larger number of the very common and much diffused or dominant species. many varieties or incipient species ought. we expect to find saplings.

inasmuch as geology plainly tells us that small genera have in the lapse of time often increased greatly in size. for if this had been so. or that no small genera are now varying and increasing. the species of the large genera which present any varieties. we ought generally to find the manufactory still in action. that. and this certainly holds good. on an average many are still forming. for wherever many species of the same genus have been formed. are altogether excluded from the tables. These facts are of plain signification on the view that species are only strongly marked and permanent varieties. wherever many species of a genus have been formed. and disappeared. It is not that all large genera are now varying much. Moreover. or where. Both these results follow when another division is made. And this certainly holds true if varieties be looked at as incipient species. the manufactory of species has been active. declined. All that we want to show is. if we may use the expression. the species of that genus present a number of varieties. with from only one to four species. of incipient species. and that large genera have often come to their maxima. for my tables clearly show. than on the side of the smaller genera. 64 . beyond the average.those of the smaller genera on the other side. and when all the least genera. more especially as we have every reason to believe the process of manufacturing new species to be a slow one. that where many species of a genus have been formed. it would have been fatal to my theory. and are thus increasing in the number of their species. 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. that is. as a general rule. invariably present a larger average number of varieties than do the species of the small genera.

As Fries has well remarked. and clustered round certain forms—that is. and. in the same manner as the varieties of any one species are related to each other. No naturalist pretends that all the species of a genus are equally distinct from each other. when compared with each other or with their parent-species. Related To Each Other. of divergence of character. that in large genera the amount of difference between the species is often exceedingly small. for they differ from each other by a less than the usual amount of difference. namely. Now Fries has remarked in regard to plants. But when we come to discuss the principle. and when intermediate links have not been found between doubtful forms. and it may be said. I have also consulted some sagacious and experienced observers. that the amount of difference between varieties. Or the case may be put in another way. as far as my imperfect results go. and Westwood in regard to insects. But Unequally. And In Having Restricted Ranges There are other relations between the species of large genera and their recorded varieties which deserve notice. Moreover. We have seen that there is no infallible criterion by which to distinguish species and well-marked varieties. we shall see how this may be explained. they concur in this view. and. the species of the larger genera resemble varieties. therefore. little groups of species are generally clustered like satellites around other species. round their parent-species. the species of the larger genera are related to each other. that in the larger genera. or sections. they may generally be divided into sub-genera. many of the species already manufactured still to a certain extent resemble varieties. Undoubtedly there is one most important point of difference between varieties and species. In this respect. Many Of The Species Included Within The Larger Genera Resemble Varieties In Being Very Closely. as I call it. they confirm the view. or lesser groups.6. Hence the amount of difference is one very important criterion in settling whether two forms should be ranked as species or varieties. judging by analogy whether or not the amount suffices to raise one or both to the rank of species. is much less than that between the species of the same genus. and how the lesser differences 65 . And what are varieties but groups of forms. I have endeavoured to test this numerically by averages. in which a number of varieties or incipient species greater than the average are now manufacturing. unequally related to each other. naturalists are compelled to come to a determination by the amount of difference between them. more than do the species of the smaller genera. after deliberation.

and in so far resemble varieties. Watson has marked for me in the wellsifted London catalogue of Plants (4th edition) sixty-three plants which are therein ranked as 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. Now. their denominations would be reversed. fifty-three acknowledged varieties are recorded.9 of the provinces into which Mr. Watson as doubtful species. Watson has divided Great Britain.C. This statement is indeed scarcely more than a truism. whereas. Mr. often have much restricted ranges. in this same catalogue.3 provinces. Varieties generally have much restricted ranges. For instance.between varieties tend to increase into the greater differences between species. 66 . H. the species to which these varieties belong range over 14. and these range over 7. for if a variety were found to have a wider range than that of its supposed parent-species. as have the closely allied forms. marked for me by Mr. So that the acknowledged varieties have very nearly the same restricted average range. There is one other point which is worth notice. but which are almost universally ranked by British botanists as good and true species. But there is reason to believe that the species which are very closely allied to other species.7 provinces.

And thus. whereas. And we can clearly understand these analogies. secondly. varieties cannot be distinguished from species—except. and thus originated. the species of these genera have more than the average number of varieties. In genera having more than the average number of species in any country. Summary Finally. if species once existed as varieties. as we shall hereafter see. but the amount of difference considered necessary to give to any two forms the rank of species cannot be defined. if differing very little. 67 . and. But. by a certain indefinite amount of difference between them. for two forms. tend to become converted into new and distinct species. by the discovery of intermediate linking forms. forming little clusters round other species. are generally ranked as varieties. first. by steps hereafter to be explained. the forms of life throughout the universe become divided into groups subordinate to groups. In large genera the species are apt to be closely but unequally allied together. 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. these analogies are utterly inexplicable if species are independent creations. notwithstanding that they cannot be closely connected. Thus the larger genera tend to become larger. In all these respects the species of large genera present a strong analogy with varieties.7. the larger genera also tend to break up into smaller genera. and varieties. and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. Species very closely allied to other species apparently have restricted ranges.

Chapter 3 Struggle For Existence 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. 68 .

1. for instance. as we shall more fully see in the next chapter. is preserved. become ultimately converted into good and distinct species. The offspring. arise? All these results. if useful. we see beautiful adaptations everywhere and in every part of the organic world. for. what rank. follow from the struggle for life. Again. will thus have a better chance of surviving. also. in the structure of the beetle which dives through the water. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties. of the Survival of the Fittest. in short. and to the conditions of life and of one organic being to another being. of the many individuals of any species which are periodically born. in their infinitely complex relations to other organic beings and to their physical conditions of life. which I have called incipient species. Owing to this struggle. by the term natural selection. by which each slight variation. been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe. 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. it may be asked. though necessary as the foundation for the work. will tend to the preservation of such individuals. 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. 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. and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird. 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. helps us but little in understanding how species arise in nature. but a small number can survive. if they be in any degree profitable to the individuals of a species. variations. But the mere existence of individual variability and of some few well-marked varieties. in order to mark its relation to man's power of selection. the two or three hundred doubtful forms of British plants are entitled to hold. how is it that varieties. I have called this principle. if the existence of any well-marked varieties be admitted. in the plumed seed which is wafted by the gentlest breeze. is 69 . How have all those exquisite adaptations of one part of the organisation to another part. however slight and from whatever cause proceeding. But the expression often used by Mr. Herbert Spencer.

In regard to plants. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. extinction. as it well deserves. we shall hereafter see. abundance. and is sometimes equally convenient. we often see superabundance of food. or their eggs. it is not so at all seasons of each recurring year. or more difficult—at least I found it so—than constantly to bear this conclusion in mind. with every fact on distribution. we do not always bear in mind. no one has treated this subject with more spirit and ability than W. Nothing is easier than to admit in words the truth of the universal struggle for life. that. Yet unless it be thoroughly engrained in the mind. rarity. But Natural Selection. we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds. and is as immeasurably superior to man's feeble efforts. We have seen that man by selection can certainly produce great results. We will now discuss in a little more detail the struggle for existence. 70 . will be dimly seen or quite misunderstood. is a power incessantly ready for action. and variation. In my future work this subject will be treated. We behold the face of nature bright with gladness. as the works of Nature are to those of Art. the whole economy of nature. or their nestlings. through the accumulation of slight but useful variations. or we forget how largely these songsters. and are thus constantly destroying life. are destroyed by birds and beasts of prey. given to him by the hand of Nature. Herbert. at greater length. though food may be now superabundant. evidently the result of his great horticultural knowledge. and can adapt organic beings to his own uses.more accurate. Dean of Manchester.

I use for convenience sake the general term of Struggle for Existence. But a plant on the edge of a desert is said to struggle for life against the drought. though more properly it should be said to be dependent on the moisture. in a time of dearth. Used In A Large Sense I should premise that I use this term in a large and metaphorical sense. may more truly be said to struggle with each other. which pass into each other. if too many of these parasites grow on the same tree. Two canine animals. and it may metaphorically be said to struggle with other fruit-bearing plants. may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. in tempting the birds to devour and thus disseminate its seeds. Struggle For Existence. its existence depends on them. for. it languishes and dies. In these several senses. including dependence of one being on another. but success in leaving progeny. may be truly said to struggle with each other which shall get food and live. The mistletoe is dependent on the apple and a few other trees. but can only in a far-fetched sense be said to struggle with these trees.2. of which only one of an average comes to maturity. and including (which is more important) not only the life of the individual. growing close together on the same branch. A plant which annually produces a thousand seeds. The Term. As the mistletoe is disseminated by birds. But several seedling mistletoes. 71 .

when circumstances have been favourable to them during two or three following seasons. which during its natural lifetime produces several eggs or seeds. for the world would not hold them. the numerous recorded cases of the astonishingly rapid increase of various animals in a state of nature. Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world. after a period of from 740 to 750 years there would be nearly nineteen million elephants alive descended from the first pair. and I have taken some pains to estimate its probable minimum rate of natural increase. bringing forth six young in the interval. more or less rapidly. all cannot do so. there must in every case be a struggle for existence. There is no exception to the rule that every organic being naturally increases at so high a rate. as more individuals are produced than can possibly survive. Although some species may be now increasing. and during some season or occasional year. and so on. in numbers. then in twenty years there would be a million plants. if this be so. It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms. in less than a thousand years. Every being. for in this case there can be no artificial increase of food. on the principle of geometrical increase. and goes on breeding till ninety years old. Even slow-breeding man has doubled in twenty-five years. if not destroyed. must suffer destruction during some period of its life.3. or with the individuals of distinct species. namely. the earth would soon be covered by the progeny of a single pair. that. there would literally not be standing room for his progeny. otherwise. Hence. 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. But we have better evidence on this subject than mere theoretical calculations. or with the physical conditions of life. it will be safest to assume that it begins breeding when thirty years old. either one individual with another of the same species. The elephant is reckoned the slowest breeder of all known animals. and no prudential restraint from marriage. Geometrical Ratio Of Increase A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. its numbers would quickly become so inordinately great that no country could support the product. and at this rate. if the statements of the rate of increase of slow-breeding cattle and horses in South 72 . and surviving till one hundred years old.

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. Our familiarity with the larger domestic animals tends. cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years. and those which produce extremely few. under favourable conditions. But this difference does not determine how many individuals of the two species can be supported in a district. and endless others could be given. simply explains their extraordinarily rapid increase and wide diffusion in their new homes. as I hear from Dr. let it be ever so large. and that there has consequently been less destruction of the old and young and that nearly all the young have been enabled to breed. they would have been incredible. to mislead us. Their geometrical ratio of increase. which are now the commonest over the wide plains of La Plata. Several of the plants. like the hippobosca. Falconer. So it is with plants. One fly deposits hundreds of eggs. and we do not keep in mind that thousands are annually slaughtered for food. had not been well authenticated. we see no great destruction falling on them. the result of which never fails to be surprising. and that this geometrical tendency to increase must be checked by destruction at some period of life. The only difference between organisms which annually produce eggs or seeds by the thousand. clothing square leagues of surface almost to the exclusion of every other plant. that the slow breeders would require a few more years to people.America. from Cape Comorin to the Himalaya. and latterly in Australia. In a state of nature almost every full-grown plant annually produces seed. and among animals there are very few which do not annually pair. The obvious explanation is that the conditions of life have been highly favourable. such as the cardoon and a tall thistle. is. A large number of eggs is of some importance to those species which depend on a fluctuating amount of food. which have been imported from America since its discovery. and another. and that in a state of nature an equal number would have somehow to be disposed of. The condor lays a couple of eggs and the ostrich a score. a whole district. a single one. and yet in the same country the condor may be the more numerous of the two. I think. The Fulmar petrel lays but one egg. no one supposes that the fertility of the animals or plants has been suddenly and temporarily increased in any sensible degree. In such cases. have been introduced from Europe. yet it is believed to be the most numerous bird in the world. for it allows them rapidly to increase in 73 . and there are plants which now range in India.

In looking at Nature. that heavy destruction inevitably falls either on the young or old during each generation or at recurrent intervals. 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 the number of the species will almost instantaneously increase to any amount. the average number of any animal or plant depends only indirectly on the number of its eggs or seeds. if a single seed were produced once in a thousand years. so that. many must be produced or the species will become extinct. but if many eggs or young are destroyed. 74 . a small number may be produced. It would suffice to keep up the full number of a tree. and this period in the great majority of cases is an early one. in all cases. which lived on an average for a thousand years. and yet the average stock be fully kept up.number. 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. supposing that this seed were never destroyed and could be ensured to germinate in a fitting place. that each lives by a struggle at some period of its life. If an animal can in any way protect its own eggs or young. Lighten any check. mitigate the destruction ever so little.

With plants there is a vast destruction of seeds. chiefly by slugs and insects. This subject of the checks to increase has been ably treated by several authors. on a piece of ground three feet long and two wide. Look at the most vigorous species. but this is not invariably the case. as with the elephant. by so much will it tend to increase still further. there seems to be little doubt that the stock of partridges. for even the tiger in India most rarely dares to attack a young elephant protected by its dam. Seedlings. for instance. by as much as it swarms in numbers. and where there could be no choking from other plants. and the case would be the same with turf closely browsed by quadrupeds. The amount of food for each species. which determines the average number of a species. grouse. in some cases. in all probability. Here I will make only a few remarks. also. just to recall to the reader's mind some of the chief points. 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. If turf which has long been mown. 75 . there would. but the serving as prey to other animals. We know not exactly what the checks are even in a single instance. although hundreds of thousands of game animals are now annually shot. On the other hand. Nor will this surprise any one who reflects how ignorant we are on this head. but very frequently it is not the obtaining food. dug and cleared. even in regard to mankind. thus out of twenty species grown on a little plot of mown turf (three feet by four) nine species perished. more especially in regard to the feral animals of South America. the more vigorous plants gradually kill the less vigorous. from the other species being allowed to grow up freely.4. of course. are destroyed in vast numbers by various enemies. Nature Of The Checks To Increase The causes which check the natural tendency of each species to increase are most obscure. be let to grow. If not one head of game were shot during the next twenty years in England. and out of 357 no less than 295 were destroyed. although so incomparably better known than any other animal. be less game than at present. I marked all the seedlings of our native weeds as they came up. if no vermin were destroyed. none are destroyed by beasts of prey. and hares on any large estate depends chiefly on the destruction of vermin. and. gives the extreme limit to which each can increase. Thus. at the same time. though fully grown plants. Eggs or very young animals seem generally to suffer most.

acts directly. 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. is an extraordinarily severe mortality from epidemics with man. and finally disappearing. and the change of climate being conspicuous. but which never become naturalised. but in so far as climate chiefly acts in reducing food. and this is a tremendous destruction. which subsist on the same kind of food. But this is a false view. for the number of species of all kinds. and therefore of competitors. or absolute deserts.Climate plays an important part in determining the average numbers of a species. Even when climate. as in this one being hurt. hence in going northward. When we reach the Arctic regions. The action of climate seems at first sight to be quite independent of the struggle for existence. is constantly suffering enormous destruction at some period of its life. for they cannot compete with our native plants nor resist destruction by our native animals. the other species must decrease. than we do in proceeding southward or in descending a mountain. we may feel sure that the cause lies quite as much in other species being favoured. whether of the same or of distinct species. 76 . we are tempted to attribute the whole effect to its direct action. the struggle for life is almost exclusively with the elements. but in a somewhat lesser degree. when we remember that ten per cent. which will suffer the most. we invariably see some species gradually getting rarer and rarer. from enemies or from competitors for the same place and food. we far oftener meet with stunted forms. it brings on the most severe struggle between the 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. and periodical seasons of extreme cold or drought seem to be the most effective of all checks. due to the DIRECTLY injurious action of climate. and if these enemies or competitors be in the least degree favoured by any slight change of climate. or in ascending a mountain. even where it most abounds. it will be the least vigorous individuals. decreases northward. So it is when we travel northward. and as each area is already fully stocked with inhabitants. When we travel from south to north. or from a damp region to a dry. When we travel southward and see a species decreasing in numbers. they will increase in numbers. or snow-capped summits. or those which have got least food through the advancing winter. for instance. extreme cold. we forget that each species.

that is abounding in individuals. and thus save the species from utter destruction. this seems generally to occur with our game animals—often ensue. is absolutely necessary for its preservation. epidemics—at least. 77 . I have in this case lost every single seed.. This view of the necessity of a large stock of the same species for its preservation. possibly in part through facility of diffusion among the crowded animals. in our fields. and here we have a limiting check independent of the struggle for life. because the seeds are in great excess compared with the number of birds which feed on them. and the ill effects of close interbreeding. no doubt come into play in many of these cases. as their numbers are checked during the winter.When a species. some singular facts in nature such as that of very rare plants being sometimes extremely abundant. which have from some cause. but I will not here enlarge on this subject. I should add that the good effects of intercrossing. increase in number proportionally to the supply of seed. we may believe. owing to highly favourable circumstances. that a plant could exist only where the conditions of its life were so favourable that many could exist together. 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. and that of some social plants being social. even on the extreme verge of their range. relatively to the numbers of its enemies. etc. For in such cases. explains. in the few spots where they do exist. though having a superabundance of food at this one season. But even some of these so-called epidemics appear to be due to parasitic worms. I believe. On the other hand. a large stock of individuals of the same species. in many cases. increases inordinately in numbers in a small tract. Thus we can easily raise plenty of corn and rape-seed. nor can the birds. been disproportionally favoured: and here comes in a sort of struggle between the parasite and its prey.

and the heath was frequented by two or three distinct insectivorous birds. whence I could examine hundreds of acres of the unenclosed heath. I found a multitude of seedlings and little trees. which had been perpetually browsed down by the cattle. Here there are extensive heaths. though a simple one. and one of them. which have to struggle together in the same country. but twelve species of plants (not counting grasses and carices) flourished in the plantations. except the old planted clumps. But on looking closely between the stems of the heath.5. nothing whatever else having been done. In one square yard. 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. I counted thirty-two little trees. during many years tried to raise its head above the stems of the heath. as soon as the land was enclosed. 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. there was a large and extremely barren heath. in Surrey. and had failed. which. which had never been touched by the hand of man. The change in the native vegetation of the planted part of the heath was most remarkable. for six insectivorous birds were very common in the plantations. it became thickly clothed with vigorously growing young firs. Yet the heath was so extremely barren 78 . I will give only a single instance. with the exception of the land having been enclosed. In Staffordshire. and self-sown firs are now springing up in multitudes. 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. which were not to be seen on the heath. had. Complex Relations Of All Animals And Plants To Each Other In The Struggle For Existence Many cases are on record showing how complex and unexpected are the checks and relations between organic beings. I plainly saw near Farnham. The effect on the insects must have been still greater. with a few clumps of old Scotch firs on the distant hill-tops: within the last ten years large spaces have been enclosed. But how important an element enclosure is. interested me. with twenty-six rings of growth. so close together that all cannot live. No wonder that. where I had ample means of investigation. and literally I could not see a single Scotch fir. which could not be found on the heath. so that cattle could not enter. on the estate of a relation. Here we see how potent has been the effect of the introduction of a single tree.

produced not one. Not that under nature the relations will ever be as simple as this. that we marvel when we hear of the extinction of an organic being. Hence. I find from experiments that humblebees are almost indispensable to the fertilisation of the heartsease (Viola tricolor).700 seeds. and Azara and Rengger have shown that this is caused by the greater number in Paraguay of a certain fly. must be habitually checked by some means. and this. if certain insectivorous birds were to decrease in Paraguay. or invent laws on the duration of the forms of life! I am tempted to give one more instance showing how plants and animals. though assuredly the merest trifle would give the victory to one organic being over another. 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. protected from bees. Battle within battle must be continually recurring with varying success. 100 heads of red clover (T. probably by other parasitic insects. so profound is our ignorance. 79 . but in several parts of the world insects determine the existence of cattle. from its peculiar structure.and so extensive that no one would ever have imagined that cattle would have so closely and effectually searched it for food. as we have just seen in Staffordshire. for instance twenty heads of Dutch clover (Trifolium repens) yielded 2. the insectivorous birds.290 seeds. Nearly all our orchidaceous plants absolutely require the visits of insects to remove their pollenmasses and thus to fertilise them. I shall hereafter have occasion to show that the exotic Lobelia fulgens is never visited in my garden by insects. for here neither cattle nor horses nor dogs have ever run wild. the parasitic insects would probably increase. numerous as they are. and so high our presumption. The increase of these flies. and so onwards in ever-increasing circles of complexity. 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. Nevertheless. Again. are bound together by a web of complex relations. remote in the scale of nature. Here we see that cattle absolutely determine the existence of the Scotch fir. which lays its eggs in the navels of these animals when first born. but twenty other heads. Perhaps Paraguay offers the most curious instance of this. and consequently. though they swarm southward and northward in a feral state. and this would lessen the number of the navel-frequenting flies—then cattle and horses would become feral. I have also found that the visits of bees are necessary for the fertilisation of some kinds of clover. pratense) produced 2. and as we do not see the cause. for other bees do not visit this flower.

The number of humble-bees in any district depends in a great measure upon the number of field-mice. and Colonel Newman. believes that "more than two-thirds of them are thus destroyed all over England. what war between insect and insect—between insects. Throw up a handful of feathers. or even the existence of the species. What a struggle must have gone on during long centuries between the several kinds of trees. snails. on the number of cats. It has been suggested that moths may fertilise the clovers. from their weight not being sufficient to depress the wing petals. each annually scattering its seeds by the thousand. but it has been observed that ancient Indian ruins in the Southern United States. and other animals with birds and beasts of prey—all striving to increase. but all will concur in determining the average number. which must formerly have been cleared of trees. now display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. or wholly disappear. as other bees cannot reach the nectar. Hence we may infer as highly probable that. But how false a view is this! Every one has heard that when an American forest is cut down. their seeds and seedlings. if the whole genus of humble-bees became extinct or very rare in England. acting at different periods of life. which I attribute to the number of cats that destroy the mice. we are tempted to attribute their proportional numbers and kinds to what we call chance. In some cases it can be shown that widely-different checks act on the same species in different districts. and all fall to the ground according to definite laws. many different checks. as every one knows." Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine. probably come into play. but how simple is the problem where each shall 80 . some one check or some few being generally the most potent. all feeding on each other. which destroy their combs and nests. who has long attended to the habits of humble-bees. and during different seasons or years. the frequency of certain flowers in that district! In the case of every species. "Near villages and small towns I have found the nests of humble-bees more numerous than elsewhere. When we look at the plants and bushes clothing an entangled bank. the heartsease and red clover would become very rare. but I doubt whether they could do so in the case of the red clover." Now the number of mice is largely dependent. or on the other plants which first clothed the ground and thus checked the growth of the trees. Humble bees alone visit red clover. through the intervention first of mice and then of bees. and Colonel Newman says. a very different vegetation springs up. or on the trees.but the same number of protected heads produced not a single seed.

This is likewise sometimes the case with those which may strictly be said to struggle with each other for existence. lies generally between beings remote in the scale of nature. some of the varieties which best suit the soil or climate. for they frequent the same districts. or are naturally the most fertile. 81 . the proportional numbers and kinds of trees now growing on the old Indian ruins! The dependency of one organic being on another. as of a parasite on its prey. To keep up a mixed stock of even such extremely close varieties as the variously coloured sweet-peas. and we sometimes see the contest soon decided: for instance. and are exposed to the same dangers. It may even be doubted whether the varieties of any of our domestic plants or animals have so exactly the same strength. they must be each year harvested separately. habits. and will consequently in a few years supplant the other varieties. So again with the varieties of sheep: it has been asserted that certain mountain-varieties will starve out other mountain-varieties. so that they cannot be kept together. the struggle will generally be almost equally severe.fall compared to that of the action and reaction of the innumerable plants and animals which have determined. In the case of varieties of the same species. and the seed then mixed in due proportion. will beat the others and so yield more seed. otherwise the weaker kinds will steadily decrease in number and disappear. and the mixed seed be resown. if several varieties of wheat be sown together. But the struggle will almost invariably be most severe between the individuals of the same species. require the same food. and if the seed or young were not annually preserved in due proportion. and constitution. The same result has followed from keeping together different varieties of the medicinal leech. that the original proportions of a mixed stock (crossing being prevented) could be kept up for half-a-dozen generations. in the same manner as beings in a state of nature. in the course of centuries. if they were allowed to struggle together. as in the case of locusts and grassfeeding quadrupeds.

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. which fill nearly the same place in the economy of nature. so well adapted for diving. with which it comes into competition for food or residence. and so in other cases.6. stingless native bee. as peas and beans. and in the flattened and fringed legs of the water-beetle. 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. to that of all other organic beings. Struggle For Life Most Severe Between Individuals And Varieties Of The Same Species As the species of the same genus usually have. the struggle will generally be more severe between them. or on which it preys. But from the strong growth of young plants produced from such seeds. Yet the advantage of the plumed seeds no doubt stands in the closest relation to the land being already thickly clothed with other plants. so that the seeds may be widely distributed and fall on unoccupied ground. This is obvious in the structure of the teeth and talons of the tiger. but probably in no one case could we precisely say why one species has been victorious over another in the great battle of life. that the structure of every organic being is related. In the water-beetle. if they come into competition with each other. But in the beautifully plumed seed of the dandelion. and in that of the legs and claws of the parasite which clings to the hair on the tiger's body. or from which it has to escape. it may be suspected that 82 . than between the species of distinct genera. and to escape serving as prey to other animals. in the most essential yet often hidden manner. when sown in the midst of long grass. The recent increase of the missel-thrush in parts of Scotland has caused the decrease of the song-thrush. namely. the structure of its legs. In Australia the imported hive-bee is rapidly exterminating the small. We can dimly see why the competition should be most severe between allied forms. much similarity in habits and constitution. 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. though by no means invariably. One species of charlock has been known to supplant another species. allows it to compete with other aquatic insects. the relation seems at first confined to the elements of air and water. A corollary of the highest importance may be deduced from the foregoing remarks. to hunt for its own prey. and always in structure.

or at intervals. as it is difficult to acquire. It is good thus to try in imagination to give any one species an advantage over another. yet there will be competition between some few species. we should have to modify it in a different way to what we should have had to do in its native country.the chief use of the nutriment in the seed is to favour the growth of the seedlings. that they are destroyed exclusively by the rigour of the climate. Not until we reach the extreme confines of life. for elsewhere it ranges into slightly hotter or colder. a conviction as necessary. will competition cease. that no fear is felt. for we should have to give it some advantage over a different set of competitors or enemies. dampness or dryness. damper or drier districts. Hence we can see that when a plant or animal is placed in a new country. during some season of the year. at some period of its life. or over the animals which prey on it. that death is generally prompt. has to struggle for life and to suffer great destruction. during each generation. among new competitors. in the Arctic regions or on the borders of an utter desert. we should have to give it some advantage over its competitors. If its average numbers are to increase in its new home. that each. On the confines of its geographical range. Probably in no single instance should we know what to do. This ought to convince us of our ignorance on the mutual relations of all organic beings. for the warmest or dampest spots. The land may be extremely cold or dry. or between the individuals of the same species. the conditions of its life will generally be changed in an essential manner. When we reflect on this struggle we may console ourselves with the full belief that the war of nature is not incessant. whilst struggling with other plants growing vigorously all around. 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. and the happy survive and multiply. and that the vigorous. the healthy. although the climate may be exactly the same as in its former home. but we have reason to believe that only a few plants or animals range so far. In this case we can clearly see that if we wish in imagination to give the plant the power of increasing in numbers. All that we can do is to keep steadily in mind that each organic being is striving to increase in a geometrical ratio. 83 . a change of constitution with respect to climate would clearly be an advantage to our plant.

Or The Survival Of The Fittest Natural Selection — its power compared with man's selection — its power on characters of trifling importance — its power at all ages and on both sexes — Sexual Selection — On the generality of intercrosses between individuals of the same species — Circumstances favourable and unfavourable to the results of Natural Selection. intercrossing. through Divergence of Character and Extinction. 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. number of individuals — Slow action — Extinction caused by Natural Selection — Divergence of Character. isolation. 84 .Chapter 4 Natural Selection. related to the diversity of inhabitants of any small area and to naturalisation — Action of Natural Selection. namely.

seeing that variations useful to man have undoubtedly occurred. act in regard to variation? Can the principle of selection. I have called Natural Selection. we may feel sure that any variation in the least degree injurious would be rigidly destroyed. No one 85 . Several writers have misapprehended or objected to the term Natural Selection. owing to the nature of the organism and the nature of the conditions. which we almost universally meet with in our domestic productions is not directly produced. he can neither originate varieties nor prevent their occurrence. This preservation of favourable individual differences and variations. Can it then be thought improbable. however slight. or the Survival of the Fittest. and consequently what infinitely varied diversities of structure might be of use to each being under changing conditions of life. and the destruction of those which are injurious. whereas it implies only the preservation of such variations as arise and are beneficial to the being under its conditions of life. Variations neither useful nor injurious would not be affected by natural selection. and variability ensues. which we have seen is so potent in the hands of man. briefly discussed in the last chapter. can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage. that other variations useful in some way to each being in the great and complex battle of life. as Hooker and Asa Gray have well remarked. would have the best chance of surviving and procreating their kind? On the other hand. and would be left either a fluctuating element. and. over others. or would ultimately become fixed.1. should occur in the course of many successive generations? If such do occur. But the variability. Let the endless number of slight variations and individual differences occurring in our domestic productions. it may truly be said that the whole organisation becomes in some degree plastic. as perhaps we see in certain polymorphic species. Some have even imagined that natural selection induces variability. but similar changes of conditions might and do occur under nature. by man. Under domestication. in those under nature. he can only preserve and accumulate such as do occur. How will the struggle for existence. 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. Unintentionally he exposes organic beings to new and changing conditions of life. as well as the strength of the hereditary tendency. be borne in mind. in a lesser degree. apply under nature? I think we shall see that it can act most efficiently.

We may conclude. that any change in the numerical proportions of the inhabitants. In such cases. and by laws the sequence of events as ascertained by us. no doubt. natural selection is a false term. of climate. these same places would have been seized on by intruders. for. would tend to be preserved. So again it is difficult to avoid personifying the word Nature. 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. 86 . from what we have seen of the intimate and complex manner in which the inhabitants of each country are bound together. would seriously affect the others. It has been said that I speak of natural selection as an active power or Deity. and they are almost necessary for brevity. which in any way favoured the individuals of any species. must of necessity first occur. 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. We shall best understand the probable course of natural selection by taking the case of a country undergoing some slight physical change. and this would likewise seriously disturb the relations of some of the former inhabitants.objects to agriculturists speaking of the potent effects of man's selection. but I mean by nature. and it has even been urged that. If the country were open on its borders. natural selection is not applicable to them! In the literal sense of the word. had the area been open to immigration. Let it be remembered how powerful the influence of a single introduced tree or mammal has been shown to be. which man for some object selects. or of a country partly surrounded by barriers. by better adapting them to their altered conditions. independently of the change of climate itself. 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. into which new and better adapted forms could not freely enter. slight modifications. for instance. With a little familiarity such superficial objections will be forgotten. Others have objected that the term selection implies conscious choice in the animals which become modified. But in the case of an island. as plants have no volition. The proportional numbers of its inhabitants will almost immediately undergo a change. and some species will probably become extinct. only the aggregate action and product of many natural laws. new forms would certainly immigrate. and natural selection would have free scope for the work of improvement. and in this case the individual differences given by nature.

except in so far as they are useful to any being. Man keeps the natives of many climates in the same country. a great result by his methodical and unconscious means of selection. the natives have been so far conquered by naturalised productions that they have allowed some foreigners to take firm possession of the land. for in all countries. cares nothing for appearances. Nature only for that of the being which she tends. As man can produce. so could natural selection. Man selects only for his own good. Unless such occur. And as foreigners have thus in every country beaten some of the natives. Every selected character is fully exercised by her. and in the foregoing cases the conditions the changed. For as all the inhabitants of each country are struggling together with nicely balanced forces. that none of them could be still better adapted or improved. by affording a better chance of the occurrence of profitable variations. to check immigration. extremely slight modifications in the structure or habits of one species would often give it an advantage over others. Nor do I believe that any great physical change. Under the term of "variations. we may safely conclude that the natives might have been modified with advantage. but far more easily from having incomparably longer time for action. and still further modifications of the same kind would often still further increase the advantage. as shown in the first chapter. he feeds a long and a short-beaked pigeon 87 . He seldom exercises each selected character in some peculiar and fitting manner. as of climate. As man can produce a great result with his domestic animals and plants by adding up in any given direction individual differences." it must never be forgotten that mere individual differences are included. as long as the species continued under the same conditions of life and profited by similar means of subsistence and defence. what may not natural selection effect? Man can act only on external and visible characters: Nature. that changes in the conditions of life give a tendency to increased variability. natural selection can do nothing. She can act on every internal organ. 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.We have good reason to believe. as is implied by the fact of their selection. 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. on the whole machinery of life. or any unusual degree of isolation. so as to have better resisted the intruders. and this would manifestly be favourable to natural selection. and certainly has produced. on every shade of constitutional difference. if I may be allowed to personify the natural preservation or survival of the fittest.

he does not rigidly destroy all inferior animals. this can hardly be considered as an unwarrantable assumption.on the same food. does not allow the most vigorous males to struggle for the females. He often begins his selection by some half-monstrous form. throughout the world. When we see leaf- 88 . until the hand of time has marked the long lapse of ages. Although natural selection can act only through and for the good of each being. compared with those accumulated by Nature during whole geological periods! Can we wonder. he exposes sheep with long and short wool to the same climate. as far as lies in his power. then. at the improvement of each organic being in relation to its organic and inorganic conditions of life. that they should be infinitely better adapted to the most complex conditions of life. and should plainly bear the stamp of far higher workmanship? It may metaphorically be said that natural selection is daily and hourly scrutinising. But whether it is true. and so onward. yet characters and structures. In order that any great amount of modification should be effected in a species. or at least by some modification prominent enough to catch the eye or to be plainly useful to him. we can judge only by seeing how far the hypothesis accords with and explains the general phenomena of nature. is likewise a simple assumption. We see nothing of these slow changes in progress. may thus be acted on. vary or present individual differences of the same favourable nature as before. the ordinary belief that the amount of possible variation is a strictly limited quantity. step by step. preserving and adding up all that are good. that Nature's productions should be far "truer" in character than man's productions. the slightest variations. WHENEVER AND WHEREVER OPPORTUNITY OFFERS. which we are apt to consider as of very trifling importance. Under nature. On the other hand. and consequently how poor will be his results. and these must again be preserved. How fleeting are the wishes and efforts of man! How short his time. but protects during each varying season. silently and insensibly working. all his productions. perhaps after a long interval of time. the slightest differences of structure or constitution may well turn the nicely-balanced scale in the struggle for life. a variety. rejecting those that are bad. when once formed must again. Seeing that individual differences of the same kind perpetually recur. he does not exercise a long-backed or long-legged quadruped in any peculiar manner. and so be preserved. and then so imperfect is our view into long-past geological ages that we see only that the forms of life are now different from what they formerly were.

In plants. Grouse. owing to the law of correlation. in the eggs of poultry. the red-grouse the colour of heather. such differences would effectually settle which variety. as being the most liable to destruction. as far as our ignorance permits us to judge. In looking at many small points of difference between species. seem quite unimportant. other modifications. will ensue. a Curculio. It is also necessary to bear in mind that. and hawks are guided by eyesight to their prey. determines whether they shall live or die. that purple plums suffer far more from a certain disease than yellow plums. if not destroyed at some period of their lives. whether a smooth or downy. we should remember how essential it is in a flock of white sheep to destroy a lamb with the faintest trace of black. and bark-feeders mottled-grey. size and flavour of the seeds of the many varieties of our culinary and agricultural plants. food. these slight differences make a great difference in cultivating the several varieties. Hence natural selection might be effective in giving the proper colour to each kind of grouse. the alpine ptarmigan white in winter. where the trees would have to struggle with other trees and with a host of enemies. with all the aids of art. have no doubt produced some direct effect. which. assuredly. for instance. should succeed. they are known to suffer largely from birds of prey. and in keeping that colour. often of the most unexpected nature. we must believe that these tints are of service to these birds and insects in preserving them from danger. than those with down. whereas another disease attacks yellow-fleshed peaches far more than those with other coloured flesh. true and constant.. when once acquired.—so much so that on parts of the continent persons are warned not to keep white pigeons. a yellow or a purple-fleshed fruit. we must not forget that climate. yet we hear from an excellent horticulturist. etc. Downing. tend to reappear in the offspring at the same period. appear at any particular period of life. If. in the horns of our sheep 89 . and in the colour of the down of their chickens. that in the United States smooth-skinned fruits suffer far more from a beetle.eating insects green. in a state of nature. which feed on the "paint-root" in Virginia. would increase in countless numbers. As we see that those variations which. Nor ought we to think that the occasional destruction of an animal of any particular colour would produce little effect. in the caterpillar and cocoon stages of the varieties of the silkworm. We have seen how the colour of hogs. in the shape. when one part varies and the variations are accumulated through natural selection. under domestication. the down on the fruit and the colour of the flesh are considered by botanists as characters of the most trifling importance.

for instance. wholly different from those which concern the mature insect. and by their inheritance at a corresponding age. Now. the process of modification would be very slow.and cattle when nearly adult. I can see no greater difficulty in this being effected through natural selection. more delicate and more easily broken shells might be selected. which can have little or no influence on the course of natural selection. Natural selection will modify the structure of the young in relation to the parent and of the parent in relation to the young. if of high importance to it. used exclusively for opening the cocoon—or the hard tip to the beak of unhatched birds. In social animals it will adapt the structure of each individual for the benefit of the whole community. which had the most powerful and hardest beaks. that of the best short-beaked tumbler-pigeons a greater number perish in the egg than are able to get out of it. For instance. without giving it any advantage. I cannot find one case which will bear investigation. is to modify the structure of one species. and there would be simultaneously the most rigorous selection of all the young birds within the egg. If it profit a plant to have its seeds more and more widely disseminated by the wind. so that fanciers assist in the act of hatching. but in all cases natural selection will ensure that they shall not be injurious: for if they were so. What natural selection cannot do. than in the cotton-planter increasing and improving by selection the down in the pods on his cottontrees. a vast number of eggs or seeds are annually devoured. used for breaking the eggs. the species would become extinct. and these modifications may affect. for the good of another species. A structure used only once in an animal's life. for all with weak beaks would inevitably perish: or. if nature had to make the beak of a full-grown pigeon very short for the bird's own advantage. if the community profits by the selected change. Natural selection may modify and adapt the larva of an insect to a score of contingencies. the thickness of the shell being known to vary like every other structure. and these could be modified through natural selection 90 . conversely. and though statements to this effect may be found in works of natural history. It may be well here to remark that with all beings there must be much fortuitous destruction. through correlation. might be modified to any extent by natural selection. So. modifications in the adult may affect the structure of the larva. It has been asserted. the structure of the adult. the great jaws possessed by certain insects. so in a state of nature natural selection will be enabled to act on and modify organic beings at any age. by the accumulation of variations profitable at that age.

So again a vast number of mature animals and plants. but this is no valid objection to its efficiency at other times and in other ways. supposing that there is any variability in a favourable direction. 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. must be annually destroyed by accidental causes. But let the destruction of the adults be ever so heavy. will tend to propagate their kind in larger numbers than the less well adapted. Yet many of these eggs or seeds would perhaps. as will often have been the case. If the numbers be wholly kept down by the causes just indicated. whether or not they be the best adapted to their conditions. 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. have yielded individuals better adapted to their conditions of life than any of those which happened to survive. 91 . natural selection will be powerless in certain beneficial directions. the best adapted individuals. if not destroyed. 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.only if they varied in some manner which protected them from their enemies.

the most vigorous males. confined to the male sex. for the shield may be as important for victory as the sword or spear. How low in the scale of nature the law of battle descends I know not. male stag-beetles sometimes bear wounds from the huge mandibles of other males. perhaps. as is sometimes the case. but on having special weapons. in nearly the same manner as does the brutal cockfighter by the careful selection of his best cocks. but on a struggle between the individuals of one sex. The males of carnivorous animals are already well armed. not on a struggle for existence in relation to other organic beings or to external conditions.2. Sexual selection. fighting for a particular female who sits by. so no doubt it will be under nature. Sexual selection is. for the possession of the females. and these seem oftenest provided with special weapons. This leads me to say a few words on what I have called sexual selection. by singing. This form of selection depends. as commonly occurs. the males of certain hymenopterous insects have been frequently seen by that inimitable observer M. But in many cases victory depends not so much on general vigour. generally the males. severest between the males of polygamous animals. for the possession of the other sex. A hornless stag or spurless cock would have a poor chance of leaving numerous offspring. and strength to the wing to strike in the spurred leg. The result is not death to the unsuccessful competitor. believe that there is the severest rivalry between the males of many species to attract. less rigorous than natural selection. male salmons have been observed fighting all day long. and whirling round. All those who have attended to the subject. and the hooked jaw to the male salmon. male alligators have been described as fighting. as the mane of the lion. Sexual Selection Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex. the 92 . might surely give indomitable courage. will leave most progeny. The war is. Fabre. bellowing. special means of defence may be given through means of sexual selection. those which are best fitted for their places in nature. Thus it is rendered possible for the two sexes to be modified through natural selection in relation to different habits of life. by always allowing the victor to breed. Generally. but few or no offspring. or for one sex to be modified in relation to the other sex. Among birds. length of spur. an apparently unconcerned beholder of the struggle. therefore. like Indians in a war-dance. though to them and to others. and then retires with the conqueror. the contest is often of a more peaceful character.

means of defence. by selecting. which they have transmitted to their male offspring alone. according to his standard of beauty. Yet. they likewise perform strange antics before the females. the most melodious or beautiful males. I can see no good reason to doubt that female birds.females. as I believe. some slight advantage over other males. and show off in the best manner. birds of paradise. and transmitted to the males alone or to both sexes at corresponding ages. with respect to the plumage of male and female birds. standing by as spectators. might produce a marked effect. The tuft of hair on the breast of the wild turkey-cock cannot be of any use. their gorgeous plumage. that when the males and females of any animal have the same general habits of life. Heron has described how a pied peacock was eminently attractive to all his hen birds. in comparison with the plumage of the young. Those who have closely attended to birds in confinement well know that they often take individual preferences and dislikes: thus Sir R. according to their standard of beauty. and it is doubtful whether it can be ornamental in the eyes of the female bird. in successive generations. at last choose the most attractive partner. can partly be explained through the action of sexual selection on variations occurring at different ages. which. congregate. colour. 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. I cannot here enter on the necessary details. Thus it is. which apparently have not been augmented through selection by man. 93 . and successive males display with the most elaborate care. The rock-thrush of Guiana. but I have not space here to enter on this subject. and some others. indeed. or charms. such differences have been mainly caused by sexual selection: that is. in their weapons. had the tuft appeared under domestication it would have been called a monstrosity. but if man can in a short time give beauty and an elegant carriage to his bantams. by individual males having had. but differ in structure. or ornament. Some well-known laws. during thousands of generations.

securing some by craft. another mice. taking to catch rats. than that man should be able to improve the fleetness of his greyhounds by careful and methodical selection. Even without any change in the proportional numbers of the animals on which our wolf preyed. I must beg permission to give one or two imaginary illustrations. and by the repetition of this process. if any slight innate change of habit or of structure benefited an individual wolf. and another hunting on marshy ground and almost nightly catching woodcocks or snipes. according to Mr. and so be preserved or selected. Or. one cat. The tendency to catch rats rather than mice is known to be inherited. when they were compelled to prey on other animals. bringing home winged game. one with a light greyhound-like form. I can see no more reason to doubt that this would be the result. there are two varieties of the wolf inhabiting the Catskill Mountains. a cub might be born with an innate tendency to pursue certain kinds of prey. one cat. with shorter legs. which preys on various animals. Some of its young would probably inherit the same habits or structure. according to Mr. another hares or rabbits. for instance. provided always that they retained strength to master their prey at this or some other period of the year. the wolves inhabiting a mountainous district. which more frequently attacks the shepherd's flocks. and those frequenting the lowlands. Under such circumstances the swiftest and slimmest wolves have the best chance of surviving. again. I may add that. Pierce. in the United States. Now. Let us take the case of a wolf. which pursues deer. and let us suppose that the fleetest prey. and the other more bulky. would naturally be forced to hunt different prey. during that season of the year when the wolf was hardest pressed for food. it would have the best chance of surviving and of leaving offspring. St. 94 . and from the continued preservation of the individuals best fitted for the two sites. natural selection acts. had from any change in the country increased in numbers. and some by fleetness.3. as I believe. Nor can this be thought very improbable. or that other prey had decreased in numbers. a new variety might be formed which would either supplant or coexist with the parent-form of wolf. John. for we often observe great differences in the natural tendencies of our domestic animals. some by strength. a deer for instance. Or The Survival Of The Fittest In order to make it clear how. 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. Illustrations Of The Action Of Natural Selection.

This is rather an extreme estimate for most of the higher animals. I saw the great importance of individual differences. would be a rare event. which depends on the preservation of all the more or less valuable individuals. which varied in some manner. still. from various causes of destruction. but there can hardly be a doubt. judging by what we see taking place under domestication.two varieties might slowly be formed. a bird of some kind could procure its food more easily by having its beak curved. I think. that this result would follow from the preservation during many generations of a large number of individuals 95 . such as a monstrosity. These varieties would cross and blend where they met. giving it twice as good a chance of life as that of the other individuals. be disputed. which more frequently attacks the shepherd's flocks. nevertheless there would be a very poor chance of this one individual perpetuating its kind to the exclusion of the common form. Nevertheless. only two on an average survive to pro-create their kind. Supposing it to survive and to breed. and this led me fully to discuss the results of unconscious selection by man. of which. producing during their lifetime two hundred offspring. that. I did not appreciate how rarely single variations. which pursues deer. The justice of these remarks cannot. and if one were born with its beak strongly curved. yet the chances would be strongly against its survival. and this chance would go on decreasing in the succeeding generations. but to this subject of intercrossing we shall soon have to return. there are two varieties of the wolf inhabiting the Catskill Mountains in the United States. and which consequently flourished. He then shows that if a single individual were born. also. The author takes the case of a pair of animals. I may add. if at first preserved. I speak of the slimmest individual wolves. In former editions of this work I sometimes spoke as if this latter alternative had frequently occurred. for instance. Pierce. one with a light greyhound-like form. and the other more bulky. and that half its young inherited the favourable variation. If. but by no means so for many of the lower organisms. as the Reviewer goes onto show. I saw. could be perpetuated. it would generally be lost by subsequent intercrossing with ordinary individuals. the young would have only a slightly better chance of surviving and breeding. according to Mr. and on the destruction of the worst. that the preservation in a state of nature of any occasional deviation of structure. It should be observed that in the above illustration. with shorter legs. and that. whether slight or strongly marked. until reading an able and valuable article in the "North British Review" (1867). and not of any single strongly marked variation having been preserved.

I shall have to recur. Consequently each newly-formed variety would generally be at first local. but their visits do not in any way benefit the plant. and from the destruction of a still larger number with the straightest beaks. Insects in seeking the nectar would get 96 . This juice. is greedily sought by insects. so that similarly modified individuals would soon exist in a small body together. It may be worth while to give another and more complex illustration of the action of natural selection. if the variation were of a beneficial nature. fifth. competing with and conquering the unchanged individuals on the margins of an ever-increasing circle. frequently recur owing to a similar organisation being similarly acted on— of which fact numerous instances could be given with our domestic productions. of which fact several instances could be given. There can also be little doubt that the tendency to vary in the same manner has often been so strong that all the individuals of the same species have been similarly modified without the aid of any form of selection. if the varying individual did not actually transmit to its offspring its newly-acquired character. If the new variety were successful in its battle for life. and would often breed together. the original form would soon be supplanted by the modified form. Now.with more or less strongly curved beaks. 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. It should not. we see this even with migratory birds. through the survival of the fittest. by glands at the base of the stipules in some Leguminosae. which no one would rank as mere individual differences. though small in quantity. In cases of this kind. it would slowly spread from a central district. To the effects of intercrossing in eliminating variations of all kinds. a still stronger tendency to vary in the same manner. apparently for the sake of eliminating something injurious from the sap: this is effected. for instance. however. but it may be here remarked that most animals and plants keep to their proper homes. Or only a third. that it was formerly ranked as a distinct species under the name of Uria lacrymans. and do not needlessly wander about. which almost always return to the same spot. as seems to be the common rule with varieties in a state of nature. In such cases. be overlooked that certain rather strongly marked variations. Certain plants excrete sweet juice. as long as the existing conditions remained the same. Thus Graba estimates that about one-fifth of the guillemots in the Faroe Islands consist of a variety so well marked. it would undoubtedly transmit to them. and at the backs of the leaves of the common laurel. or tenth part of the individuals may have been thus affected.

would likewise be favoured. and on some a profusion. unintentionally on their part. and a cross thus effected. which had their stamens and pistils placed. at first occasionally and then habitually. and four stamens with shrivelled anthers. also. the pollen could not thus have been carried. Having found a female tree exactly sixty yards from a male tree. gives rise to vigorous seedlings. The plants which produced flowers with the largest glands or nectaries. Some holly-trees bear only male flowers. and that they do this effectually I could easily show by many striking facts. nevertheless every female flower which I examined had been effectually fertilised by the bees. and on all. The flowers of two distinct individuals of the same species would thus get crossed. taken from different branches.dusted with pollen. The weather had been cold and boisterous and therefore not favourable to bees. I will give only one. there were a few pollen-grains. and so in the long-run would gain the upper hand and form a local variety. as likewise illustrating one step in the separation of the sexes of plants. As the wind had set for several days from the female to the male tree. would oftenest be visited by insects. by the above process long continued. they would. and a rudimentary pistil. had been rendered highly attractive to insects. yet if a little pollen were carried. by the pollen-devouring insects from flower to flower. We might have taken the case of insects visiting flowers for the sake of collecting pollen instead of nectar. its destruction appears to be a simple loss to the plant. would be selected. and would often transport it from one flower to another. which consequently would have the best chance of flourishing and surviving. The flowers. another process 97 . which have four stamens producing a rather small quantity of pollen. these have a full-sized pistil. under the microscope. other holly-trees bear only female flowers. which had flown from tree to tree in search of nectar. I put the stigmas of twenty flowers. and as pollen is formed for the sole purpose of fertilisation. and the act of crossing. regularly carry pollen from flower to flower. and would oftenest be crossed. in relation to the size and habits of the particular insect which visited them. and had larger anthers. excreting most nectar. although nine-tenths of the pollen were destroyed it might still be a great gain to the plant to be thus robbed. and the individuals which produced more and more pollen. But to return to our imaginary case. without exception. When our plant. in which not a grain of pollen can be detected. so as to favour in any degree the transportal of the pollen. as can be fully proved. as soon as the plant had been rendered so highly attractive to insects that pollen was regularly carried from flower to flower.

it may be believed that under certain circumstances individual differences in the curvature or length of the proboscis. but only in the autumn. That this nectar is much liked by the hive-bee is certain. yet the hive-bee can easily suck the nectar out of the incarnate clover. and pistils alone in another flower or on another plant. too slight to be appreciated by us. etc. in an exactly intermediate condition. and as a more complete separation of the sexes of our plant would be advantageous on the principle of the division of labour. Bearing such facts in mind. their habit of cutting holes and sucking the nectar at the bases of certain flowers. by which the separation of the sexes in plants of various kinds is apparently now in progress. so that whole fields of the red clover offer in vain an abundant supply of precious nectar to the hive-bee. No naturalist doubts the advantage of what has been called the "physiological division of labour. might profit a bee or other insect. which is visited by humble-bees alone.might commence. would be continually favoured or selected. The difference in the length of the corolla in the 98 ." hence we may believe that it would be advantageous to a plant to produce stamens alone in one flower or on one whole plant. Let us now turn to the nectar-feeding insects. as he expresses it. I could give many facts showing how anxious bees are to save time: for instance. according to Asa Gray. for I have repeatedly seen. In plants under culture and placed under new conditions of life. but I may add that some of the species of holly in North America are. It would take up too much space to show the various steps. to be a common plant. now if we suppose this to occur in ever so slight a degree under nature. we may suppose the plant of which we have been slowly increasing the nectar by continued selection. then. as pollen is already carried regularly from flower to flower. or. through dimorphism and other means. and that certain insects depended in main part on its nectar for food. but not out of the common red clover.. many hive-bees sucking the flowers through holes bitten in the base of the tube by humble bees. so that certain individuals would be able to obtain their food more quickly than others. are more or less dioeciously polygamous. 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. individuals with this tendency more and more increased. which with a very little more trouble they can enter by the mouth. and thus the communities to which they belonged would flourish and throw off many swarms inheriting the same peculiarities. sometimes the male organs and sometimes the female organs become more or less impotent. until at last a complete separation of the sexes might be effected.

when used in explaining the excavation of the deepest valleys or the formation of long lines of inland cliffs. or of any great and sudden modification in their structure. each profitable to the preserved being. either simultaneously or one after the other. if humble-bees were to become rare in any country. I am well aware that this doctrine of natural selection. it might be a great advantage to the plant to have a shorter or more deeply divided corolla. by the continued preservation of all the individuals which presented slight deviations of structure mutually favourable to each other. modified and adapted to each other in the most perfect manner. must be very trifling. so that the hive-bees should be enabled to suck its flowers. 99 . spoken of as trifling and insignificant. and as modern geology has almost banished such views as the excavation of a great valley by a single diluvial wave.two kinds of clover. that the Ligurian bee. and that these are visited by many hive-bees. which is generally considered a mere variety of the common hive-bee. it might be a great advantage to the hivebee to have a slightly longer or differently constructed proboscis. the flowers of the second crop are somewhat smaller. namely. Thus I can understand how a flower and a bee might slowly become. and which freely crosses with it." but we now seldom hear the agencies which we see still at work. for I have been assured that when red clover has been mown. exemplified in the above imaginary instances. which determines the visits of the hive-bee. I do not know whether this statement is accurate. Thus. as the fertility of this clover absolutely depends on bees visiting the flowers. as illustrative of geology. so will natural selection banish the belief of the continued creation of new organic beings. is open to the same objections which were first urged against Sir Charles Lyell's noble views on "the modern changes of the earth. On the other hand. is able to reach and suck the nectar of the red clover. Natural selection acts only by the preservation and accumulation of small inherited modifications. nor whether another published statement can be trusted. in a country where this kind of clover abounded.

that CLOSE interbreeding diminishes vigour and fertility. though I have the materials prepared for an ample discussion. all insects and some other large groups of animals. On The Intercrossing Of Individuals I must here introduce a short digression. either occasionally or habitually. On the belief that this is a law of nature. which is all that concerns us. notwithstanding that the plant's own anthers and pistil stand so near each other as almost to ensure selffertilisation. yet what a multitude of flowers have their anthers and stigmas fully exposed to the weather! If an occasional cross be indispensable. Knight and Kolreuter. and made so many experiments. that is. In the first place. This view was long ago doubtfully suggested by Sprengel. I think. or between individuals of the same variety but of another strain. the fullest freedom for the entrance of pollen from another 100 . gives vigour and fertility to the offspring. All vertebrate animals. 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 of generations. and on the other hand. but I must here treat the subject with extreme brevity. but in the case of hermaphrodites this is far from obvious. it may be asked. concur for the reproduction of their kind. In the case of animals and plants with separated sexes. understand several large classes of facts.4. such as the following. But still there are many hermaphrodite animals which certainly do not habitually pair. We shall presently see its importance. What reason. that with animals and plants a cross between different varieties. Every hybridizer knows how unfavourable exposure to wet is to the fertilisation of a flower. two individuals regularly unite for reproduction. which on any other view are inexplicable. showing. but that a cross with another individual is occasionally—perhaps at long intervals of time—indispensable. is there for supposing in these cases that two individuals ever concur in reproduction? As it is impossible here to enter on details. I must trust to some general considerations alone. pair for each birth. 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. Nevertheless there is reason to believe that with all hermaphrodites two individuals. and a vast majority of plants are hermaphrodites. we can. Modern research has much diminished the number of supposed hermaphrodites and of real hermaphrodites a large number pair. I have collected so large a body of facts. in accordance with the almost universal belief of breeders.

to the great good of the plant. which is visited by bees. or slowly move one after the other towards it. the contrivance seems adapted solely to ensure self-fertilisation. and no doubt it is useful for this end: but the agency of insects is often required to cause the stamens to spring forward. I raise plenty of seedlings. In very many other cases. as has been shown by Gartner. at least in my garden. and more recently Hildebrand and others have shown. it is scarcely possible for insects to fly from flower to flower. by insects. have their organs of fructification closely enclosed. Now. the former is so prepotent that it invariably and completely destroys. yet. either the anthers burst before the stigma is ready for fertilisation. it is hardly possible to raise pure seedlings. and not to carry pollen from one to the other. When the stamens of a flower suddenly spring towards the pistil. there are special contrivances which effectually prevent the stigma receiving pollen from its own flower. so largely do they naturally cross. as Kolreuter has shown to be the case with the barberry. Many flowers. just to touch with the same brush the anthers of one flower and then the stigma of another. it is well known that. seeds freely in my garden. far from selffertilisation being favoured. and it is sufficient. on the other hand. or the stigma is 101 . before the stigma of that individual flower is ready to receive them. and as I can confirm. Insects act like a camel-hair pencil. Another species of Lobelia. though there is no special mechanical contrivance to prevent the stigma receiving pollen from the same flower. but these almost invariably present beautiful and curious adaptations in relation to the visits of insects. there is a really beautiful and elaborate contrivance by which all the infinitely numerous pollengranules are swept out of the conjoined anthers of each flower. as well as from my own observations: for instance. for if a plant's own pollen and that from another species are placed on the same stigma. as in the great papilionaceous or pea-family. if closely-allied forms or varieties are planted near each other. it never sets a seed. So necessary are the visits of bees to many papilionaceous flowers. but it must not be supposed that bees would thus produce a multitude of hybrids between distinct species.individual will explain the above state of exposure of the organs. the influence of the foreign pollen. though by placing pollen from one flower on the stigma of another. in Lobelia fulgens. that their fertility is greatly diminished if these visits be prevented. which seems to have a special contrivance for self-fertilisation. to ensure fertilisation. as Sprengel. and in this very genus. In numerous other cases. as I could show from the works of Sprengel and others. and as this flower is never visited.

I raised 233 seedling cabbages from some plants of different varieties growing near each other. it may be objected that pollen could seldom be carried from tree to tree. 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. and must habitually be crossed. then. 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. I find to be the case in this country. but that nature has largely provided against it by giving to trees a strong tendency to bear flowers with separated sexes. and the result was as I anticipated. as if for the very purpose of self-fertilisation. as I found. for I have found that plants carefully protected from insects produce the full number of pods. 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. and at my request Dr. onion. be allowed to seed near each other. Dr. and that this is part of the general law of good being derived from the intercrossing of distinct individuals of the same species.ready before the pollen of that flower is ready. and of some other plants. Asa Gray those of the United States. but to this subject we shall return in a future chapter. Hooker tabulated the trees of New Zealand. so that these so-named dichogamous plants have in fact separated sexes. When distinct SPECIES are crossed the case is reversed. pollen must be regularly carried from flower to flower. On the other hand. When the sexes are separated. though placed so close together. How. although the male and female flowers may be produced on the same tree. and the pollen of each flower readily gets on its stigma without insect agency. mongrels: for instance. How strange are these facts! How strange that the pollen and stigmatic surface of the same flower. Hooker 102 . radish. That trees belonging to all orders have their sexes more often separated than other plants. and some even of these were not perfectly true. So it is with the reciprocally dimorphic and trimorphic plants previously alluded to. In the case of a large tree covered with innumerable flowers. for a plant's own pollen is always prepotent over foreign pollen. a large majority of the seedlings thus raised turn out. and at most only from flower to flower on the same tree. and this will give a better chance of pollen being occasionally carried from tree to tree. and Dr. and of these only 78 were true to their kind. I believe this objection to be valid. and flowers on the same tree can be considered as distinct individuals only in a limited sense.

though both are self-fertilising hermaphrodites. as far as function is concerned. Turning for a brief space to animals: various terrestrial species are hermaphrodites. but I have been enabled. the difference between them and unisexual species is. for owing to the nature of the fertilising element there are no means. Professor Huxley. it appears that with animals and plants an occasional intercross between distinct individuals is a very general. can be shown to be physically impossible. by which an occasional cross could be effected with terrestrial animals without the concurrence of two individuals. there are many self-fertilising hermaphrodites. but these all pair. namely. by a fortunate chance. under this point of view. the same result would follow as if they bore flowers with separated sexes. to discover a single hermaphrodite animal with the organs of reproduction so perfectly enclosed that access from without. though agreeing closely with each other in their whole organisation. such as the land-mollusca and earth-worms. But if. do sometimes cross. From these several considerations and from the many special facts which I have collected. but here the currents of water offer an obvious means for an occasional cross. a case of great difficulty. but which I am unable here to give. and the occasional influence of a distinct individual. are hermaphrodites. It must have struck most naturalists as a strange anomaly that. and some unisexual. As yet I have not found a single terrestrial animal which can fertilise itself. some species of the same family and even of the same genus. to prove that two individuals. if not universal. in fact. which offers so strong a contrast with terrestrial plants. I have as yet failed. very small. As in the case of flowers. both with animals and plants. Of aquatic animals. is intelligible on the view of an occasional cross being indispensable. after consultation with one of the highest authorities. 103 .informs me that the rule does not hold good in Australia: but if most of the Australian trees are dichogamous. law of nature. Cirripedes long appeared to me to present. I have made these few remarks on trees simply to call attention to the subject. all hermaphrodites do occasionally intercross. This remarkable fact. analogous to the action of insects and of the wind with plants.

But we shall see in the sixth chapter that intermediate varieties. Thus it will be under nature. under which term individual differences are always included. for as all organic beings are striving to seize on each place in the economy of nature. a highly important element of success. Unless favourable variations be inherited by some at least of the offspring. Intercrossing will chiefly affect those animals which unite for each birth and wander much. though in different degrees. the newly formed varieties will intercross on the confines of each. if the same species undergoes modification in different districts. and is. a breeder selects for some definite object. why should it prevail against natural selection? In the case of methodical selection. inhabiting intermediate districts. and which do not breed at a very quick rate. and this I find to be the case. if any one species does not become modified and improved in a corresponding degree with its competitors it will be exterminated. for instance birds. A great amount of variability. I believe. but as this tendency has not prevented man from forming by selection numerous domestic races. will evidently be favourable. Though nature grants long periods of time for the work of natural selection. she does not grant an indefinite period. notwithstanding that there is no separation of selected individuals. without intending to alter the breed. its several districts will almost certainly present different conditions of life. have a nearly common standard of perfection. his work will completely fail. Hence with animals of this nature. and if the individuals be allowed freely to intercross. for within a confined area. Circumstances Favourable For The Production Of New Forms Through Natural Selection This is an extremely intricate subject. The tendency to reversion may often check or prevent the work. by giving a better chance within any given period for the appearance of profitable variations. But if the area be large. improvement surely but slowly follows from this unconscious process of selection. But when many men. and then. with some place in the natural polity not perfectly occupied. will in the long run generally be supplanted by one of the adjoining varieties. varieties will generally be confined to separated countries. and all try to procure and breed from the best animals. With hermaphrodite organisms which cross only occasionally. A large number of individuals. nothing can be effected by natural selection. and likewise with animals which unite for each 104 . will tend to be preserved. all the individuals varying in the right direction.5. will compensate for a lesser amount of variability in each individual.

only through the principle of inheritance. and thus in the long run the influence of crosses. Intercrossing plays a very important part in nature by keeping the individuals of the same species. we have reason to believe that occasional intercrosses take place with all animals and plants. if not very large. we must not assume that free intercrossing would always eliminate the effects of natural selection. true and uniform in character. will also be thus prevented. a new and improved variety might be quickly formed on any one spot. solely by natural selection preserving similar favourable variations. as already stated. uniformity of character can be retained by them under the same conditions of life. Even with animals which unite for each birth. and through natural selection which will destroy any individuals departing from the proper type. from haunting different stations. nor conjugate. will be great. With respect to organic beings extremely low in the scale. so that natural selection will tend to modify all the varying individuals of the same species in the same manner. Moritz Wagner has lately published an interesting essay on this subject. and might there maintain itself in a body and afterward spread. the organic and inorganic conditions of life will generally be almost uniform. In a confined or isolated area. but. 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. and which cannot possibly intercross. On this principle nurserymen always prefer saving seed from a large body of plants. and has shown that the service rendered by isolation in preventing crosses between newly-formed varieties is probably greater even than I 105 . Even if these take place only at long intervals of time. even at rare intervals. If the conditions of life change and the form undergoes modification. so that the individuals of the new variety would chiefly cross together. or of the same variety. and which do not propagate rapidly. the young thus produced will gain so much in vigour and fertility over the offspring from long-continued self-fertilisation.birth. uniformity of character can be given to the modified offspring. Isolation also is an important element in the modification of species through natural selection. It will obviously thus act far more efficiently with those animals which unite for each birth. Intercrossing with the inhabitants of the surrounding districts. but which wander little and can increase at a rapid rate. or from the individuals of each variety preferring to pair together. which do not propagate sexually. that they will have a better chance of surviving and propagating their kind. as the chance of intercrossing is thus lessened. from breeding at slightly different seasons.

however. Although isolation is of great importance in the production of new species. has been most favourable for the production of new organic forms. and this may sometimes be of much importance. It likewise tends to increase the direct action of the physical conditions of life. and of spreading widely. isolation will give time for a new variety to be improved at a slow rate. such as an oceanic island. especially for the production of species which shall prove capable of enduring for a long period. that it gives a better chance of beneficial variations arising and of their being selected. elevation of the land. the immigration of better adapted organisms. as if all the forms of life were necessarily undergoing change through some innate law. for to ascertain whether a small isolated area. an isolated area be very small.. on the whole I am inclined to believe that largeness of area is still more important. in relation to the constitution of each organism. But from reasons already assigned I can by no means agree with this naturalist. Hence an oceanic island at first sight seems to have been highly favourable for the production of new species. If we turn to nature to test the truth of these remarks. either for or against natural selection. have been produced there and nowhere else in the world. Lapse of time is only so far important.supposed. and fixed. The importance of isolation is likewise great in preventing. and this we are incapable of doing. and this will retard the production of new species through natural selection. If. or a large open area like a continent. and its importance in this respect is great. Throughout a great and open area. that migration and isolation are necessary elements for the formation of new species. as we shall see in our chapter on Geographical Distribution. either from being surrounded by barriers. accumulated. by decreasing the chances of favourable variations arising. But we may thus deceive ourselves. we ought to make the comparison within equal times. although the number of the species inhabiting it is small. and look at any small isolated area. or from having very peculiar physical conditions. such as of climate.—that is. etc. Lastly. the total number of the inhabitants will be small. after any physical change in the conditions. yet of these species a very large proportion are endemic. not only will there be a better chance 106 . I state this because it has been erroneously asserted that the element of time has been assumed by me to play an all-important part in modifying species. 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. The mere lapse of time by itself does nothing.

In accordance with this view. They will thus play a more important part in the changing history of the organic world. Moreover. understand some facts which will be again alluded to in our chapter on Geographical Distribution. make a small area compared with that of the sea or of the land. we can understand how it is that the flora of Madeira. will often. although small isolated areas have been in some respects highly favourable for the production of new species. the race for life will have been less severe. like fossils. and will give rise to the greatest number of new varieties and species. All fresh water basins. as soon as it has been much improved.of favourable variations. to a certain extent. and if some of these many species become modified and improved. new forms will have been more slowly produced. and there will have been less modification and less extermination. will be those that will spread most widely. Consequently. 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. yet that the course of modification will generally have been more rapid on large areas. for instance. have concurred. will be able to spread over the open and continuous area. connect to a certain extent orders at present widely separated in the natural scale. also. so that the good effects of isolation will generally. resembles to a certain extent the extinct tertiary flora of Europe. that the new forms produced on large areas. perhaps. the competition between fresh water productions will have been less severe than elsewhere. also. which. but the conditions of life are much more complex from the large number of already existing species. And it is in fresh water basins that we find seven genera of Ganoid fishes. it is that continental productions have everywhere become so largely naturalised on islands. Each new form. Thus. as the Ornithorhynchus and Lepidosiren. remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world. or they will be exterminated. taken together. arising from the large number of individuals of the same species there supported. and what is more important. These anomalous forms may be called living fossils. have existed in a broken condition. from 107 . others will have to be improved in a corresponding degree. according to Oswald Heer. Finally. though now continuous. and old forms more slowly exterminated. and will thus come into competition with many other forms. Hence. they have endured to the present day. I conclude that. On a small island. we can. great areas. owing to former oscillations of level.

which generally take place very slowly. there will again have been very severe competition. as far as the extreme intricacy of the subject permits. and will have been subjected to severe competition. The occurrence of such places will often depend on physical changes. and on the immigration of better adapted forms being prevented. When. the most favoured or improved varieties will have been enabled to spread.having inhabited a confined area. so that new places in the polity of each island will have had to be filled up by the modification of the old inhabitants. it would often be long before differences of the right nature in various parts of the organisation might occur. and from having been exposed to less varied. I conclude that for terrestrial productions a large continental area. The result would often be greatly retarded by free intercrossing. fitted to endure for a long time and to spread widely. there will have been much extinction of the less improved forms. which has undergone many oscillations of level. competition. 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 therefore less severe. To sum up. and the relative proportional numbers of the various inhabitants of the reunited continent will again have been changed. and this will create new places. While the area existed as a continent the inhabitants will have been numerous in individuals and kinds. and time will have been allowed for the varieties in each to become well modified and perfected. 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. As some few of the old inhabitants become modified the mutual relations of others will often be disturbed. and thus to produce new species. by renewed elevation. will have been the most favourable for the production of many new forms of life. That natural selection generally act with extreme slowness I fully admit. and again there will have been a fair field for natural selection to improve still further the inhabitants. Many will exclaim that these several causes are amply sufficient to neutralise the power of natural selection. Although all the individuals of the same species differ in some slight degree from each other. but all this will take place very slowly. the islands were reconverted into a continental area. ready to be filled up by better adapted forms. the circumstances favourable and unfavourable for the production of new species through natural selection. But I do believe that natural selection will generally act 108 . I do not believe so.

I can see no limit to the amount of change. I further believe that these slow. only at long intervals of time. that is by the survival of the fittest. one with another and with their physical conditions of life. Slow though the process of selection may be. if feeble man can do much by artificial selection. intermittent results accord well with what geology tells us of the rate and manner at which the inhabitants of the world have changed. and only on a few of the inhabitants of the same region. to the beauty and complexity of the coadaptations between all organic beings. which may have been effected in the long course of time through nature's power of selection.very slowly. 109 .

But we may go further than this. which consequently endure. and tend to exterminate them. for as new forms are produced. That the number of specific forms has not indefinitely increased. Extinction Caused By Natural Selection This subject will be more fully discussed in our chapter on Geology.6. rare species will be less quickly modified or improved within any given period. Natural selection acts solely through the preservation of variations in some way advantageous. that as new species in the course of time are formed through natural selection. will the less favoured decrease and become rare. Consequently. generally come into the severest competition with each other. We see the same process of extermination 110 . We have seen that the species which are most numerous in individuals have the best chance of producing favourable variations within any given period. showing that it is the common and diffused or dominant species which offer the greatest number of recorded varieties.—which. We can see that any form which is represented by few individuals will run a good chance of utter extinction. The forms which stand in closest competition with those undergoing modification and improvement. many old forms must become extinct. they will consequently be beaten in the race for life by the modified and improved descendants of the commoner species. geology plainly tells us. And we have seen in the chapter on the Struggle for Existence that it is the most closely-allied forms. Rarity. during the progress of its formation. Owing to the high geometrical rate of increase of all organic beings. unless we admit that specific forms can go on indefinitely increasing in number. in the facts stated in the second chapter. will generally press hardest on its nearest kindred. during great fluctuations in the nature or the seasons. generally. others will become rarer and rarer. and species of the same genus or related genera.—varieties of the same species. will naturally suffer most. or from a temporary increase in the number of its enemies. that as the favoured forms increase in number. so. from having nearly the same structure. as geology tells us. We have evidence of this. constitution and habits. and we shall presently attempt to show why it is that the number of species throughout the world has not become immeasurably great. is the precursor to extinction. and it follows from this. Hence. but it must here be alluded to from being intimately connected with natural selection. From these several considerations I think it inevitably follows. each area is already fully stocked with inhabitants. each new variety or species. and finally extinct.

it is historically known that the ancient black cattle were displaced by the longhorns. In Yorkshire. and varieties of flowers. take the place of older and inferior kinds. through the selection of improved forms by man. Many curious instances could be given showing how quickly new breeds of cattle. sheep and other animals. and that these "were swept away by the short-horns" (I quote the words of an agricultural writer) "as if by some murderous pestilence.among our domesticated productions." 111 .

but like extremes. varieties are species in the process of formation. we may suppose that at an early period of history. but this alone would never account for so habitual and large a degree of difference as that between the species of the same genus. could never have been effected by the mere chance accumulation of similar variations during many successive generations. The early differences would be very slight. the differences would become greater. or are. Divergence Of Character The principle." 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. might cause one variety to differ in some character from its parents. the several breeds of pigeons. we must infer from most of the innumerable species throughout nature presenting well-marked differences. as I believe. incipient species. another fancier is struck by a pigeon having a rather longer beak. several important facts. but. 112 . I have sought light on this head from our domestic productions. struck by a pigeon having a slightly shorter beak. present slight and ill-defined differences. though having somewhat of the character of species—as is shown by the hopeless doubts in many cases how to rank them—yet certainly differ far less from each other than do good and distinct species. We shall here find something analogous. and on the acknowledged principle that "fanciers do not and will not admire a medium standard. does the lesser difference between varieties become augmented into the greater difference between species? That this does habitually happen. It will be admitted that the production of races so different as short-horn and Hereford cattle. and would be noted as forming two sub-breeds. even strongly-marked ones. Nevertheless according to my view. etc. Mere chance. for instance. race and cart horses. as we may call it. is of high importance. as I have called them. In practice. whereas varieties. in the course of time. and of stronger ones in the other. while those of another required stronger and bulkier horses. and the offspring of this variety again to differ from its parent in the very same character and in a greater degree. then. from the continued selection of swifter horses in the one case.. these sub-breeds would become converted into two well-established and distinct breeds. a fancier is. How. the men of one nation or district required swifter horses. or with shorter and shorter beaks.7. Ultimately after the lapse of centuries. Again. In the first place. the supposed prototypes and parents of future well-marked species. varieties. As has always been my practice. and explains. which I have designated by this term.

would not have been used for breeding. and some perhaps becoming less carnivorous. So it will be with plants. being enabled to feed on new kinds of prey. and so be enabled to increase in numbers. 113 . some inhabiting new stations. causing differences. constitution. frequenting water. from the simple circumstance that the more diversified the descendants from any one species become in structure. a greater number of plants and a greater weight of dry herbage can be raised in the latter than in the former case. a greater number of individual plants of this species. then. The more diversified in habits and structure the descendants of our carnivorous animals become.As the differences became greater. we see in man's productions the action of what may be called the principle of divergence. and is thus striving. if any one species of grass were to go on varying. It has been experimentally proved. 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). would succeed in living on the same piece of ground. Hence. And we know that each species and each variety of grass is annually sowing almost countless seeds. including its modified descendants. being neither very swift nor very strong. and the breeds to diverge in character. the inferior animals with intermediate characters. if they vary—for otherwise natural selection can effect nothing. either dead or alive. But how. 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. of which the number that can be supported in any country has long ago arrived at its full average. and a similar plot be sown with several distinct genera of grasses. for instance. both from each other and from their common parent. as do the distinct species and genera of grasses. and will thus have tended to disappear. climbing trees. Take the case of a carnivorous quadruped. steadily to increase. Here. 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. 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. If its natural power of increase be allowed to act. at first barely appreciable. though in a very slight degree. the more places they will be enabled to occupy. We can clearly discern this in the case of animals with simple habits. and habits. by so much will they be better enabled to seize on many and widely diversified places in the polity of nature. it may be asked. and the varieties were continually selected which differed from each other in the same manner.

we always find great diversity in its inhabitants. de Candolle has well remarked. Most of the animals and plants which live close round any small piece of ground. So it is with the plants and insects on small and uniform islets: also in small ponds of fresh water. three feet by four in size. to the utmost to increase in number. take the rank of species. Asa Gray's "Manual of the Flora of the Northern United States. and may be said to be striving to the utmost to live there. and where the contest between individual and individual must be very severe. that where they come into the closest competition. 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. which shows how much these plants differed from each other. determine that the inhabitants. It might also. especially if freely open to immigration. is seen under many natural circumstances." 260 naturalised plants are enumerated. perhaps. that floras gain by naturalisation. which thus jostle each other most closely. the advantages of diversification of structure. have been expected that naturalised plants would have belonged to a few groups more especially adapted to certain stations in their new homes. and varieties. for these are commonly looked at as specially created and adapted for their own country. 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. shall. proportionally with the number of the native genera and species. the most distinct varieties of any one species of grass would have the best chance of succeeding and of increasing in numbers. I found that a piece of turf. The same principle is seen in the naturalisation of plants through man's agency in foreign lands. when rendered very distinct from each other. Consequently. in his great and admirable work. In an extremely small area. belong to what we call different genera and orders. with the accompanying differences of habit and constitution. To give a single instance: in the last edition of Dr. it is seen. and thus of supplanting the less distinct varieties. and these belonged to eighteen genera and to eight orders.as it may be said. and Alph. far more in new genera than in new species. 114 . in the course of many thousand generations. which had been exposed for many years to exactly the same conditions. as a general rule. supported twenty species of plants. could live on it (supposing its nature not to be in any way peculiar). For instance. but. But the case is very different. The truth of the principle that the greatest amount of life can be supported by great diversification of structure.

We thus see that these naturalised plants are of a highly diversified nature. our carnivorous. 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. and have there become naturalised. It may be doubted. the more widely and perfectly the animals and plants are diversified for different habits of life. draws most nutriment from these substances. 115 . to a large extent. So in the general economy of any land. no less than 100 genera are not there indigenous. and we may at least infer that diversification of structure. from the indigenes. The advantage of diversification of structure in the inhabitants of the same region is. could successfully compete with these well-developed orders. could hardly compete with a set more perfectly diversified in structure. amounting to new generic differences. or flesh alone. and feebly representing. which are divided into groups differing but little from each other. in fact. as Mr. would be profitable to them. moreover. for out of the 162 naturalised genera. and rodent mammals. Waterhouse and others have remarked. They differ. In the Australian mammals.and these belong to 162 genera. so will a greater number of individuals be capable of there supporting themselves. with their organisation but little diversified. By considering the nature of the plants or animals which have in any country struggled successfully with the indigenes. for instance. whether the Australian marsupials. we see the process of diversification in an early and incomplete stage of development. No physiologist doubts that a stomach by being adapted to digest vegetable matter alone. 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. ruminant. and thus a large proportional addition is made to the genera now living in the United States. A set of animals.

Only those variations which are in some way profitable will be preserved or naturally selected. such as would be thought worthy of record in a systematic work. they are not supposed all to appear simultaneously. these species are supposed to resemble each other in unequal degrees.8. 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 branching and diverging dotted lines of unequal lengths proceeding from (A). Now let us see how this principle of benefit being derived from divergence of character. belonging to a genus large in its own country. nor are they all supposed to endure for equal periods. and are thus enabled to encroach on places occupied by other beings. 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. also. as is so generally the case in nature. The Probable Effects Of The Action Of Natural Selection Through Divergence Of Character And Extinction. Let (A) be a common. but of the most diversified nature. seen that the species. species (A) is supposed to have produced two fairly well-marked 116 . And here the importance of the principle of benefit derived from divergence of character comes in. on an average more species vary in large genera than in small genera. combined with the principles of natural selection and of extinction. On The Descendants Of A Common Ancestor After the foregoing discussion. may represent its varying offspring. tends to act. vary more than do the rare and restricted species. The intervals between the horizontal lines in the diagram. The accompanying diagram will aid us in understanding this rather perplexing subject. and as is represented in the diagram by the letters standing at unequal distances. The variations are supposed to be extremely slight. because as we saw in the second chapter. and the varying species of the large genera present a greater number of varieties. and varying species. When a dotted line reaches one of the horizontal lines. After a thousand generations. may represent each a thousand or more generations. which has been much compressed. which are the commonest and most widely-diffused. but often after long intervals of time. We have. a sufficient amount of variation is supposed to have been accumulated to form it into a fairly well-marked variety. and is there marked by a small numbered letter. Let A to L represent the species of a genus large in its own country. widely-diffused. I have said a large genus.

producing only a single variety. 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 more considerably from their common parent (A). these two varieties be variable. namely m2 and s2. a large genus in its own country. Moreover.varieties. and under a condensed and simplified form up to the fourteen-thousandth generation. these two varieties. being only slightly modified forms. and some failing to produce any. owing to the principle of divergence. then. variety a1 is supposed in the diagram to have produced variety a2. and may or may not produce more than one modified descendant. But I must here remark that I do not suppose that the process ever goes on so regularly as is represented in the diagram. Variety m1 is supposed to have produced two varieties. If. We may continue the process by similar steps for any length of time. the most divergent of their variations will generally be preserved during the next thousand generations. and the tendency to variability is in itself hereditary. and commonly in nearly the same manner as did their parents. which will. the more places they will be enabled to seize on. In the diagram the process is represented up to the ten-thousandth generation. consequently they will likewise tend to vary. Nor do I suppose that the most divergent varieties are invariably preserved: a medium form may often long endure. And after this interval. Thus the varieties or modified descendants of the common parent (A). but in a more and more modified condition. But as a general rule. and then again undergoes modification. And all these circumstances are favourable to the production of new varieties. some of the varieties. differing from each other. and this will depend on infinitely complex relations. after each thousand generations. it is far more probable that each form remains for long periods unaltered. the more diversified in structure the descendants from any one species can be rendered. In our diagram the 117 . namely a1 and m1. These two varieties will generally still be exposed to the same conditions which made their parents variable. nor that it goes on continuously. they will also partake of those more general advantages which made the genus to which the parent-species belonged. will tend to inherit those advantages which made their parent (A) more numerous than most of the other inhabitants of the same country. though in itself made somewhat irregular. will generally go on increasing in number and diverging in character. some producing two or three varieties. differ more from (A) than did variety a1. and the more their modified progeny will increase.

species are multiplied and genera are formed. f10. If we suppose the amount of change between each horizontal line in our diagram to be excessively small. Thus. and m10. 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). if all the lines proceeding from (A) were removed. But these breaks are imaginary. and so destroy. we get eight 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. will. it is probable. and the number of modified descendants will not be increased. as I believe. belonging to a large genus. species (A) is supposed to have produced three forms. from each other and from their common parent. As all the modified descendants from a common and widely-diffused species. although the amount of divergent modification may have been augmented. after intervals long enough to allow the accumulation of a considerable amount of divergent variation. from having diverged in character during the successive generations. but perhaps unequally. a10. these three forms may still be only well-marked varieties. In the same way the English racehorse and English pointer have apparently both gone on slowly diverging in character from their original stocks. After ten thousand generations. 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. In some cases no doubt the process of modification will be confined to a single line of descent. will have come to differ largely.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. often take the place of. all descended from (A). This case would be represented in the diagram. The modified offspring from the later and more highly improved branches in the lines of descent. but we have only to suppose the steps in the process of modification to be more numerous or greater in amount. excepting that from a1 to a10. marked by the letters between a14 and m14. and might have been inserted anywhere. 118 . By continuing the same process for a greater number of generations (as shown in the diagram in a condensed and simplified manner). without either having given off any fresh branches or races. which. will tend to partake of the same advantages which made their parent successful in life.

another of our principles. and the nearly extreme species (I). the modified offspring of a species get into some distinct country. after ten thousand generations. as well as the original parent-species itself. species (A) and all the earlier varieties will have become extinct. 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. If. by analogous steps. In the diagram I have assumed that a second species (I) has produced. being replaced by eight new species (a14 to m14). will generally tend to become extinct. and have given rise to new varieties and species. In any genus. 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. will generally tend to produce the greatest number of modified descendants. The other nine species (marked by capital letters) of our original genus. 119 . either two well-marked varieties (w10 and z10) or two species. may for long but unequal periods continue to transmit unaltered descendants. according to the amount of change supposed to be represented between the horizontal lines. 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). however. then. So it probably will be with many whole collateral lines of descent. marked by the letters n14 to z14.In a large genus it is probable that more than one species would vary. six new species. our diagram be assumed to represent a considerable amount of modification. Hence all the intermediate forms between the earlier and later states. After fourteen thousand generations. represented in the diagram. constitution and structure. will have played an important part. If. and species (I) will be replaced by six (n14 to z14) new species. that is between the less and more improved states of a the same species. But during the process of modification. both may continue to exist. and this is shown in the diagram by the dotted lines unequally prolonged upwards. the species which are already very different in character from each other. which will be conquered by later and improved lines. are supposed to have been produced. in which offspring and progenitor do not come into competition. 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. namely that of extinction. or become quickly adapted to some quite new station. as those which have largely varied.

b14 and f14. These two species (A and I). has transmitted descendants to this late stage of descent. K. than to the others. 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. standing nearly at the extreme end of the original genus. But as the original species (I) differed largely from (A). therefore. o14. from having diverged at an earlier period from a5. H. The six descendants from (I) will form two sub-genera or genera. also (and this is a very important consideration). and lastly. and m14. The original species of our genus were supposed to resemble each other in unequal degrees. e14. 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. but likewise some of the original species which were most nearly related to their parents. Hence very few of the original species will have transmitted offspring to the fourteen-thousandth generation. descended from the original eleven species. will be nearly related from having recently branched off from a10. C. owing to inheritance alone. The new species in our diagram. the six descendants from (I) will. and may constitute a sub-genus or a distinct genus. 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. the two groups. and species (I) more to G. but. moreover. are supposed to have gone on diverging in different directions. not only their parents (A) and (I). were also supposed to be very common and widely diffused species. Their modified descendants. from having diverged at the first commencement of the process of modification. will be allied to each other in a widely different manner. species (A) being more nearly related to B. It seems. fourteen in number at the fourteen-thousandth generation. will be nearly related one to the other. will be in some degree distinct from the three first-named species. which connected the original species 120 . extremely probable that they will have taken the places of. moreover. p14. L. The intermediate species. will be widely different from the other five species. differ considerably from the eight descendants from (A). Of the eight descendants from (A) the three marked a14.But we may go further than this. as is so generally the case in nature. q14. will now be fifteen in number. Owing to the divergent tendency of natural selection. so that they must originally have had some advantage over most of the other species of the genus. and D than to the other species. and thus exterminated. The new species. so as to have become adapted to many related places in the natural economy of their country.

converging in sub-branches downwards towards a single point. for the extinct species lived at various remote epochs when the branching lines of descent had diverged less. We shall. and the eight descendants from (A). We shall also have two very distinct genera descended from (I). have to refer again to this subject. and we can understand this fact. will form three very distinct genera. from two or more species of the same genus. or even as distinct sub-families. the forms marked a14 to p14. but rather between types of the two groups. which is supposed not to have diverged much in character. which. Hence the six new species descended from (I). to the formation of genera alone. have all become. In the diagram each horizontal line has hitherto been supposed to represent a thousand generations. except (F). And the two or more parent-species are supposed to be descended from some one species of an earlier genus. intermediate in character between existing groups. and have left no descendants. the supposed progenitor of our several new sub-genera and genera. this point represents a species. with those now living. it may also represent a section of the successive strata of the earth's crust including extinct remains.(A) and (I). and I think we shall then see that the diagram throws light on the affinities of extinct beings. as I believe. or genera. It is worth while to reflect for a moment on the character of the new species F14. but each may represent a million or more generations. 121 . those marked b14 and f14. we suppose the amount of change represented by each successive group of diverging dotted lines to be great. Being descended from a form that stood between the parent-species (A) and (I). yet are often. either unaltered or altered only in a slight degree. In this case its affinities to the other fourteen new species will be of a curious and circuitous nature. I see no reason to limit the process of modification. that two or more genera are produced by descent with modification. But as these two groups have gone on diverging in character from the type of their parents. but to have retained the form of (F). will have to be ranked as very distinct genera. Thus it is. in the diagram. extinct. it will be in some degree intermediate in character between the two groups descended from these two species. If. though generally belonging to the same orders. when we come to our chapter on geology. now supposed to be extinct and unknown. in some degree. families. and those marked o14 to m14. and every naturalist will be able to call such cases before his mind. In our diagram this is indicated by the broken lines beneath the capital letters. the new species (F14) will not be directly intermediate between them. as now explained.

according to this view. Looking still more remotely to the future. as all the descendants of the same species form a class. which are all trying to increase in number. for as natural selection acts through one form having some advantage over other forms in the struggle for existence. for a long period. Although few of the most ancient species have left modified descendants. we can predict that the groups of organic beings which are now large and triumphant. This. or orders. which have as yet suffered least extinction. at remote geological periods. it will chiefly act on those which already have some advantage. We have seen that in each country it is the species belonging to the larger genera which oftenest present varieties or incipient species. and leave no modified descendants. Looking to the future. formerly most extensively developed. But which groups will ultimately prevail. will constantly tend to supplant and destroy the earlier and less improved sub-groups. have now become extinct. Small and broken groups and sub-groups will finally disappear. And the two new families. Within the same large group. the struggle for the production of new and modified descendants will mainly lie between the larger groups. will. One large group will slowly conquer another large group. continue to increase. and these are supposed to be descended from some still more ancient and unknown form. These two groups of genera will thus form two distinct families. the earth may 122 .differing widely from the descendants of (A). might have been expected. I shall have to return to this subject in the chapter on classification. owing to the continued and steady increase of the larger groups. are descended from two species of the original genus. indeed. but I may add that as. and. and which are least broken up. we may predict that. extremely few of the more ancient species have transmitted descendants to the present day. a multitude of smaller groups will become utterly extinct. of the species living at any one period. the later and more highly perfected sub-groups. and consequently that. from branching out and seizing on many new places in the polity of nature. Hence. and thus lessen its chance of further variation and improvement. or orders. for we know that many groups. we can understand how it is that there exist so few classes in each main division of the animal and vegetable kingdoms. extremely few will transmit descendants to a remote futurity. yet. that is. no man can predict. and the largeness of any group shows that its species have inherited from a common ancestor some advantage in common. reduce its number. according to the amount of divergent modification supposed to be represented in the diagram.

as at the present day. families. orders and classes.have been almost as well peopled with species of many genera. 123 .

among which some naturalists rank those as highest which. namely. Among the vertebrata the degree of intellect and an approach in structure to man clearly come into play. so that the mature animal cannot be called higher than its larva. On The Degree To Which Organisation Tends To Advance Natural selection acts exclusively by the preservation and accumulation of variations. which are beneficial under the organic and inorganic conditions to which each creature is exposed at all periods of life. 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). as I should be inclined to add. 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. the amount of differentiation of the parts of the same organic being. but there are cases. Von Baer's standard seems the most widely applicable and the best. in which several parts of the structure become less perfect.9. while other naturalists rank the common bony or teleostean fishes as the highest. is an advantage to each being. 124 . This improvement inevitably leads to the gradual advancement of the organisation of the greater number of living beings throughout the world. inasmuch as they are most strictly fish. and differ most from the other vertebrate classes. whereas other botanists. and here some botanists rank those plants as highest which have every organ.like. look at the plants which have their several organs much modified and reduced in number as the highest. approach nearest to amphibians. probably with more truth. inasmuch as in this state they perform their functions better. or. We see still more plainly the obscurity of the subject by turning to plants. fully developed in each flower. natural selection clearly leads towards this standard: for all physiologists admit that the specialisation of organs. If we take as the standard of high organisation. stamens and pistils. among which the standard of intellect is of course quite excluded. for instance. for naturalists have not defined to each other's satisfaction what is meant by an advance in organisation. like the sharks. petals. in the adult state. and their specialisation for different functions. the completeness of the division of physiological labour. The ultimate result is that each creature tends to become more and more improved in relation to its conditions. as Milne Edwards would express it. But we shall see how obscure this subject is if we look. to fishes. as with certain parasitic crustaceans. as sepals. But here we enter on a very intricate subject.

Nearly the same remarks are applicable. On the other hand. to the co-existence of man and the ornithorhynchus—among fishes. and might remain for indefinite ages in their present lowly condition. have remained for an enormous period in nearly their present state. by natural selection. we can see. 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. unimproved or but little improved. these forms would be left. Science has not as yet proved the truth of this belief. for every naturalist who has dissected some of the beings now ranked as very low in the scale. as the infusoria and rhizopods. And it may be asked what advantage. And geology tells us that some of the lowest forms. On our theory the continued existence of lowly organisms offers no difficulty. who believed in an innate and inevitable tendency towards perfection in all organic beings. to the co-existence of mammals and fish—among mammalia. as far as we can see. must have been struck with their really wondrous and beautiful organisation. for instance. if we look to the different grades of organisation within the same great group. to 125 . 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. 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.and hence the accumulation of variations tending towards specialisation is within the scope of natural selection. whatever the future may reveal. 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. If it were no advantage. 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. for natural selection. 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. 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. or the survival of the fittest. in the vertebrata. But it may be objected that if all organic beings thus tend to rise in the scale. would it be to an infusorian animalcule—to an intestinal worm—or even to an earth-worm. how is it that throughout the world a multitude of the lowest forms still exist. to be highly organised.

and where their scanty numbers have retarded the chance of favourable variations arising. co-exist in South America in the same region with numerous monkeys. members of the shark family would not tend to supplant the lancelet. it has been asked. probably. presented the simplest structure. which latter fish in the extreme simplicity of its structure approaches the invertebrate classes. I believe that many lowly organised forms now exist throughout the world. for the lancelet. so that warm-blooded mammals when inhabiting the water lie under a disadvantage in having to come continually to the surface to breathe. has as sole companion and competitor on the barren sandy shore of South Brazil. for the high advancement of certain whole classes. from various causes. lowly organised forms appear to have been preserved to the present day. edentata. could the first step in the advancement or differentiation of parts have arisen? Mr. to the highest grade would not lead to their taking the place of fishes. or of certain members of each class. In some cases.the co. and this requires aerial respiration. from inhabiting confined or peculiar stations. on the whole. namely. and rodents. and probably interfere little with each other. But mammals and fish hardly come into competition with each other. Finally. an anomalous annelid. With fishes.existence of the shark and the lancelet (Amphioxus). does not at all necessarily lead to the extinction of those groups with which they do not enter into close competition. as we shall hereafter see. The three lowest orders of mammals. the advancement of the whole class of mammals. as I hear from Fritz Muller. In some cases variations or individual differences of a favourable nature may never have arisen for natural selection to act on and accumulate. as we may believe. may have advanced and be still advancing throughout the world. where they have been subjected to less severe competition. In no case. 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. Although organisation. and more liable to be put out of order and injured. Looking to the first dawn of life. or of certain members in this class. how. as soon as simple unicellular organisms came by growth or division to be compounded of 126 . In some few cases there has been what we must call retrogression or organisation. as being of a more delicate nature. Herbert Spencer would probably answer that. yet the scale will always present many degrees of perfection. has time sufficed for the utmost possible amount of development. when all organic beings. marsupials. Physiologists believe that the brain must be bathed by warm blood to be highly active.

127 . until many forms had been produced: variations in a single species inhabiting an isolated station might be beneficial. however. as I remarked towards the close of the introduction. no natural selection. no one ought to feel surprise at much remaining as yet unexplained on the origin of species. consequently. But as we have no facts to guide us. and thus the whole mass of individuals might be modified. if we make due allowance for our profound ignorance on the mutual relations of the inhabitants of the world at the present time. his law "that homologous units of any order become differentiated in proportion as their relations to incident forces become different" would come into action. and. speculation on the subject is almost useless. It is.several cells. and still more so during past ages. an error to suppose that there would be no struggle for existence. or two distinct forms might arise. But. or became attached to any supporting surface.

these being due to causes far too intricate to be followed out—on the nature of the variations which have been preserved or selected. and it is not surprising that dissimilar substances should sometimes assume the same form. which had originally differed in a marked manner. If this had occurred. on inheritance (in itself a fluctuating element) from innumerable progenitors. but I fully admit that the mutual relations of organic beings are more important. and the balance of evidence is opposed to any such an admission. should ever afterwards converge so closely as to lead to a near approach to identity throughout their whole organisation. it is conceivable that these might approach each other so closely that they would have all to be classed under the same genus. etc. namely on the variations which have arisen. It is incredible that the descendants of two organisms. the organic conditions of life must become more and more complex. and in a still higher degree on the surrounding organisms with which each being has come into competition—and lastly. but with organic beings we should bear in mind that the form of each depends on an infinitude of complex relations. As far as mere inorganic conditions are concerned. and this depends on the surrounding physical conditions. Convergence Of Character Mr. he apparently believes). moisture. independently of genetic connection. it seems probable that a sufficient number of species would soon become adapted to all considerable diversities of heat. together with divergence of character. The shape of a crystal is determined solely by the molecular forces. H. Watson has also objected that the continued action of natural selection.. we should meet with the same form. 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.10. Watson thinks that I have overrated the importance of divergence of character (in which. If two species belonging to two distinct though allied genera. and therefore no limit to the 128 . and that convergence. as it may be called. had both produced a large number of new and divergent forms. Consequently there seems at first no limit to the amount of profitable diversification of structure. Mr. however. all of which have had their forms determined through equally complex relations.C. and as the number of species in any country goes on increasing. would tend to make an indefinite number of specific forms. has likewise played a part. recurring in widely separated geological formations. and thus the descendants of two distinct genera would converge into one.

The process of extermination in such cases would be rapid. therefore. will. the process of giving birth to new specific forms would thus be retarded. depending so largely as it does on physical conditions. Lastly. and each species will become rare if the number of species in any country becomes indefinitely increased. and that from the middle part of this same period. will tend to spread and supplant many others. Hooker has recently shown that in the southeast corner of Australia. on the principal often explained. of red deer in Scotland and of bears in Norway. consequently. many European plants have become naturalised. Rare species. each or nearly each species will be represented by few individuals. that from an early part of the tertiary period the number of species of shells. Dr. When any species becomes very rare. 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. if an area be inhabited by very many species. Imagine the extreme case of as many species as individuals in England. which has already beaten many competitors in its own home. and such species will be liable to extermination from accidental fluctuations in the nature of the seasons or in the number of their enemies. consequently they will tend to supplant and exterminate several species in several areas. there are many invaders from different quarters of the globe. the endemic Australian species have been greatly reduced in number. 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. de Candolle has shown that those species which spread widely tend generally to spread VERY widely. Alph.number of species which might be produced. But geology shows us. How much weight to attribute to these several considerations I will not pretend to say. the number of mammals has not greatly or at all increased. whereas the production of new species must always be slow. but conjointly they must limit in each country the tendency to an indefinite augmentation of specific forms. 129 . a dominant species. and this I am inclined to think is the most important element. where. and the first severe winter or very dry summer would exterminate thousands on thousands of species. authors have thought that this comes into play in accounting for the deterioration of the aurochs in Lithuania. etc. which support such an astonishing number of species. and thus check the inordinate increase of specific forms throughout the world. close interbreeding will help to exterminate it. present within a given period few favourable variations. apparently.

It leads to the improvement of each creature in relation to its organic and inorganic conditions of life. and this cannot be disputed. by so much the more can a large number be supported on the area. also. season or year. geology plainly declares. constitution. But if variations useful to any organic being ever do occur. for the more organic beings diverge in structure. or the survival of the fittest. and how largely extinction has acted in the world's history. Therefore. on the principle of qualities being inherited at corresponding ages. to be advantageous to them. in the same manner as so many variations have occurred useful to man. Natural selection. and from the strong principle of inheritance. But we have already seen how it entails extinction. of which we see proof by looking to the inhabitants of any small spot. then. and consequently. and habits. habits and constitution. must be judged by the general tenour and balance of evidence given in the following chapters. and these characters will be transmitted to one sex or to both sexes. I have called natural selection. This principle of preservation. causing an infinite diversity in structure. Nevertheless. seed. leads to divergence of character. and to the productions naturalised in foreign lands. these will tend to produce offspring similarly characterised. in most cases. to what must be regarded as an advance in organisation. if there be. Natural selection. assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life. it would be a most extraordinary fact if no variations had ever occurred useful to each being's own welfare. Whether natural selection has really thus acted in adapting the various forms of life to their several conditions and stations. considering the infinite complexity of the relations of all organic beings to each other and to their conditions of life. owing to their geometrical rate of increase. during the modification of the descendants of any one species. or young as easily as the adult. 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. low and simple forms will long endure if well fitted for their simple conditions of life. and this certainly cannot be disputed. a severe struggle for life at some age. and during the incessant 130 . Sexual selection will also give characters useful to the males alone in their struggles or rivalry with other males. Summary Of Chapter If under changing conditions of life organic beings present individual differences in almost every part of their structure.11. according to the form of inheritance which prevails. can modify the egg.

Natural selection. but it is explained through inheritance and the complex action of natural selection. in the same manner as species and groups of species have at all times overmastered other species in the great battle for life. and to overtop and kill the surrounding twigs and branches. in the manner which we everywhere behold—namely. and the generally well defined distinctions between the innumerable organic beings in each class throughout the world. We have seen that it is the common. and this connexion of the former and present buds by ramifying 131 . may be explained. and those produced during former years may represent the long succession of extinct species. species of distinct genera much less closely related. The green and budding twigs may represent existing species. when the tree was young. as has just been remarked. forming sections and sub-genera. as we have seen illustrated in the diagram. I believe this simile largely speaks the truth. but seem clustered round points. were themselves once. Thus the small differences distinguishing varieties of the same species. species of the same genus less closely and unequally related.struggle of all species to increase in numbers. and widely ranging species. forming sub-families. varieties of the same species most closely related. entailing extinction and divergence of character. belonging to the larger genera within each class. the better will be their chance of success in the battle for life. till they equal the greater differences between species of the same genus. leads to divergence of character and to much extinction of the less improved and intermediate forms of life. budding twigs. and classes. The limbs divided into great branches. the widely diffused. and these tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. and genera related in different degrees. the more diversified the descendants become. On these principles. orders. and these into lesser and lesser branches. The affinities of all the beings of the same class have sometimes been represented by a great tree. steadily tend to increase. and these round other points. If species had been independently created. the nature of the affinities. sub-classes. no explanation would have been possible of this kind of classification. At each period of growth all the growing twigs have tried to branch out on all sides. or even of distinct genera. The several subordinate groups in any class cannot be ranked in a single file. 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. which vary most. and so on in almost endless cycles. families. subordinate to groups.

and covers the surface with its ever-branching and beautiful ramifications. which in some small degree connects by its affinities two large branches of life. As we here and there see a thin. so with the species which lived during long-past geological periods. and which by some chance has been favoured and is still alive on its summit. yet survive and bear the other branches. families. and these. As buds give rise by growth to fresh buds. which fills with its dead and broken branches the crust of the earth. now grown into great branches. branch out and overtop on all sides many a feebler branch.branches may well represent the classification of all extinct and living species in groups subordinate to groups. so by generation I believe it has been with the great Tree of Life. very few have left living and modified descendants. only two or three. and which are known to us only in a fossil state. Of the many twigs which flourished when the tree was a mere bush. 132 . From the first growth of the tree. straggling branch springing from a fork low down in a tree. many a limb and branch has decayed and dropped off. if vigorous. and which has apparently been saved from fatal competition by having inhabited a protected station. and these fallen branches of various sizes may represent those whole orders. so we occasionally see an animal like the Ornithorhynchus or Lepidosiren. and genera which have now no living representatives.

133 . 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. organs of flight and of vision — Acclimatisation — Correlated variation — Compensation and economy of growth — False correlations — Multiple. rudimentary. combined with natural selection.Chapter 5 Laws Of Variation Effects of changed conditions — Use and disuse.

cannot be attributed simply to such action. I have hitherto sometimes spoken as if the variations—so common and multiform with organic beings under domestication. The direct action of changed conditions leads to definite or indefinite results. Some authors believe it to be as much the function of the reproductive system to produce individual differences. as to make the child like its parents. and indirectly through the reproductive system. In the following cases the conditions seem to have produced some slight definite effect: E. In the former case the nature of the organism is such that it yields readily. but it serves to acknowledge plainly our ignorance of the cause of each particular variation. In the latter case the organisation seems to become plastic. and Wollaston is convinced that residence near the sea affects the colours of insects. when growing near the sea-shore. when subjected to certain conditions. Forbes asserts that shells at their southern limit. the nature of the organism. MoquinTandon gives a list of plants which. Mr. In the first chapter I attempted to show that changed conditions act in two ways. such as of climate. food. and the nature of the conditions. But we may safely conclude that the innumerable complex co-adaptations of structure. which we see throughout nature between various organic beings. than when living near the coast or on islands. or slight deviations of structure.. have acted in a definite manner. But the fact of variations and monstrosities occurring much more frequently under domestication than under nature. Gould believes that birds of the same species are more brightly coloured under a clear atmosphere. of course is a wholly incorrect expression. This. are more brightly coloured than those of the same species from further north or from a greater depth.1. and in a lesser degree with those under nature—were due to chance. In all cases there are two factors. etc. 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 when living in shallow water. and the greater variability of species having wide ranges than of those with restricted ranges. or nearly all. the individuals become modified in the same way. which is much the most important of the two. It is very difficult to decide how far changed conditions. and all. directly on the whole organisation or on certain parts alone. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence. but this certainly does not always hold good. and we have much fluctuating variability. 134 .

In one sense the conditions of life may be said. Thus. than on a tendency to vary. not only to cause variability. 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.have their leaves in some degree fleshy. but likewise to include natural selection. either directly or indirectly. and how much to the definite action of the conditions of life. 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. 135 . Again. although living under the most opposite climates. it is well known to furriers that animals of the same species have thicker and better fur the further north they live. 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. and it is this latter agency which answers to the survival of the fittest under nature. we clearly see that the two elements of change are distinct. but it is the will of man which accumulates the variations in certain direction. or not varying at all. variability is in some manner excited. When a variation is of the slightest use to any being. of dissimilar varieties being produced under apparently the same external conditions. though not elsewhere fleshy. and. 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. But when man is the selecting agent. we cannot tell how much to attribute to the accumulative action of natural selection. due to causes of which we are quite ignorant. of species keeping true. innumerable instances are known to every naturalist. on the other hand. for the conditions determine whether this or that variety shall survive. Such considerations as these incline me to lay less weight on the direct action of the surrounding conditions.

and not one had even a relic left. We may believe that the progenitor of the ostrich genus had habits like those of the bustard. now inhabiting or which lately inhabited several oceanic islands. and their rudimentary condition in some other genera. As Professor Owen has remarked. and that such modifications are inherited. as the size and weight of its body were increased during successive generations. as efficiently as many quadrupeds. In the Onites apelles the tarsi are so habitually lost that the insect has been described as not having them.2. for we know not the parent-forms. but many animals possess structures which can be best explained by the effects of disuse. The ostrich indeed inhabits continents. of many male dung-feeding beetles are often broken off. As Controlled By Natural Selection From the facts alluded to in the first chapter. Cunningham. Effects Of The Increased Use And Disuse Of Parts. should make us cautious in denying this tendency. In the Ateuchus or sacred beetle of the Egyptians. yet there are several in this state. and is exposed to danger from which it cannot escape by flight. Kirby has remarked (and I have observed the same fact) that the anterior tarsi. and that. or feet. and disuse diminished them. Hence. they are totally deficient. tenanted by no beasts of prey. but the remarkable cases observed by Brown-Sequard in guinea-pigs. The logger-headed duck of South America can only flap along the surface of the water. The evidence that accidental mutilations can be inherited is at present not decisive. but in a rudimentary condition. there is no greater anomaly in nature than a bird that cannot fly. until they became incapable of flight. while the adults have lost this power. of the inherited effects of operations. can fly. In some other genera they are present. by kicking its enemies. not as cases of inherited mutilations. for as many dung-feeding beetles are 136 . it will perhaps be safest to look at the entire absence of the anterior tarsi in Ateuchus. but as due to the effects of long-continued disuse. it is probable that the nearly wingless condition of several birds. but it can defend itself. has been caused by disuse. Under free nature we have no standard of comparison by which to judge of the effects of long-continued use or disuse. As the larger ground-feeding birds seldom take flight except to escape danger. he examined seventeen specimens in his own collection. I think there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts. its legs were used more and its wings less. according to Mr. and has its wings in nearly the same condition as the domestic Aylesbury duck: it is a remarkable fact that the young birds.

out of the 550 species (but more are now known) inhabiting Madeira. These several considerations make me believe that the wingless condition of so many Madeira beetles is mainly due to the action of natural selection. Wollaston. and in some cases are quite covered by skin and fur. combined probably with disuse. until the wind lulls and the sun shines. have. or by giving up the attempt and rarely or never flying. either from its wings having been ever so little less perfectly developed or from indolent habit. on the other hand. This is quite compatible with the action of natural selection. Wollaston has discovered the remarkable fact that 200 beetles. For during many successive generations each individual beetle which flew least. Wollaston. are here almost entirely absent. and that. namely. lie much concealed. elsewhere excessively numerous. that the proportion of wingless beetles is larger on the exposed Desertas than in Madeira itself. or mainly due to natural selection. This state of 137 . and. of the twenty-nine endemic genera.generally found with their tarsi lost. Wollaston suspects. their wings not at all reduced. Mr. the tendency of natural selection to enlarge or to reduce the wings. whereas it would have been better for the bad swimmers if they had not been able to swim at all and had stuck to the wreck. must habitually use their wings to gain their subsistence. that beetles in many parts of the world are very frequently blown to sea and perish. as certain flower-feeding coleoptera and lepidoptera. as observed by Mr. therefore the tarsi cannot be of much importance or be much used by these insects. this must happen early in life. so strongly insisted on by Mr. as Mr. are so far deficient in wings that they cannot fly. but even enlarged. no less than twenty-three have all their species in this condition! Several facts. that the beetles in Madeira. and especially the extraordinary fact. those beetles which most readily took to flight would oftenest have been blown to sea. and thus destroyed. which absolutely require the use of their wings. that certain large groups of beetles. would depend on whether a greater number of individuals were saved by successfully battling with the winds. will have had the best chance of surviving from not being blown out to sea. it would have been better for the good swimmers if they had been able to swim still further. The eyes of moles and of some burrowing rodents are rudimentary in size. For when a new insect first arrived on the island. As with mariners shipwrecked near a coast. The insects in Madeira which are not ground-feeders. and which. In some cases we might easily put down to disuse modifications of structure which are wholly.

their loss may be attributed to disuse. slowly migrated by successive generations from the outer world into the deeper and deeper recesses of the Kentucky caves. and the similarity which is exhibited in a few forms between the Mammoth Cave (in Kentucky) and the caves in Carniola. and if so. natural selection would aid the effects of disuse. two of which were captured by Professor Silliman at above half a mile distance from the mouth of the cave. very close similarity in their organisation and affinities might have been expected. is even more subterranean in its habits than the mole. or Ctenomys. having been inflammation of the nictitating membrane." On my view we must suppose that American animals. It is difficult to imagine conditions of life more similar than deep limestone caverns under a nearly similar climate. belonging to the most different classes. as appeared on dissection. It is well known that several animals. with the adhesion of the eyelids and growth of fur over them. the cause. In some of the crabs the foot-stalk for the eye remains. as did 138 . namely. acquired a dim perception of objects. a reduction in their size. and these animals. so that. the tuco-tuco. This is certainly not the case if we look at the two whole faunas.the eyes is probably due to gradual reduction from disuse. the cave-rat (Neotoma). As it is difficult to imagine that eyes. One which I kept alive was certainly in this condition. with respect to the insects alone. are blind. that they were frequently blind. though useless. As frequent inflammation of the eyes must be injurious to any animal. could be in any way injurious to animals living in darkness. in accordance with the old view of the blind animals having been separately created for the American and European caverns. the stand for the telescope is there. might in such case be an advantage. a burrowing rodent. and as eyes are certainly not necessary to animals having subterranean habits. otherwise than as a very plain expression of that analogy which subsists generally between the fauna of Europe and of North America. who had often caught them. after having been exposed for about a month to a graduated light. In one of the blind animals. but aided perhaps by natural selection. In South America. the eyes were lustrous and of large size. and therefore not in the profoundest depths. and I was assured by a Spaniard. which inhabit the caves of Carniola and Kentucky. as I am informed by Professor Silliman. though the eye is gone. though the telescope with its glasses has been lost. having in most cases ordinary powers of vision. Schiodte has remarked: "We are accordingly prevented from considering the entire phenomenon in any other light than something purely local.

may formerly have ranged over both continents. affinities to the other inhabitants of that continent." These remarks of Schiodte's it should be understood. the Amblyopsis. 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. and natural selection will often have effected other changes. By the time that an animal had reached. disuse will on this view have more or less perfectly obliterated its eyes. have been accommodated to surrounding circumstances. after numberless generations. for. and some of the European cave-insects are very closely allied to those of the surrounding country. Notwithstanding such modifications. and as is the case with the blind Proteus. prepare the transition from light to darkness. with reference to the reptiles of 139 . but to distinct species. Animals not far remote from ordinary forms. 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. that the species. while they were furnished with eyes. and then have become extinct. but it is possible that the progenitors of these several species. That several of the inhabitants of the caves of the Old and New Worlds should be closely related. and which. we might expect from the well-known relationship of most of their other productions. and. excepting in their present secluded abodes. as Agassiz has remarked in regard to the blind fish. Next follow those that are constructed for twilight. for it is natural that an insect already deprived of vision should readily become adapted to dark caverns. last of all. And this is the case with some of the American cave-animals. as they extended themselves into darkness. have not as yet been found anywhere except in caves. Far from feeling surprise that some of the cave-animals should be very anomalous. we might expect still to see in the caveanimals of America. the deepest recesses. as a compensation for blindness. those destined for total darkness. such as an increase in the length of the antennae or palpi. yet those which inhabit the several caves of Europe and America are distinct. apply not to the same. As a blind species of Bathyscia is found in abundance on shady rocks far from caves. Murray observes. the loss of vision in the cave species of this one genus has probably had no relation to its dark habitation. and in those of Europe to the inhabitants of the European continent. as Mr. Another blind genus (Anophthalmus) offers this remarkable peculiarity. We have some evidence of this gradation of habit. as I hear from Professor Dana.European animals into the caves of Europe. and whose formation is quite peculiar.

140 .Europe. owing to the less severe competition to which the scanty inhabitants of these dark abodes will have been exposed. I am only surprised that more wrecks of ancient life have not been preserved.

we have evidence with some few plants. But whether or not this adaptation is in most cases very close. or conversely. naturally habituated to different temperatures. In regard to animals. or more than. and from the number of plants and animals brought from different countries which are here perfectly healthy. in the amount of rain requisite for seeds to germinate. raised from seed collected by Dr. and this leads me to say a few words on acclimatisation. and not because they were subsequently found capable of far-extended transportation. As we may infer that our domestic animals were originally chosen by uncivilised man because they were useful. and I could give other cases. As it is extremely common for distinct species belonging to the same genus to inhabit hot and cold countries. Watson on European species of plants brought from the Azores to England. But the degree of adaptation of species to the climates under which they live is often overrated. Mr. H. were found to possess in this country different constitutional powers of resisting cold. So again. by adaptation to particular climates. acclimatisation must be readily effected during a long course of descent. of their becoming. if it be true that all the species of the same genus are descended from a single parent-form. in the time of sleep. the common and extraordinary 141 . analogous observations have been made by Mr.. and because they bred readily under confinement. within historical times. Thwaites informs me that he has observed similar facts in Ceylon. We may infer this from our frequent inability to predict whether or not an imported plant will endure our climate. several authentic instances could be adduced of species having largely extended. 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. and conversely. to a certain extent. etc. nor do we know that they have subsequently become specially acclimatised to their new homes. Acclimatisation Habit is hereditary with plants. so as to be better fitted for them than they were at first. that is. their range from warmer to colder latitudes. Hooker from the same species growing at different heights on the Himalayas. as in the period of flowering.C. but we do not positively know that these animals were strictly adapted to their native climate. 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. they become acclimatised: thus the pines and rhododendrons.3. many succulent plants cannot endure a damp climate. though in all ordinary cases we assume such to be the case.

Hence adaptation to any special climate may be looked at as a quality readily grafted on an innate wide flexibility of constitution. however. the capacity of enduring the most different climates by man himself and by his domestic animals. In treatises on many kinds of cultivated plants. both from analogy and from the incessant advice given in agricultural works. be due to habit. natural selection would inevitably tend to preserve those individuals which were born with constitutions best adapted to any country which they inhabited. On the other hand. push the foregoing argument too far. and 142 . to be very cautious in transporting animals from one district to another. and as most of these varieties are of recent origin. even in the ancient Encyclopaedias of China. That habit or custom has some influence. but as examples of a very common flexibility of constitution. and now have a far wider range than any other rodent. ought not to be looked at as anomalies.capacity in our domestic animals of not only withstanding the most different climates. this is strikingly shown in works on fruit-trees published in the United States. common to most animals. and the fact of the extinct elephant and rhinoceros having formerly endured a glacial climate. and how much to the natural selection of varieties having different innate constitutions. they cannot owe their constitutional differences to habit. on account of the probable origin of some of our domestic animals from several wild stocks: the blood. certain varieties are said to withstand certain climates better than others. which is never propagated in England by seed. of a tropical and arctic wolf may perhaps be mingled in our domestic breeds. 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. but they have been transported by man to many parts of the world. into action. brought. How much of the acclimatisation of species to any peculiar climate is due to mere habit. We must not. whereas the living species are now all tropical or subtropical in their habits. and on many an island in the torrid zones. for instance. and how much to both means combined. in which certain varieties are habitually recommended for the northern and others for the southern states. I must believe. is an obscure question. the result must. under peculiar circumstances. The rat and mouse cannot be considered as domestic animals. 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. The case of the Jerusalem artichoke. for they live under the cold climate of Faroe in the north and of the Falklands in the south. but of being perfectly fertile (a far severer test) under them. On this view. I think.

for it is now as tender as ever it was! The case. and then again get seed from these seedlings. of the kidney-bean has been often cited for a similar purpose. and of this fact I have myself observed striking instances. and then collect seed from the few survivors. and sometimes overmastered by. his kidneybeans so early that a very large proportion are destroyed by frost. new varieties have not been produced. also. the experiment cannot be said to have been even tried. have.of which. and with much greater weight. On the whole. we may conclude that habit. as proving that acclimatisation cannot be effected. consequently. Nor let it be supposed that differences in the constitution of seedling kidneybeans never appear. played a considerable part in the modification of the constitution and structure. has even been advanced. but until some one will sow. in some cases. 143 . during a score of generations. but that the effects have often been largely combined with. or use and disuse. with care to prevent accidental crosses. with the same precautions. the natural selection of innate variations. for an account has been published how much more hardy some seedlings are than others.

the shape of the body and the manner of swallowing determine the position and form of several of the most important viscera. most imperfectly understood. this is often seen in monstrous plants: and nothing is more common than the union of homologous parts in normal structures. Is. The nature of the bond is frequently quite obscure. as in the union of the petals into a tube. between their feathered feet and skin betwixt the outer toes. 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. at an early embryonic period. In snakes. other parts become modified. or between the presence of more or 144 . and if this had been of any great use to the breed. and which are necessarily exposed to similar conditions. that when slight variations in any one part occur and are accumulated through natural selection. that variations of structure arising in the young or larvae naturally tend to affect the structure of the mature animal. Homologous parts. or in pigeons. Others believe that the shape of the pelvis in the human mother influences by pressure the shape of the head of the child. Hilaire has forcibly remarked that certain malconformations frequently. varying together. and which. coexist without our being able to assign any reason. are identical in structure. I do not doubt. it might probably have been rendered permanent by natural selection. This is a very important subject.4. Hard parts seem to affect the form of adjoining soft parts. or between the tortoiseshell colour and the female sex. 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. 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. tend to cohere. These tendencies. M. Geoffroy St. One of the most obvious real cases is. during its growth and development. as has been remarked by some authors. according to Schlegel. Correlated Variation I mean by this expression that the whole organisation is so tied together. for the lower jaw is believed by some anatomists to be homologous with the limbs. in the front and hind legs. may be mastered more or less completely by natural selection: thus a family of stags once existed with an antler only on one side. and no doubt wholly different classes of facts may be here easily confounded together. The several parts which are homologous. and even in the jaws and limbs. and that others rarely.

but there are so many exceptions to this rule. or. the central flower thus becoming peloric or regular. that in many pelargoniums the two upper petals in the central flower of the truss often lose their patches of darker colour. and this difference is often accompanied with the partial or complete abortion of the reproductive organs. I think it can hardly be accidental that the two orders of mammals which are most abnormal in their dermal covering. the nectary is not quite aborted but is much shortened. It might have been thought that the development of the raypetals. are likewise on the whole the most abnormal in their teeth. Sprengel's idea that the ray-florets serve to attract insects. with the future colour of its plumage. scaly ant-eaters. These differences have sometimes been attributed to the pressure of the involucra on the florets. than that of the difference between the outer and inner flowers in some Compositous and Umbelliferous plants. and as a striking case of correlation. again. We know. and the shape of the seeds in the rayflorets of some Compositae countenances this idea. But in some of these plants the seeds also differ in shape and sculpture. and therefore of natural selection. for in some Compositae the seeds of the outer and inner florets differ. I know of no case better adapted to show the importance of the laws of correlation and variation. as an instance of this fact. Possibly these several differences may be connected with the different flow of nutriment towards the central and external flowers.. without any difference in the corolla. With respect to the development of the corolla. but with the Umbelliferae it is by no means. the adherent nectary is quite aborted. is highly probable. Cetacea (whales) and Edentata (armadilloes. viz. Mivart has remarked. I may add. But with respect to the 145 . for instance. or necessary for the fertilisation of these plants. that with irregular flowers those nearest to the axis are most subject to peloria. the relation between the hair and the teeth in the naked Turkish dog. that it has little value. independently of utility. at least. natural selection may have come into play. though here no doubt homology comes into play? With respect to this latter case of correlation. etc.less down on the young pigeon when first hatched. by drawing nourishment from the reproductive organs causes their abortion. When the colour is absent from only one of the two upper petals. the daisy. or to their mutual pressure. that is to become abnormally symmetrical. Everyone is familiar with the difference between the ray and central florets of. Hooker informs me. but this can hardly be the sole case. whose agency is highly advantageous. and if so. the species with the densest heads which most frequently differ in their inner and outer flowers. and when this occurs.). as Dr. as Mr.

can be in any way beneficial. De Candolle has remarked that winged seeds are never found in fruits which do not open. Some other correlations are apparently due to the manner in which natural selection can alone act. for an ancient progenitor may have acquired through natural selection some one modification in structure. gain an advantage over others less well fitted for wide dispersal. which are not always correlated with any difference in the corolla. 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. which were a little better adapted to be wafted by the wind. viewed by systematists as of high value. would naturally be thought to be in some necessary manner correlated. Alph. as far as we can judge. of the slightest service to the species. For instance. and which in truth are simply due to inheritance. and these two modifications. for in this case alone could the seeds. Hence modifications of structure. I should explain this rule by the impossibility of seeds gradually becoming winged through natural selection. some other and independent modification. may be wholly due to the laws of variation and correlation. after thousands of generations.seeds. and. We may often falsely attribute to correlated variation structures which are common to whole groups of species. unless the capsules were open. 146 . without being. having been transmitted to a whole group of descendants with diverse habits. it seems impossible that their differences in shape.

" I think this holds true to a certain extent with our domestic productions: if nourishment flows to one part or organ in excess. but many good observers. and furnished with great nerves and muscles. would 147 . namely. believe in its truth. Now the saving of a large and complex structure. but in the parasitic and protected Proteolepas. a large tuft of feathers on the head is generally accompanied by a diminished comb. at least in excess. more especially botanists. and. at about the same time. as Goethe expressed it. I will not. the actual withdrawal of nutriment from one part owing to the excess of growth in another and adjoining part. Compensation And Economy Of Growth The elder Geoffroy and Goethe propounded. This is the case with the male Ibla. When the seeds in our fruits become atrophied. I can thus only understand a fact with which I was much struck when examining cirripedes. their law of compensation or balancement of growth. on the one hand. to another part. it loses more or less completely its own shell or carapace. the whole anterior part of the head is reduced to the merest rudiment attached to the bases of the prehensile antennae.5. becomes less useful. that natural selection is continually trying to economise in every part of the organisation. that when a cirripede is parasitic within another cirripede and is thus protected. for I see hardly any way of distinguishing between the effects. With species in a state of nature it can hardly be maintained that the law is of universal application. also. and likewise some other facts. 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. for it will profit the individual not to have its nutriment wasted in building up a useless structure. it rarely flows. that some of the cases of compensation which have been advanced. or. may be merged under a more general principle. If under changed conditions of life a structure. the fruit itself gains largely in size and quality. its diminution will be favoured. when rendered superfluous. of a part being largely developed through natural selection and another and adjoining part being reduced by the same process or by disuse. In our poultry. and of which many other instances could be given: namely. and a large beard by diminished wattles. nature is forced to economise on the other side. however. thus it is difficult to get a cow to give much milk and to fatten readily. "in order to spend on one side. here give any instances. The same varieties of the cabbage do not yield abundant and nutritious foliage and a copious supply of oil-bearing seeds. on the other hand. before useful. I suspect.

that natural selection may perfectly well succeed in largely developing an organ without requiring as a necessary compensation the reduction of some adjoining part.be a decided advantage to each successive individual of the species. And conversely. natural selection will tend in the long run to reduce any part of the organisation. superfluous. as soon as it becomes. each would have a better chance of supporting itself. by less nutriment being wasted. as I believe. through changed habits. for in the struggle for life to which every animal is exposed. 148 . Thus. without by any means causing some other part to be largely developed in a corresponding degree.

can act solely through and for the advantage of each being. Natural selection. that beings which stand low in the scale of nature are more variable than those which are higher. and I will here only add that their variability seems to result from their uselessness. Multiple. Geoffroy St. and consequently from natural selection having had no power to check deviations in their structure. is a sign of low organisation. Rudimentary parts.6. the foregoing statements accord with the common opinion of naturalists. 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. have further remarked that multiple parts are extremely liable to vary in structure. Rudimentary. as is generally admitted. when it occurs in lesser numbers. 149 . Hilaire. and as long as the same part has to perform diversified work. I presume that lowness here means that the several parts of the organisation have been but little specialised for particular functions. As "vegetative repetition." to use Professor Owen's expression. is constant. we can perhaps see why it should remain variable. and the stamens in polyandrous flowers) the number is variable. both with varieties and species. it should never be forgotten. that is. as remarked by Is. whereas the number of the same part or organ. In the same way that a knife which has to cut all sorts of things may be of almost any shape. that when any part or organ is repeated many times in the same individual (as the vertebrae in snakes. The same author as well as some botanists. And Lowly-Organised Structures Are Variable It seems to be a rule. We shall have to recur to this subject. whilst a tool for some particular purpose must be of some particular shape. are apt to be highly variable.

but I hope that I have made due allowances for them. also. I shall. but the rule would not apply here. I particularly attended to Mr. The rule being so plainly applicable in the case of secondary sexual characters. the wing of the bat is a most abnormal structure in the class of mammals. when displayed in any unusual manner. whilst investigating this order. secondary sexual characters. used by Hunter. and which cannot possibly be here introduced. But that our rule is not confined to secondary sexual characters is clearly shown in the case of hermaphrodite cirripedes. The rule applies to males and females. It should be understood that the rule by no means applies to any part. as it illustrates the rule in its largest application. 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. Tends To Be Highly Variable Several years ago I was much struck by a remark to the above effect made by Mr. as they seldom offer remarkable secondary sexual characters. and I am fully convinced that the rule almost always holds good. I am aware of several causes of error. may be due to the great variability of these characters. Waterhouse's remark. A Part Developed In Any Species In An Extraordinary Degree Or Manner.7. these valves present a marvellous amount of diversification. I can only state my conviction that it is a rule of high generality. give a list of all the more remarkable cases. in a future work. but in the several species of one genus. however unusually developed. 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. unless it be unusually developed in one species or in a few species in comparison with the same part in many closely allied species. Professor Owen. The rule applies very strongly in the case of secondary sexual characters. in every sense of the word. Thus. In Comparison With The Same Part In Allied Species. the homologous valves in the different species being sometimes wholly unlike in shape. I will here give only one. Waterhouse. Pyrgoma. and they differ extremely little even in distinct genera. but more rarely to females. very important structures. The term. relates to characters which are attached to one sex. seems to have come to a nearly similar conclusion. because the whole group of bats possesses wings. whether or not displayed in any unusual manner—of which fact I think there can be little doubt. but are not directly connected with the act of reproduction. The opercular valves of sessile cirripedes (rock barnacles) are. and the amount of variation in the individuals of the same species is so great that 150 .

as in that of the short-faced tumbler. that part (for instance. and have been modified through natural selection. I cannot make out that it applies to plants. and thus the organisation is left in a fluctuating condition. First let me make some preliminary remarks. I think we can obtain some light.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. 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. it is notoriously difficult to breed nearly perfect birds. There may truly be said to be a constant struggle going on between. and in those which have been but little specialised for any particular purpose. with all its parts as we now see them. had not the great variability in plants made it particularly difficult to compare their relative degrees of variability. vary extremely little. these being the points now mainly attended to by English fanciers. and this would have seriously shaken my belief in its truth. on the one hand. In rudimentary organs. are also eminently liable to variation. and the rule certainly seems to hold good in this class. we see a nearly parallel case. and perhaps in polymorphic groups. the power of steady selection to keep the breed true. and see what a prodigious amount of difference there is in the beak of tumblers. the tendency to reversion to a less perfect state. When we see any part or organ developed in a remarkable degree or manner in a species. any part or the whole animal be neglected.. the fair presumption is that it is of high importance to that species: nevertheless it is in this case eminently liable to variation. for in such cases natural selection either has not or cannot come into full play. But what here more particularly concerns us is. in the beak and wattle of carriers. As with birds the individuals of the same species. Look at the individuals of the same breed of the pigeon. in our domestic animals. and no selection be applied. and. in the carriage and tail of fantails. etc. I have particularly attended to them. as well as an innate tendency to new variations. I can see no explanation. In the long run selection gains the day. inhabiting the same country. which at the present time are undergoing rapid change by continued selection. 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- 151 . than do the species belonging to other distinct genera. Even in the same sub-breed. Why should this be so? On the view that each species has been independently created. that those points in our domestic animals. But on the view that groups of species are descended from some other species. If. many departing widely from the standard. on the other hand.

But as long as selection is rapidly going on. And this. is the case. Hence. as a general rule. it must have existed. has been transmitted in approximately the same condition to many modified descendants. for an immense period in nearly the same state. and the tendency to reversion and variability on the other hand. as species rarely endure for more than one geological period. This period will seldom be remote in any extreme degree. compared with the other species of the same genus. That the struggle between natural selection on the one hand. When a part has been developed in an extraordinary manner in any one species. as in the case of the wing of the bat. An extraordinary amount of modification implies an unusually large and long-continued amount of variability. when an organ. I see no reason to doubt. and that the most abnormally developed organs may be made constant. Now let us turn to nature. as it may be called. much variability in the parts undergoing modification may always be expected. 152 . 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. 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. which has continually been accumulated by natural selection for the benefit of the species. 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. and by the continued rejection of those tending to revert to a former and less modified condition. according to our theory. will in the course of time cease. still present in a high degree.faced strain. however abnormal it may be. But as the variability of the extraordinarily developed part or organ has been so great and long-continued within a period not excessively remote. and thus it has come not to be more variable than any other structure. we might. I am convinced. 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.

It would be almost superfluous to adduce evidence in support of the statement. On the ordinary view of each species having been independently created. DIFFERS considerably in closely-allied species. it is often VARIABLE in the individuals of the same species. and its variation would be a more unusual circumstance. that when an author remarks with surprise that some important organ or part. that the more an organ normally differs in the different species of the same group. but with respect to important characters. Or to state the case in another manner: the points in which all the species of a genus resemble each other. when it sinks in value and becomes only of specific value. I have chosen this example because the explanation which most naturalists would advance is not here applicable. 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. that specific characters are more variable than generic. the colour would be only a specific character.8. be more variable than those parts which are closely alike in the several species? I do not see that any explanation can be given. true. and in which they 153 . But on the view that species are only strongly marked and fixed varieties. Specific Characters More Variable Than Generic Characters The principle discussed under the last heading may be applied to our present subject. have to return to this point in the chapter on Classification. the colour would become a generic character. Hilaire apparently entertains no doubt. Geoffroy St. I believe this explanation is partly. namely. And this fact shows that a character. though its physiological importance may remain the same. Something of the same kind applies to monstrosities: at least Is. or conversely. why should that part of the structure. but if all the species had blue flowers. however. yet only indirectly. because they are taken from parts of less physiological importance than those commonly used for classing genera. and which have thus come to differ. I shall. It is notorious that specific characters are more variable than generic. I have repeatedly noticed in works on natural history. which is generally very constant throughout a large group of species. 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. which differs from the same part in other independently created species of the same genus. the more subject it is to anomalies in the individuals. and no one would be surprised at one of the blue species varying into red. that ordinary specific characters are more variable than generic. which is generally of generic value. often becomes variable.

and as these specific characters have varied and come to differ since the period when the species branched off from a common progenitor. the points in which species differ from other species of the same genus are called specific characters. it is not probable that they should vary at the present day. for it can rarely have happened that natural selection will have modified several distinct species. fitted to more or less widely different habits. 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. 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. 154 . and subsequently have not varied or come to differ in any degree. and these characters may be attributed to inheritance from a common progenitor. On the other hand. or only in a slight degree. are called generic characters.differ from allied genera.

Secondary Sexual Characters Variable I think it will be admitted by naturalists. sexual selection will have had a wide scope for action. The cause of the original variability of these characters is not manifest. for instance. but in certain genera the neuration differs in the different species. in which secondary sexual characters are strongly displayed. and as the differences in these cases are of a very unusual nature. "In Pontella. It is a remarkable fact. whatever part of the structure of the common progenitor. 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. as Westwood has remarked. It will also be admitted that species of the same group differ from each other more widely in their secondary sexual characters. that secondary sexual characters are highly variable. as they are highly variable." 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. variations of this part would. as have the two sexes of any one species. the neuration of the wings is a character of the highest importance. or of its early descendants. for they are accumulated by sexual selection. compare. that several minute crustaceans offer excellent illustrations of this law. the relation can hardly be accidental. the amount of difference between the males of gallinaceous birds. but only gives fewer offspring to the less favoured males. as it does not entail death. Consequently. which is less rigid in its action than ordinary selection. with the amount of difference between the females. 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.9. without my entering on details. be 155 . The same number of joints in the tarsi is a character common to very large groups of beetles. Lubbock has recently remarked. became variable. for instance. because common to large groups. it is highly probable. than in other parts of their organisation. the number varies greatly and the number likewise differs in the two sexes of the same 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. but in the Engidae. Whatever the cause may be of the variability of secondary sexual characters. Again in the fossorial hymenoptera. but we can see why they should not have been rendered as constant and uniform as others. Sir J. Of this fact I will give in illustration the first two instances which happen to stand on my list. and likewise in the two sexes of the same species.

and for ordinary purposes. that secondary sexual and ordinary specific differences are generally displayed in the same parts of the organisation. and thus having been adapted for secondary sexual. if it be common to a whole group of species. are all principles closely connected together. All being mainly due to the species of the same group being the descendants of a common progenitor. 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. however extraordinarily it may be developed.taken advantage of by natural and sexual selection. 156 . to sexual selection being less rigid than ordinary selection. or to fit the males to struggle with other males for the possession of the females. and the slight degree of variability in a part. and to variations in the same parts having been accumulated by natural and sexual selection. I conclude that the greater variability of specific characters. to natural selection having more or less completely. according to the lapse of time. than of generic characters. or those which are possessed by all the species. then. and likewise to fit the two sexes of the same species to each other. from whom they have inherited much in common. or those which distinguish species from species. Finally. that the great variability of secondary sexual characters and their great difference in closely allied species. overmastered the tendency to reversion and to further variability. 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.

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. white loins. when acted on by similar unknown influences. I think. plants which several botanists rank as varieties produced by cultivation from a common parent: if this be not so. We may. characters not possessed by the aboriginal rock-pigeon. As all these marks are characteristic of the parent rock-pigeon. According to the ordinary view of each species having been independently created. these coloured marks are eminently liable to appear in the crossed offspring of two distinct and 157 . because. and to these a third may be added. not to the vera causa of community of descent. the case will then be one of analogous variation in two so-called distinct species. Or Reverts To Some Of The Characters Of An Early Progenitor These propositions will be most readily understood by looking to our domestic races. a bar at the end of the tail. of slaty-blue birds with two black bars on the wings. confidently come to this conclusion. however. Distinct Species Present Analogous Variations. Similar cases occurring with insects under natural conditions have lately been discussed with much ability by Mr. but to three separate yet closely related acts of creation. 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. of the Swedish turnip and ruta-baga. Walsh. and not of a new yet analogous variation appearing in the several breeds. namely. So That A Variety Of One Species Often Assumes A Character Proper To An Allied Species. Many similar cases of analogous variation have been observed by Naudin in the great gourd family. and with feathers on the feet. in the enlarged stems. the occasional appearance in all the breeds. With pigeons. present sub-varieties with reversed feathers on the head. who has grouped them under his law of equable variability.10. The most distinct breeds of the pigeon. I presume that no one will doubt that this is a case of reversion. we have another case. we should have to attribute this similarity in the enlarged stems of these three plants. the fantail. with the outer feathers externally edged near their bases with white. in countries widely apart. or as commonly called roots. In the vegetable kingdom we have a case of analogous variation. these then are analogous variations in two or more distinct races. and by various authors in our cereals. and a consequent tendency to vary in a like manner. namely. as we have seen. the common turnip.

probably for hundreds of generations. reappears after a great number of generations. but in which BOTH parents have lost some character which their progenitor possessed. it might be expected that they would occasionally vary in an analogous manner. it is generally believed that a tendency to reversion is retained by this remnant of foreign blood. A mere tendency to produce a rudiment is indeed sometimes thus inherited. under unknown favourable conditions. is only 1 in 2048. which very rarely produces a blue bird. for a dozen or even a score of generations. so that the varieties of two or more species would resemble each other. In a breed which has not been crossed. the offspring occasionally show for many generations a tendency to revert in character to the foreign breed—some say. But when a breed has been crossed only once by some other breed. it is probable that there is a latent tendency in each generation to produce blue plumage. It might further be expected that the species of the same genus would 158 . and at last. to reproduce the lost character might. With the barb-pigeon. the proportion of blood. As all the species of the same genus are supposed to be descended from a common progenitor. only a well-marked and permanent variety. is not greater than that of quite useless or rudimentary organs being similarly transmitted.differently coloured breeds. the most probable hypothesis is. for all that we can see to the contrary. After twelve generations. as was formerly remarked. for the preservation of all functionally important characters will have been determined through natural selection. and yet. When a character which has been lost in a breed. whether strong or weak. for instance. from one ancestor. or that a variety of one species would resemble in certain characters another and distinct species. The abstract improbability of such a tendency being transmitted through a vast number of generations. this other species being. not that one individual suddenly takes after an ancestor removed by some hundred generations. to use a common expression. No doubt it is a very surprising fact that characters should reappear after having been lost for many. But characters exclusively due to analogous variation would probably be of an unimportant nature. the tendency. in accordance with the different habits of the species. with the several marks. beyond the influence of the mere act of crossing on the laws of inheritance. as we see. but that in each successive generation the character in question has been lying latent. according to our view. and in this case there is nothing in the external conditions of life to cause the reappearance of the slaty-blue. be transmitted for almost any number of generations. is developed.

not indeed as affecting any important character. as it were. and which would not probably have all appeared together from simple variation. sometimes to find the varying offspring of a species assuming characters which are already present in other members of the same group. The stripe on the shoulder is sometimes double. give one curious and complex case. we cannot distinguish between reversionary and analogous characters. we did not know that the parent rock-pigeon was not feather-footed or turn-crowned. I believe this to be true. It has been asserted that these are plainest in the foal. But the best evidence of analogous variations is afforded by parts or organs which are generally constant in character. whether such characters in our domestic breeds were reversions or only analogous variations. I have collected a long list of such cases. And this undoubtedly is the case. I can only repeat that such cases certainly occur. has been described without either spinal or shoulder stripe. but from occurring in several species of the same genus. and is very variable in length and outline. 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. and from inquiries which I have made. but NOT an albino. Hence. and what are new but analogous variations. The ass sometimes has very distinct transverse bars on its legs. but which occasionally vary so as to resemble. that they have in varying assumed some of the characters of the others. unless all these closely allied forms be considered as independently created species. or actually quite 159 . which are correlated with this tint. It is a case almost certainly of reversion. I lie under the great disadvantage of not being able to give them. as before. like those on the legs of a zebra.occasionally exhibit reversions to long-lost characters. the same part or organ in an allied species. other species of the same genus. A white ass. for instance. and this shows. If. A considerable catalogue. and these stripes are sometimes very obscure. The difficulty in distinguishing variable species is largely due to the varieties mocking. although under nature it must generally be left doubtful. partly under domestication and partly under nature. yet we ought. but we might have inferred that the blue colour was a case of reversion from the number of the markings. I will. but here. in some degree. could be given of forms intermediate between two other forms. and seem to me very remarkable. we do not know the common ancestor of any natural group. however. which themselves can only doubtfully be ranked as species. what cases are reversions to formerly existing characters. also. As. we could not have told. on our theory.

who examined this breed for the Indian Government. and a small dun Welsh pony has been carefully described to me. and of ALL colours. as I hear from Colonel Poole. from one between brown and black to a close approach to cream colour. In all parts of the world these stripes occur far oftenest in duns and mouse-duns. though it properly has none. and I have been informed by Colonel Poole that foals of this species are generally striped on the legs and faintly on the shoulder. the legs are generally barred. was marked on its hinder quarters and on its forehead with numerous very narrow. I have also reason to suspect. I have collected cases in England of the spinal stripe in horses of the most distinct breeds. mouse-duns. when a week old. 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. and its legs were feebly striped. The koulan of Pallas is said to have been seen with a double shoulder-stripe. Blyth has seen a specimen of the hemionus with a distinct shoulder-stripe.lost. and I have seen a trace in a bay horse. dark. I have myself seen a dun Devonshire pony. and in one instance in a chestnut. and from Norway in the north to the Malay Archipelago in the south. that with the English race-horse the spinal stripe is much commoner in the foal than in the full-grown animal. which is sometimes double and sometimes treble. In the northwest part of India the Kattywar breed of horses is so generally striped. both with THREE parallel stripes on each shoulder. and the shoulder-stripe. The stripes are often plainest in the foal. the side of the face. 160 . The spine is always striped. from information given me by Mr. Edwards. 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. that. W. 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. All the stripes soon disappeared completely. by the term dun a large range of colour is included. With respect to the horse. though so plainly barred like a zebra over the body. is without bars on the legs. Mr. Gray has figured one specimen with very distinct zebralike bars on the hocks. but Dr. a horse without stripes is not considered as purely bred. The quagga. is sometimes striped.W. Colonel Poole has seen both gray and bay Kattywar horses striped when first foaled. and sometimes quite disappear in old horses. a faint shoulder-stripe may sometimes be seen in duns. zebra-like bars. is common. This foal. moreover. in dark-coloured asses. transverse bars on the legs are not rare in duns.

like those on the dun Devonshire and Welsh ponies. 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. a hybrid has been figured by Dr. which I have seen. In Lord Morton's famous hybrid. I was so convinced that not even a stripe of colour appears from what is commonly called chance. of hybrids between the ass and zebra. one of which. by simple variation. inhabiting the most distant parts of the world.I am aware that Colonel Hamilton Smith. was striped. the hybrid and even the pure offspring subsequently produced from the same mare by a black Arabian sire. I once saw a mule with its legs so much striped that any one might have thought that it was a hybrid zebra. Gosse. and this hybrid. Welsh ponies. Martin. What now are we to say to these several facts? We see several distinct species of the horse genus becoming. etc. and had three short shoulder-stripes. has given a figure of a similar mule. But this view may be safely rejected. or striped on the shoulders like an ass.C. and this is another most remarkable case. In four coloured drawings. and in one of them there was a double shoulder-stripe. for it is highly improbable that the heavy Belgian cart-horse. Norwegian cobs. were much more plainly barred across the legs than is even the pure quagga. 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. according to Mr. who has written on this subject. Lastly. the legs were much more plainly barred than the rest of the body. in certain parts of the United States. The appearance of the stripes is not accompanied by any change 161 . believes that the several breeds of the horse are descended from several aboriginal species. nevertheless had all four legs barred.. and even had some zebra-like stripes on the sides of its face. W. the dun. striped on the legs like a zebra. Rollin asserts that the common mule from the ass and horse is particularly apt to have bars on its legs. Gray (and he informs me that he knows of a second case) from the ass and the hemionus. in his excellent treatise on the horse. from a chestnut mare and male quagga. Now let us turn to the effects of crossing the several species of the horse genus. as we have seen. and that the above-described appearances are all due to ancient crosses with the dun stock. and was. about nine out of ten mules have striped legs. the lanky Kattywar race. should have all have been crossed with one supposed aboriginal stock. and Mr. answered in the affirmative. With respect to this last fact. though the ass only occasionally has stripes on his legs and the hemionus has none and has not even a shoulder-stripe.

some of which have bred true for centuries. or by any other new character. or at least for an unknown cause. is—that there is a TENDENCY in the young of each successive generation to produce the long-lost character. and I see an animal striped like a zebra. when crossed with species inhabiting distant quarters of the world. When the oldest and truest breeds of various colours are crossed. so as often to become striped like the other species of the genus. I have stated that the most probable hypothesis to account for the reappearance of very ancient characters. 162 . to reject a real for an unreal. but had been created in stone so as to mock the shells now living on the sea-shore. the hemionus. in this particular manner. assert that each species has been created with a tendency to vary. quagga. 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. these bars and other marks invariably reappear. and zebra. but perhaps otherwise very differently constructed. He who believes that each equine species was independently created. species. and how exactly parallel is the case with that of the species of the horse genus! For myself. but other species of the genus. To admit this view is. that fossil shells had never lived.of form. as it seems to me. and that each has been created with a strong tendency. We see this tendency to become striped most strongly displayed in hybrids from between several of the most distinct species. to produce hybrids resembling in their stripes. I venture confidently to look back thousands on thousands of generations. we see a strong tendency for the blue tint and bars and marks to reappear in the mongrels. and when any breed assumes by simple variation a bluish tint. from unknown causes. Call the breeds of pigeons. not their own parents. Now observe the case of the several breeds of pigeons: they are descended from a pigeon (including two or three subspecies or geographical races) of a bluish colour. but without any other change of form or character. It makes the works of God a mere mockery and deception. I would almost as soon believe with the old and ignorant cosmogonists. sometimes prevails. both under nature and under domestication. with certain bars and other marks. I presume. will. and that this tendency. the common parent of our domestic horse (whether or not it be descended from one or more wild stocks) of the ass.

Modifications in hard parts and in external parts sometimes affect softer and internal parts.11. 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. from being useless. When one part is largely developed. Changed conditions generally induce mere fluctuating variability. but sometimes they cause direct and definite effects. Specific characters—that is. are not regulated by natural selection. the same laws appear to have acted in producing the lesser differences between varieties of the same species. that organic beings low in the scale are more variable than those standing higher in the scale. the nature of which we are unable to understand. where there has been much former variation and differentiation. But whenever we have the means of instituting a comparison. most 163 . and which have their whole organisation more specialised. or where the manufactory of new specific forms has been actively at work—in that district and among these species. Summary Our ignorance of the laws of variation is profound. and disuse in weakening and diminishing organs. and these may become strongly marked in the course of time. or those which have long been inherited. Changes of structure at an early age may affect parts subsequently developed. perhaps arising from such parts not having been closely specialised for any particular function. for in a district where many species of a genus are found—that is. and every part of the structure which can be saved without detriment will be saved. on an average. and have not differed within this same period. but we have also seen in the second chapter that the same principle applies to the whole individual. and homologous parts tend to cohere. and hence are variable. though we have not sufficient evidence on this head. and many cases of correlated variation. we now find. and the greater differences between species of the same genus. In these remarks we have referred to special parts or organs being still variable. so that their modifications have not been closely checked by natural selection. because they have recently varied and thus come to differ. It follows probably from this same cause. Not in one case out of a hundred can we pretend to assign any reason why this or that part has varied. Rudimentary organs. Homologous parts tend to vary in the same manner. perhaps it tends to draw nourishment from the adjoining parts. undoubtedly occur. Habit in producing constitutional peculiarities. Multiple parts are variable in number and in structure. appear in many cases to have been potent in their effects. and use in strengthening.

164 . and exposed to similar influences. naturally tend to present analogous variations. Any part or organ developed to an extraordinary size or in an extraordinary manner. and such characters differ much in the species of the same group. such modifications will add to the beautiful and harmonious diversity of nature. in comparison with the same part or organ in the allied species. natural selection has succeeded in giving a fixed character to the organ. in however extraordinary a manner it may have been developed. for variation is a longcontinued and slow process.varieties. and thus we can understand why it should often still be variable in a much higher degree than other parts. 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. 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. must have gone through an extraordinary amount of modification since the genus arose. Species inheriting nearly the same constitution from a common parent. and specific differences to the several species of the same genus. 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. Although new and important modifications may not arise from reversion and analogous variation. Whatever the cause may be of each slight difference between the 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. Secondary sexual characters are highly variable. or these same species may occasionally revert to some of the characters of their ancient progenitors.

Chapter 6 Difficulties Of The Theory 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. 165 .

Long before the reader has arrived at this part of my work. why. an organ so wonderful as the eye? Thirdly. 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. how can we account for species. as we see them. and those that are real are not. on the other hand. can instincts be acquired and modified through natural selection? What shall we say to the instinct which leads the bee to make cells. being sterile and producing sterile offspring. if species have descended from other species by fine gradations. These difficulties and objections may be classed under the following heads: First. a crowd of difficulties will have occurred to him. well defined? Secondly. Some of them are so serious that to this day I can hardly reflect on them without being in some degree staggered. I think.1. on the one hand. but. instead of the species being. which serves as a flyflapper. and which has practically anticipated the discoveries of profound mathematicians? Fourthly. some miscellaneous objections in the following chapter. 166 . when crossed. for instance. whereas. do we not everywhere see innumerable transitional forms? Why is not all nature in confusion. such as the tail of a giraffe. the structure and habits of a bat. fatal to the theory. an organ of trifling importance. their fertility is unimpaired? The two first heads will be here discussed. the greater number are only apparent. is it possible that an animal having. to the best of my judgment. and. when varieties are crossed. Instinct and Hybridism in the two succeeding chapters.

and during the process of modification. we generally meet at successive intervals with closely allied or representative species. but the natural collections have been imperfectly made. and as the one becomes rarer and rarer. But in the intermediate region. Hence. if we look at each species as descended from some unknown form. But I think it can be in large part explained. each has become adapted to the conditions of life of its own region. On The Absence Or Rarity Of Transitional Varieties As natural selection acts solely by the preservation of profitable modifications. Thus extinction and natural selection go hand in hand. till the one replaces the other. and finally to exterminate. and I will here only state that I believe the answer mainly lies in the record being incomparably less perfect than is generally supposed. Let us take a simple case: in travelling from north to south over a continent. having intermediate conditions of life. 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. Hence we ought not to expect at the present time to meet with numerous transitional varieties in each region. 167 . But it may be urged that when several closely allied species inhabit the same territory. The crust of the earth is a vast museum. as by this theory innumerable transitional forms must have existed. But if we compare these species where they intermingle. why do we not now find closely-linking intermediate varieties? This difficulty for a long time quite confounded me. they are generally as absolutely distinct from each other in every detail of structure as are specimens taken from the metropolis inhabited by each. By my theory these allied species are descended from a common parent. These representative species often meet and interlock. its own less improved parent-form and other less-favoured forms with which it comes into competition. we surely ought to find at the present time many transitional forms.2. though they must have existed there. the other becomes more and more frequent. each new form will tend in a fully-stocked country to take the place of. and only at long intervals of time. But. evidently filling nearly the same place in the natural economy of the land. 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. and may be embedded there in a fossil condition. and has supplanted and exterminated its original parentform and all the transitional varieties between its past and present states.

that nearly all either prey on or serve as prey for others. then becoming somewhat abruptly rarer and rarer on the confines. the range of any one species. in short. not blending one into another by insensible gradations. will. and in such islands distinct species might have been separately formed without the possibility of intermediate varieties existing in the intermediate zones. these facts ought to cause surprise. though I do not doubt that the formerly broken condition of areas now continuous. were it not for other competing species. that it has been continuous during a long period. be extremely liable 168 . depending as it does on the range of others. But when we bear in mind that almost every species. as climate and height or depth graduate away insensibly. would increase immensely in numbers. Forbes in sounding the depths of the sea with the dredge. or by which it is destroyed. Hence the neutral territory between two representative species is generally narrow in comparison with the territory proper to each. and finally disappearing. In looking at species as they are now distributed over a wide area. Moreover. where it exists in lessened numbers. and sometimes it is quite remarkable how abruptly. We see the same fact in ascending mountains. By changes in the form of the land and of climate. 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. but in large part on the presence of other species. De Candolle has observed. or in the nature of the seasons. even in its metropolis.In the first place we should be extremely cautious in inferring. on which it lives. we generally find them tolerably numerous over a large territory. But I will pass over this way of escaping from the difficulty. more especially with freely-crossing and wandering animals. marine areas now continuous must often have existed within recent times in a far less continuous and uniform condition than at present. Geology would lead us to believe that most continents have been broken up into islands even during the later tertiary periods. for I believe that many perfectly defined species have been formed on strictly continuous areas. each species on the confines of its range. during fluctuations in the number of its enemies or of its prey. as Alph. The same fact has been noticed by E. has played an important part in the formation of new species. and as these species are already defined objects. 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. because an area is now continuous. will tend to be sharply defined. a common alpine species disappears.

as already remarked. from inhabiting larger areas.to utter extermination. by which two varieties are supposed to be converted and perfected into two distinct species. the more common forms. run a greater chance of being exterminated than one existing in large numbers. Hence. as varieties do not essentially differ from species. then. and thus its geographical range will come to be still more sharply defined. sooner than the forms which they originally linked together. will have a great advantage over the intermediate variety. then we can understand why intermediate varieties should not endure for very long periods: why. For any form existing in lesser numbers would. Watson. will tend to beat and supplant the less common forms. And it would appear from information given me by Mr. Wollaston. they should be exterminated and disappear. As allied or representative species. are generally distributed in such a manner that each has a wide range. they are much rarer numerically than the forms which they connect. 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. as a general rule. than will the rarer forms which exist in lesser numbers. the two which exist in larger numbers. It is the same principle which. for these will be more slowly modified and improved. that during the process of further modification. But it is a far more important consideration. and practically. when varieties intermediate between two other forms occur. within any given period. of presenting further favourable variations for natural selection to seize on. this rule holds good with varieties in a state of nature. Asa Gray. with a comparatively narrow neutral territory between them. if we may trust these facts and inferences. and Mr. consequently. For forms existing in larger numbers will have a better chance. Dr. in which they become rather suddenly rarer and rarer. Now. and if we take a varying species inhabiting a very large area. will exist in lesser numbers from inhabiting a narrow and lesser area. that generally. as far as I can make out. The intermediate variety. as I believe. when inhabiting a continuous area. accounts for the common species in each 169 . I have met with striking instances of the rule in the case of varieties intermediate between well-marked varieties in the genus Balanus. in the race for life. which exists in smaller numbers in a narrow and intermediate zone. and a third variety to a narrow intermediate zone. the same rule will probably apply to both. and conclude that varieties linking two other varieties together generally have existed in lesser numbers than the forms which they connect. we shall have to adapt two varieties to two large areas.

and that the inhabitants are all trying with equal steadiness and skill to improve their stocks by selection. So that. intermediate hill variety. I may illustrate what I mean by supposing three varieties of sheep to be kept. but these links during the process of natural selection will have been supplanted and exterminated. so that they will no longer be found in a living state. intermediate varieties will. or on the occasional immigration of new inhabitants. In this case. and consequently the improved mountain or plain breed will soon take the place of the less improved hill breed. it is probable. may have separately been rendered sufficiently distinct to rank as representative species. one adapted to an extensive mountainous region. on some of the old inhabitants becoming slowly modified. hilly tract. at first have been formed in the intermediate zones. and a third to the wide plains at the base. in a still more important degree. Thirdly. and. with the new forms thus produced and the old ones acting and reacting on each other. 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. more especially among the classes which unite for each birth and wander much. without the interposition of the supplanted. To sum up. and thus the two breeds. because new varieties are very slowly formed. and this assuredly we do see. when two or more varieties have been formed in different portions of a strictly continuous area. probably. in which many forms. as shown in the second chapter. I believe that species come to be tolerably well-defined objects. for variation is a slow process. 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. intermediate varieties between the several representative species and their common parent.country. and do not at any one period present an inextricable chaos of varying and intermediate links: first. hilly tract. in any one region and at any one time. but they 170 . And such new places will depend on slow changes of climate. and natural selection can do nothing until favourable individual differences or variations occur. we ought to see only a few species presenting slight modifications of structure in some degree permanent. presenting on an average a greater number of well-marked varieties than do the rarer species. will come into close contact with each other. must formerly have existed within each isolated portion of the land. areas now continuous must often have existed within the recent period as isolated portions. Secondly. which originally existed in greater numbers. a second to a comparatively narrow.

present more varieties.will generally have had a short duration. linking closely together all the species of the same group. from existing in greater numbers will. 171 . as we shall attempt to show in a future chapter. to exterminate the parent forms and the intermediate links. in an extremely imperfect and intermittent record. exist in the intermediate zones in lesser numbers than the varieties which they tend to connect. they will almost certainly be beaten and supplanted by the forms which they connect. looking not to any one time. and thus be further improved through natural selection and gain further advantages. in the aggregate. numberless intermediate varieties. Lastly. from reasons already assigned (namely from what we know of the actual distribution of closely allied or representative species. but at all time. Consequently evidence of their former existence could be found only among fossil remains. but the very process of natural selection constantly tends. for these. which are preserved. For these intermediate varieties will. and likewise of acknowledged varieties). and during the process of further modification through natural selection. as has been so often remarked. must assuredly have existed. if my theory be true. From this cause alone the intermediate varieties will be liable to accidental extermination.

and from others. to the so-called flying squirrels. and of diversified habits. and flying squirrels have their limbs and even the base of the tail united by a broad expanse of skin. by enabling it to escape birds or beasts of prey. at least. in the same species. like other polecats on mice and land animals. let other competing rodents or new beasts of prey immigrate. I lie under a heavy disadvantage. with the posterior part of their bodies rather wide and with the skin on their flanks rather full. as there is reason to believe. I can give only one or two instances of transitional habits and structures in allied species. either constant or occasional. for. Let the climate and vegetation change. which has webbed feet. Richardson has remarked. and all analogy would lead us to believe that some. 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. On The Origin And Transition Of Organic Beings With Peculiar Habits And Structure It has been asked by the opponents of such views as I hold. and preys.3. could a land carnivorous animal have been converted into one with aquatic habits. or. Look at the Mustela vison of North America. here we have the finest gradation from animals with their tails only slightly flattened. which serves as a parachute and allows them to glide through the air to an astonishing distance from tree to tree. short legs. and it had been asked how an insectivorous quadruped could possibly have been converted into a flying bat. the question would have been far more difficult to answer. or to collect food more quickly. to lessen the danger from occasional falls. out of the many striking cases which I have collected. and form of tail. We cannot doubt that each structure is of use to each kind of squirrel in its own country. but during the long winter it leaves the frozen waters. 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. or old ones become modified. it is clear that each must be well adapted to its place in nature. during summer this animal dives for and preys on fish. Here. and as each exists by a struggle for life. 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. and which resembles an otter in its fur. as Sir J. Look at the family of squirrels. Yet I think such difficulties have little weight. as on other occasions. for instance. how. If a different case had been taken. of the squirrels would decrease in numbers or become 172 .

in the continued preservation of individuals with fuller and fuller flank-membranes. as sails. which perhaps may all be the result of disuse. and seeing that 173 . each grade of structure having been useful to its possessor. If about a dozen genera of birds were to become extinct. yet there is no difficulty in supposing that such links formerly existed. and this. Although no graduated links of structure. each being propagated. would have converted the animal into a bat. This flank. as far as the organs of flight are concerned. for each has to live by a struggle: but it is not necessarily the best possible under all possible conditions. Now look at the Galeopithecus or so-called flying lemur. In certain bats in which the wing-membrane extends from the top of the shoulder to the tail and includes the hind-legs. more especially under changing conditions of life. which was formerly ranked among bats. fitted for gliding through the air. we perhaps see traces of an apparatus originally fitted for gliding through the air rather than for flight. Seeing that a few members of such water-breathing classes as the Crustacea and Mollusca are adapted to live on the land. a perfect so-called flying squirrel was produced. unless they also become modified and improved in structure in a corresponding manner. under the conditions of life to which it is exposed. each modification being useful. by the accumulated effects of this process of natural selection.membrane is furnished with an extensor muscle. indicate the steps by which birds actually acquired their perfect power of flight. like the logger headed duck (Micropterus of Eyton). but is now believed to belong to the Insectivora. who would have ventured to surmise that birds might have existed which used their wings solely as flappers. and that each was developed in the same manner as with the less perfectly gliding squirrels. and functionally for no purpose. now connect the Galeopithecus with the other Insectivora. An extremely wide flank-membrane stretches from the corners of the jaw to the tail. but they serve to show what diversified means of transition are at least possible. until. and includes the limbs with the elongated fingers. as fins in the water and as front legs on the land. It must not be inferred from these remarks that any of the grades of wingstructure here alluded to. 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. like the penguin. I can see no difficulty.exterminated. like the ostrich. Therefore. like the apteryx? Yet the structure of each of these birds is good for it.

whether habits generally change first and structure afterwards. In our own country the larger titmouse (Parus major) 174 . difficult to decide and immaterial for us. I will now give two or three instances. than in the case of species with fully developed structures.we have flying birds and mammals. Hence the chance of discovering species with transitional grades of structure in a fossil condition will always be less. In either case it would be easy for natural selection to adapt the structure of the animal to its changed habits. Of cases of changed habits it will suffice merely to allude to that of the many British insects which now feed on exotic plants. who would have ever imagined that in an early transitional state they had been inhabitants of the open ocean. to escape being devoured by other fish? When we see any structure highly perfected for any particular habit. until their organs of flight had come to a high stage of perfection. It is. and had used their incipient organs of flight exclusively. like a kestrel. and formerly had flying reptiles. on the land and in the water. it is conceivable that flying-fish. or exclusively on artificial substances. to return to our imaginary illustration of the flying-fish. If this had been effected. which were gradually rendered more perfect through natural selection. both probably often occurring almost simultaneously. for taking prey of many kinds in many ways. 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. Of diversified habits innumerable instances could be given: I have often watched a tyrant flycatcher (Saurophagus sulphuratus) in South America. however. so far as we know. might have been modified into perfectly winged animals. slightly rising and turning by the aid of their fluttering fins. we should bear in mind that animals displaying early transitional grades of the structure will seldom have survived to the present day. as the wings of a bird for flight. or whether slight modifications of structure lead to changed habits. both of diversified and of changed habits. Furthermore. or exclusively to one of its several habits. it does not seem probable that fishes capable of true flight would have been developed under many subordinate forms. which now glide far through the air. so as to have given them a decided advantage over other animals in the battle for life. flying insects of the most diversified types. and then dashing into it like a kingfisher at a fish. and at other times standing stationary on the margin of water. for they will have been supplanted by their successors. from their having existed in lesser numbers. Thus. hovering over one spot and then proceeding to another. in the individuals of the same species.

and undulatory flight. almost like a whale. like a shrike. and it makes its nest in holes in banks! In certain other districts. is a woodpecker. in all the essential parts of its structure. this same woodpecker. Hudson states. having anomalous habits. almost like a creeper. not only from my own observations. kills small birds by blows on the head. but. but with many parts of its organisation profoundly modified in relation to its new habits of life. Even in such trifling characters as the colouring. the harsh tone of the voice. and a straight. and I have many times seen and heard it hammering the seeds of the yew on a branch. the Puffinuria berardi. would be mistaken by any one for an auk or a grebe. however. pointed tail-feathers. and thus breaking them like a nuthatch. however. in certain large districts it does not climb trees. whereas the woodpecker of La Plata has had its structure only slightly 175 . sufficiently stiff to support the bird in a vertical position on a post. as Mr. as I can assert. Petrels are the most aerial and oceanic of birds. we might expect that such individuals would occasionally give rise to new species. yet. insects in the water. And such instances occur in nature. On the plains of La Plata. in its general habits. I may mention as another illustration of the varied habits of this genus. and with their structure either slightly or considerably modified from that of their type. 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. nevertheless. The beak. strong beak. thus catching. and bores holes in the trunk for its nest. but from those of the accurate Azara. it sometimes. is not so straight or so strong as in the typical woodpeckers but it is strong enough to bore into wood. in its astonishing power of diving. it is essentially a petrel. a long. there is a woodpecker (Colaptes campestris) which has two toes before and two behind.pointed tongue. but not so stiff as in the typical wood-peckers.may be seen climbing branches. where hardly a tree grows. its close blood-relationship to our common woodpecker is plainly declared. 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. in its manner of swimming and of flying when made to take flight. In North America the black bear was seen by Hearne swimming for hours with widely open mouth. and others with elongated wings which chase insects on the wing. As we sometimes see individuals following habits different from those proper to their species and to the other species of the same genus. in the quiet sounds of Tierra del Fuego. frequents trees. Hence this Colaptes.

habits have changed without a corresponding change of structure. In such cases. however different that may be from its own place. He who believes in the struggle for existence and in the principle of natural selection. it will seize on the place of that inhabitant. excepting the genus Proctotrupes. though not in structure.—using its wings under water and grasping stones with its feet. and thus gains an advantage over some other inhabitant of the same country. would never have suspected its sub-aquatic habits. 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. All the members of the great order of Hymenopterous insects are terrestrial. and no one except Audubon. it often enters the water and dives about by the use not of its legs but of its wings. grebes and coots are eminently aquatic. either in habits or structure. will acknowledge that every organic being is constantly endeavouring to increase in numbers. of the Grallatores. yet it exhibits no modification in structure in accordance with its abnormal habits. the acutest observer. the deeply scooped membrane between the toes shows that structure has begun to change. and that if any one being varies ever so little. living on the dry land and rarely alighting on the water. but this seems to me only restating the fact in dignified language. and the second is nearly as terrestrial as the quail or partridge. In the case of the water-ouzel. What seems plainer than that the long toes. yet this bird. must occasionally have felt surprise when he has met with an animal having habits and structure not in agreement. has seen the frigate-bird. are formed for walking over swamps and floating plants. 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. and remains as long as four hours beneath the surface. In the frigate-bird. On the other hand. He who believes in separate and innumerable acts of creation may say. and many others could be given. 176 . which is allied to the thrush family. yet the first is nearly as aquatic as the coot.modified. subsists by diving. He who believes that each being has been created as we now see it. not furnished with membrane. The webbed feet of the upland goose may be said to have become almost rudimentary in function. Hence it will cause him no surprise that there should be geese and frigate-birds with webbed feet. which Sir John Lubbock has discovered to be aquatic in its habits. alight on the surface of the ocean. although their toes are only bordered by membrane. by examining its dead body. The water-hen and landrail are members of this order. which has all its four toes webbed.

living in meadows instead of in swamps. that there should be woodpeckers where hardly a tree grows. that there should be diving thrushes and diving Hymenoptera.that there should be long-toed corncrakes. and petrels with the habits of auks. 177 .

could have been formed by natural selection. When it was first said that the sun stood still and the world turned round. We may. as is likewise certainly the case. however. that is to the collateral descendants from the same parent-form. cannot be trusted in science. for admitting different amounts of light. as every philosopher knows. and we are forced to look to other species and genera of the same group. Reason tells me. and resting merely on sarcodic tissue. though insuperable by our imagination. and for the chance of some gradations having been transmitted in an unaltered or little altered condition. the eye ever varies and the variations be inherited. but this is scarcely ever possible. as is certainly the case. surrounded by pigment-cells and covered by translucent skin. are capable of perceiving light. each grade being useful to its possessor. and for the correction of spherical and chromatic aberration. in order to see what gradations are possible. should not be considered as subversive of the theory. the common sense of mankind declared the doctrine false. but without any lens or other refractive body. absurd in the highest degree. without any nerves. vox Dei. as some of the lowest organisms in which nerves cannot be detected. and serve only to distinguish 178 . if further.4. we ought to look exclusively to its lineal progenitors. according to M. that if numerous gradations from a simple and imperfect eye to one complex and perfect can be shown to exist. The simplest organ which can be called an eye consists of an optic nerve. seems. but the old saying of Vox populi. and if such variations should be useful to any animal under changing conditions of life. Jourdain. but I may remark that. it does not seem impossible that certain sensitive elements in their sarcode should become aggregated and developed into nerves. descend even a step lower and find aggregates of pigment-cells. then the difficulty of believing that a perfect and complex eye could be formed by natural selection. Eyes of the above simple nature are not capable of distinct vision. apparently serving as organs of vision. I freely confess. How a nerve comes to be sensitive to light. But the state of the same organ in distinct classes may incidentally throw light on the steps by which it has been perfected. In searching for the gradations through which an organ in any species has been perfected. endowed with this special sensibility. hardly concerns us more than how life itself originated. Organs Of Extreme Perfection And Complication To suppose that the eye with all its inimitable contrivances for adjusting the focus to different distances.

if they were extremely slight 179 . and that the cones include curiously modified nervous filaments. at the right distance from the concentrating apparatus. and an image will be formed on it. He who will go thus far. here given much too briefly. In this concentration of the rays we gain the first and by far the most important step towards the formation of a true. and graduated range of structure in the eyes of the lower animals. With insects it is now known that the numerous facets on the cornea of their great compound eyes form true lenses. besides a fourth main class of aggregated simple eyes. although in this case he does not know the transitional states. picture-forming eye. can be explained by the theory of modification through natural selection. and in some near the surface. which. projecting with a convex surface. we may start from an optic nerve simply coated with pigment. small depressions in the layer of pigment which surrounds the nerve are filled. like the cornea in the higher animals. for we have only to place the naked extremity of the optic nerve. as described by the author just quoted. the latter sometimes forming a sort of pupil. But these organs in the Articulata are so much diversified that Muller formerly made three main classes with seven subdivisions. When we reflect on these facts. In certain star-fishes. ought not to hesitate to go one step further. he ought to admit that a structure even as perfect as an eagle's eye might thus be formed. with transparent gelatinous matter. could not be done through natural selection. It has been objected that in order to modify the eye and still preserve it as a perfect instrument. it is not necessary to suppose that the modifications were all simultaneous. but destitute of lens or other optical contrivance. many changes would have to be effected simultaneously. it is assumed. otherwise inexplicable. if he finds on finishing this volume that large bodies of facts. but only to concentrate the luminous rays and render their perception more easy. the difficulty ceases to be very great in believing that natural selection may have converted the simple apparatus of an optic nerve. and when we bear in mind how small the number of all living forms must be in comparison with those which have become extinct. but as I have attempted to show in my work on the variation of domestic animals.light from darkness. He suggests that this serves not to form an image. with respect to the wide. into an optical instrument as perfect as is possessed by any member of the Articulata class. coated with pigment and invested by transparent membrane. which in some of the lower animals lies deeply buried in the body. diversified. In the great class of the Articulata.

and gradual. Different kinds of modification would, also, serve for the same general purpose: as Mr. Wallace has remarked, "If a lens has too short or too long a focus, it may be amended either by an alteration of curvature, or an alteration of density; if the curvature be irregular, and the rays do not converge to a point, then any increased regularity of curvature will be an improvement. So the contraction of the iris and the muscular movements of the eye are neither of them essential to vision, but only improvements which might have been added and perfected at any stage of the construction of the instrument." Within the highest division of the animal kingdom, namely, the Vertebrata, we can start from an eye so simple, that it consists, as in the lancelet, of a little sack of transparent skin, furnished with a nerve and lined with pigment, but destitute of any other apparatus. In fishes and reptiles, as Owen has remarked, "The range of gradation of dioptric structures is very great." It is a significant fact that even in man, according to the high authority of Virchow, the beautiful crystalline lens is formed in the embryo by an accumulation of epidermic cells, lying in a sack-like fold of the skin; and the vitreous body is formed from embryonic subcutaneous tissue. To arrive, however, at a just conclusion regarding the formation of the eye, with all its marvellous yet not absolutely perfect characters, it is indispensable that the reason should conquer the imagination; 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. It is scarcely possible to avoid comparing the eye with a telescope. We know that this instrument has been perfected by the long-continued efforts of the highest human intellects; and we naturally infer that the eye has been formed by a somewhat analogous process. 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, we ought in imagination to take a thick layer of transparent tissue, with spaces filled with fluid, and with a nerve sensitive to light beneath, 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, placed at different distances from each other, and with the surfaces of each layer slowly changing in form. Further we must suppose that there is a power, represented by natural selection or the survival of the fittest, always intently watching each slight alteration in the transparent layers; and carefully preserving each which, under varied circumstances, in any way or degree, tends to produce a distincter image. We must suppose each new state of the

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instrument to be multiplied by the million; each to be preserved until a better is produced, and then the old ones to be all destroyed. In living bodies, variation will cause the slight alteration, generation will multiply them almost infinitely, and natural selection will pick out with unerring skill each improvement. Let this process go on for millions of years; and during each year on millions of individuals of many kinds; and may we not believe that a living optical instrument might thus be formed as superior to one of glass, as the works of the Creator are to those of man?

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5. Modes Of Transition
If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case. No doubt many organs exist of which we do not know the transitional grades, more especially if we look to much-isolated species, around which, according to the theory, there has been much extinction. Or again, if we take an organ common to all the members of a class, for in this latter case the organ must have been originally formed at a remote period, since which all the many members of the class have been developed; and in order to discover the early transitional grades through which the organ has passed, we should have to look to very ancient ancestral forms, long since become extinct. We should be extremely cautious in concluding that an organ could not have been formed by transitional gradations of some kind. Numerous cases could be given among the lower animals of the same organ performing at the same time wholly distinct functions; thus in the larva of the dragon-fly and in the fish Cobites the alimentary canal respires, digests, and excretes. In the Hydra, the animal may be turned inside out, and the exterior surface will then digest and the stomach respire. In such cases natural selection might specialise, if any advantage were thus gained, the whole or part of an organ, which had previously performed two functions, for one function alone, and thus by insensible steps greatly change its nature. Many plants are known which regularly produce at the same time differently constructed flowers; and if such plants were to produce one kind alone, a great change would be effected with comparative suddenness in the character of the species. It is, however, probable that the two sorts of flowers borne by the same plant were originally differentiated by finely graduated steps, which may still be followed in some few cases. Again, two distinct organs, or the same organ under two very different forms, may simultaneously perform in the same individual the same function, 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, at the same time that they breathe free air in their swim-bladders, this latter organ being divided by highly vascular partitions and having a ductus pneumaticus for the supply of air. To give another instance from the vegetable kingdom: plants climb by three distinct means, by spirally twining, by clasping a support with their sensitive

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tendrils, and by the emission of aerial rootlets; these three means are usually found in distinct groups, but some few species exhibit two of the means, or even all three, combined in the same individual. In all such cases one of the two organs might readily be modified and perfected so as to perform all the work, being aided during the progress of modification by the other organ; and then this other organ might be modified for some other and quite distinct purpose, or be wholly obliterated. The illustration of the swim-bladder in fishes is a good one, because it shows us clearly the highly important fact that an organ originally constructed for one purpose, namely flotation, may be converted into one for a widely different purpose, namely respiration. The swim-bladder has, also, been worked in as an accessory to the auditory organs of certain fishes. All physiologists admit that the swim-bladder is homologous, or "ideally similar" in position and structure with the lungs of the higher vertebrate animals: hence there is no reason to doubt that the swimbladder has actually been converted into lungs, or an organ used exclusively for respiration. 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. We can thus, as I infer from Professor Owen's interesting description of these parts, understand the strange fact that every particle of food and drink which we swallow has to pass over the orifice of the trachea, with some risk of falling into the lungs, notwithstanding the beautiful contrivance by which the glottis is closed. 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. 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, Landois has shown that the wings of insects are developed from the trachea; it is therefore highly probable that in this great class organs which once served for respiration have been actually converted into organs for flight. In considering transitions of organs, it is so important to bear in mind the probability of conversion from one function to another, that I will give another instance. Pedunculated cirripedes have two minute folds of skin, called by me the ovigerous frena, which serve, through the means of a sticky secretion, to retain the eggs until they are hatched within the sack. These cirripedes have no branchiae, the whole surface of the body and of the sack, together with the small frena, serving for respiration.

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The Balanidae or sessile cirripedes, on the other hand, have no ovigerous frena, the eggs lying loose at the bottom of the sack, within the well-enclosed shell; but they have, in the same relative position with the frena, large, much-folded membranes, which freely communicate with the circulatory lacunae of the sack and body, and which have been considered by all naturalists to act as branchiae. 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; indeed, they graduate into each other. Therefore it need not be doubted that the two little folds of skin, which originally served as ovigerous frena, but which, likewise, very slightly aided in the act of respiration, have been gradually converted by natural selection into branchiae, simply through an increase in their size and the obliteration of their adhesive glands. If all pedunculated cirripedes had become extinct, and they have suffered far more extinction than have sessile cirripedes, 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, namely, through the acceleration or retardation of the period of reproduction. 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, before they have acquired their perfect characters; and if this power became thoroughly well developed in a species, it seems probable that the adult stage of development would sooner or later be lost; and in this case, especially if the larva differed much from the mature form, the character of the species would be greatly changed and degraded. Again, not a few animals, after arriving at maturity, go on changing in character during nearly their whole lives. With mammals, for instance, the form of the skull is often much altered with age, of which Dr. Murie has given some striking instances with seals. Every one knows how the horns of stags become more and more branched, and the plumes of some birds become more finely developed, as they grow older. Professor Cope states that the teeth of certain lizards change much in shape with advancing years. With crustaceans not only many trivial, but some important parts assume a new character, as recorded by Fritz Muller, after maturity. In all such cases- -and many could be given—if the age for reproduction were retarded, the character of the species, at least in its adult state, would be modified; nor is it improbable that the previous and earlier stages of development would in some cases be hurried through and finally lost. Whether species have often or ever been modified through this

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comparatively sudden mode of transition, I can form no opinion; but if this has occurred, it is probable that the differences between the young and the mature, and between the mature and the old, were primordially acquired by graduated steps.

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6. Special Difficulties Of The Theory Of Natural Selection
Although we must be extremely cautious in concluding that any organ could not have been produced by successive, small, transitional gradations, yet undoubtedly serious cases of difficulty occur. One of the most serious is that of neuter insects, which are often differently constructed from either the males or fertile females; but this case will be treated of in the next chapter. The electric organs of fishes offer another case of special difficulty; for it is impossible to conceive by what steps these wondrous organs have been produced. But this is not surprising, for we do not even know of what use they are. In the gymnotus and torpedo they no doubt serve as powerful means of defence, and perhaps for securing prey; yet in the ray, as observed by Matteucci, an analogous organ in the tail manifests but little electricity, even when the animal is greatly irritated; so little that it can hardly be of any use for the above purposes. Moreover, in the ray, besides the organ just referred to, there is, as Dr. R. McDonnell has shown, another organ near the head, not known to be electrical, but which appears to be the real homologue of the electric battery in the torpedo. It is generally admitted that there exists between these organs and ordinary muscle a close analogy, in intimate structure, in the distribution of the nerves, and in the manner in which they are acted on by various reagents. It should, also, be especially observed that muscular contraction is accompanied by an electrical discharge; and, as Dr. Radcliffe insists, "in the electrical apparatus of the torpedo during rest, there would seem to be a charge in every respect like that which is met with in muscle and nerve during the rest, and the discharge of the torpedo, instead of being peculiar, may be only another form of the discharge which attends upon the action of muscle and motor nerve." Beyond this we cannot at present go in the way of explanation; but as we know so little about the uses of these organs, and as we know nothing about the habits and structure of the progenitors of the existing electric fishes, it would be extremely bold to maintain that no serviceable transitions are possible by which these organs might have been gradually developed. These organs appear at first to offer another and far more serious difficulty; for they occur in about a dozen kinds of fish, of which several are widely remote in their affinities. When the same organ is found in several members of the same class, especially if in members having very different habits of life, we may generally attribute its presence to inheritance from a common ancestor; and its absence in some of the members to

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loss through disuse or natural selection. So that, if the electric organs had been inherited from some one ancient progenitor, we might have expected that all electric fishes would have been specially related to each other; but this is far from the case. Nor does geology at all lead to the belief that most fishes formerly possessed electric organs, which their modified descendants have now lost. But when we look at the subject more closely, we find in the several fishes provided with electric organs, that these are situated in different parts of the body, that they differ in construction, as in the arrangement of the plates, and, according to Pacini, in the process or means by which the electricity is excited—and lastly, in being supplied with nerves proceeding from different sources, and this is perhaps the most important of all the differences. Hence in the several fishes furnished with electric organs, these cannot be considered as homologous, but only as analogous in function. Consequently there is no reason to suppose that they have been inherited from a common progenitor; for had this been the case they would have closely resembled each other in all respects. Thus the difficulty of an organ, apparently the same, arising in several remotely allied species, disappears, leaving only the lesser yet still great difficulty: namely, by what graduated steps these organs have been developed in each separate group of fishes. The luminous organs which occur in a few insects, belonging to widely different families, and which are situated in different parts of the body, offer, under our present state of ignorance, a difficulty almost exactly parallel with that of the electric organs. Other similar cases could be given; for instance in plants, the very curious contrivance of a mass of pollen-grains, borne on a foot-stalk with an adhesive gland, is apparently the same in Orchis and Asclepias, genera almost as remote as is possible among flowering plants; but here again the parts are not homologous. In all cases of beings, far removed from each other in the scale of organisation, which are furnished with similar and peculiar organs, it will be found that although the general appearance and function of the organs may be the same, yet fundamental differences between them can always be detected. For instance, the eyes of Cephalopods or cuttle-fish and of vertebrate animals appear wonderfully alike; and in such widely sundered groups no part of this resemblance can be due to inheritance from a common progenitor. Mr. Mivart has advanced this case as one of special difficulty, but I am unable to see the force of his argument. An organ for vision must be formed of transparent tissue, and must include some sort of lens for throwing an image at the back of a darkened chamber. Beyond this superficial resemblance, there is hardly any real

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similarity between the eyes of cuttle-fish and vertebrates, as may be seen by consulting Hensen's admirable memoir on these organs in the Cephalopoda. It is impossible for me here to enter on details, but I may specify a few of the points of difference. The crystalline lens in the higher cuttlefish consists of two parts, placed one behind the other like two lenses, both having a very different structure and disposition to what occurs in the vertebrata. The retina is wholly different, with an actual inversion of the elemental parts, and with a large nervous ganglion included within the membranes of the eye. The relations of the muscles are as different as it is possible to conceive, and so in other points. Hence it is not a little difficult to decide how far even the same terms ought to be employed in describing the eyes of the Cephalopoda and Vertebrata. It is, of course, open to any one to deny that the eye in either case could have been developed through the natural selection of successive slight variations; but if this be admitted in the one case it is clearly possible in the other; and fundamental differences of structure in the visual organs of two groups might have been anticipated, in accordance with this view of their manner of formation. As two men have sometimes independently hit on the same invention, so in the several foregoing cases it appears that natural selection, working for the good of each being, and taking advantage of all favourable variations, has produced similar organs, as far as function is concerned, in distinct organic beings, which owe none of their structure in common to inheritance from a common progenitor. Fritz Muller, in order to test the conclusions arrived at in this volume, has followed out with much care a nearly similar line of argument. Several families of crustaceans include a few species, possessing an airbreathing apparatus and fitted to live out of the water. In two of these families, which were more especially examined by Muller, and which are nearly related to each other, the species agree most closely in all important characters: namely in their sense organs, circulating systems, in the position of the tufts of hair within their complex stomachs, and lastly in the whole structure of the water-breathing branchiae, even to the microscopical hooks by which they are cleansed. Hence it might have been expected that in the few species belonging to both families which live on the land, the equally important air-breathing apparatus would have been the same; for why should this one apparatus, given for the same purpose, have been made to differ, while all the other important organs were closely similar, or rather, identical. Fritz Muller argues that this close similarity in so many points of structure must, in accordance with the views advanced by me, be accounted

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for by inheritance from a common progenitor. But as the vast majority of the species in the above two families, as well as most other crustaceans, are aquatic in their habits, it is improbable in the highest degree that their common progenitor should have been adapted for breathing air. Muller was thus led carefully to examine the apparatus in the air-breathing species; and he found it to differ in each in several important points, as in the position of the orifices, in the manner in which they are opened and closed, and in some accessory details. Now such differences are intelligible, and might even have been expected, on the supposition that species belonging to distinct families had slowly become adapted to live more and more out of water, and to breathe the air. For these species, from belonging to distinct families, would have differed to a certain extent, and in accordance with the principle that the nature of each variation depends on two factors, viz., the nature of the organism and that of the surrounding conditions, their variability assuredly would not have been exactly the same. Consequently natural selection would have had different materials or variations to work on, in order to arrive at the same functional result; and the structures thus acquired would almost necessarily have differed. On the hypothesis of separate acts of creation the whole case remains unintelligible. This line of argument seems to have had great weight in leading Fritz Muller to accept the views maintained by me in this volume. Another distinguished zoologist, the late Professor Claparede, has argued in the same manner, and has arrived at the same result. He shows that there are parasitic mites (Acaridae), belonging to distinct sub-families and families, which are furnished with hair-claspers. These organs must have been independently developed, as they could not have been inherited from a common progenitor; and in the several groups they are formed by the modification of the fore legs, of the hind legs, of the maxillae or lips, and of appendages on the under side of the hind part of the body. In the foregoing cases, we see the same end gained and the same function performed, in beings not at all or only remotely allied, by organs in appearance, though not in development, closely similar. On the other hand, it is a common rule throughout nature that the same end should be gained, even sometimes in the case of closely related beings, by the most diversified means. How differently constructed is the feathered wing of a bird and the membrane-covered wing of a bat; and still more so the four wings of a butterfly, the two wings of a fly, and the two wings with the elytra of a beetle. Bivalve shells are made to open and shut, but on what

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and sometimes capable of neatly adapted movements through irritability or elasticity. and these carry the pollen from the anthers to the stigma. the pollen does not spontaneously fall on the stigma. formed of the most diverse parts. and when the bucket is half-full.a number of patterns is the hinge constructed. by being embedded in pulp or flesh. From such structures we may advance till we come to such a case of extraordinary adaptation as that lately described by Dr. by their capsule being converted into a light balloon-like envelope. being blown by the wind through mere chance on to the stigma. but such a view of nature is incredible. The basal part of the labellum stands over the bucket. With several kinds this is effected by the pollen-grains. which are light and incoherent. some aid is necessary for their fertilisation. and is itself hollowed out into a sort of chamber with two lateral entrances. so as to adhere to the fur of quadrupeds. as well as conspicuously coloured. From this simple stage we may pass through an inexhaustible number of contrivances. Cruger saw crowds of large 190 . from the long row of neatly interlocking teeth in a Nucula to the simple ligament of a Mussel! Seeds are disseminated by their minuteness. by having hooks and grapnels of many kinds and serrated awns. all for the same purpose and effected in essentially the same manner. though hermaphrodites. though very different plan occurs in many plants in which a symmetrical flower secretes a few drops of nectar. so as to attract and be devoured by birds. and with those in which. and this is the simplest plan which can well be conceived. But Dr. almost like toys in a shop. into which drops of almost pure water continually fall from two secreting horns which stand above it. and is consequently visited by insects. with the stamens and pistils modified in many ways. and rendered nutritious. The nectar may be stored in variously shaped receptacles. sometimes forming trap-like contrivances. so as to be wafted by every breeze. but entailing changes in every part of the flower. I will give one other instance: for this subject of the same end being gained by the most diversified means well deserves attention. With plants having separated sexes. The most ingenious man. An almost equally simple. Cruger in the Coryanthes. if he had not witnessed what takes place. the water overflows by a spout on one side. and by being furnished with wings and plumes. as different in shape as they are elegant in structure. Some authors maintain that organic beings have been formed in many ways for the sake of mere variety. This orchid has part of its labellum or lower lip hollowed out into a great bucket. could never have imagined what purpose all these parts serve. within this chamber there are curious fleshy ridges.

Dr. sensitive projection. and retain the pollen. first rubs its back against the viscid stigma and then against the viscid glands of the pollenmasses. but to gnaw off the ridges within the chamber above the bucket. like those of the Coryanthes. the variability will not be of the same 191 . The construction of the flower in another closely allied orchid. as I have called it. like an arrow. The passage is narrow. 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. not in order to suck nectar. transmits a sensation or vibration to a certain membrane which is instantly ruptured. namely. and rub against the properly placed viscid pollen-masses and the viscid stigma. or to the same flower a second time. in the foregoing and in innumerable other instances. How. thus provided. that when two forms vary.humble-bees visiting the gigantic flowers of this orchid. The answer no doubt is. though serving the same end. in doing this they frequently pushed each other into the bucket. can we understand the graduated scale of complexity and the multifarious means for gaining the same end. and adheres to it. or. and their wings being thus wetted they could not fly away. so that a bee. Dr. this sets free a spring by which the pollen-mass is shot forth. 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. Cruger saw a "continual procession" of bees thus crawling out of their involuntary bath. and adheres by its viscid extremity to the back of the bee. When the bee. but were compelled to crawl out through the passage formed by the spout or overflow. Now at last we see the full use of every part of the flower. and forces them to crawl out through the spout. and is pushed by its comrades into the bucket and then crawls out by the passage. This antenna. in order to gnaw the labellum. Bees visit these flowers. the Catasetum. tapering. with a pollen-mass still fastened to its back. of the water-secreting horns of the bucket half-full of water. and the flower is fertilised. in the right direction. as already remarked. it may be asked. flies to another flower. and is equally curious. and is roofed over by the column. is widely different. where it is brought into contact with the stigma. which is viscid enough to break certain elastic threads. when touched. the antenna. and are thus carried away. thus effecting fertilisation. with a bee which he had killed before it had quite crawled out. Cruger sent me a flower in spirits of wine. in doing this they inevitably touch a long. which prevents the bees from flying away. which already differ from each other in some slight degree. in forcing its way out.

We should also bear in mind that every highly developed organism has passed through many changes. It is certainly true. so that each modification will not readily be quite lost. for whatever purpose it may serve. yet. but somewhat exaggerated. Hence." Why. as Milne Edwards has well expressed it. for natural selection acts only by taking advantage of slight successive variations. on the theory of Creation. we can clearly understand why she should not." We meet with this admission in the writings of almost every experienced naturalist. then. but must advance by the short and sure. is the sum of many inherited changes. canon in natural history of "Natura non facit saltum. although in many cases it is most difficult even to conjecture by what transitions organs could have arrived at their present state. the structure of each part of each species.exact nature. or. Finally. 192 . though slow steps. considering how small the proportion of living and known forms is to the extinct and unknown. and consequently the results obtained through natural selection for the same general purpose will not be the same. and that each modified structure tends to be inherited. as indeed is shown by that old. she can never take a great and sudden leap. I have been astonished how rarely an organ can be named. but niggard in innovation. towards which no transitional grade is known to lead. through which the species has passed during its successive adaptations to changed habits and conditions of life. each supposed to have been separately created for its own proper place in nature. 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. should there be so much variety and so little real novelty? Why should all the parts and organs of many independent beings. that new organs appearing as if created for some special purpose rarely or never appear in any being. but may be again and again further altered. "Nature is prodigal in variety.

and it seems at first incredible that this could have been adapted for its present purpose by successive slight modifications. or from determining the attacks of insects. although now of very slight use. but they are incessantly harassed and their strength reduced. such as the down on fruit and the colour of its flesh. 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. or not so well enabled in a coming dearth to search for food. 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. its general presence and use for many purposes in so many land animals. I have sometimes felt great difficulty in understanding the origin or formation of parts of little importance. by the survival of the fittest. so that they are more subject to disease. have been transmitted to existing species in nearly the same state. Organs now of trifling importance have probably in some cases been of high importance to an early progenitor. as in the case of the most perfect and complex organs.7. Seeing how important an organ of locomotion the tail is in most aquatic animals. In a former chapter I have given instances of very trifling characters. would be able to range into new pastures and thus gain a great advantage. which in their lungs or modified swim-bladders betray their aquatic origin. or to escape from beasts of prey. might assuredly be acted on by natural selection. almost as great. and. and by the destruction of the less well-fitted individuals. after having been slowly perfected at a former period. but any actually injurious deviations in their structure would of course have been checked by natural selection. It is not that the larger quadrupeds are actually destroyed (except in some rare cases) by flies. As Affected By Natural Selection As natural selection acts by life and death. Organs Of Little Apparent Importance. the colour of the skin and hair of quadrupeds. from being correlated with constitutional differences. may perhaps be thus accounted for. yet we should pause before being too positive even in this case. The tail of the giraffe looks like an artificially constructed fly-flapper. though of a very different kind. it might subsequently come to be worked in for all sorts 193 . which. A well-developed tail having been formed in an aquatic animal. In the first place. for so trifling an object as to drive away flies. each better and better fitted.

bearing species in Africa and South America. may subsequently have been taken advantage of by its modified descendants. and which. is of the highest service to the plant. If green woodpeckers alone had existed. of so-called spontaneous variations. can double still more quickly. We must by no means overlook the effects of the definite action of changed conditions of life. or it may possibly be due to the direct action of putrid matter. the colour is probably in chief part due to sexual selection. for the hare. etc. as a fly-flapper. though the aid in this latter respect must be slight. of the tendency to reversion to long-lost characters. But structures thus indirectly gained. or may be indispensable for this act. as there is reason to believe from the distribution of the thorn. such as of correlation. we may easily err in attributing importance to characters. with hardly any tail. The naked skin on the head of a vulture is generally considered as a direct adaptation for wallowing in putridity. as it is. as it underwent further modification and became a climber.. comprehension. by which characters of use to one sex are often gained and then transmitted more or less perfectly to the other sex. but as we see nearly similar hooks on many trees which are not climbers. and had been acquired through natural selection. under new conditions of life and newly acquired habits. of the pressure of one part on another. and we did not know that there were many black and pied kinds. and in believing that they have been developed through natural selection.of purposes. and consequently that it was a character of importance. and so it may be. no doubt. 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. as in the case of the dog. but we should be very cautious in drawing any such inference. although at first of no advantage to a species. and this contrivance. an organ of prehension. 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 no doubt they facilitate. of the complex laws of growth. though of no use to the sex. and subsequently have been improved and taken advantage of by the plant. when we see that the skin on the head of the clean-feeding male turkey is likewise naked. In the second place. The sutures in the skulls of young mammals have been advanced as a beautiful adaptation for aiding parturition. so the spikes on the palm may at first have been developed for this object. and finally of sexual selection. which seem to depend in a quite subordinate degree on the nature of the conditions. which have only to 194 . but as sutures occur in the skulls of young birds and reptiles. or as an aid in turning. serve as a defence against browsing quadrupeds.

as we have good reason to believe. and we are immediately made conscious of this by reflecting on the differences between the breeds of our domesticated animals in different countries. if we are unable to account for the characteristic differences of our several domestic breeds. and I have made these remarks only to show that. where there has been but little methodical selection. Mountain breeds always differ from lowland breeds. the front limbs and the head would probably be affected. But we are far too ignorant to speculate on the relative importance of the several known and unknown causes of variation. The laborious breathing necessary in high regions tends. as H.escape from a broken egg. so that even colour would be thus subjected to the action of natural selection. and are exposed to a certain extent to natural selection. We are profoundly ignorant of the cause of each slight variation or individual difference. 195 . von Nathusius has lately shown in his excellent Treatise. and a mountainous country would probably affect the hind limbs from exercising them more. and again correlation would come into play. and that with the hair the horns are correlated. and possibly even the form of the pelvis. of the pelvis might affect by pressure the shape of certain parts of the young in the womb. The shape. which nevertheless are generally admitted to have arisen through ordinary generation from one or a few parent-stocks. to increase the size of the chest. Some observers are convinced that a damp climate affects the growth of the hair. together with abundant food. we may infer that this structure has arisen from the laws of growth. also. With cattle susceptibility to the attacks of flies is correlated with colour. and then by the law of homologous variation. on the whole organisation is probably still more important. and individuals with slightly different constitutions would succeed best under different climates. and this. The effects of lessened exercise. as is the liability to be poisoned by certain plants. is apparently one chief cause of the great modification which the breeds of swine have undergone. and has been taken advantage of in the parturition of the higher animals. we ought not to lay too much stress on our ignorance of the precise cause of the slight analogous differences between true species. Animals kept by savages in different countries often have to struggle for their own subsistence. more especially in the less civilized countries.

So we may believe that the progenitor of the seal did not possess a flipper. and consequently. and may never have been of any use to their progenitors. in the fore leg of the horse. but a foot with five toes fitted for walking or grasping. as they now are to the most aquatic of living birds. or was formerly. as the definite action of external conditions. we may conclude that the structure of every living creature either now is. probably a great effect. we can hardly believe that the webbed feet of the upland goose. Thus. were originally developed. probably through the reduction of more numerous bones in the fin of some ancient fish-like progenitor of the whole class. have all produced an effect. lately specified. a view already discussed. No doubt the definite action of changed conditions.bird. in the wing of the bat. Such doctrines. It is scarcely possible to decide how much allowance ought to be made for such causes of change. They believe that many structures have been created for the sake of beauty. or for the sake of mere variety. if true. Utilitarian Doctrine. How Far True: Beauty. on the principle of utility. are of special use to these animals. independently of any advantage thus gained. and we may further venture to believe that the several bones in the limbs of the monkey. we cannot believe that the similar bones in the arm of the monkey. horse and bat. to delight man or the Creator (but this latter point is beyond the scope of scientific discussion). so-called spontaneous variations. would be absolutely fatal to my theory. though each being assuredly is well fitted for its place in nature. but with these important exceptions. and the various causes of modifications. or of the frigate. and the complex laws of growth. and in the flipper of the seal. 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 196 . I fully admit that many structures are now of no direct use to their possessors. How Acquired 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. of some direct or indirect use to its possessor. are of special use to these birds.8. but this does not prove that they were formed solely for beauty or variety. But a still more important consideration is that the chief part of the organisation of every living creature is due to inheritance. But webbed feet no doubt were as useful to the progenitor of the upland goose and of the frigate-bird. We may safely attribute these structures to inheritance. many structures have now no very close and direct relation to present habits of life.

not coloured. we may conclude that. reptiles. docks and nettles. A similar line of argument holds good with fruits. destitute of nectar. 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. nut and ash trees. the other closed. If beautiful objects had been created solely for man's gratification. 197 . is apparently wholly due to symmetry of growth. 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). and mammals. in order that the fruit may be devoured and the matured seeds disseminated. We see this.subversive of my whole theory—I may first remark that the sense of beauty obviously depends on the nature of the mind. But this beauty serves merely as a guide to birds and beasts. and in consequence at the same time beautiful. if it be coloured of any brilliant tint. On the other hand. and that the idea of what is beautiful. 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. some fishes. is not innate or unalterable. our plants would not have been decked with beautiful flowers. I willingly admit that a great number of male animals. for instance. Hence. 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. Were the beautiful volute and cone shells of the Eocene epoch. irrespective of any real quality in the admired object. as all our most gorgeous birds. and in many others. and the gracefully sculptured ammonites of the Secondary period. in the men of different races admiring an entirely different standard of beauty in their women. spinach. on grasses. so that they may be easily observed by insects. if insects had not been developed on the face of the earth. 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. or rendered conspicuous by being white or black. Several plants habitually produce two kinds of flowers. but they have been rendered conspicuous in contrast with the green leaves. which are all fertilised through the agency of the wind. Flowers rank among the most beautiful productions of nature. but would have produced only such poor flowers as we see on our fir. one kind open and coloured so as to attract insects. and never visited by insects. oak.

instead of to the males alone. 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. I cannot find even one which seems to me of any weight. though throughout nature one species incessantly takes advantage of. I would almost as soon believe that the cat curls the end of its tail when preparing to spring. Although many statements may be found in works on natural history to this effect. and others displeasure. namely. as we see in the fang of the adder.and a host of magnificently coloured butterflies. How the sense of beauty in its simplest form—that is. It is a much more probable view that the rattlesnake uses its rattle. it would annihilate my theory. in order to warn the doomed mouse. But this has been effected through sexual selection. and not for the delight of man. in order to alarm the many birds and beasts which are known to attack even the most venomous species. which is not rarely the case with birds and butterflies. is a very obscure subject. and profits by the structures of others. but some authors suppose that at the same time it is furnished with a rattle for its own injury. to warn its prey. but there must be some fundamental cause in the constitution of the nervous system in each species. Natural selection cannot possibly produce any modification in a species exclusively for the good of another species. So it is with the music of birds. the cause apparently lies in the colours acquired through sexual selection having been transmitted to both sexes. When the female is as beautifully coloured as the male. forms and sounds—was first developed in the mind of man and of the lower animals. by which its eggs are deposited in the living bodies of other insects. Snakes act on the same principle which makes the hen ruffle her feathers and expand her wings when a dog 198 . the cobra expands its frill and the puff-adder swells while hissing so loudly and harshly. Habit in all these cases appears to have come to a certain extent into play. and in the ovipositor of the ichneumon. have been rendered beautiful for beauty's sake. the reception of a peculiar kind of pleasure from certain colours. for such could not have been produced through natural selection. But natural selection can and does often produce structures for the direct injury of other animals. It is admitted that the rattlesnake has a poison-fang for its own defence and for the destruction of its prey. that is. 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. 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.

If a fair balance be struck between the good and evil caused by each part. owing to the backward serratures. are perfect. as myriads have become extinct. with this high standard under nature. Natural selection will never produce in a being any structure more injurious than beneficial to that being. The endemic productions of New Zealand. this same reason tells us. as a boring and serrated instrument. or if it be not so. it will be modified. for natural selection acts solely by and for the good of each. No organ will be formed. After the lapse of time. whose judgment no one will dispute. cannot be withdrawn. The correction for the aberration of light is said by Muller not to be perfect even in that most perfect organ. under changing conditions of life. like that in so many members of the same great order. nor do we always meet. Helmholtz. And we see that this is the standard of perfection attained under nature. as having existed in a remote progenitor. one compared with another. when used against many kinds of enemies. with the poison originally adapted for 199 . Natural selection will not produce absolute perfection. But I have not space here to enlarge on the many ways by which animals endeavour to frighten away their enemies. though we may easily err on both sides. Natural selection tends only to make each organic being as perfect as. and that it has since been modified but not perfected for its present purpose.approaches her chickens. as far as we can judge. each will be found on the whole advantageous. and thus inevitably causes the death of the insect by tearing out its viscera? If we look at the sting of the bee. Can we consider the sting of the bee as perfect. as Paley has remarked. the being will become extinct. that some other contrivances are less perfect. is as nothing in comparison with the incongruities which we have just come across in the domain of the sensations." If our reason leads us to admire with enthusiasm a multitude of inimitable contrivances in nature. for the purpose of causing pain or for doing an injury to its possessor. 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. but they are now rapidly yielding before the advancing legions of plants and animals introduced from Europe. for instance. or slightly more perfect than the other inhabitants of the same country with which it comes into competition. 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. which. if any part comes to be injurious. after describing in the strongest terms the wonderful powers of the human eye. the human eye.

such as to produce galls. her daughters. If we admire the truly wonderful power of scent by which the males of many insects find their females. for undoubtedly this is for the good of the community. If we admire the several ingenious contrivances by which orchids and many other plants are fertilised through insect agency. is all the same to the inexorable principles of natural selection. which urges her to destroy the young queens. as soon as they are born. can we consider as equally perfect the elaboration of dense clouds of pollen by our fir-trees. or to perish herself in the combat. 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. though the latter fortunately is most rare.some other object. since intensified. and which are ultimately slaughtered by their industrious and sterile sisters? It may be difficult. which are utterly useless to the community for any other purpose. so that a few granules may be wafted by chance on to the ovules? 200 . but we ought to admire the savage instinctive hatred of the queen-bee. it will fulfil all the requirements of natural selection. can we admire the production for this single purpose of thousands of drones. and maternal love or maternal hatred. though it may cause the death of some few members.

and petrels with the habits of auks. will have a great advantage over the less numerous intermediate variety. In the cases in which we know of 201 . 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. during the course of further modification. now living on a continuous area. ground woodpeckers. We have seen that species at any one period are not indefinitely variable. Hence we can understand. bearing in mind that each organic being is trying to live wherever it can live. and partly because the very process of natural selection implies the continual supplanting and extinction of preceding and intermediate gradations. yet in the case of any organ. but I think that in the discussion light has been thrown on several facts. Many of them are serious. that a bat. which on the belief of independent acts of creation are utterly obscure. how it has arisen that there are upland geese with webbed feet. Although the belief that an organ so perfect as the eye could have been formed by natural selection. from existing in greater numbers. When two varieties are formed in two districts of a continuous area. Closely allied species. fitted for an intermediate zone. is enough to stagger any one. with some very unlike those of its nearest congeners. and when the conditions of life did not insensibly graduate away from one part to another. must often have been formed when the area was not continuous. and are not linked together by a multitude of intermediate gradations. We have seen that a species under new conditions of life may change its habits. each good for its possessor. there is no logical impossibility in the acquirement of any conceivable degree of perfection through natural selection. or it may have diversified habits. for instance. could not have been formed by natural selection from an animal which at first only glided through the air. partly because the process of natural selection is always very slow. Summary: The Law Of Unity Of Type And Of The Conditions Of Existence Embraced By The Theory Of Natural Selection We have in this chapter discussed some of the difficulties and objections which may be urged against the theory. if we know of a long series of gradations in complexity. but from reasons assigned. and will thus generally succeed in supplanting and exterminating it. diving thrushes. consequently the two latter. then under changing conditions of life. the intermediate variety will usually exist in lesser numbers than the two forms which it connects. an intermediate variety will often be formed. and at any one time acts only on a few forms.9.

On the other hand. and this naturally follows from the principle of natural selection. a swim-bladder has apparently been converted into an air-breathing lung. independently of any good having been thus gained. the one having been perfected whilst aided by the other. We have seen that in two beings widely remote from each other in the natural scale. only in accordance with the standard of that particular country. but when such organs are closely examined. organs. We may. modifications are probably the direct result of the laws of variation or of growth. and two distinct organs having performed at the same time the same function. For instance. In each well-stocked country natural selection acts through the competition of the inhabitants and consequently leads to success in the battle for life. The same organ having performed simultaneously very different functions. for the metamorphoses of many organs show what wonderful changes in function are at least possible. but in all cases at the same time useful to the possessor. essential differences in their structure can almost always be detected. we should be extremely cautious in concluding that none can have existed. and excretions highly useful or even indispensable.no intermediate or transitional states. have been acquired by means of natural selection. organs serving for the same purpose and in external appearance closely similar may have been separately and independently formed. Hence the inhabitants of one country. the common rule throughout nature is infinite diversity of structure for gaining the same end. as we may feel assured. Natural selection can produce nothing in one species for the exclusive good or injury of another. But even such structures have often. 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. and this again naturally follows from the same great principle. or highly injurious to another species. generally the smaller one. been subsequently taken advantage of. that modifications in its structure could not have been slowly accumulated by means of natural selection. and then having been in part or in whole specialised for one function. must often have largely facilitated transitions. and still further modified. for the good of species under new conditions of life. though it may well produce parts. In many other cases. also. in its present state. believe that a part formerly of high importance has frequently been retained (as the tail of an aquatic animal by its terrestrial descendants). though it has become of such small importance that it could not. often yield to the inhabitants of another and generally the larger country. For in the 202 .

nor. Natural selection will not necessarily lead to absolute perfection. if we look to the present inhabitants alone of the world. On the theory of natural selection we can clearly understand the full meaning of that old canon in natural history. unity of type is explained by unity of descent. "Natura non facit saltum. that of Unity of Type. Hence.larger country there will have existed more individuals. and thus the standard of perfection will have been rendered higher. as it includes. being affected by the direct action of external conditions of life. and which is quite independent of their habits of life. in fact. and the Conditions of Existence. The expression of conditions of existence. 203 . but if we include all those of past times. the law of the Conditions of Existence is the higher law. whether known or unknown. 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. can absolute perfection be everywhere predicated. so often insisted on by the illustrious Cuvier. On my theory. By unity of type is meant that fundamental agreement in structure which we see in organic beings of the same class." This canon. it must on this theory be strictly true. as far as we can judge by our limited faculties. is fully embraced by the principle of natural selection. and more diversified forms. is not strictly correct. and the competition will have been severer. through the inheritance of former variations and adaptations. and subjected in all cases to the several laws of growth and variation. 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.

with some parade of mathematical accuracy. Thus a distinguished German naturalist has asserted that the weakest part of my theory is. A critic has lately insisted. as some of the previous discussions may thus be made clearer. as many have been made by writers who have not taken the trouble to understand the subject. that all are not as perfect as they might have been in relation to their conditions. I will devote this chapter to the consideration of various miscellaneous objections which have been advanced against my views. when their conditions changed. and no one will dispute that the physical conditions of each country. 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. Nor can organic beings. that longevity is a great advantage to all species. 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. developed from one and the same source — Reasons for disbelieving in great and abrupt modifications. have undergone many mutations. 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! 204 . have remained so.Chapter 7 Miscellaneous Objections To The Theory Of Natural Selection Longevity — Modifications not necessarily simultaneous — Modifications apparently of no direct service — Progressive development — Characters of small functional importance. even if they were at any one time perfectly adapted to their conditions of life. but it would be useless to discuss all of them. unless they themselves likewise changed. as well as the number and kinds of its inhabitants. that I consider all organic beings as imperfect: what I have really said is.

survive from year to year by means of its seeds or ova? Mr. are closely similar or even identical with those now living. during the last several thousand years. the more permanent varieties are generally found. and all mere temporary variations. and if we lay on one side polymorphic species. The many animals which have remained unchanged since the commencement of the glacial period. as far as I can discover.. such as high land or low land. such as size. Lewes has remarked. for the ancient domestic races figured on the Egyptian monuments. in the case of 205 . have remained absolutely uniform. The fact of little or no modification having been effected since the glacial period. have changed during the last three or four thousand years. as none of the animals and plants of Egypt. as well as to the amount of expenditure in reproduction and in general activity. can a variety live side by side with the parent species? If both have become fitted for slightly different habits of life or conditions. as Mr. dry or moist districts. of which we know anything. Bronn. been largely determined through natural selection. but is powerless against the doctrine of natural selection or the survival of the fittest. or embalmed. inhabiting distinct stations. but this will be effected only under certain favourable circumstances. they might live together. that longevity is generally related to the standard of each species in the scale of organisation. on the principle of natural selection. the conditions of life. so probably have none in any part of the world. as far as we know. But. in Egypt. it is probable. these will be preserved. would have been an incomparably stronger case. The celebrated palaeontologist. and yet. albinism. E. and he concludes. for these have been exposed to great changes of climate and have migrated over great distances. this line of argument proves too much. at the close of his German translation of this work. whereas. as far as its extreme complexity allows him to form a judgment. Ray Lankester has recently discussed this subject. asks how. It has been argued that. G. owing to advantages gained through natural selection. yet all naturalists admit that such races have been produced through the modification of their original types. which implies that when variations or individual differences of a beneficial nature happen to arise. in which the variability seems to be of a peculiar nature. would have been of some avail against those who believe in an innate and necessary law of development. etc. Moreover.H.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. And these conditions have.

whether it be the flower. be acquired by successive variations. or at the greyhound and mastiff. almost all the other parts have been slightly modified. has been greatly changed. but in many parts. and a multitude of analogous cases. and partly to so-called spontaneous variation. they would appear to us as if they had been simultaneously developed. namely. as was formerly remarked. or leaves. but he insists that the families of plants differ chiefly from each other in morphological characters. 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. that many characters appear to be of no service whatever to their possessors. if slight. excellently adapted for some purpose. 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. chiefly through man's power of selection. This may be attributed partly to the principle of correlated growth. and he asks. might. which appear to be quite unimportant for the welfare of the species. and even their mental characteristics. and therefore cannot have been influenced through natural selection.animals which wander much about and cross freely. The most striking modifications. He specifies the arrangement of the cells in the tissues. this subject has been discussed by Nageli in an admirable essay. as cases in which natural selection could not have acted. their varieties seem to be generally confined to distinct regions. 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. and recently by Broca. and of the leaves on the axis. and as they would be transmitted all together. 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. to the above objection is afforded by those domestic races which have been modified. To these may be added the 206 . Look at the race and dray-horse. for some special purpose. He admits that natural selection has effected much. first in one part and then in another. but first one part and then another slightly modified and improved. The best answer. Their whole frames. Bronn also insists that distinct species never differ from each other in single characters. A much more serious objection has been urged by Bronn. With respect to plants. fruit. have been modified. He consequently believes in an innate tendency towards progressive and more perfect development. however.

through certain dimly seen causes. I can no more believe in this than that the well-adapted form of a racehorse or greyhound. 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. which we do not in the least understand. In the second place. Nevertheless. of use to each species. for the sake of brevity. it should always be borne in mind that when one part is modified. under the expression of the laws of growth. which are so highly developed that no one doubts that they are of importance. the position of the ovules. There is much force in the above objection. so will be other parts. In the third place. to be extremely cautious in pretending to decide what structures now are. the frequency and importance of modifications due to spontaneous variability. ascertained.numerical divisions in the parts of the flower. due to some change in the conditions. In the earlier editions of this work I underrated. as it now seems probable. It may be worth while to illustrate some of the foregoing remarks. etc. in the first place. and so forth —as well as through other causes which lead to the many mysterious cases of correlation. or has only recently been. offer good instances of spontaneous variations. such as the appearance of a moss-rose on a common rose. These agencies may be all grouped together. when not of any use for dissemination. as well as for more strongly marked variations which occasionally arise. mutual pressure. and if the unknown cause were to act persistently. which before the principle of selection by man was well understood. in which the nature of the conditions apparently plays a quite subordinate part. it is almost certain that all the individuals of the species would be similarly modified. or have formerly been. we have to allow for the direct and definite action of changed conditions of life. yet their use has not been. it is hardly necessary to observe that even in the higher and best-known animals many structures exist. the shape of the seed. Bud-variations. or of a nectarine on a peach-tree. There must be some efficient cause for each slight individual difference. can thus be explained. an early developed part affecting one subsequently developed. if we bear in mind the power of a minute drop of poison in producing complex galls. 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. and for so-called spontaneous variations. such as an increased or diminished flow of nutriment to a part. excited so much surprise in the minds of the older naturalists. With respect to the assumed inutility of various parts and organs. but even in these cases. As Bronn gives the length of the ears and 207 . we ought.

and in others the lower ones alone are fertilised. and their arrangement. With respect to plants. Several plants belonging to distinct orders habitually produce flowers of two kinds—the one open. of the ordinary structure. We shall. but Dr. one ovule holds the former and a second ovule the latter position. yet may be seen to graduate into each other on the same plant. according to Dr. could have been of any service. Schobl. and its use would be much influence by its length. but now we know this to be the case. of differences in structure which can be of no special use.tail in the several species of mice as instances. also. No one until lately would have imagined that in dimorphic and trimorphic plants the different lengths of the stamens and pistils. The closed and imperfect flowers are. the other closed and imperfect. as they yield with the utmost safety a large stock of seed. it will be admitted that the flowers of the orchids present a multitude of curious structures. with the expenditure of wonderfully little pollen. the external ears of the common mouse are supplied in an extraordinary manner with nerves. and in others they are suspended. though trifling ones. These positions seem at first purely morphological. If so. The petals in the imperfect 208 . and the benefits which certainly are derived from this process are thus secured. and have probably been gained through natural selection. even when one is erect and the other suspended within the same ovarium. which a few years ago would have been considered as mere morphological differences without any special function. to which on account of Nageli's essay I shall confine myself in the following remarks. Hooker informs me that within the same ovarium the upper ovules alone in some cases. manifestly of high importance. would follow the selection of any slight deviations in position which favoured their fertilisation. The ordinary and open flowers can be intercrossed. These two kinds of flowers sometimes differ wonderfully in structure. the position of the ovules. I may mention that. in structure. presently see that the tail is a highly useful prehensile organ to some of the species. and the production of seed. The two kinds of flowers often differ much. but they are now known to be of the highest importance for the fertilisation of the species through the aid of insects. and within the same ovarium of some few plants. or of no physiological signification. In certain whole groups of plants the ovules stand erect. as just stated. so that they no doubt serve as tactile organs. hence the length of the ears can hardly be quite unimportant. however. and he suggests that this probably depends on the direction in which the pollen-tubes enter the ovarium.

the sepals are reduced from the normal number of five to three. are still further modified. one flower.trees. the angles of divergence of the leaves differ. In many Compositae and Umbelliferae (and in some other plants) the circumferential flowers have their corollas much more developed than those of the centre. In one section of the Malpighiaceae the closed flowers. colour and other characters. and five divisions to the ovarium. while the surrounding flowers generally have three calyx-lobes with the other organs pentamerous. opens first.flowers almost always consist of mere rudiments. and this seems often connected with the abortion of the reproductive organs. while all the other flowers on the plant are tetramerous. due to differences in relative position on the same plant. and has five sepals and petals. It is a more curious fact. a sixth stamen standing opposite to a petal being alone developed. Now although natural selection may well have had the power to prevent some of the flowers from expanding. the style is aborted. In Carthamus and some other Compositae the central achenes alone are furnished with a pappus. according to Tausch. but must have followed from the laws of growth. during the progress of the reduction of the pollen and the closure of the flowers. In six out of thirty of the closed flowers in an Indian violet (name unknown. that I will give some additional cases of another kind. for the five stamens which stand opposite to the sepals are all aborted. that the achenes or seeds of the circumference and centre sometimes differ greatly in form. previously referred to. and this stamen is not present in the ordinary flowers of this species. for the plants have never produced with me perfect flowers). according to A. according to Schacht. In the common rue and some other plants. are 209 . and the pollen-grains are reduced in diameter. and in certain fir. and to reduce the amount of pollen. and in some species of Viola three stamens are in this state. In the British Adoxa the uppermost flower generally has two calyx-lobes with the other organs tetramerous. and in Hyoseris the same head yields achenes of three different forms. including the functional inactivity of parts. de Jussieu. in the nearly horizontal and in the upright branches. In Ononis columnae five of the alternate stamens are rudimentary. It is so necessary to appreciate the important effects of the laws of growth. two retaining their proper function. and the ovaria are reduced from three to two. when rendered by the closure of the flowers superfluous. namely of differences in the same part or organ. yet hardly any of the above special modifications can have been thus determined. In the Spanish chestnut. In certain Umbelliferae the exterior seeds. but being of very small size. usually the central or terminal one.

s'etend entre le pericarpe et le placenta. but Professor Asa Gray states that with some species of Mimulus. Of their cause we are quite ignorant. one-seeded nutlets. the flowers of Papaver bracteatum offer either two sepals with four petals (which is the common type with poppies). in which the flowers in the lower part of the spike bear oval. to any proximate agency. St. affecting only some of the flowers on the same plant. flowers indifferently tetramerous. etc. St. as in the last class of cases. In Helianthemum the capsule has been described as unilocular or tri-locular. 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. which are considered by botanists to be generally of a highly important nature. that I need not give examples. have come into play. Lastly. or only in a quite subordinate manner. with the exception of that of the well-developed ray. Professor Braun mentions a Fumariaceous genus. with either one or two ovaries. or occurring on distinct plants. I will give only a few instances. As these variations seem of no special use to the plants.orthospermous. The manner in which the petals are folded in the bud is in most groups a very constant morphological character. and even in the same panicle. as far as we can judge. and this is a character which was considered by De Candolle to be in other species of the highest systematic importance. "Une lame PLUS OU MOINS LARGE. which are of service in making the flowers conspicuous to insects. or three sepals with six petals. In numerous other cases we find modifications of structure. two-valved and two-seeded siliques. as are the flowers and leaves in certain positions.. Aug. but in some species flowers may be found on the same plant. Hilaire found 210 . All these modifications follow from the relative position and inter-action of the parts. pentamerous. we cannot even attribute them. which grow close together under the same conditions. such as relative position. to which latter tribe the genus belongs. Masters has observed instances of both marginal and free central placentation. lanceolate. It is so common to observe on the same plant. they cannot have been influenced by natural selection. according to De Candolle. the aestivation is almost as frequently that of the Rhinanthideae as of the Antirrhinideae. but as numerical variations are comparatively rare when the parts are few." In the flowers of Saponaria officinalis Dr. In these several cases. and in H. natural selection cannot. I may mention that. mutabile.florets. Hilaire gives the following cases: the genus Zanthoxylon belongs to a division of the Rutaceae with a single ovary. ribbed. and in the upper part of the spike. and the central one coelospermous. all would have been modified in the same manner.

towards the southern extreme of the range of Gomphia oleaeformis two forms which he did not at first doubt were distinct species. It cannot have been of much importance to the greater number of mammals. independently of natural selection. feathers to all birds. any slight variations which occurred in them would not have been accumulated and augmented through natural selection. and consequently is often 211 . we may infer that the result would be almost uniform. A structure. closed flowers. and so it must be with many parasitic and degraded animals. The acquisition of a useless part can hardly be said to raise an organism in the natural scale. of whatever importance they may generally be to us for our classifications. and in this case all the individuals of the species would be modified in the same manner. But with respect to Nageli's doctrine of an innate tendency towards perfection or progressive development. "Voila donc dans un meme individu des loges et un style qui se rattachent tantot a un axe verticale et tantot a un gynobase. I should infer from the mere fact of the parts in question differing or varying greatly on the same plant. when it ceases to be of service to a species. birds. But when. for it will no longer be regulated by this same power of selection. as we see with rudimentary organs." We thus see that with plants many morphological changes may be attributed to the laws of growth and the inter-action of parts. generally becomes variable. that such modifications were of extremely small importance to the plants themselves. and scales to all true reptiles. descendants. is ranked by us as of high systematic importance. that the plants have been caught in the act of progressing towards a higher state of development? On the contrary. from the nature of the organism and of the conditions. A structure which has been developed through long-continued selection. if any new principle has to be invoked. but if the unknown cause were to act almost uniformly for a length of time. it must be one of retrogression rather than of progression. but he subsequently saw them growing on the same bush. We are ignorant of the exciting cause of the above specified modifications. otherwise modified. whatever it may be. feathers or scales. they may be. yet hair has been transmitted to almost all mammals. can it be said in the case of these strongly pronounced variations. and in the case of the imperfect. which is common to many allied forms. and apparently often have been. From the fact of the above characters being unimportant for the welfare of the species. modifications have been induced which are unimportant for the welfare of the species. or reptiles. transmitted in nearly the same state to numerous. and he then adds. above described. whether they were clothed with hair.

Thus. that characters of slight vital importance to the species. any slight deviations in them could not have been governed or accumulated through this latter agency. For the best definition which has ever been given of a high standard of organisation. the position of the ovules. It is a strange result which we thus arrive at. as I have attempted to show in the fourth chapter. etc. as I am inclined to believe.assumed to be of high vital importance to the species. morphological differences. but not through natural selection. Mivart's book. Mivart passes over the effects of the increased use and disuse of parts. but. no slight effort of reason and memory is left to the reader. A distinguished zoologist. and have treated in my "Variation under Domestication" at greater length than. Although we have no good evidence of the existence in organic beings of an innate tendency towards progressive development. 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. as I believe. George Mivart. which we consider as important—such as the arrangement of the leaves. My judgment may not be trustworthy. St. and natural selection tends towards this end. this is by no means so paradoxical as it may at first appear. any other writer. Mr. first appeared in many cases as fluctuating variations. has recently collected all the objections which have ever been advanced by myself and others against the theory of natural selection. is the degree to which the parts have been specialised or differentiated. namely. for as these morphological characters do not affect the welfare of the species. inasmuch as the parts are thus enabled to perform their functions more efficiently. When discussing special cases. and comparing each section with what I have said on the same head. through the continued action of natural selection. as propounded by Mr. When thus marshalled. and has illustrated them with admirable art and force. which I have always maintained to be highly important. Mivart's plan to give the various facts and considerations opposed to his conclusions. I never before felt so strongly convinced of the general 212 . are the most important to the systematist. and as it forms no part of Mr. Wallace and myself. independently of natural selection. Mr. as well as through the intercrossing of distinct individuals. as we shall hereafter see when we treat of the genetic principle of classification. He likewise often assumes that I attribute nothing to variation.. yet this necessarily follows. which sooner or later became constant through the nature of the organism and of the surrounding conditions. the divisions of the flower or of the ovarium. who may wish to weigh the evidence on both sides. but after reading with care Mr. they make a formidable array.

These slight proportional differences. the conversion of a swim-bladder into lungs. to much partial error. These cattle can browse as well as others on grass. of course. in which careful measurements are given. I will here consider in some detail several of the cases advanced by Mr. during such periods. browse on the twigs of trees. and this must be a great advantage to it during dearths. for they will have roamed over the whole country in search of food. It can thus obtain food beyond the reach of the other Ungulata or hoofed animals inhabiting the same country. for those individuals which had some one part or several parts of their bodies rather more 213 . points which were discussed in the last chapter under two headings. etc. but from the projection of the lower jaw they cannot. during the often recurrent droughts. without necessarily having attended to special points of structure. The Niata cattle in South America show us how small a difference in structure may make. for instance. subject. as want of space prevents me from considering all. in so intricate a subject. considered in the present volume. by breeding from the victorious birds. 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.. much elongated neck. by simply preserving and breeding from the fleetest individuals. Mivart's objections." This subject is intimately connected with that of the gradation of the characters. head and tongue. "That natural selection is incompetent to account for the incipient stages of useful structures. has its whole frame beautifully adapted for browsing on the higher branches of trees. or have been.truth of the conclusions here arrived at. due to the laws of growth and variation. the individuals which were the highest browsers and were able during dearths to reach even an inch or two above the others. reeds. Before coming to Mr. The giraffe. Man has modified some of his animals. considering its probable habits of life. Mivart's objections will be. The one new point which appears to have struck many readers is. often accompanied by a change of function. to which food the common cattle and horses are then driven. Nevertheless. so that at these times the Niatas perish. a great difference in preserving an animal's life. it may be well to explain once again how natural selection will act in all ordinary cases. if not fed by their owners. fore legs. All Mr. But it will have been otherwise with the nascent giraffe. So under nature with the nascent giraffe. will often have been preserved. Mivart. or as with the game-cock. as with the race-horse and greyhound. by its lofty stature. are not of the slightest use or importance to most species. selecting those which are the most illustrative.

more than counterbalance the advantages. which exactly corresponds with what I have called unconscious selection by man. The preservation of each species can rarely be determined by any one advantage. and. Mivart brings forward two objections. Mr. by violently swinging its head armed with stump-like horns. in a lesser degree. or with a tendency to vary again in the same manner. taller than an ox. why has not any other hoofed quadruped acquired a long neck and lofty stature. again. would generally have survived. and will destroy all the inferior individuals. while the individuals less favoured in the same respects will have been the most liable to perish. Chauncey Wright has remarked. By this process long-continued. We here see that there is no need to separate single pairs. as Mr. These will have intercrossed and left offspring. to a supply of food. but by the union of all. as Sir S. serve as a watch-tower. Baker remarks. the camel. and as some of the largest antelopes in the world. would have been of some advantage to the nascent giraffe. when he methodically improves a breed: natural selection will preserve and thus separate all the superior individuals. if natural selection be so potent. combined. that no animal is more difficult to stalk than the giraffe. and he considers it as "very problematical whether the disadvantages thence arising would not. Assuredly the being able to reach. it seems to me almost certain that an ordinary hoofed quadruped might be converted into a giraffe. and against this animal. why should we doubt that. intermediate gradations could formerly have existed there. left untouched by the other hoofed quadrupeds of the country. abound there. subjected as now to severe dearths. in times of scarcity.elongated than usual. This animal also uses its long neck as a means of offence or defence. that increased bulk would act as a protection against almost all beasts of prey excepting the lion. its tall neck—and the taller the better— would. allowing them freely to intercross. at each stage of increased size." But as the giraffe does actually exist in large numbers in Africa. in a most important manner with the inherited effects of the increased use of parts. either inheriting the same bodily peculiarities. It is from this cause. besides the giraffe. great and small. and if high browsing be so great an advantage. why has not any member of the group 214 . as far as size is concerned. To this conclusion Mr. no doubt. Nor must we overlook the fact. Mivart then asks (and this is his second objection). as man does. guanaco and macrauchenia? Or. One is that the increased size of the body would obviously require an increased supply of food.

In order that an animal should acquire some structure specially and largely developed. to sheep. Although every part of the body varies slightly. We can. it does not follow that the necessary parts should always vary in the right direction and to the right degree. and we know that some areas support singularly few large quadrupeds. for which hoofed animals are singularly ill-constructed) implies greatly increased bulk of body. though it is so luxuriant. in which trees grow. nor why the later tertiary periods should have been much more favourable for their existence than the present time. which was formerly inhabited by numerous herds of the giraffe. to acquire slightly longer necks? In every district some one kind of animal will almost certainly be able to browse higher than the others. the answer is not difficult. and not with the other ungulate animals. In South Africa the competition for browsing on the higher branches of the acacias and other trees must be between giraffe and giraffe. it is almost indispensable that several other parts should be modified and coadapted. cannot be distinctly answered. and that some species are much more variable 215 . for instance South America. for instance. To reach the foliage at a considerable height (without climbing. Why. 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. through natural selection and the effects of increased use. and we cannot even conjecture what changes of structure would be favourable to its increase in some new country. 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. various animals belonging to this same order have not acquired either an elongated neck or a proboscis. in other quarters of the world. With the different species of our domesticated animals we know that the parts vary in a different manner and degree. In every meadow in England. Whatever the causes may have been. and it is almost equally certain that this one kind alone could have its neck elongated for this purpose. however. we see the lower branches trimmed or planed to an exact level by the browsing of the horses or cattle. Why this should be so we do not know.acquired a long proboscis? With respect to South Africa. see in a general manner that various causes might have interfered with the development of a long neck or proboscis. We are ignorant with respect to the conditions which determine the numbers and range of each species. and what advantage would it be. while South Africa abounds with them to an unparalleled degree. and can best be given by an illustration. if kept there.

or will be greatly retarded. will be favoured only under certain peculiar conditions. they must have long inhabited their present homes. for the sake of escaping from their enemies. Gradations of structure.than others. it does not follow that natural selection would be able to act on them and produce a structure which apparently would be beneficial to the species. if the number of individuals existing in a country is determined chiefly through destruction by beasts of prey—by external or internal parasites. for the bats groundinsects would serve as food. which it is thought would be beneficial to certain species. have probably interfered with the acquisition through natural selection of structures. A strictly terrestrial animal. yet as some of these bats are peculiar species. in many quarters of the world. might at last be converted into an animal so thoroughly aquatic as to brave the open ocean. which first colonise and abound on most oceanic islands. then in streams or lakes. Lyell asks. natural selection is a slow process. then natural selection will be able to do little. But seals would not find on oceanic islands the conditions favourable to their gradual reconversion into a terrestrial form. and the same favourable conditions must long endure in order that any marked effect should thus be produced. in modifying any particular structure for obtaining food. Oceanic islands are inhabited by bats and seals. In each case various causes. and assigns certain reasons in answer. Lastly. as formerly shown. Bats. For instance. Objections of the same nature as the foregoing have been advanced by many writers. besides the general ones just indicated. etc. probably acquired their wings by at first gliding through the air from tree to tree. hoofed quadrupeds have not acquired much elongated necks or other means for browsing on the higher branches of trees. and bats into terrestrial insectivorous animals. by occasionally hunting for food in shallow water. but by no terrestrial mammals. 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.—as seems often to be the case. but these would already be largely preyed on by the reptiles or birds. but when the power of true 216 . Except by assigning such general and vague reasons. for the former there would be no prey. we cannot explain why. Therefore Sir C. with each stage beneficial to a changing species. One writer asks. Even if the fitting variations did arise. or for avoiding falls. 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. like the so-called flying squirrels.

The caterpillars which project motionless like dead twigs from the bushes on which they feed. as such development would be advantageous to all? Why have not apes acquired the intellectual powers of man? Various causes could be assigned. are rare and exceptional. Darwin's theory. and as the minute incipient variations will be in ALL DIRECTIONS. These conjectural remarks have been made merely to show that a transition of structure. Insects often resemble for the sake of protection various objects. have had their wings greatly reduced in size. The cases of the imitation of such objects as the excrement of birds. there is a constant tendency to indefinite variation. for natural selection to seize upon and perpetuate. A definite answer to the latter question ought not to be expected. bits of lichen. is a highly complex affair. and living insects. Mr. by the aid of their hind legs alone. they must tend to neutralize each other. spines. "As. We will return to Mr. more than one writer has asked why have some animals had their mental powers more highly developed than others. at least for the above purposes. seeing that no one can solve the simpler problem." But in all the foregoing cases the insects in their original state no doubt presented some rude and accidental resemblance to an object commonly 217 . bamboo. so as to compete with birds or other ground animals. or other object. and their relative probability cannot be weighed. Mivart's other objections. it would be useless to give them. and is not confined to colour. excrement of birds. through disuse. flowers. like many birds. might. and at first to form such unstable modifications that it is difficult. according to Mr. Lastly. indeed. one has risen higher in the scale of civilisation than the other. or completely lost. and this apparently implies increased brain power.flight had once been acquired. but in this case it would be necessary that they should first have acquired the power of running quickly on the ground. of two races of savages. offer an excellent instance of a resemblance of this kind. but to this latter point I shall hereafter recur. if not impossible. into the less efficient power of gliding through the air. but extends to form. The resemblance is often wonderfully close. Bats. but as they are conjectural. Mivart remarks. such as green or decayed leaves. with each step beneficial. to see how such indefinite oscillations of infinitesimal beginnings can ever build up a sufficiently appreciable resemblance to a leaf. why. On this head. it would never be reconverted back. dead twigs. and for such a change a bat seems singularly ill-fitted. and even to the manner in which the insects hold themselves. and that there is nothing strange in a transition not having occurred in any particular case.

or those common to all the species. we can understand how it is that the larger and higher animals do not (with the exception." So close was this resemblance. through mere fluctuating variability. of a walking-stick insect (Ceroxylus laceratus). there is nothing improbable in this insect having varied in the irregularities on its surface. Within the main row there are some subsidiary rows. which clothe the whole gigantic palate.found in the stations frequented by them. of one fish) resemble for the sake of protection special objects. Wallace. then all the variations which rendered the insect at all more like any such object. which resembles "a stick grown over by a creeping moss or jungermannia. on each side of the upper jaw. The extremities and inner margins of all the plates are frayed into stiff bristles. would be preserved. Assuming that an insect originally happened to resemble in some degree a dead twig or a decayed leaf. one of its greatest peculiarities. and in these having become more or less green-coloured. or. of about 300 plates or laminae. Mivart's objection. and thus favoured its escape. which stand close together transversely to the longer axis of the mouth. or whalebone. independently of natural selection. as far as I know. Nor is this at all improbable. while other variations would be neglected and ultimately lost. but as the case stands there is none. that a native Dyak maintained that the foliaceous excrescences were really moss." as in the case given by Mr. and that it varied slightly in many ways. for in every group the characters which differ in the several species are the most apt to vary. would tend towards its preservation. are the most constant. and this chiefly in colour. and the more perfect the resemblance so much the better for the insect. but only the surface which commonly surrounds them. considering the almost infinite number of surrounding objects and the diversity in form and colour of the hosts of insects which exist. and serve to strain or sift 218 . The baleen consists of a row. The Greenland whale is one of the most wonderful animals in the world. they would be eliminated. Mivart's difficulty with respect to "the last touches of perfection in the mimicry. if they rendered the insect at all less like the imitated object. and every grade in resemblance which aided an insect to escape notice or detection. There would indeed be force in Mr. if we were to attempt to account for the above resemblances. while the generic characters. Nor can I see any force in Mr. As some rude resemblance is necessary for the first start. and the baleen. Insects are preyed on by birds and other enemies whose sight is probably sharper than ours. Considering the nature of the differences between the species in the group which includes the above Ceroxylus.

obliquely bevelled so as to be pointed. but in the different species of Cetaceans there are gradations in length. The entire head of the shoveller. though incomparably less bulky. 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. and placed transversely to the longer axis of the mouth. so that if we were to make the head of the shoveller as long as that of the Balaenoptera. as they project inward. and they project fourteen onehundredths of an inch beneath the edge.the water. two-thirds of the length of the baleen in this species of whale. according to Scoresby. each of service to its possessor. 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 upper mandible is furnished on each side (in the specimen examined by me) with a row or comb formed of 188 thin. The lower mandible of the shoveller-duck is furnished with 219 . subsist by sifting the mud and water. or even fifteen feet in length. then its preservation and augmentation within serviceable limits would be promoted by natural selection alone. With respect to the baleen. it may be asked. 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. The quality of the whalebone also differs in the different species. In these several respects they resemble the plates of baleen in the mouth of a whale. But how to obtain the beginning of such useful development?" In answer. Mivart remarks that if it "had once attained such a size and development as to be at all useful. instead of straight downward. the middle lamina being in one species. They arise from the palate. At their bases there is a short subsidiary row of obliquely transverse lamellae. Those standing towards the middle are the longest. The middle and longest lamina in the Greenland whale is ten. four feet. being about one-third of an inch in length. and in the Balaenoptera rostrata only about nine inches in length. in another three. elastic lamellae. or sifters. and the family has sometimes been called Criblatores. and thus to secure the minute prey on which these great animals subsist. is about one-eighteenth of the length of the head of a moderately large Balaenoptera rostrata. in another eighteen inches. But towards the extremity of the beak they differ much. that is. the lamellae would be six inches in length. Mr. and that the immense plates of baleen in the Greenland whale might have been developed from such lamellae by finely graduated steps. twelve. like whales. I wish only to show that this is not incredible. in which species the baleen is only nine inches long.

almost confluent. to the beak of the common duck. but the lamellae are not so numerous. as if for crushing food. in which the lamellae are less developed than in the common goose.lamellae of equal length with these above. "uses its bill like a duck by throwing the water out at the corners." Its chief food. as every one knows. but uses its beak exclusively for tearing or cutting herbage. so that the beak of this bird resembles in this respect the mouth of a whale. and in being thus furnished it differs conspicuously from the lower jaw of a whale. as I am informed by Mr. yet this goose. the upper mandible alone is furnished with lamellae. and are firmly attached to the sides of the mandible. and are edged with translucent. yet this bird. The edges of the lower mandible are serrated with teeth much more prominent. Turning to another group of the same family. 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. Although the beak is thus very inferior as a sifter to that of a shoveller. The edges of the lower mandible are crossed by numerous fine ridges. There are other species. which it crops like the common goose. so that they thus curiously resemble the plates of baleen. Bartlett. without any great break. a member of the distinct family of the Petrels. E. which project very little. as I hear from Mr. The palate is also covered with hard rounded knobs. however. as I hear from Mr. nor do they project so much inward. Bartlett. In this latter bird the lamellae of the upper mandible are much coarser than in the common duck. They are square-topped. constantly uses it for this purpose. but finer. The common goose does not sift the water. which are well developed and project beneath the margin. There are other species of geese. the extremities of these lower lamellae are frayed into fine bristly points. in which the lamellae are considerably less developed than in the common duck. is grass. which is destitute of baleen. and terminating upward in teeth-like knobs. On the other hand. In this latter species the lamellae are much coarser than in the shoveller. and in some respects through that of the Aix sponsa. Salvin). through the beak of the Merganetta armata. In the genus Prion. for which purpose it is so well fitted that it can crop grass closer than almost any other animal. coarser and sharper than in the duck. about twentyseven in number on each side. they are only about fifty in number on each side. as far as fitness for sifting is concerned. hardish tissue. 220 . In the Egyptian goose (Chenalopex) the beak closely resembles that of the common duck. nor so distinct from each other. but I do not know whether they use their beaks for sifting the water. and do not project at all beneath the margin. Salvin.

which may be observed in still-existing Cetaceans. less developed 221 . in which case they would have been used both for seizing objects and for sifting the water. in which the lamellae would be two-thirds of the length of the plates of baleen in the Balaenoptera rostrata.We thus see that a member of the duck family. and that the structure of every part of its frame must be well adapted to its conditions of life. but its palate is roughened. There is. like the knobs on the beak of the goose. 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 goose. The lower. might also be converted by small changes into one provided with prominent. it is of a white colour. We should bear in mind that each species of duck is subjected to a severe struggle for existence. and so onward. lead us onward to the enormous plates of baleen in the Greenland whale. according to Lacepede. Returning to the whales. in which case they would have served exclusively as a sifting apparatus. but rather more regularly placed. but in some on the right side. as I may add. lastly. and occasionally reversed adult specimens occur. From this stage. hard points of horn. are remarkable for their asymmetrical bodies. or resting-surface. resembles at first sight the ventral surface of an ordinary fish. The beak of a goose. with a beak constructed like that of a common goose and adapted solely for grazing. They rest on one side—in the greater number of species on the left. with the functions of the parts slowly changing during the progress of development. therefore. aided it in seizing or tearing its food. then into lamellae like those of the domestic duck. or Flat-fish. into one like the shoveller. Nor is there the least reason to doubt that each step in this scale might have been as serviceable to certain ancient Cetaceans. might be converted by small changes into a species like the Egyptian goose—this into one like the common duck—and. If so. and which. nothing improbable in supposing that some early Cetacean form was provided with similar points of horn on the palate. with small unequal. The Pleuronectidae. for seizing or tearing solid food. as are the gradations in the beaks of the different existing members of the duck-family. provided with a beak almost exclusively adapted for sifting the water. The Hyperoodon bidens is destitute of true teeth in an efficient condition. or even a member with a beak having less well-developed lamellae. except the hooked tip. serving for the widely different purpose of securing live fish. until they became as well constructed as those of the shoveller. for this bird could hardly use any part of its beak. recurved teeth. gradations. like those of the Merganser (a member of the same family).

being extremely common. such as soles. The forehead between the eyes consequently becomes. The Pleuronectidae. with both sides equally coloured. It is obvious that unless the lower eye did thus travel round. 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. which are entirely thrown to one side. however. cannot long retain a vertical position." But he might have found an answer to this objection in the excellent observations published in 1867 by Malm. also. far from clear. with their eyes standing on opposite sides of the head. flounders. It seems. so that it readily yields to muscular action. and the whole body is then symmetrical. etc. During early youth. and facility for feeding on the ground. On one occasion Malm saw a young fish raise and depress the lower eye through an angular distance of about seventy degrees.in many ways than the upper side. We should remember that the skull at this early age is cartilaginous and flexible. it could not be used by the fish while lying in its habitual position on one side. owing to the excessive depth of their bodies. is manifest from several species. they stand opposite to each other. which does not in any considerable degree alter the shape in which it leaves the ovum. even. soon growing tired. Mivart has taken up this case. While thus at rest they often twist. The lower eye would. It is also known 222 . they fall to the bottom on one side. while very young and still symmetrical. as Malm observed. to see above them. The different members. that such an incipient transformation must rather have been injurious. the lower eye upward.. "a long series of forms exhibiting a gradual transition from Hippoglossus pinguis. Soon the eye proper to the lower side begins to glide slowly round the head to the upper side. and they do this so vigorously that the eye is pressed hard against the upper part of the orbit. for they are both placed on the upper side of the head. and remarks that a sudden spontaneous transformation in the position of the eyes is hardly conceivable. the small size of their lateral fins. and to their being destitute of a swim-bladder. indeed. as could be plainly seen. The chief advantages thus gained seem to be protection from their enemies. to the soles. as was formerly thought to be the case. however. with the lateral fins often of smaller size. Hence. temporarily contracted in breadth. That the Pleuronectidae are admirably adapted by their flattened and asymmetrical structure for their habits of life. but does not pass right through the skull. He then adds: "If the transit was gradual." Mr. in which I quite agree with him. as Schiodte remarks. But the eyes offer the most remarkable peculiarity. of the family present. have been liable to be abraded by the sandy bottom.

and if this be so. and no permanent effect is thus produced. with the jaw bones stronger and more effective on this. of feeding with ease on the ground. on the other hand. that the Pleuronectidae are not quite symmetrical even in the embryo. and he has observed that they often then strain their lower eyes so as to look upward. and other species on the right side. that the skull yields and is altered in shape. salmon. Mivart considers would be injurious. These fishes. are soon able to hold themselves in a vertical position. may be attributed to the habit. With the Pleuronectidae. no doubt beneficial to the individual and to the species. the eyeless side of the head. by remarking that "the author gives a very simple explanation of the abnormal condition of the Pleuronectoids. 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. in opposition to some other naturalists. Dr. and swims diagonally through the water. than on the other.with the higher animals. and in this fish. Disuse. for the sake. habitually fall over and rest on the left side. Traquair supposes." We thus see that the first stages of the transit of the eye from one side of the head to the other. Malm adds. the older they grow the more habitually they rest on one side. rests on its left side at the bottom. and several other symmetrical fishes. the tendency to distortion would no doubt be increased through the principle of inheritance. which is not a member of the Pleuronectidae. if one ear flops forward and downward. which Mr. Judging from analogy. the two sides of the head are said to be somewhat dissimilar. concludes his abstract of Malm's paper. Malm states that the newly-hatched young of perches. we could understand how it is that certain species. even after early youth. including the lateral fins. and on the position of the eyes. that the adult Trachypterus arcticus. Our great authority on Fishes. and their skulls are thus rendered rather crooked. owing to the increasing flatness of their bodies. With long-eared rabbits. as "there is so much less room for their action than with the larger fins 223 . of endeavouring to look upward with both eyes. though Yarrel thinks that the reduced size of these fins is advantageous to the fish. as Dr. while young. and a permanent effect is thus produced on the form of the head. Schiodte believes. in confirmation of the above view. on the other hand. of which I have given a figure. Gunther. its weight drags forward all the bones of the skull on the same side. have the habit of occasionally resting on one side at the bottom. if the skin or muscles be permanently contracted through disease or some accident. will account for the less developed condition of the whole inferior half of the body. while resting on one side at the bottom. however.

and serves as a fifth hand. 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. of changing their colour in accordance with the surrounding surface. are due to the action of the light. and this almost implies that some benefit great or small is thus derived." But there is no necessity for any such belief. will be strengthened by natural selection. we may reasonably suppose that the absence of colour in flatfish on the side. as recently shown by Pouchet. would in all probability suffice for the work. and perhaps of their disuse. I have received an analogous account from Dr. which is under. Mivart in every detail. A reviewer. How much to attribute in each particular case to the effects of use. Gunther. or favour their chance of having and of rearing offspring. so like the sandy bed of the sea. But it cannot be supposed that the peculiar speckled appearance of the upper side of the sole. to their habits of life. and many other peculiarities. The extremity of the tail in some American monkeys has been converted into a wonderfully perfect prehensile organ. whether it be the right or left. If the harvest mouse had been more strictly arboreal. but he frequently observed that they curled their tails round the branches of a bush placed in the cage. I may give another instance of a structure which apparently owes its origin exclusively to use or habit. as well as in adapting the general shape of the body of these fishes. For all spontaneous variations in the right direction will thus be preserved. Professor Henslow kept in confinement some harvest mice (Mus messorius) which do not possess a structurally prehensive tail. that the inherited effects of the increased use of parts. as will those individuals which inherit in the highest degree the effects of the increased and beneficial use of any part. and at the same time they hooked their little tails round that of their mother. may likewise be accounted for by disuse. From the colourless state of the ventral surface of most fishes and of many other animals. it would 224 . who has seen a mouse thus suspend itself. who agrees with Mr. We should keep in mind. Here natural selection has probably come into play. Habit.above. and how much to natural selection. or the power in some species.most. or the presence of bony tubercles on the upper side of the turbot. to twenty-five to thirty in the lower halves. as I have before insisted. and thus aided themselves in climbing." Perhaps the lesser number of teeth in the proportion of four to seven in the upper halves of the two jaws of the plaice. is due to the exclusion of light. Brehm saw the young of an African monkey (Cercopithecus) clinging to the under surface of their mother by their hands. it seems impossible to decide.

I will not pretend to decide. 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. therefore. the effects of use. whether in part through compensation of growth. have been developed at an extremely remote period. Through what agency the glands over a certain space became more highly specialised than the others. In the case of the fish (Hippocampus) the eggs are hatched. unless 225 . the mammary glands will have been at first developed within the marsupial sack. The development of the mammary glands would have been of no service. would have been improved or rendered more effective. than as a prehensile organ. It accords with the widely extended principle of specialisation. as is the case with some members of the same order. in some degree or manner the most nutritious. than would the individuals which secreted a poorer fluid. The mammary glands are common to the whole class of mammals. Now. at the base of the mammalian series. It is. considering its habits while young. so as to partake of the nature of milk. Mr. 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 could not have been affected through natural selection. which are the homologues of the mammary glands. that the glands over a certain space of the sack should have become more highly developed than the remainder. and we can know nothing positively about their manner of development. almost before they deserved to be thus designated. as we see in the Ornithorhyncus. and are indispensable for their existence. and if so. however. has not become thus provided. or of natural selection. with the early progenitors of mammals. that they are nourished by a secretion from the cutaneous glands of the sack. and the young are reared for a time. the individuals which secreted a fluid. and they would then have formed a breast. what chance was there of the perpetuation of such a variation?" But the case is not here put fairly. is it not at least possible that the young might have been similarly nourished? And in this case. Why Cercopithecus. and an American naturalist. and thus the cutaneous glands. within a sack of this nature. Mr. they must. It is admitted by most evolutionists that mammals are descended from a marsupial form. Lockwood. it would be difficult to say.perhaps have had its tail rendered structurally prehensile. but at first without a nipple. believes from what he has seen of the development of the young. would in the long run have reared a larger number of well-nourished offspring.

who has the power of injecting milk into the mouth of her helpless. With respect to these organs. and is thus enabled to give free entrance to the air for the lungs. 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. 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. On this head Mr. "this at least perfectly innocent and harmless structure?" It may be suggested in answer that the voice. asks: "What would be the utility of the FIRST RUDIMENTARY BEGINNINGS of such structures.) are furnished with remarkable organs. In such cases the most probable solution seems to be.the young at the same time were able to partake of the secretion. which consist. or how a few hours after leaving the shell they have learned to pick up grains of food. and afterwards transmitted to the offspring at an earlier age. We will now turn for a short space to the lower divisions of the animal kingdom. than in understanding how unhatched chickens have learned to break the egg-shell by tapping against it with their specially adapted beaks. could hardly have been used with full force as long as the larynx entered the nasal passage. "not 226 . of a tridactyle forceps—that is. sea-urchins. and Professor Flower has suggested to me that this structure would have greatly interfered with an animal swallowing solid food. Mivart. The larynx is so elongated that it rises up into the posterior end of the nasal passage. Mr. and one of these apparently is defence. half-formed offspring. while the milk passes harmlessly on each side of this elongated larynx. moved by muscles. that the habit was at first acquired by practice at a more advanced age. they subserve other functions. and how could such insipient buddings have ever preserved the life of a single Echinus?" He adds. But there is no doubt that besides removing dirt of all kinds. as on so many previous occasions." Mr. which is certainly of high importance to many animals. The Echinodermata (star-fishes. called pedicellariae. neatly fitting together and placed on the summit of a flexible stem. Mivart then asks how did natural selection remove in the adult kangaroo (and in most other mammals. the young one must infallibly be choked by the intrusion of the milk into the wind-pipe. But there IS a special provision. of one formed of three serrated arms. But the young kangaroo is said not to suck. etc. These forceps can seize firmly hold of any object. on the assumption that they are descended from a marsupial form). Mivart remarks: "Did no special provision exist. when well developed. and so safely attains the gullet behind it.

even the SUDDEN development of the snapping action would have been beneficial without the freely movable stalk, nor could the latter have been efficient without the snapping jaws, yet no minute, nearly indefinite variations could simultaneously evolve these complex co-ordinations of structure; to deny this seems to do no less than to affirm a startling paradox." Paradoxical as this may appear to Mr. Mivart, tridactyle forcepses, immovably fixed at the base, but capable of a snapping action, certainly exist on some star- fishes; and this is intelligible if they serve, at least in part, as a means of defence. Mr. Agassiz, to whose great kindness I am indebted for much information on the subject, informs me that there are other star- fishes, in which one of the three arms of the forceps is reduced to a support for the other two; and again, other genera in which the third arm is completely lost. In Echinoneus, the shell is described by M. Perrier as bearing two kinds of pedicellariae, one resembling those of Echinus, and the other those of Spatangus; and such cases are always interesting as affording the means of apparently sudden transitions, through the abortion of one of the two states of an organ. With respect to the steps by which these curious organs have been evolved, Mr. Agassiz infers from his own researches and those of Mr. Muller, that both in star-fishes and sea-urchins the pedicellariae must undoubtedly be looked at as modified spines. This may be inferred from their manner of development in the individual, as well as from a long and perfect series of gradations in different species and genera, from simple granules to ordinary spines, to perfect tridactyle pedicellariae. The gradation extends even to the manner in which ordinary spines and the pedicellariae, with their supporting calcareous rods, are articulated to the shell. In certain genera of star-fishes, "the very combinations needed to show that the pedicellariae are only modified branching spines" may be found. Thus we have fixed spines, with three equi-distant, serrated, movable branches, articulated to near their bases; and higher up, on the same spine, three other movable branches. Now when the latter arise from the summit of a spine they form, in fact, a rude tridactyle pedicellariae, and such may be seen on the same spine together with the three lower branches. In this case the identity in nature between the arms of the pedicellariae and the movable branches of a spine, is unmistakable. It is generally admitted that the ordinary spines serve as a protection; and if so, there can be no reason to doubt that those furnished with serrated and movable branches likewise serve for the same purpose; and they would thus serve still more effectively as soon as by meeting together they acted as a prehensile or snapping apparatus. Thus

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every gradation, from an ordinary fixed spine to a fixed pedicellariae, would be of service. In certain genera of star-fishes these organs, instead of being fixed or borne on an immovable support, are placed on the summit of a flexible and muscular, though short, stem; and in this case they probably subserve some additional function besides defence. In the sea-urchins the steps can be followed by which a fixed spine becomes articulated to the shell, and is thus rendered movable. I wish I had space here to give a fuller abstract of Mr. Agassiz's interesting observations on the development of the pedicellariae. All possible gradations, as he adds, may likewise be found between the pedicellariae of the star-fishes and the hooks of the Ophiurians, another group of the Echinodermata; and again between the pedicellariae of sea-urchins and the anchors of the Holothuriae, also belonging to the same great class. Certain compound animals, or zoophytes, as they have been termed, namely the Polyzoa, are provided with curious organs called avicularia. These differ much in structure in the different species. In their most perfect condition they curiously resemble the head and beak of a vulture in miniature, seated on a neck and capable of movement, as is likewise the lower jaw or mandible. In one species observed by me, all the avicularia on the same branch often moved simultaneously backwards and forwards, with the lower jaw widely open, through an angle of about 90 degrees, in the course of five seconds; and their movement caused the whole polyzoary to tremble. When the jaws are touched with a needle they seize it so firmly that the branch can thus be shaken. Mr. Mivart adduces this case, chiefly on account of the supposed difficulty of organs, namely the avicularia of the Polyzoa and the pedicellariae of the Echinodermata, which he considers as "essentially similar," having been developed through natural selection in widely distinct divisions of the animal kingdom. But, as far as structure is concerned, I can see no similarity between tridactyle pedicellariae and avicularia. The latter resembles somewhat more closely the chelae or pincers of Crustaceans; and Mr. Mivart might have adduced with equal appropriateness this resemblance as a special difficulty, or even their resemblance to the head and beak of a bird. The avicularia are believed by Mr. Busk, Dr. Smitt and Dr. Nitsche—naturalists who have carefully studied this group—to be homologous with the zooids and their cells which compose the zoophyte, the movable lip or lid of the cell corresponding with the lower and movable mandible of the avicularium. Mr. Busk, however, does not know of any gradations now existing between a zooid and an

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avicularium. It is therefore impossible to conjecture by what serviceable gradations the one could have been converted into the other, but it by no means follows from this that such gradations have not existed. As the chelae of Crustaceans resemble in some degree the avicularia of Polyzoa, both serving as pincers, it may be worth while to show that with the former a long series of serviceable gradations still exists. In the first and simplest stage, the terminal segment of a limb shuts down either on the square summit of the broad penultimate segment, or against one whole side, and is thus enabled to catch hold of an object, but the limb still serves as an organ of locomotion. We next find one corner of the broad penultimate segment slightly prominent, sometimes furnished with irregular teeth, and against these the terminal segment shuts down. By an increase in the size of this projection, with its shape, as well as that of the terminal segment, slightly modified and improved, the pincers are rendered more and more perfect, until we have at last an instrument as efficient as the chelae of a lobster. And all these gradations can be actually traced. Besides the avicularia, the polyzoa possess curious organs called vibracula. These generally consist of long bristles, capable of movement and easily excited. In one species examined by me the vibracula were slightly curved and serrated along the outer margin, and all of them on the same polyzoary often moved simultaneously; so that, acting like long oars, they swept a branch rapidly across the object-glass of my microscope. When a branch was placed on its face, the vibracula became entangled, and they made violent efforts to free themselves. They are supposed to serve as a defence, and may be seen, as Mr. Busk remarks, "to sweep slowly and carefully over the surface of the polyzoary, removing what might be noxious to the delicate inhabitants of the cells when their tentacula are protruded." The avicularia, like the vibracula, probably serve for defence, but they also catch and kill small living animals, which, it is believed, are afterwards swept by the currents within reach of the tentacula of the zooids. Some species are provided with avicularia and vibracula, some with avicularia alone and a few with vibracula alone. It is not easy to imagine two objects more widely different in appearance than a bristle or vibraculum, and an avicularium like the head of a bird; yet they are almost certainly homologous and have been developed from the same common source, namely a zooid with its cell. Hence, we can understand how it is that these organs graduate in some cases, as I am informed by Mr. Busk, into each other. Thus, with the avicularia of

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several species of Lepralia, 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 vibracula may have been directly developed from the lips of the cells, without having passed through the avicularian stage; but it seems more probable that they have passed through this stage, as during the early stages of the transformation, the other parts of the cell, with the included zooid, could hardly have disappeared at once. In many cases the vibracula have a grooved support at the base, which seems to represent the fixed beak; though this support in some species is quite absent. This view of the development of the vibracula, if trustworthy, is interesting; for supposing that all the species provided with avicularia had become extinct, no one with the most vivid imagination would ever have thought that the vibracula had originally existed as part of an organ, resembling a bird's head, or an irregular box or hood. It is interesting to see two such widely different organs developed from a common origin; and as the movable lip of the cell serves as a protection to the zooid, there is no difficulty in believing that all the gradations, by which the lip became converted first into the lower mandible of an avicularium, and then into an elongated bristle, likewise served as a protection in different ways and under different circumstances. In the vegetable kingdom Mr. Mivart only alludes to two cases, namely the structure of the flowers of orchids, and the movements of climbing plants. With respect to the former, he says: "The explanation of their ORIGIN is deemed thoroughly unsatisfactory—utterly insufficient to explain the incipient, infinitesimal beginnings of structures which are of utility only when they are considerably developed." As I have fully treated this subject in another work, I will here give only a few details on one alone of the most striking peculiarities of the flowers of orchids, namely, their pollinia. A pollinium, when highly developed, consists of a mass of pollen-grains, affixed to an elastic foot-stalk or caudicle, and this to a little mass of extremely viscid matter. The pollinia are by this means transported by insects from one flower to the stigma of another. In some orchids there is no caudicle to the pollen-masses, and the grains are merely tied together by fine threads; but as these are not confined to orchids, they need not here be considered; yet I may mention that at the base of the orchidaceous series, in Cypripedium, we can see how the threads were probably first developed. In other orchids the threads cohere at one end of the pollen-masses; and this forms the first or nascent trace of a caudicle. That this is the origin of the caudicle, even when of

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considerable length and highly developed, we have good evidence in the aborted pollen-grains which can sometimes be detected embedded within the central and solid parts. With respect to the second chief peculiarity, namely, the little mass of viscid matter attached to the end of the caudicle, a long series of gradations can be specified, each of plain service to the plant. In most flowers belonging to other orders the stigma secretes a little viscid matter. Now, in certain orchids similar viscid matter is secreted, but in much larger quantities by one alone of the three stigmas; and this stigma, perhaps in consequence of the copious secretion, is rendered sterile. When an insect visits a flower of this kind, it rubs off some of the viscid matter, and thus at the same time drags away some of the pollen-grains. From this simple condition, which differs but little from that of a multitude of common flowers, there are endless gradations—to species in which the pollenmass terminates in a very short, free caudicle—to others in which the caudicle becomes firmly attached to the viscid matter, with the sterile stigma itself much modified. In this latter case we have a pollinium in its most highly developed and perfect condition. 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, with the stigma differing but little from that of the ordinary flowers, to a highly complex pollinium, admirably adapted for transportal by insects; 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. In this, and in almost every other case, the enquiry may be pushed further backwards; and it may be asked how did the stigma of an ordinary flower become viscid, but as we do not know the full history of any one group of beings, it is as useless to ask, as it is hopeless to attempt answering, such questions. We will now turn to climbing plants. These can be arranged in a long series, from those which simply twine round a support, to those which I have called leaf-climbers, and to those provided with tendrils. In these two latter classes the stems have generally, but not always, lost the power of twining, though they retain the power of revolving, which the tendrils likewise possess. The gradations from leaf-climbers to tendril bearers are wonderfully close, and certain plants may be differently placed in either class. But in ascending the series from simple twiners to leaf-climbers, an important quality is added, namely sensitiveness to a touch, by which means the foot-stalks of the leaves or flowers, or these modified and converted into tendrils, are excited to bend round and

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clasp the touching object. He who will read my memoir on these plants will, I think, 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. For instance, it is clearly a great advantage to a twining plant to become a leaf-climber; and it is probable that every twiner which possessed leaves with long foot-stalks would have been developed into a leaf-climber, if the foot-stalks had possessed in any slight degree the requisite sensitiveness to a touch. As twining is the simplest means of ascending a support, and forms the basis of our series, it may naturally be asked how did plants acquire this power in an incipient degree, afterwards to be improved and increased through natural selection. The power of twining depends, firstly, on the stems while young being extremely flexible (but this is a character common to many plants which are not climbers); and, secondly, on their continually bending to all points of the compass, one after the other in succession, in the same order. By this movement the stems are inclined to all sides, and are made to move round and round. As soon as the lower part of a stem strikes against any object and is stopped, the upper part still goes on bending and revolving, and thus necessarily twines round and up the support. The revolving movement ceases after the early growth of each shoot. As in many widely separated families of plants, single species and single genera possess the power of revolving, and have thus become twiners, they must have independently acquired it, and cannot have inherited it from a common progenitor. Hence, 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 not climb; and that this had afforded the basis for natural selection to work on and improve. When I made this prediction, I knew of only one imperfect case, namely, 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. 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, though irregularly, and he states that he has reason to suspect that this occurs with some other plants. These slight movements appear to be of no service to the plants in question; anyhow, they are not of the least use in the way of climbing, which is the point that concerns us. Nevertheless we can see that if the stems of these plants had been flexible, and if under the conditions to which they are exposed it had profited them to ascend to a height, then the habit of slightly and

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irregularly revolving might have been increased and utilised through natural selection, until they had become converted into well-developed twining species. With respect to the sensitiveness of the foot-stalks of the leaves and flowers, and of tendrils, nearly the same remarks are applicable as in the case of the revolving movements of twining plants. As a vast number of species, belonging to widely distinct groups, are endowed with this kind of sensitiveness, it ought to be found in a nascent condition in many plants which have not become climbers. 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. Morren found in several species of Oxalis that the leaves and their foot-stalks moved, especially after exposure to a hot sun, when they were gently and repeatedly touched, or when the plant was shaken. I repeated these observations on some other species of Oxalis with the same result; in some of them the movement was distinct, but was best seen in the young leaves; in others it was extremely slight. It is a more important fact that according to the high authority of Hofmeister, the young shoots and leaves of all plants move after being shaken; and with climbing plants it is, as we know, only during the early stages of growth that the foot-stalks and tendrils are sensitive. It is scarcely possible that the above slight movements, due to a touch or shake, in the young and growing organs of plants, can be of any functional importance to them. But plants possess, in obedience to various stimuli, powers of movement, which are of manifest importance to them; for instance, towards and more rarely from the light—in opposition to, and more rarely in the direction of, the attraction of gravity. When the nerves and muscles of an animal are excited by galvanism or by the absorption of strychnine, the consequent movements may be called an incidental result, for the nerves and muscles have not been rendered specially sensitive to these stimuli. So with plants it appears that, from having the power of movement in obedience to certain stimuli, they are excited in an incidental manner by a touch, or by being shaken. Hence there is no great difficulty in admitting that in the case of leaf-climbers and tendril-bearers, it is this tendency which has been taken advantage of and increased through natural selection. It is, however, probable, from reasons which I have assigned in my memoir, that this will have occurred only with plants which had already acquired the power of revolving, and had thus become twiners.

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I have already endeavoured to explain how plants became twiners, namely, by the increase of a tendency to slight and irregular revolving movements, which were at first of no use to them; this movement, as well as that due to a touch or shake, being the incidental result of the power of moving, gained for other and beneficial purposes. Whether, during the gradual development of climbing plants, natural selection has been aided by the inherited effects of use, I will not pretend to decide; but we know that certain periodical movements, for instance the socalled sleep of plants, are governed by habit. I have now considered enough, perhaps more than enough, of the cases, selected with care by a skilful naturalist, to prove that natural selection is incompetent to account for the incipient stages of useful structures; and I have shown, as I hope, that there is no great difficulty on this head. A good opportunity has thus been afforded for enlarging a little on gradations of structure, often associated with strange functions—an important subject, which was not treated at sufficient length in the former editions of this work. I will now briefly recapitulate the foregoing cases. With the giraffe, the continued preservation of the individuals of some extinct high-reaching ruminant, which had the longest necks, legs, etc., and could browse a little above the average height, and the continued destruction of those which could not browse so high, would have sufficed for the production of this remarkable quadruped; but the prolonged use of all the parts, together with inheritance, will have aided in an important manner in their co-ordination. With the many insects which imitate various objects, 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, since perfected through the occasional preservation of slight variations which made the resemblance at all closer; and this will have been carried on as long as the insect continued to vary, and as long as a more and more perfect resemblance led to its escape from sharp-sighted enemies. In certain species of whales there is a tendency to the formation of irregular little points of horn on the palate; and it seems to be quite within the scope of natural selection to preserve all favourable variations, until the points were converted, first into lamellated knobs or teeth, like those on the beak of a goose—then into short lamellae, like those of the domestic ducks—and then into lamellae, as perfect as those of the shoveller-duck—and finally into the gigantic plates of baleen, as in the mouth of the Greenland whale. In the family of the ducks, the lamellae are first used as teeth,

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then partly as teeth and partly as a sifting apparatus, and at last almost exclusively for this latter purpose. With such structures as the above lamellae of horn or whalebone, habit or use can have done little or nothing, as far as we can judge, towards their development. On the other hand, the transportal of the lower eye of a flat-fish to the upper side of the head, and the formation of a prehensile tail, may be attributed almost wholly to continued use, together with inheritance. With respect to the mammae of the higher animals, the most probable conjecture is that primordially the cutaneous glands over the whole surface of a marsupial sack secreted a nutritious fluid; and that these glands were improved in function through natural selection, and concentrated into a confined area, in which case they would have formed a mamma. There is no more difficulty in understanding how the branched spines of some ancient Echinoderm, which served as a defence, became developed through natural selection into tridactyle pedicellariae, than in understanding the development of the pincers of crustaceans, through slight, serviceable modifications in the ultimate and penultimate segments of a limb, which was at first used solely for locomotion. In the avicularia and vibracula of the Polyzoa we have organs widely different in appearance developed from the same source; and with the vibracula we can understand how the successive gradations might have been of service. With the pollinia of orchids, the threads which originally served to tie together the pollen-grains, can be traced cohering into caudicles; and the steps can likewise be followed by which viscid matter, such as that secreted by the stigmas of ordinary flowers, and still subserving nearly but not quite the same purpose, became attached to the free ends of the caudicles—all these gradations being of manifest benefit to the plants in question. With respect to climbing plants, I need not repeat what has been so lately said. It has often been asked, if natural selection be so potent, why has not this or that structure been gained by certain species, to which it would apparently have been advantageous? But it is unreasonable to expect a precise answer to such questions, considering our ignorance of the past history of each species, and of the conditions which at the present day determine its numbers and range. In most cases only general reasons, but in some few cases special reasons, can be assigned. Thus to adapt a species to new habits of life, many co-ordinated modifications are almost indispensable, and it may often have happened that the requisite parts did not vary in the right manner or to the right degree. Many species must have been prevented from increasing in numbers through

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destructive agencies, which stood in no relation to certain structures, which we imagine would have been gained through natural selection from appearing to us advantageous to the species. In this case, as the struggle for life did not depend on such structures, they could not have been acquired through natural selection. In many cases complex and long-enduring conditions, often of a peculiar nature, are necessary for the development of a structure; and the requisite conditions may seldom have concurred. The belief that any given structure, which we think, often erroneously, would have been beneficial to a species, would have been gained under all circumstances through natural selection, is opposed to what we can understand of its manner of action. Mr. Mivart does not deny that natural selection has effected something; but he considers it as "demonstrably insufficient" to account for the phenomena which I explain by its agency. His chief arguments have now been considered, and the others will hereafter be considered. They seem to me to partake little of the character of demonstration, and to have little weight in comparison with those in favour of the power of natural selection, aided by the other agencies often specified. I am bound to add, that some of the facts and arguments here used by me, have been advanced for the same purpose in an able article lately published in the "Medico-Chirurgical Review." At the present day almost all naturalists admit evolution under some form. Mr. Mivart believes that species change through "an internal force or tendency," about which it is not pretended that anything is known. That species have a capacity for change will be admitted by all evolutionists; but there is no need, as it seems to me, to invoke any internal force beyond the tendency to ordinary variability, which through the aid of selection, by man has given rise to many well-adapted domestic races, and which, through the aid of natural selection, would equally well give rise by graduated steps to natural races or species. The final result will generally have been, as already explained, an advance, but in some few cases a retrogression, in organisation. Mr. Mivart is further inclined to believe, and some naturalists agree with him, that new species manifest themselves "with suddenness and by modifications appearing at once." For instance, he supposes that the differences between the extinct three-toed Hipparion and the horse arose suddenly. He thinks it difficult to believe that the wing of a bird "was developed in any other way than by a comparatively sudden modification of a marked and important kind;" and apparently he would extend the same view to the wings of bats and pterodactyles. This conclusion,

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etc. as are known occasionally to arise under domestication. is avoided on the theory of gradual evolution. A still greater number must be called monstrosities. According to our experience. and the characters which thus reappear were. If such occurred under nature. Hence. that several wonderfully changed individuals appeared simultaneously within the same district. This difficulty.. That many species have been evolved in an extremely gradual manner. abrupt and strongly marked variations occur in our domesticated productions. 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 for entirely disbelieving that they have changed in the wonderful manner indicated by Mr. from lowland to upland. etc. Mivart. Mivart. are as follows. as in the case of unconscious selection by man. if found in a state of nature. On every continent. But as species are more variable when domesticated or cultivated than under their natural conditions. we meet with a host of 237 . through the preservation of a large number of individuals. in proceeding from north to south. and so it is known to be under domestication. they would be liable. 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. in order that a new species should suddenly appear in the manner supposed by Mr. it is not probable that such great and abrupt variations have often occurred under nature. or even under domestication. in opposition to all analogy. Excluding such cases of abrupt variations. Of these latter variations several may be attributed to reversion.. it is probable. in many cases at first gained in a gradual manner. My reasons for doubting whether natural species have changed as abruptly as have occasionally domestic races. such as six-fingered men. there can hardly be a doubt. it is almost necessary to believe. appears to me improbable in the highest degree. and as they are widely different in character from natural species. unless abrupt variations of this kind are specially preserved and separated by the care of man. which varied more or less in any favourable direction. Ancon sheep. to be lost by accidental causes of destruction and by subsequent intercrossing.which implies great breaks or discontinuity in the series. the few which remain would at best constitute. doubtful species. porcupine men. they throw very little light on our subject. and of the destruction of a large number which varied in an opposite manner. Niata cattle. as formerly explained. closely related to their parental types. Everyone who believes in slow and gradual evolution. singly and at rather long intervals of time.

could be added. So it is if we look to past times. like varieties round species. Nor should it be forgotten. all pointing in the same direction. such as the sudden crystallisation of inorganic substances. One class of facts. or the falling of a facetted spheroid from one facet to another. the sudden appearance of new and distinct forms of life in our geological formations supports at first sight the belief in abrupt development. as they have been shown to be by Mr. yet it may be maintained that some have been developed in a different and abrupt manner. Although very many species have almost certainly been produced by steps not greater than those separating fine varieties. connecting together widely different structures. in relation to periods remote in the history of the 238 . or have lately existed. I am compelled to allude to subjects hereafter to be discussed. Many analogous facts. namely. and compare the species which have just passed away with those still living within the same areas. and they present other analogies with varieties. and how the parts which are developed in an extraordinary degree or manner are more variable than other parts of the same species. and present a greater number of varieties than do the species in the smaller genera. that numerous and wonderfully fine gradations can be traced. or if we compare the fossil species embedded in the sub-stages of the same geological formation. and see how many of their inhabitants can be raised only to the rank of doubtful species. For instance. The former are also grouped in little clusters. hardly deserve consideration. Such an admission. Chauncey Wright. however. which we have reason to believe were formerly connected. 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. Many large groups of facts are intelligible only on the principle that species have been evolved by very small steps. as we likewise do on certain distinct continents. But the value of this evidence depends entirely on the perfection of the geological record. Look at the many outlying islands round a continent.closely related or representative species. however. But in making these and the following remarks. ought not to be made without strong evidence being assigned. The vague and in some respects false analogies. which have been advanced in favour of this view. and it will hardly be maintained that such species have been developed in an abrupt or sudden manner. the fact that the species included in the larger genera are more closely related to each other. instead of to distinct species. as was shown in our second chapter. It is indeed manifest that multitudes of species are related in the closest manner to other species that still exist.

and that they become differentiated by insensibly fine steps. for instance. To admit all this is. every detail in its structure being developed by insensibly fine steps. there is nothing strange in new forms appearing as if suddenly developed. 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. Embryological resemblances of all kinds can be accounted for. one furnished with wings. Mivart. and serves as a record of the past condition of the species. He will be forced to admit that these great and sudden transformations have left no trace of their action on the embryo. embryology enters a strong protest. The embryo is thus left almost unaffected. he will not be able to assign a shadow of an explanation. It cannot be denied that such abrupt and great changes of structure are widely different from those which most species apparently have undergone. But against the belief in such abrupt changes. as it seems to me. and yet should not bear even a trace in its embryonic condition of any sudden modification. by the progenitors of our existing species having varied after early youth. 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.world. will be almost compelled to assume. and of such complex and wonderful co-adaptations. have been suddenly produced. it is incredible that an animal should have undergone such momentous and abrupt transformations as those above indicated. He who believes that some ancient form was transformed suddenly through an internal force or tendency into. as we shall hereafter see. at a corresponding age. or the sudden conversion of a Hipparion into a horse. On this view of the meaning of embryological resemblances. and to leave those of science. and the legs of horses or other quadrupeds. to enter into the realms of miracle. Unless we admit transformations as prodigious as those advocated by Mr. in opposition to all analogy. and indeed on any view. and having transmitted their newly-acquired characters to their offspring. It is notorious that the wings of birds and bats. If the record is as fragmentary as many geologists strenuously assert. 239 . that many individuals varied simultaneously. are undistinguishable at an early embryonic period. hardly any light is thrown by the belief in abrupt modifications on the deficiency of connecting links in our geological formations. such as the sudden development of the wings of birds or bats.

240 . molothrus. their origin — Natural instincts of the cuckoo. but different in their origin — Instincts graduated — Aphides and ants — Instincts variable — Domestic instincts. and parasitic bees — Slave-making ants — Hive-bee. ostrich. 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.Chapter Instinct 8 Instincts comparable with habits.

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. fifth. without their knowing for what purpose it is performed. but every one understands what is meant. for instance. An action. the caterpillar simply re-performed the fourth. Habits easily become associated with other habits. A little dose of judgment or reason. when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds' nests. so in instincts. Huber found it was with a caterpillar. one action follows another by a sort of rhythm. is usually said to be instinctive. which we ourselves require experience to enable us to perform. Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. and sixth stages of construction. and were put into one finished up to the sixth stage.1. I think. How unconsciously many habitual actions are performed. but not necessarily of its origin. But I could show that none of these characters are universal. so that much of its 241 . as Pierre Huber expresses it. even with animals low in the scale of nature. As in repeating a well-known song. It would be easy to show that several distinct mental actions are commonly embraced by this term. to the third stage. We are concerned only with the diversities of instinct and of the other mental faculties in animals of the same class. When once acquired. and when performed by many individuals in the same way. he is generally forced to go back to recover the habitual train of thought: so P. a caterpillar were taken out of a hammock made up. This comparison gives. say. for if he took a caterpillar which had completed its hammock up to. without experience. Several other points of resemblance between instincts and habits could be pointed out. often comes into play. more especially by a very young one. Many instincts are so wonderful that their development will probably appear to the reader a difficulty sufficient to overthrow my whole theory. which makes a very complicated hammock. with certain periods of time and states of the body. if a person be interrupted in a song. however. If. they often remain constant throughout life. an accurate notion of the frame of mind under which an instinctive action is performed. the sixth stage of construction. I may here premise. and put it into a hammock completed up only to the third stage. or in repeating anything by rote. any more than I have with that of life itself. that I have nothing to do with the origin of the mental powers. when performed by an animal.

yet profitable.—that is of variations produced by the same unknown causes which produce slight deviations of bodily structure. and thus tried to complete the already finished work. we ought to find in nature. or we ought at least to be able to show that gradations of some kind are possible. those of the hive-bee and of many ants. namely. and. so I do not doubt it has been with instincts. 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. under its present conditions of life. as in the case of corporeal structures. then I can see no difficulty in natural selection preserving and continually accumulating variations of instinct to any extent that was profitable. and then transmitted by inheritance to succeeding generations. as I believe. 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. far from deriving any benefit from this. except by the slow and gradual accumulation of numerous. it is at least possible that slight modifications of instinct might be profitable to a species. It is thus. it was much embarrassed. and if it can be shown that instincts do vary ever so little. use or habit. could not possibly have been acquired by habit. be might truly be said to have done so instinctively. instead of playing the pianoforte at three years old with wonderfully little practice. But it would be a serious error to suppose that the greater number of instincts have been acquired by habit in one generation. It will be universally admitted that instincts are as important as corporeal structures for the welfare of each species. It can be clearly shown that the most wonderful instincts with which we are acquainted. Under changed conditions of life.work was already done for it. 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. slight. and this we certainly can do. in order to complete its hammock. No complex instinct can possibly be produced through natural selection. that all the most complex and wonderful instincts have originated. Hence. seemed forced to start from the third stage. I have been surprised to find. As modifications of corporeal structure arise from. variations. had played a tune with no practice at all. and are diminished or lost by disuse. where it had left off. making 242 . and are increased by.

yet each tries to take advantage of the instincts of others. and each. immediately lifted up its abdomen and excreted a limpid drop of sweet juice. with which I am acquainted. After this interval. and prevented their attendance during several hours. Changes of instinct may sometimes be facilitated by the same species having different instincts at different periods of life. I removed all the ants from a group of about a dozen aphides on a dockplant. Afterwards. the instinct of each species is good for itself. leading to the most complex instincts. as far as we can judge. as soon as it felt the antennae. as was first observed by Huber. except in Europe and North America. but not one excreted. showing that the action was instinctive. or when placed under different circumstances. their sweet excretion to ants: that they do so voluntarily. been produced for the exclusive good of others. I watched them for some time through a lens. that the aphides show no dislike to the ants: if the latter be not present they are at last compelled to eject their excretion. I felt sure that the aphides would want to excrete. therefore probably they do not excrete solely for the good of the ants. can be discovered. or at different seasons of the year. as well as I could. from the observations of Huber. But as the excretion is extremely viscid. I then tickled and stroked them with a hair in the same manner. in which case either the one or the other instinct might be preserved by natural selection. Even the quite young aphides behaved in this manner. One of the strongest instances of an animal apparently performing an action for the sole good of another. And such instances of diversity of instinct in the same species can be shown to occur in nature. Again. I allowed an ant to visit them. as each takes advantage of the weaker bodily structure of other species.. the following facts show. how very generally gradations. Although there is no evidence that any animal performs an action for the exclusive good of another species. by its eager way of running about to be well aware what a rich flock it had discovered. as the ants do with their antennae. which was eagerly devoured by the ant. as in the case of corporeal structure. and not the result of experience. It is certain. is that of aphides voluntarily yielding. etc. it then began to play with its antennae on the abdomen first of one aphis and then of another.allowance for the instincts of animals having been but little observed. and for no instinct being known among extinct species. but has never. and conformably to my theory. So again certain instincts cannot be considered as 243 . and it immediately seemed. but not one excreted. it is no doubt a convenience to the aphides to have it removed.

but want of space prevents me. but often from causes wholly unknown to us. The fear of man is slowly acquired. both in extent and direction. by the various animals which inhabit desert islands. with wax hardened with vermilion or softened with lard. even in England. so wary in England. namely. It has lately been shown that bees. That the mental qualities of animals of the same kind. with which he had covered decorticated trees. and the inheritance of such variations. it has been asked. and on the nature and temperature of the country inhabited. as may be seen in nestling birds. as many instances as possible ought to be given. as I have elsewhere shown. are indispensable for the action of natural selection. as I have seen. used a cement of wax and turpentine. and the magpie. but as details on this and other such points are not indispensable. So it is with the nests of birds. We may safely attribute the greater wildness of our large birds to this cause. As some degree of variation in instincts under a state of nature. Why. the migratory instinct. without the facts in detail. instead of searching for pollen. 244 . is tame in Norway. vary much. though it is strengthened by experience. in the greater wildness of all our large birds in comparison with our small birds. and in its total loss. will gladly use a very different substance. 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.absolutely perfect. which. and we see an instance of this. But I am well aware that these general statements. if instinct be variable. might have given rise. they may be here passed over. for in uninhabited islands large birds are not more fearful than small. and by the sight of fear of the same enemy in other animals. if advantageous to the species. to new instincts. Andrew Knight observed that his bees. I can only repeat my assurance. Several cases could also be adduced of occasional and strange habits in wild animals. Audubon has given several remarkable cases of differences in the nests of the same species in the northern and southern United States. that I do not speak without good evidence. for the large birds have been most persecuted by man. instead of laboriously collecting propolis. can produce but a feeble effect on the reader's mind. oatmeal. as is the hooded crow in Egypt. I can only assert that instincts certainly do vary—for instance. could be shown by many facts. born in a state of nature. Fear of any particular enemy is certainly an instinctive quality. which vary partly in dependence on the situations chosen. through natural selection.

and these tendencies are known to be inherited. when young and without any training. a herd of deer. St. and without the end being known—for the young pointer can no more know that he points to aid his master. habits. for instance. as they may be called. and have been transmitted for an incomparably shorter period. we should assuredly call these actions instinctive. is well shown when different breeds of dogs are crossed. and in nearly the same manner by each individual. 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. A number of curious and authentic instances could be given of various shades of disposition and taste. and another kind of wolf rushing round. one naturally takes to catching rats. instead of at. and likewise of the oddest tricks. by shepherd-dogs. Thus it is known that a cross with a bull-dog has affected for many generations the courage and obstinacy of 245 . John. and another mice. I cannot see that these actions. It is notorious how much domestic animals vary in their mental qualities. instead of at. performed with eager delight by each breed. and then slowly crawl forward with a peculiar gait. and a tendency to run round. 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. are certainly far less fixed than natural instincts. 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. If we were to behold one kind of wolf. One cat. of inherited variations of instinct in a state of nature will be strengthened by briefly considering a few cases under domestication. Inherited Changes Of Habit Or Instinct In Domesticated Animals The possibility. and how curiously they become mingled. retrieving is certainly in some degree inherited by retrievers. How strongly these domestic instincts. With cats. and driving them to a distant point. and dispositions are inherited. Domestic instincts. and another hunted on marshy ground and almost nightly caught woodcocks or snipes. performed without experience by the young. always brought home game birds. associated with certain frames of mind or periods of time. under less fixed conditions of life. or even probability. according to Mr.2. but they have been acted on by far less rigorous selection. another hares or rabbits. stand motionless like a statue. a flock of sheep. as soon as it scented its prey.

hardly any animal is more difficult to tame than the young of the wild rabbit. but this is not true. only the exaggerated pause of an animal preparing to spring on its prey. It is scarcely possible to doubt that 246 . and unconscious selection is still in progress. dogs which stand and hunt best. as many have thought. and this is known occasionally to happen. which cannot fly eighteen inches high without going head over heels. On the other hand. habit alone in some cases has sufficed. the tumbler-pigeon to tumble—an action which. and near Glasgow there are house-tumblers. and this dog showed a trace of its wild parentage only in one way. Familiarity alone prevents our seeing how largely and how permanently the minds of our domestic animals have been modified. as each man tries to procure. methodical selection and the inherited effects of compulsory training in each successive generation would soon complete the work. whose great-grandfather was a wolf. in a pure terrier: the act of pointing is probably. by not coming in a straight line to his master.greyhounds. resemble natural instincts. Brent. never wish to sit on their eggs. Domestic instincts are sometimes spoken of as actions which have become inherited solely from long-continued and compulsory habit. and a cross with a greyhound has given to a whole family of shepherd-dogs a tendency to hunt hares. had not some one dog naturally shown a tendency in this line. but I can hardly suppose that domestic rabbits have often been selected for tameness alone. as I once saw. or probably could have taught. which in a like manner become curiously blended together. These domestic instincts. as I have witnessed. Le Roy describes a dog. No one would ever have thought of teaching." that is. and for a long period exhibit traces of the instincts of either parent: for example. It may be doubted whether any one would have thought of training a dog to point. is performed by young birds. We may believe that some one pigeon showed a slight tendency to this strange habit. 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. that have never seen a pigeon tumble. and that the long-continued selection of the best individuals in successive generations made tumblers what they now are. when called. scarcely any animal is tamer than the young of the tame rabbit. so that we must attribute at least the greater part of the inherited change from extreme wildness to extreme tameness. without intending to improve the breed. to habit and long-continued close confinement. When the first tendency to point was once displayed. when thus tested by crossing. as I hear from Mr.

on the other hand. they are destroyed. It is not that chickens have lost all fear. in other cases compulsory habit has done nothing. and this is evidently done for the instinctive purpose of allowing. young chickens have lost wholly by habit. In some cases compulsory habit alone has sufficed to produce inherited mental changes. So it is with young pheasants reared in England under a hen. foxes. are most eager to attack poultry. which at first appeared from what we must in our ignorance call an accident. for I am informed by Captain Hutton that the young chickens of the parent stock. so that habit and some degree of selection have probably concurred in civilising by inheritance our dogs. sheep. where the savages do not keep these domestic animals. do our civilised dogs. and if not cured. But this instinct retained by our chickens has become useless under domestication. peculiar mental habits and actions. during successive generations. Hence. 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. we may conclude that under domestication instincts have been acquired and natural instincts have been lost. that fear of the dog and cat which no doubt was originally instinctive in them.the love of man has become instinctive in the dog. partly by habit and partly by man selecting and accumulating. when kept tame. On the other hand. sheep and pigs. All wolves. are at first excessively wild. as we see in wild ground-birds. but in most cases habit and selection have probably concurred. How rarely. and pigs! No doubt they occasionally do make an attack. when reared in India under a hen. but fear only of dogs and cats. their mother to fly away. even when quite young. the Gallus bankiva. jackals and species of the cat genus. require to be taught not to attack poultry. pursued both methodically and unconsciously. for the mother-hen has almost lost by disuse the power of flight. and all has been the result of selection. and are then beaten. 247 . 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.

the slave-making instinct of certain ants. perhaps. 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. I will select only three. best understand how instincts in a state of nature have become modified by selection by considering a few cases. the instinct which leads the cuckoo to lay her eggs in other birds' nests. Special Instincts We shall. namely. 248 .3.

sits on them and feeds her young. but at intervals of two or three days. 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. and as both were nearly full feathered. But in all cases. By a continued process of this nature. not daily. there could be no mistake in their identification.4. and in their turn would be apt to lay their eggs in other birds' nests. and the first hatched young would probably have to be fed by the male alone. 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. If this were the case the process of laying and hatching might be inconveniently long. Now let us suppose that the ancient progenitor of our European cuckoo had the habits of the American cuckoo. if she were to make her own nest and sit on her own eggs. of Iowa. by Adolf Muller. all at the same time. then the old birds or the fostered young would gain an advantage. that the cuckoo occasionally lays her eggs on the bare ground. 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. But the American cuckoo is in this predicament. that he once found in Illinois a young cuckoo. and thus be more successful in rearing their young. more especially as she migrates at a very early period. It has been both asserted and denied that the American cuckoo occasionally lays her eggs in other birds' nests. together with a young jay in the nest of a blue jay (Garrulus cristatus). so that. aboriginal instinct of nidification. for she makes her own nest and has eggs and young successively hatched. It has been objected that I have not noticed other related instincts and adaptations of structure in the cuckoo. also recently been ascertained on sufficient evidence. This rare event is probably a case of reversion to the long-lost. I believe that the strange instinct of our cuckoo has been generated. but I have lately heard from Dr. Merrill. Instincts Of The Cuckoo It is supposed by some naturalists that the more immediate cause of the instinct of the cuckoo is that she lays her eggs. If the old bird profited by this occasional habit through being enabled to emigrate earlier or through any other cause. It has. I could also give several instances of various birds which have been known occasionally to lay their eggs in other birds' nests. and that she occasionally laid an egg in another bird's nest. speculation on an instinct 249 . which are spoken of as necessarily co-ordinated. encumbered as she could hardly fail to be by having eggs and young of different ages at the same time.

to an extraordinary degree in colour. as well as in size. is useless. so that in this respect. now. for we have hitherto had no facts to guide us. Now. with rare exceptions. Ramsay remarks that two of the Australian cuckoos. Ramsay. that the common cuckoo. 250 . as is shown by her laying her dull and pale-coloured eggs in the nest of the hedgewarbler with bright greenish-blue eggs. soon after birth. Thirdly. Had our cuckoo invariably displayed the above instinct. or. 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). it is not rare to find two and even three eggs in the same nest. has the instinct. so as to have deceived certain foster-parents.parasitic American cuckoo alone were known. in order that the young cuckoo may get sufficient food. natural selection might have secured and fixed any advantageous variation. for these would have been more safely hatched and reared. that the eggs are remarkably small. that the young cuckoo. when they lay their eggs in an open nest. as is more probable. 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. The eggs of the Australian bronze cuckoo vary. if it had been of an advantage to this species to have laid eggs even smaller than those now laid. we have learned something about three Australian species. In the bronze cuckoo the eggs vary greatly in size. from eight to ten lines in length. it would assuredly have been added to those which it is assumed must all have been acquired together. Secondly. according to Mr. then there is no difficulty in believing that a race or species might have been formed which would have laid smaller and smaller eggs. lays only one egg in a nest. the strength and a properly shaped back for ejecting its foster. The European species apparently manifests some tendency towards a similar instinct. Ramsay's observations. The chief points to be referred to are three: first. which then perish from cold and hunger. This has been boldly called a beneficent arrangement. owing to Mr. manifest a decided preference for nests containing eggs similar in colour to their own. Until recently the instincts of the European and of the non.known to us only in a single species. which lay their eggs in other birds' nests. 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. not exceeding those of the skylark—a bird about one-fourth as large as the cuckoo. but not rarely departs from it. so that the large and voracious young bird receives ample food.brothers. Mr.

and was again thrown out. in the act of ejecting its foster-brothers. Gould was formerly inclined to believe that the act of ejection was performed by the foster-parents themselves. when somewhat advanced in age and strength. Hudson says it is probable that they are occasionally parasitic. But he has now received a trustworthy account of a young cuckoo which was actually seen. for those cuckoos which had such habits and structure best developed would be the most securely reared. during successive generations. allied to our starlings. and as the latter at this age is in a most helpless condition. and structure necessary for the work of ejection. They either lay their eggs in the nest thus appropriated. 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 strength.In the case of the European cuckoo. a widely distinct genus of American birds. the blind desire. while still blind and not able even to hold up its own head. or than in young snakes acquiring in their upper jaws. or oddly enough build one for themselves on the top of it. if it were of great importance for the young cuckoo. the habit having been afterwards improved. and the species present an interesting gradation in the perfection of their instincts. have parasitic habits like those of the cuckoo. The sexes of Molothrus badius are stated by an excellent observer. 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. Some species of Molothrus. I can see no special difficulty in its having gradually acquired. Mr. the offspring of the foster-parents are commonly ejected from the nest within three days after the cuckoo is hatched. One of these was replaced in the nest by the observer. With respect to the means by which this strange and odious instinct was acquired. as Owen has remarked. The first step towards the acquisition of the proper instinct might have been mere unintentional restlessness on the part of the young bird. Hudson. They usually sit on their own eggs and rear their own young. a transitory sharp tooth for cutting through the tough egg-shell. but Mr. and transmitted to an earlier age. sometimes to live promiscuously together in flocks. For if each part is liable to individual variations at all ages. for he has seen the young of 251 . and sometimes to pair. Mr. They either build a nest of their own or seize on one belonging to some other bird. occasionally throwing out the nestlings of the stranger. and both cases must stand or fall together with the whole theory of natural selection. as is probably the case. to receive as much food as possible soon after birth.

and M. Many bees are parasitic. the M. however. for they do not possess the pollen-collecting apparatus which would have been indispensable if they had stored up food for their own young. as with the cuckoo.this species following old birds of a distinct kind and clamouring to be fed by them. They often lay so many eggs—from fifteen to twenty—in the same foster-nest. for these bees have not only had their instincts but their structure modified in accordance with their parasitic habits. This habit is not very uncommon with the Gallinaceae. but it is remarkable that several together sometimes commence to build an irregular untidy nest of their own. and throws some light on the singular instinct of the ostrich. however. transition?" Various birds. Mr. invariably lays its eggs in the nests of strangers. the extraordinary habit of pecking holes in the eggs. A third species. They never. whether of their own species or of their foster parents. as far as it is known. although the Tachytes nigra generally 252 . pecoris of North America. bonariensis. that few or none can possibly be hatched. which are thus wasted. moreover. but. has acquired instincts as perfect as those of the cuckoo. but as small consequences of one general law. the M. for a surprising number of eggs lie strewed over the plains. The parasitic habits of another species of Molothrus. "Must we consider these habits. but are still far from perfect. as has already been remarked. The instinct. are much more highly developed than those of the last. so that the young bird is securely reared. This case is more remarkable than that of the cuckoo. They have. of the American ostrich. Some species of Sphegidae (wasp-like insects) are likewise parasitic. has not as yet been perfected. occasionally lay their eggs in the nests of other birds. and regularly lay their eggs in the nests of other kinds of bees. In this family several hen birds unite and lay first a few eggs in one nest and then in another. Hudson has ascertained. Fabre has lately shown good reason for believing that. and asks. as far as Mr. namely. at intervals of two or three days. which they find in the appropriated nests. Hudson is a strong disbeliever in evolution. They drop also many eggs on the bare ground. and these are hatched by the males. not as especially endowed or created instincts. complete a nest for themselves. for it never lays more than one egg in a foster-nest. This instinct may probably be accounted for by the fact of the hens laying a large number of eggs. but he appears to have been so much struck by the imperfect instincts of the Molothrus bonariensis that he quotes my words. This bird. so that in one day's hunting I picked up no less than twenty lost and wasted eggs. as in the case of the Molothrus bonariensis. as on the leaves of a large thistle. placed in singular ill-adapted situations.

makes its own burrow and stores it with paralysed prey for its own larvae. 253 . yet that. it takes advantage of the prize. as with that of the Molothrus or cuckoo. In this case. and becomes for the occasion parasitic. I can see no difficulty in natural selection making an occasional habit permanent. if of advantage to the species. and if the insect whose nest and stored food are feloniously appropriated. be not thus exterminated. when this insect finds a burrow already made and stored by another sphex.

F. fusca). Formica sanguinea. or of feeding their own larvae. though most energetic and courageous in capturing slaves. fusca) are found only in their own proper communities. but with plenty of the food which they like best. and like their masters are much agitated and defend the nest: when the nest is much disturbed. it would have been hopeless to speculate how so wonderful an instinct could have been perfected. Although fully trusting to the statements of Huber and Mr. do no other work. This ant is absolutely dependent on its slaves. Another species. the slaves occasionally come out. they could not even feed themselves. it is clear that the slaves feel quite at home. and many perished of hunger. so that the contrast in their appearance is great. Smith. that when Huber shut up thirty of them without a slave. I will give the observations which I made in some little detail. of the British Museum. as any one may well be excused for doubting the existence of so extraordinary an instinct as that of making slaves. Hence. Males and fertile females of the slave-species (F. During the months of June and 254 . they did nothing. and its habits have been attended to by Mr. So utterly helpless are the masters. What can be more extraordinary than these well-ascertained facts? If we had not known of any other slave-making ant. without their aid. The males and fertile females do no work of any kind. fed and saved the survivors. a better observer even than his celebrated father. I tried to approach the subject in a sceptical frame of mind. Smith. The slaves are black and not above half the size of their red masters. When the nest is slightly disturbed. and with their larvae and pupae to stimulate them to work. and the larvae and pupae are exposed. it is the slaves which determine the migration. Huber to be a slave-making ant. Huber then introduced a single slave (F. the species would certainly become extinct in a single year. Hence. They are incapable of making their own nests. sanguinea. and they have to migrate. and have never been observed in the nests of F. I opened fourteen nests of F. to whom I am much indebted for information on this and other subjects. This species is found in the southern parts of England. and the workers or sterile females. was likewise first discovered by P. and found a few slaves in all. Slave-Making Instinct This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber. sanguinea. and she instantly set to work. and put all to rights. and actually carry their masters in their jaws. the slaves work energetically together with their masters in carrying them away to a place of safety. When the old nest is found inconvenient. made some cells and tended the larvae.5.

July. and it was a most interesting spectacle to behold the masters carefully carrying their slaves in their jaws instead of being carried by them. and has never seen the slaves. I came across a community with an unusually large stock of slaves. during these months. At the same time I laid on the same place a small parcel of the pupae of another species. they were eagerly seized and carried off by the tyrants. on the other hand. probably depends merely on the slaves being captured in greater numbers in Switzerland than in England. One day I fortunately witnessed a migration of F. though present in large numbers in August. the slaves are very few in number. and put them down on a bare spot near the place of combat. and food of all kinds. rufescens. in the month of July. he considers them as strictly household slaves. 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. twenty-nine yards distant. they had been victorious in their late combat. but Mr. sanguinea. Another day my attention was struck by about a score of the slave-makers haunting the same spot. F. as Huber expressly states. as in the case of F. According to Huber. Hence. however. I watched for many hours several nests in Surrey and Sussex. The masters. after all. flava. with a few of these little yellow ants still clinging to the fragments of their nest. their principal office is to search for aphides. twenty-five yards distant. The latter ruthlessly killed their small opponents and carried their dead bodies as food to their nest. they approached and were vigorously repulsed by an independent community of the slave species (F. sanguinea from one nest to another. June and August. may be constantly seen bringing in materials for the nest. I then dug up a small parcel of the pupae of F. but they were prevented from getting any pupae to rear as slaves. I thought that they might behave differently when more numerous. and evidently not in search of food. During the year 1860. and. and they alone open and close the doors in the morning and evening. the slaves in Switzerland habitually work with their masters in making the nest. either leave or enter the nest. and I observed a few slaves mingled with their masters leaving the nest. which they ascended together. As. who had ample opportunities for observation. and marching along the same road to a tall Scotch-fir tree. both in Surrey and Hampshire. fusca). on three successive years. This species is sometimes. who perhaps fancied that. Smith informs me that he has watched the nests at various hours during May. fusca from another nest. probably in search of aphides or cocci. This difference in the usual habits of the masters and slaves in the two countries. though 255 .

rufescens. sanguinea emerge. and when they migrate. fusca. Now I was curious to ascertain whether F. flava. it is very courageous. The latter does not build its own nest. or even the earth from the nest. and found a number of these ants returning home and entering their nests. In one instance I found to my surprise an independent community of F. from those of the little and furious F. tend and milk as it may be called. whence I saw the last individual of F. for about forty yards back. flava. they took heart and carried off the pupae. making and bringing materials for the nest: both. I traced a long file of ants burdened with booty. sanguinea present with those of the continental F. Although so small a species. in regard to the wonderful instinct of making slaves. and in the early part of the summer extremely few. which they habitually make into slaves. an image of despair over its ravaged home. and cannot even feed itself: it is absolutely dependent on its numerous slaves. the masters carry the slaves. and it was evident that they did at once distinguish them. The nest. but in about a quarter of an hour. shortly after all the little yellow ants had crawled away. does not collect food for itself or its young. carrying a pupa. Let it be observed what a contrast the instinctive habits of F.rarely. for we have seen that they eagerly and instantly seized the pupae of F. possesses much fewer slaves. carrying the dead bodies of F. One evening I visited another community of F. of F. fusca were rushing about in the greatest agitation. and I have seen it ferociously attack other ants. for two or three individuals of F. whereas they were much terrified when they came across the pupae. and the masters alone go on slave-making expeditions. does not determine its own migrations. as has been described by Mr. fusca. sanguinea could distinguish the pupae of F. but I was not able to find the desolated nest in the thick heath. which they rarely capture. must have been close at hand. fusca (showing that it was not a migration) and numerous pupae. but chiefly the slaves. Formica sanguinea. The masters determine when and where a new nest shall be formed. flava under a stone beneath a nest of the slave-making F. and quickly ran away. Both in Switzerland and England the slaves seem to have the exclusive care of the larvae. the little ants attacked their big neighbours with surprising courage. however. to a very thick clump of heath. on the other hand. their 256 . In Switzerland the slaves and masters work together. Such are the facts. though they did not need confirmation by me. sanguinea. and when I had accidentally disturbed both nests. Smith. made into slaves. and one was perched motionless with its own pupa in its mouth on the top of a spray of heath. sanguinea.

if scattered near their nests. their slaves and larvae. 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. if carried out to a much less extent even than in our British F. But as ants which are not slave-makers. will. and thus both collect food for the community. If their presence proved useful to the species which had seized them—if it were more advantageous to this species. to capture workers than to procreate them—the habit of collecting pupae.aphides. sanguinea. as I have seen. sanguinea originated I will not pretend to conjecture. and the foreign ants thus unintentionally reared would then follow their proper instincts. it is possible that such pupae originally stored as food might become developed. might by natural selection be strengthened and rendered permanent for the very different purpose of raising slaves. and do what work they could. By what steps the instinct of F. In England the masters alone usually leave the nest to collect building materials and food for themselves. carry off pupae of other species. is less aided by its slaves than the same species in Switzerland. So that the masters in this country receive much less service from their slaves than they do in Switzerland. 257 . originally for food. as we have seen. which. When the instinct was once acquired.

with the basal edges of its six sides bevelled so as to join an inverted pyramid. We hear from mathematicians that bees have practically solved a recondite problem. Waterhouse. would find it very difficult to make cells of wax of the true form. These latter cells are nearly spherical and of nearly equal sizes.6. In the series between the extreme perfection of the cells of the hivebee and the simplicity of those of the humble-bee. I was led to investigate this subject by Mr. of three rhombs. it seems at first quite inconceivable how they can make all the necessary angles and planes. and the three which form the pyramidal base of a single cell on one side of the comb. is an hexagonal prism. At one end of a short series we have humble-bees. but will merely give an outline of the conclusions at which I have arrived. or even perceive when they are correctly made. with the least possible consumption of precious wax in their construction. though this is effected by a crowd of bees working in a dark hive. sometimes adding to them short tubes of wax. But the difficulty is not nearly so great as at first appears: all this beautiful work can be shown. and are aggregated into an irregular mass. enter into the composition of the bases of three adjoining cells on the opposite side. But the important point to notice is. so beautifully adapted to its end. It has been remarked that a skilful workman. carefully described and figured by Pierre Huber. and. as is well known. 258 . and see whether Nature does not reveal to us her method of work. to follow from a few simple instincts. At the other end of the series we have the cells of the hive-bee. Let us look to the great principle of gradation. in which the young are hatched. I think. some large cells of wax for holding honey. Granting whatever instincts you please. These rhombs have certain angles. but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells. which use their old cocoons to hold honey. and have made their cells of the proper shape to hold the greatest possible amount of honey. placed in a double layer: each cell. He must be a dull man who can examine the exquisite structure of a comb. who has shown that the form of the cell stands in close relation to the presence of adjoining cells. in addition. without enthusiastic admiration. and likewise making separate and very irregular rounded cells of wax. with fitting tools and measures. The Melipona itself is intermediate in structure between the hive and humble bee. we have the cells of the Mexican Melipona domestica. and the following view may. be considered only as a modification of his theory. perhaps. Cell-Making Instinct Of The Hive-Bee I will not here enter on minute details on this subject.

each cell consists of an outer spherical portion. is manifestly a gross imitation of the three-sided pyramidal base of the cell of the hive-bee. from the centres of the six surrounding spheres in the same layer. Hence we may safely conclude that. and this pyramid. and what is more important.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. and of two. it occurred to me that if the Melipona had made its spheres at some given distance from each other. the three flat surfaces are united into a pyramid. by this manner of building. three. the resulting structure would have been as perfect as the comb of the hive-bee. then. and tells me that it is strictly correct:If a number of equal spheres be described with their centres placed in two parallel layers.41421 (or at some lesser distance). but are of the same thickness as the outer spherical portions. of Cambridge. 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. if planes of intersection between the several spheres in both layers be formed. and yet each flat portion forms a part of two cells. or more flat surfaces. that whatever the typical form of the cell may be. As in the cells of the hive-bee. from the spheres being nearly of the same size. and this geometer has kindly read over the following statement. Hence. so much so. with the centre of each sphere at the distance of radius x sqrt(2) or radius x 1. so here. and in themselves not very 259 . But I hear from Professor Wyman. realised. is very frequently and necessarily the case. Reflecting on this case. if we could slightly modify the instincts already possessed by the Melipona. labour. that the accuracy of the workmanship of the bee has been greatly exaggerated. it is rarely. and at the same distance from the centres of the adjoining spheres in the other and parallel layer. if ever. the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. It is obvious that the Melipona saves wax. drawn up from his information. Accordingly I wrote to Professor Miller. for the flat walls between the adjoining cells are not double. who has made numerous careful measurements. three or more other cells. there will result a double layer of hexagonal prisms united together by pyramidal bases formed of three rhombs. which. according as the cell adjoins two. as Huber has remarked. When one cell rests on three other cells. but this is never permitted. and had made them of equal sizes and had arranged them symmetrically in a double layer.

seeing that she already does so to a certain extent. and were in depth about one sixth of the diameter of the sphere of which they formed a part. Following the example of Mr. so that each hexagonal prism was built upon the scalloped edge of a smooth basin. that she always describes her spheres so as to intersect to a certain extent. As soon as this occurred. and began to build up flat walls of wax on the lines of intersection between the basins. Tegetmeier. that the bottoms of the basins. instead of a thick. We must suppose the Melipona to arrange her cells in level layers. would have broken into each other from the opposite 260 . and of equal sizes. but the ridge of wax was so thin. I separated two combs. but she is already so far enabled to judge of distance. if they had been excavated to the same depth as in the former experiment. I then put into the hive. wherever several bees had begun to excavate these basins near together. rectangular strip of wax: the bees instantly began to excavate minute circular pits in it. through natural selection. apparently by turning round on a fixed point.e. they made them wider and wider until they were converted into shallow basins. and then she unites the points of intersection by perfectly flat surfaces. 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. this bee would make a structure as wonderfully perfect as that of the hive-bee. instead of on the straight edges of a three-sided pyramid as in the case of ordinary cells. and as they deepened these little pits. that she can somehow judge accurately at what distance to stand from her fellow-labourers when several are making their spheres. in the same way as before.wonderful. and put between them a long. and this is the greatest difficulty. as she already does her cylindrical cells. and of about the diameter of a cell. about the width of an ordinary cell). But this theory can be tested by experiment. and this would not be very surprising. thick. the rims of the basins intersected or broke into each other. the bees ceased to excavate. knife-edged ridge. The bees instantly began on both sides to excavate little basins near to each other. 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. and we must further suppose. We must suppose the Melipona to have the power of forming her cells truly spherical. rectangular piece of wax. It was most interesting to observe that. and seeing what perfectly cylindrical burrows many insects make in wood. appearing to the eye perfectly true or parts of a sphere. her inimitable architectural powers. coloured with vermilion. a thin and narrow.

so that the basins. In one wellmarked instance. and convex on the opposed side where the bees had worked less quickly. but never allowing the spheres to break into each other. and allowed the bees to go on working for a short time. Considering how flexible thin wax is. were situated. and thus flatten it. large portions of a rhombic plate were thus left between the opposed basins. as soon as they had been a little deepened. The bees. 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. 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. where I suppose that the bees had excavated too quickly. and then stopping their work. and I found that the rhombic plate had been completed. for I have noticed half-completed rhombs at the base of a just-commenced cell.sides. and had become PERFECTLY FLAT: it was absolutely impossible. by standing at the proper distance from each other. whilst at work on the two sides of a strip of wax. which were slightly concave on one side. Now bees. perceiving when they have gnawed the wax away to the proper thinness. that they could have effected this by gnawing away the convex side. they could make their cells of the proper shape. do make a rough. always working circularly as they deepen each cell. however. I put the comb back into the hive. from the extreme thinness of the little plate. if the bees were to build for themselves a thin wall of wax. came to have flat bases. I do not see that there is any difficulty in the bees. exactly along the planes of imaginary intersection between the basins on the opposite side of the ridge of wax. as may be clearly seen by examining the edge of a growing comb. and by endeavouring to make equal spherical hollows. circumferential wall or rim all round the comb. did not suffer this to happen. by stopping work at the planes of intersection. in other parts. 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. From the experiment of the ridge of vermilion wax we can see that. from the unnatural state of things. by excavating at the same rate. only small portions. in order to have thus succeeded in leaving flat plates between the basins. had not been neatly performed. and these flat bases. and they stopped their excavations in due time. and they gnaw this away from the opposite sides. as far as the eye could judge. but the work. In some parts. and again examined the cell. They do not make the 261 . formed by thin little plates of the vermilion wax left ungnawed.

very thin wall is left in the middle. until the hexagonal walls are commenced. The manner in which the bees build is curious. that the very first cell is excavated out of a little parallel-sided wall of wax. but only that one rhombic plate which stands on the extreme growing margin. on an average of twelve measurements made near the border of the comb. corresponding in position to the planes of the rhombic basal plates of future cells. but I will not here enter on details. By the above singular manner of building. whereas the basal rhomboidal plates are thicker. they always make the first rough wall from ten to twenty times thicker than the excessively thin finished wall of the cell. as far as I have seen. and then to begin cutting it away equally on both sides near the ground. that a multitude of bees all work together. which will ultimately be left. as the case may be. is not. from twenty-one measurements. I have several specimens showing clearly that they can do this. Even in the rude circumferential rim or wall of wax round a growing comb. the first commencement having always been a little hood of wax. Some of these statements differ from those made by the justly celebrated elder Huber. I could show that they are conformable with my theory. being. 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. But the rough wall of wax has in every case to be finished off. being thus crowned by a strong coping of wax. We see how important a part excavation plays in the construction of the cells. the bees can cluster and crawl over the comb without injuring the delicate hexagonal walls. by being largely gnawed away on both sides. flexures may sometimes be observed. strictly correct.whole three-sided pyramidal base of any one cell at the same time. We shall understand how they work. nearly in the proportion of three to two. From all the cells. Huber's statement. one bee after 262 . These walls. having a mean thickness. the masons always piling up the cut-away cement. along the plane of intersection between two adjoining spheres. of 1/229 of an inch. both those just commenced and those completed. as Professor Miller has kindly ascertained for me. or the two plates. We shall thus have a thin wall steadily growing upward but always crowned by a gigantic coping. strength is continually given to the comb. vary greatly in thickness. by supposing masons first to pile up a broad ridge of cement. till a smooth. 1/352 of an inch in thickness. and if I had space. with the utmost ultimate economy of wax. and adding fresh cement on the summit of the ridge. It seems at first to add to the difficulty of understanding how the cells are made. and they never complete the upper edges of the rhombic plates. but I am convinced of their accuracy.

When bees have a place on which they can stand in their proper positions for working—for instance. It was really curious to note in cases of difficulty. all trying to sweep equal spheres. sweeping spheres or cylinders. or leaving ungnawed. 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. as Huber has stated. This capacity in bees of laying down under certain circumstances a rough wall in its proper place between two justcommenced cells. as when two pieces of comb met at an angle. which seems at first subversive of the foregoing theory. with an extremely thin layer of melted vermilion wax. on a slip of wood. I was able practically to show this fact. the planes of intersection between these spheres. or the extreme margin of the circumferential rim of a growing comb. placed directly under the middle of a comb growing downwards. and then building up. 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. if she were to work alternately on the inside and outside of two or three cells commenced at the same time. as it bears on a fact. 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. as far as I have seen. by covering the edges of the hexagonal walls of a single cell. is important. always standing at the proper relative distance from the parts of the cells just begun. projecting beyond the other completed cells. and building up intermediate planes. 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.working a short time at one cell going to another. 263 . 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. that the cells on the extreme margin of wasp-combs are sometimes strictly hexagonal. all instinctively standing at the same relative distance from each other. they can build up a wall intermediate between two adjoining spheres. in its strictly proper place. sometimes recurring to a shape which they had at first rejected. and then. a score of individuals work even at the commencement of the first cell. and worked into the growing edges of the cells all round. by striking imaginary spheres. but. 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. but I have not space here to enter on this subject. so that.

must be an important element of success any family of bees. the 264 . as it probably often has determined. each profitable to the individual under its conditions of life. 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. how a long and graduated succession of modified architectural instincts.As natural selection acts only by the accumulation of slight modifications of structure or instinct. for a wall in common even to two adjoining cells would save some little labour and wax. Hence the saving of wax by largely saving honey. and so be altogether independent of the quantity of honey which the bees can collect. from the same cause. like the cells of the Melipona. for in this case a large part of the bounding surface of each cell would serve to bound the adjoining cells. and the security of the hive is known mainly to depend on a large number of bees being supported. Again. if she were to make her cells closer together. Of course the success of the species may be dependent on the number of its enemies. or parasites. it is known that bees are often hard pressed to get sufficient nectar. for then. all tending towards the present perfect plan of construction. and I am informed by Mr. it would be advantageous to the Melipona. or on quite distinct causes. it would continually be more and more advantageous to our humble-bees. Moreover. whether a bee allied to our humble-bees could exist in large numbers in any country. But let us suppose that this latter circumstance determined. it may reasonably be asked. and the time consumed in collecting the honey. and more regular in every way than at present. A large store of honey is indispensable to support a large stock of bees during the winter. many bees have to remain idle for many days during the process of secretion. 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. With respect to the formation of wax. and aggregated into a mass. 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. and save much in labour and space. and much labour and wax would be saved. 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 we have seen. so as to intersect a little. Hence. and in the materials of which they are constructed. if they were to make their cells more and more regular. nearer together.

more and more perfectly led the bees to sweep equal spheres at a given distance from each other in a double layer. slight modifications of simpler instincts. no more knowing that they swept their spheres at one particular distance from each other. The bees. by slow degrees. 265 . 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. as far as we can see. and least waste of honey in the secretion of wax. successive. than they know what are the several angles of the hexagonal prisms and of the basal rhombic plates. that individual swarm which thus made the best cells with least labour. is absolutely perfect in economising labour and wax. which in their turn will have had the best chance of succeeding in the struggle for existence. and having transmitted their newly-acquired economical instincts to new swarms. Thus. as I believe. and to build up and excavate the wax along the planes of intersection. can be explained by natural selection having taken advantage of numerous. Beyond this stage of perfection in architecture. this being effected with the greatest possible economy of labour and wax. of course. and the Melipona would make a comb as perfect as that of the hive-bee. that of the hive-bee. having succeeded best. natural selection could not lead. the most wonderful of all known instincts. natural selection having. for the comb of the hive-bee.spherical surfaces would wholly disappear and be replaced by plane surfaces.

believed to be moistened with saliva. Is it then very improbable that the natural selection of individual swifts. 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. No doubt many instincts of very difficult explanation could be opposed to the theory of natural selection—cases.7. until a beak was formed. at the same time that habit. but in accordance with. It must. or spontaneous variations of taste. however. as well constructed for this purpose as that of the nuthatch. (Parus major) alluded to in a previous chapter. To take as an illustration the case of the larger titmouse. subsequently to. and one of the swifts of North America makes its nest (as I have seen) of sticks agglutinated with saliva." The force of this objection rests entirely on the assumption that the changes in the instincts and structure are abrupt. but let the feet of the titmouse vary and grow larger from correlation with the beak. slowly changing habits or taste. which were better and better adapted to break open the seeds. as a modification in the one without an immediate corresponding change in the other would have been fatal. or from any other unknown cause. be admitted that in many instances we cannot conjecture whether it was instinct or structure which first varied. which secreted more and more saliva. Some birds build their nests of mud. 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. Now what special difficulty would there be in natural selection preserving all the slight individual variations in the shape of the beak. or compulsion. and hammers with its beak till it gets at the kernel. cases. and even with flakes of this substance. in which no intermediate gradations are known to exist. In this case a gradual change of structure is supposed to lead to changed instinctive habits. cases of instincts of such trifling 266 . 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. in which we cannot see how an instinct could have originated. this bird often holds the seeds of the yew between its feet on a branch. Objections To The Theory Of Natural Selection As Applied To Instincts: Neuter And Sterile Insects 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. 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.

from being sterile. 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 should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection. As far as instinct alone is concerned. which were inherited by the offspring. which at first appeared to me insuperable. and in instinct. If a working ant or other neuter insect had been an ordinary animal. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure. and yet. I will not here enter on these several cases. as in the shape of the thorax. by individuals having been born with slight profitable modifications. and if such insects had been social. but I will here take only a single case. yet absolutely sterile. and in being destitute of wings and sometimes of eyes. so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny. 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.importance. that of working or sterile ants. But with the working ant we have an insect differing greatly from its parents. The subject well deserves to be discussed at great length. and that these again varied and again were selected. both in our domestic productions and in those in a state of nature. and consequently must believe that they were independently acquired through natural selection. How the workers have been rendered sterile is a difficulty. for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile. and it had been profitable to the community that a number should have been annually born capable of work. but incapable of procreation. that they could hardly have been acted on by natural selection. the wonderful difference in this respect between the workers and the perfect females would have been better exemplified by the hivebee. We have differences correlated not only with one 267 . of all sorts of differences of inherited structure which are correlated with certain ages and with either sex. and so onwards. but not much greater than that of any other striking modification of structure. they cannot propagate their kind. but will confine myself to one special difficulty. and actually fatal to the whole theory. But I must pass over this preliminary difficulty. It may well be asked how it is possible to reconcile this case with the theory of natural selection? First. let it be remembered that we have innumerable instances. namely.

consequently the fertile males and females have flourished. These latter. Breeders of cattle wish the flesh and fat to be well marbled together. when matched. and may thus gain the desired end. though appearing insuperable. have proved advantageous. and transmitted to their fertile offspring a tendency to produce sterile members with the same modifications. Verlot. from having been long and carefully selected to the right degree. but with that short period when the reproductive system is active. some varieties of the double annual stock. disappears. 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. the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection. Hence. and not to the individual. and in the hooked jaws of the male salmon. as in the nuptial plumage of many birds. as I believe. correlated with the sterile condition of certain members of the community. selection has been applied to the family. Such faith may be placed in the power of selection that a breed of cattle. produced oxen with the longest horns. as well as to the individual. always yielding oxen with extraordinarily long horns. 268 . but they likewise yield some single and fertile plants. We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male sex. it is probable. could. we may conclude that slight modifications of structure or of instinct.sex. so with social insects. be formed by carefully watching which individual bulls and cows. always produce a large proportion of seedlings bearing double and quite sterile flowers. or. An animal thus characterized has been slaughtered. Here is a better and real illustration: According to M. but the breeder has gone with confidence to the same stock and has succeeded. relatively to the length of the horns in both the bulls and cows of these same breeds. This difficulty. I can see no great difficulty in any character becoming correlated with the sterile condition of certain members of insect communities. and the double sterile plants with the neuters of the same community. This process must have been repeated many times. by which alone the variety can be propagated. and yet no one ox would ever have propagated its kind. for oxen of certain breeds have longer horns than the oxen of other breeds. As with the varieties of the stock. is lessened. may be compared with the fertile male and female ants. for the sake of gaining a serviceable end. Hence. when it is remembered that selection may be applied to the family.

Thus. the fact that the neuters of several ants differ. namely. F. which. can be plainly distinguished. or the domestic cattle as they may be called. It will indeed be thought that I have an overweening confidence in the principle of natural selection. presenting gradations of structure. or that both large and small are numerous. though small. and. when I do not admit that such wonderful and well-established facts at once annihilate the theory. the use of which is quite unknown: in the Mexican Myrmecocystus. According to this view we ought occasionally to find in the same nest neuter-insects. the larger workers have simple eyes (ocelli).But we have not as yet touched on the acme of the difficulty. In the simpler case of neuter insects all of one caste. with some few of intermediate size. which. sometimes to an almost incredible degree. we may conclude from the analogy of ordinary variations. the workers of one caste never leave the nest. whereas the smaller workers have 269 . Smith has shown that the neuters of several British ants differ surprisingly from each other in size and sometimes in colour. Smith has observed. and this we do find. in this species. F. and that by the survival of the communities with females which produced most neuters having the advantageous modification. or rather as any two genera of the same family. Mr. not only from the fertile females and males. even not rarely. but in some few alone. that the successive. It sometimes happens that the larger or the smaller sized workers are the most numerous. have been rendered different from the fertile males and females through natural selection. Formica flava has larger and smaller workers. but are perfectly well defined. the workers of one caste alone carry a wonderful sort of shield on their heads. as I believe. supplying the place of that excreted by the aphides. they are fed by the workers of another caste. and are thus divided into two or even three castes. which our European ants guard and imprison. with jaws and instincts extraordinarily different: in Cryptocerus. all the neuters ultimately came to be thus characterized. slight. as Mr. moreover. profitable modifications did not first arise in all the neuters in the same nest. and they have an enormously developed abdomen which secretes a sort of honey. there are working and soldier neuters. while those of an intermediate size are scanty in numbers. 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 Eciton. being as distinct from each other as are any two species of the same genus. The castes. but from each other. considering how few neuter-insects out of Europe have been carefully examined. do not generally graduate into each other.

that I gladly availed myself of Mr. I speak confidently on this latter point. in his interesting "Naturalist on the Amazons. Lubbock made drawings for me. by acting on the fertile ants or parents. of the working ants of the several sizes differed wonderfully in shape. until all the workers were in this condition. with the camera lucida. as does the widely-different structure of their jaws. all of large size with one form of jaw. For the workers of Myrmica have not even rudiments of ocelli. that the workers of intermediate size have their ocelli in an exactly intermediate condition. yet connected by some few members in an intermediate condition. The reader will perhaps best appreciate the amount of difference in these workers by my giving. though the male and female ants of this genus have well-developed ocelli. differing not only in size. or lastly. and jaws nearly five times as big. but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house. not the actual measurements. With these facts before me. and this is the greatest difficulty. F. Smith's offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa. could form a species which should regularly produce neuters. and those males and females had been continually selected. So that here we have two bodies of sterile workers in the same nest. I believe that natural selection. 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. Mr. and many sixteen feet high. that if the smaller workers had been the most useful to the community. The jaws. we should then have had a species of ant with neuters in nearly the same condition as those of Myrmica. which produced more and more of the smaller workers. 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. though I dare not assert so positively." has described analogous cases. or all of small size with widely different jaws. Bates. I may digress by adding. though the workers can be grouped into castes of different sizes. and I fully believe. Having carefully dissected several specimens of these workers. as Sir J. of the jaws which I dissected from the workers of the several sizes. yet they graduate insensibly into each other.their ocelli rudimentary. but in their organs of vision. of whom many were five feet four inches high. moreover. 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. But the important fact for us is that. 270 . and in the form and number of the teeth.

until none with an intermediate structure were produced. I should never have anticipated that this principle could have been efficient in so high a degree. while man works by acquired knowledge and manufactured instruments. But I must confess. confined to the workers of sterile females. of certain Brazilian crustaceans likewise appearing under two widely distinct male forms. The case. on the same principle that the division of labour is useful to civilised man. is very interesting. as with plants. any amount of modification may be effected by the accumulation of numerous. and then the extreme forms having been produced in greater and greater numbers. I have. We can see how useful their production may have been to a social community of ants. I believe. against the well-known doctrine of inherited habit. For peculiar habits. also. slight. Wallace. and simultaneously another set of workers of a different size and structure. the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest. I have now explained how. without exercise or habit having been brought into play. as advanced by Lamarck. as in the case of the driver ant. But this subject need not here be discussed. and likewise because this is by far the most serious special difficulty which my theory has encountered. has originated. that. spontaneous variations. therefore. which are in any way profitable. in order to show the power of natural selection. through the survival of the parents which generated them. however. could not possibly affect the males and fertile females. Ants.one set of workers of one size and structure. with all my faith in natural selection. however long they might be followed. had not the case of these neuter insects led me to this conclusion. which alone leave descendants. a graduated series having first been formed. and by Fritz Muller. I am surprised that no one has advanced this demonstrative case of neuter insects. at some little but wholly insufficient length. as it proves that with animals. An analogous explanation has been given by Mr. discussed this case. of certain Malayan butterflies regularly appearing under two or even three distinct female forms. work by inherited instincts and by inherited organs or tools. of the equally complex case. both widely different from each other and from their parents. 271 .

but none of the cases of difficulty. there is no real difficulty. Summary I have endeavoured in this chapter briefly to show that the mental qualities of our domestic animals vary. but as small consequences of one general law leading to the advancement of all organic beings—namely. let the strongest live and the weakest die. it may not be a logical deduction. like the males of our Kittywrens. in natural selection accumulating to any extent slight modifications of instinct which are in any way useful. in the same peculiar manner as does our British thrush. we can understand." to roost in. how it is that the thrush of tropical South America lines its nest with mud. the larvae of ichneumonidae feeding within the live bodies of caterpillars.8. on the principle of inheritance. On the other hand. when inhabiting distant parts of the world and living under considerably different conditions of life. that the canon in natural history. species. yet often retaining nearly the same instincts. with only a small hole left in the plaster through which the males feed them and their young when hatched. This theory is also strengthened by some few other facts in regard to instincts. as by that common case of closely allied. and is plainly explicable on the foregoing views. In many cases habit or use and disuse have probably come into play. but is otherwise inexplicable—all tend to corroborate the theory of natural selection. No one will dispute that instincts are of the highest importance to each animal. that no instinct can be shown to have been produced for the good of other animals. but distinct. Finally. though animals take advantage of the instincts of others. annihilate it. ants making slaves." is applicable to instincts as well as to corporeal structure. how it is that the male wrens (Troglodytes) of North America. and that the variations are inherited. but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its fosterbrothers. to the best of my judgment. For instance. 272 . build "cock-nests. of "Natura non facit saltum. under changing conditions of life. 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. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. multiply. not as specially endowed or created instincts. vary.—a habit wholly unlike that of any other known bird. I do not pretend that the facts given in this chapter strengthen in any great degree my theory. Therefore. the fact that instincts are not always absolutely perfect and are liable to mistakes.

273 . removed by domestication — Laws governing the sterility of hybrids — Sterility not a special endowment. but incidental on other differences.Chapter 9 Hybridism Distinction between the sterility of first crosses and of hybrids — Sterility various in degree. affected by close interbreeding. not universal. 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.

two classes of facts. of equal importance with the sterility of species. more especially as the sterility of species when first crossed. This view certainly seems at first highly probable. have been specially endowed with sterility. as I shall show. on the other hand. Pure species have of course their organs of reproduction in a perfect condition. and likewise the fertility of their mongrel offspring. with reference to my theory. though the formative organs themselves are perfect in structure. owing to the sterility in both cases being looked on as a special endowment. by the preservation of successive profitable degrees of sterility. yet when intercrossed they produce either few or no offspring. It is an incidental result of differences in the reproductive systems of the parentspecies. is. when intercrossed. In treating this subject. in order to prevent their confusion. Hybrids. The fertility of varieties. in the second case they are either not at all developed. and that of their hybrid offspring. which is common to the two cases. when the cause of the sterility. The distinction probably has been slurred over. the sterility of species when first crossed. In the first case the two sexual elements which go to form the embryo are perfect. namely. for species living together could hardly have been kept distinct had they been capable of freely crossing. for it seems to make a broad and clear distinction between varieties and species. The subject is in many ways important for us. have their reproductive organs functionally impotent. have generally been confounded. cannot have been acquired. as may be clearly seen in the state of the male element in both plants and animals. or are imperfectly developed.1. This distinction is important. 274 . that is of the forms known or believed to be descended from common parents. as far as the microscope reveals. has to be considered. The view commonly entertained by naturalists is that species. to a large extent fundamentally different. and the sterility of the hybrids produced from them. when crossed. beyond the province of our reasoning powers.

Moreover. when intercrossed. It is certain. I think no better evidence of this can be required than that the two most experienced observers who have ever lived. and (excluding all cases such as the Leguminosae. as Gartner repeatedly crossed some forms. without being deeply impressed with the high generality of some degree of sterility. makes the rule equally universal. and artificially fertilised with their own pollen. for the sterility of species when crossed and of their hybrid offspring. namely Kolreuter and Gartner. in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. quite fertile together.2. Nearly all the plants experimented on by Gartner were potted. Gartner is obliged carefully to count the seeds. Degrees Of Sterility First. he unhesitatingly ranks them as varieties. and were kept in a chamber in his house. must be secluded in order to prevent pollen being brought to it by insects from other plants. in order to show that there is any degree of sterility. and the maximum produced by their hybrid offspring. Kolreuter and Gartner. Gartner. But causes of serious error here intervene: a plant. which the best botanists rank as varieties. on the one hand. That these processes are often injurious to the fertility of a plant cannot be doubted. and. as he believed. for Gartner gives in his table about a score of cases of plants which he castrated. but then he cuts the knot. also. who almost devoted their lives to this subject. arrived at diametrically opposite conclusions in regard to some of the very same 275 . for in ten cases in which he found two forms. to be hybridised. must be castrated. with the average number produced by both pure parent-species in a state of nature. Kolreuter makes the rule universal. on the other hand. considered by most authors as distinct species. and found them absolutely sterile. and he disputes the entire fertility of Kolreuter's ten cases. It is impossible to study the several memoirs and works of those two conscientious and admirable observers. we may doubt whether many species are really so sterile. But in these and in many other cases. that the sterility of various species when crossed is so different in degree and graduates away so insensibly. such as the common red and blue pimpernels (Anagallis arvensis and coerulea). that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins. that the fertility of pure species is so easily affected by various circumstances. He always compares the maximum number of seeds produced by two species when first crossed. and. what is often more important.

Moreover. namely. whenever complicated 276 . namely. Now. if left to themselves. I am strengthened in this conviction by a remarkable statement repeatedly made by Gartner. will generally be fertilised during each generation by pollen from the same flower. But I believe that their fertility has been diminished in nearly all these cases by an independent cause. notwithstanding the frequent ill effects from manipulation. already lessened by their hybrid origin. as from the anthers of the flower itself which is to be fertilised. would be thus effected. The evidence from this source graduates away. It can thus be shown that neither sterility nor fertility affords any certain distinction between species and varieties. that if even the less fertile hybrids be artificially fertilised with hybrid pollen of the same kind. for six or seven. that I cannot doubt the correctness of this conclusion. With respect to this decrease. or other allied hybrids. generally grow in the same garden. with the evidence from fertility adduced by different hybridisers. In regard to the sterility of hybrids in successive generations. Hybrids are seldom raised by experimentalists in great numbers. the visits of insects must be carefully prevented during the flowering season: hence hybrids. sometimes decidedly increases. pollen is as often taken by chance (as I know from my own experience) from the anthers of another flower. yet he asserts positively that their fertility never increases. I have made so many experiments and collected so many facts. and goes on increasing. 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. so that a cross between two flowers. and both sexual elements in hybrid plants are already affected in some degree. or by the same observer from experiments made during different years. their fertility. and as the parent-species. it may first be noticed that when any deviation in structure or constitution is common to both parents. by too close interbreeding. in the process of artificial fertilisation. and in one case for ten generations. though probably often on the same plant. carefully guarding them from a cross with either pure parent. and this would probably be injurious to their fertility. this is often transmitted in an augmented degree to the offspring. but generally decreases greatly and suddenly. though Gartner was enabled to rear some hybrids. showing on the one hand that an occasional cross with a distinct individual or variety increases the vigour and fertility of the offspring. and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences. and on the other hand that very close interbreeding lessens their vigour and fertility.forms.

three were fertilised by Herbert with their own pollen. 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. that individual plants of certain species of Lobelia. Herbert. so careful an observer as Gartner would have castrated his hybrids. that "every ovule in a pod of Crinum capense fertilised by C. This case of the Crinum leads me to refer to a singular fact. namely. and by his having hot-houses at his command. W. and bore good seed. In the genus Hippeastrum." So that here we have perfect.experiments are in progress. Now let us turn to the results arrived at by a third most experienced hybridiser. and Rev. Of his many important statements I will here give only a single one as an example. though this pollen can be proved to be perfectly sound by fertilising other plants or species. in Corydalis as shown by Professor Hildebrand. 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. The difference in their results may. in a first cross between two distinct species. Herbert tried similar 277 . So that with some species. be accounted for by too close interbreeding having been avoided. a bulb of Hippeastrum aulicum produced four flowers. and in other species all the individuals. 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. in contrast with those spontaneously self. Verbascum and Passiflora. namely. can easily be fertilised by the pollen from a distinct species. Scott and Fritz Muller. revolutum produced a plant. may. and after a few days perished entirely. And thus. be in part accounted for by Herbert's great horticultural skill. whereas the pod impregnated by the pollen of the hybrid made vigorous growth and rapid progress to maturity. or even more than commonly perfect fertility. the strange fact of an increase of fertility in the successive generations of ARTIFICIALLY FERTILISED hybrids.fertilised. as I believe. can actually be hybridised much more readily than they can be fertilised by pollen from the same individual plant! To give one instance. in various orchids as shown by Mr. I think. but not by pollen from the same plant." Mr. He experimented on some of the very same species as did Gartner. namely. all the individuals are in this peculiar condition. the Hon. either from the same plant or from another plant of the same hybrid nature. certain abnormal individuals. which vegetated freely.

"reproduces itself as perfectly as if it had been a natural species from the mountains of Chile. For instance. 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. when fairly treated. C. so as to avoid the ill effects of close 278 . but. Herbert asserts that a hybrid from Calceolaria integrifolia and plantaginea. Horticulturists raise large beds of the same hybrid. species most widely dissimilar in general habit. I hardly know of an instance in which two families of the same hybrid have been raised at the same time from different parents. informs me that he raises stocks for grafting from a hybrid between Rhod. few experiments have been fairly tried: for instance. the fact would have been notorious to nurserymen. Again. It should. for he will find on their stigmas plenty of pollen brought from other flowers. etc.. Fuchsia. more sterile. which produce no pollen. ponticum and catawbiense. Any one may readily convince himself of the efficiency of insect agency by examining the flowers of the more sterile kinds of hybrid Rhododendrons. and that this hybrid "seeds as freely as it is possible to imagine.experiments during many years. as not one of these breeds freely in confinement. much fewer experiments have been carefully tried than with plants. These cases serve to show on what slight and mysterious causes the lesser or greater fertility of a species sometimes depends. have been crossed. we have no right to expect that the first crosses between them and the canary. or that their hybrids. but the hybrids themselves are. always gone on decreasing in fertility in each successive generation. for instance. yet many of these hybrids seed freely. as Gartner believed to be the case. the canary-bird has been crossed with nine distinct species of finches. Rhododendron. and always with the same result. owing to few animals breeding freely under confinement." I have taken some pains to ascertain the degree of fertility of some of the complex crosses of Rhododendrons. be borne in mind that. and the injurious influence of close interbreeding is thus prevented. and I am assured that many of them are perfectly fertile. It is notorious in how complicated a manner the species of Pelargonium. Mr. Petunia. Calceolaria. In regard to animals. Noble. I think. though not made with scientific precision. The practical experiments of horticulturists. that is. however. should be perfectly fertile. for by insect agency the several individuals are allowed to cross freely with each other. deserve some notice. and such alone are fairly treated. If our systematic arrangements can be trusted." Had hybrids. if the genera of animals are as distinct from each other as are the genera of plants. with respect to the fertility in successive generations of the more fertile hybrid animals.

in opposition to the constantly repeated admonition of every breeder. however. which was first propounded by Pallas. This latter alternative. This was effected by Mr. It is. cygnoides). species which are so different that they are generally ranked in distinct genera. namely Mr. It has lately been asserted that two such distinct species as the hare and rabbit. and as they are kept for profit. brothers and sisters have usually been crossed in each successive generation. the various races when crossed together are quite fertile. for instance. M. or that the hybrids subsequently reared under domestication became quite fertile. indeed. that whole flocks of these crossed geese are kept in various parts of the country. all are quite fertile together. who raised two hybrids from the same parents. On the contrary. seems by far the most probable. With our domesticated animals. I have reason to believe that the hybrids from Cervulus vaginalis and Reevesii. Eyton. yet in many cases they are descended from two or more wild species. are perfectly fertile. they must certainly be highly or perfectly fertile. yet. Quatrefages states that the hybrids from two moths (Bombyx cynthia and arrindia) were proved in Paris to be fertile inter se for eight generations. In India. The hybrids from the common and Chinese geese (A. So again I have lately acquired decisive evidence that the crossed offspring from the Indian humped and common cattle are inter se perfectly 279 . with perhaps the exception of certain indigenous domestic dogs of South America. for I am assured by two eminently capable judges. but analogy makes me greatly doubt. have often bred in this country with either pure parent. torquatus. and from these two birds he raised no less than eight hybrids (grandchildren of the pure geese) from one nest. and in one single instance they have bred inter se. these cross-bred geese must be far more fertile. whether the several aboriginal species would at first have freely bred together and have produced quite fertile hybrids. where neither pure parent-species exists. almost certain that our dogs are descended from several wild stocks. which are highly fertile when crossed with one of the parent-species. it is not at all surprising that the inherent sterility in the hybrids should have gone on increasing. Although I know of hardly any thoroughly well-authenticated cases of perfectly fertile hybrid animals. but from different hatches. Blyth and Captain Hutton. hardly be doubted. produce offspring.interbreeding. From this fact we must conclude either that the aboriginal parent-species at first produced perfectly fertile hybrids. when they can be got to breed together. and can. and from Phasianus colchicus with P. And in this case.

be considered as absolutely universal. either give up the belief of the universal sterility of species when crossed. The same remarks may be extended to the two chief races of the pig. voice. as well as from those by Mr. 280 . constitution. it may be concluded that some degree of sterility. and from the observations by Rutimeyer on their important osteological differences. but as one capable of being removed by domestication. under our present state of knowledge. or we must look at this sterility in animals. etc. Finally. not as an indelible characteristic.fertile. but that it cannot.. We must. therefore. both in first crosses and in hybrids. Blyth on their differences in habits. is an extremely general result. these two forms must be regarded as good and distinct species. considering all the ascertained facts on the intercrossing of plants and animals.

So in hybrids themselves. The hybrids raised from two species which are very difficult to cross. as we have seen. as in the genus Verbascum. graduates from zero to perfect fertility. beyond that which the plant's own pollen produces. The following conclusions are drawn up chiefly from Gartner's admirable work on the hybridisation of plants. both of first crosses and of hybrids. and the sterility of the hybrids thus produced—two classes of facts which are generally confounded together—is by no means strict. From this absolute zero of fertility. Our chief object will be to see whether or not these laws indicate that species have been specially endowed with this quality. even to an excess of fertility. It has been already remarked. can be united with unusual facility. are generally very sterile. but only the barest outline of the facts can here be given. in which two pure species.3. there are some which never have produced. There are many cases. From this extreme degree of sterility we have self-fertilised hybrids producing a greater and greater number of seeds up to perfect fertility. yet these hybrids are remarkably sterile. When pollen from a plant of one family is placed on the stigma of a plant of a distinct family. and. and probably never would produce. I have been surprised to find how generally the same rules apply to both kingdoms. and the early withering of the flower is well known to be a sign of incipient fertilisation. and. On the other hand. considering how scanty our knowledge is in regard to hybrid animals. that the degree of fertility. but the parallelism between the difficulty of making a first cross. in certain abnormal cases. Laws Governing The Sterility Of First Crosses And Of Hybrids We will now consider a little more in detail the laws governing the sterility of first crosses and of hybrids. even with the pollen of the pure parents. I have taken much pains to ascertain how far they apply to animals. there are 281 . it exerts no more influence than so much inorganic dust. a single fertile seed: but in some of these cases a first trace of fertility may be detected. 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. It is surprising in how many curious ways this gradation can be shown. in order to prevent their crossing and blending together in utter confusion. and produce numerous hybrid offspring. and which rarely produce any offspring. the pollen of different species applied to the stigma of some one species of the same genus. up to nearly complete or even quite complete fertility. yields a perfect gradation in the number of seeds produced.

obstinately failed to fertilise. Now the fertility of first crosses. when at last produced. The fertility. or with extreme difficulty. is largely governed by their systematic affinity. the general resemblance between species in structure and constitution. Even within the limits of the same genus. and on the other hand. In the same family there may be a genus. and on the other hand of very distinct species which unite with the utmost facility. but the hybrids. Even within the limits of the same genus. for it is not always the same in degree when the same two species are crossed under the same circumstances. It can be shown that plants most widely different in habit and general appearance. Many analogous facts could be given. are very fertile. for instance in Dianthus. by no less than eight other species of Nicotiana. in which very many species can most readily be crossed. But the correspondence between systematic affinity and the facility of crossing is by no means strict. is sufficient to prevent two species crossing. as Silene. and in the cotyledons. these two opposite cases occur. as Dianthus. and having strongly marked differences in every part of the flower. or to be fertilised. 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. So it is with hybrids. A multitude of cases could be given of very closely allied species which will not unite. No one has been able to point out what kind or what amount of difference. we meet with this same difference.species which can be crossed very rarely. in any recognisable character. for instance. than is that of pure species. This is clearly shown by hybrids never having been raised between species ranked by systematists in distinct families. and another genus. the many species of Nicotiana have been more largely crossed than the species of almost any other genus. is more easily affected by unfavourable conditions. but Gartner found that N. even in the pollen. But the fertility of first crosses is likewise innately variable. By the term systematic affinity is meant. it depends in part upon the constitution of the individuals which happen to have been chosen for the experiment. and of the hybrids produced from them. in which the most persevering efforts have failed to produce between extremely close species a single hybrid. or only with extreme difficulty. in the fruit. which is not a particularly distinct species. can be crossed. acuminata. Annual and perennial plants. both of first crosses and of hybrids. by very closely allied species generally uniting with facility. deciduous and evergreen 282 .

to fertilise reciprocally M. excepting in their reproductive systems. To give an instance: Mirabilis jalapa can easily be fertilised by the pollen of M. jalapa. yet generally differ in fertility in a small. It is also a remarkable fact that hybrids raised from reciprocal crosses. Several other equally striking cases could be given. an intermediate character between their two parents. I mean the case. Gartner. though externally so like one of their pure parent-species. longiflora with the pollen of M. other species of the same genus have a remarkable power of impressing their likeness on their hybrid offspring. and the hybrids thus produced are sufficiently fertile. He has observed it even between closely related forms (as Matthiola annua and glabra) which many botanists rank only as varieties. of a female-ass being first crossed by a stallion. the one species having first been used as the father and then as the mother. By a reciprocal cross between two species. longiflora. though they rarely differ in external characters. exceptional and abnormal individuals sometimes are born. as is usual. Thuret has observed the same fact with certain sea-weeds or Fuci. though of course compounded of the very same two species.trees. plants inhabiting different stations and fitted for extremely different climates. Several other singular rules could be given from Gartner: for instance. always closely resemble one of them. So again among hybrids which are usually intermediate in structure between their parents. for instance. There is often the widest possible difference in the facility of making reciprocal crosses. and these hybrids are almost always utterly sterile. The diversity of the result in reciprocal crosses between the same two species was long ago observed by Kolreuter. that is of any difference in their structure or constitution. can often be crossed with ease. There are certain hybrids which. instead of having. and such hybrids. and utterly failed. and then a mare by a male-ass: these two species may then be said to have been reciprocally crossed. found that this difference of facility in making reciprocal crosses is extremely common in a lesser degree. are with rare exceptions extremely sterile. but these two powers do not at all necessarily go together. and occasionally in a high degree. some species have a remarkable power of crossing with other species. Such cases are highly important. but Kolreuter tried more than two hundred times. moreover. which closely resemble one of their pure parents. during eight following years. even when the other hybrids raised from seed from the same capsule have a considerable 283 . for they prove that the capacity in any two species to cross is often completely independent of their systematic affinity.

These facts show how completely the fertility of a hybrid may be independent of its external resemblance to either pure parent. is simply incidental or dependent on unknown differences in their reproductive systems. seems a strange arrangement. the male sexual 284 . in the facility of effecting an union. and then to stop their further propagation by different degrees of sterility.degree of fertility. for. 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. we see that when forms. that. that the fertility of hybrids is not related to the degree in which they resemble in external appearance either parent. both of first crosses and of hybrids. in reciprocal crosses between the same two species. is innately variable. there is generally some difference. according as the one species or the other is used as the father or the mother. which must be considered as good and distinct species. moreover. produced from reciprocal crosses often differ in fertility. their fertility graduates from zero to perfect fertility. it may even be asked. when various species are crossed. The hybrids. besides being eminently susceptible to favourable and unfavourable conditions. This latter statement is clearly proved by the difference in the result of reciprocal crosses between the same two species. the differences being of so peculiar and limited a nature. that their fertility. Considering the several rules now given. that it is by no means always the same in degree in the first cross and in the hybrids produced from this cross. not strictly related to the facility of the first union between their parents. 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. has the production of hybrids been permitted? To grant to species the special power of producing hybrids. or even to fertility under certain conditions in excess. and occasionally the widest possible difference. The foregoing rules and facts. 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 lastly. on the other hand. For why should the sterility be so extremely different in degree. which govern the fertility of first crosses and of hybrids. 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. are united. and other species cross with extreme difficulty.

in the period of the flow or nature of their sap. so it sometimes is in grafting. one being evergreen and the other deciduous. As Gartner found that there was sometimes an innate difference in different INDIVIDUALS of the same two species in crossing. on the other hand. Even different varieties of the pear take with different degrees of facility on the quince. can usually. cannot be grafted on the currant. The pear can be grafted far more readily on the quince. whereas the currant will take. The common gooseberry. But this capacity. is a different case from the difficulty of uniting two pure species.. I presume that no one will suppose that this capacity is a SPECIALLY endowed quality. which is a member of the same genus. be grafted with ease. the capacity is limited by systematic affinity. one being woody and the other herbaceous. As the capacity of one plant to be grafted or budded on another is unimportant for their welfare in a state of nature. the facility of effecting an union is often very far from equal. but in a multitude of cases we can assign no reason whatever. for no one has been able to graft together trees belonging to quite distinct families. so with grafting. and not a specially endowed quality. We can sometimes see the reason why one tree will not take on another from differences in their rate of growth. As in reciprocal crosses. Although many distinct genera within the same family have been grafted together. so Sagaret believes this to be the case with different individuals of the same two species in being grafted together. so do different varieties of the apricot and peach on certain varieties of the plum. but not in a reversed direction. than on the apple. in other cases species of the same genus will not take on each other. is by no means absolutely governed by systematic affinity. As in hybridisation. on the gooseberry. though with difficulty. It will be advisable to explain a little more fully. and. We have seen that the sterility of hybrids which have their reproductive organs in an imperfect condition. etc. in the hardness of their wood. Great diversity in the size of two plants. closely allied species and varieties of the same species. by an example. but will admit that it is incidental on differences in the laws of growth of the two plants. Something analogous occurs in grafting. what I mean by sterility being incidental on other differences. which have their reproductive organs perfect. for instance. yet these two distinct classes of cases run to a large extent parallel.element of the one will often freely act on the female sexual element of the other. which is ranked as a distinct genus. does not always prevent the two grafting together. as in hybridisation. but not invariably. for Thouin found that three 285 . and adaptation to widely different climates.

certain species of Sorbus. when thus grafted were rendered barren. 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. etc. to a certain extent. We thus see that.. Passiflora. We are reminded by this latter fact of the extraordinary cases of Hippeastrum. when grafted on other species. systematic affinity. 286 . although in the case of crossing. so I believe that the still more complex laws governing the facility of first crosses are incidental on unknown differences in their reproductive systems. yet that there is a rude degree of parallelism in the results of grafting and of crossing distinct species. by which term every kind of resemblance and dissimilarity between organic beings is attempted to be expressed. 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. 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. These differences in both cases follow. 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.species of Robinia. which seed much more freely when fertilised with the pollen of a distinct species than when fertilised with pollen from the same plant. yielded twice as much fruit as when on their own roots. as might have been expected. and which could be grafted with no great difficulty on a fourth species. which seeded freely on their own roots. On the other hand.

while at the same time the male element of this second form is enabled freely to fertilise the first form. In the first place. spontaneously appeared in certain individuals of one variety when crossed with those of another variety. now it could clearly have been of no advantage to such separated species to have been rendered mutually sterile. that.4. 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. will find the subject extraordinarily complex. when man is selecting at the same time two varieties. Take the case of any two species which. in rendering species mutually sterile. and consequently this could not have been effected through natural selection. for this peculiar state of the reproductive system could hardly have been advantageous to either species. for thus fewer bastardised and deteriorated offspring would be produced to commingle their blood with the new species in process of formation. when crossed. that in reciprocal crosses the male element of one form should have been rendered utterly impotent on a second form. which. After mature reflection. In the second place. from slightly lessened fertility to absolute sterility. but it may perhaps be argued. it seems to me that this could not have been effected through natural selection. it may be remarked that species inhabiting distinct regions are often sterile when crossed. Origin And Causes Of The Sterility Of First Crosses And Of Hybrids 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 fertility. For it would clearly be advantageous to two varieties or incipient species if they could be kept from blending. It may be admitted that it would profit an incipient species. if it were rendered in some slight degree sterile when crossed with its parent form or with some other variety. sterility with other species would follow as a necessary contingency. now. if a species was rendered sterile with some one compatriot. 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. the greatest difficulty will be found to lie in the existence of many graduated steps. In considering the probability of natural selection having come into action. as it has to others. it is almost as much opposed to the theory of natural selection as to that of special creation. produced few and sterile offspring. like any other variation. what is there which could favour the 287 . on the same principle that.

so that this acme of sterility. if rendered slightly sterile when crossed with some other variety. With sterile neuter insects we have reason to believe that modifications in their structure and fertility have been slowly accumulated by natural selection. but be incapable of causing an embryo to be developed. for the germen swells. must have incessantly occurred with many species. Both Gartner and Kolreuter have proved that in genera including numerous species. from an advantage having been thus indirectly given to the community to which they belonged over other communities of the same species. cannot have been gained through selection. the male element may reach the female element. they do not penetrate the stigmatic surface. as would be the case with a plant having a pistil too long for the pollen-tubes to reach the ovarium. and from the laws governing the various grades of sterility being so uniform throughout the animal and vegetable kingdoms. quite independent of natural selection. whatever it may be. but an individual animal not belonging to a social community. if the theory of natural selection be brought to bear. for a multitude are mutually quite barren.survival of those individuals which happened to be endowed in a slightly higher degree with mutual infertility. though the pollen-tubes protrude. and which thus approached by one small step towards absolute sterility? Yet an advance of this kind. It has also been observed that when the pollen of one species is placed on the stigma of a distantly allied species. There must sometimes be a physical impossibility in the male element reaching the ovule. 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. when the germen alone is effected. to species which never produce a single seed. a series can be formed from species which when crossed yield fewer and fewer seeds. would not thus itself gain any advantage or indirectly give any advantage to the other individuals of the same variety. the greater or less difficulty in effecting a union and in obtaining offspring apparently depends on several distinct causes. In the case of first crosses. we may infer that the cause. but yet are affected by the pollen of certain other species. It is here manifestly impossible to select the more sterile individuals. which have already ceased to yield seeds. Again. thus leading to their preservation. as seems to have been the case with some of Thuret's 288 . 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. is the same or nearly the same in all cases.

more especially as all very young beings are eminently sensitive to injurious or unnatural conditions of life. any more than why certain trees cannot be grafted on others. however. Until becoming acquainted with these facts." so that from the 500 eggs only twelve chickens were reared. 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. or had become nearly mature. at least it is known that hybrids raised from very distinct species are sometimes weak and dwarfed. that the early death of the embryo is a very frequent cause of sterility in first crosses. for hybrids. causing the embryo to be imperfectly developed. I was unwilling to believe in the frequent early death of hybrid embryos. are generally healthy and long-lived. hybridized embryos probably often perish in a like manner. the majority of these eggs had been fertilised. as we see in the case of the common mule. an embryo may be developed. apparently from mere inability to live. it may therefore. Hewitt. more than four-fifths died within the first few days. the embryos had either been partially developed and had then perished. or at latest weeks. 289 . and consequently be liable to perish at an early period. Lastly. and perish at an early age. when once born. as long as it is nourished within its mother's womb. Of the chickens which were born. But a hybrid partakes of only half of the nature and constitution of its mother. they are generally placed under suitable conditions of life. With plants. It may be here worth noticing that in some cases of parthenogenesis. the embryos within the eggs of silk moths which had not been fertilised. Mr. rather than in the conditions to which it is subsequently exposed. the cause more probably lies in some imperfection in the original act of impregnation. who has had great experience in hybridising pheasants and fowls. and in the majority of the fertilised eggs. are differently circumstanced before and after birth: when born and living in a country where their two parents live. but the young chickens had been unable to break through the shell. or within the egg or seed produced by the mother. "without any obvious cause. be exposed to conditions in some degree unsuitable. Hybrids. of which fact Max Wichura has recently given some striking cases with hybrid willows. but I believe.experiments on Fuci. No explanation can be given of these facts. and then perish at an early period. But after all. pass through their early stages of development and then perish like the embryos produced by a cross between distinct species. before birth. This latter alternative has not been sufficiently attended to. from observations communicated to me by Mr.

the external conditions have remained the same. when organic beings are placed during several generations under conditions not natural to them. In the one case. for their offspring in successive generations are eminently liable to vary. or any exotic plant seed freely under culture. is affected in a very similar manner. the case is somewhat different. without some disturbance occurring in the development. though in a lesser degree than when sterility ensues. In both cases the sterility occurs in various degrees. Lastly. in the other case. they are extremely liable to have their reproductive systems seriously affected. though often in so slight a degree as to be inappreciable by us. In both. or periodical action. independently of the general state of health. and when hybrids are produced by the unnatural crossing of two species. No one can tell till he tries. nor can he tell till he tries. For it is scarcely possible that two organisations should be compounded into one. or that of hybrids. I have more than once alluded to a large body of facts showing that. the male element is the most liable to be affected. On the other hand. whether any two species of a genus will produce more or less sterile hybrids. and whole groups of species tend to produce sterile hybrids. the tendency goes to a certain extent with systematic affinity. when animals and plants are removed from their natural conditions. in fact. they are extremely liable to vary. So it is with hybrids. including of course the reproductive systems. which seems to be partly due to their reproductive systems having been specially affected. there are many points of similarity. Thus we see that when organic beings are placed under new and unnatural conditions. having been blended into one. In both cases the sterility is independent of general health. When hybrids are able to breed inter se. for whole groups of animals and plants are rendered impotent by the same unnatural conditions. and is often accompanied by excess of size or great luxuriance. they transmit to their offspring from generation to generation the same 290 . in both. This. is the great bar to the domestication of animals. in which the sexual elements are imperfectly developed. Between the sterility thus superinduced and that of hybrids. the reproductive system. as every experimentalist has observed.In regard to the sterility of hybrids. the conditions of life have been disturbed. or mutual relations of the different parts and organs one to another or to the conditions of life. and certain species in a group will produce unusually fertile hybrids. one species in a group will sometimes resist great changes of conditions with unimpaired fertility. but the organisation has been disturbed by two distinct structures and constitutions. whether any particular animal will breed under confinement. but sometimes the female more than the male.

All that I have attempted to show is. this being generally the result. that slight crosses. it is even apt to increase. be owned that we cannot understand. But. or which have slightly varied. there is the clearest evidence that a cross between individuals of the same species. of too close interbreeding.compounded organisation. as we have seen. when subjected. or the increased sterility in those hybrids which occasionally and exceptionally resemble closely either pure parent. slight changes in the conditions of life benefit all organic beings. if these be kept under the same conditions of life. when placed under unnatural conditions. several facts with respect to the sterility of hybrids. however. the unequal fertility of hybrids produced from reciprocal crosses. does not diminish.. on the above or any other view. as under confinement. which I have elsewhere given. organic beings long habituated to certain uniform conditions under a state of nature. crosses between the males and females of the same species. is rendered sterile. and that close interbreeding continued during several generations between the nearest relations. and hence we need not be surprised that their sterility. that in two cases. weakness. in some respects allied. to a considerable change in their conditions. which differ to a certain extent. give vigour and fertility to the offspring. almost always leads to decreased size. gives vigour and fertility to the offspring. both with plants and animals. or sterility. sterility is the common result—in the one case from the conditions of life having been disturbed. Hence it seems that. that slight changes in the conditions of life are beneficial to all living things. that is. A similar parallelism holds good with an allied yet very different class of facts. It must. Nor do I pretend that the foregoing remarks go to the root of the matter: no explanation is offered why an organism. etc. and on the other hand. as before explained. great benefit is derived from almost any change in their habits of life. from one soil or climate to another. though in some degree variable. which have been subjected to slightly different conditions. Again. We see this acted on by farmers and gardeners in their frequent exchanges of seed. During the convalescence of animals. in the other case from the organisation having been disturbed by two organisations being compounded into one. tubers. founded on a considerable body of evidence. on the one hand. very frequently are rendered more 291 . for instance. The above view of the sterility of hybrids being caused by two constitutions being compounded into one has been strongly maintained by Max Wichura. and back again. It is an old and almost universal belief.

are incapable of breeding when kept under only partial confinement in their native country. I am fully persuaded that this double parallelism is by no means an accident or an illusion. He who is able to explain why the elephant.or less sterile. 292 . which is essentially related to the principle of life. which. are always tending towards an equilibrium. this principle. Herbert Spencer. will be able to explain the primary cause of hybrids being so generally sterile. as throughout nature. or consists in. being that life depends on. according to Mr. which would probably have been sterile if aboriginally crossed. the vital forces gain in power. and when this tendency is slightly disturbed by any change. which have often been subjected to new and not uniform conditions. and we know that a cross between two forms that have become widely or specifically different. although they are descended from distinct species. the incessant action and reaction of various forces. The above two parallel series of facts seem to be connected together by some common but unknown bond. are quite fertile together. produce hybrids which are almost always in some degree sterile. and a multitude of other animals. He will at the same time be able to explain how it is that the races of some of our domesticated animals.

that is by pollen taken from stamens not corresponding in height with the pistil. which exist in about equal numbers and which differ in no respect except in their reproductive organs. are fully fertile. and all the seedlings were similarly coloured. or more or less infertile. 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 as in each of the three forms there are two sets of stamens. and two. the two having differently sized pollen-grains. differs much in degree. and its own pollen be afterwards. 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. I ascertained this by fertilising several flowers. and twelve are illegitimate. first illegitimately. which may be called illegitimate. in the size and colour of the pollen-grains. even after a considerable interval of time. placed on the same stigma. had wholly destroyed 293 . It is well known that if pollen of a distinct species be placed on the stigma of a flower. Reciprocal Dimorphism And Trimorphism This subject may be here briefly discussed. are more or less infertile. so I have found it with illegitimate unions. though applied twenty-four hours subsequently. or fully fertile. for legitimate pollen is strongly prepotent over illegitimate pollen. which may be called legitimate. and in some other respects. its action is so strongly prepotent that it generally annihilates the effect of the foreign pollen. that in order to obtain full fertility with these plants. With trimorphic species six unions are legitimate. when both are placed on the same stigma. so it is with the pollen of the several forms of the same species. the other a short pistil with long stamens. So that with dimorphic species two unions. and the result has been confirmed by other observers. With trimorphic plants there are three forms likewise differing in the lengths of their pistils and stamens. The infertility which may be observed in various dimorphic and trimorphic plants. These organs are so proportioned in length to each other that half the stamens in two of the forms stand on a level with the stigma of the third form. the three forms possess altogether six sets of stamens and three kinds of pistils. when they are illegitimately fertilised. Several plants belonging to distinct orders present two forms. Now I have shown. one form having a long pistil with short stamens. and will be found to throw some light on hybridism. this shows that the legitimate pollen. with pollen taken from a peculiarly coloured variety. just in the same manner as occurs in crossing distinct species.5. up to absolute and utter sterility. and twenty-four hours afterwards legitimately.

But such is not the case. and from trimorphic plants all three illegitimate forms. there is occasionally a great difference in the result. in various degrees. 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. exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants. when united with each other in a legitimate manner. When this is done. the forms of the same undoubted species. when illegitimately united. produced within the limits of the same species by the improper union of certain forms. are not fully fertile. This led me carefully to observe during four years many seedlings. and yielded many seeds. while 294 . and are weak. These can then be properly united in a legitimate manner. They are all infertile. 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 parentspecies. miserable dwarfs. some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even seed-capsule. raised from several illegitimate unions. Lastly. It is hardly an exaggeration to maintain that illegitimate plants are hybrids. With hybrids raised from the same seed. both long-styled and short-styled illegitimate plants. Altogether there is the closest identity in character and behaviour between illegitimate plants and hybrids. while other and more sterile hybrids produce few flowers. for instance. and in others which might be added. a hybrid is crossed with either pure parent-species. behave in exactly the same manner as do two distinct species when crossed. on the other hand. so the same thing occurs with trimorphic plants.or prevented the action of the previously applied illegitimate pollen. so it is in a marked manner with illegitimate plants. It is possible to raise from dimorphic species. In all these respects.capsule the degree of sterility is innately variable. The chief result is that these illegitimate plants. the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. many hybrids are profuse and persistent flowerers. but the latter form did not yield a single seed when fertilised by the longer stamens of the mid-styled form. there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. Again. The sterility of these illegitimate plants. as in making reciprocal crosses between the same two species. as they may be called. while the sterility of the union from which they were derived was by no means great. If. so that sterility of certain illegitimate plants was unusually great. may be strictly compared with that of hybrids when crossed inter se.

that his two varieties were as good and as distinct species as any in the world. we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lythrum salicaria. thirdly. but I will not enlarge on this obscure subject. infer as probable from the consideration of dimorphic and trimorphic plants. doubtful whether this is really so.styled forms. first. and that they behaved in all other respects like ordinary hybrids. and yet be sterile when united in certain ways. Hence the case appears at first sight exactly the reverse of what occurs. We are also led to this same conclusion by 295 . either in structure or in constitution. however. is no safe criterion of specific distinction. and we are led to extend the same view to first crosses and hybrids. It is. But to make the case sure. secondly. We have also already seen that there is the closest similarity in all respects between first illegitimate unions and first crosses between distinct species. for instance. and this seems to me of especial importance. and not on any difference in their structure or general constitution. We may. He might then maintain that he had actually proved. depends exclusively on the nature of their sexual elements. but he would be completely mistaken. because we may conclude that there is some unknown bond which connects the infertility of illegitimate unions with that of their illegitimate offspring. because they show us. and that they behaved in all the other above specified respects as if they had been two distinct species. The facts now given on dimorphic and trimorphic plants are important. in the ordinary unions of the individuals of the same species and with crosses between distinct species. and he would find that the seedlings were miserably dwarfed and utterly sterile.ordinary hybrids are produced from an improper union between socalled distinct species. that the physiological test of lessened fertility. This will perhaps be made more fully apparent by an illustration. which results in sterility. For we must remember that it is the union of the sexual elements of individuals of the same form. in accordance with the common view. He would find that they yielded only about one-fifth of the proper number of seed. of two long. relatively to external conditions. that two or three forms of the same species may exist and may differ in no respect whatever. both in first crosses and in hybrids. he would raise plants from his supposed hybridised seed. because we find. that the sterility of distinct species when crossed and of their hybrid progeny. while it is the union of the sexual elements proper to two distinct forms which is fertile. however. and that he determined to try by crossing whether they were specifically distinct.

while the converse cross can be effected with perfect facility. likewise concluded that species when crossed are sterile owing to differences confined to their reproductive systems. in which the male of one species cannot be united. Gartner. or can be united with great difficulty. with the female of a second species.considering reciprocal crosses. 296 . That excellent observer.

they are at once ranked by most naturalists as species. differing widely from each other in appearance. I fully admit that this is the rule. Several considerations. which are considered by most botanists as varieties. produced. for instance. Not Universal It may be urged as an overwhelming argument that there must be some essential distinction between species and varieties inasmuch as the latter. presently to be given. if two forms hitherto reputed to be varieties be found in any degree sterile together. we are still involved in some doubt. more especially when we reflect how many species there are. are utterly sterile when intercrossed. so that the domesticated descendants of species. and yield perfectly fertile offspring. For when it is stated. are said by Gartner to be quite sterile when crossed. have had so little tendency towards modifying the reproductive system in a manner leading to mutual sterility. so that similar differences in the case of varieties would be no sure guide. those of the pigeon. or supposed to have been produced. and he consequently ranks them as undoubted species. or of the cabbage. And Of Their Mongrel Offspring. For instance. the fertility of all varieties produced under nature will assuredly have to be granted. With plants. and probably the true one. which. Now the varying conditions to which domesticated animals and cultivated plants have been subjected. looking to varieties produced under nature. In the first place. however. become perfectly fertile together. that certain South American indigenous domestic dogs do not readily unite with European dogs. under domestication. that we have good grounds for admitting the directly opposite doctrine of Pallas. render the fertility of domestic varieties less remarkable. for. however much they may differ from each other in external appearance. for instance. namely. the explanation which will occur to everyone. cross with perfect facility. Fertility Of Varieties When Crossed. is that they are descended from aboriginally distinct species.6. the blue and red pimpernel. it may be observed that the amount of external difference between two species is no sure guide to their degree of mutual sterility. It is certain that with species the cause lies exclusively in differences in their sexual constitution. though resembling each other most closely. If we thus argue in a circle. is a remarkable fact. that such conditions generally eliminate this tendency. With some exceptions. Nevertheless the perfect fertility of so many domestic races. so far is 297 . But the subject is surrounded by difficulties. which in their natural state probably would have been in some degree sterile when crossed. If we turn to varieties.

as I believe. why domestic varieties have not become mutually infertile when crossed. as soon as they have been permanently modified in a sufficient degree to take rank as species. when taken from their natural conditions and subjected to captivity. we can understand why. and being fertilised by. and it can hardly be rejected. The real difficulty in our present subject is not. than have domestic varieties. We are far from precisely knowing the cause. But we can see that species. on the other hand. while still retaining the capacity of fertilising. 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. The evidence is at least as good as that from which we believe in the sterility of a multitude of species. and why with plants only a few such cases. other species.cultivation from giving a tendency towards sterility between distinct species. that in several well. with species having a peculiar constitution.authenticated cases already alluded to. and this may well make a wide difference in the result. but why this has so generally occurred with natural varieties. for they have become self-impotent. seeing how profoundly ignorant we are in regard to the normal and abnormal action of the reproductive system. varieties have not been produced which are mutually sterile. will have been exposed during long periods of time to more uniform conditions. though in certain cases. are rendered sterile. have been observed. nor is this surprising. 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. For we know how commonly wild animals and plants. Domesticated productions.continued should likewise induce this tendency. as shown by the mere fact of their domestication. and which can now generally resist with undiminished fertility repeated changes of conditions. sterility might occasionally be thus caused. might be expected to produce varieties. as it appears to me. I have as yet spoken as if the varieties of the same species were invariably fertile when intercrossed. which I will briefly abstract. Thus. owing to their struggle for existence with numerous competitors. If the Pallasian doctrine of the elimination of sterility through long-continued domestication be admitted. with domesticated animals. immediately to be given. The evidence 298 . certain plants have been affected in an opposite manner. But it is impossible to resist the evidence of the existence of a certain amount of sterility in the few following cases. it becomes in the highest degree improbable that similar conditions long. were not originally highly sensitive to changes in their conditions of life. which.

which like the maize has separated sexes. he finds that the dissimilarly coloured varieties of the same species yield fewer seeds. and one variety can sometimes be raised from the seed of another. whose accuracy has been confirmed by every subsequent observer. more seed is produced by the crosses between the similarly coloured flowers. than the similarly coloured varieties. Yet these varieties differ in no respect. but only a single head produced any seed. Manipulation in this case could not have been injurious. and it is important to notice that the hybrid plants thus raised were themselves PERFECTLY fertile. but it is the result of an astonishing number of experiments made during many years on nine species of Verbascum. Moreover. and which he tested by the 299 . How far these experiments may be trusted. in the proportion of eighty-six to 100. has suspected that these varieties of maize are distinct species. they never naturally crossed. I know not. and seems at first incredible. He experimented on five forms which are commonly reputed to be varieties. Girou de Buzareingues crossed three varieties of gourd. Kolreuter. when yellow and white varieties of one species are crossed with yellow and white varieties of a DISTINCT species. as the plants have separated sexes. and Naudin has come to the same conclusion. a dwarf kind of maize with yellow seeds. except in the colour of their flowers. No one. The following case is far more remarkable. who mainly founds his classification by the test of infertility. as varieties. I believe. when crossed with a widely distinct species. than between those which are differently coloured. and he asserts that their mutual fertilisation is by so much the less easy as their differences are greater. and this one head produced only five grains. but the forms experimented on are ranked by Sagaret. that the yellow and white varieties when crossed produce less seed than the similarly coloured varieties of the same species. he asserts that. during several years. so that even Gartner did not venture to consider the two varieties as specifically distinct.is also derived from hostile witnesses. Gartner kept. and a tall variety with red seeds growing near each other in his garden. and although unable to confirm Gartner's results on the crossing of the distinct species. He then fertilised thirteen flowers of the one kind with pollen of the other. Mr. by so good an observer and so hostile a witness as Gartner: namely. and although these plants have separated sexes. Scott also has experimented on the species and varieties of Verbascum. who in all other cases consider fertility and sterility as safe criterions of specific distinction. has proved the remarkable fact that one particular variety of the common tobacco was more fertile than the other varieties.

when used either as the father or mother.severest trial. from man attending only to external characters in his domestic varieties. and crossed with the Nicotiana glutinosa. for a supposed variety. Hence the reproductive system of this one variety must have been in some manner and in some degree modified. But one of these five varieties. but as incidental on changes of an unknown nature in their sexual elements. 300 . by reciprocal crosses. glutinosa. would almost universally be ranked as a species. From the great difficulty of ascertaining the infertility of varieties in a state of nature. from these several considerations we may conclude that fertility does not constitute a fundamental distinction between varieties and species when crossed. and he found their mongrel offspring perfectly fertile. not as a special acquirement or endowment. The general sterility of crossed species may safely be looked at. always yielded hybrids not so sterile as those which were produced from the four other varieties when crossed with N. namely. and from such varieties not having been exposed for very long periods to uniform conditions of life. if proved to be infertile in any degree. From these facts it can no longer be maintained that varieties when crossed are invariably quite fertile.

but hybrids themselves have their reproductive systems seriously affected. 301 . For the parents of mongrels are varieties. When mongrels and the more fertile hybrids are propagated for several generations. The variability. and this implies that there has been recent variability. and this shows that the difference in the degree of variability graduates away. quite unimportant differences between the so-called hybrid offspring of species. Now. that in the first generation mongrels are more variable than hybrids. in contrast with that in the succeeding generations. but some few instances of both hybrids and mongrels long retaining a uniform character could be given. Hybrids And Mongrels Compared. I shall here discuss this subject with extreme brevity. This greater variability in mongrels than in hybrids does not seem at all surprising. and their descendants are highly variable.7. that the reproductive system. hybrids in the first generation are descended from species (excluding those long cultivated) which have not had their reproductive systems in any way affected. could find very few. they agree most closely in many important respects. and mostly domestic varieties (very few experiments having been tried on natural varieties). however. Gartner. and. from being eminently sensitive to changed conditions of life. whose strong wish it was to draw a distinct line between species and varieties. fails under these circumstances to perform its proper function of producing offspring closely similar in all respects to the parent-form. as it seems to me. an extreme amount of variability in the offspring in both cases is notorious. and the socalled mongrel offspring of varieties. the offspring of species and of varieties when crossed may be compared in several other respects. perhaps. and I have myself seen striking instances of this fact. but Gartner admits that hybrids from species which have long been cultivated are often variable in the first generation. greater than in hybrids. The most important distinction is. The slight variability of hybrids in the first generation. For it bears on the view which I have taken of one of the causes of ordinary variability. And. is a curious fact and deserves attention. namely. Independently Of Their Fertility Independently of the question of fertility. and they are not variable. in the successive generations of mongrels is. on the other hand. Gartner further admits that hybrids between very closely allied species are more variable than those from very distinct species. which would often continue and would augment that arising from the act of crossing.

Moreover. one has sometimes a prepotent power of impressing its likeness on the hybrid. is founded on a single experiment. the hybrids are widely different from each other. is certainly only a difference in degree. Hybrid plants produced from a reciprocal cross generally resemble each other closely. Gartner expressly states that the hybrids from long cultivated plants are more subject to reversion than hybrids from species in their natural state. So I believe it to be with varieties of plants. Both hybrids and mongrels can be reduced to either pure parent form. For instance. according to Gartner the same laws. are crossed with a third species. while Naudin. but more especially owing to prepotency in transmitting likeness running more strongly in one sex than in the other. These several remarks are apparently applicable to animals. both when one species is crossed with another and when one variety is crossed with another variety. Thus Max Wichura doubts whether hybrids ever revert to their parent forms. On the other hand. so that both the mule and the hinny resemble more closely the ass than the horse. the degrees and kinds of resemblance in mongrels and in hybrids to their respective parents. one variety certainly often has this prepotent power over another variety. and he experimented chiefly on cultivated plants. and so it is with mongrel plants from a reciprocal cross. but that the prepotency runs more strongly in the male than in the 302 . Gartner further states that when any two species. more especially in hybrids produced from nearly related species. but this. whereas if two very distinct varieties of one species are crossed with another species. but the subject is here much complicated.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. and with animals. if it be true. the hybrids do not differ much. partly owing to the existence of secondary sexual characters. insists in the strongest terms on the almost universal tendency to reversion in hybrids. on the other hand. and he experimented on uncultivated species of willows. and seems directly opposed to the results of several experiments made by Kolreuter. Such alone are the unimportant differences which Gartner is able to point out between hybrid and mongrel plants. But this conclusion. When two species are crossed. I think those authors are right who maintain that the ass has a prepotent power over the horse. by repeated crosses in successive generations with either parent. follow. as far as I can make out. and this probably explains the singular difference in the results arrived at by different observers. although most closely allied to each other.

melanism. which is the offspring of the female-ass and stallion. Prosper Lucas. that it is only with mongrels that the offspring are not intermediate in character. Independently of the question of fertility and sterility. which are descended from varieties often suddenly produced and semi-monstrous in character. which is an offspring of the male ass and mare. or additional fingers and toes.female ass. On the whole. deficiency of tail or horns. but this does sometimes occur with hybrids. and of crossed varieties. comes to the conclusion that the laws of resemblance of the child to its parents are the same. than with hybrids. Looking to the cases which I have collected of cross-bred animals closely resembling one parent. but closely resemble one of their parents. so that the mule. which are descended from species slowly and naturally produced. be much more likely to occur with mongrels. who. or of distinct species. and do not relate to characters which have been slowly acquired through selection. If we look at species as having been specially created. in the union of individuals of the same variety. whether the two parents differ little or much from each other. this similarity would be an astonishing fact. But it harmonises perfectly with the view that there is no essential distinction between species and varieties. and which have suddenly appeared—such as albinism. or of different varieties. namely. after arranging an enormous body of facts with respect to animals. yet I grant much less frequently than with mongrels. the resemblances seem chiefly confined to characters almost monstrous in their nature. is more like an ass than is the hinny. in all other respects there seems to be a general and close similarity in the offspring of crossed species. Much stress has been laid by some authors on the supposed fact. 303 . A tendency to sudden reversions to the perfect character of either parent would. I entirely agree with Dr. also. and at varieties as having been produced by secondary laws.

that the crossing of forms. but is governed by several curious and complex laws. In the case of first crosses it seems to depend on several circumstances. the capacity in one species or variety to take on another. and sometimes widely different in reciprocal crosses between the same two species. and secondly. and is often so slight that the most careful experimentalists have arrived at diametrically opposite conclusions in ranking forms by this test. generally of an unknown nature. sterile. which have been exposed to slightly different conditions of life. In the same manner as in grafting trees. slight changes in the conditions of life add to the vigour and fertility of all organic beings. firstly. is incidental on differences. The sterility of first crosses and of their hybrid progeny has not been acquired through natural selection. the sterility being closely allied to that which so frequently affects pure species. The sterility is of all degrees. It is generally different. or which have varied. that. sufficiently distinct to be ranked as species. when exposed to new and unnatural conditions of life.8. 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. The degree of sterility does not strictly follow systematic affinity. The facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny. In the case of hybrids. The sterility is innately variable in individuals of the same species. He who will explain these latter cases will be able to explain the sterility of hybrids. so in crossing. but not universally. the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. it apparently depends on their whole organisation having been disturbed by being compounded from two distinct forms. favours the size. and their hybrids. and is eminently susceptible to action of favourable and unfavourable conditions. Summary Of Chapter First crosses between forms. in their vegetative systems. This view is strongly supported by a parallelism of another kind: namely. 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. vigour and fertility of their offspring. are very generally. in some instances in chief part on the early death of the embryo. It is not always equal in degree in a first cross and in the hybrids produced from this cross. perhaps render it probable that some unknown bond in all cases connects the degree of fertility of first unions with that 304 .

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. should in most cases correspond. we do not know. Nor is this almost universal and perfect fertility surprising. First crosses between forms known to be varieties. parallel with the systematic affinity of the forms subjected to experiment. 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. and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences. but not. 305 . But why. when it is remembered how liable we are to argue in a circle with respect to varieties in a state of nature. The consideration of these facts on dimorphism. to a certain extent. invariably fertile. in the case of distinct species. yet the facts given in this chapter do not seem to me opposed to the belief that species aboriginally existed as varieties. and the sterility of their hybrid offspring. Nor is it surprising that the facility of effecting a first cross. and the capacity of being grafted together—though this latter capacity evidently depends on widely different circumstances—should all run. in all other respects there is the closest general resemblance between hybrids and mongrels. It should also be especially kept in mind. in their variability.of their offspring. even if due to distinct causes: for both depend on the amount of difference between the species which are crossed. that longcontinued domestication tends to eliminate sterility. the sexual elements should so generally have become more or less modified. and the fertility of the hybrids thus produced. clearly leads to the conclusion that the primary cause of the sterility of crossed species is confined to differences in their sexual elements. leading to their mutual infertility. and that they have not been long exposed to uniform conditions of life. and their mongrel offspring. as well as of the results of reciprocal crosses. Independently of the question of fertility. in their power of absorbing each other by repeated crosses. as is so often stated. are very generally. Finally. then. It is not surprising that the difficulty in crossing any two species. or sufficiently alike to be considered as varieties. for systematic affinity includes resemblances of all kinds. and is therefore little likely to induce this same quality. and in their inheritance of characters from both parent-forms.

on their number — On the lapse of time. 306 . 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.Chapter 10 On The Imperfection Of The Geological Record On the absence of intermediate varieties at the present day — On the nature of extinct intermediate varieties.

on the very process of natural selection. I endeavoured. that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. through which new varieties continually take the places of and supplant their parent-forms. to show that intermediate varieties. of innumerable intermediate links not now occurring everywhere throughout nature depends. in the extreme imperfection of the geological record. namely. is a very obvious difficulty. that the life of each species depends in a more important manner on the presence of other already defined organic forms. I assigned reasons why such links do not commonly occur at the present day under the circumstances apparently most favourable for their presence. To give a simple illustration: the fantail and pouter pigeons are both descended from the rock-pigeon. is the most obvious and serious objection which can be urged against my theory. if we possessed all the intermediate varieties which have ever existed. Most of them have now been discussed. for 307 . and this. perhaps. be truly enormous. In the first place. none. In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. 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. we should always look for forms intermediate between each species and a common but unknown progenitor. The main cause. which have formerly existed. to avoid picturing to myself forms DIRECTLY intermediate between them. I have found it difficult. than on climate. so must the number of intermediate varieties. One. on the theory. The explanation lies.1. we should have an extremely close series between both and the rock-pigeon. I endeavoured to show. have formerly existed. on an extensive and continuous area with graduated physical conditions. and the progenitor will generally have differed in some respects from all its modified descendants. the distinctness of specific forms and their not being blended together by innumerable transitional links. when looking at any two species. and. from existing in lesser numbers than the forms which they connect. will generally be beaten out and exterminated during the course of further modification and improvement. But this is a wholly false view. but we should have no varieties directly intermediate between the fantail and pouter. namely. it should always be borne in mind what sort of intermediate forms must. But just in proportion as this process of extermination has acted on an enormous scale. as I believe. also. however.

now generally extinct. have become so much modified. such have lived upon the earth. have in their turn been similarly connected with more ancient forms. But such a case would imply that one form had remained for a very long period unaltered. it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon. we have no reason to suppose that links directly intermediate between them ever existed. By the theory of natural selection all living species have been connected with the parent-species of each genus. that one of two living forms might have descended from the other. we should be unable to recognise the parent-form of any two or more species. such as C. combining a tail somewhat expanded with a crop somewhat enlarged. even perhaps more than they differ from each other. between child and parent. So that the number of intermediate and transitional links. if we look to forms very distinct. whether they had descended from this species or from some other allied species. if we had no historical or indirect evidence regarding their origin. unless at the same time we had a nearly perfect chain of the intermediate links. So with natural species. for in all cases the new and improved forms of life tend to supplant the old and unimproved forms. and these parent-species. will render this a very rare event. must have been inconceivably great. a horse from a tapir. and so on backwards. between all living and extinct species. moreover.instance. by differences not greater than we see between the natural and domestic varieties of the same species at the present day. C. always converging to the common ancestor of each great class. in all such cases. It is just possible. even if we closely compared the structure of the parent with that of its modified descendants. But assuredly. These two breeds. 308 . oenas. that. the characteristic features of these two breeds. for instance to the horse and tapir. whilst its descendants had undergone a vast amount of change. but between each and an unknown common parent. The common parent will have had in its whole organisation much general resemblance to the tapir and to the horse. for instance. by the theory. and the principle of competition between organism and organism. but in some points of structure may have differed considerably from both. livia. and in this case DIRECT intermediate links will have existed between them. if this theory be true. Hence.

and yet does not admit how vast have been the past periods of time. or mud. At last the base of the cliff is undermined. or even of each stratum. along a short length or round a promontory. The appearance of the surface and 309 . huge fragments fall down. all thickly clothed by marine productions. and the waves eat into them only when they are charged with sand or pebbles. It is hardly possible for me to recall to the reader who is not a practical geologist. and these remaining fixed. Therefore a man should examine for himself the great piles of superimposed strata. and learning how deeply the surface of the land has been denuded. and to mark how each author attempts to give an inadequate idea of the duration of each formation. if we follow for a few miles any line of rocky cliff. It is good to wander along the coast. We can best gain some idea of past time by knowing the agencies at work. But how often do we see along the bases of retreating cliffs rounded boulders. and then they are more quickly ground into pebbles. showing how little they are abraded and how seldom they are rolled about! Moreover.2. and watch the rivulets bringing down mud. He who can read Sir Charles Lyell's grand work on the Principles of Geology. that the cliffs are at the present time suffering. and how much sediment has been deposited. and mark the process of degradation. and the waves wearing away the sea-cliffs. 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. the facts leading the mind feebly to comprehend the lapse of time. when formed of moderately hard rocks. all changes having been effected slowly. until after being reduced in size they can be rolled about by the waves. Not that it suffices to study the Principles of Geology. which is undergoing degradation. As Lyell has well remarked. sand. we find that it is only here and there. As Inferred From The Rate Of Deposition And Extent Of Denudation Independently of our not finding fossil remains of such infinitely numerous connecting links. On The Lapse Of Time. it may be objected that time cannot have sufficed for so great an amount of organic change. or to read special treatises by different observers on separate formations. may at once close this volume. for there is good evidence that pure water effects nothing in wearing away rock. the monuments of which we see all around us. in order to comprehend something about the duration of past time. have to be worn away atom by atom. which the future historian will recognise as having produced a revolution in natural science. The tides in most cases reach the cliffs only for a short time twice a day.

for each line is composed of one and the same formation. in order to appreciate the past duration of time. and triturate the fragments. on the one hand. and in colder countries to frost. due to their formerly liquid state. Ramsay and Whitaker have shown. we are compelled to admit that the escarpments owe their origin in chief part to the rocks of which they are composed. I remember having been much struck when viewing volcanic islands. according to our ideas of time. or the power of the waves. The same story is told still more plainly by faults—those great cracks along which the strata have been upheaved on one side. that the great lines of escarpment in the Wealden district and those ranging across England. Geikie. This being the case. which have been worn by the waves and pared all round into perpendicular cliffs of one or two thousand feet in height. recently learned from the observations of Ramsay. showed at a glance how far the hard. and to a greater extent than might be supposed. by the wind. when rapid deepen their channels. while our sea-cliffs are everywhere formed by the intersection of various formations. it is then transported by the streams and rivers. in the van of many excellent observers—of Jukes. rocky beds had once extended into the open ocean. this surface consequently has been gradually lowered.the vegetation show that elsewhere years have elapsed since the waters washed their base. On a rainy day. even in a gently undulating country. We have. to consider. we see the effects of subaerial degradation in the muddy rills which flow down every slope. which formerly were looked at as ancient seacoasts. and which seem to work so slowly. which. Nothing impresses the mind with the vast duration of time. and on the other hand the thickness of our sedimentary formations. for the gentle slope of the lavastreams. When thus impressed with the slow rate at which the land is worn away through subaerial and littoral action. which apparently have so little power. having resisted subaerial denudation better than the surrounding surface. the disintegrated matter is carried down even gentle slopes during heavy rain. however. or thrown down on the 310 . Messrs. that subaerial degradation is a much more important agency than coast-action. with the lines of harder rock left projecting. The whole surface of the land is exposed to the chemical action of the air and of the rainwater. with its dissolved carbonic acid. more forcibly than the conviction thus gained that subaerial agencies. especially in arid districts. cannot have been thus formed. have produced great results. and the observation is a most striking one. Croll and others. it is good. the masses of rock which have been removed over many extensive areas.

in an interesting paper. and although conglomerates have probably been accumulated at a quicker rate than finer sediments. So that the lofty pile of sedimentary rocks in Britain gives but an inadequate idea of the time which has elapsed during their accumulation. In the Cordillera. blank periods of enormous length. which are represented in England by thin beds. from actual measurement in most cases. for instance. When geologists look at large and complicated phenomena. Mr. 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. Some of these formations. are thousands of feet in thickness on the Continent. as most geologists now believe. was slow and effected by many starts. in the opinion of most geologists. Moreover.154 Secondary strata… … … … … … … … … … .240 —making altogether 72. Nevertheless this impression is partly false. I estimated one mass of conglomerate at ten thousand feet. Croll. yet in these cases there is nothing on the surface of the land to show such prodigious movements. Professor Ramsay has published an account of a downthrow in Anglesea of 2.000 feet. and along this line the vertical displacement of the strata varies from 600 to 3. in all parts of the world the piles of sedimentary strata are of wonderful thickness. remarks that we do not err "in forming too great a conception of the length of geological periods.2. very nearly thirteen and threequarters British miles. the pile of rocks on either side of the crack having been smoothly swept away. On the other hand. they are good to show how slowly the mass must have been heaped together. to the height or depth of thousands of feet. Professor Ramsay has given me the maximum thickness. and then at the figures representing several million years. Croll shows." but in estimating them by years. each of which bears the stamp of time..13. for since the crust cracked. and it makes no great difference whether the upheaval was sudden. the two produce a totally different effect on the mind. the surface of the land has been so completely planed down that no trace of these vast dislocations is externally visible. Mr. The Craven fault. and the figures are at once pronounced too small.other. and this is the result: Feet Palaeozoic strata (not including igneous beds).. or.190 Tertiary strata… … … … … … … … … … … .584 feet. of the successive formations in DIFFERENT parts of Great Britain. yet from being formed of worn and rounded pebbles.57. between each successive formation we have. by calculating the 311 . extends for upward of thirty miles.300 feet. and he informs me that he fully believes that there is one in Merionethshire of 12.000 feet. that is. In regard to subaerial denudation.

Croll gives the following illustration: Take a narrow strip of paper. or through the immigration of new forms. But let it be borne in mind. that is. Moreover. according to the standard of years. and within the same country only a few change at the same time. know what a million really means: Mr. with no intention of modifying the breed. This seems an astonishing result. but to the subject of time we must return. Several eminent breeders. Few men have attended with due care to any one strain for more than half a century. but by this process of unconscious selection. but if halved or quartered it is still very surprising. eighty-three feet four inches in length. have so largely modified some of the higher animals. by which some of the inhabitants might be better fitted to their new places under the altered circumstance. however. so that a hundred years represents the work of two breeders in succession. how long a period it takes to modify a species. during a single lifetime.known amount of sediment annually brought down by certain rivers. and stretch it along the wall of a large hall. would thus be removed from the mean level of the whole area in the course of six million years. This tenth of an inch will represent one hundred years. Few of us. that new places in the polity of nature do not occur until after long intervals. that 1. 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. however. Species. 312 . relatively to their areas of drainage. and the entire strip a million years. the preservation of the most useful or beautiful animals. that they have formed what well deserves to be called a new sub-breed. due to the occurrence of physical changes of some kind. The comparison would be in every way fairer with the effects which follow from unconscious selection. Unfortunately we have no means of determining. probably change much more slowly. which propagate their kind much more slowly than most of the lower animals. would not always occur at once. various breeds have been sensibly changed in the course of two or three centuries. variations or individual differences of the right nature. as it became gradually disintegrated. then mark off at one end the tenth of an inch.000 feet of solid rock. represented as it is by a measure utterly insignificant in a hall of the above dimensions. in relation to the subject of this work. and some considerations lead to the suspicion that it may be too large. This slowness follows from all the inhabitants of the same country being already so well adapted to each other. what a hundred years implies.

with the exception of a single Mediterranean species. many great deposits. occasionally even six thousand feet in thickness. and although this great mass has been most carefully searched. at a rate sufficiently quick to embed and preserve fossil remains. For instance. Throughout an enormously large proportion of the ocean. no fossils. 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 the beds are upraised. seem explicable only on the view of the bottom of the sea not rarely lying for ages in an unaltered condition. The remains which do become embedded. by another and later formation. which inhabits deep water and this has been found fossil in Sicily. and extending for at least 300 miles from Vienna to Switzerland. Edward Forbes. after an immense interval of time. where sediment is not accumulating. The remark of that admirable palaeontologist. and what a paltry display we behold! That our collections are imperfect is admitted by every one. without the underlying bed having suffered in the interval any wear and tear.3. We probably take a quite erroneous view. Some of the many kinds of animals which live on the beach between high and low water mark seem to be rarely preserved. without our being able to assign any reason: one of the most striking instances is that of the Flysch formation. have been found. namely. 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. The many cases on record of a formation conformably covered. will. No organism wholly soft can be preserved. the bright blue tint of the water bespeaks its purity. Only a small portion of the surface of the earth has been geologically explored. On The Poorness Of Palaeontological Collections Now let us turn to our richest museums. should never be forgotten. requiring a vast length of time for their accumulation. which consists of shale and sandstone. Lastly. when we assume that sediment is being deposited over nearly the whole bed of the sea. generally be dissolved by the percolation of rain water charged with carbonic acid. or from a few specimens collected on some one spot. if in sand or gravel. that very many fossil species are known and named from single and often broken specimens. several thousand. and no part with sufficient care. as the important discoveries made every year in Europe prove. Shells and bones decay and disappear when left on the bottom of the sea. 313 . are entirely destitute of organic remains. except a few vegetable remains.

entirely disbelieve in the change of species. But the imperfection in the geological record largely results from another and more important cause than any of the foregoing.With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods. when we remember how large a proportion of the bones of tertiary mammals have been discovered either in caves or in lacustrine deposits. a glance at the historical table published in Lyell's Manual. we may infer that this could nowhere be ascertained. had elsewhere been accumulated. When we see the formations tabulated in written works. see why the geological formations of each region are almost invariably intermittent. so it is in North America. Scarcely any fact struck me more when examining many hundred miles of the South American coasts. Murchison's great work on Russia. which have been upraised 314 . whence the sediment was derived. and in many other parts of the world. until recently not a land-shell was known belonging to either of these vast periods. Nor is their rarity surprising. what wide gaps there are in that country between the superimposed formations. I think. will bring home the truth. generally implying great changes in the geography of the surrounding lands. hardly any idea can be formed of the length of time which has elapsed between the consecutive formations. it is difficult to avoid believing that they are closely consecutive. with the exception of one species discovered by Sir C. Dawson in the carboniferous strata of North America. if his attention had been confined exclusively to these large territories. For instance. accord with the belief of vast intervals of time having elapsed between each formation. have not followed each other in close sequence. The most skilful geologist. And if. Forbes. charged with new and peculiar forms of life. great piles of sediment. for instance. but now land-shells have been found in the lias. like E. This doctrine has been emphatically admitted by many geologists and palaeontologists. or when we follow them in nature. that is. from Sir R. Lyell and Dr. would never have suspected that during the periods which were blank and barren in his own country. namely. it is superfluous to state that our evidence is fragmentary in an extreme degree. far better than pages of detail. from the several formations being separated from each other by wide intervals of time. and that not a cave or true lacustrine bed is known belonging to the age of our secondary or palaeozoic formations. in every separate territory. how accidental and rare is their preservation. But we know. We can. who. In regard to mammiferous remains. The frequent and great changes in the mineralogical composition of consecutive formations.

and have been surprised to note how author after author. conclude that sediment must be accumulated in extremely thick. I think. solid. in order to withstand the incessant action of the waves. The explanation. that the only ancient tertiary formation on the west coast of South America. We may. but which will hardly last to a 315 . than the absence of any recent deposits sufficiently extensive to last for even a short geological period. when upraised. and the mass when upraised will give an imperfect record of the organisms which existed in the neighbourhood during the period of its accumulation. I may add. Along the whole west coast. no extensive formations with recent or tertiary remains can anywhere be found. I am convinced that nearly all our ancient formations. though the supply of sediment must for ages have been great. the sea will remain shallow and favourable for many and varied forms. Such thick and extensive accumulations of sediment may be formed in two ways. tertiary beds are so poorly developed that no record of several successive and peculiar marine faunas will probably be preserved to a distant age. Since publishing my views on this subject in 1845. in which case the bottom will not be inhabited by so many and such varied forms of life as the more shallow seas. from the enormous degradation of the coast rocks and from the muddy streams entering the sea. no doubt. is that the littoral and sub-littoral deposits are continually worn away. Or sediment may be deposited to any thickness and extent over a shallow bottom. thick enough. have thus been formed during subsidence. I have watched the progress of geology.several hundred feet within the recent period. and thus a rich fossiliferous formation. to resist a large amount of denudation. A little reflection will explain why. when first upraised and during subsequent oscillations of level. which has been bulky enough to resist such degradation as it has as yet suffered. which is inhabited by a peculiar marine fauna. as soon as they are brought up by the slow and gradual rising of the land within the grinding action of the coast-waves. as long as the rate of subsidence and supply of sediment nearly balance each other. may be formed. if it continue slowly to subside. In this latter case. as well as the subsequent subaerial degradation. or extensive masses. in treating of this or that great formation. either in profound depths of the sea. along the rising coast of the western side of South America. has come to the conclusion that it was accumulated during subsidence. which are throughout the greater part of their thickness RICH IN FOSSILS.

that if one part of the area. mica-schist. Still less can this have happened during the alternate periods of elevation. Hopkins also expresses his belief that sedimentary beds of considerable horizontal extent have rarely been completely destroyed. Admitting then that gneiss. but the thickness of the formation could not be great. which are the most favourable to life. however. were once 316 . and yet not suffer excessively from denudation during its slow upheaval. a widely extended formation might be formed during a period of elevation. the beds which were then accumulated will generally have been destroyed by being upraised and brought within the limits of the coast-action. after rising and before being denuded. diorite. and thus gained considerable thickness. as long as the bed of the sea remained stationary. might afterwards become protected by fresh accumulations. but if the metamorphic action occurred at profound depths of the ocean. excepting the few who believe that our present metamorphic schists and plutonic rocks once formed the primordial nucleus of the globe. and thus be preserved for a long period. granite. such as that within a large part of the Malay Archipelago. All geological facts tell us plainly that each area has undergone numerous slow oscillations of level. the former protecting mantle of rock may not have been very thick. nor would the deposit be much consolidated. 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. though not thick. Consequently. THICK deposits cannot have been accumulated in the shallow parts. Mr. will admit that these latter rocks have been stripped of their covering to an enormous extent. On the other hand. formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation. will have been formed over wide spaces during periods of subsidence. or.. Hopkins. subsided. 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 the case of an extensive and shallow sea. 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 apparently these oscillations have affected wide spaces. to speak more accurately. For it is scarcely possible that such rocks could have been solidified and crystallised while uncovered.distant geological age. where the depth varies from thirty or forty to sixty fathoms. These remarks apply chiefly to littoral and sublittoral deposits. nor be capped by overlying formations. It has. But all geologists. was deposited during a downward oscillation of level. etc. been suggested by Mr.

but from the concurrent testimony of travellers. from near Rio de Janeiro to the mouth of the Plata. as shown in Professor H. the inhabited area and number of inhabitants will decrease (excepting on the shores of a continent when first broken up into an archipelago). I saw. and it is during these very 317 . 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. and the British Islands. the granitic area is very large: thus Von Eschwege gives a detailed section of these rocks. with not a wreck left behind. besides modern tertiary beds. to the United States and Canada.D. during subsidence. it is probable that in some parts of the world whole formations have been completely denuded. few new varieties or species will be formed. In many regions the metamorphic and granite rocks would be found much more widely extended than they appear to be. if all the sedimentary beds were removed which rest unconformably on them. and saw nothing but granitic rocks.100 geographical miles. On the other hand. how can we account for the naked and extensive areas of such rocks in many parts of the world. a distance of 1. the whole of the newer Palaeozoic formations. though there will be much extinction. collected along the whole coast. Boue colours an area composed of rocks of this nature as equal to that of Spain. and consequently during subsidence. Hence. along the whole northern bank of the Plata. only one small patch of slightly metamorphosed rock.necessarily covered up. Numerous specimens. Italy. in the proportion of 19 to 12. One remark is here worth a passing notice.5. South of the Amazon. During periods of elevation the area of the land and of the adjoining shoal parts of the sea will be increased and new stations will often be formed—all circumstances favourable. and which could not have formed part of the original mantle under which they were crystallised. but during such periods there will generally be a blank in the geological record. Inland. I have estimated the areas by cutting out and weighing the paper. for the formation of new varieties and species. France. namely. stretching from Rio de Janeiro for 260 geographical miles inland in a straight line. Turning to a wellknown region. as previously explained. which alone could have formed a part of the original capping of the granitic series. part of Germany. and they all belonged to this class. all conjoined. and I find that the metamorphic (excluding the "semi-metamorphic") and granite rocks exceed. This region has not been carefully explored. and I travelled for 150 miles in another direction. were examined by me. Rogers' beautiful map.

periods of subsidence that the deposits which are richest in fossils have been accumulated. 318 .

is extremely imperfect. and when we see a species first appearing in any formation. it would be rash in the extreme to infer that it had not elsewhere previously existed. whose opinions are worthy of much deference. 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. I am aware that two palaeontologists. On The Absence Of Numerous Intermediate Varieties In Any Single Formation From these several considerations it cannot be doubted that the geological record. 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. When we see a species first appearing in the middle of any formation. 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. viewed as a whole. It is well known. it would be equally rash to suppose that it then became extinct. Although each formation may mark a very long lapse of years. But insuperable difficulties. for instance. namely Bronn and Woodward. prevent us from coming to any just conclusion on this head. We forget how small the area of Europe is compared with the rest of the world. Although each formation has indisputably required a vast number of years for its deposition. when we find a species disappearing before the last layers have been deposited. time having apparently been required for their migration from the American to the European 319 .4. Several cases are on record of the same species presenting varieties in the upper and lower parts of the same formation. 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. but I cannot assign due proportional weight to the following considerations. but if we confine our attention to any one formation. have concluded that the average duration of each formation is twice or thrice as long as the average duration of specific forms. Thus Trautschold gives a number of instances with Ammonites. the probability is that it only then first immigrated into that area. as it seems to me. So again. each probably is short compared with the period requisite to change one species into another. that several species appear somewhat earlier in the palaeozoic beds of North America than in those of Europe. nor have the several stages of the same formation throughout Europe been correlated with perfect accuracy.

and the species undergoing change must have lived in the same district throughout the whole time. but have become extinct in the immediately surrounding sea. probable that sediment was deposited during the whole of the glacial period near the mouth of the Mississippi. on the extreme change of climate. for instance. 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. conversely. But we have seen that a thick formation. hence. In examining the latest deposits. It is an excellent lesson to reflect on the ascertained amount of migration of the inhabitants of Europe during the glacial epoch. all included within this same glacial period. And in the distant future. examining these beds. INCLUDING FOSSIL REMAINS. and to keep the depth approximately the same. and on the great lapse of time. which is necessary that the same marine species may live on the same space. and likewise to reflect on the changes of level. fossiliferous throughout its entire thickness. which forms only a part of one whole geological period. the deposit must be a very thick one. in various quarters of the world. that some are now abundant in the neighbouring sea. sedimentary deposits. Yet it may be doubted whether. that some few still existing species are common in the deposit.seas. It is not. the supply of sediment must nearly counterbalance the amount of subsidence. organic remains will probably first appear and disappear at different levels. sufficient for the slow process of modification. or. would be tempted to conclude that the average duration of life of the embedded fossils had been less than that of the glacial period. this nearly exact balancing between the supply of sediment and the amount of subsidence is probably a rare contingency. but are rare or absent in this particular deposit. in any quarter of the world. extending from before the glacial epoch to the present day. the deposit must have gone on continuously accumulating during a long period. and thus diminish the supply. have gone on accumulating within the same area during the whole of this period. it has everywhere been noted. owing to the migrations of species and to geographical changes. whilst the downward movement continues. But this same movement of subsidence will tend to submerge the area whence the sediment is derived. a geologist. that is. In fact. In order to get a perfect gradation between two forms in the upper and lower parts of the same formation. instead of having been really far greater. 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. can accumulate only during a period of subsidence. for it has been observed by more 320 .

a formation composed of beds of widely different mineralogical composition. It is all-important to remember that naturalists have no golden rule by which to distinguish species and varieties. and this. showing what wide. we can seldom hope to effect in any one geological section. only a few feet in thickness. with ancient root-bearing strata. but abrupt. which would not have been suspected. we may reasonably suspect that the process of deposition has been more or less interrupted. even if 321 . except near their upper or lower limits. It would seem that each separate formation. intervals have occurred in its accumulation. Many instances could be given of beds. as is so often the case.than one palaeontologist that very thick deposits are usually barren of organic remains. Hence. middle. In other cases we have the plainest evidence in great fossilised trees. they rank both as species. Many cases could be given of the lower beds of a formation having been upraised. yet no one ignorant of this fact would have even suspected the vast lapse of time represented by the thinner formation. they grant some little variability to each species. yet easily overlooked. unless they are enabled to connect them together by the closest intermediate gradations. when the same species occurs at the bottom. but has disappeared and reappeared. the probability is that it has not lived on the same spot during the whole period of deposition. perhaps many times. changes of form.400 feet thick in Nova Scotia. though perhaps slight. a section would not include all the fine intermediate gradations which must on our theory have existed. like the whole pile of formations in any country. and which must have required an enormous period for their accumulation. A. but when they meet with a somewhat greater amount of difference between any two forms. Lyell and Dr. and top of a formation. to be found in an older and underlying bed. from the reasons just assigned. representing formations which are elsewhere thousands of feet in thickness. Supposing B and C to be two species. during the same geological period. Consequently if it were to undergo a considerable amount of modification during the deposition of any one geological formation. of many long intervals of time and changes of level during the process of deposition. had not the trees been preserved: thus Sir C. When we see. has generally been intermittent in its accumulation. and a third. submerged. Dawson found carboniferous beds 1. still standing upright as they grew. denuded. and then re-covered by the upper beds of the same formation—facts. at no less than sixty-eight 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. one above the other.

so that here again we have undoubted evidence of change in the direction required by the theory. maintain that all these tertiary species are specifically distinct. that A might be the actual progenitor of B and C.A were strictly intermediate between B and C. 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. unless we believe that these eminent naturalists have been misled by their imaginations. as before explained. If we look to rather wider intervals of time. unless at the same time it could be closely connected by intermediate varieties with either one or both forms. that these tertiary species are all truly distinct from the recent. and unless we obtained numerous transitional gradations. though the distinction is admitted to be very slight. and yet would not necessarily be strictly intermediate between them in all respects. It is notorious on what excessively slight differences many palaeontologists have founded their species. Some experienced conchologists are now sinking many of the very fine species of D'Orbigny and others into the rank of varieties. that their varieties are generally at first local. though universally ranked as specifically different. and they do this the more readily if the specimens come from different sub-stages of the same formation. Look again at the later tertiary deposits. we have evidence of the frequent occurrence of slight modifications of the kind required. or unless we admit. it would simply be ranked as a third and distinct species. but to this latter subject I shall return in the following chapter. So that we might obtain the parentspecies and its several modified descendants from the lower and upper beds of the same formation. the chance of discovering in a formation in any one country all the early 322 . and on this view we do find the kind of evidence of change which on the theory we ought to find. namely. According to this view. as Agassiz and Pictet. we should not recognise their blood-relationship. yet are far more closely related to each other than are the species found in more widely separated formations. we find that the embedded fossils. With animals and plants that propagate rapidly and do not wander much. which include many shells believed by the majority of naturalists to be identical with existing species. and that these late tertiary species really present no difference whatever from their living representatives. and should consequently rank them as distinct species. Nor should it be forgotten. in opposition to the judgment of most naturalists. so that here. as we have formerly seen. but some excellent naturalists. there is reason to suspect. to distinct but consecutive stages of the same great formation.

was probably short in comparison with that during which it remained without undergoing any change. horses.stages of transition between any two forms. for instance. namely. that geology yields no linking forms. and destroy four-fifths of them. that oftenest present varieties. by writers who believe in the immutability of species. It is a more important consideration. geologists at some future period will be able to prove that our different breeds of cattle. This assertion. with shells and other marine animals. it is probable that those which had the widest range." If we take a genus having a score of species. which are ranked by some conchologists as distinct species from their European representatives. fine. and by other conchologists as only varieties. no one doubts that the remainder will stand much more distinct from each other. Most marine animals have a wide range. Lubbock has remarked. sheep. and dogs are descended from a single stock or from several aboriginal stocks. far exceeding the limits of the known geological formations in Europe. as lately insisted on by Dr. intermediate. perhaps. and with fossil species this can rarely be done. with perfect specimens for examination. As Sir J. so that. as we shall see in the next chapter. and we have seen that with plants it is those which have the widest range. or. until many specimens are collected from many places. and thus proved to be the same species. We shall. leading to the same result. best perceive the improbability of our being enabled to connect species by numerous. What geological research has not 323 . and this again would greatly lessen the chance of our being able to trace the stages of transition in any one geological formation. by asking ourselves whether. though long as measured by years. This could be effected by the future geologist only by his discovering in a fossil state numerous intermediate gradations. It should not be forgotten. that the period during which each species underwent modification. is small. for the successive changes are supposed to have been local or confined to some one spot. "Every species is a link between other allied forms. as it is called. is certainly erroneous. recent and extinct. It has been asserted over and over again. fossil links. are really varieties. first to local varieties and ultimately to new species. or are. the genus itself will stand more distinct from other allied genera. specifically distinct. that at the present day. whether certain sea-shells inhabiting the shores of North America. and such success is improbable in the highest degree. again. two forms can seldom be connected by intermediate varieties. If the extreme forms in the genus happen to have been thus destroyed. have oftenest given rise. Falconer.

These periods of subsidence would be separated from each other by immense intervals of time. During the periods of subsidence. Wherever sediment did not accumulate on the bed of the sea. the fossiliferous formations on the steeper shores would be destroyed. But this ought not to be expected. during which the area would be either stationary or rising. connecting together nearly all existing and extinct species. I fully agree with Mr. Godwin. by the incessant coast-action. and of thickness sufficient to last to an age as distant in futurity as the secondary formations lie in the past. or of those which lived on naked submarine rocks. yet if all the species were to be collected which have ever lived there. almost as soon as accumulated. that the present condition of the Malay Archipelago. with its numerous large islands separated by wide and shallow seas. yet this has been repeatedly advanced as a most serious objection against my views. Formations rich in fossils of many kinds. as fine as existing varieties. sedimentary beds could hardly be accumulated of great thickness during the periods of elevation. and from Britain to Russia. and those embedded in gravel or sand would not endure to a distant epoch. 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. would generally be formed in the archipelago only during periods of subsidence. no remains could be preserved. The Malay Archipelago is about the size of Europe from the North Cape to the Mediterranean.revealed. there would probably 324 . Even throughout the extensive and shallow seas within the archipelago. is the former existence of infinitely numerous gradations. would be embedded. whilst rising. as we now see on the shores of South America.Austen. 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. and therefore equals all the geological formations which have been examined with any accuracy. excepting those of the United States of America. while most of our formations were accumulating. or where it did not accumulate at a sufficient rate to protect organic bodies from decay. or become capped and protected by subsequent deposits. Not many of the strictly littoral animals. probably represents the former state of Europe. so as to have a good chance of enduring to a very distant future. The Malay Archipelago is one of the richest regions in organic beings.

transitional varieties would merely appear as so many new. 325 . and no closely consecutive record of their modifications could be preserved in any one formation. and such assuredly we do find—some more distantly. on our theory. according to the principles followed by many palaeontologists. as they would differ from their former state in a nearly uniform. if found in different stages of the same formation. which would oftenest produce new varieties. they would. pressed so hardly on my theory. If then there be some degree of truth in these remarks. though only some of them. and analogy plainly leads to the belief that it would be chiefly these far-ranging species.be much extinction of life. and these links. some more closely. If such gradations were not all fully preserved. they would slowly spread and supplant their parent-forms. an infinite number of those fine transitional forms. there would be much variation. related to each other. be ranked as distinct species. have connected all the past and present species of the same group into one long and branching chain of life. be ranked as new and distinct species. by many palaeontologists. would. but the geological record would then be less perfect. We ought only to look for a few links. and in these cases the inhabitants of the archipelago would migrate. But I do not pretend that I should ever have suspected how poor was the record in the best preserved geological sections. It may be doubted whether the duration of any one great period of subsidence over the whole or part of the archipelago. but if possessed of any decided advantage. we have no right to expect to find. though closely allied species. and that slight climatical changes would intervene during such lengthy periods. let them be ever so close. and as they would be found embedded in slightly different substages of the same formation. together with a contemporaneous accumulation of sediment. would EXCEED the average duration of the same specific forms. and the varieties would at first be local or confined to one place. and these contingencies are indispensable for the preservation of all the transitional gradations between any two or more species. or when further modified and improved. had not the absence of innumerable transitional links between the species which lived at the commencement and close of each formation. It is also probable that each great period of subsidence would be interrupted by oscillations of level. in our geological formations. When such varieties returned to their ancient homes. which. though perhaps extremely slight degree. during the periods of elevation. Very many of the marine inhabitants of the archipelago now range thousands of miles beyond its confines.

longer perhaps in many cases than the time required for the accumulation of each formation. and have slowly multiplied. and Sedgwick. If numerous species. that they did not exist before that stage. belonging to the same genera or families. has been urged by several palaeontologists—for instance. have really started into life at once. in his excellent Review of this work. must have been an extremely slow process. all of which are descended from some one progenitor. and the progenitors must have lived long before their modified descendants. to fly through the air. Professor Pictet. These intervals will have given time for the multiplication of species from some one parent-form: and in the succeeding formation. Pictet. and a few species had thus acquired a great advantage over other organisms. as experience has so often shown. the fact would be fatal to the theory of evolution through natural selection. But look at the 326 . but that. which would spread rapidly and widely throughout the world. a comparatively short time would be necessary to produce many divergent forms. We do not make due allowance for the enormous intervals of time which have elapsed between our consecutive formations. cannot see how the successive modifications of the anterior limbs of a supposed prototype could possibly have been of any advantage. we forget that groups of species may elsewhere have long existed. negative evidence is worthless. compared with the area over which our geological formations have been carefully examined. I may here recall a remark formerly made. On The Sudden Appearance Of Whole Groups Of Allied Species The abrupt manner in which whole groups of species suddenly appear in certain formations. in commenting on early transitional forms. that it might require a long succession of ages to adapt an organism to some new and peculiar line of life. before they invaded the ancient archipelagoes of Europe and the United States. namely. as a fatal objection to the belief in the transmutation of species. and consequently that the transitional forms would often long remain confined to some one region. and falsely infer. In all cases positive palaeontological evidence may be implicitly trusted. But we continually overrate the perfection of the geological record. For the development by this means of a group of forms. for instance. We continually forget how large the world is.5. because certain genera or families have not been found beneath a certain stage. by Agassiz. and taking birds as an illustration. such groups or species will appear as if suddenly created. when this adaptation had once been effected.

And now one of the richest known accumulations of fossil mammals belongs to the middle of the secondary series. as far back as the miocene stage. published not many years ago. who would have ventured to suppose that no less than at least thirty different bird-like animals. some of gigantic size. the conclusions on the first appearance and disappearance of several groups of animals have been considerably modified. I may recall the well-known fact that in geological treatises. I may give another instance. Had it not been for the rare accident of the preservation of footsteps in the new red sandstone of the United States. and ultimately to rise from its surface and glide through the air? I will now give a few examples to illustrate the foregoing remarks. I 327 . and to show how liable we are to error in supposing that whole groups of species have suddenly been produced. and true mammals have been discovered in the new red sandstone at nearly the commencement of this great series. mammals were always spoken of as having abruptly come in at the commencement of the tertiary series. Not long ago. that strange bird.penguins of the Southern Ocean. palaeontologists maintained that the whole class of birds came suddenly into existence during the eocene period. South America and in Europe. 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. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world. with a long lizard-like tail. which. and with its wings furnished with two free claws. on the authority of Professor Owen. existed during that period? Not a fragment of bone has been discovered in these beds. but now we know. but now extinct species have been discovered in India. bearing a pair of feathers on each joint. and still more recently. Even in so short an interval as that between the first and second editions of Pictet's great work on Palaeontology. that a bird certainly lived during the deposition of the upper greensand. but what special difficulty is there in believing that it might profit the modified descendants of the penguin. and a third edition would require still further changes. for they exist in infinite numbers and of many kinds. published in 1844-46 and in 1853-57. from having passed under my own eyes has much struck me. Cuvier used to urge that no monkey occurred in any tertiary stratum. I do not suppose that we here see the real transitional grades through which the wings of birds have passed. first to become enabled to flap along the surface of the sea like the logger-headed duck. has been discovered in the oolitic slates of Solenhofen. In a memoir on Fossil Sessile Cirripedes. the Archeopteryx.

I inferred that. but it would not have formed an insuperable difficulty. low down. from the extraordinary abundance of the individuals of many species all over the world. Some few families of fish now have a confined range. adding. Woodward in the upper chalk. from all these circumstances. according to Agassiz. a Pyrgoma. this cirripede was a Chthamalus. Still more recently. inhabiting various zones of depths. as I then thought. from the Arctic regions to the equator. and ubiquitous genus. This group includes the large majority of existing species. and as not one species had then been discovered in beds of this age. And. M. one more instance of the abrupt appearance of a great group of species. and even some palaeozoic forms have thus been classed by one high authority. had sessile cirripedes existed during the secondary periods. as if to make the case as striking as possible. and after having been largely developed in 328 . from the ease with which even a fragment of a valve can be recognised. large. The case most frequently insisted on by palaeontologists of the apparently sudden appearance of a whole group of species. so that we now have abundant evidence of the existence of this group of animals during the secondary period. This was a sore trouble to me.stated that. Bosquet. 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. when a skilful palaeontologist. and by running through Pictet's Palaeontology it will be seen that very few species are known from several formations in Europe. has been discovered by Mr. I concluded that this great group had been suddenly developed at the commencement of the tertiary series. they would certainly have been preserved and discovered. If the teleosteans had really appeared suddenly in the northern hemisphere at the commencement of the chalk formation. is that of the teleostean fishes. the teleostean fishes might formerly have had a similarly confined range. a very common. the fact would have been highly remarkable. of which not one species has as yet been found even in any tertiary stratum. in the Chalk period. a member of a distinct subfamily of sessile cirripedes. But my work had hardly been published. It is almost superfluous to remark that hardly any fossil-fish are known from south of the equator. But certain Jurassic and Triassic forms are now commonly admitted to be teleostean. from the upper tidal limits to fifty fathoms. from the large number of existing and extinct tertiary species. from the perfect manner in which specimens are preserved in the oldest tertiary beds. sent me a drawing of a perfect specimen of an unmistakable sessile cirripede. which he had himself extracted from the chalk of Belgium.

if the Malay Archipelago were converted into land. from our ignorance of the geology of other countries beyond the confines of Europe and the United States. 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. until some of the species became adapted to a cooler climate. and thus reach other and distant seas. the tropical parts of the Indian Ocean would form a large and perfectly enclosed basin. From these considerations. Even at this day.some one sea. and here they would remain confined. as it would be for a naturalist to land for five minutes on a barren point in Australia. in which any great group of marine animals might be multiplied. have spread widely. 329 . and were enabled to double the southern capes of Africa or Australia. it seems to me to be about as rash to dogmatize on the succession of organic forms throughout the world. and then to discuss the number and range of its productions. and from the revolution in our palaeontological knowledge effected by the discoveries of the last dozen years.

and which probably differed greatly from any known animal. as the Nautilus. and other elements may have hereafter to be introduced into the problem. appears a very short time for the many and great mutations of life. It is. probable. that these old species were the progenitors of all the species belonging to the same groups which have subsequently appeared. Sir W. Most of the arguments which have convinced me that all the existing species of the same group are descended from a single progenitor. and it cannot on our theory be supposed. Lingula.. do not differ much from living species. which must have lived long before the Cambrian age. etc. Thompson concludes that the consolidation of the crust can hardly have occurred less than twenty or more than four hundred million years ago. and such 330 . which have certainly occurred since the Cambrian formation. and that during these vast periods the world swarmed with living creatures. For instance. 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. Croll estimates that about sixty million years have elapsed since the Cambrian period. however. judging from the small amount of organic change since the commencement of the Glacial epoch. but this. These very wide limits show how doubtful the data are. Some of the most ancient animals. in a fit state for the habitation of living creatures.6. Mr. as long as. it is indisputable that before the lowest Cambrian stratum was deposited long periods elapsed. or probably far longer than. for it seems doubtful whether the earth. if the theory be true. as Sir William Thompson insists. which is much more serious. Consequently. has lasted long enough. for they are not in any degree intermediate in character. it cannot be doubted that all the Cambrian and Silurian trilobites are descended from some one crustacean. that the world at a very early period was subjected to more rapid and violent changes in its physical conditions than those now occurring. but probably not less than ninety-eight or more than two hundred million years. 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. the whole interval from the Cambrian age to the present day. Here we encounter a formidable objection. On The Sudden Appearance Of Groups Of Allied Species In The Lowest Known Fossiliferous Strata There is another and allied difficulty. apply with equal force to the earliest known species.

and the existence of the Eozoon in the Laurentian formation of Canada is generally admitted. and these would always have existed in a partially metamorphosed condition. probably indicates life at these periods. still lower down in the Lower Cambrian formation. and. that the appearance of the so-called primordial fauna (of Barrande) may by some be considered as a comparatively modern event. I can give no satisfactory answer. Not very long ago M. from the base of the palaeozoic series to the present time. certainly preyed on other minute organic beings. and containing various molluscs and annelids. as Lyell and E. have proved true. We should not forget that only a small portion of the world is known with accuracy. To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system. Thus the words. 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. and which are almost the same with those since used by Sir W. The presence of phosphatic nodules and bituminous matter. about the existence of living beings long before the Cambrian period. in the lowest of which the Eozoon is found. We are thus carried back to a period so remote. Mr Hicks has found South Wales beds rich in trilobites. There are three great series of strata beneath the Silurian system in Canada. Murchison at their head. But the descriptions which we possess of the Silurian deposits over immense territories in Russia and in North 331 . Logan. Forbes. and now. Other highly competent judges. beneath the then known Silurian system. but is highly organised for its class. even in some of the lowest azotic rocks. have disputed this conclusion. were until recently convinced that we beheld in the organic remains of the lowest Silurian stratum the first dawn of life. Logan states that their "united thickness may possibly far surpass that of all the succeeding rocks. as Dr. Barrande added another and lower stage. which I wrote in 1859. It does not seem probable that the most ancient beds have been quite worn away by denudation.changes would have tended to induce changes at a corresponding rate in the organisms which then existed. abounding with new and peculiar species. or that their fossils have been wholly obliterated by metamorphic action." The Eozoon belongs to the most lowly organised of all classes of animals. Dawson has remarked. for if this had been the case we should have found only small remnants of the formations next succeeding them in age. it existed in countless numbers. Nevertheless. which must have lived in great numbers. with Sir R. Several eminent geologists. Sir W.

But we do not know what was the state of things in the intervals between the several successive formations. then. which must have intervened during these enormously long periods. do not support the view that the older a formation is the more invariably it has suffered extreme denudation and metamorphism. large tracts of land have existed. To show that it may hereafter receive some explanation. and on the other hand. oceans have extended from the remotest period of which we have any record. and would have been at least partially upheaved by the oscillations of level. whence the sediment was derived. or as the bed of an open and unfathomable sea. of the force of elevation. Our continents seem to have been formed by a preponderance. we see them studded with many islands. the great archipelagoes still areas of oscillations of level. subjected. if this can be called a truly oceanic island) is as yet known to afford even a remnant of any palaeozoic or secondary formation. on which sediment was not deposited. and may be truly urged as a valid argument against the views here entertained. I will give the following hypothesis. for had they existed. But we have no reason to assume that things have thus remained from the beginning of the world. The case at present must remain inexplicable. Hence. in the several formations of Europe and of the United States. which are thrice as extensive as the land. whether Europe and the United States during these intervals existed as dry land. where our oceans now extend. This same view has since been maintained by Agassiz and others. since the Cambrian period. neither continents nor continental islands existed where our oceans now extend. The coloured map appended to my volume on Coral Reefs. and the continents areas of elevation. miles in thickness. of which the formations are composed.America. Looking to the existing oceans. and from the amount of sediment. no doubt. that where continents now exist. but hardly one truly oceanic island (with the exception of New Zealand. to great oscillations of level. occurred in the neighbourhood of the now existing continents of Europe and North America. we may infer anything from these facts. If. From the nature of the organic remains which do not appear to have inhabited profound depths. led me to conclude that the great oceans are still mainly areas of subsidence. we may infer that. palaeozoic and secondary formations would in all probability have been accumulated from sediment derived from their wear and tear. But may not the areas of preponderant movement have changed in 332 . during many oscillations of level. or as a submarine surface near land. that during the palaeozoic and secondary periods. we may infer that from first to last large islands or tracts of land. we may perhaps infer.

might have undergone far more metamorphic action than strata which have always remained nearer to the surface. more or less different in the successive chapters. etc. which are entombed in our consecutive formations. Each word of the slowly-changing language. The several difficulties here discussed. and most geologists and palaeontologists are much shaken in their former belief. will undoubtedly at once reject my theory. The sudden manner in which several groups of species first appear in our European formations. 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. Forbes.. maintained the immutability of species. Sedgwick. may represent the forms of life. which must have been heated under great pressure. But Sir Charles Lyell now gives the support of his high authority to the opposite side. Agassiz. only here and there a short chapter has been preserved. Falconer. relating only to two or three countries. The immense areas in some parts of the world. we do not find infinitely numerous fine transitional forms closely joining them all together. of naked metamorphic rocks. We see this in the fact that the most eminent palaeontologists. as at present known. for instance. Of this history we possess the last volume alone. though we find in our geological formations many links between the species which now exist and which formerly existed. Pictet. continents may have existed where oceans are now spread out. E. the almost entire absence. Nor should we be justified in assuming that if. have always seemed to me to require some special explanation. only here and there a few lines. and which falsely appear to have been 333 . have unanimously. often vehemently.. the bed of the Pacific Ocean were now converted into a continent we should there find sedimentary formations. that. namely. for it might well happen that strata which had subsided some miles nearer to the centre of the earth. older than the Cambrian strata. For my part. and clear and open oceans may have existed where our continents now stand. in recognisable condition. and of each page. Those who believe that the geological record is in any degree perfect. Murchison. Barrande. as Lyell.the lapse of ages? At a period long antecedent to the Cambrian epoch. following out Lyell's metaphor. namely. I look at the geological record as a history of the world imperfectly kept and written in a changing dialect. Of this volume. of formations rich in fossils beneath the Cambrian strata. supposing such to have been formerly deposited. for instance in South America. etc. are all undoubtedly of the most serious nature. Cuvier. and which had been pressed on by an enormous weight of superincumbent water. and all our greatest geologists.

abruptly introduced. 334 . On this view the difficulties above discussed are greatly diminished or even disappear.

Chapter 11 On The Geological Succession Of Organic Beings 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. 335 .

as far as we know. and every year tends to fill up the blanks between the stages. and then allow the pre-existing fauna to reappear. or. of which a striking instance has been observed in Switzerland. we have no reason to believe that the same identical form ever reappears. or with that of their slow and gradual modification. change more quickly than those that are low: though there are exceptions to this rule. and only one or two are new. neither the appearance nor disappearance of the many species embedded in each formation has been simultaneous. seems satisfactory. that it is a case of temporary migration from a distinct geographical province. Species belonging to different genera and classes have not changed at the same rate. Falconer has given a striking instance of a similar fact. The Silurian Lingula differs but little from the living species of this genus. namely. Barrande. The productions of the land seem to have changed at a quicker rate than those of the sea. as Bronn has remarked. though undoubtedly of high antiquity if measured by years. only one or two species are extinct. New species have appeared very slowly. is not the same in each successive so-called formation. 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. which intrude for a period in the midst of an older formation. one after another. having appeared there for the first time. The amount of organic change. Yet if we compare any but the most closely related formations. as Pictet has remarked. or in the same degree. but. 336 . There is some reason to believe that organisms high in the scale. Lyell has shown that it is hardly possible to resist the evidence on this head in the case of the several tertiary stages. all the species will be found to have undergone some change. both on the land and in the waters. through variation and natural selection. but Lyell's explanation. In the older tertiary beds a few living shells may still be found in the midst of a multitude of extinct forms. The strongest apparent exception to this latter rule is that of the so-called "colonies" of M. The secondary formations are more broken. When a species has once disappeared from the face of the earth.1. for an existing crocodile is associated with many lost mammals and reptiles in the subHimalayan deposits. whereas most of the other Silurian Molluscs and all the Crustaceans have changed greatly. on the face of the earth. and to make the proportion between the lost and existing forms more gradual. In some of the most recent beds. either locally.

These several facts accord well with our theory, which includes no fixed law of development, causing all the inhabitants of an area to change abruptly, or simultaneously, or to an equal degree. The process of modification must be slow, and will generally affect only a few species at the same time; for the variability of each species is independent of that of all others. Whether such variations or individual differences as may arise will be accumulated through natural selection in a greater or less degree, thus causing a greater or less amount of permanent modification, will depend on many complex contingencies—on the variations being of a beneficial nature, on the freedom of intercrossing, on the slowly changing physical conditions of the country, on the immigration of new colonists, and on the nature of the other inhabitants with which the varying species come into competition. Hence it is by no means surprising that one species should retain the same identical form much longer than others; or, if changing, should change in a less degree. We find similar relations between the existing inhabitants of distinct countries; for instance, the land-shells and coleopterous insects of Madeira have come to differ considerably from their nearest allies on the continent of Europe, whereas the marine shells and birds have remained unaltered. We can perhaps understand the apparently quicker rate of change in terrestrial and in more highly organised productions compared with marine and lower productions, by the more complex relations of the higher beings to their organic and inorganic conditions of life, as explained in a former chapter. When many of the inhabitants of any area have become modified and improved, we can understand, on the principle of competition, and from the all-important relations of organism to organism in the struggle for life, that any form which did not become in some degree modified and improved, would be liable to extermination. Hence, we see why all the species in the same region do at last, if we look to long enough intervals of time, become modified; for otherwise they would become extinct. In members of the same class the average amount of change, during long and equal periods of time, may, perhaps, be nearly the same; but as the accumulation of enduring formations, rich in fossils, depends on great masses of sediment being deposited on subsiding areas, our formations have been almost necessarily accumulated at wide and irregularly intermittent intervals of time; consequently the amount of organic change exhibited by the fossils embedded in consecutive formations is not equal. Each formation, on this view, does not mark a new and complete act of creation, but only an occasional scene, taken almost at hazard, in an ever slowly changing drama.

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We can clearly understand why a species when once lost should never reappear, even if the very same conditions of life, organic and inorganic, should recur. 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, and thus supplant it; yet the two forms—the old and the new—would not be identically the same; for both would almost certainly inherit different characters from their distinct progenitors; and organisms already differing would vary in a different manner. For instance, it is possible, if all our fantail-pigeons were destroyed, that fanciers might make a new breed hardly distinguishable from the present breed; but if the parent rock-pigeon were likewise destroyed, and under nature we have every reason to believe that parent forms are generally supplanted and exterminated by their improved offspring, it is incredible that a fantail, identical with the existing breed, could be raised from any other species of pigeon, or even from any other well established race of the domestic pigeon, for the successive variations would almost certainly be in some degree different, and the newly-formed variety would probably inherit from its progenitor some characteristic differences. Groups of species, that is, genera and families, follow the same general rules in their appearance and disappearance as do single species, changing more or less quickly, and in a greater or lesser degree. A group, when it has once disappeared, never reappears; that is, its existence, as long as it lasts, is continuous. I am aware that there are some apparent exceptions to this rule, but the exceptions are surprisingly few, so few that E. Forbes, Pictet, and Woodward (though all strongly opposed to such views as I maintain) admit its truth; and the rule strictly accords with the theory. For all the species of the same group, however long it may have lasted, are the modified descendants one from the other, and all from a common progenitor. In the genus Lingula, for instance, the species which have successively appeared at all ages must have been connected by an unbroken series of generations, from the lowest Silurian stratum to the present day. We have seen in the last chapter that whole groups of species sometimes falsely appear to have been abruptly developed; and I have attempted to give an explanation of this fact, which if true would be fatal to my views. But such cases are certainly exceptional; the general rule being a gradual increase in number, until the group reaches its maximum, and then, sooner or later, a gradual decrease. If the number of the species included within a genus, or the number of the genera within a family, be

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represented by a vertical line of varying thickness, ascending through the successive geological formations, in which the species are found, the line will sometimes falsely appear to begin at its lower end, not in a sharp point, but abruptly; it then gradually thickens upwards, often keeping of equal thickness for a space, and ultimately thins out in the upper beds, marking the decrease and final extinction of the species. This gradual increase in number of the species of a group is strictly conformable with the theory; for the species of the same genus, and the genera of the same family, can increase only slowly and progressively; the process of modification and the production of a number of allied forms necessarily being a slow and gradual process, one species first giving rise to two or three varieties, these being slowly converted into species, which in their turn produce by equally slow steps other varieties and species, and so on, like the branching of a great tree from a single stem, till the group becomes large.

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2. On Extinction
We have as yet only spokesn incidentally of the disappearance of species and of groups of species. On the theory of natural selection, the extinction of old forms and the production of new and improved forms are intimately connected together. The old notion of all the inhabitants of the earth having been swept away by catastrophes at successive periods is very generally given up, even by those geologists, as Elie de Beaumont, Murchison, Barrande, etc., whose general views would naturally lead them to this conclusion. On the contrary, we have every reason to believe, from the study of the tertiary formations, that species and groups of species gradually disappear, one after another, first from one spot, then from another, and finally from the world. In some few cases, however, as by the breaking of an isthmus and the consequent irruption of a multitude of new inhabitants into an adjoining sea, or by the final subsidence of an island, the process of extinction may have been rapid. Both single species and whole groups of species last for very unequal periods; some groups, as we have seen, have endured from the earliest known dawn of life to the present day; some have disappeared before the close of the palaeozoic period. No fixed law seems to determine the length of time during which any single species or any single genus endures. 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, as before, by a vertical line of varying thickness the line is found to taper more gradually at its upper end, which marks the progress of extermination, than at its lower end, which marks the first appearance and the early increase in number of the species. In some cases, however, the extermination of whole groups, as of ammonites, towards the close of the secondary period, has been wonderfully sudden. The extinction of species has been involved in the most gratuitous mystery. Some authors have even supposed that, as the individual has a definite length of life, so have species a definite duration. No one can have marvelled more than I have done at the extinction of species. When I found in La Plata the tooth of a horse embedded with the remains of Mastodon, Megatherium, Toxodon and other extinct monsters, which all co-existed with still living shells at a very late geological period, I was filled with astonishment; for, seeing that the horse, since its introduction by the Spaniards into South America, has run wild over the whole country and has increased in numbers at an unparalleled rate, I asked myself

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what could so recently have exterminated the former horse under conditions of life apparently so favourable. But my astonishment was groundless. Professor Owen soon perceived that the tooth, though so like that of the existing horse, belonged to an extinct species. Had this horse been still living, but in some degree rare, no naturalist would have felt the least surprise at its rarity; for rarity is the attribute of a vast number of species of all classes, in all countries. If we ask ourselves why this or that species is rare, we answer that something is unfavourable in its conditions of life; but what that something is, we can hardly ever tell. On the supposition of the fossil horse still existing as a rare species, we might have felt certain, from the analogy of all other mammals, even of the slow-breeding elephant, and from the history of the naturalisation of the domestic horse in South America, that under more favourable conditions it would in a very few years have stocked the whole continent. But we could not have told what the unfavourable conditions were which checked its increase, whether some one or several contingencies, and at what period of the horse's life, and in what degree they severally acted. If the conditions had gone on, however slowly, becoming less and less favourable, we assuredly should not have perceived the fact, yet the fossil horse would certainly have become rarer and rarer, and finally extinct—its place being seized on by some more successful competitor. It is most difficult always to remember that the increase of every living creature is constantly being checked by unperceived hostile agencies; and that these same unperceived agencies are amply sufficient to cause rarity, and finally extinction. So little is this subject understood, that I have heard surprise repeatedly expressed at such great monsters as the Mastodon and the more ancient Dinosaurians having become extinct; as if mere bodily strength gave victory in the battle of life. Mere size, on the contrary, would in some cases determine, as has been remarked by Owen, quicker extermination, from the greater amount of requisite food. Before man inhabited India or Africa, some cause must have checked the continued increase of the existing elephant. A highly capable judge, Dr. Falconer, believes that it is chiefly insects which, from incessantly harassing and weakening the elephant in India, check its increase; and this was Bruce's conclusion with respect to the African elephant in Abyssinia. It is certain that insects and blood-sucking bats determine the existence of the larger naturalised quadrupeds in several parts of South America. We see in many cases in the more recent tertiary formations that rarity precedes extinction; and we know that this has been the progress of events with those animals which have been exterminated, either locally

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or wholly, through man's agency. I may repeat what I published in 1845, namely, that to admit that species generally become rare before they become extinct—to feel no surprise at the rarity of a species, and yet to marvel greatly when the species ceases to exist, is much the same as to admit that sickness in the individual is the forerunner of death—to feel no surprise at sickness, but, when the sick man dies, to wonder and to suspect that he died by some deed of violence. The theory of natural selection is grounded on the belief that each new variety and ultimately each new species, is produced and maintained by having some advantage over those with which it comes into competition; and the consequent extinction of less-favoured forms almost inevitably follows. It is the same with our domestic productions: when a new and slightly improved variety has been raised, it at first supplants the less improved varieties in the same neighbourhood; when much improved it is transported far and near, like our short-horn cattle, and takes the place of other breeds in other countries. Thus the appearance of new forms and the disappearance of old forms, both those naturally and artificially produced, are bound together. In flourishing groups, 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; but we know that species have not gone on indefinitely increasing, at least during the later geological epochs, so that, looking to later times, we may believe that the production of new forms has caused the extinction of about the same number of old forms. The competition will generally be most severe, as formerly explained and illustrated by examples, between the forms which are most like each other in all respects. Hence the improved and modified descendants of a species will generally cause the extermination of the parent-species; and if many new forms have been developed from any one species, the nearest allies of that species, i.e. the species of the same genus, will be the most liable to extermination. Thus, as I believe, a number of new species descended from one species, that is a new genus, comes to supplant an old genus, belonging to the same family. But it must often have happened that a new species belonging to some one group has seized on the place occupied by a species belonging to a distinct group, and thus have caused its extermination. If many allied forms be developed from the successful intruder, many will have to yield their places; and it will generally be the allied forms, which will suffer from some inherited inferiority in common. But whether it be species belonging to the same or

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to a distinct class, which have yielded their places to other modified and improved species, a few of the sufferers may often be preserved for a long time, from being fitted to some peculiar line of life, or from inhabiting some distant and isolated station, where they will have escaped severe competition. For instance, some species of Trigonia, a great genus of shells in the secondary formations, survive in the Australian seas; and a few members of the great and almost extinct group of Ganoid fishes still inhabit our fresh waters. Therefore, the utter extinction of a group is generally, as we have seen, a slower process than its production. With respect to the apparently sudden extermination of whole families or orders, as of Trilobites at the close of the palaeozoic period, and of Ammonites at the close of the secondary period, we must remember what has been already said on the probable wide intervals of time between our consecutive formations; and in these intervals there may have been much slow extermination. Moreover, when, by sudden immigration or by unusually rapid development, many species of a new group have taken possession of an area, many of the older species will have been exterminated in a correspondingly rapid manner; and the forms which thus yield their places will commonly be allied, for they will partake of the same inferiority in common. Thus, as it seems to me, the manner in which single species and whole groups of species become extinct accords well with the theory of natural selection. We need not marvel at extinction; if we must marvel, let it be at our presumption in imagining for a moment that we understand the many complex contingencies on which the existence of each species depends. If we forget for an instant that each species tends to increase inordinately, and that some check is always in action, yet seldom perceived by us, the whole economy of nature will be utterly obscured. Whenever we can precisely say why this species is more abundant in individuals than that; why this species and not another can be naturalised in a given country; then, and not until then, we may justly feel surprise why we cannot account for the extinction of any particular species or group of species.

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3. On The Forms Of Life Changing Almost Simultaneously Throughout The World
Scarcely any palaeontological discovery is more striking than the fact that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant regions, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the organic remains in certain beds present an unmistakable resemblance to those of the Chalk. It is not that the same species are met with; for in some cases not one species is identically the same, but they belong to the same families, genera, and sections of genera, and sometimes are similarly characterised in such trifling points as mere superficial sculpture. Moreover, other forms, which are not found in the Chalk of Europe, but which occur in the formations either above or below, occur in the same order at these distant points of the world. In the several successive palaeozoic formations of Russia, Western Europe and North America, a similar parallelism in the forms of life has been observed by several authors; so it is, according to Lyell, with the European and North American tertiary deposits. Even if the few fossil species which are common to the Old and New Worlds were kept wholly out of view, the general parallelism in the successive forms of life, in the palaeozoic and tertiary stages, would still be manifest, and the several formations could be easily correlated. These observations, however, 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. We may doubt whether they have thus changed: if the Megatherium, Mylodon, Macrauchenia, and Toxodon had been brought to Europe from La Plata, without any information in regard to their geological position, no one would have suspected that they had co-existed with sea-shells all still living; but as these anomalous monsters co-existed with the Mastodon and Horse, it might at least have been inferred that they had lived during one of the later tertiary stages. When the marine forms of life are spoken of as having changed simultaneously throughout the world, it must not be supposed that this expression relates to the same year, or even to the same century, or even that it has a very strict geological sense; for if all the marine animals now

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living in Europe, and all those that lived in Europe during the pleistocene period (a very remote period as measured by years, including the whole glacial epoch) were compared with those now existing in South America or in Australia, 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. So, again, 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, than to the present inhabitants of Europe; and if this be so, 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. Nevertheless, looking to a remotely future epoch, there can be little doubt that all the more modern MARINE formations, namely, the upper pliocene, the pleistocene and strictly modern beds of Europe, North and South America, and Australia, from containing fossil remains in some degree allied, and from not including those forms which are found only in the older underlying deposits, would be correctly ranked as simultaneous in a geological sense. The fact of the forms of life changing simultaneously in the above large sense, at distant parts of the world, has greatly struck those admirable observers, MM. de Verneuil and d'Archiac. After referring to the parallelism of the palaeozoic forms of life in various parts of Europe, they add, "If struck by this strange sequence, we turn our attention to North America, and there discover a series of analogous phenomena, it will appear certain that all these modifications of species, their extinction, and the introduction of new ones, cannot be owing to mere changes in marine currents or other causes more or less local and temporary, but depend on general laws which govern the whole animal kingdom." M. Barrande has made forcible remarks to precisely the same effect. It is, indeed, quite futile to look to changes of currents, climate, or other physical conditions, as the cause of these great mutations in the forms of life throughout the world, under the most different climates. We must, as Barrande has remarked, look to some special law. We shall see this more clearly when we treat of the present distribution of organic beings, and find how slight is the relation between the physical conditions of various countries and the nature of their inhabitants. This great fact of the parallel succession of the forms of life throughout the world, is explicable on the theory of natural selection. New species are formed by having some advantage over older forms; and the forms, which are already dominant, or have some advantage over the other

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forms in their own country, give birth to the greatest number of new varieties or incipient species. We have distinct evidence on this head, in the plants which are dominant, that is, which are commonest and most widely diffused, producing the greatest number of new varieties. It is also natural that the dominant, varying and far-spreading species, which have already invaded, to a certain extent, the territories of other species, should be those which would have the best chance of spreading still further, and of giving rise in new countries to other new varieties and species. The process of diffusion would often be very slow, depending on climatal and geographical changes, on strange accidents, and on the gradual acclimatization of new species to the various climates through which they might have to pass, but in the course of time the dominant forms would generally succeed in spreading and would ultimately prevail. The diffusion would, it is probable, be slower with the terrestrial inhabitants of distinct continents than with the marine inhabitants of the continuous sea. We might therefore expect to find, as we do find, a less strict degree of parallelism in the succession of the productions of the land than with those of the sea. Thus, as it seems to me, the parallel, and, taken in a large sense, simultaneous, succession of the same forms of life throughout the world, accords well with the principle of new species having been formed by dominant species spreading widely and varying; the new species thus produced being themselves dominant, owing to their having had some advantage over their already dominant parents, as well as over other species; and again spreading, varying, and producing new forms. The old forms which are beaten and which yield their places to the new and victorious forms, will generally be allied in groups, from inheriting some inferiority in common; and, therefore, as new and improved groups spread throughout the world, old groups disappear from the world; and the succession of forms everywhere tends to correspond both in their first appearance and final disappearance. There is one other remark connected with this subject worth making. I have given my reasons for believing that most of our great formations, rich in fossils, were deposited during periods of subsidence; and that blank intervals of vast duration, as far as fossils are concerned, occurred during the periods when the bed of the sea was either stationary or rising, and likewise when sediment was not thrown down quickly enough to embed and preserve organic remains. During these long and blank intervals I suppose that the inhabitants of each region underwent a considerable amount of modification and extinction, and that there was

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and immigration. When two formations have been deposited in two regions during nearly. As we have reason to believe that large areas are affected by the same movement. 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. but not exactly. but contemporaneous faunas. the same period. in his admirable Memoirs on the eocene deposits of England and France. yet the species themselves differ in a manner very difficult to account for considering the proximity of the two areas. it is probable that strictly contemporaneous formations have often been accumulated over very wide spaces in the same quarter of the world. in accordance with the general succession of the forms of life. is able to draw a close general parallelism between the successive stages in the two countries. although he finds in both a curious accordance in the numbers of the species belonging to the same genera. and that large areas have invariably been affected by the same movements. but when he compares certain stages in England with those in France. nevertheless the species would not all be the same in the apparently corresponding stages in the two regions. nevertheless he finds a surprising amount of difference in the species. but we are very far from having any right to conclude that this has invariably been the case. Barrande. the same general succession in the forms of life. for there will have been a little more time in the one region than in the other for modification. Mr. unless. it be assumed that an isthmus separated two seas inhabited by distinct. extinction.much migration from other parts of the world. from the causes explained in the foregoing paragraphs. we should find in both. indeed. and the order would falsely appear to be strictly parallel. in this case the several formations in the two regions could be arranged in the same order. also. but the species would not exactly correspond. shows that there is a striking general parallelism in the successive Silurian deposits of Bohemia and Scandinavia. Prestwich. I suspect that cases of this nature occur in Europe. 347 . Lyell has made similar observations on some of the later tertiary formations.

The 348 . M.4. in fact. 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. extinct species can all be classed either in still existing groups. but the extinct Halitherium had. Cuvier ranked the Ruminants and Pachyderms as two of the most distinct orders of mammals. without even a rudiment being left. and these terms imply that such forms are. but as this statement has often been ignored or even denied. as applied to extinct animals. If we confine our attention either to the living or to the extinct species of the same class. the series is far less perfect than if we combine both into one general system. he dissolves by gradations the apparently wide interval between the pig and the camel. for example. as a general rule. That the extinct forms of life help to fill up the intervals between existing genera. as the name given to it by Professor Gervais expresses. and orders. Gaudry. it may be well to make some remarks on this subject. But. The more ancient any form is. it differs from living forms. to which the Sirenia are in other respects allied. and to give some instances. but so many fossil links have been disentombed that Owen has had to alter the whole classification. of prophetic or synthetic types. or between them. No one will deny that the Hipparion is intermediate between the existing horse and certain other ungulate forms. and has placed certain Pachyderms in the same sub-order with ruminants. The Sirenia form a very distinct group of the mammals. Another distinguished palaeontologist. an ossified thigh-bone "articulated to a well-defined acetabulum in the pelvis. and this fact is at once explained on the principle of descent." and it thus makes some approach to ordinary hoofed quadrupeds. the more. is certainly true. and in the writings of Agassiz. What a wonderful connecting link in the chain of mammals is the Typotherium from South America. but the Macrauchenia of South America connects to a certain extent these two grand divisions. The Ungulata or hoofed quadrupeds are now divided into the even-toed or odd-toed divisions. intermediate or connecting links. according to Professor Flower. as Buckland long ago remarked. and which cannot be placed in any existing order. On The Affinities Of Extinct Species To Each Other. In the writings of Professor Owen we continually meet with the expression of generalised forms. And To Living Forms Let us now look to the mutual affinities of extinct and living species. families. All fall into a few grand classes. and one of the most remarkable peculiarities in existing dugong and lamentin is the entire absence of hind limbs.

as the Lepidosiren. It is a common belief that the more ancient a form is. is discovered having affinities directed towards very distinct groups. a higher authority could not be named. yet that at this ancient period the groups were not so distinctly separated from each other as they now are. and it would be difficult to prove the truth of the proposition." 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.cetaceans or whales are widely different from all other mammals. are considered by Professor Huxley to be undoubtedly cetaceans. Some writers have objected to any extinct species. on the one hand. Turning to the Invertebrata. The most common case. one of the Dinosaurians—that group which includes the most gigantic of all terrestrial reptiles. especially with respect to very distinct groups. and some extinct genera between living genera. with the recent members of the same classes. or group of species. and the eocene Mammals. being considered as intermediate between any two living species. we must admit that there is truth in the remark. but the tertiary Zeuglodon and Squalodon. that he is every day taught that. although palaeozoic animals can certainly be classed under existing groups. the older Cephalopods. This remark no doubt must be restricted to those groups which have undergone much change in the course of geological ages. As the subject is somewhat 349 . supposing them to be distinguished at the present day by a score of characters. which have been placed by some naturalists in an order by themselves. Barrande asserts. the objection is probably valid. for every now and then even a living animal. seems to be that. such as fish and reptiles. But in a natural classification many fossil species certainly stand between living species. the older Fish. so that the two groups formerly made a somewhat nearer approach to each other than they now do. by so much the more it tends to connect by some of its characters groups now widely separated from each other. and on the other hand by the Compsognathus. even between genera belonging to distinct families. or groups of species. Yet if we compare the older Reptiles and Batrachians. If by this term it is meant that an extinct form is directly intermediate in all its characters between two living forms or groups. "and to constitute connecting links with the aquatic carnivora. the ancient members are separated by a somewhat lesser number of characters. Let us see how far these several facts and inferences accord with the theory of descent with modification. by the ostrich and extinct Archeopteryx.

would be partly justified. make. Yet he who objected to consider as intermediate the extinct genera. the three existing families on the uppermost line would be rendered less distinct from each other. a closely allied family or subfamily. the genera a1. but this is unimportant for us. has become divided into several sub-families and families. that a species might go on being slightly modified in relation to its slightly altered conditions of life. which thus link together the living genera of three families. some of which are supposed to have perished at different periods. The diagram is much too simple. Therefore it is quite possible. for instance. f8. were disinterred. This is represented in the diagram by the letter F14. m3. i14. b14 and f14. These three families. and the dotted lines diverging from them the species in each genus. m14. together with the many extinct genera on the several lines of descent diverging from the parent form (A) will form an order. and o14. and some to have endured to the present day. descended from (A). these three families would be so closely linked together that they probably would have to be united into one great family. I must request the reader to turn to the diagram in the fourth chapter. but only by a long and circuitous 350 . too few genera and too few species being given. for they are intermediate. assume that divergence of character is a necessary contingency. in nearly the same manner as has occurred with ruminants and certain pachyderms. were discovered at several points low down in the series. q14. it depends solely on the descendants from a species being thus enabled to seize on many and different places in the economy of nature. extinct and recent. m6. p14. All the many forms. a third family. and all the forms beneath the uppermost line may be considered as extinct. which was formerly illustrated by this diagram. Hence. and this order. We must not. as before remarked. however. the more recent any form is the more it will generally differ from its ancient progenitor. The three existing genera. one order. m9. We may suppose that the numbered letters in italics represent genera. a5. for all will have inherited something in common from their ancient progenitor. If. as we have seen in the case of some Silurian forms. will form a small family. not directly. By looking at the diagram we can see that if many of the extinct forms supposed to be embedded in the successive formations. from the continued effects of extinction and divergence of character.complex. a14. On the principle of the continued tendency to divergence of character. The horizontal lines may represent successive geological formations. a10. and yet retain throughout a vast period the same general characteristics. we can understand the rule that the most ancient fossils differ most from existing forms.

that those groups which have. be supposed to differ from each other by half-a-dozen important characters. 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. are explained in a satisfactory manner. Under nature the process will be far more complicated than is represented in the diagram. Thus it comes that ancient and extinct genera are often in a greater or less degree intermediate in character between their modified descendants. above No. As we possess only the last volume of the geological record. on the theory of descent with modification. allow for 351 . VI. and are the parents of those which became still more modified at the seventh stage. and this by the concurrent evidence of our best palaeontologists is frequently the case. Thus. they will have endured for extremely unequal lengths of time. then the families which existed at a period marked VI would certainly have differed from each other by a less number of characters. however. 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. and thus to unite distinct families or orders. And they are wholly inexplicable on any other view. then only two of the families (those on the left hand a14. etc. So again. etc. and that in a very broken condition. we have no right to expect. on the uppermost line. and there would remain two families which would be less distinct from each other than they were before the discovery of the fossils. If many extinct forms were to be discovered above one of the middle horizontal lines or geological formations—for instance. On this same theory. hence they could hardly fail to be nearly intermediate in character between the forms of life above and below.. to fill up the wide intervals in the natural system.) would have to be united into one. undergone much modification. We must. except in rare cases. 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. within known geological periods. the main facts with respect to the mutual affinities of the extinct forms of life to each other and to living forms. for the groups will have been more numerous. and will have been modified in various degrees. and b14. should in the older formations make some slight approach to each other. or between their collateral relations. if the three families formed of eight genera (a14 to m14). All that we have a right to expect is.course through many widely different forms. for they would at this early stage of descent have diverged in a less degree from their common progenitor.—but none from beneath this line.

and for a large amount of modification during the long and blank intervals between the successive formations. the fauna of each geological period undoubtedly is intermediate in character. if the principal living and extinct races of the domestic pigeon were arranged in serial affinity. especially in the case of terrestrial productions inhabiting separated districts. For instance. the species of mastodons and elephants. for the parent rock-pigeon still lives. which are at the opposite end of the series in this respect. the manner in which the fossils of the Devonian system. we have no reason to believe that forms successively produced necessarily endure for corresponding lengths of time.the entire extinction of some preceding forms. nor are those which are intermediate in character. and in the second place according to their periods of existence—do not accord in arrangement. is the fact. 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. that the organic remains from an intermediate formation are in some degree intermediate in character. insisted on by all palaeontologists. and in any one region for the immigration of new forms from other regions. were at once recognised by palaeontologists as intermediate in character between those of the overlying carboniferous and underlying Silurian systems. as unequal intervals of time have elapsed between consecutive formations. The species extreme in character are not the oldest or the most recent. between the preceding and succeeding faunas. A very ancient form may occasionally have lasted much longer than a form elsewhere subsequently produced. than are 352 . that fossils from two consecutive formations are far more closely related to each other. and many varieties between the rock-pigeon and the carrier have become extinct. intermediate in age. and carriers which are extreme in the important character of length of beak originated earlier than short-beaked tumblers. namely. To compare small things with great. Subject to these allowances. and even less with the order of their disappearance. when this system was first discovered. that certain genera offer exceptions to the rule. I need give only one instance. Closely connected with the statement. Falconer in two series—in the first place according to their mutual affinities. which is far from the case. that the record of the first appearance and disappearance of the species was complete. in this and other such cases. this arrangement would not closely accord with the order in time of their production. when arranged by Dr. But supposing for an instant. But each fauna is not necessarily exactly intermediate.

As the accumulation of each formation has often been interrupted. Let it be remembered that the forms of life. as I attempted to show in the last chapter. though the species are distinct in each stage. have changed almost simultaneously throughout the world. This fact alone.the fossils from two remote formations. very long as measured by years. which includes the whole glacial epoch. seems to have shaken Professor Pictet in his belief in the immutability of species. He who is acquainted with the distribution of existing species over the globe. being closely related. and note how little the specific forms of the inhabitants of the sea have been affected. Pictet gives as a well-known instance. closely allied forms. and these assuredly we do find. at least those inhabiting the sea. but only moderately long as measured geologically. as they have been called by some authors. or. and as long blank intervals have intervened between successive formations. we ought not to expect to find. We find. by the physical conditions of the ancient areas having remained nearly the same. such evidence of the slow and scarcely sensible mutations of specific forms. will not attempt to account for the close resemblance of distinct species in closely consecutive formations. Consider the prodigious vicissitudes of climate during the pleistocene period. all the intermediate varieties between the species which appeared at the commencement and close of these periods: but we ought to find after intervals. the full meaning of the fossil remains from closely consecutive formations. as we have the right to expect. representative species. in short. though ranked as distinct species. and therefore under the most different climates and conditions. in any one or in any two formations. 353 . is obvious. from its generality. the general resemblance of the organic remains from the several stages of the Chalk formation. On the theory of descent.

as insisted on by Dr. the former would be beaten and exterminated by the latter. modern forms ought. 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. they first appeared. and it seems that this answer must be admitted as true. could be proved to have first come into existence during the Laurentian epoch. there would not have been time sufficient for the development 354 . as well as by the standard of the specialisation of organs. We have also seen that. though difficult of proof. and in this sense higher. that certain Brachiopods have been but slightly modified from an extremely remote geological epoch. not but that it may leave many creatures with simple and unimproved structures fitted for simple conditions of life. so natural selection will tend to render the organisation of each being more specialised and perfect. on the theory of natural selection. In another and more general manner. when arrived at maturity. or the above Brachiopods during the Cambrian formation. from the time when. of their degree of perfection or highness. with which they come into close competition. new species become superior to their predecessors. So that by this fundamental test of victory in the battle for life. It is not an insuperable difficulty that Foraminifera have not. as far as is known. if it included advance in organisation as a necessary contingent. to stand higher than ancient forms. and the palaeozoic by the secondary forms. as yet suggested. Is this the case? A large majority of palaeontologists would answer in the affirmative. On The State Of Development Of Ancient Compared With Living Forms We have seen in the fourth chapter that the degree of differentiation and specialisation of the parts in organic beings. is the best standard. yet leaving such degraded beings better fitted for their new walks of life. and that certain land and fresh-water shells have remained nearly the same. They would likewise be fatal. if the above Foraminifera.5. Carpenter. for they have to beat in the struggle for life all the older forms. It is no valid objection to this conclusion. and in some cases will even degrade or simplify the organisation. We may therefore conclude that if under a nearly similar climate the eocene inhabitants of the world could be put into competition with the existing inhabitants. for instance. as would the secondary by the eocene. progressed in organisation since even the Laurentian epoch. for in this case. as the specialisation of parts is an advantage to each being. for some organisms would have to remain fitted for simple conditions of life.

might beat cephalopods. there is no necessity. The foregoing objections hinge on the question whether we really know how old the world is. and this may well be disputed. as the highest fish. at the present time both groups are greatly reduced. although upon another type?" In the complex struggle for life it is quite credible that crustaceans. at all times imperfect. some look at the selaceans or sharks. high and low. though they will. namely. but formerly selaceans and ganoids alone existed. but a stronger case can 355 . according to the standard of highness chosen. if judged by the most decisive of all trials—the law of battle. At an ancient epoch the highest and lowest molluscoidal animals. 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. so as to hold their places in relation to slight changes in their conditions. does not extend far enough back to show with unmistakable clearness that within the known history of the world organisation has largely advanced. from their approach in some important points of structure to reptiles. during each successive age.of these organisms up to the standard which they had then reached. looking to members of the same class. and in this case. naturalists are not unanimous which forms ought to be ranked as highest: thus. Even at the present day. on the theory of natural selection. others look at the teleosteans as the highest. To attempt to compare members of distinct types in the scale of highness seems hopeless. Beside these inherent difficulties in deciding which forms are the most advanced in organisation. while others. the latter at the present day are largely preponderant in number. The problem whether organisation on the whole has advanced is in many ways excessively intricate. 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 ganoids stand intermediate between the selaceans and teleosteans. have largely increased. and at what period the various forms of life first appeared. The geological record. at two periods. and such crustaceans. would stand very high in the scale of invertebrate animals. cephalopods and brachiopods. the highest molluscs. for their further continued process. When advanced up to any given point. have to be slightly modified. not very high in their own class. so will it be said that fishes have advanced or retrograded in organisation. though not highly developed. intermediate in organisation. consequently some naturalists maintain that molluscs were formerly more highly developed than at present. swarmed in numbers.

if all the productions of New Zealand were set free in Great Britain. the productions of Great Britain stand much higher in the scale than those of New Zealand. Under this point of view. under such extremely complex relations. in the course of successive generations. as a decided advance in the organisation of the world. a multitude of British forms would in the course of time become thoroughly naturalized there. of the former 356 . which implies a great displacement of lower forms. and would exterminate many of the natives. we must believe. at any two periods. are more highly organised than their ancient representatives. could not have foreseen this result. continually adds. by considering the vast reduction of brachiopods. 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. the standard of organisation of the imperfectly-known faunas of successive periods. and have seized on places which must have been previously occupied by the indigenes. any considerable number would be enabled to seize on places now occupied by our native plants and animals. at the present day fifty thousand kinds of vertebrate animals exist. This process. Thus the embryo comes to be left as a sort of picture. from the fact that hardly a single inhabitant of the southern hemisphere has become wild in any part of Europe. we may well doubt whether. whilst it leaves the embryo almost unaltered. and if we knew that at some former period only ten thousand kinds existed. we ought to look at this increase in number in the highest class. We ought also to compare the relative proportional numbers. preserved by nature. more and more difference to the adult. though few in number. We shall appreciate this difficulty more clearly by looking to certain existing faunas and floras. and that the geological succession of extinct forms is nearly parallel with the embryological development of existing forms. In a future chapter I shall attempt to show that the adult differs from its embryo. of the high and low classes throughout the world: if. from an examination of the species of the two countries. owing to variations having supervened at a not early age. for instance. that if all the animals and plants of Great Britain were set free in New Zealand. We thus see how hopelessly difficult it is to compare with perfect fairness. From the extraordinary manner in which European productions have recently spread over New Zealand. Yet the most skilful naturalist. This view accords admirably well with our theory.be made out on the opposite side. and the fact that our existing cephalopods. On the other hand. and having been inherited at a corresponding age.

and yet may never be capable of proof. 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. reptiles. 357 . This view may be true. for instance. it would be vain to look for animals having the common embryological character of the Vertebrata. Seeing. and fishes strictly belong to their proper classes.and less modified condition of the species. that the oldest known mammals. until beds rich in fossils are discovered far beneath the lowest Cambrian strata—a discovery of which the chance is small.

On The Succession Of The Same Types Within The Same Areas. and between the extinct and living brackish water-shells of the Aralo-Caspian Sea. For we know that Europe in ancient times was peopled by numerous marsupials." Professor Owen has subsequently extended the same generalisation to the mammals of the Old World. We see it also in the birds of the caves of Brazil. or that Edentata and other American types should have been solely produced in South America. Other cases could be added. a similar relationship is manifest. what does this remarkable law of the succession of the same types within the same areas mean? He would be a bold man who. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent. in 1839 and 1845. and Professor Owen has shown in the most striking manner that most of the fossil mammals. that in America the law of distribution of terrestrial mammals was formerly different from what it now is. and I have shown in the publications above alluded to. Lund and Clausen in the caves of Brazil. are related to South American types. Mr. for the dissimilarity of the inhabitants of these two continents. We see the same law in this author's restorations of the extinct and gigantic birds of New Zealand. from the wide distribution of most molluscs. even to an uneducated eye. and. on this "law of the succession of types. under the same latitude. and the southern half was formerly more 358 . Now. on the other hand through similarity of conditions.6. North America formerly partook strongly of the present character of the southern half of the continent. In South America. on the one hand through dissimilar physical conditions. I was so much impressed with these facts that I strongly insisted. Woodward has shown that the same law holds good with sea-shells. would attempt to account. it is not well displayed by them. like those of the armadillo. Nor can it be pretended that it is an immutable law that marsupials should have been chiefly or solely produced in Australia. after comparing the present climate of Australia and of parts of South America."—on "this wonderful relationship in the same continent between the dead and the living. in the gigantic pieces of armour. This relationship is even more clearly seen in the wonderful collection of fossil bones made by MM. During The Later Tertiary Periods Mr. for the uniformity of the same types in each continent during the later tertiary periods. as the relation between the extinct and living land-shells of Madeira. buried there in such numbers. found in several parts of La Plata. but.

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. to the northern half. with the genera and species decreasing in numbers as is the case with the Edentata of South America. from Falconer and Cautley's discoveries. as their degenerate descendants. and in a succeeding formation there be six other allied or representative genera. which formerly lived in South America. so that. In a similar manner we know. during the next succeeding period of time. and some of these fossils may have been the actual progenitors of the living species. 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. be found in one geological formation. which constitute the new genera containing the several species. But after very long intervals of time. then we may conclude that generally only one species of each of the older genera has left modified descendants. all the species of the same genus are the descendants of some one species. closely allied though in some degree modified descendants. permitting much intermigration. 359 . but not immutable. is at once explained. have left behind them the sloth. each having eight species. succession of the same types within the same areas. These huge animals have become wholly extinct. and after great geographical changes. armadillo. for the inhabitants of each quarter of the world will obviously tend to leave in that quarter. the feebler will yield to the more dominant forms. If the inhabitants of one continent formerly differed greatly from those of another continent. than it is at present. and anteater. Analogous facts could be given in relation to the distribution of marine animals. On the theory of descent with modification. and this will be a far commoner case. and have left no progeny. so will their modified descendants still differ in nearly the same manner and degree. if six genera. This cannot for an instant be admitted. that Northern India was formerly more closely related in its mammals to Africa than it is at the present time. It must not be forgotten that. each with the same number of species. the great law of the long enduring. In failing orders.closely allied. Or. and there will be nothing immutable in the distribution of organic beings. still fewer genera and species will leave modified blood-descendants. the other seven species of each old genus having died out and left no progeny. on our theory. It may be asked in ridicule whether I suppose that the megatherium and other allied huge monsters.

unless the whole chain could be perfectly restored. will to a large extent explain why—though we do find many links—we do not find interminable varieties. and lastly. and thick enough to outlast future degradation. would be ranked. that only a small portion of the globe has been geologically explored with care. is absolutely as nothing compared with the number of generations which must have passed away even during a single formation. that the duration of each formation is probably short compared with the average duration of specific forms. that each single formation has not been continuously deposited. connecting together all extinct and existing forms by the finest graduated steps. he may overlook how important a part migration has played. preserved in our museums. though many and long as measured by years.7. as a new and distinct species. and during the latter the record will have been least perfectly kept. great intervals of time must have elapsed between most of our successive formations. have been short in comparison with the periods during which each remained in an unchanged condition. He who rejects this view of the imperfection of the geological record. owing to subsidence being almost necessary for the accumulation of deposits rich in fossil species of many kinds. These causes. although each species must have passed through numerous transitional stages. and have oftenest given rise to new species. when the 360 . and more variation during the periods of elevation. Summary Of The Preceding And Present Chapters I have attempted to show that the geological record is extremely imperfect. For he may ask in vain where are the numberless transitional links which must formerly have connected the closely allied or representative species. it is probable that the periods. will rightly reject the whole theory. that only certain classes of organic beings have been largely preserved in a fossil state. which might be found. that widely ranging species are those which have varied most frequently. for it is not pretended that we have any sure criterion by which species and varieties can be discriminated. that the number both of specimens and of species. It should also be constantly borne in mind that any linking variety between two forms. 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. that. that varieties have at first been local. that migration has played an important part in the first appearance of new forms in any one area and formation. that there has probably been more extinction during the periods of subsidence. during which each underwent modification. taken conjointly.

The extinction of old forms is the almost inevitable consequence of the production of new forms. long before that epoch. from their inferiority inherited from a common progenitor. But the utter extinction of a whole group of species has sometimes been a slow process. for the process of modification is necessarily slow. or lie still buried under the ocean. tend to become extinct together. and these will generally succeed in displacing the groups which are their inferiors in the struggle for existence. it never reappears. from the survival of a few descendants. but often falsely apparent. he may urge the apparent. When a group has once wholly disappeared. Passing from these difficulties. The dominant species belonging to large and dominant groups tend to leave many modified descendants. but modified. for the link of generation has been broken. lingering in protected and isolated situations. As these are formed. but I can answer this last question only by supposing that where our oceans now extend they have extended for an enormous period. the world presented a widely different aspect. how species of different classes do not necessarily change together. descendants. sudden coming in of whole groups of species. 361 . as those of Europe. 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 other great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and natural selection. it does not reappear. formed of formations older than any known to us. and to leave no modified offspring on the face of the earth. and that the older continents. yet in the long run that all undergo modification to some extent. after long intervals of time. 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. Groups of species increase in numbers slowly. and endure for unequal periods of time. which form new sub-groups and groups. the species of the less vigorous groups. exist now only as remnants in a metamorphosed condition. or in the same degree. the productions of the world appear to have changed simultaneously. when a species has once disappeared. but that.formations of any one great region. We can understand why. Hence. are considered. and depends on many complex contingencies. and where our oscillating continents now stand they have stood since the commencement of the Cambrian system. or at the same rate. We can thus understand how it is that new species come in slowly and successively.

and are. the more it generally differs from those now living. Extinct and ancient animals resemble to a certain extent the embryos of the more recent animals belonging to the same classes. the geological record be as imperfect as many believe. for the more ancient a form is. but more commonly bringing them only a little closer together. the main objections to the theory of natural selection are greatly diminished or disappear. since become widely divergent. previously classed as distinct into one. Why ancient and extinct forms often tend to fill up gaps between existing forms. and consequently resemble. as it seems to me. and their structure has generally become more specialised. 362 . make together a few grand classes. The more ancient a form is. The inhabitants of the world at each successive period in its history have beaten their predecessors in the race for life. and is intelligible on the principle of inheritance. ancient and recent. all the chief laws of palaeontology plainly proclaim. that species have been produced by ordinary generation: old forms having been supplanted by new and improved forms of life. We can clearly see why the organic remains of closely consecutive formations are closely allied. and it may at least be asserted that the record cannot be proved to be much more perfect. If. On the other hand. that organisation on the whole has progressed. in so far. the products of variation and the survival of the fittest. higher in the scale. for they are closely linked together by generation. from the continued tendency to divergence of character. the common progenitor of groups. the more often it stands in some degree intermediate between groups now distinct. and this wonderful fact receives a simple explanation according to our views. the more nearly it will be related to. why the more ancient a form is. We can clearly see why the remains of an intermediate formation are intermediate in character. but are intermediate only by a long and circuitous course through other extinct and different forms. and this may account for the common belief held by so many palaeontologists. then. sometimes blending two groups. Extinct forms are seldom directly intermediate between existing forms. We can understand. The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious.We can understand how it is that all the forms of life.

Chapter 12 Geographical Distribution 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. and by occasional means — Dispersal during the Glacial period — Alternate Glacial periods in the North and South. 363 .

we meet with the most diversified conditions. 364 . between latitudes 25 and 35 degrees. or obstacles to free migration. humid districts. and western South America. marshes.1. 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. arid deserts. excepting in the northern parts. under almost every temperature. yet it would not be possible to point out three faunas and floras more utterly dissimilar. The case of America alone would almost suffice to prove its truth. South Africa. are related in a close and important manner to the differences between the productions of various regions. if we compare large tracts of land in Australia. the first great fact which strikes us is. almost every author who has studied the subject has come to this conclusion. we may compare the productions of South America south of latitude 35 degrees with those north of 25 degrees. 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. we shall find parts extremely similar in all their conditions. Analogous facts could be given with respect to the inhabitants of the sea. In considering the distribution of organic beings over the face of the globe. yet if we travel over the vast American continent. No doubt small areas can be pointed out in the Old World hotter than any in the New World. Of late. of which the conditions are peculiar in only a slight degree. We see this in the great difference in nearly all the terrestrial productions of the New and Old Worlds. for if we exclude the arctic and northern temperate parts. grassy plains. from the central parts of the United States to its extreme southern point. how widely different are their living productions! In the southern hemisphere. forests. lakes and great rivers. which consequently are separated by a space of ten degrees of latitude. Notwithstanding this general parallelism in the conditions of Old and New Worlds. for it is rare to find a group of organisms confined to a small area. that barriers of any kind. again. but these are not inhabited by a fauna different from that of the surrounding districts. Or. and are exposed to considerably different conditions. that neither the similarity nor the dissimilarity of the inhabitants of various regions can be wholly accounted for by climatal and other physical conditions. A second great fact which strikes us in our general review is. lofty mountains. all authors agree that one of the most fundamental divisions in geographical distribution is that between the New and Old Worlds.

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. as there now is for the strictly arctic productions. and as soon as this is passed we meet in the eastern islands of the Pacific with another and totally distinct fauna. or echinodermata in common. On the other hand. here we have a barrier of another kind. The marine inhabitants of the eastern and western shores of South America are very distinct.where the land almost joins. as the oceans separating continents. a wide space of open ocean extends. we find different productions. are not as impassable. we see the same fact. 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. though the species themselves are distinct at different points and stations. Although so few marine animals are common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands. until. and South America under the same latitude. with extremely few shells. Westward of the shores of America. crustacea. is the affinity of the productions of the same continent or of the same sea. It is a law of the widest generality. also. So that three marine faunas range northward and southward in parallel lines not far from each other. deserts. and every continent offers innumerable 365 . or continuous coasts. and this fact has led naturalists to believe that the isthmus was formerly open. the differences are very inferior in degree to those characteristic of distinct continents. under corresponding climate.. and we have innumerable islands as halting-places. but from being separated from each other by impassable barriers. we encounter no impassable barriers. etc. under a slightly different climate. either of land or open sea. On each continent. though as mountain chains. with not an island as a halting-place for emigrants. We see the same fact in the great difference between the inhabitants of Australia. we come to the shores of Africa. Africa. and over this vast space we meet with no well-defined and distinct marine faunas. but Dr. and where. after travelling over a hemisphere. partly included in the foregoing statement. yet many fishes range from the Pacific into the Indian Ocean. they are almost wholly distinct. or likely to have endured so long. A third great fact. for these countries are almost as much isolated from each other as is possible. Turning to the sea. and of great deserts and even of large rivers. Gunther has recently shown that about thirty per cent of the fishes are the same on the opposite sides of the isthmus of Panama. for on the opposite sides of lofty and continuous mountain-ranges. we find the same law.

the naturalist. specifically distinct. for instance. We see in these facts some deep organic bond. but not quite alike. as far as we positively know.instances. over the same areas of land and water. in travelling. The naturalist must be dull who is not led to inquire what this bond is. as does time for the slow process of modification through natural selection. never fails to be struck by the manner in which successive groups of beings. but they plainly display an American type of structure. as shown in the last chapter. and not by a true ostrich or emu. and sees their nests similarly constructed. He hears from closely allied. that cause which alone. 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. On these same plains of La Plata we see the agouti and bizcacha. the relation of organism to organism in the struggle for life being. The plains near the Straits of Magellan are inhabited by one species of Rhea (American ostrich). nearly alike. from north to south. We may look back to past ages. notes nearly similar. and probably in a subordinate degree to the definite influence of different physical conditions. The bond is simply inheritance. We ascend the lofty peaks of the Cordillera. Nevertheless. replace each other. or. produces organisms quite like each other. however much they may differ in geological structure. and we find an alpine species of bizcacha. and we find American types then prevailing on the American continent and in the American seas. as we see in the case of varieties. If we look to the islands off the American shore. the most important of all relations. animals having nearly the same habits as our hares and rabbits. 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. and belonging to the same order of Rodents. the inhabitants are essentially American. Innumerable other instances could be given. though they may be all peculiar species. though nearly related. with eggs coloured in nearly the same manner. as I have already often remarked. The dissimilarity of the inhabitants of different regions may be attributed to modification through variation and natural selection. throughout space and time. and northward the plains of La Plata by another species of the same genus. Thus the high importance of barriers comes into play by checking migration. and we do not find the beaver or muskrat. independently of physical conditions. yet distinct kinds of birds. we look to the waters. like those inhabiting Africa and Australia under the same latitude. rodents of the South American type. Widely-ranging 366 . but the coypu and capybara.

and have not become greatly or at all modified. In their new homes they will be exposed to new conditions. It is also obvious that the individuals of the same species. when they spread out into new countries. so certain species have migrated over vast spaces. almost any amount of migration is possible. as is so commonly and notoriously the case. little modification. as they are descended from the same progenitor. In the case of those species which have undergone. during whole geological periods. will have the best chance of seizing on new places. must have proceeded from one spot. after having long competed with each other in their old home. If a number of species. for neither migration nor isolation in themselves effect anything. for during the vast geographical and climatical changes which have supervened since ancient times. and will produce groups of modified descendants. only so far as it profits each individual in its complex struggle for life. so the amount of modification in different species will be no uniform quantity. whole genera. According to these views. and even families. and will frequently undergo further modification and improvement. of the existence of any law of necessary development. are confined to the same areas.species. it is incredible that 367 . As we have seen in the last chapter that some forms have retained nearly the same character from an enormously remote geological period. There is no evidence. On this principle of inheritance with modification we can understand how it is that sections of genera. in which we have reason to believe that the species of a genus have been produced within comparatively recent times. there is great difficulty on this head. though now inhabiting distant and isolated regions. there is not much difficulty in believing that they have migrated from the same region. As the variability of each species is an independent property. were to migrate in a body into a new and afterwards isolated country. abounding in individuals. But in many other cases. where their parents were first produced: for. and will be taken advantage of by natural selection. which have already triumphed over many competitors in their own widely-extended homes. it is obvious that the several species of the same genus. though inhabiting the most distant quarters of the world. as has been explained. 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. and thus they will become still further victorious. as was remarked in the last chapter. must originally have proceeded from the same source. they would be little liable to modification.

368 .individuals identically the same should have been produced from parents specifically distinct.

and have not been able to migrate to the opposite side. accordingly. namely. The incapacity of migrating across a wide sea is more clear in the case of terrestrial mammals than perhaps with any other organic beings. and that when a plant or animal inhabits two points so distant from each other. and it has been observed by several naturalists that the most natural genera. But if the same species can be produced at two separate points. the fact is given as something remarkable and exceptional. a still greater number of sections of genera. and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres? The answer. or if they have a wide range that their range is continuous. Single Centres Of Supposed Creation We are thus brought to the question which has been largely discussed by naturalists. The great and striking influence of barriers of all kinds. is intelligible only on the view that the great majority of species have been produced on one side. have migrated across the wide and broken interspaces. very many genera. Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. No geologist feels any difficulty in Great Britain possessing the same quadrupeds with the rest of Europe. that mammals have not been able to migrate. whether species have been created at one or more points of the earth's surface. He who rejects it. and. for they were no doubt once united.2. Some few families. 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. are generally confined to the same country. as I believe. are confined to a single region. whereas some plants. and calls in the agency of a miracle. so that a multitude of European animals and plants have become naturalised in America and Australia. that the space could not have been easily passed over by migration. we find no inexplicable instances of the same mammals inhabiting distant points of the world. that in most cases the area inhabited by a species is continuous. rejects the vera causa of ordinary generation with subsequent migration. or those genera in which the species are most closely related to each other. where now found. why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same. many subfamilies. or with an interval of such a nature. What a strange anomaly it would be if a directly opposite rule were to prevail when we go down one step lower in the 369 . from their varied means of dispersal. is. It is universally admitted.

namely. our general view will be much strengthened. undergoing modification during their migration from some one area. confined to some one region! Hence. and these had not been. when most of the species inhabiting one region are different from those of another region. can have migrated. and having subsequently migrated from that area as far as its powers of migration and subsistence under past and present conditions permitted. is the most probable. the occurrence of the same terrestrial species on islands and on the nearest mainland. considering our ignorance with respect to former climatical and geographical changes. for the explanation is obvious on the 370 . as it has to many other naturalists. and to the various occasional means of transport. But the geographical and climatical changes which have certainly occurred within recent geological times. must have rendered discontinuous the formerly continuous range of many species. that the view of each species having been produced in one area alone. it can be shown that migration from the one region to the other has probably occurred at some former period. In discussing this subject we shall be enabled at the same time to consider a point equally important for us. and has migrated thence as far as it could. though separated by hundreds of miles of open sea. now living at distant and separated points. then. rendered probable by general considerations. nor do I for a moment pretend that any explanation could be offered of many instances. But. and at distant points in the Arctic and Antarctic regions. the wide distribution of fresh water productions. the existence of the same species on the summits of distant mountain ranges. after some preliminary remarks. whether the several species of a genus which must on our theory all be descended from a common progenitor. 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. namely to the individuals of the same species. I will discuss a few of the most striking classes of facts. If. that we ought to give up the belief. that each species has been produced within one area. though closely allied to them. namely. it seems to me. and of so grave a nature. So that we are reduced to consider whether the exceptions to continuity of range are so numerous.series. and secondly (in the following chapter). the belief that a single birthplace is the law seems to me incomparably the safest. It would be hopelessly tedious to discuss all the exceptional cases of the same species. Undoubtedly many cases occur in which we cannot explain how the same species could have passed from one point to the other. and thirdly. at least at first.

at each successive stage of modification. but have never blended with other individuals or varieties of the same species. A volcanic island. and are. all the individuals of the same form will be descended from a single parent. as some authors suppose. the individuals of the same species inhabiting the same area will be kept nearly uniform by intercrossing." I must say a few words on the means of dispersal. and the whole amount of modification at each stage will not be due to descent from a single parent. or whether. as we shall hereafter see. or which occasionally intercross. But in the great majority of cases. does not differ much from that advanced by Mr. To illustrate what I mean: our English race-horses differ from the horses of every other breed. Wallace. but they do not owe their difference and superiority to descent from any single pair. and their descendants. upheaved and formed at the distance of a few hundreds of miles from a continent. so that many individuals will go on simultaneously changing. for instance. which I have selected as presenting the greatest amount of difficulty on the theory of "single centres of creation." And it is now well known that he attributes this coincidence to descent with modification. Before discussing the three classes of facts. 371 . With organic beings which never intercross. The question of single or multiple centres of creation differs from another though allied question. each species. would still be related by inheritance to the inhabitants of that continent. Cases of this nature are common. though modified. with all organisms which habitually unite for each birth. so that.principle of descent with modification. who concludes that "every species has come into existence coincident both in space and time with a pre-existing closely allied species. if such exist. namely. that have supplanted each other. must be descended from a succession of modified varieties. but to continued care in the selecting and training of many individuals during each generation. inexplicable on the theory of independent creation. whether all the individuals of the same species are descended from a single pair. This view of the relation of the species of one region to those of another. from many individuals simultaneously created. or single hermaphrodite. namely. would probably receive from it in the course of time a few colonists.

and thus have allowed terrestrial productions to pass from one to the other. and united almost every island with some mainland. A region now impassable to certain organisms from the nature of its climate. Means Of Dispersal Sir C. and the two faunas will now blend together. It seems to me that we have abundant evidence of great oscillations in the level of the land or sea.3. or may formerly have blended. when the climate was different. presently have to discuss this branch of the subject in some detail. 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. Edward Forbes insisted that all the islands in the Atlantic must have been recently connected with Europe or Africa. but not of such vast changes in the position and extension of our continents. I freely admit the former existence of many islands. we shall be enabled to speculate with security on the former extension of the land. and removes many a difficulty. however. now buried beneath the sea. have been 372 . but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. Change of climate must have had a powerful influence on migration. as it will some day be. If. Where the sea now extends. the arguments used by Forbes are to be trusted. I can give here only the briefest abstract of the more important facts. Other authors have thus hypothetically bridged over every ocean. 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 islands. land may at a former period have connected islands or possibly even continents together. it must be admitted that scarcely a single island exists which has not recently been united to some continent. that each species has proceeded from a single birthplace. indeed. Whenever it is fully admitted. and when in the course of time we know something definite about the means of distribution. In the coral-producing oceans such sunken islands are now marked by rings of coral or atolls standing over them. This view cuts the Gordian knot of the dispersal of the same species to the most distant points. which may have served as halting places for plants and for many animals during their migration. I shall. and Europe likewise with America. or let it formerly have been submerged. Lyell and other authors have ably treated this subject. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas. submerge it. might have been a high road for migration.

For convenience sake I chiefly tried small seeds without the capsules or fruit.continuously. Nor does the almost universally volcanic composition of such islands favour the admission that they are the wrecks of sunken continents. it was not even known how far seeds could resist the injurious action of sea-water. this or that plant is often stated to be ill adapted for wide dissemination. capsules. metamorphic schists. and some of these floated for a long time. sixtyfour germinated after an immersion of twenty-eight days. Hence I was led to dry the stems and branches of ninety-four plants with ripe 373 . with Mr. they resisted the salt-water badly. but which more properly should be called occasional means of distribution. like other mountain summits. and with the many existing oceanic islands. Berkeley's aid. To my surprise I found that out of eighty-seven kinds. with one exception. 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. were all killed by a month's immersion. The nature and relative proportions of the inhabitants of oceanic islands are likewise opposed to the belief of their former continuity of continents. and a few survived an immersion of 137 days. and. and as all of these sank in a few days. Afterwards I tried some larger fruits. they could not have been floated across wide spaces of the sea. a few experiments. In botanical works. It deserves notice that certain orders were far more injured than others: nine Leguminosae were tried. 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.. of granite. instead of consisting of mere piles of volcanic matter. It is well known what a difference there is in the buoyancy of green and seasoned timber. but the greater or less facilities for transport across the sea may be said to be almost wholly unknown. Until I tried. old fossiliferous and other rocks. Hydrophyllaceae and Polemoniaceae. 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. some at least of the islands would have been formed. as are necessary on the view advanced by Forbes and admitted by his followers. I shall here confine myself to plants. or almost continuously united with each other. seven species of the allied orders. whether or not they were injured by salt water. I must now say a few words on what are called accidental means. etc. if they had originally existed as continental mountain ranges.

after being dried. Subsequently to my experiments. mostly different from mine. and this. The fact of the larger fruits often floating longer than the small. from plants which live near the sea. but in a much better manner. the seeds of 14/100 plants belonging to one country might be floated across 924 miles of sea to another country. he did not previously dry the plants or branches with the fruit. as we have seen. when dried it floated for eighty-five days. as far as anything can be inferred from these scanty facts. for instance. The majority sank quickly. and to place them on sea-water. M. it would perhaps be safer to assume that the seeds of about 10/100 plants of a flora. after having been dried. Altogether. on this average. and when stranded. and were then capable of germination. we may conclude. an asparagus plant with ripe berries floated for twenty-three days. The result was that 18/98 of his seeds of different kinds floated for forty-two days. 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). On the other hand. In Johnston's Physical Atlas. and would then germinate. So that as 64/87 kinds of seeds germinated after an immersion of twenty-eight days. He tried ninety-eight seeds. if blown by an inland gale to a favourable spot. whilst green. that the seeds of 14/ 100 kinds of plants of any country might be floated by sea-currents during twenty-eight days. and as 18/94 distinct species with ripe fruit (but not all the same species as in the foregoing experiment) floated. and some of the eighteen floated for a very much longer period. so that they were alternately wet and exposed to the air like really floating plants. but some which.fruit. for he placed the seeds in a box in the actual sea. and likewise seeds. would germinate. and afterwards germinated. and afterwards when planted germinated. is 374 . for above twenty-eight days. but he chose many large fruits. could be floated across a space of sea 900 miles in width. and this would have favoured both the average length of their flotation and their resistance to the injurious action of the salt-water. when dried floated much longer. and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days. eighteen floated for above twenty-eight days. ripe hazel-nuts sank immediately. Therefore. and would retain their power of germination. would have caused some of them to have floated much longer. 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. floated for a very short time. out of the ninety-four dried plants. when dried they floated for above ninety days. but when dried they floated for ninety days. Martens tried similar ones.

out of the excrement of small birds. In the course of two months. Again. now. as Alph. when floating on the sea. I can show that the carcasses of birds. Some hawks and owls bolt their prey whole. Living birds can hardly fail to be highly effective agents in the transportation of seeds. de Candolle has shown. could hardly be transported by any other means. Seeds may be occasionally transported in another manner. after a bird has found and devoured a large supply of food. and these seemed perfect. it is positively asserted that all the grains do not pass into the gizzard for twelve or even eighteen hours. are killed by even a few days' immersion in sea-water. I have never seen an instance of nutritious seeds passing through the intestines of a bird. three dicotyledonous plants germinated: I am certain of the accuracy of this observation. solely from the roots of drifted trees. as plants with large seeds or fruit which. sometimes escape being immediately devoured. these stones being a valuable royal tax. for instance. disgorge pellets. I find that when irregularly shaped stones are embedded in the roots of trees. and the natives of the coral islands in the Pacific procure stones for their tools. to my surprise nearly all germinated. which had floated on artificial sea-water for thirty days. generally have restricted ranges. But the following fact is more important: the crops of birds do not secrete gastric juice. Drift timber is thrown up on most islands. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the ocean. A bird in this interval might easily be blown to the distance of five hundred miles. injure in the least the germination of seeds. and hawks are known to look out for tired birds. and many kinds of seeds in the crops of floating birds long retain their vitality: peas and vetches. and some authors have given a far higher estimate. 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. germinated.interesting. which were tried. as I know from experiments made 375 . and do not. and the contents of their torn crops might thus readily get scattered. but hard seeds of fruit pass uninjured through even the digestive organs of a turkey. which. but some taken out of the crop of a pigeon. We may safely assume that under such circumstances their rate of flight would often be thirty-five miles an hour. even on those in the midst of the widest oceans. small parcels of earth are very frequently enclosed in their interstices and behind them. as I know by trial. I picked up in my garden twelve kinds of seeds. and after an interval of from twelve to twenty hours. and some of them.

T. Although the beaks and feet of birds are generally clean. Mr. Lyell that in November. The Rev. with a little cake of dry earth attached to the shank. I forced many kinds of seeds into the stomachs of dead fish. storks. though on insufficient evidence. Now. 376 . and this contained a seed of the toad-rush (Juncus bufonius) which germinated and flowered. canary. such as that which visited Madeira. of two genera. Fresh-water fish. I find. in parts of Natal it is believed by some farmers. after an interval of many hours. belonging to two species. Weale sent me in a letter a small packet of the dried pellets. that injurious seeds are introduced into their grass-land in the dung left by the great flights of locusts which often visit that country. Hence a swarm of locusts. and at night alighted on the taller trees. which were completely coated with them.in the Zoological Gardens. eat seeds of many land and water plants. Some seeds of the oat. and two seeds of beet grew after having been thus retained for two days and fourteen hours. swarms of locusts visited the island of Madeira. wheat. these birds. and have heard of others caught at greater distances. might readily be the means of introducing several kinds of plants into an island lying far from the mainland. of Brighton. They were in countless numbers. either rejected the seeds in pellets or passed them in their excrement. include seeds capable of germination. millet. at least five or six miles in diameter. clover. and in the earth there was a pebble as large as the seed of a vetch. R. In consequence of this belief Mr. and in another case twenty-two grains of dry argillaceous earth from the foot of a partridge. and raised from them seven grass plants. as suddenly as they had appeared. fish are frequently devoured by birds. out of which I extracted under the microscope several seeds. however. and pelicans. 1844. Certain seeds. were always killed by this process. and have not since visited the island. and then gave their bodies to fishing-eagles. During two or three days they slowly careered round and round in an immense ellipse. hemp. and extended upward as far as could be seen with a telescope. I myself caught one 370 miles from the coast of Africa. and several of these seeds retained the power of germination. They then disappeared over the sea. Swaysland. as thick as the flakes of snow in the heaviest snowstorm. weighing only nine grains. Locusts are sometimes blown to great distances from the land. and thus the seeds might be transported from place to place. earth sometimes adheres to them: in one case I removed sixty-one grains. Here is a better case: the leg of a woodcock was sent to me by a friend. and beet germinated after having been from twelve to twenty-one hours in the stomachs of different birds of prey. Lowe informed Sir C.

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. bones. including the common oat. judging from the young leaves. Hence we may safely infer that icebergs formerly landed their rocky burdens on the shores of these mid-ocean islands. and at least one kind of grass. I suspected that these islands had been partly stocked by ice-borne seeds during the Glacial epoch. 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. but when broken. and (as remarked by Mr.C. As icebergs are known to be sometimes loaded with earth and stones. and of seventy dicotyledons. Hartung to inquire whether he had observed erratic boulders on these islands. H. Many facts could be given showing how generally soil is charged with seeds. have been in action year after year for tens of thousands of years. and which annually migrate—for instance. and that other means. Professor Newton sent me the leg of a red-legged partridge (Caccabis rufa) which had been wounded and could not fly. from the large number of plants common to Europe. which consisted. which do not occur in the archipelago. wheatears. in comparison with the species on the other islands of the Atlantic. In the Azores. informs me that he has often shot wagtails (Motacillae). and whinchats (Saxicolae). and the nest of a land-bird. and weighing six and a half ounces. no less than eighty-two plants sprung from it: these consisted of twelve monocotyledons. and he has several times noticed little cakes of earth attached to their feet. watered and placed under a bell glass. be a 377 . and he answered that he had found large fragments of granite and other rocks. Considering that these several means of transport. With such facts before us. At my request Sir C. which stand nearer to the mainland. which without doubt remain to be discovered. of at least three distinct species. and it is at least possible that they may have brought thither the seeds of northern plants. and have even carried brushwood. can we doubt that the many birds which are annually blown by gales across great spaces of ocean. before they had alighted. it would. Watson) from their somewhat northern character. I think. on their first arrival on our shores. with a ball of hard earth adhering to it.who during the last forty years has paid close attention to our migratory birds. Lyell wrote to M. it can hardly be doubted that they must occasionally. in comparison with the latitude. For instance. The earth had been kept for three years.

immigrants from Europe or any other continent. as to become naturalised. like Great Britain. though standing more remote from the mainland. namely. These means. would germinate and survive. would suffice for occasional transport across tracts of sea some hundred miles in breadth. or from island to island. where. but seeds could be transported by these rare wanderers only by one means. if fitted for the climate. The currents. but this is not strictly correct: the currents of the sea are not accidental. during the long lapse of geological time. if not killed by their very long immersion in salt water. It should be observed that scarcely any means of transport would carry seeds for very great distances. and coming to maturity! But it would be a great error to argue that because a well-stocked island. one or two land-birds are blown across the whole Atlantic Ocean. however. On almost bare land. and before it had become fully stocked with inhabitants. though they might and do bring seeds from the West Indies to our western shores. would never bring seeds from North America to Britain. Out of a hundred kinds of seeds or animals transported to an island. perhaps not more than one would be so well fitted to its new home.marvellous fact if many plants had not thus become widely transported. has not. nearly every seed which chanced to arrive. whilst the island was being upheaved. 378 . But this is no valid argument against what would be effected by occasional means of transport. received within the last few centuries. Even in this case. These means of transport are sometimes called accidental. even if far less well-stocked than Britain. or from a continent to a neighbouring island. nor could they be long carried in the crops or intestines of birds. by dirt adhering to their feet or beaks. for seeds do not retain their vitality when exposed for a great length of time to the action of sea water. but not from one distant continent to another. but would remain as distinct as they now are. how small would be the chance of a seed falling on favourable soil. from their course. Almost every year. through occasional means of transport. would not receive colonists by similar means. they could not endure our climate. as far as is known (and it would be very difficult to prove this). which is in itself a rare accident. The floras of distant continents would not by such means become mingled. that a poorlystocked island. nor is the direction of prevalent gales of wind. from North America to the western shores of Ireland and England. with few or no destructive insects or birds living there.

covering the central 379 . with their scored flanks. as we shall immediately see. with those on the loftiest mountains of Europe. and perched boulders. but it is far more remarkable. polished surfaces. as explained by Edward Forbes. by supposing a new glacial period slowly to come on. Dispersal During The Glacial Period The identity of many plants and animals. as formerly occurred. So greatly has the climate of Europe changed. separated from each other by hundreds of miles of lowlands. left by old glaciers. we should have an arctic fauna and flora. By the time that the cold had reached its maximum. As the cold came on. affords a simple explanation of these facts. that the plants on the White Mountains. The mountains would become covered with snow and ice. in the United States of America. are now clothed by the vine and maize. within a very recent geological period. at the same time would travel further and further southward. is one of the most striking cases known of the same species living at distant points. Even as long ago as 1747. is substantially as follows. without the apparent possibility of their having migrated from one point to the other. gigantic moraines. erratic boulders and scored rocks plainly reveal a former cold period. central Europe and North America suffered under an Arctic climate. unless they were stopped by barriers. We have evidence of almost every conceivable kind. The ruins of a house burnt by fire do not tell their tale more plainly than do the mountains of Scotland and Wales. But we shall follow the changes more readily. such facts led Gmelin to conclude that the same species must have been independently created at many distinct points. organic and inorganic. and we might have remained in this same belief.4. on mountain-summits. that in Northern Italy. and then pass away. where Alpine species could not possibly exist. of the icy streams with which their valleys were lately filled. had not Agassiz and others called vivid attention to the Glacial period. The latter. 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. and in the extreme northern parts of Europe. as we hear from Asa Gray. in which case they would perish. that. which. and nearly all the same. Throughout a large part of the United States. 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. are all the same with those of Labrador. and their former Alpine inhabitants would descend to the plains. The former influence of the glacial climate on the distribution of the inhabitants of Europe.

that a colder climate formerly permitted their migration across the intervening lowlands. As the arctic forms moved first southward and afterwards backward to the north. These views. those of the United States to Labrador. 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. in unison with the changing climate. without other evidence. as remarked by Mr. Hence. which had lately lived together on the European and North American lowlands. that when in other regions we find the same species on distant mountain-summits. 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. and those of the Pyrenees. H. as the warmth increased and the snow still further disappeared. grounded as they are on the perfectly well-ascertained occurrence of a former Glacial period. those of the mountains of Siberia to the arctic regions of that country. when the warmth had fully returned. their mutual relations 380 . are remarkably uniform round the world.C. the arctic forms would retreat northward. we may almost conclude. seem to me to explain in so satisfactory a manner the present distribution of the Alpine and Arctic productions of Europe and America. as far south as the Alps and Pyrenees. Watson. and as they all migrated in a body together. whilst their brethren were pursuing their northern journey.parts of Europe. and on many isolated mountain-summits far distant from each other. The Alpine plants. closely followed up in their retreat by the productions of the more temperate regions. now become too warm for their existence. for example. of Scotland. for the present circumpolar inhabitants. 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. As the warmth returned. are more especially allied to the plants of northern Scandinavia. And as the snow melted from the bases of the mountains. would again be found in the arctic regions of the Old and New Worlds. which we suppose to have everywhere travelled southward. they will not have been exposed during their long migrations to any great diversity of temperature. would generally have been due south and north. and the re-migration on the returning warmth. the same species. as remarked by Ramond. always ascending. and even stretching into Spain. higher and higher. the arctic forms would seize on the cleared and thawed ground.

first at the bases and ultimately on the summits of the mountains. these forms will not have been liable to much modification. in all probability. which must have existed on the mountains before the commencement of the Glacial epoch. lies in the nature of the climate before the commencement of the Glacial period. The explanation. and have survived there ever since. the case will have been somewhat different. Their mutual relations will thus have been in some degree disturbed. have been subsequently exposed to somewhat different climatical influences. consequently they will have been liable to modification. left isolated from the moment of the returning warmth. and it may be asked how I account for this degree of uniformity of the sub-arctic and temperate forms round the world. and which during the coldest period will have been temporarily driven down to the plains. in accordance with the principles inculcated in this volume. 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. 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. Hence. some exist as varieties. and they have been modified. have become mingled with ancient Alpine species. In the foregoing illustration. At the present day. one with another. and we have good reason to believe that the climate was 381 . the newer Pliocene period. so that it may well be asked how the same species could then or previously have entered the two continents. I believe. they must have been still more completely separated from each other by wider spaces of ocean. they will. for it is not likely that all the same arctic species will have been left on mountain ranges far distant from each other. when the inhabitants of the Old and New Worlds lived further southwards than they do at present. they will also. During the Glacial period. some as doubtful forms or sub-species and some as distinct yet closely allied species representing each other on the several ranges. at the commencement of the real Glacial period. But with the Alpine productions. the majority of the inhabitants of the world were specifically the same as now. At this. I have assumed that at the commencement of our imaginary Glacial period. also. But it is also necessary to assume that many sub-arctic and some few temperate forms were the same round the world. the arctic productions were as uniform round the polar regions as they are at the present day.will not have been much disturbed. for if we compare the present Alpine plants and animals of the several great European mountain ranges. though many of the species remain identically the same.

to eastern America. from reasons before alluded to. and that the present arctic productions then lived on the broken land still nearer to the pole. at a period anterior to the Glacial epoch. as far as the more temperate productions are concerned. as soon as the species in common. migrated south of the Polar Circle. And this continuity of the circumpolar land. they will have been completely cut off from each other. mostly in a modified condition. will account for the supposed uniformity of the sub-arctic and temperate productions of the Old and New Worlds. During the slowly decreasing warmth of the Pliocene period. 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. and to infer that during some earlier and still warmer period. serving as a bridge. Now. began slowly to migrate southwards as the climate became less warm. in latitude 66-67 degrees. and that these plants and animals. and their separation by the whole Atlantic Ocean. we see under the Polar Circle that there is almost continuous land from western Europe through Siberia. both in the Old and New Worlds. they will have become mingled in the one great region with the native American productions. long before the commencement of the Glacial period. Believing. under the Polar Circle. and in the other great region. lived during the Pliocene period further north. As the plants and animals migrated southward. that our continents have long remained in nearly the same relative position. since rendered impassable by cold. though subjected to great oscillations of level. we may suppose that the organisms which now live under latitude 60 degrees.warmer than at the present day. as I believe. if we look at a terrestrial globe. for the intermigration of their inhabitants. We now see. such as the older Pliocene period. This separation. with the consequent freedom under a more favourable climate for intermigration. must have taken place long ages ago. and would have had to compete with them. with those of 382 . which inhabited the New and Old Worlds. for during these warmer periods the northern parts of the Old and New Worlds will have been almost continuously united by land. considering the distance of the two areas. On this view we can understand the relationship with very little identity. their descendants. a large number of the same plants and animals inhabited the almost continuous circumpolar land. in the central parts of Europe and the United States. Hence. I am strongly inclined to extend the above view. between the productions of North America and Europe—a relationship which is highly remarkable.

and a host of closely allied or representative forms which are ranked by all naturalists as specifically distinct. still existing and extinct tertiary forms. will account. on the several mountain ranges and on the arctic lands of Europe and North America. for instance. that when we compare the now living productions of the temperate regions of the New and Old Worlds. I think. a slow southern migration of a marine fauna. some fish and other marine animals. within a much more recent period. left isolated. Consequently we have here everything favourable for much modification—for far more modification than with the Alpine productions. We cannot maintain that such species have been created alike. As on the land. These cases of close relationship in species either now or formerly inhabiting the seas on the eastern and western shores of North America.the Old World. we can understand the presence of some closely allied. so in the waters of the sea. the Mediterranean and Japan. are inexplicable on the theory of creation. and the temperate lands of North America and Europe. Hence. on the theory of modification. it has come. we find very few identical species (though Asa Gray has lately shown that more plants are identical than was formerly supposed). and others as distinct species. during the Pliocene or even a somewhat earlier period. which. we see countries closely similar in all their physical conditions. for many closely allied forms now living in marine areas completely sundered. for if we compare. was nearly uniform along the continuous shores of the Polar Circle. which some naturalists rank as geographical races. certain parts of South America with parts of South Africa or Australia. 383 . in correspondence with the nearly similar physical conditions of the areas. with their inhabitants utterly dissimilar. Thus. and the still more striking fact of many closely allied crustaceans (as described in Dana's admirable work). on the eastern and western shores of temperate North America. 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. but we find in every great class many forms.

Dr. once formed a huge moraine. that in New Zealand immense glaciers formerly descended to a low level. it appears also that there are traces of former glacial action on the mountains of the south.5.000 feet down the valleys. at points 900 miles apart. including grooved pebbles. at about the height of 12. In the Cordillera of South America. we have the clearest evidence of former glacial action. W. glaciers have left the marks of their former low descent. In Europe we meet with the plainest evidence of the Glacial period. nearly under the equator. Hooker on widely separated mountains in this island tell the same story of a former cold period. crossing the Portillo valley. Forbes informs me that he found in various parts of the Cordillera. from the excellent researches of Dr. 384 . glaciers once extended far below their present level. also. and likewise great masses of detritus. resembling those with which he was familiar in Norway. and the same plants. from latitude 41 degrees to the southernmost extremity. we know. Along the Himalaya. Hooker. according to Dr. Southward of the Asiatic continent.000 feet. and southward to the Pyrenees. in numerous immense boulders transported far from their parent source. J. there can hardly be a doubt. from latitude 13 to 30 degrees south. In the Lebanon. been noticed on the Rocky Mountains. Looking to America: in the northern half. as far south as latitude 46 degrees. perpetual snow formerly covered the central axis. Further south. on both sides of the continent. and fed glaciers which rolled 4. Hector. deeply-furrowed rocks. as far south as latitude 36 and 37 degrees. found by Dr. D. on the opposite side of the equator. where the climate is now so different. Alternate Glacial Periods In The North And South But we must return to our more immediate subject. and in Sikkim. and Mr. Clarke. In central Chile I examined a vast mound of detritus with great boulders. Hooker saw maize growing on ancient and gigantic moraines. Erratic boulders have.eastern corner of Australia. Along this whole space of the Cordillera true glaciers do not now exist even at much more considerable heights. I am convinced that Forbes's view may be largely extended. from the western shores of Britain to the Ural range. We may infer from the frozen mammals and nature of the mountain vegetation. Haast and Dr. ice-borne fragments of rock have been observed on the eastern side of the continent.B. From facts communicated to me by the Rev. The same observer has recently found great moraines at a low level on the Atlas range in North Africa. that Siberia was similarly affected. and on the shores of the Pacific. which.

In South America. On the Silla of Caraccas 385 . as may be inferred from the amount of work effected—and lastly. cold periods regularly recur every ten or fifteen thousand years. between forty and fifty of the flowering plants of Tierra del Fuego. but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. some Antarctic and some Andean genera were found by Gardner which do not exist in the low intervening hot countries. owing to certain contingencies. Dr. chiefly through changes in the direction of the ocean currents. brought into operation by an increase in the eccentricity of the earth's orbit. as Sir C. arrived at by Mr. and these at long intervals are extremely severe. Mr. But now. Croll believes that the last great glacial period occurred about 240. from direct evidence. and endured. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur. not to mention still more ancient formations. Croll.000 years ago. with slight alterations of climate. while the southern passes through a glacial period. but I will first give the facts which demand an explanation. According to Mr.From these several facts. forming no inconsiderable part of its scanty flora.000 years. several geologists are convinced. 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. that such occurred during the miocene and eocene formations. 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. in a series of admirable memoirs. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it. 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. for about 160. are common to North America and Europe. Hooker has shown that besides many closely allied species. But the most important result for us. of which the most important. So conversely it will be with the northern hemisphere. namely. has attempted to show that a glacial condition of climate is the result of various physical causes. enormously remote as these areas in opposite hemispheres are from each other. Croll. On the Organ Mountains of Brazil some few temperate European. All these causes tend towards the same end. Lyell has shown. With respect to more ancient glacial periods. from glaciers having recently descended to a low level along the whole line of the Cordillera. Croll. Mr. is the relative position of the land and water.

Muller has discovered several European species. and a long list can be given. several forms characteristic of Europe. It now also appears. raises a picture of a collection made on a hillock in Europe. and likewise to those of temperate Europe. Still more striking is the fact that peculiar Australian forms are represented by certain plants growing on the summits of the mountains of Borneo. Lowe on the mountains of the Cape Verde Islands. but not in the intermediate torrid regions. and some few representatives of the flora of the Cape of Good Hope. occur on the lowlands. as I hear from Dr.T. and at the same time representing plants of Europe not found in the intervening hot lowlands. It should. and on the temperate plains of the north and south. Dr. Some of these Australian forms. F. as I hear from Dr. in the Gulf of Guinea. across the whole continent of Africa and to the mountains of the Cape Verde archipelago. are either the same species or varieties of the same species. Hence. be observed that 386 . we see that certain plants growing on the more lofty mountains of the tropics in all parts of the world. Hooker. analogous and striking facts are given in regard to the plants of that large island. Hooker. that some of these same temperate plants have been discovered by the Rev. of European genera. many plants occur either identically the same or representing each other. as I am informed by Dr. on the heights of Ceylon. and are thinly scattered on the one hand over India. and on the isolated mountain ranges of the peninsula of India. In the admirable "Introduction to the Flora of New Zealand. however. not introduced by man.the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera. Hooker has also lately shown that several of the plants living on the upper parts of the lofty island of Fernando Po. On the southern mountains of Australia. Hooker. R. and on the volcanic cones of Java. This extension of the same temperate forms. other species. believed not to have been introduced by man. A list of the genera of plants collected on the loftier peaks of Java. found in Australia. are closely related to those on the mountains of Abyssinia." by Dr. and on the neighbouring Cameroon Mountains. occur on the mountains of Abyssinia. almost under the equator. Hooker. and on the other hand as far north as Japan. In Africa. At the Cape of Good Hope a very few European species. On the Himalaya. Dr. is one of the most astonishing facts ever recorded in the distribution of plants. extend along the heights of the peninsula of Malacca. and on the mountains several representative European forms are found which have not been discovered in the intertropical parts of Africa.

the lowlands of these great continents were everywhere tenanted under the equator by a considerable number of temperate forms. of northern forms of fish. in the Gulf of Guinea. it appears almost certain that at some former period. Prof. Dr. "with forms of the torrid zone harmoniously blended with those of the temperate. 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. During this. Seemann found the vegetation like that of Mexico. but some few analogous facts could be given in regard to terrestrial animals. its antipode. but have not been found in the intermediate tropical seas. 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. etc. Croll's conclusion that when the northern hemisphere suffered from the extreme cold of the great Glacial period. the coldest period. the southern hemisphere was actually warmer." Sir J. belong to genera not now found in the intermediate tropical lowlands." Besides these identical and closely allied forms. Mann found temperate European forms beginning to appear at the height of about five thousand feet. at the height of only two thousand feet. no doubt during the most severe part of a Glacial period. Richardson. than to any other part of the world. Hooker informs me that twenty-five species of Algae are common to New Zealand and to Europe." Now let us see whether Mr. Tasmania. and along the Peninsula of India. Watson has remarked.. Mr. the Alpine or mountain flora really become less and less Arctic. many species inhabiting the same widely sundered areas. to Ceylon and the Malay Archipelago. "in receding from polar toward equatorial latitudes. for. In marine productions. but with perhaps a still greater preponderance of temperate forms. Dr. Dana.these plants are not strictly arctic forms. I may quote a statement by the highest authority. namely. These brief remarks apply to plants alone. On the mountains of Panama. and in a less wellmarked manner across the wide expanse of tropical South America. the presence of temperate forms on the highlands across the whole of equatorial Africa. From the foregoing facts. H.C. also. as an example. So again in the mountainous island of Fernando Po. speaks of the reappearance on the shores of New Zealand. similar cases likewise occur. or perhaps even rather cooler. the lowlands under the equator must have been clothed with a mingled tropical and temperate vegetation. as Mr. that "it is certainly a wonderful fact that New Zealand should have a closer resemblance in its crustacea to Great Britain. throws any clear light on 387 .

Some. of the northern temperate forms would almost certainly have ascended any adjoining high land. we know that Arctic forms invaded the temperate regions. they would be eminently liable to modification. they would have long survived like the Arctic forms on the mountains of Europe. and would generally 388 . as measured by years. dominant and widest-spreading temperate forms invaded the equatorial lowlands. These latter. the northern temperate forms living on the lowlands under the equator. even if the climate was not perfectly fitted for them. for the change of temperature must have been very slow. where. The inhabitants of these hot lowlands would at the same time have migrated to the tropical and subtropical regions of the south. there can hardly be a doubt that some of the more vigorous. In the regular course of events the southern hemisphere would in its turn be subjected to a severe Glacial period. however. and then the southern temperate forms would invade the equatorial lowlands. They might have survived. would have been driven to their former homes or have been destroyed. and from the facts just given. would have to compete with many new forms and would be exposed to somewhat different physical conditions. leaving some few species on the mountains. As the cold became more and more intense. must have been very long. 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. and on the mountains of the tropics. would return to their former homes. with the northern hemisphere rendered warmer. as both hemispheres gradually recovered their former temperature. But the species left during a long time on these mountains. The northern forms which had before been left on the mountains would now descend and mingle with the southern forms. hence. this period will have been ample for any amount of migration. being replaced by the equatorial forms returning from the south. or in opposite hemispheres.the present apparently inexplicable distribution of various organisms in the temperate parts of both hemispheres. and plants undoubtedly possess a certain capacity for acclimatisation. for the southern hemisphere was at this period warmer. and carrying southward with them some of the northern temperate forms which had descended from their mountain fastnesses. as shown by their transmitting to their offspring different constitutional powers of resisting heat and cold. Thus. when the warmth returned. On the decline of the Glacial period. The Glacial period. and when we remember over what vast spaces some naturalised plants and animals have spread within a few centuries. if sufficiently lofty.

I suspect that this preponderant migration from the north to the south is due to the greater extent of land in the north. by those which have become naturalised. offer a partial exception. and this is the case. in accordance with the same principles. We must. at the present day. a few southern forms on the mountains of Borneo and Abyssinia. for the many quite distinct species inhabiting the same widely separated areas. Mountains are islands on the land. And thus. or dominating power. The Neilgherrie Mountains in India. for these will account. wool. strongly insisted on by Hooker in regard to America. de Candolle in regard to Australia. New Zealand. than in a reversed direction. but not so the southern in regard to the northern forms. Hooker. during the alternations of the Glacial periods. for here. and by Alph. and belonging to genera not now found in the intermediate torrid zones. and to the northern forms having existed in their own homes in greater numbers. also. It is a remarkable fact. we see that very many European productions cover the ground in La Plata. than the southern forms. In the same manner. and have beaten the natives. however. Australian forms are rapidly sowing themselves and becoming naturalised. but these have almost everywhere yielded to the more dominant forms generated in the larger areas and more efficient workshops of the north. the northern forms were the more powerful and were able to hold their places on the mountains. and to a lesser degree in Australia. and their inhabitants have yielded to those produced within the larger areas of the north. 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. whereas extremely few southern forms have become naturalised in any part of the northern hemisphere. that many more identical or slightly modified species have migrated from the north to the south. and afterwards migrate southward with the southern forms. In many islands the native productions are nearly equalled.now exist as varieties or as representative species. just in the same way as the inhabitants of real islands have everywhere 389 . Before the last great Glacial period. or even outnumbered. however. no doubt the intertropical mountains were stocked with endemic Alpine forms. bear in mind the occurrence in both hemispheres of former Glacial periods. and having consequently been advanced through natural selection and competition to a higher stage of perfection. though hides. when the two sets became commingled in the equatorial regions. We see. as I hear from Dr. and this is the first stage towards their extinction.

during the height of the Glacial period. which now live so widely separated in the north and south. The existence at these and other distant points of the southern hemisphere. New Zealand. by keeping to the cooler currents. may have been concerned in their dispersal. but icebergs. is a more remarkable case. according to Forbes. supported a highly peculiar and isolated 390 .yielded and are still yielding to continental forms naturalised through man's agency. are removed on the views above given. I am far from supposing that all the difficulties in regard to the distribution and affinities of the identical and allied species. We cannot hope to explain such facts. of the same plants at points so enormously remote as Kerguelen Land. of these a few would perhaps at once be able to migrate southwards. 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. The facts seem to indicate that distinct species belonging to the same genera have migrated in radiating lines from a common centre. until we can say why one species and not another becomes naturalised by man's agency in a foreign land. why one species ranges twice or thrice as far. Various special difficulties also remain to be solved. and sometimes on the intermediate mountain ranges. which. in nearly the same manner as. and on the intertropical mountains. The same principles apply to the distribution of terrestrial animals and of marine productions. the ocean-currents were widely different to what they now are. in the northern and southern temperate zones. now covered with ice. while others have remained unaltered. 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. and Fuegia. as suggested by Lyell. why certain species have been modified and have given rise to new forms. The exact lines of migration cannot be indicated. as in the northern hemisphere. for instance. and is twice or thrice as common. the occurrence. though distinct. when the Antarctic lands. as shown by Dr. of species. When. to a former and warmer period. isolated spaces inhabited by Arctic productions exist to the present day in the deeper parts of the northern temperate seas. and I am inclined to look in the southern. Some of these species are so distinct. before the commencement of the last Glacial period. Hooker. as another species within their own homes. We cannot say why certain species and not others have migrated. belong to genera exclusively confined to the south. some of the inhabitants of the temperate seas might have reached the equator.

which serves as a record. and in both cases have reached the equator. driven up and surviving in the mountain fastnesses of almost every land. Croll's conclusion that successive Glacial periods in the one hemisphere coincide with warmer periods in the opposite hemisphere. It may be suspected that before this flora was exterminated during the last Glacial epoch. as halting-places. The various beings thus left stranded may be compared with savage races of man. on the effects of great alternations of climate throughout the world on geographical distribution. and New Zealand may have become slightly tinted by the same peculiar forms of life. 391 . and has consequently more freely inundated the south. and by the aid. of the former inhabitants of the surrounding lowlands. rising higher on the shores where the tide rises highest. explains a multitude of facts in the distribution of the same and of the allied forms of life in all parts of the globe. Sir C. Australia. And we have now seen that Mr. Thus the southern shores of America. together with the admission of the slow modification of species. The living waters have flowed during one period from the north and during another from the south. so have the living waters left their living drift on our mountain summits. full of interest to us. As the tide leaves its drift in horizontal lines. Lyell in a striking passage has speculated. a few forms had been already widely dispersed to various points of the southern hemisphere by occasional means of transport. in language almost identical with mine.flora. of now sunken islands. in a line gently rising from the Arctic lowlands to a great latitude under the equator. but the stream of life has flowed with greater force from the north than in the opposite direction.

392 .Chapter 13 Geographical Distribution--continued 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.

Gunther has lately shown that the Galaxias attenuatus inhabits Tasmania. This case. But the wide ranging power of fresh-water productions can. be mainly attributed to changes in the level of the land within the recent period. Thus fishes still alive are not very rarely dropped at distant points by whirlwinds. Not only have many fresh-water species. the Falkland Islands and the mainland of South America. and liability to wide dispersal would follow from this capacity as an almost necessary consequence. or from stream to stream. for short and frequent migrations from pond to pond. belonging to different classes. etc. and as the sea is apparently a still more formidable barrier.. compared with those of Britain. and as if capriciously. however. Their dispersal may. in a manner highly useful to them. but allied species prevail in a remarkable manner throughout the world. in most cases be explained by their having become fitted. It is probable that they are occasionally transported by what may be called accidental means. and at the dissimilarity of the surrounding terrestrial beings. When first collecting in the fresh waters of Brazil. Instances. But Dr. for in two adjoining river systems some of the species may be the same and some wholly different. it might have been thought that fresh-water productions would not have ranged widely within the same country. shells. an enormous range. This is a wonderful case. I think. I well remember feeling much surprise at the similarity of the fresh-water insects. that they would never have extended to distant countries. some of the most difficult to explain are presented by fish. within their own countries. causing rivers to flow into each other. also. It was formerly believed that the same fresh-water species never existed on two continents distant from each other. and probably indicates dispersal from an Antarctic centre during a former warm period.1. and it is known that the ova retain their vitality for a considerable time after removal from the water. We can here consider only a few cases. however. of these. Fresh-Water Productions As lakes and river-systems are separated from each other by barriers of land. But the case is exactly the reverse. On the same continent fresh-water fish often range widely. though separated by a distance of about 230 miles. could be given of this having occurred during 393 . 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. New Zealand.

Salt-water fish can with care be slowly accustomed to live in fresh water.floods. without any change of level. and allied species which. prevail throughout the world. though aquatic in their nature. Gunther has recently been led by several considerations to infer that with fishes the same forms have a long endurance. and the ova. that I have unintentionally stocked the one with fresh-water shells from the other. The wide differences of the fish on the opposite sides of most mountain-ranges. in damp air. and must have proceeded from a single source. and. But two facts. Some fresh-water fish belong to very ancient forms. are descended from a common parent. When ducks suddenly emerge from a pond covered with duck-weed. and consequently time and means for much migration. and it has happened to me. though at a somewhat more advanced age they would voluntarily drop off. which I have observed—and many others no doubt will be discovered—throw some light on this subject. according to Valenciennes. and could. as well as the adults. Their distribution at first perplexed me much. Dr. and if blown across the sea to an oceanic island. I have twice seen these little plants adhering to their backs. Some species of fresh-water shells have very wide ranges. become adapted without much difficulty to the fresh waters of a distant land. would be sure to alight on a pool or rivulet. Sir Charles Lyell informs me that a Dyticus has been caught with an Ancylus (a fresh-water shell like a 394 . and in such cases there will have been ample time for great geographical changes. so that a marine species belonging to a fresh-water group might travel far along the shores of the sea. from an early period. or to any other distant point. which are continuous and consequently must. are immediately killed by sea-water. and in this length of time a duck or heron might fly at least six or seven hundred miles. and I found that numbers of the extremely minute and just-hatched shells crawled on the feet. I could not even understand how some naturalised species have spread rapidly throughout the same country. as their ova are not likely to be transported by birds. it is probable. These just-hatched molluscs. in removing a little duck-weed from one aquarium to another. where many ova of fresh-water shells were hatching. have completely prevented the inosculation of the river systems on the two sides. there is hardly a single group of which all the members are confined to fresh water. from twelve to twenty hours. on our theory. and clung to them so firmly that when taken out of the water they could not be jarred off. Moreover. survived on the duck's feet. leads to the same conclusion. But another agency is perhaps more effectual: I suspended the feet of a duck in an aquarium.

according to Alph. would be the most likely to have muddy feet. which frequent the muddy edges of ponds. in those large groups of terrestrial plants. they then take flight and go to other waters. de Candolle. century after century. Other and unknown agencies probably have also played a part. when rejected many hours afterwards in pellets or in the excrement. Herons and other birds. beneath water.limpet) firmly adhering to it. they would be sure to fly to their natural fresh-water haunts. once flew on board the "Beagle. I think favourable means of dispersal explain this fact. and when gaining the land. which have very few aquatic members. it has long been known what enormous ranges many fresh-water. a Colymbetes. have. The same agency may have come into play with the eggs of some of the smaller fresh-water animals. situated at very distant points. and yet the viscid mud was all contained in a breakfast cup! Considering these facts. pulling up and counting each plant as it grew. With respect to plants. 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. and are occasionally found on the most remote and barren islands of the open ocean. both over continents and to the most remote oceanic islands. this mud when dry weighed only 6 and 3/4 ounces. a wide range. the plants were of many kinds. so that any dirt on their feet would not be washed off. on the edge of a little pond. and a water-beetle of the same family. but will here give only the most striking case: I took in February three tablespoonfuls of mud from three different points. I have before mentioned that earth occasionally adheres in some quantity to the feet and beaks of birds. have gone on daily devouring fish. I do not believe that botanists are aware how charged the mud of ponds is with seeds: I have tried several little experiments. the 395 . When I saw the great size of the seeds of that fine water-lily. for the latter seem immediately to acquire. and we have seen that seeds retain their power of germination. Wading birds. I have stated that fresh-water fish eat some kinds of seeds. as of the yellow water-lily and Potamogeton. Birds of this order wander more than those of any other. I kept it covered up in my study for six months." 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. though they reject many other kinds after having swallowed them. they would not be likely to alight on the surface of the sea. or are blown across the sea. even small fish swallow seeds of moderate size. This is strikingly illustrated. and were altogether 537 in number. if suddenly flushed. as if in consequence. and even marsh-species.

apparently depends in main part on the wide dispersal of their seeds and eggs by animals. and. than in the case of terrestrial colonists. the competition between them will probably be less severe than between terrestrial species. In considering these several means of distribution. or in some degree modified. Hooker. and of the lower animals. Now this bird must often have flown with its stomach thus well stocked to distant ponds. 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. more especially by fresh-water birds. then getting a hearty meal of fish. consequently an intruder from the waters of a foreign country would have a better chance of seizing on a new place. de Candolle's remarks on the distribution of this plant. which have great powers of flight. and remembered Alph. and a single seed or egg will have a good chance of succeeding. it should be remembered that when a pond or stream is first formed. the Nelumbium luteum) in a heron's stomach. but Audubon states that he found the seeds of the great southern water-lily (probably according to Dr. however few in kind. I thought that the means of its dispersal must remain inexplicable. Although there will always be a struggle for life between the inhabitants of the same pond. 396 . whether retaining the same identical form. it will be unoccupied. and then having become extinct at intermediate points. But the wide distribution of fresh-water plants. and this will give time for the migration of aquatic species. analogy makes me believe that it would have rejected the seeds in the pellet in a fit state for germination. for instance on a rising islet. We should also remember that many fresh-water productions are low in the scale of nature. We should not forget the probability of many fresh-water forms having formerly ranged continuously over immense areas. and we have reason to believe that such beings become modified more slowly than the high. and naturally travel from one piece of water to another.Nelumbium.

for instance. 397 . on the view that not only all the individuals of the same species have migrated from some one area. but it does not accord with all the facts in regard to the productions of islands. with its lofty mountains and diversified stations. the birthplace of their early progenitors. has given rise to so great a difference in number. In St. On The Inhabitants Of Oceanic Islands We now come to the last of the three classes of facts. we must admit that some cause.2. together with the outlying islands of Auckland. as they have in New Zealand and on every other oceanic island which can be named. This view removes many difficulties. although now inhabiting the most distant points. de Candolle admits this for plants. Even the uniform county of Cambridge has 847 plants. but a few ferns and a few introduced plants are included in these numbers. but that allied species. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions. Campbell and Chatham. will have to admit that a sufficient number of the best adapted plants and animals were not created for oceanic islands. yet many species have now become naturalised on it. We have evidence that the barren island of Ascension aboriginally possessed less than half-a-dozen flowering plants. for man has unintentionally stocked them far more fully and perfectly than did nature. New Zealand. and Wollaston for insects. He who admits the doctrine of the creation of each separate species. have proceeded from a single area. independently of different physical conditions. extending over 780 miles of latitude. but shall consider some other cases bearing on the truth of the two theories of independent creation and of descent with modification. 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. and the comparison in some other respects is not quite fair. if we compare this moderate number with the species which swarm over equal areas in Southwestern Australia or at the Cape of Good Hope. contain altogether only 960 kinds of flowering plants. and the little island of Anglesea 764. which I have selected as presenting the greatest amount of difficulty with respect to distribution. The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. In the following remarks I shall not confine myself to the mere question of dispersal.

I will give a few illustrations of the foregoing remarks: in the Galapagos Islands there are twenty-six land birds. for. on the other hand. Jones's admirable account of Bermuda. does not possess a single endemic land bird. we shall see that this is true. and then compare the area of the island with that of the continent. Almost every year. that very many North American birds occasionally or even frequently visit this island. species occasionally arriving. J.Although in oceanic islands the species are few in number. If we compare.e. which for long ages have there struggled together. those found nowhere else in the world) is often extremely large. with which the insular forms have intercrossed. Hence. with the number found on any continent. because in an island nearly all the species of one class are peculiar. are peculiar. and have become mutually co-adapted. and which has a very peculiar soil. Any tendency to 398 . which lies at about the same distance from North America as the Galapagos Islands do from South America. of which one alone is peculiar. or of endemic birds in the Galapagos Archipelago. this island is inhabited by ninetynine kinds. as I am informed by Mr. or of another section of the same class. after long intervals of time in the new and isolated district. and partly on the frequent arrival of unmodified immigrants from the mother-country. E.V. and it is obvious that marine birds could arrive at these islands much more easily and frequently than land-birds.M. and will consequently have been but little liable to modification. so that their mutual relations have not been much disturbed. for instance. of these twenty-one (or perhaps twentythree) are peculiar. whereas of the eleven marine birds only two are peculiar. and having to compete with new associates. would be eminently liable to modification. those of another class. It should be borne in mind that the offspring of such crosses would certainly gain in vigour. each kind will have been kept by the others to its proper place and habits. and three or four other species are confined to this island and to the Canaries. though very closely related to a European form. Harcourt. so that even an occasional cross would produce more effect than might have been anticipated. when settled in their new homes. But it by no means follows that. and this difference seems to depend partly on the species which are not modified having immigrated in a body. and we know from Mr. Bermuda. and would often produce groups of modified descendants. as already explained. the number of endemic land-shells in Madeira. This fact might have been theoretically expected. the proportion of endemic kinds (i. many European and African birds are blown to Madeira. So that the islands of Bermuda and Madeira have been stocked from the neighbouring continents with birds.

from its geological character and the direction of its mountain ranges. Dr. and is not separated from Australia by a profoundly deep sea. and their places are occupied by other classes. should be considered as appurtenances of Australia. still retaining its hooks. For instance. might be transported across three or four hundred miles of open sea far more easily than land-shells. thus in the Galapagos Islands reptiles. it is in some degree doubtful whether it should be so ranked. though we do not know how sea-shells are dispersed. perhaps attached to seaweed or floating timber. Facility of immigration seems to have been fully as important as the nature of the conditions. or to the feet of wading birds. the Rev. W. whatever the cause may be. Many remarkable little facts could be given with respect to the inhabitants of oceanic islands.modification will also have been checked by intercrossing with the unmodified immigrants. Again. often arriving from the mother-country. it is of large size. which had no chance of successfully competing with the many 399 . yet we can see that their eggs or larvae. All such differences in number. as Alph. yet few relations are more manifest than that hooks serve for the transportal of seeds in the wool or fur of quadrupeds. the place of mammals. take. now trees. Madeira again is inhabited by a wonderful number of peculiar land-shells. The different orders of insects inhabiting Madeira present nearly parallel cases. in certain islands not tenanted by a single mammal.B. Clarke has lately maintained that this island. Turning to plants. islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species. or recently took. and in New Zealand gigantic wingless birds. Hence trees would be little likely to reach distant oceanic islands. and the plant then becoming modified would form an endemic species. generally have. but this explanation is not a little doubtful. de Candolle has shown. and the absence of certain whole groups of animals and plants. like the shrivelled wings under the soldered wing-covers of many insular beetles. But a hooked seed might be carried to an island by other means. which would form a useless appendage. are generally accounted for by supposed differences in the physical conditions of the islands. whereas not one species of sea-shell is peculiar to its shores: now. Oceanic islands are sometimes deficient in animals of certain whole classes. some of the endemic plants have beautifully hooked seeds. as well as New Caledonia. Although New Zealand is here spoken of as an oceanic island. confined ranges. and an herbaceous plant. Hooker has shown that in the Galapagos Islands the proportional numbers of the different orders are very different from what they are elsewhere.

fully developed trees growing on a continent. 400 . might. when established on an island. gain an advantage over other herbaceous plants by growing taller and taller and overtopping them. natural selection would tend to add to the stature of the plant. and thus first convert it into a bush and then into a tree. In this case. to whatever order it belonged.

and many islands situated at a much less distance are equally barren. The Falkland Islands. icebergs formerly brought boulders to its western shores. and hardly an island can be named on which our smaller quadrupeds have not become naturalised and greatly multiplied. with the exception of New Zealand. It cannot be said. many volcanic islands are sufficiently 401 . the Azores. come nearest to an exception. which are inhabited by a wolf-like fox. I have carefully searched the oldest voyages. New Caledonia. But why. on the ordinary view of creation. indeed it seems that islands are peculiarly fitted for these animals. This general absence of frogs. for frogs have been introduced into Madeira. newts) are never found on any of the many islands with which the great oceans are studded. and have found it true. moreover. it would be very difficult to explain. toads and newts on so many true oceanic islands cannot be accounted for by their physical conditions. 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. the Andaman Islands. free from doubt. But as these animals and their spawn are immediately killed (with the exception. and therefore we can see why they do not exist on strictly oceanic islands. as it lies on a bank in connection with the mainland at a distance of about 280 miles. they should not have been created there. but this group cannot be considered as oceanic. and this is still more doubtful with respect to the Andaman and Solomon groups and the Seychelles. as far as known. on the theory of creation. toads. and perhaps the Solomon Islands and the Seychelles. as now frequently happens in the arctic regions. But I have already remarked that it is doubtful whether New Zealand and New Caledonia ought to be classed as oceanic islands. when lying close to a continent. Yet it cannot be said that small islands will not support at least small mammals. of one Indian species) by sea-water. Absence Of Batrachians And Terrestrial Mammals On Oceanic Islands With respect to the absence of whole orders of animals on oceanic islands. there would be great difficulty in their transportal across the sea. that there has not been time for the creation of mammals. and have multiplied so as to become a nuisance. I have taken pains to verify this assertion.3. and have not found a single instance. and Mauritius. Vincent long ago remarked that Batrachians (frogs. and they may have formerly transported foxes. for they occur in many parts of the world on very small islands. Mammals offer another and similar case. Bory St.

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

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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,

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after having been thus treated, and again hybernating, was put into seawater 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.

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4. 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 twentyone, 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

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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

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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 bestfitted 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

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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

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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

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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.

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5. 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

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to those of the nearest continent. On these same principles we can understand. there would be an almost endless amount of organic action and reaction. should be closely related to each other. and another group. some existing in scanty numbers—and this we do find in the several great geographical provinces of the world. and how it is that under different latitudes. and the depth of the sea between such islands and the mainland. in a more or less modified condition. as batrachians and terrestrial mammals. the inhabitants of the plains and mountains. having nearly the same physical conditions. but less closely. are linked together in so mysterious a manner. we can see why two areas. should be absent from oceanic islands. if there exist very closely allied or 412 . there would ensue in the to or more regions. for according to the length of time which has elapsed since the colonists entered one of the regions. whilst the most isolated islands should possess their own peculiar species of aerial mammals or bats. in South America.botanical provinces. should have all its species the same with those in an adjoining quarter of the world. We can see why whole groups of organisms. even within the same class. according or not as those which entered happened to come into more or less direct competition with each other and with the aborigines. and deserts. why oceanic islands should have few inhabitants. and are likewise linked to the extinct beings which formerly inhabited the same continent. We can see why. of the forests. as I have endeavoured to show. We can thus understand the concentration of related species within the same areas. We can clearly see why all the inhabitants of an archipelago. in relation to the means of migration. in islands. independently of their physical conditions. and should likewise be related. and why. or other source whence immigrants might have been derived. and we should find some groups of beings greatly. We can see why. and some only slightly modified. and according as the immigrants were capable of varying more or less rapidly. or both. should often be inhabited by very different forms of life. infinitely diversified conditions of life. there should be some relation between the presence of mammals. marshes. a large proportion should be endemic or peculiar. either in greater or lesser numbers. one group of beings should have all its species peculiar. Bearing in mind that the mutual relation of organism to organism is of the highest importance. though specifically distinct on the several islets. but that of these. according to the nature of the communication which allowed certain forms and not others to enter. some developed in great force. for instance.

it certainly is the general rule that the area inhabited by a single species. some identical species will almost always there be found. but there are in both cases marked exceptions to the rule. or by the species having become extinct in the intermediate tracts. 413 . or by a group of species. and modifications have been accumulated by the same means of natural selection. may. As the late Edward Forbes often insisted. but which occur above and below: so in space. living during the same period of time. differ greatly from each other. and the exceptions. as in looking to distant provinces throughout the world. 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. or only in a different section of the same order. be accounted for by former migrations under different circumstances. we find that species in certain classes differ little from each other. In both time and space the lowly organised members of each class generally change less than the highly organised. or through occasional means of transport. in both cases the laws of variation have been the same. there is a striking parallelism in the laws of life throughout time and space. which are not rare. as I have attempted to show. 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. In looking to the long succession of past ages. According to our theory. The endurance of each species and group of species is continuous in time. these several relations throughout time and space are intelligible. Groups of species. Both in time and space species and groups of species have their points of maximum development. or to those which have changed after having migrated into distant quarters. in both cases they are connected by the same bond of ordinary generation. whilst those in another class. is continuous. however distant from each other. or living within the same area. are often characterised by trifling features in common. for whether we look to the allied forms of life which have changed during successive ages.representative species in two areas. as of sculpture or colour. We see this in many facts.

laws of. their origin explained — Summary.Chapter 14 Mutual Affinities Of Organic Beings: Morphology -Embryology -. general. explained by variations not supervening at an early age. explained on the theory of descent with modification — Classification of varieties — Descent always used in classification — Analogical or adaptive characters — Affinities. between parts of the same individual — Embryology. 414 . groups subordinate to groups — Natural system — Rules and difficulties in classification. complex and radiating — Extinction separates and defines groups — Morphology.Rudimentary Organs Classification. between members of the same class. and being inherited at a corresponding age — Rudimentary organs.

In the diagram each letter on the uppermost line may represent a genus including several species. In the second and fourth chapters. ultimately become converted into new and distinct species. and he will see that the inevitable result is. This latter conclusion is supported by observing the great diversity of forms. which. I have attempted to show that within each country it is the widely ranging. if one group had been exclusively fitted to inhabit the land. which diverged from a common parent at the fifth stage of descent. The varieties. and these. and another the water. as formerly explained. in any small area. so that they can be classed in groups under groups. which vary most. one to feed on flesh. and by certain facts in naturalisation. consequently. I further attempted to show that from the varying descendants of each species trying to occupy as many and as different places as possible in the economy of nature. The existence of groups would have been of simple significance. and which generally include many dominant species.1. belonging to the larger genera in each class. on Variation and on Natural Selection. for all are descended from one ancient parent. These five genera have also much in common. thus produced. Consequently the groups which are now large. on the principle of inheritance. much in common. the much diffused and common. another on vegetable matter. they constantly tend to diverge in character. on this same principle. to supplant and exterminate the preceding. have inherited something in common. though 415 . This classification is not arbitrary like the grouping of the stars in constellations. or incipient species. distinct from that containing the next two genera on the right hand. but the case is widely different. and form a subfamily. tend to produce other new and dominant species. tend to go on increasing in size. But the three genera on the left hand have. that is the dominant species. come into the closest competition. and the whole of the genera along this upper line form together one class. I attempted also to show that there is a steady tendency in the forms which are increasing in number and diverging in character. for it is notorious how commonly members of even the same subgroup have different habits. that the modified descendants proceeding from one progenitor become broken up into groups subordinate to groups. I request the reader to turn to the diagram illustrating the action. and. less divergent and less improved forms. Classification From the most remote period in the history of the world organic beings have been found to resemble each other in descending degrees. and so on. of these several principles.

all under one great class. namely. Naturalists. but unless it be specified whether order in time or space. and no cause can at present be assigned for their falling into groups. for instance. And all these genera. which. that minerals and the elemental substances can be thus arranged. The grand fact of the natural subordination of organic beings in groups under groups.less than when grouped in subfamilies. as we have seen. by one sentence to give the characters common. But what is meant by this system? Some authors look at it merely as a scheme for arranging together those living objects which are most alike. seem to imply that some deeper bond is included in our classifications than mere resemblance. and the view above given accords with their natural arrangement in group under group. and that community of descent—the one known cause of close similarity in organic beings—is the bond. genera and families in each class. In this case there is of course no relation to genealogical succession. a full description is given of each kind of dog. by another those common to the doggenus. or what else is meant by the plan of the Creator. Expressions such as that famous one by Linnaeus. they believe that it reveals the plan of the Creator. can be classed in many ways. try to arrange the species. that the characters do not make the genus. or both. descended from (A). So that we here have many species descended from a single progenitor grouped into genera. by adding a single sentence. or more naturally by a number of characters. either artificially by single characters. No doubt organic beings. but that the genus gives the characters. and they form a family distinct from that containing the three genera still further to the right hand. and for separating those which are most unlike. which diverged at an earlier period. is partially revealed to us by our classifications. for instance. like all other objects. and the genera into subfamilies. The ingenuity and utility of this system are indisputable. does not always sufficiently strike us. and then. to all mammals. on what is called the Natural System. form an order distinct from the genera descended from (I). it seems to me that nothing is thus added to our knowledge. or as an artificial method of enunciating. But with organic beings the case is different. which we often meet with in a more or less concealed form. from its familiarity. though observed by various degrees of modification. as briefly as possible. and no other explanation has ever been attempted. general propositions—that is. We know. I believe that this is the case. 416 . by another those common to all carnivora. is in my judgment thus explained. But many naturalists think that something more is meant by the Natural System. families and orders. which.

in another work he says. we have seen that they are often of the highest service in classification. This depends on their constancy throughout many allied groups. as of any importance. has nearly the same physiological value. on which their nutrition and life depend. Robert Brown. Nothing can be more false. It may even be given as a general rule. not only in this. and the general place of each being in the economy of nature. "The generative organs." Again. is almost proved by the fact. No one regards the external similarity of a mouse to a shrew. "like that of all their parts. That the mere physiological importance of an organ does not determine its classificatory value. and it has been fully acknowledged in the writings of almost every author. but. its classificatory value is widely different. It might have been thought (and was in ancient times thought) that those parts of the structure which determined the habits of life. who. in formerly discussing certain morphological characters which are not functionally important. We are least likely in the modifications of these organs to mistake a merely adaptive for an essential character.Let us now consider the rules followed in classification. or is simply a scheme for enunciating general propositions and of placing together the forms most like each other. the 417 . the more important it becomes for classification. being those which are most remotely related to the habits and food of an animal. of a dugong to a whale." With plants how remarkable it is that the organs of vegetation. These resemblances. are ranked as merely "adaptive or analogical characters." but to the consideration of these resemblances we shall recur. and in some cases seems to be entirely lost. which acts only on serviceable characters. and the difficulties which are encountered on the view that classification either gives some unknown plan of creation. No naturalist can have worked at any group without being struck with this fact. that in allied groups. as we have every reason to suppose. As an instance: Owen. that the less any part of the organisation is concerned with special habits. says their generic importance. It will suffice to quote the highest authority. I have always regarded as affording very clear indications of its true affinities. are of little signification. would be of very high importance in classification. says. is very unequal. whereas the organs of reproduction. in speaking of the dugong. in speaking of certain organs in the Proteaceae. and their constancy chiefly depends on any slight deviations not having been preserved and accumulated by natural selection. as I apprehend in every natural family. though so intimately connected with the whole life of the being. of a whale to a fish. are of paramount importance! So again. in which the same organ. with their product the seed and embryo.

as hair or feathers. the antennae." To give an example among insects: in one great division of the Hymenoptera. as has often been remarked. also. and certain rudimentary bones of the leg. and of almost universal prevalence. in another division they differ much. both of high physiological importance. are most constant in structure. according to Owen. are highly serviceable in exhibiting the close affinity between Ruminants and Pachyderms. this external and trifling character would have been considered by naturalists as an important aid in determining the degree of affinity of this strange creature to birds. though here even. for classification. and the differences are of quite subordinate value in classification. no one will say that rudimentary or atrophied organs are of high physiological or vital importance. in the imbricate or valvular aestivation. The value indeed of an aggregate of characters is very evident in natural history. Hence. they appear insufficient to separate Cnestis from Connarus.genera of the Connaraceae &q