On the Origin of Species, 6th Edition

Darwin, Charles

Published: 1872 Type(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".


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


in the organic, as well as in the inorganic world, being the result of law, and not of miraculous interposition. Lamarck seems to have been chiefly led to his conclusion on the gradual change of species, by the difficulty of distinguishing species and varieties, by the almost perfect gradation of forms in certain groups, and by the analogy of domestic productions. With respect to the means of modification, he attributed something to the direct action of the physical conditions of life, something to the crossing of already existing forms, and much to use and disuse, that is, to the effects of habit. To this latter agency he seems to attribute all the beautiful adaptations in nature; such as the long neck of the giraffe for browsing on the branches of trees. But he likewise believed in a law of progressive development, and as all the forms of life thus tend to progress, in order to account for the existence at the present day of simple productions, he maintains that such forms are now spontaneously generated. (I have taken the date of the first publication of Lamarck from Isidore Geoffroy Saint- Hilaire's ("Hist. Nat. Generale", tom. ii. page 405, 1859) excellent history of opinion on this subject. In this work a full account is given of Buffon's conclusions on the same subject. It is curious how largely my grandfather, Dr. Erasmus Darwin, anticipated the views and erroneous grounds of opinion of Lamarck in his "Zoonomia" (vol. i. pages 500-510), published in 1794. According to Isid. Geoffroy there is no doubt that Goethe was an extreme partisan of similar views, as shown in the introduction to a work written in 1794 and 1795, but not published till long afterward; he has pointedly remarked ("Goethe als Naturforscher", von Dr. Karl Meding, s. 34) that the future question for naturalists will be how, for instance, cattle got their horns and not for what they are used. It is rather a singular instance of the manner in which similar views arise at about the same time, that Goethe in Germany, Dr. Darwin in England, and Geoffroy Saint-Hilaire (as we shall immediately see) in France, came to the same conclusion on the origin of species, in the years 1794-5.) Geoffroy Saint-Hilaire, as is stated in his "Life", written by his son, suspected, as early as 1795, that what we call species are various degenerations of the same type. It was not until 1828 that he published his conviction that the same forms have not been perpetuated since the origin of all things. Geoffroy seems to have relied chiefly on the conditions of life, or the "monde ambiant" as the cause of change. He was cautious in drawing conclusions, and did not believe that existing species are now undergoing modification; and, as his son adds, "C'est donc un probleme a reserver entierement a l'avenir, suppose meme que l'avenir doive avoir prise sur lui."


In 1813 Dr. W.C. Wells read before the Royal Society "An Account of a White Female, part of whose skin resembles that of a Negro"; but his paper was not published until his famous "Two Essays upon Dew and Single Vision" appeared in 1818. In this paper he distinctly recognises the principle of natural selection, and this is the first recognition which has been indicated; but he applies it only to the races of man, and to certain characters alone. After remarking that negroes and mulattoes enjoy an immunity from certain tropical diseases, he observes, firstly, that all animals tend to vary in some degree, and, secondly, that agriculturists improve their domesticated animals by selection; and then, he adds, but what is done in this latter case "by art, seems to be done with equal efficacy, though more slowly, by nature, in the formation of varieties of mankind, fitted for the country which they inhabit. Of the accidental varieties of man, which would occur among the first few and scattered inhabitants of the middle regions of Africa, some one would be better fitted than others to bear the diseases of the country. This race would consequently multiply, while the others would decrease; not only from their in ability to sustain the attacks of disease, but from their incapacity of contending with their more vigorous neighbours. The colour of this vigorous race I take for granted, from what has been already said, would be dark. But the same disposition to form varieties still existing, a darker and a darker race would in the course of time occur: and as the darkest would be the best fitted for the climate, this would at length become the most prevalent, if not the only race, in the particular country in which it had originated." He then extends these same views to the white inhabitants of colder climates. I am indebted to Mr. Rowley, of the United States, for having called my attention, through Mr. Brace, to the above passage of Dr. Wells' work. The Hon. and Rev. W. Herbert, afterward Dean of Manchester, in the fourth volume of the "Horticultural Transactions", 1822, and in his work on the "Amaryllidaceae" (1837, pages 19, 339), declares that "horticultural experiments have established, beyond the possibility of refutation, that botanical species are only a higher and more permanent class of varieties." He extends the same view to animals. The dean believes that single species of each genus were created in an originally highly plastic condition, and that these have produced, chiefly by inter-crossing, but likewise by variation, all our existing species. In 1826 Professor Grant, in the concluding paragraph in his wellknown paper ("Edinburgh Philosophical Journal", vol. XIV, page 283) on the Spongilla, clearly declares his belief that species are descended from


other species, and that they become improved in the course of modification. This same view was given in his Fifty-fifth Lecture, published in the "Lancet" in 1834. In 1831 Mr. Patrick Matthew published his work on "Naval Timber and Arboriculture", in which he gives precisely the same view on the origin of species as that (presently to be alluded to) propounded by Mr. Wallace and myself in the "Linnean Journal", and as that enlarged in the present volume. Unfortunately the view was given by Mr. Matthew very briefly in scattered passages in an appendix to a work on a different subject, so that it remained unnoticed until Mr. Matthew himself drew attention to it in the "Gardeners' Chronicle", on April 7, 1860. The differences of Mr. Matthew's views from mine are not of much importance: he seems to consider that the world was nearly depopulated at successive periods, and then restocked; and he gives as an alternative, that new forms may be generated "without the presence of any mold or germ of former aggregates." I am not sure that I understand some passages; but it seems that he attributes much influence to the direct action of the conditions of life. He clearly saw, however, the full force of the principle of natural selection. The celebrated geologist and naturalist, Von Buch, in his excellent "Description Physique des Isles Canaries" (1836, page 147), clearly expresses his belief that varieties slowly become changed into permanent species, which are no longer capable of intercrossing. Rafinesque, in his "New Flora of North America", published in 1836, wrote (page 6) as follows: "All species might have been varieties once, and many varieties are gradually becoming species by assuming constant and peculiar characters;" but further on (page 18) he adds, "except the original types or ancestors of the genus." In 1843-44 Professor Haldeman ("Boston Journal of Nat. Hist. U. States", vol. iv, page 468) has ably given the arguments for and against the hypothesis of the development and modification of species: he seems to lean toward the side of change. The "Vestiges of Creation" appeared in 1844. In the tenth and much improved edition (1853) the anonymous author says (page 155): "The proposition determined on after much consideration is, that the several series of animated beings, from the simplest and oldest up to the highest and most recent, are, under the providence of God, the results, FIRST, of an impulse which has been imparted to the forms of life, advancing them, in definite times, by generation, through grades of organisation


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


"enumerated by the zoologist as evidence of distinct creation of the bird in and for such islands, he chiefly expresses that he knows not how the Red Grouse came to be there, and there exclusively; signifying also, by this mode of expressing such ignorance, his belief that both the bird and the islands owed their origin to a great first Creative Cause." If we interpret these sentences given in the same address, one by the other, it appears that this eminent philosopher felt in 1858 his confidence shaken that the Apteryx and the Red Grouse first appeared in their respective homes "he knew not how," or by some process "he knew not what." This address was delivered after the papers by Mr. Wallace and myself on the Origin of Species, presently to be referred to, had been read before the Linnean Society. When the first edition of this work was published, I was so completely deceived, as were many others, by such expressions as "the continuous operation of creative power," that I included Professor Owen with other palaeontologists as being firmly convinced of the immutability of species; but it appears ("Anat. of Vertebrates", vol. iii, page 796) that this was on my part a preposterous error. In the last edition of this work I inferred, and the inference still seems to me perfectly just, from a passage beginning with the words "no doubt the type- form," etc.(Ibid., vol. i, page xxxv), that Professor Owen admitted that natural selection may have done something in the formation of a new species; but this it appears (Ibid., vol. iii. page 798) is inaccurate and without evidence. I also gave some extracts from a correspondence between Professor Owen and the editor of the "London Review", from which it appeared manifest to the editor as well as to myself, that Professor Owen claimed to have promulgated the theory of natural selection before I had done so; and I expressed my surprise and satisfaction at this announcement; but as far as it is possible to understand certain recently published passages (Ibid., vol. iii. page 798) I have either partially or wholly again fallen into error. It is consolatory to me that others find Professor Owen's controversial writings as difficult to understand and to reconcile with each other, as I do. As far as the mere enunciation of the principle of natural selection is concerned, it is quite immaterial whether or not Professor Owen preceded me, for both of us, as shown in this historical sketch, were long ago preceded by Dr. Wells and Mr. Matthews. M. Isidore Geoffroy Saint-Hilaire, in his lectures delivered in 1850 (of which a Resume appeared in the "Revue et Mag. de Zoolog.", Jan., 1851), briefly gives his reason for believing that specific characters "sont fixes, pour chaque espece, tant qu'elle se perpetue au milieu des memes circonstances: ils se modifient, si les circonstances ambiantes viennent a


changer. En resume, L'OBSERVATION des animaux sauvages demontre deja la variabilite LIMITEE des especes. Les EXPERIENCES sur les animaux sauvages devenus domestiques, et sur les animaux domestiques redevenus sauvages, la demontrent plus clairment encore. Ces memes experiences prouvent, de plus, que les differences produites peuvent etre de VALEUR GENERIQUE." In his "Hist. Nat. Generale" (tom. ii, page 430, 1859) he amplifies analogous conclusions. From a circular lately issued it appears that Dr. Freke, in 1851 ("Dublin Medical Press", page 322), propounded the doctrine that all organic beings have descended from one primordial form. His grounds of belief and treatment of the subject are wholly different from mine; but as Dr. Freke has now (1861) published his Essay on the "Origin of Species by means of Organic Affinity", the difficult attempt to give any idea of his views would be superfluous on my part. Mr. Herbert Spencer, in an Essay (originally published in the "Leader", March, 1852, and republished in his "Essays", in 1858), has contrasted the theories of the Creation and the Development of organic beings with remarkable skill and force. He argues from the analogy of domestic productions, from the changes which the embryos of many species undergo, from the difficulty of distinguishing species and varieties, and from the principle of general gradation, that species have been modified; and he attributes the modification to the change of circumstances. The author (1855) has also treated Psychology on the principle of the necessary acquirement of each mental power and capacity by gradation. In 1852 M. Naudin, a distinguished botanist, expressly stated, in an admirable paper on the Origin of Species ("Revue Horticole", page 102; since partly republished in the "Nouvelles Archives du Museum", tom. i, page 171), his belief that species are formed in an analogous manner as varieties are under cultivation; and the latter process he attributes to man's power of selection. But he does not show how selection acts under nature. He believes, like Dean Herbert, that species, when nascent, were more plastic than at present. He lays weight on what he calls the principle of finality, "puissance mysterieuse, indeterminee; fatalite pour les uns; pour les autres volonte providentielle, dont l'action incessante sur les etres vivantes determine, a toutes les epoques de l'existence du monde, la forme, le volume, et la duree de chacun d'eux, en raison de sa destinee dans l'ordre de choses dont il fait partie. C'est cette puissance qui harmonise chaque membre a l'ensemble, en l'appropriant a la fonction qu'il doit remplir dans l'organisme general de la nature, fonction qui est pour lui sa raison d'etre." (From references in Bronn's


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

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

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

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

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

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

17 .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.

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

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

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

the organism in determining each particular form of variation. 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. 21 .

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

so rich in species. individuals of the same species. who cultivated the best pears which they could procure. I think. Man can hardly select. never thought what splendid fruit we should eat. at least during certain seasons. No man would ever try to make a fantail till he saw a pigeon with a tail developed in some slight degree in an unusual manner." as will hereafter be more fully explained. In regard to the domestic animals kept by uncivilised man. the wild parentstocks of the plants which have been longest cultivated in our flower and kitchen gardens. that in a number of cases we cannot recognise. would often succeed better in the one country than in the other. and relatively so slight in internal parts or organs.onwards. or only with much difficulty. any deviation of structure excepting such as is externally visible. 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. the Cape of Good Hope. has afforded us a single plant worth culture. and indeed he rarely cares for what is internal. it should not be overlooked that they almost always have to struggle for their own food. We can. excepting on variations which are first given to him in some slight degree by nature. perhaps. the well-known fact. as I believe. have more of the character of true species than the varieties kept in civilised countries. explains. On the view here given of the important part which selection by man has played. And in two countries very differently circumstanced. This. it becomes at once obvious. and likewise their differences being so great in external characters. though we owe our excellent fruit in some small degree to their having naturally chosen and preserved the best varieties they could anywhere find. partly explains why the varieties kept by savages. we can understand how it is that neither Australia. But the gardeners of the classical period. and thus by a process of "natural selection. It is not that these countries. do not by a strange chance possess the aboriginal stocks of any useful plants. how it is that our domestic races show adaptation in their structure or in their habits to man's wants or fancies. further understand the frequently abnormal character of our domestic races. A large amount of change. or a 43 . having slightly different constitutions or structure. two sub-breeds might be formed. thus slowly and unconsciously accumulated. as has been remarked by some authors. nor any other region inhabited by quite uncivilised man. 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. He can never act by selection. and therefore do not know.

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

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

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

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

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

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

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

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

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

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

should produce from the same seed.capsule three distinct hermaphrodite forms. 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. 54 .

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

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

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

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

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

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

is also applied arbitrarily. 61 .forms. The term variety. again. for convenience sake. in comparison with mere individual differences.

soon convinced me that there were many difficulties. I shall reserve for a future work the discussion of these difficulties. in order to become in any degree permanent. and are the most numerous in individuals—which oftenest produce well-marked varieties. for. and the species which are most widely diffused within their own country (and this is a different consideration from wide range. as did subsequently Dr. Watson. even in stronger terms. hereafter to be discussed. it should be understood that reference is made only to the forms which come into competition with each 62 . treated as it necessarily here is with much brevity. Dr. And this. and the tables of the proportional numbers of the varying species. and this might have been expected. incipient species. Hence it is the most flourishing.C. still inherit those advantages that enabled their parents to become dominant over their compatriots. as we shall hereafter see. And Common Species Vary Most Guided by theoretical considerations. or. are the most diffused in their own country. and as they come into competition (which. But my tables further show that. as they are exposed to diverse physical conditions. he thinks that the following statements are fairly well established.4. I thought that some interesting results might be obtained in regard to the nature and relations of the species which vary most. however. but Mr. Hooker permits me to add that after having carefully read my manuscript. perhaps. in any limited country. or. which. as I consider them. that is abound most in individuals. Wide-Ranging. to whom I am much indebted for valuable advice and assistance on this subject. as varieties. and examined the tables. the species which are already dominant will be the most likely to yield offspring. The whole subject. the dominant species—those which range widely. and to a certain extent from commonness)." "divergence of character. and allusions cannot be avoided to the "struggle for existence. is rather perplexing. oftenest give rise to varieties sufficiently well-marked to have been recorded in botanical works. At first this seemed a simple task. H." and other questions. the species which are the most common. Hooker. is a far more important circumstance) with different sets of organic beings. though in some slight degree modified. necessarily have to struggle with the other inhabitants of the country. Much Diffused. Alphonse de Candolle and others have shown that plants which have very wide ranges generally present varieties. might have been anticipated. In these remarks on predominence. by tabulating all the varieties in several well-worked floras. as they may be called.

With respect to the number of individuals or commonness of species. 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. and more especially to the members of the same genus or class having nearly similar habits of life. it will then be dominant within its own class. and more widely diffused. 63 .other. which live under nearly the same conditions. But if the conferva or parasitic fungus exceeds its allies in the above respects. 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. the comparison of course relates only to the members of the same group.

than the species of the smaller genera. but this seems to be connected with the nature of the stations inhabited by them. consequently. into two nearly equal masses. I was led to anticipate that the species of the larger genera in each country would oftener present varieties. Fresh water and salt-loving plants generally have very wide ranges and are much diffused. and 64 . This might have been anticipated. 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.e. many varieties or incipient species ought.. that I am surprised that my tables show even a small majority on the side of the larger genera. a larger proportional number of dominant species. plants low in the scale of organisation are generally much more widely diffused than plants higher in the scale. On the other hand.5. for the mere fact of many species of the same genus inhabiting any country. we expect to find saplings. circumstances have been favourable for variation. all those in the larger genera (i. The cause of lowly-organised plants ranging widely will be discussed in our chapter on Geographical Distribution. the species of the larger genera on one side. if we look at each species as a special act of creation. From looking at species as only strongly marked and well-defined varieties. and all those in the smaller genera on the other side. and hence we might expect that the circumstances would generally still be favourable to variation. and has little or no relation to the size of the genera to which the species belong. those including many species) being placed on one side. for wherever many closely related species (i. But so many causes tend to obscure this result. and here again there is no close relation to the size of the genera. Where many species of a genus have been formed through variation. we might have expected to have found in the larger genera. and. To test the truth of this anticipation I have arranged the plants of twelve countries.e.. than in one having few. or those including many species. the former will be found to include a somewhat larger number of the very common and much diffused or dominant species. Where many large trees grow. there is no apparent reason why more varieties should occur in a group having many species. shows that there is something in the organic or inorganic conditions of that country favourable to the genus. Again. as a general rule. species of the same genus) have been formed. and the coleopterous insects of two districts. to be now forming. I will here allude to only two causes of obscurity. be divided into two equal masses.

invariably present a larger average number of varieties than do the species of the small genera. it would have been fatal to my theory. are altogether excluded from the tables. the species of that genus present a number of varieties.those of the smaller genera on the other side. more especially as we have every reason to believe the process of manufacturing new species to be a slow one. And this certainly holds true if varieties be looked at as incipient species. on an average many are still forming. for wherever many species of the same genus have been formed. It is not that all large genera are now varying much. declined. beyond the average. that is. and are thus increasing in the number of their species. than on the side of the smaller genera. that where many species of a genus have been formed. and it has invariably proved to be the case that a larger proportion of the species on the side of the larger genera presented varieties. for if this had been so. Moreover. or where. for my tables clearly show. the manufactory of species has been active. and disappeared. Both these results follow when another division is made. wherever many species of a genus have been formed. we ought generally to find the manufactory still in action. of incipient species. the species of the large genera which present any varieties. as a general rule. that. All that we want to show is. These facts are of plain signification on the view that species are only strongly marked and permanent varieties. and when all the least genera. and this certainly holds good. or that no small genera are now varying and increasing. and that large genera have often come to their maxima. with from only one to four species. 65 . inasmuch as geology plainly tells us that small genera have in the lapse of time often increased greatly in size. if we may use the expression.

and Westwood in regard to insects. that in large genera the amount of difference between the species is often exceedingly small. they concur in this view. and when intermediate links have not been found between doubtful forms. that in the larger genera. they confirm the view. the species of the larger genera resemble varieties. Moreover. more than do the species of the smaller genera.6. Now Fries has remarked in regard to plants. judging by analogy whether or not the amount suffices to raise one or both to the rank of species. But when we come to discuss the principle. namely. and clustered round certain forms—that is. or sections. as far as my imperfect results go. naturalists are compelled to come to a determination by the amount of difference between them. therefore. for they differ from each other by a less than the usual amount of difference. many of the species already manufactured still to a certain extent resemble varieties. But Unequally. And what are varieties but groups of forms. round their parent-species. after deliberation. We have seen that there is no infallible criterion by which to distinguish species and well-marked varieties. of divergence of character. they may generally be divided into sub-genera. that the amount of difference between varieties. unequally related to each other. little groups of species are generally clustered like satellites around other species. as I call it. As Fries has well remarked. Related To Each Other. is much less than that between the species of the same genus. or lesser groups. we shall see how this may be explained. Many Of The Species Included Within The Larger Genera Resemble Varieties In Being Very Closely. I have endeavoured to test this numerically by averages. in the same manner as the varieties of any one species are related to each other. and. Hence the amount of difference is one very important criterion in settling whether two forms should be ranked as species or varieties. and how the lesser differences 66 . I have also consulted some sagacious and experienced observers. when compared with each other or with their parent-species. and it may be said. the species of the larger genera are related to each other. And In Having Restricted Ranges There are other relations between the species of large genera and their recorded varieties which deserve notice. No naturalist pretends that all the species of a genus are equally distinct from each other. Undoubtedly there is one most important point of difference between varieties and species. in which a number of varieties or incipient species greater than the average are now manufacturing. Or the case may be put in another way. In this respect. and.

and in so far resemble varieties.between varieties tend to increase into the greater differences between species. Now. This statement is indeed scarcely more than a truism. Watson has divided Great Britain. their denominations would be reversed.9 of the provinces into which Mr. for if a variety were found to have a wider range than that of its supposed parent-species. but which he considers as so closely allied to other species as to be of doubtful value: these sixty-three reputed species range on an average over 6. Varieties generally have much restricted ranges. but which are almost universally ranked by British botanists as good and true species. fifty-three acknowledged varieties are recorded.3 provinces. For instance. Mr. the species to which these varieties belong range over 14. H. So that the acknowledged varieties have very nearly the same restricted average range. 67 . often have much restricted ranges. in this same catalogue. as have the closely allied forms.7 provinces. and these range over 7. Watson as doubtful species. whereas. But there is reason to believe that the species which are very closely allied to other species. Watson has marked for me in the wellsifted London catalogue of Plants (4th edition) sixty-three plants which are therein ranked as species. There is one other point which is worth notice.C. marked for me by Mr.

if species once existed as varieties. But. and varieties. by the discovery of intermediate linking forms. tend to become converted into new and distinct species. notwithstanding that they cannot be closely connected. varieties cannot be distinguished from species—except. And thus. are generally ranked as varieties. the species of these genera have more than the average number of varieties. In all these respects the species of large genera present a strong analogy with varieties. as we shall hereafter see. if differing very little. secondly. by steps hereafter to be explained. the larger genera also tend to break up into smaller genera.7. In large genera the species are apt to be closely but unequally allied together. And we can clearly understand these analogies. 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. and. whereas. forming little clusters round other species. Thus the larger genera tend to become larger. but the amount of difference considered necessary to give to any two forms the rank of species cannot be defined. In genera having more than the average number of species in any country. and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. 68 . Summary Finally. these analogies are utterly inexplicable if species are independent creations. for two forms. first. the forms of life throughout the universe become divided into groups subordinate to groups. by a certain indefinite amount of difference between them. Species very closely allied to other species apparently have restricted ranges. and thus originated.

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. 69 .

what rank. become ultimately converted into good and distinct 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. which constitute what are called distinct genera and which differ from each other more than do the species of the same genus. and to the conditions of life and of one organic being to another being. in their infinitely complex relations to other organic beings and to their physical conditions of life. also. and will generally be inherited by the offspring. Again. the two or three hundred doubtful forms of British plants are entitled to hold. 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. which I have called incipient species. in the structure of the beetle which dives through the water. by which each slight variation. for. follow from the struggle for life. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties.1. 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. been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe. of the Survival of the Fittest. arise? All these results. I have called this principle. if useful. But the mere existence of individual variability and of some few well-marked varieties. and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird. of the many individuals of any species which are periodically born. helps us but little in understanding how species arise in nature. though necessary as the foundation for the work. How have all those exquisite adaptations of one part of the organisation to another part. however slight and from whatever cause proceeding. in the plumed seed which is wafted by the gentlest breeze. is preserved. if they be in any degree profitable to the individuals of a species. Herbert Spencer. for instance. will tend to the preservation of such individuals. in order to mark its relation to man's power of selection. Owing to this struggle. it may be asked. by the term natural selection. if the existence of any well-marked varieties be admitted. But the expression often used by Mr. in short. will thus have a better chance of surviving. The offspring. is 70 . as we shall more fully see in the next chapter. variations. how is it that varieties. but a small number can survive. we see beautiful adaptations everywhere and in every part of the organic world.

In regard to plants. at greater length. 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. and variation. are destroyed by birds and beasts of prey. that. as it well deserves. or their nestlings.more accurate. we often see superabundance of food. we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds. and can adapt organic beings to his own uses. rarity. given to him by the hand of Nature. we shall hereafter see. 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. will be dimly seen or quite misunderstood. extinction. or we forget how largely these songsters. In my future work this subject will be treated. 71 . The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. We have seen that man by selection can certainly produce great results. We behold the face of nature bright with gladness. We will now discuss in a little more detail the struggle for existence. Herbert. abundance. the whole economy of nature. Nothing is easier than to admit in words the truth of the universal struggle for life. and is as immeasurably superior to man's feeble efforts. and are thus constantly destroying life. But Natural Selection. through the accumulation of slight but useful variations. is a power incessantly ready for action. Dean of Manchester. though food may be now superabundant. evidently the result of his great horticultural knowledge. as the works of Nature are to those of Art. or their eggs. and is sometimes equally convenient. Yet unless it be thoroughly engrained in the mind.

Struggle For Existence. The Term. growing close together on the same branch. and including (which is more important) not only the life of the individual. The mistletoe is dependent on the apple and a few other trees. In these several senses. but success in leaving progeny. 72 . may be truly said to struggle with each other which shall get food and live. including dependence of one being on another. for. in a time of dearth. if too many of these parasites grow on the same tree. it languishes and dies. may more truly be said to struggle with each other. 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. but can only in a far-fetched sense be said to struggle with these trees. in tempting the birds to devour and thus disseminate its seeds. its existence depends on them. 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. I use for convenience sake the general term of Struggle for Existence. which pass into each other. But several seedling mistletoes.2. Two canine animals. of which only one of an average comes to maturity. A plant which annually produces a thousand seeds. As the mistletoe is disseminated by birds. Used In A Large Sense I should premise that I use this term in a large and metaphorical sense.

and goes on breeding till ninety years old. in numbers. It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms. all cannot do so. if not destroyed. and surviving till one hundred years old. for in this case there can be no artificial increase of food. Although some species may be now increasing. must suffer destruction during some period of its life. which during its natural lifetime produces several eggs or seeds. for the world would not hold them. there must in every case be a struggle for existence. if the statements of the rate of increase of slow-breeding cattle and horses in South 73 . that. and at this rate. the numerous recorded cases of the astonishingly rapid increase of various animals in a state of nature. Geometrical Ratio Of Increase A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. there would literally not be standing room for his progeny. Even slow-breeding man has doubled in twenty-five years. Every being. But we have better evidence on this subject than mere theoretical calculations. more or less rapidly.3. when circumstances have been favourable to them during two or three following seasons. as more individuals are produced than can possibly survive. Linnaeus has calculated that if an annual plant produced only two seeds—and there is no plant so unproductive as this—and their seedlings next year produced two. bringing forth six young in the interval. There is no exception to the rule that every organic being naturally increases at so high a rate. after a period of from 740 to 750 years there would be nearly nineteen million elephants alive descended from the first pair. or with the physical conditions of life. on the principle of geometrical increase. Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world. it will be safest to assume that it begins breeding when thirty years old. then in twenty years there would be a million plants. otherwise. or with the individuals of distinct species. either one individual with another of the same species. in less than a thousand years. and so on. its numbers would quickly become so inordinately great that no country could support the product. namely. if this be so. Hence. and during some season or occasional year. and no prudential restraint from marriage. and I have taken some pains to estimate its probable minimum rate of natural increase. The elephant is reckoned the slowest breeder of all known animals. the earth would soon be covered by the progeny of a single pair.

is. 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. like the hippobosca. I think. and there are plants which now range in India. One fly deposits hundreds of eggs. and among animals there are very few which do not annually pair. and that there has consequently been less destruction of the old and young and that nearly all the young have been enabled to breed. and yet in the same country the condor may be the more numerous of the two. So it is with plants. Our familiarity with the larger domestic animals tends. Several of the plants. simply explains their extraordinarily rapid increase and wide diffusion in their new homes. to mislead us. and that in a state of nature an equal number would have somehow to be disposed of. and we do not keep in mind that thousands are annually slaughtered for food. no one supposes that the fertility of the animals or plants has been suddenly and temporarily increased in any sensible degree. In a state of nature almost every full-grown plant annually produces seed. had not been well authenticated. yet it is believed to be the most numerous bird in the world. a whole district. The condor lays a couple of eggs and the ostrich a score. and endless others could be given. Their geometrical ratio of increase. The Fulmar petrel lays but one egg. the result of which never fails to be surprising. In such cases. A large number of eggs is of some importance to those species which depend on a fluctuating amount of food. such as the cardoon and a tall thistle. as I hear from Dr. under favourable conditions. and those which produce extremely few. and latterly in Australia. and another. for it allows them rapidly to increase in 74 . we see no great destruction falling on them. let it be ever so large. and that this geometrical tendency to increase must be checked by destruction at some period of life. Falconer. The only difference between organisms which annually produce eggs or seeds by the thousand. a single one. which have been imported from America since its discovery.America. 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. they would have been incredible. which are now the commonest over the wide plains of La Plata. have been introduced from Europe. from Cape Comorin to the Himalaya. But this difference does not determine how many individuals of the two species can be supported in a district. cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years. The obvious explanation is that the conditions of life have been highly favourable.

many must be produced or the species will become extinct. a small number may be produced. It would suffice to keep up the full number of a tree. and yet the average stock be fully kept up. but if many eggs or young are destroyed. if a single seed were produced once in a thousand years. But the real importance of a large number of eggs or seeds is to make up for much destruction at some period of life. 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. in all cases. that each lives by a struggle at some period of its life. and the number of the species will almost instantaneously increase to any amount. Lighten any check. and this period in the great majority of cases is an early one. supposing that this seed were never destroyed and could be ensured to germinate in a fitting place. which lived on an average for a thousand years. In looking at Nature. the average number of any animal or plant depends only indirectly on the number of its eggs or seeds. If an animal can in any way protect its own eggs or young.number. that heavy destruction inevitably falls either on the young or old during each generation or at recurrent intervals. so that. mitigate the destruction ever so little. 75 .

grouse. on a piece of ground three feet long and two wide. but this is not invariably the case. If not one head of game were shot during the next twenty years in England. at the same time. chiefly by slugs and insects. Nature Of The Checks To Increase The causes which check the natural tendency of each species to increase are most obscure. if no vermin were destroyed. If turf which has long been mown. more especially in regard to the feral animals of South America. but the serving as prey to other animals. in some cases. which determines the average number of a species. even in regard to mankind. Seedlings. On the other hand. We know not exactly what the checks are even in a single instance. are destroyed in vast numbers by various enemies. and. dug and cleared. also. Here I will make only a few remarks. for instance. gives the extreme limit to which each can increase. be less game than at present. there seems to be little doubt that the stock of partridges. by so much will it tend to increase still further. none are destroyed by beasts of prey. and out of 357 no less than 295 were destroyed.4. 76 . though fully grown plants. but very frequently it is not the obtaining food. as with the elephant. the more vigorous plants gradually kill the less vigorous. of course. by as much as it swarms in numbers. although so incomparably better known than any other animal. 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. although hundreds of thousands of game animals are now annually shot. and hares on any large estate depends chiefly on the destruction of vermin. and the case would be the same with turf closely browsed by quadrupeds. in all probability. The amount of food for each species. With plants there is a vast destruction of seeds. and where there could be no choking from other plants. This subject of the checks to increase has been ably treated by several authors. there would. Nor will this surprise any one who reflects how ignorant we are on this head. thus out of twenty species grown on a little plot of mown turf (three feet by four) nine species perished. I marked all the seedlings of our native weeds as they came up. Thus. Eggs or very young animals seem generally to suffer most. be let to grow. and I hope in a future work to discuss it at considerable length. Look at the most vigorous species. from the other species being allowed to grow up freely. just to recall to the reader's mind some of the chief points. for even the tiger in India most rarely dares to attack a young elephant protected by its dam.

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. the other species must decrease. The action of climate seems at first sight to be quite independent of the struggle for existence. it brings on the most severe struggle between the individuals. but in a somewhat lesser degree. which subsist on the same kind of food. acts directly. due to the DIRECTLY injurious action of climate. Even when climate. and therefore of competitors. we invariably see some species gradually getting rarer and rarer. hence in going northward. they will increase in numbers. than we do in proceeding southward or in descending a mountain. the struggle for life is almost exclusively with the elements. we may feel sure that the cause lies quite as much in other species being favoured. But this is a false view.Climate plays an important part in determining the average numbers of a species. for the number of species of all kinds. and the change of climate being conspicuous. is an extraordinarily severe mortality from epidemics with man. we forget that each species. we are tempted to attribute the whole effect to its direct action. whether of the same or of distinct species. for they cannot compete with our native plants nor resist destruction by our native animals. or from a damp region to a dry. decreases northward. 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. When we reach the Arctic regions. extreme cold. or snow-capped summits. So it is when we travel northward. 77 . or absolute deserts. and this is a tremendous destruction. or in ascending a mountain. or those which have got least food through the advancing winter. When we travel from south to north. for instance. and if these enemies or competitors be in the least degree favoured by any slight change of climate. even where it most abounds. we far oftener meet with stunted forms. but which never become naturalised. and periodical seasons of extreme cold or drought seem to be the most effective of all checks. When we travel southward and see a species decreasing in numbers. from enemies or from competitors for the same place and food. which will suffer the most. and as each area is already fully stocked with inhabitants. when we remember that ten per cent. as in this one being hurt. is constantly suffering enormous destruction at some period of its life. it will be the least vigorous individuals. but in so far as climate chiefly acts in reducing food. and finally disappearing.

relatively to the numbers of its enemies. I should add that the good effects of intercrossing. which have from some cause. I have in this case lost every single seed. this seems generally to occur with our game animals—often ensue. possibly in part through facility of diffusion among the crowded animals.. explains. nor can the birds. in many cases. I believe. is absolutely necessary for its preservation. in the few spots where they do exist. that a plant could exist only where the conditions of its life were so favourable that many could exist together. This view of the necessity of a large stock of the same species for its preservation. but I will not here enlarge on this subject. that is abounding in individuals.When a species. in our fields. some singular facts in nature such as that of very rare plants being sometimes extremely abundant. though having a superabundance of food at this one season. and that of some social plants being social. no doubt come into play in many of these cases. owing to highly favourable circumstances. and thus save the species from utter destruction. as their numbers are checked during the winter. even on the extreme verge of their range. But even some of these so-called epidemics appear to be due to parasitic worms. epidemics—at least. 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. been disproportionally favoured: and here comes in a sort of struggle between the parasite and its prey. etc. On the other hand. because the seeds are in great excess compared with the number of birds which feed on them. and here we have a limiting check independent of the struggle for life. a large stock of individuals of the same species. Thus we can easily raise plenty of corn and rape-seed. we may believe. 78 . For in such cases. increase in number proportionally to the supply of seed. increases inordinately in numbers in a small tract. and the ill effects of close interbreeding.

Here we see how potent has been the effect of the introduction of a single tree. so close together that all cannot live. but twelve species of plants (not counting grasses and carices) flourished in the plantations. and self-sown firs are now springing up in multitudes. which have to struggle together in the same country. The change in the native vegetation of the planted part of the heath was most remarkable. In one square yard. it became thickly clothed with vigorously growing young firs. I found a multitude of seedlings and little trees. 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. and had failed. nothing whatever else having been done. whence I could examine hundreds of acres of the unenclosed heath. 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. on the estate of a relation. But how important an element enclosure is. had. which had been perpetually browsed down by the cattle. during many years tried to raise its head above the stems of the heath. The effect on the insects must have been still greater. with twenty-six rings of growth. 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 will give only a single instance. No wonder that. in Surrey. Here there are extensive heaths. which were not to be seen on the heath. and one of them. for six insectivorous birds were very common in the plantations. with the exception of the land having been enclosed. In Staffordshire. as soon as the land was enclosed. which had never been touched by the hand of man. which. at a point some hundred yards distant from one of the old clumps. But on looking closely between the stems of the heath.5. I counted thirty-two little trees. and the heath was frequented by two or three distinct insectivorous birds. I plainly saw near Farnham. and literally I could not see a single Scotch fir. where I had ample means of investigation. but several hundred acres of exactly the same nature had been enclosed twenty-five years previously and planted with Scotch fir. with a few clumps of old Scotch firs on the distant hill-tops: within the last ten years large spaces have been enclosed. though a simple one. interested me. there was a large and extremely barren heath. Yet the heath was so extremely barren 79 . except the old planted clumps. which could not be found on the heath. so that cattle could not enter.

I find from experiments that humblebees are almost indispensable to the fertilisation of the heartsease (Viola tricolor). and yet in the long-run the forces are so nicely balanced that the face of nature remains for long periods of time uniform. if certain insectivorous birds were to decrease in Paraguay. and consequently. and so high our presumption. for here neither cattle nor horses nor dogs have ever run wild. which lays its eggs in the navels of these animals when first born. 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. though they swarm southward and northward in a feral state. and Azara and Rengger have shown that this is caused by the greater number in Paraguay of a certain fly.700 seeds. Again. as we have just seen in Staffordshire. that we marvel when we hear of the extinction of an organic being. and as we do not see the cause. and this would lessen the number of the navel-frequenting flies—then cattle and horses would become feral. for other bees do not visit this flower. and this. or invent laws on the duration of the forms of life! I am tempted to give one more instance showing how plants and animals. numerous as they are. but twenty other heads. pratense) produced 2. the parasitic insects would probably increase.and so extensive that no one would ever have imagined that cattle would have so closely and effectually searched it for food. produced not one.290 seeds. 100 heads of red clover (T. probably by other parasitic insects. the insectivorous birds. remote in the scale of nature. and so onwards in ever-increasing circles of complexity. 80 . Not that under nature the relations will ever be as simple as this. so profound is our ignorance. for instance twenty heads of Dutch clover (Trifolium repens) yielded 2. I have also found that the visits of bees are necessary for the fertilisation of some kinds of clover. we invoke cataclysms to desolate the world. must be habitually checked by some means. but in several parts of the world insects determine the existence of cattle. Perhaps Paraguay offers the most curious instance of this. Nevertheless. are bound together by a web of complex relations. Nearly all our orchidaceous plants absolutely require the visits of insects to remove their pollenmasses and thus to fertilise them. Hence. though assuredly the merest trifle would give the victory to one organic being over another. I shall hereafter have occasion to show that the exotic Lobelia fulgens is never visited in my garden by insects. Here we see that cattle absolutely determine the existence of the Scotch fir. protected from bees. never sets a seed. Battle within battle must be continually recurring with varying success. The increase of these flies. from its peculiar structure.

each annually scattering its seeds by the thousand. many different checks. and Colonel Newman says. acting at different periods of life. believes that "more than two-thirds of them are thus destroyed all over England. but I doubt whether they could do so in the case of the red clover. snails. as other bees cannot reach the nectar. which destroy their combs and nests. from their weight not being sufficient to depress the wing petals. which must formerly have been cleared of trees. the heartsease and red clover would become very rare. In some cases it can be shown that widely-different checks act on the same species in different districts. what war between insect and insect—between insects." Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine. or even the existence of the species. through the intervention first of mice and then of bees. all feeding on each other. Throw up a handful of feathers. The number of humble-bees in any district depends in a great measure upon the number of field-mice. but all will concur in determining the average number. as every one knows. a very different vegetation springs up. the frequency of certain flowers in that district! In the case of every species.but the same number of protected heads produced not a single seed. and during different seasons or years." Now the number of mice is largely dependent. Humble bees alone visit red clover. or on the trees. some one check or some few being generally the most potent. and Colonel Newman. and other animals with birds and beasts of prey—all striving to increase. which I attribute to the number of cats that destroy the mice. "Near villages and small towns I have found the nests of humble-bees more numerous than elsewhere. It has been suggested that moths may fertilise the clovers. their seeds and seedlings. But how false a view is this! Every one has heard that when an American forest is cut down. now display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. but it has been observed that ancient Indian ruins in the Southern United States. or wholly disappear. we are tempted to attribute their proportional numbers and kinds to what we call chance. and all fall to the ground according to definite laws. When we look at the plants and bushes clothing an entangled bank. or on the other plants which first clothed the ground and thus checked the growth of the trees. on the number of cats. Hence we may infer as highly probable that. if the whole genus of humble-bees became extinct or very rare in England. but how simple is the problem where each shall 81 . What a struggle must have gone on during long centuries between the several kinds of trees. who has long attended to the habits of humble-bees. probably come into play.

the proportional numbers and kinds of trees now growing on the old Indian ruins! The dependency of one organic being on another. in the same manner as beings in a state of nature. otherwise the weaker kinds will steadily decrease in number and disappear. or are naturally the most fertile. So again with the varieties of sheep: it has been asserted that certain mountain-varieties will starve out other mountain-varieties. they must be each year harvested separately. that the original proportions of a mixed stock (crossing being prevented) could be kept up for half-a-dozen generations. if they were allowed to struggle together. for they frequent the same districts. But the struggle will almost invariably be most severe between the individuals of the same species. and we sometimes see the contest soon decided: for instance. and constitution. It may even be doubted whether the varieties of any of our domestic plants or animals have so exactly the same strength. if several varieties of wheat be sown together. require the same food. and are exposed to the same dangers. in the course of centuries. the struggle will generally be almost equally severe. some of the varieties which best suit the soil or climate. 82 . as in the case of locusts and grassfeeding quadrupeds. so that they cannot be kept together. This is likewise sometimes the case with those which may strictly be said to struggle with each other for existence. and the mixed seed be resown. and will consequently in a few years supplant the other varieties. habits. and if the seed or young were not annually preserved in due proportion. as of a parasite on its prey.fall compared to that of the action and reaction of the innumerable plants and animals which have determined. lies generally between beings remote in the scale of nature. The same result has followed from keeping together different varieties of the medicinal leech. will beat the others and so yield more seed. To keep up a mixed stock of even such extremely close varieties as the variously coloured sweet-peas. In the case of varieties of the same species. and the seed then mixed in due proportion.

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. in the most essential yet often hidden manner. and in the flattened and fringed legs of the water-beetle. the struggle will generally be more severe between them. 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. which fill nearly the same place in the economy of nature. that the structure of every organic being is related. so that the seeds may be widely distributed and fall on unoccupied ground. stingless native bee. the structure of its legs. In the water-beetle.6. or on which it preys. so well adapted for diving. 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 hunt for its own prey. with which it comes into competition for food or residence. In Australia the imported hive-bee is rapidly exterminating the small. A corollary of the highest importance may be deduced from the foregoing remarks. and so in other cases. or from which it has to escape. though by no means invariably. and to escape serving as prey to other animals. namely. Yet the advantage of the plumed seeds no doubt stands in the closest relation to the land being already thickly clothed with other plants. the relation seems at first confined to the elements of air and water. if they come into competition with each other. much similarity in habits and constitution. This is obvious in the structure of the teeth and talons of the tiger. 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. as peas and beans. But from the strong growth of young plants produced from such seeds. allows it to compete with other aquatic insects. We can dimly see why the competition should be most severe between allied forms. and always in structure. it may be suspected that 83 . but probably in no one case could we precisely say why one species has been victorious over another in the great battle of life. than between the species of distinct genera. to that of all other organic beings. One species of charlock has been known to supplant another species. 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. Struggle For Life Most Severe Between Individuals And Varieties Of The Same Species As the species of the same genus usually have.

dampness or dryness. When we reflect on this struggle we may console ourselves with the full belief that the war of nature is not incessant. damper or drier districts. and that the vigorous. for elsewhere it ranges into slightly hotter or colder. for the warmest or dampest spots. and the happy survive and multiply. 84 . 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. the healthy. a conviction as necessary. On the confines of its geographical range. we should have to give it some advantage over its competitors. or at intervals. In this case we can clearly see that if we wish in imagination to give the plant the power of increasing in numbers. Not until we reach the extreme confines of life. All that we can do is to keep steadily in mind that each organic being is striving to increase in a geometrical ratio. as it is difficult to acquire. we should have to modify it in a different way to what we should have had to do in its native country. for we should have to give it some advantage over a different set of competitors or enemies. This ought to convince us of our ignorance on the mutual relations of all organic beings. a change of constitution with respect to climate would clearly be an advantage to our plant. during some season of the year. although the climate may be exactly the same as in its former home. has to struggle for life and to suffer great destruction. that no fear is felt. at some period of its life. among new competitors. in the Arctic regions or on the borders of an utter desert. The land may be extremely cold or dry. It is good thus to try in imagination to give any one species an advantage over another.the chief use of the nutriment in the seed is to favour the growth of the seedlings. Probably in no single instance should we know what to do. during each generation. whilst struggling with other plants growing vigorously all around. or over the animals which prey on it. will competition cease. the conditions of its life will generally be changed in an essential manner. but we have reason to believe that only a few plants or animals range so far. that they are destroyed exclusively by the rigour of the climate. or between the individuals of the same species. that death is generally prompt. that each. If its average numbers are to increase in its new home. Hence we can see that when a plant or animal is placed in a new country. yet there will be competition between some few species.

intercrossing. 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. 85 . isolation. through Divergence of Character and Extinction. 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.Chapter 4 Natural Selection. related to the diversity of inhabitants of any small area and to naturalisation — Action of Natural Selection. number of individuals — Slow action — Extinction caused by Natural Selection — Divergence of Character. namely.

apply under nature? I think we shall see that it can act most efficiently. which we have seen is so potent in the hands of man. I have called Natural Selection. or would ultimately become fixed. in a lesser degree. Some have even imagined that natural selection induces variability. as well as the strength of the hereditary tendency. over others. which we almost universally meet with in our domestic productions is not directly produced. as perhaps we see in certain polymorphic species. Let the endless number of slight variations and individual differences occurring in our domestic productions. briefly discussed in the last chapter. and would be left either a fluctuating element.1. as Hooker and Asa Gray have well remarked. How will the struggle for existence. This preservation of favourable individual differences and variations. should occur in the course of many successive generations? If such do occur. and consequently what infinitely varied diversities of structure might be of use to each being under changing conditions of life. he can neither originate varieties nor prevent their occurrence. and the destruction of those which are injurious. in those under nature. But the variability. or the Survival of the Fittest. Unintentionally he exposes organic beings to new and changing conditions of life. Variations neither useful nor injurious would not be affected by natural selection. that other variations useful in some way to each being in the great and complex battle of life. but similar changes of conditions might and do occur under nature. seeing that variations useful to man have undoubtedly occurred. would have the best chance of surviving and procreating their kind? On the other hand. 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. it may truly be said that the whole organisation becomes in some degree plastic. act in regard to variation? Can the principle of selection. and variability ensues. by man. however slight. and. whereas it implies only the preservation of such variations as arise and are beneficial to the being under its conditions of life. owing to the nature of the organism and the nature of the conditions. Under domestication. Can it then be thought improbable. we may feel sure that any variation in the least degree injurious would be rigidly destroyed. he can only preserve and accumulate such as do occur. Several writers have misapprehended or objected to the term Natural Selection. can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage. No one 86 . be borne in mind.

no doubt. would tend to be preserved. from what we have seen of the intimate and complex manner in which the inhabitants of each country are bound together. by better adapting them to their altered conditions. of climate. must of necessity first occur. In such cases. which in any way favoured the individuals of any species. It has been said that I speak of natural selection as an active power or Deity. and in this case the individual differences given by nature. We shall best understand the probable course of natural selection by taking the case of a country undergoing some slight physical change. these same places would have been seized on by intruders. The proportional numbers of its inhabitants will almost immediately undergo a change. for. which man for some object selects. into which new and better adapted forms could not freely enter. new forms would certainly immigrate. and they are almost necessary for brevity. 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. Others have objected that the term selection implies conscious choice in the animals which become modified. as plants have no volition. that any change in the numerical proportions of the inhabitants. 87 . but who objects to an author speaking of the attraction of gravity as ruling the movements of the planets? Every one knows what is meant and is implied by such metaphorical expressions. and natural selection would have free scope for the work of improvement. would seriously affect the others. So again it is difficult to avoid personifying the word Nature. independently of the change of climate itself. 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. for instance. With a little familiarity such superficial objections will be forgotten. and it has even been urged that. 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. or of a country partly surrounded by barriers. natural selection is not applicable to them! In the literal sense of the word.objects to agriculturists speaking of the potent effects of man's selection. and by laws the sequence of events as ascertained by us. natural selection is a false term. and some species will probably become extinct. We may conclude. But in the case of an island. If the country were open on its borders. but I mean by nature. only the aggregate action and product of many natural laws. and this would likewise seriously disturb the relations of some of the former inhabitants. slight modifications.

He seldom exercises each selected character in some peculiar and fitting manner. for in all countries. so as to have better resisted the intruders. as shown in the first chapter." it must never be forgotten that mere individual differences are included. on the whole machinery of life. cares nothing for appearances. As man can produce a great result with his domestic animals and plants by adding up in any given direction individual differences. we may safely conclude that the natives might have been modified with advantage. as is implied by the fact of their selection. as long as the species continued under the same conditions of life and profited by similar means of subsistence and defence. or any unusual degree of isolation. if I may be allowed to personify the natural preservation or survival of the fittest. the natives have been so far conquered by naturalised productions that they have allowed some foreigners to take firm possession of the land. Unless such occur. except in so far as they are useful to any being.We have good reason to believe. Nature only for that of the being which she tends. Every selected character is fully exercised by her. and still further modifications of the same kind would often still further increase the advantage. what may not natural selection effect? Man can act only on external and visible characters: Nature. extremely slight modifications in the structure or habits of one species would often give it an advantage over others. so could natural selection. he feeds a long and a short-beaked pigeon 88 . that changes in the conditions of life give a tendency to increased variability. 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. Man selects only for his own good. but far more easily from having incomparably longer time for action. She can act on every internal organ. For as all the inhabitants of each country are struggling together with nicely balanced forces. and certainly has produced. 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. by affording a better chance of the occurrence of profitable variations. and this would manifestly be favourable to natural selection. And as foreigners have thus in every country beaten some of the natives. as of climate. and in the foregoing cases the conditions the changed. on every shade of constitutional difference. to check immigration. Nor do I believe that any great physical change. As man can produce. a great result by his methodical and unconscious means of selection. Under the term of "variations. Man keeps the natives of many climates in the same country. that none of them could be still better adapted or improved. natural selection can do nothing.

Seeing that individual differences of the same kind perpetually recur. as far as lies in his power. But whether it is true. we can judge only by seeing how far the hypothesis accords with and explains the general phenomena of nature. is likewise a simple assumption. When we see leaf- 89 . or at least by some modification prominent enough to catch the eye or to be plainly useful to him. Although natural selection can act only through and for the good of each being. 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. he exposes sheep with long and short wool to the same climate. until the hand of time has marked the long lapse of ages. he does not rigidly destroy all inferior animals. then.on the same food. compared with those accumulated by Nature during whole geological periods! Can we wonder. may thus be acted on. which we are apt to consider as of very trifling importance. rejecting those that are bad. preserving and adding up all that are good. a variety. does not allow the most vigorous males to struggle for the females. On the other hand. all his productions. he does not exercise a long-backed or long-legged quadruped in any peculiar manner. the ordinary belief that the amount of possible variation is a strictly limited quantity. throughout the world. this can hardly be considered as an unwarrantable assumption. perhaps after a long interval of time. and should plainly bear the stamp of far higher workmanship? It may metaphorically be said that natural selection is daily and hourly scrutinising. and so onward. Under nature. yet characters and structures. the slightest differences of structure or constitution may well turn the nicely-balanced scale in the struggle for life. and so be preserved. step by step. We see nothing of these slow changes in progress. at the improvement of each organic being in relation to its organic and inorganic conditions of life. that Nature's productions should be far "truer" in character than man's productions. and these must again be preserved. He often begins his selection by some half-monstrous form. and consequently how poor will be his results. that they should be infinitely better adapted to the most complex conditions of life. vary or present individual differences of the same favourable nature as before. silently and insensibly working. In order that any great amount of modification should be effected in a species. WHENEVER AND WHEREVER OPPORTUNITY OFFERS. but protects during each varying season. the slightest variations. How fleeting are the wishes and efforts of man! How short his time. when once formed must again.

where the trees would have to struggle with other trees and with a host of enemies. whereas another disease attacks yellow-fleshed peaches far more than those with other coloured flesh. We have seen how the colour of hogs. these slight differences make a great difference in cultivating the several varieties. the down on the fruit and the colour of the flesh are considered by botanists as characters of the most trifling importance. such differences would effectually settle which variety. a Curculio. as far as our ignorance permits us to judge. if not destroyed at some period of their lives. that in the United States smooth-skinned fruits suffer far more from a beetle. appear at any particular period of life. should succeed. determines whether they shall live or die. the alpine ptarmigan white in winter. yet we hear from an excellent horticulturist. when one part varies and the variations are accumulated through natural selection. the red-grouse the colour of heather. assuredly. tend to reappear in the offspring at the same period. that purple plums suffer far more from a certain disease than yellow plums. and in keeping that colour.—so much so that on parts of the continent persons are warned not to keep white pigeons. under domestication. and bark-feeders mottled-grey. Downing. It is also necessary to bear in mind that. In looking at many small points of difference between species. when once acquired. would increase in countless numbers. As we see that those variations which. for instance. In plants. Nor ought we to think that the occasional destruction of an animal of any particular colour would produce little effect. true and constant. other modifications. and in the colour of the down of their chickens. often of the most unexpected nature. which. whether a smooth or downy. and hawks are guided by eyesight to their prey. with all the aids of art. If. food. Grouse. in the eggs of poultry. in the caterpillar and cocoon stages of the varieties of the silkworm. than those with down. we must believe that these tints are of service to these birds and insects in preserving them from danger. owing to the law of correlation. etc. which feed on the "paint-root" in Virginia. have no doubt produced some direct effect. in a state of nature. we should remember how essential it is in a flock of white sheep to destroy a lamb with the faintest trace of black.eating insects green. seem quite unimportant. they are known to suffer largely from birds of prey. we must not forget that climate. will ensue. as being the most liable to destruction. in the shape. in the horns of our sheep 90 . Hence natural selection might be effective in giving the proper colour to each kind of grouse. size and flavour of the seeds of the many varieties of our culinary and agricultural plants.. a yellow or a purple-fleshed fruit.

the thickness of the shell being known to vary like every other structure. is to modify the structure of one species. which can have little or no influence on the course of natural selection. which had the most powerful and hardest beaks. but in all cases natural selection will ensure that they shall not be injurious: for if they were so. I can see no greater difficulty in this being effected through natural selection. Natural selection may modify and adapt the larva of an insect to a score of contingencies. for all with weak beaks would inevitably perish: or. Now. For instance. If it profit a plant to have its seeds more and more widely disseminated by the wind. through correlation. the structure of the adult. 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. So. for instance. the species would become extinct. and by their inheritance at a corresponding age. used for breaking the eggs. for the good of another species. by the accumulation of variations profitable at that age. so that fanciers assist in the act of hatching. It has been asserted.and cattle when nearly adult. so in a state of nature natural selection will be enabled to act on and modify organic beings at any age. without giving it any advantage. and though statements to this effect may be found in works of natural history. wholly different from those which concern the mature insect. and these could be modified through natural selection 91 . conversely. if of high importance to it. that of the best short-beaked tumbler-pigeons a greater number perish in the egg than are able to get out of it. if nature had to make the beak of a full-grown pigeon very short for the bird's own advantage. more delicate and more easily broken shells might be selected. the great jaws possessed by certain insects. the process of modification would be very slow. It may be well here to remark that with all beings there must be much fortuitous destruction. and there would be simultaneously the most rigorous selection of all the young birds within the egg. I cannot find one case which will bear investigation. and these modifications may affect. might be modified to any extent by natural selection. What natural selection cannot do. used exclusively for opening the cocoon—or the hard tip to the beak of unhatched birds. A structure used only once in an animal's life. if the community profits by the selected change. modifications in the adult may affect the structure of the larva. In social animals it will adapt the structure of each individual for the benefit of the whole community. Natural selection will modify the structure of the young in relation to the parent and of the parent in relation to the young.

have yielded individuals better adapted to their conditions of life than any of those which happened to survive. but this is no valid objection to its efficiency at other times and in other ways. So again a vast number of mature animals and plants.only if they varied in some manner which protected them from their enemies. the best adapted individuals. Yet many of these eggs or seeds would perhaps. 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. if not destroyed. 92 . will tend to propagate their kind in larger numbers than the less well adapted. must be annually destroyed by accidental causes. whether or not they be the best adapted to their conditions. supposing that there is any variability in a favourable direction. 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. But let the destruction of the adults be ever so heavy. as will often have been the case. natural selection will be powerless in certain beneficial directions. 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. If the numbers be wholly kept down by the causes just indicated.

confined to the male sex. male alligators have been described as fighting. and these seem oftenest provided with special weapons. might surely give indomitable courage. though to them and to others. less rigorous than natural selection. the males of certain hymenopterous insects have been frequently seen by that inimitable observer M. so no doubt it will be under nature. and strength to the wing to strike in the spurred leg. as the mane of the lion. special means of defence may be given through means of sexual selection. fighting for a particular female who sits by. for the possession of the females. Sexual selection. Fabre. The males of carnivorous animals are already well armed.2. and the hooked jaw to the male salmon. by always allowing the victor to breed. male salmons have been observed fighting all day long. length of spur. This leads me to say a few words on what I have called sexual selection. the contest is often of a more peaceful character. for the possession of the other sex. therefore. believe that there is the severest rivalry between the males of many species to attract. But in many cases victory depends not so much on general vigour. and then retires with the conqueror. as is sometimes the case. those which are best fitted for their places in nature. generally the males. Sexual selection is. This form of selection depends. but few or no offspring. male stag-beetles sometimes bear wounds from the huge mandibles of other males. How low in the scale of nature the law of battle descends I know not. The result is not death to the unsuccessful competitor. but on a struggle between the individuals of one sex. in nearly the same manner as does the brutal cockfighter by the careful selection of his best cocks. not on a struggle for existence in relation to other organic beings or to external conditions. like Indians in a war-dance. perhaps. by singing. The war is. the most vigorous males. will leave most progeny. but on having special weapons. bellowing. Among birds. an apparently unconcerned beholder of the struggle. A hornless stag or spurless cock would have a poor chance of leaving numerous offspring. and whirling round. the 93 . as commonly occurs. for the shield may be as important for victory as the sword or spear. or for one sex to be modified in relation to the other sex. severest between the males of polygamous animals. Generally. Sexual Selection Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex. Thus it is rendered possible for the two sexes to be modified through natural selection in relation to different habits of life. All those who have attended to the subject.

and it is doubtful whether it can be ornamental in the eyes of the female bird. which apparently have not been augmented through selection by man. 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. which they have transmitted to their male offspring alone. and transmitted to the males alone or to both sexes at corresponding ages. and some others. during thousands of generations. according to his standard of beauty. or ornament. might produce a marked effect. Yet. The rock-thrush of Guiana. their gorgeous plumage. colour. by individual males having had. had the tuft appeared under domestication it would have been called a monstrosity. but I have not space here to enter on this subject. at last choose the most attractive partner. which. in their weapons. Heron has described how a pied peacock was eminently attractive to all his hen birds. they likewise perform strange antics before the females. The tuft of hair on the breast of the wild turkey-cock cannot be of any use. congregate. some slight advantage over other males. and successive males display with the most elaborate care. and show off in the best manner. or charms. such differences have been mainly caused by sexual selection: that is. but if man can in a short time give beauty and an elegant carriage to his bantams. standing by as spectators. means of defence. with respect to the plumage of male and female birds. indeed. Some well-known laws. 94 . I cannot here enter on the necessary details. that when the males and females of any animal have the same general habits of life. I can see no good reason to doubt that female birds. in successive generations. birds of paradise. in comparison with the plumage of the young. as I believe. 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.females. can partly be explained through the action of sexual selection on variations occurring at different ages. by selecting. but differ in structure. Thus it is. the most melodious or beautiful males.

John. and from the continued preservation of the individuals best fitted for the two sites. one with a light greyhound-like form. with shorter legs. another mice. as I believe. would naturally be forced to hunt different prey. Illustrations Of The Action Of Natural Selection. which pursues deer. for instance. one cat. Even without any change in the proportional numbers of the animals on which our wolf preyed. Now. Pierce. and some by fleetness. in the United States. had from any change in the country increased in numbers. 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. there are two varieties of the wolf inhabiting the Catskill Mountains. if any slight innate change of habit or of structure benefited an individual wolf. I may add that. some by strength.3. it would have the best chance of surviving and of leaving offspring. St. a new variety might be formed which would either supplant or coexist with the parent-form of wolf. which preys on various animals. for we often observe great differences in the natural tendencies of our domestic animals. when they were compelled to prey on other animals. which more frequently attacks the shepherd's flocks. securing some by craft. or that other prey had decreased in numbers. The tendency to catch rats rather than mice is known to be inherited. and so be preserved or selected. taking to catch rats. Under such circumstances the swiftest and slimmest wolves have the best chance of surviving. and those frequenting the lowlands. Or The Survival Of The Fittest In order to make it clear how. and another hunting on marshy ground and almost nightly catching woodcocks or snipes. the wolves inhabiting a mountainous district. and let us suppose that the fleetest prey. another hares or rabbits. according to Mr. a cub might be born with an innate tendency to pursue certain kinds of prey. Nor can this be thought very improbable. than that man should be able to improve the fleetness of his greyhounds by careful and methodical selection. I can see no more reason to doubt that this would be the result. provided always that they retained strength to master their prey at this or some other period of the year. one cat. I must beg permission to give one or two imaginary illustrations. and the other more bulky. during that season of the year when the wolf was hardest pressed for food. Or. according to Mr. Let us take the case of a wolf. natural selection acts. bringing home winged game. again. Some of its young would probably inherit the same habits or structure. 95 . and by the repetition of this process. a deer for instance.

and that. and if one were born with its beak strongly curved. yet the chances would be strongly against its survival. I saw. of which. I did not appreciate how rarely single variations. I may add. it would generally be lost by subsequent intercrossing with ordinary individuals. Pierce. from various causes of destruction. one with a light greyhound-like form. This is rather an extreme estimate for most of the higher animals. giving it twice as good a chance of life as that of the other individuals. and on the destruction of the worst. that. with shorter legs. also. and not of any single strongly marked variation having been preserved. but to this subject of intercrossing we shall soon have to return. but by no means so for many of the lower organisms. In former editions of this work I sometimes spoke as if this latter alternative had frequently occurred. according to Mr. but there can hardly be a doubt. and the other more bulky. would be a rare event. that this result would follow from the preservation during many generations of a large number of individuals 96 . be disputed. that the preservation in a state of nature of any occasional deviation of structure. which more frequently attacks the shepherd's flocks. which pursues deer. until reading an able and valuable article in the "North British Review" (1867). I think. still. producing during their lifetime two hundred offspring. if at first preserved. I saw the great importance of individual differences. and which consequently flourished. He then shows that if a single individual were born. could be perpetuated. for instance. The justice of these remarks cannot. only two on an average survive to pro-create their kind. a bird of some kind could procure its food more easily by having its beak curved. as the Reviewer goes onto show. The author takes the case of a pair of animals. Nevertheless. which varied in some manner. judging by what we see taking place under domestication.two varieties might slowly be formed. I speak of the slimmest individual wolves. and that half its young inherited the favourable variation. and this led me fully to discuss the results of unconscious selection by man. and this chance would go on decreasing in the succeeding generations. If. whether slight or strongly marked. such as a monstrosity. the young would have only a slightly better chance of surviving and breeding. Supposing it to survive and to breed. It should be observed that in the above illustration. nevertheless there would be a very poor chance of this one individual perpetuating its kind to the exclusion of the common form. there are two varieties of the wolf inhabiting the Catskill Mountains in the United States. These varieties would cross and blend where they met. which depends on the preservation of all the more or less valuable individuals.

competing with and conquering the unchanged individuals on the margins of an ever-increasing circle. though small in quantity. frequently recur owing to a similar organisation being similarly acted on— of which fact numerous instances could be given with our domestic productions. This juice. apparently for the sake of eliminating something injurious from the sap: this is effected. and would often breed together. which no one would rank as mere individual differences. It should not. it would slowly spread from a central district. but it may be here remarked that most animals and plants keep to their proper homes. as seems to be the common rule with varieties in a state of nature. we see this even with migratory birds. the original form would soon be supplanted by the modified form. through the survival of the fittest. it would undoubtedly transmit to them. In such cases. however. is greedily sought by insects. if the varying individual did not actually transmit to its offspring its newly-acquired character. as long as the existing conditions remained the same. and at the backs of the leaves of the common laurel. Now. Or only a third. that it was formerly ranked as a distinct species under the name of Uria lacrymans. I shall have to recur. of which fact several instances could be given. fifth. Consequently each newly-formed variety would generally be at first local. Certain plants excrete sweet juice. if the variation were of a beneficial nature. so that similarly modified individuals would soon exist in a small body together. for instance. In cases of this kind. 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. and do not needlessly wander about. It may be worth while to give another and more complex illustration of the action of natural selection. 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. be overlooked that certain rather strongly marked variations. but their visits do not in any way benefit the plant. by glands at the base of the stipules in some Leguminosae. a still stronger tendency to vary in the same manner. Thus Graba estimates that about one-fifth of the guillemots in the Faroe Islands consist of a variety so well marked. To the effects of intercrossing in eliminating variations of all kinds. or tenth part of the individuals may have been thus affected. and from the destruction of a still larger number with the straightest beaks. which almost always return to the same spot.with more or less strongly curved beaks. If the new variety were successful in its battle for life. Insects in seeking the nectar would get 97 .

excreting most nectar. and had larger anthers. there were a few pollen-grains. and the act of crossing. and so in the long-run would gain the upper hand and form a local variety. which had flown from tree to tree in search of nectar. Some holly-trees bear only male flowers. unintentionally on their part. would likewise be favoured. nevertheless every female flower which I examined had been effectually fertilised by the bees. and on some a profusion. and four stamens with shrivelled anthers. and would oftenest be crossed. regularly carry pollen from flower to flower. But to return to our imaginary case. as can be fully proved. The flowers. We might have taken the case of insects visiting flowers for the sake of collecting pollen instead of nectar. taken from different branches. yet if a little pollen were carried. The plants which produced flowers with the largest glands or nectaries. and would often transport it from one flower to another. The weather had been cold and boisterous and therefore not favourable to bees. although nine-tenths of the pollen were destroyed it might still be a great gain to the plant to be thus robbed. I will give only one. under the microscope. in relation to the size and habits of the particular insect which visited them. which consequently would have the best chance of flourishing and surviving. would oftenest be visited by insects. The flowers of two distinct individuals of the same species would thus get crossed. and a cross thus effected. which had their stamens and pistils placed. which have four stamens producing a rather small quantity of pollen. another process 98 . and a rudimentary pistil. would be selected. When our plant. by the pollen-devouring insects from flower to flower. and on all. its destruction appears to be a simple loss to the plant. other holly-trees bear only female flowers. at first occasionally and then habitually. as soon as the plant had been rendered so highly attractive to insects that pollen was regularly carried from flower to flower. Having found a female tree exactly sixty yards from a male tree. gives rise to vigorous seedlings.dusted with pollen. and that they do this effectually I could easily show by many striking facts. As the wind had set for several days from the female to the male tree. I put the stigmas of twenty flowers. the pollen could not thus have been carried. they would. as likewise illustrating one step in the separation of the sexes of plants. and as pollen is formed for the sole purpose of fertilisation. had been rendered highly attractive to insects. in which not a grain of pollen can be detected. also. so as to favour in any degree the transportal of the pollen. and the individuals which produced more and more pollen. without exception. by the above process long continued. these have a full-sized pistil.

I could give many facts showing how anxious bees are to save time: for instance. then. That this nectar is much liked by the hive-bee is certain. but I may add that some of the species of holly in North America are. but not out of the common red clover. their habit of cutting holes and sucking the nectar at the bases of certain flowers. or. which is visited by humble-bees alone. sometimes the male organs and sometimes the female organs become more or less impotent. so that certain individuals would be able to obtain their food more quickly than others. until at last a complete separation of the sexes might be effected. Let us now turn to the nectar-feeding insects. according to Asa Gray.. for I have repeatedly seen. through dimorphism and other means. and thus the communities to which they belonged would flourish and throw off many swarms inheriting the same peculiarities. and as a more complete separation of the sexes of our plant would be advantageous on the principle of the division of labour. we may suppose the plant of which we have been slowly increasing the nectar by continued selection. It would take up too much space to show the various steps. In plants under culture and placed under new conditions of life. now if we suppose this to occur in ever so slight a degree under nature. and pistils alone in another flower or on another plant. No naturalist doubts the advantage of what has been called the "physiological division of labour. in an exactly intermediate condition. too slight to be appreciated by us. yet the hive-bee can easily suck the nectar out of the incarnate clover." hence we may believe that it would be advantageous to a plant to produce stamens alone in one flower or on one whole plant. but only in the autumn. are more or less dioeciously polygamous. to be a common plant. Bearing such facts in mind.might commence. it may be believed that under certain circumstances individual differences in the curvature or length of the proboscis. etc. The difference in the length of the corolla in the 99 . as pollen is already carried regularly from flower to flower. many hive-bees sucking the flowers through holes bitten in the base of the tube by humble bees. might profit a bee or other insect. which with a very little more trouble they can enter by the mouth. so that whole fields of the red clover offer in vain an abundant supply of precious nectar to the hive-bee. by which the separation of the sexes in plants of various kinds is apparently now in progress. 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. as he expresses it. would be continually favoured or selected. individuals with this tendency more and more increased. and that certain insects depended in main part on its nectar for food.

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

and made so many experiments. All vertebrate animals. and a vast majority of plants are hermaphrodites. Knight and Kolreuter. and on the other hand. that these facts alone incline me to believe that it is a general law of nature that no organic being fertilises itself for a perpetuity of generations. Modern research has much diminished the number of supposed hermaphrodites and of real hermaphrodites a large number pair. understand several large classes of facts. Nevertheless there is reason to believe that with all hermaphrodites two individuals. it may be asked. that CLOSE interbreeding diminishes vigour and fertility. What reason. I think. 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. two individuals regularly unite for reproduction. yet what a multitude of flowers have their anthers and stigmas fully exposed to the weather! If an occasional cross be indispensable. In the case of animals and plants with separated sexes. pair for each birth. I must trust to some general considerations alone. which on any other view are inexplicable. either occasionally or habitually. But still there are many hermaphrodite animals which certainly do not habitually pair. but I must here treat the subject with extreme brevity. is there for supposing in these cases that two individuals ever concur in reproduction? As it is impossible here to enter on details. In the first place. concur for the reproduction of their kind.4. which is all that concerns us. gives vigour and fertility to the offspring. This view was long ago doubtfully suggested by Sprengel. but that a cross with another individual is occasionally—perhaps at long intervals of time—indispensable. On The Intercrossing Of Individuals I must here introduce a short digression. I have collected so large a body of facts. We shall presently see its importance. though I have the materials prepared for an ample discussion. On the belief that this is a law of nature. notwithstanding that the plant's own anthers and pistil stand so near each other as almost to ensure selffertilisation. that with animals and plants a cross between different varieties. all insects and some other large groups of animals. or between individuals of the same variety but of another strain. in accordance with the almost universal belief of breeders. that is. such as the following. but in the case of hermaphrodites this is far from obvious. Every hybridizer knows how unfavourable exposure to wet is to the fertilisation of a flower. we can. showing. the fullest freedom for the entrance of pollen from another 101 .

as in the great papilionaceous or pea-family. that their fertility is greatly diminished if these visits be prevented. by insects. in Lobelia fulgens. and more recently Hildebrand and others have shown. to ensure fertilisation. and it is sufficient. though by placing pollen from one flower on the stigma of another. which seems to have a special contrivance for 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. to the great good of the plant. In very many other cases. but these almost invariably present beautiful and curious adaptations in relation to the visits of insects. before the stigma of that individual flower is ready to receive them. as I could show from the works of Sprengel and others. the former is so prepotent that it invariably and completely destroys. it is hardly possible to raise pure seedlings. just to touch with the same brush the anthers of one flower and then the stigma of another. and not to carry pollen from one to the other. Now. 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. which is visited by bees. Many flowers. I raise plenty of seedlings. as Sprengel.individual will explain the above state of exposure of the organs. if closely-allied forms or varieties are planted near each other. or the stigma is 102 . on the other hand. at least in my garden. as Kolreuter has shown to be the case with the barberry. Another species of Lobelia. Insects act like a camel-hair pencil. as well as from my own observations: for instance. it never sets a seed. for if a plant's own pollen and that from another species are placed on the same stigma. far from selffertilisation being favoured. the influence of the foreign pollen. so largely do they naturally cross. or slowly move one after the other towards it. the contrivance seems adapted solely to ensure self-fertilisation. it is scarcely possible for insects to fly from flower to flower. When the stamens of a flower suddenly spring towards the pistil. and as I can confirm. there are special contrivances which effectually prevent the stigma receiving pollen from its own flower. and in this very genus. though there is no special mechanical contrivance to prevent the stigma receiving pollen from the same flower. but it must not be supposed that bees would thus produce a multitude of hybrids between distinct species. and as this flower is never visited. either the anthers burst before the stigma is ready for fertilisation. as has been shown by Gartner. In numerous other cases. seeds freely in my garden. it is well known that. So necessary are the visits of bees to many papilionaceous flowers. yet. have their organs of fructification closely enclosed.

and Dr. and of some other plants. for I have found that plants carefully protected from insects produce the full number of pods. Asa Gray those of the United States. and the result was as I anticipated. be allowed to seed near each other. as I found. and that this is part of the general law of good being derived from the intercrossing of distinct individuals of the same species. When the sexes are separated. On the other hand. So it is with the reciprocally dimorphic and trimorphic plants previously alluded to. pollen must be regularly carried from flower to flower. then. and flowers on the same tree can be considered as distinct individuals only in a limited sense. although the male and female flowers may be produced on the same tree. and must habitually be crossed. Hooker 103 . mongrels: for instance. I raised 233 seedling cabbages from some plants of different varieties growing near each other. How strange are these facts! How strange that the pollen and stigmatic surface of the same flower. but that nature has largely provided against it by giving to trees a strong tendency to bear flowers with separated sexes. and at my request Dr. and some even of these were not perfectly true. 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. for a plant's own pollen is always prepotent over foreign pollen. and the pollen of each flower readily gets on its stigma without insect agency. When distinct SPECIES are crossed the case is reversed. I believe this objection to be valid. a large majority of the seedlings thus raised turn out. and at most only from flower to flower on the same tree.ready before the pollen of that flower is ready. and this will give a better chance of pollen being occasionally carried from tree to tree. radish. Dr. as if for the very purpose of self-fertilisation. it may be objected that pollen could seldom be carried from tree to tree. 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 of these only 78 were true to their kind. How. onion. but to this subject we shall return in a future chapter. so that these so-named dichogamous plants have in fact separated sexes. In the case of a large tree covered with innumerable flowers. Hooker tabulated the trees of New Zealand. though placed so close together. I find to be the case in this country. 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. That trees belonging to all orders have their sexes more often separated than other plants.

law of nature. but which I am unable here to give. I have made these few remarks on trees simply to call attention to the subject. some species of the same family and even of the same genus. As yet I have not found a single terrestrial animal which can fertilise itself. but here the currents of water offer an obvious means for an occasional cross. it appears that with animals and plants an occasional intercross between distinct individuals is a very general. to discover a single hermaphrodite animal with the organs of reproduction so perfectly enclosed that access from without. but I have been enabled. the same result would follow as if they bore flowers with separated sexes. if not universal. As in the case of flowers. Turning for a brief space to animals: various terrestrial species are hermaphrodites. though both are self-fertilising hermaphrodites. It must have struck most naturalists as a strange anomaly that. a case of great difficulty. Professor Huxley. and some unisexual. do sometimes cross. after consultation with one of the highest authorities.informs me that the rule does not hold good in Australia: but if most of the Australian trees are dichogamous. but these all pair. is intelligible on the view of an occasional cross being indispensable. though agreeing closely with each other in their whole organisation. the difference between them and unisexual species is. are hermaphrodites. under this point of view. as far as function is concerned. very small. I have as yet failed. in fact. both with animals and plants. Cirripedes long appeared to me to present. there are many self-fertilising hermaphrodites. and the occasional influence of a distinct individual. But if. by which an occasional cross could be effected with terrestrial animals without the concurrence of two individuals. all hermaphrodites do occasionally intercross. can be shown to be physically impossible. to prove that two individuals. From these several considerations and from the many special facts which I have collected. which offers so strong a contrast with terrestrial plants. namely. such as the land-mollusca and earth-worms. 104 . Of aquatic animals. by a fortunate chance. analogous to the action of insects and of the wind with plants. for owing to the nature of the fertilising element there are no means. This remarkable fact.

But when many men. and is. all the individuals varying in the right direction. if any one species does not become modified and improved in a corresponding degree with its competitors it will be exterminated. with some place in the natural polity not perfectly occupied. if the same species undergoes modification in different districts. and then. But if the area be large. A great amount of variability. With hermaphrodite organisms which cross only occasionally. Thus it will be under nature. But we shall see in the sixth chapter that intermediate varieties. though in different degrees. have a nearly common standard of perfection. under which term individual differences are always included. will evidently be favourable. nothing can be effected by natural selection. why should it prevail against natural selection? In the case of methodical selection. I believe. by giving a better chance within any given period for the appearance of profitable variations. its several districts will almost certainly present different conditions of life. will tend to be preserved. a highly important element of success. notwithstanding that there is no separation of selected individuals.5. Hence with animals of this nature. The tendency to reversion may often check or prevent the work. and all try to procure and breed from the best animals. Intercrossing will chiefly affect those animals which unite for each birth and wander much. for as all organic beings are striving to seize on each place in the economy of nature. without intending to alter the breed. for instance birds. his work will completely fail. but as this tendency has not prevented man from forming by selection numerous domestic races. for within a confined area. Though nature grants long periods of time for the work of natural selection. will in the long run generally be supplanted by one of the adjoining varieties. Circumstances Favourable For The Production Of New Forms Through Natural Selection This is an extremely intricate subject. varieties will generally be confined to separated countries. she does not grant an indefinite period. improvement surely but slowly follows from this unconscious process of selection. and this I find to be the case. and likewise with animals which unite for each 105 . will compensate for a lesser amount of variability in each individual. and which do not breed at a very quick rate. Unless favourable variations be inherited by some at least of the offspring. the newly formed varieties will intercross on the confines of each. A large number of individuals. a breeder selects for some definite object. inhabiting intermediate districts. and if the individuals be allowed freely to intercross.

Moritz Wagner has lately published an interesting essay on this subject. solely by natural selection preserving similar favourable variations. and might there maintain itself in a body and afterward spread. that they will have a better chance of surviving and propagating their kind. the young thus produced will gain so much in vigour and fertility over the offspring from long-continued self-fertilisation. or from the individuals of each variety preferring to pair together. In a confined or isolated area. but which wander little and can increase at a rapid rate. the organic and inorganic conditions of life will generally be almost uniform. or of the same variety. a new and improved variety might be quickly formed on any one spot. from breeding at slightly different seasons. but. so that the individuals of the new variety would chiefly cross together. Even with animals which unite for each birth. only through the principle of inheritance. Isolation also is an important element in the modification of species through natural selection. will be great. and thus in the long run the influence of crosses. will also be thus prevented. as already stated. and has shown that the service rendered by isolation in preventing crosses between newly-formed varieties is probably greater even than I 106 . we have reason to believe that occasional intercrosses take place with all animals and plants. Intercrossing plays a very important part in nature by keeping the individuals of the same species. nor conjugate. so that natural selection will tend to modify all the varying individuals of the same species in the same manner. Intercrossing with the inhabitants of the surrounding districts. uniformity of character can be retained by them under the same conditions of life. With respect to organic beings extremely low in the scale. If the conditions of life change and the form undergoes modification. and which do not propagate rapidly. even at rare intervals. which do not propagate sexually. from haunting different stations. and through natural selection which will destroy any individuals departing from the proper type. and which cannot possibly intercross. On this principle nurserymen always prefer saving seed from a large body of plants. Even if these take place only at long intervals of time. if not very large.birth. uniformity of character can be given to the modified offspring. 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. we must not assume that free intercrossing would always eliminate the effects of natural selection. true and uniform in character. as the chance of intercrossing is thus lessened. It will obviously thus act far more efficiently with those animals which unite for each birth.

as if all the forms of life were necessarily undergoing change through some innate law.—that is. and its importance in this respect is great. isolation will give time for a new variety to be improved at a slow rate. such as an oceanic island. The importance of isolation is likewise great in preventing. in relation to the constitution of each organism. especially for the production of species which shall prove capable of enduring for a long period. either from being surrounded by barriers. accumulated. Lapse of time is only so far important. and fixed. after any physical change in the conditions. that it gives a better chance of beneficial variations arising and of their being selected. Lastly. we ought to make the comparison within equal times. 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. If we turn to nature to test the truth of these remarks. such as of climate. 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.. on the whole I am inclined to believe that largeness of area is still more important. But from reasons already assigned I can by no means agree with this naturalist. and of spreading widely. and look at any small isolated area. not only will there be a better chance 107 . and this we are incapable of doing. have been produced there and nowhere else in the world. although the number of the species inhabiting it is small. that migration and isolation are necessary elements for the formation of new species. elevation of the land. has been most favourable for the production of new organic forms. It likewise tends to increase the direct action of the physical conditions of life. But we may thus deceive ourselves. etc. however. the total number of the inhabitants will be small.supposed. Although isolation is of great importance in the production of new species. If. yet of these species a very large proportion are endemic. and this may sometimes be of much importance. an isolated area be very small. either for or against natural selection. The mere lapse of time by itself does nothing. the immigration of better adapted organisms. or a large open area like a continent. and this will retard the production of new species through natural selection. Throughout a great and open area. Hence an oceanic island at first sight seems to have been highly favourable for the production of new species. for to ascertain whether a small isolated area. as we shall see in our chapter on Geographical Distribution. by decreasing the chances of favourable variations arising. or from having very peculiar physical conditions.

and there will have been less modification and less extermination. have existed in a broken condition. they have endured to the present day. These anomalous forms may be called living fossils. to a certain extent. we can understand how it is that the flora of Madeira. I conclude that. owing to former oscillations of level. also. will often. understand some facts which will be again alluded to in our chapter on Geographical Distribution. the fact of the productions of the smaller continent of Australia now yielding before those of the larger Europaeo-Asiatic area. resembles to a certain extent the extinct tertiary flora of Europe. Thus. will be those that will spread most widely. as the Ornithorhynchus and Lepidosiren. connect to a certain extent orders at present widely separated in the natural scale. the competition between fresh water productions will have been less severe than elsewhere. All fresh water basins. and what is more important. perhaps. and if some of these many species become modified and improved. taken together. which. and will thus come into competition with many other forms. though now continuous. new forms will have been more slowly produced. it is that continental productions have everywhere become so largely naturalised on islands. so that the good effects of isolation will generally. Consequently. have concurred.of favourable variations. remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world. arising from the large number of individuals of the same species there supported. They will thus play a more important part in the changing history of the organic world. which already have been victorious over many competitors. Moreover. In accordance with this view. as soon as it has been much improved. like fossils. also. make a small area compared with that of the sea or of the land. from 108 . will be able to spread over the open and continuous area. we can. yet that the course of modification will generally have been more rapid on large areas. the race for life will have been less severe. And it is in fresh water basins that we find seven genera of Ganoid fishes. and old forms more slowly exterminated. Finally. and will give rise to the greatest number of new varieties and species. for instance. Each new form. but the conditions of life are much more complex from the large number of already existing species. On a small island. great areas. according to Oswald Heer. that the new forms produced on large areas. or they will be exterminated. others will have to be improved in a corresponding degree. although small isolated areas have been in some respects highly favourable for the production of new species. Hence.

the most favoured or improved varieties will have been enabled to spread. there will again have been very severe competition. ready to be filled up by better adapted forms. by renewed elevation. and again there will have been a fair field for natural selection to improve still further the inhabitants. The result would often be greatly retarded by free intercrossing. Many will exclaim that these several causes are amply sufficient to neutralise the power of natural selection. I conclude that for terrestrial productions a large continental area. I do not believe so. as far as the extreme intricacy of the subject permits. competition. To sum up. and therefore less severe. but all this will take place very slowly. and this will create new places. The occurrence of such places will often depend on physical changes. and from having been exposed to less varied. and the relative proportional numbers of the various inhabitants of the reunited continent will again have been changed. and will have been subjected to severe competition. the islands were reconverted into a continental area. and time will have been allowed for the varieties in each to become well modified and perfected. As some few of the old inhabitants become modified the mutual relations of others will often be disturbed. which has undergone many oscillations of level. 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. the circumstances favourable and unfavourable for the production of new species through natural selection. Although all the individuals of the same species differ in some slight degree from each other. That natural selection generally act with extreme slowness I fully admit. fitted to endure for a long time and to spread widely. While the area existed as a continent the inhabitants will have been numerous in individuals and kinds. which generally take place very slowly. and thus to produce new species. there will have been much extinction of the less improved forms. and on the immigration of better adapted forms being prevented. will have been the most favourable for the production of many new forms of life.having inhabited a confined area. it would often be long before differences of the right nature in various parts of the organisation might occur. so that new places in the polity of each island will have had to be filled up by the modification of the old inhabitants. 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. When. But I do believe that natural selection will generally act 109 .

and only on a few of the inhabitants of the same region.very slowly. 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. to the beauty and complexity of the coadaptations between all organic beings. one with another and with their physical conditions of life. only at long intervals of time. which may have been effected in the long course of time through nature's power of selection. 110 . that is by the survival of the fittest. I further believe that these slow. I can see no limit to the amount of change.

—which. constitution and habits. Owing to the high geometrical rate of increase of all organic beings. We have seen that the species which are most numerous in individuals have the best chance of producing favourable variations within any given period. unless we admit that specific forms can go on indefinitely increasing in number. and species of the same genus or related genera. will the less favoured decrease and become rare.6. but it must here be alluded to from being intimately connected with natural selection. will generally press hardest on its nearest kindred. But we may go further than this. Extinction Caused By Natural Selection This subject will be more fully discussed in our chapter on Geology. showing that it is the common and diffused or dominant species which offer the greatest number of recorded varieties.—varieties of the same species. and tend to exterminate them. each new variety or species. will naturally suffer most. which consequently endure. in the facts stated in the second chapter. that as new species in the course of time are formed through natural selection. Consequently. rare species will be less quickly modified or improved within any given period. and it follows from this. from having nearly the same structure. others will become rarer and rarer. We can see that any form which is represented by few individuals will run a good chance of utter extinction. or from a temporary increase in the number of its enemies. during great fluctuations in the nature or the seasons. and we shall presently attempt to show why it is that the number of species throughout the world has not become immeasurably great. generally. That the number of specific forms has not indefinitely increased. and finally extinct. generally come into the severest competition with each other. that as the favoured forms increase in number. Hence. Natural selection acts solely through the preservation of variations in some way advantageous. From these several considerations I think it inevitably follows. each area is already fully stocked with inhabitants. geology plainly tells us. The forms which stand in closest competition with those undergoing modification and improvement. We have evidence of this. Rarity. many old forms must become extinct. We see the same process of extermination 111 . so. as geology tells us. during the progress of its formation. for as new forms are produced. And we have seen in the chapter on the Struggle for Existence that it is the most closely-allied forms. they will consequently be beaten in the race for life by the modified and improved descendants of the commoner species. is the precursor to extinction.

take the place of older and inferior kinds. through the selection of improved forms by man." 112 . and that these "were swept away by the short-horns" (I quote the words of an agricultural writer) "as if by some murderous pestilence. Many curious instances could be given showing how quickly new breeds of cattle. sheep and other animals.among our domesticated productions. and varieties of flowers. it is historically known that the ancient black cattle were displaced by the longhorns. In Yorkshire.

but. and explains. incipient species. present slight and ill-defined differences. I have sought light on this head from our domestic productions. could never have been effected by the mere chance accumulation of similar variations during many successive generations. might cause one variety to differ in some character from its parents. as I have called them. which I have designated by this term. the men of one nation or district required swifter horses. struck by a pigeon having a slightly shorter beak. and would be noted as forming two sub-breeds. In practice.. Divergence Of Character The principle. 113 . Ultimately after the lapse of centuries. for instance. How. or are. does the lesser difference between varieties become augmented into the greater difference between species? That this does habitually happen. another fancier is struck by a pigeon having a rather longer beak. As has always been my practice. race and cart horses. and on the acknowledged principle that "fanciers do not and will not admire a medium standard. varieties. or with shorter and shorter beaks. Mere chance." 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. the differences would become greater. a fancier is. while those of another required stronger and bulkier horses. these sub-breeds would become converted into two well-established and distinct breeds.7. The early differences would be very slight. It will be admitted that the production of races so different as short-horn and Hereford cattle. but this alone would never account for so habitual and large a degree of difference as that between the species of the same genus. the several breeds of pigeons. from the continued selection of swifter horses in the one case. but like extremes. and the offspring of this variety again to differ from its parent in the very same character and in a greater degree. varieties are species in the process of formation. several important facts. we may suppose that at an early period of history. etc. as we may call it. in the course of time. the supposed prototypes and parents of future well-marked species. as I believe. is of high importance. we must infer from most of the innumerable species throughout nature presenting well-marked differences. Again. then. and of stronger ones in the other. We shall here find something analogous. even strongly-marked ones. Nevertheless according to my view. In the first place. 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. whereas varieties.

being enabled to feed on new kinds of prey. The more diversified in habits and structure the descendants of our carnivorous animals become. a greater number of individual plants of this species. either dead or alive. It has been experimentally proved. then. What applies to one animal will apply throughout all time to all animals—that is. some inhabiting new stations. that if a plot of ground be sown with one species of grass. would not have been used for breeding. and the breeds to diverge in character. of which the number that can be supported in any country has long ago arrived at its full average. we see in man's productions the action of what may be called the principle of divergence. We can clearly discern this in the case of animals with simple habits. it may be asked. and will thus have tended to disappear. at first barely appreciable. constitution. if any one species of grass were to go on varying. would succeed in living on the same piece of ground. and some perhaps becoming less carnivorous. though in a very slight degree. if they vary—for otherwise natural selection can effect nothing. including its modified descendants. for instance. and a similar plot be sown with several distinct genera of grasses. as do the distinct species and genera of grasses. a greater number of plants and a greater weight of dry herbage can be raised in the latter than in the former case. being neither very swift nor very strong.As the differences became greater. steadily to increase. both from each other and from their common parent. Here. 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). and is thus striving. causing differences. climbing trees. frequenting water. Take the case of a carnivorous quadruped. from the simple circumstance that the more diversified the descendants from any one species become in structure. So it will be with plants. And we know that each species and each variety of grass is annually sowing almost countless seeds. 114 . and habits. 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. But how. and the varieties were continually selected which differed from each other in the same manner. Hence. by so much will they be better enabled to seize on many and widely diversified places in the polity of nature. 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. If its natural power of increase be allowed to act. the inferior animals with intermediate characters. the more places they will be enabled to occupy. and so be enabled to increase in numbers.

For instance. Consequently. have been expected that naturalised plants would have belonged to a few groups more especially adapted to certain stations in their new homes. to the utmost to increase in number. three feet by four in size. take the rank of species. belong to what we call different genera and orders. for these are commonly looked at as specially created and adapted for their own country. perhaps. determine that the inhabitants. is seen under many natural circumstances. and varieties. especially if freely open to immigration. So it is with the plants and insects on small and uniform islets: also in small ponds of fresh water. as a general rule. 115 . It might also. when rendered very distinct from each other. that where they come into the closest competition.as it may be said. de Candolle has well remarked. and may be said to be striving to the utmost to live there. that floras gain by naturalisation. Most of the animals and plants which live close round any small piece of ground." 260 naturalised plants are enumerated. could live on it (supposing its nature not to be in any way peculiar). Asa Gray's "Manual of the Flora of the Northern United States. the most distinct varieties of any one species of grass would have the best chance of succeeding and of increasing in numbers. which thus jostle each other most closely. and Alph. In an extremely small area. 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. and these belonged to eighteen genera and to eight orders. shall. we always find great diversity in its inhabitants. the advantages of diversification of structure. and thus of supplanting the less distinct varieties. in the course of many thousand generations. The same principle is seen in the naturalisation of plants through man's agency in foreign lands. I found that a piece of turf. far more in new genera than in new species. but. The truth of the principle that the greatest amount of life can be supported by great diversification of structure. with the accompanying differences of habit and constitution. But the case is very different. and where the contest between individual and individual must be very severe. which shows how much these plants differed from each other. which had been exposed for many years to exactly the same conditions. 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. To give a single instance: in the last edition of Dr. in his great and admirable work. it is seen. proportionally with the number of the native genera and species. supported twenty species of plants.

and feebly representing. to a large extent. 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. The advantage of diversification of structure in the inhabitants of the same region is. our carnivorous. could hardly compete with a set more perfectly diversified in structure. It may be doubted. 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. moreover. as Mr. so will a greater number of individuals be capable of there supporting themselves. Waterhouse and others have remarked. A set of animals. ruminant. amounting to new generic differences. the more widely and perfectly the animals and plants are diversified for different habits of life. We thus see that these naturalised plants are of a highly diversified nature. from the indigenes. no less than 100 genera are not there indigenous. which are divided into groups differing but little from each other. In the Australian mammals. and rodent mammals. and thus a large proportional addition is made to the genera now living in the United States. No physiologist doubts that a stomach by being adapted to digest vegetable matter alone. 116 . So in the general economy of any land. whether the Australian marsupials. or flesh alone. By considering the nature of the plants or animals which have in any country struggled successfully with the indigenes. and have there become naturalised. draws most nutriment from these substances. would be profitable to them. for out of the 162 naturalised genera. we see the process of diversification in an early and incomplete stage of development. in fact. for instance. with their organisation but little diversified.and these belong to 162 genera. could successfully compete with these well-developed orders. They differ. and we may at least infer that diversification of structure.

And here the importance of the principle of benefit derived from divergence of character comes in. The accompanying diagram will aid us in understanding this rather perplexing subject. may represent its varying offspring. The variations are supposed to be extremely slight. also. which are the commonest and most widely-diffused. and are thus enabled to encroach on places occupied by other beings. widely-diffused. After a thousand generations. When a dotted line reaches one of the horizontal lines. on an average more species vary in large genera than in small genera. and as is represented in the diagram by the letters standing at unequal distances. vary more than do the rare and restricted species. The intervals between the horizontal lines in the diagram. belonging to a genus large in its own country. nor are they all supposed to endure for equal periods. and is there marked by a small numbered letter. I have said a large genus. but often after long intervals of time. 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. which has been much compressed. tends to act. as is so generally the case in nature. they are not supposed all to appear simultaneously. seen that the species. may represent each a thousand or more generations.8. such as would be thought worthy of record in a systematic work. and the varying species of the large genera present a greater number of varieties. and varying species. The Probable Effects Of The Action Of Natural Selection Through Divergence Of Character And Extinction. species (A) is supposed to have produced two fairly well-marked 117 . Only those variations which are in some way profitable will be preserved or naturally selected. 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. a sufficient amount of variation is supposed to have been accumulated to form it into a fairly well-marked variety. combined with the principles of natural selection and of extinction. Let A to L represent the species of a genus large in its own country. we may assume that the modified descendants of any one species will succeed so much the better as they become more diversified in structure. these species are supposed to resemble each other in unequal degrees. On The Descendants Of A Common Ancestor After the foregoing discussion. Let (A) be a common. because as we saw in the second chapter. but of the most diversified nature. We have.

and this will depend on infinitely complex relations. will tend to inherit those advantages which made their parent (A) more numerous than most of the other inhabitants of the same country. Thus the varieties or modified descendants of the common parent (A). after each thousand generations. consequently they will likewise tend to vary. Moreover. Variety m1 is supposed to have produced two varieties. differing from each other. and the tendency to variability is in itself hereditary. these two varieties. some of the varieties. and then again undergoes modification. and some failing to produce any. owing to the principle of divergence. namely m2 and s2.varieties. variety a1 is supposed in the diagram to have produced variety a2. and more considerably from their common parent (A). In the diagram the process is represented up to the ten-thousandth generation. and commonly in nearly the same manner as did their parents. being only slightly modified forms. These two varieties will generally still be exposed to the same conditions which made their parents variable. will generally go on increasing in number and diverging in character. Nor do I suppose that the most divergent varieties are invariably preserved: a medium form may often long endure. they will also partake of those more general advantages which made the genus to which the parent-species belonged. But as a general rule. In our diagram the 118 . and the more their modified progeny will increase. If. the more places they will be enabled to seize on. and under a condensed and simplified form up to the fourteen-thousandth generation. it is far more probable that each form remains for long periods unaltered. a large genus in its own country. namely a1 and m1. though in itself made somewhat irregular. But I must here remark that I do not suppose that the process ever goes on so regularly as is represented in the diagram. differ more from (A) than did variety a1. which will. And all these circumstances are favourable to the production of new varieties. for natural selection will always act according to the nature of the places which are either unoccupied or not perfectly occupied by other beings. these two varieties be variable. but in a more and more modified condition. the most divergent of their variations will generally be preserved during the next thousand generations. producing only a single variety. some producing two or three varieties. then. nor that it goes on continuously. the more diversified in structure the descendants from any one species can be rendered. We may continue the process by similar steps for any length of time. And after this interval. and may or may not produce more than one modified descendant.

after intervals long enough to allow the accumulation of a considerable amount of divergent variation. and so destroy. will have come to differ largely. but we have only to suppose the steps in the process of modification to be more numerous or greater in amount. The modified offspring from the later and more highly improved branches in the lines of descent. excepting that from a1 to a10. if all the lines proceeding from (A) were removed. although the amount of divergent modification may have been augmented. By continuing the same process for a greater number of generations (as shown in the diagram in a condensed and simplified manner). This case would be represented in the diagram. without either having given off any fresh branches or races. often take the place of. 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). from each other and from their common parent. If we suppose the amount of change between each horizontal line in our diagram to be excessively small. 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 the same way the English racehorse and English pointer have apparently both gone on slowly diverging in character from their original stocks. all descended from (A). it is probable. and might have been inserted anywhere. species (A) is supposed to have produced three forms. but perhaps unequally. As all the modified descendants from a common and widely-diffused species. and the number of modified descendants will not be increased. from having diverged in character during the successive generations. we get eight species. After ten thousand generations. marked by the letters between a14 and m14. In some cases no doubt the process of modification will be confined to a single line of descent. which. and m10. 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. But these breaks are imaginary. 119 . a10. belonging to a large genus. f10. will tend to partake of the same advantages which made their parent successful in life. these three forms may still be only well-marked varieties. will. species are multiplied and genera are formed. Thus.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. as I believe.

which will be conquered by later and improved lines. The other nine species (marked by capital letters) of our original genus. 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. and have given rise to new varieties and species. after ten thousand generations. If. 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). 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. and the nearly extreme species (I). In the diagram I have assumed that a second species (I) has produced. or become quickly adapted to some quite new station. another of our principles. In any genus. and species (I) will be replaced by six (n14 to z14) new species. our diagram be assumed to represent a considerable amount of modification. will have played an important part.In a large genus it is probable that more than one species would vary. marked by the letters n14 to z14. will generally tend to produce the greatest number of modified descendants. But during the process of modification. will generally tend to become extinct. If. constitution and structure. as those which have largely varied. 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. being replaced by eight new species (a14 to m14). and this is shown in the diagram by the dotted lines unequally prolonged upwards. are supposed to have been produced. then. represented in the diagram. however. either two well-marked varieties (w10 and z10) or two species. the species which are already very different in character from each other. species (A) and all the earlier varieties will have become extinct. namely that of extinction. After fourteen thousand generations. So it probably will be with many whole collateral lines of descent. in which offspring and progenitor do not come into competition. Hence all the intermediate forms between the earlier and later states. six new species. that is between the less and more improved states of a the same species. as well as the original parent-species itself. the modified offspring of a species get into some distinct country. by analogous steps. may for long but unequal periods continue to transmit unaltered descendants. 120 . according to the amount of change supposed to be represented between the horizontal lines. both may continue to exist.

and may constitute a sub-genus or a distinct genus. so that they must originally have had some advantage over most of the other species of the genus. differ considerably from the eight descendants from (A). are supposed to have gone on diverging in different directions. p14. not only their parents (A) and (I). and D than to the other species. L. These two species (A and I). moreover. Their modified descendants. but likewise some of the original species which were most nearly related to their parents. species (A) being more nearly related to B. will be nearly related from having recently branched off from a10. will be nearly related one to the other. also (and this is a very important consideration). has transmitted descendants to this late stage of descent. and thus exterminated. than to the others. b14 and f14. It seems. therefore. moreover. o14. and m14. the two groups. but. But as the original species (I) differed largely from (A). will now be fifteen in number. K. extremely probable that they will have taken the places of. C. descended from the original eleven species. q14. were also supposed to be very common and widely diffused species. from having diverged at the first commencement of the process of modification. which connected the original species 121 . The intermediate species. 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. will be allied to each other in a widely different manner. fourteen in number at the fourteen-thousandth generation. e14. as is so generally the case in nature. will be widely different from the other five species. The original species of our genus were supposed to resemble each other in unequal degrees. and species (I) more to G. The new species in our diagram.But we may go further than this. Of the eight descendants from (A) the three marked a14. so as to have become adapted to many related places in the natural economy of their country. 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. will be in some degree distinct from the three first-named species. the six descendants from (I) will. owing to inheritance alone. The new species. Hence very few of the original species will have transmitted offspring to the fourteen-thousandth generation. from having diverged at an earlier period from a5. The six descendants from (I) will form two sub-genera or genera. and lastly. standing nearly at the extreme end of the original genus. H. 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. Owing to the divergent tendency of natural selection.

as now explained. yet are often. now supposed to be extinct and unknown. It is worth while to reflect for a moment on the character of the new species F14. In the diagram each horizontal line has hitherto been supposed to represent a thousand generations. to the formation of genera alone. with those now living. but each may represent a million or more generations. will form three very distinct 122 . And the two or more parent-species are supposed to be descended from some one species of an earlier genus. and have left no descendants. converging in sub-branches downwards towards a single point. the forms marked a14 to p14. and I think we shall then see that the diagram throws light on the affinities of extinct beings. families. Being descended from a form that stood between the parent-species (A) and (I). If. or genera. but to have retained the form of (F). this point represents a species. Hence the six new species descended from (I). have to refer again to this subject. extinct. those marked b14 and f14. and every naturalist will be able to call such cases before his mind. as I believe.(A) and (I). In our diagram this is indicated by the broken lines beneath the capital letters. it will be in some degree intermediate in character between the two groups descended from these two species. and those marked o14 to m14. But as these two groups have gone on diverging in character from the type of their parents. for the extinct species lived at various remote epochs when the branching lines of descent had diverged less. or even as distinct sub-families. that two or more genera are produced by descent with modification. either unaltered or altered only in a slight degree. which is supposed not to have diverged much in character. the new species (F14) will not be directly intermediate between them. in some degree. will have to be ranked as very distinct genera. except (F). it may also represent a section of the successive strata of the earth's crust including extinct remains. in the diagram. Thus it is. from two or more species of the same genus. but rather between types of the two groups. we suppose the amount of change represented by each successive group of diverging dotted lines to be great. the supposed progenitor of our several new sub-genera and genera. though generally belonging to the same orders. and we can understand this fact. have all become. intermediate in character between existing groups. when we come to our chapter on geology. I see no reason to limit the process of modification. We shall. which. and the eight descendants from (A). In this case its affinities to the other fourteen new species will be of a curious and circuitous nature.

and leave no modified descendants. Within the same large group. as all the descendants of the same species form a class. formerly most extensively developed. and thus lessen its chance of further variation and improvement. But which groups will ultimately prevail. continue to increase. a multitude of smaller groups will become utterly extinct. are descended from two species of the original genus. or orders. or orders. at remote geological periods. for we know that many groups. reduce its number. no man can predict. One large group will slowly conquer another large group. it will chiefly act on those which already have some advantage. differing widely from the descendants of (A). yet. the later and more highly perfected sub-groups. the struggle for the production of new and modified descendants will mainly lie between the larger groups. This. Although few of the most ancient species have left modified descendants. extremely few of the more ancient species have transmitted descendants to the present day. the earth may 123 . but I may add that as. have now become extinct. will. from branching out and seizing on many new places in the polity of nature. according to this view. We shall also have two very distinct genera descended from (I). Looking to the future. and these are supposed to be descended from some still more ancient and unknown form. I shall have to return to this subject in the chapter on classification. and which are least broken up. Hence. will constantly tend to supplant and destroy the earlier and less improved sub-groups. we can predict that the groups of organic beings which are now large and triumphant. of the species living at any one period. and consequently that. indeed. and. And the two new families. We have seen that in each country it is the species belonging to the larger genera which oftenest present varieties or incipient species. and the largeness of any group shows that its species have inherited from a common ancestor some advantage in common.genera. which are all trying to increase in number. according to the amount of divergent modification supposed to be represented in the diagram. which have as yet suffered least extinction. that is. might have been expected. Small and broken groups and sub-groups will finally disappear. extremely few will transmit descendants to a remote futurity. These two groups of genera will thus form two distinct families. we may predict that. for as natural selection acts through one form having some advantage over other forms in the struggle for existence. Looking still more remotely to the future. owing to the continued and steady increase of the larger groups. we can understand how it is that there exist so few classes in each main division of the animal and vegetable kingdoms. for a long period.

as at the present day.have been almost as well peopled with species of many genera. families. 124 . orders and classes.

stamens and pistils. which are beneficial under the organic and inorganic conditions to which each creature is exposed at all periods of life. or. inasmuch as in this state they perform their functions better. namely. for instance. Among the vertebrata the degree of intellect and an approach in structure to man clearly come into play. look at the plants which have their several organs much modified and reduced in number as the highest. whereas other botanists. But we shall see how obscure this subject is if we look. 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). natural selection clearly leads towards this standard: for all physiologists admit that the specialisation of organs. as I should be inclined to add. the amount of differentiation of the parts of the same organic being. petals. This improvement inevitably leads to the gradual advancement of the organisation of the greater number of living beings throughout the world. while other naturalists rank the common bony or teleostean fishes as the highest. and differ most from the other vertebrate classes. for naturalists have not defined to each other's satisfaction what is meant by an advance in organisation. so that the mature animal cannot be called higher than its larva. among which some naturalists rank those as highest which. Von Baer's standard seems the most widely applicable and the best. If we take as the standard of high organisation. inasmuch as they are most strictly fish. to fishes. like the sharks. and their specialisation for different functions. 125 . as sepals. On The Degree To Which Organisation Tends To Advance Natural selection acts exclusively by the preservation and accumulation of variations. in the adult state. and here some botanists rank those plants as highest which have every organ. The ultimate result is that each creature tends to become more and more improved in relation to its conditions. but there are cases. fully developed in each flower. the completeness of the division of physiological labour.9. is an advantage to each being. approach nearest to amphibians. probably with more truth. as Milne Edwards would express it. We see still more plainly the obscurity of the subject by turning to plants. among which the standard of intellect is of course quite excluded.like. 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. But here we enter on a very intricate subject. as with certain parasitic crustaceans. in which several parts of the structure become less perfect.

have remained for an enormous period in nearly their present state. how is it that throughout the world a multitude of the lowest forms still exist.and hence the accumulation of variations tending towards specialisation is within the scope of natural selection. to 126 . for natural selection. these forms would be left. 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. to the co-existence of man and the ornithorhynchus—among fishes. And geology tells us that some of the lowest forms. by natural selection. Science has not as yet proved the truth of this belief. 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. And it may be asked what advantage. for instance. 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. if we look to the different grades of organisation within the same great group. 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. as the infusoria and rhizopods. and might remain for indefinite ages in their present lowly condition. in the vertebrata. to be highly organised. On the other hand. If it were no advantage. to the co-existence of mammals and fish—among mammalia. we can see. would it be to an infusorian animalcule—to an intestinal worm—or even to an earth-worm. Nearly the same remarks are applicable. who believed in an innate and inevitable tendency towards perfection in all organic beings. does not necessarily include progressive development—it only takes advantage of such variations as arise and are beneficial to each creature under its complex relations of life. whatever the future may reveal. as far as we can see. for every naturalist who has dissected some of the beings now ranked as very low in the scale. unimproved or but little improved. 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. 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. must have been struck with their really wondrous and beautiful organisation. On our theory the continued existence of lowly organisms offers no difficulty. or the survival of the fittest. But it may be objected that if all organic beings thus tend to rise in the scale.

and where their scanty numbers have retarded the chance of favourable variations arising. Herbert Spencer would probably answer that. from inhabiting confined or peculiar stations. Although organisation. an anomalous annelid. probably. so that warm-blooded mammals when inhabiting the water lie under a disadvantage in having to come continually to the surface to breathe.existence of the shark and the lancelet (Amphioxus). to the highest grade would not lead to their taking the place of fishes. it has been asked. In some cases variations or individual differences of a favourable nature may never have arisen for natural selection to act on and accumulate. 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. co-exist in South America in the same region with numerous monkeys. lowly organised forms appear to have been preserved to the present day. and more liable to be put out of order and injured. Physiologists believe that the brain must be bathed by warm blood to be highly active. on the whole. from various causes. which latter fish in the extreme simplicity of its structure approaches the invertebrate classes. as I hear from Fritz Muller. presented the simplest structure. could the first step in the advancement or differentiation of parts have arisen? Mr. may have advanced and be still advancing throughout the world. In some cases. as we may believe. as soon as simple unicellular organisms came by growth or division to be compounded of 127 . does not at all necessarily lead to the extinction of those groups with which they do not enter into close competition. has time sufficed for the utmost possible amount of development.the co. I believe that many lowly organised forms now exist throughout the world. yet the scale will always present many degrees of perfection. for the lancelet. and this requires aerial respiration. the advancement of the whole class of mammals. In no case. The three lowest orders of mammals. as being of a more delicate nature. or of certain members of each class. has as sole companion and competitor on the barren sandy shore of South Brazil. namely. as we shall hereafter see. and rodents. Looking to the first dawn of life. where they have been subjected to less severe competition. for the high advancement of certain whole classes. when all organic beings. Finally. how. With fishes. edentata. marsupials. or of certain members in this class. and probably interfere little with each other. But mammals and fish hardly come into competition with each other. members of the shark family would not tend to supplant the lancelet. In some few cases there has been what we must call retrogression or organisation.

however. or became attached to any supporting surface. consequently. if we make due allowance for our profound ignorance on the mutual relations of the inhabitants of the world at the present time.several cells. and still more so during past ages. But. as I remarked towards the close of the introduction. no natural selection. until many forms had been produced: variations in a single species inhabiting an isolated station might be beneficial. 128 . 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. and thus the whole mass of individuals might be modified. It is. or two distinct forms might arise. speculation on the subject is almost useless. But as we have no facts to guide us. no one ought to feel surprise at much remaining as yet unexplained on the origin of species. an error to suppose that there would be no struggle for existence.

. it is conceivable that these might approach each other so closely that they would have all to be classed under the same genus. and that convergence. but with organic beings we should bear in mind that the form of each depends on an infinitude of complex relations. had both produced a large number of new and divergent forms. and it is not surprising that dissimilar substances should sometimes assume the same form. If two species belonging to two distinct though allied genera. moisture. recurring in widely separated geological formations. namely on the variations which have arisen. and therefore no limit to the 129 . which had originally differed in a marked manner. etc. The shape of a crystal is determined solely by the molecular forces.10.C. 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. and as the number of species in any country goes on increasing. If this had occurred. and thus the descendants of two distinct genera would converge into one. It is incredible that the descendants of two organisms. has likewise played a part. all of which have had their forms determined through equally complex relations. Consequently there seems at first no limit to the amount of profitable diversification of structure. Mr. As far as mere inorganic conditions are concerned. Watson has also objected that the continued action of natural selection. and in a still higher degree on the surrounding organisms with which each being has come into competition—and lastly. as it may be called. we should meet with the same form. the organic conditions of life must become more and more complex. on inheritance (in itself a fluctuating element) from innumerable progenitors. however. together with divergence of character. he apparently believes). Convergence Of Character Mr. should ever afterwards converge so closely as to lead to a near approach to identity throughout their whole organisation. independently of genetic connection. and this depends on the surrounding physical conditions. these being due to causes far too intricate to be followed out—on the nature of the variations which have been preserved or selected. 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. H. Watson thinks that I have overrated the importance of divergence of character (in which. would tend to make an indefinite number of specific forms. it seems probable that a sufficient number of species would soon become adapted to all considerable diversities of heat.

each or nearly each species will be represented by few individuals. and thus check the inordinate increase of specific forms throughout the world. 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. But geology shows us. on the principal often explained. and the first severe winter or very dry summer would exterminate thousands on thousands of species. and each species will become rare if the number of species in any country becomes indefinitely increased. 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. etc. where. close interbreeding will help to exterminate it. depending so largely as it does on physical conditions. de Candolle has shown that those species which spread widely tend generally to spread VERY widely. 130 .number of species which might be produced. Rare species. the number of mammals has not greatly or at all increased. many European plants have become naturalised. and that from the middle part of this same period. will. if an area be inhabited by very many species. apparently. Lastly. there are many invaders from different quarters of the globe. of red deer in Scotland and of bears in Norway. consequently they will tend to supplant and exterminate several species in several areas. a dominant species. Dr. When any species becomes very rare. but conjointly they must limit in each country the tendency to an indefinite augmentation of specific forms. consequently. whereas the production of new species must always be slow. the endemic Australian species have been greatly reduced in number. Imagine the extreme case of as many species as individuals in England. and such species will be liable to extermination from accidental fluctuations in the nature of the seasons or in the number of their enemies. that from an early part of the tertiary period the number of species of shells. present within a given period few favourable variations. The process of extermination in such cases would be rapid. authors have thought that this comes into play in accounting for the deterioration of the aurochs in Lithuania. which has already beaten many competitors in its own home. the process of giving birth to new specific forms would thus be retarded. which support such an astonishing number of species. Hooker has recently shown that in the southeast corner of Australia. and this I am inclined to think is the most important element. Alph. How much weight to attribute to these several considerations I will not pretend to say. therefore. will tend to spread and supplant many others.

in the same manner as so many variations have occurred useful to man. and how largely extinction has acted in the world's history. Therefore. Sexual selection will also give characters useful to the males alone in their struggles or rivalry with other males. by so much the more can a large number be supported on the area. for the more organic beings diverge in structure. in most cases. habits and constitution. if there be. and during the incessant 131 . Summary Of Chapter If under changing conditions of life organic beings present individual differences in almost every part of their structure. and these characters will be transmitted to one sex or to both sexes. constitution. and from the strong principle of inheritance. But if variations useful to any organic being ever do occur. these will tend to produce offspring similarly characterised. considering the infinite complexity of the relations of all organic beings to each other and to their conditions of life. But we have already seen how it entails extinction. or the survival of the fittest. and habits. Natural selection. and consequently. must be judged by the general tenour and balance of evidence given in the following chapters. of which we see proof by looking to the inhabitants of any small spot. Whether natural selection has really thus acted in adapting the various forms of life to their several conditions and stations. 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. and to the productions naturalised in foreign lands.11. owing to their geometrical rate of increase. and this certainly cannot be disputed. to be advantageous to them. This principle of preservation. then. according to the form of inheritance which prevails. geology plainly declares. on the principle of qualities being inherited at corresponding ages. Nevertheless. can modify the egg. or young as easily as the adult. causing an infinite diversity in structure. and this cannot be disputed. during the modification of the descendants of any one species. to what must be regarded as an advance in organisation. a severe struggle for life at some age. it would be a most extraordinary fact if no variations had ever occurred useful to each being's own welfare. also. I have called natural selection. assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life. leads to divergence of character. Natural selection. seed. season or year. It leads to the improvement of each creature in relation to its organic and inorganic conditions of life. low and simple forms will long endure if well fitted for their simple conditions of life.

the better will be their chance of success in the battle for life. forming sub-families. and these tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. At each period of growth all the growing twigs have tried to branch out on all sides. orders. budding twigs. forming sections and sub-genera. and genera related in different degrees. which vary most. We have seen that it is the common. in the manner which we everywhere behold—namely. entailing extinction and divergence of character. Thus the small differences distinguishing varieties of the same species. and to overtop and kill the surrounding twigs and branches. The limbs divided into great branches. subordinate to groups. the widely diffused. or even of distinct genera. and this connexion of the former and present buds by ramifying 132 . and classes. I believe this simile largely speaks the truth. the more diversified the descendants become. steadily tend to increase. till they equal the greater differences between species of the same genus. varieties of the same species most closely related. but seem clustered round points. sub-classes. and widely ranging species. and these round other points. no explanation would have been possible of this kind of classification. as we have seen illustrated in the diagram. when the tree was young. as has just been remarked. and those produced during former years may represent the long succession of extinct species. If species had been independently created. The green and budding twigs may represent existing species. in the same manner as species and groups of species have at all times overmastered other species in the great battle for life. species of distinct genera much less closely related. and so on in almost endless cycles. and the generally well defined distinctions between the innumerable organic beings in each class throughout the world. leads to divergence of character and to much extinction of the less improved and intermediate forms of life. On these principles. The several subordinate groups in any class cannot be ranked in a single file. The affinities of all the beings of the same class have sometimes been represented by a great tree. belonging to the larger genera within each class. families. may be explained. It is a truly wonderful fact—the wonder of which we are apt to overlook from familiarity—that all animals and all plants throughout all time and space should be related to each other in groups. and these into lesser and lesser branches. the nature of the affinities. but it is explained through inheritance and the complex action of natural selection. were themselves once. species of the same genus less closely and unequally related.struggle of all species to increase in numbers. Natural selection.

and these fallen branches of various sizes may represent those whole orders. straggling branch springing from a fork low down in a tree. which fills with its dead and broken branches the crust of the earth. many a limb and branch has decayed and dropped off. so we occasionally see an animal like the Ornithorhynchus or Lepidosiren. so by generation I believe it has been with the great Tree of Life. Of the many twigs which flourished when the tree was a mere bush. branch out and overtop on all sides many a feebler branch. and genera which have now no living representatives. and these. families. very few have left living and modified descendants. As we here and there see a thin. which in some small degree connects by its affinities two large branches of life. 133 . and which are known to us only in a fossil state. now grown into great branches. and covers the surface with its ever-branching and beautiful ramifications. yet survive and bear the other branches. and which has apparently been saved from fatal competition by having inhabited a protected station. so with the species which lived during long-past geological periods. if vigorous. From the first growth of the tree.branches may well represent the classification of all extinct and living species in groups subordinate to groups. As buds give rise by growth to fresh buds. only two or three. and which by some chance has been favoured and is still alive on its summit.

organs of flight and of vision — Acclimatisation — Correlated variation — Compensation and economy of growth — False correlations — Multiple. combined with natural selection.Chapter 5 Laws Of Variation Effects of changed conditions — Use and disuse. 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. rudimentary. 134 .

1. This. the individuals become modified in the same way. In the following cases the conditions seem to have produced some slight definite effect: E. 135 . cannot be attributed simply to such action. Some authors believe it to be as much the function of the reproductive system to produce individual differences. of course is a wholly incorrect expression. and the greater variability of species having wide ranges than of those with restricted ranges.. when subjected to certain conditions. The direct action of changed conditions leads to definite or indefinite results. or slight deviations of structure. In the latter case the organisation seems to become plastic. but it serves to acknowledge plainly our ignorance of the cause of each particular variation. or nearly all. directly on the whole organisation or on certain parts alone. In the former case the nature of the organism is such that it yields readily. are more brightly coloured than those of the same species from further north or from a greater depth. and all. the nature of the organism. and when living in shallow water. than when living near the coast or on islands. and we have much fluctuating variability. such as of climate. It is very difficult to decide how far changed conditions. In all cases there are two factors. Gould believes that birds of the same species are more brightly coloured under a clear atmosphere. and Wollaston is convinced that residence near the sea affects the colours of insects. have acted in a definite manner. which is much the most important of the two. food. which we see throughout nature between various organic beings. I have hitherto sometimes spoken as if the variations—so common and multiform with organic beings under domestication. In the first chapter I attempted to show that changed conditions act in two ways. But we may safely conclude that the innumerable complex co-adaptations of structure. and the nature of the conditions. 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. Forbes asserts that shells at their southern limit. as to make the child like its parents. MoquinTandon gives a list of plants which. But the fact of variations and monstrosities occurring much more frequently under domestication than under nature. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence. and indirectly through the reproductive system. etc. Mr. when growing near the sea-shore. but this certainly does not always hold good. and in a lesser degree with those under nature—were due to chance.

but likewise to include natural selection. 136 . on the other hand. we cannot tell how much to attribute to the accumulative action of natural selection. Again. But when man is the selecting agent. and it is this latter agency which answers to the survival of the fittest under nature. though not elsewhere fleshy. 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 it is the will of man which accumulates the variations in certain direction. for the conditions determine whether this or that variety shall survive. 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. of species keeping true. we clearly see that the two elements of change are distinct. due to causes of which we are quite ignorant. although living under the most opposite climates. variability is in some manner excited. 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. and. Thus.have their leaves in some degree fleshy. When a variation is of the slightest use to any being. of dissimilar varieties being produced under apparently the same external conditions. In one sense the conditions of life may be said. it is well known to furriers that animals of the same species have thicker and better fur the further north they live. either directly or indirectly. or not varying at all. and how much to the definite action of the conditions of life. innumerable instances are known to every naturalist. not only to cause variability. Such considerations as these incline me to lay less weight on the direct action of the surrounding conditions. than on a tendency to vary. 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.

2. and is exposed to danger from which it cannot escape by flight. The evidence that accidental mutilations can be inherited is at present not decisive. according to Mr. but in a rudimentary condition. or feet. The logger-headed duck of South America can only flap along the surface of the water. until they became incapable of flight. but it can defend itself. We may believe that the progenitor of the ostrich genus had habits like those of the bustard. it will perhaps be safest to look at the entire absence of the anterior tarsi in Ateuchus. they are totally deficient. In the Ateuchus or sacred beetle of the Egyptians. it is probable that the nearly wingless condition of several birds. In the Onites apelles the tarsi are so habitually lost that the insect has been described as not having them. and not one had even a relic left. while the adults have lost this power. The ostrich indeed inhabits continents. I think there can be no doubt that use in our domestic animals has strengthened and enlarged certain parts. tenanted by no beasts of prey. but as due to the effects of long-continued disuse. there is no greater anomaly in nature than a bird that cannot fly. As Controlled By Natural Selection From the facts alluded to in the first chapter. Cunningham. In some other genera they are present. and that. but the remarkable cases observed by Brown-Sequard in guinea-pigs. now inhabiting or which lately inhabited several oceanic islands. and that such modifications are inherited. Effects Of The Increased Use And Disuse Of Parts. and their rudimentary condition in some other genera. Kirby has remarked (and I have observed the same fact) that the anterior tarsi. has been caused by disuse. as efficiently as many quadrupeds. Hence. and has its wings in nearly the same condition as the domestic Aylesbury duck: it is a remarkable fact that the young birds. of many male dung-feeding beetles are often broken off. Under free nature we have no standard of comparison by which to judge of the effects of long-continued use or disuse. as the size and weight of its body were increased during successive generations. for we know not the parent-forms. its legs were used more and its wings less. can fly. for as many dung-feeding beetles are 137 . of the inherited effects of operations. he examined seventeen specimens in his own collection. yet there are several in this state. should make us cautious in denying this tendency. not as cases of inherited mutilations. but many animals possess structures which can be best explained by the effects of disuse. by kicking its enemies. As Professor Owen has remarked. and disuse diminished them. As the larger ground-feeding birds seldom take flight except to escape danger.

and especially the extraordinary fact. as certain flower-feeding coleoptera and lepidoptera. For during many successive generations each individual beetle which flew least. the tendency of natural selection to enlarge or to reduce the wings. those beetles which most readily took to flight would oftenest have been blown to sea. have. would depend on whether a greater number of individuals were saved by successfully battling with the winds. that the proportion of wingless beetles is larger on the exposed Desertas than in Madeira itself. and in some cases are quite covered by skin and fur. The eyes of moles and of some burrowing rodents are rudimentary in size. on the other hand. and that. Wollaston. until the wind lulls and the sun shines. and thus destroyed. 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. it would have been better for the good swimmers if they had been able to swim still further. either from its wings having been ever so little less perfectly developed or from indolent habit. therefore the tarsi cannot be of much importance or be much used by these insects. which absolutely require the use of their wings. This state of 138 . This is quite compatible with the action of natural selection. namely.generally found with their tarsi lost. Mr. that the beetles in Madeira. but even enlarged. that beetles in many parts of the world are very frequently blown to sea and perish. and. and which. of the twenty-nine endemic genera. Wollaston has discovered the remarkable fact that 200 beetles. For when a new insect first arrived on the island. are here almost entirely absent. or mainly due to natural selection. their wings not at all reduced. lie much concealed. that certain large groups of beetles. will have had the best chance of surviving from not being blown out to sea. are so far deficient in wings that they cannot fly. Wollaston. The insects in Madeira which are not ground-feeders. as observed by Mr. combined probably with disuse. this must happen early in life. must habitually use their wings to gain their subsistence. Wollaston suspects. As with mariners shipwrecked near a coast. In some cases we might easily put down to disuse modifications of structure which are wholly. so strongly insisted on by Mr. 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. or by giving up the attempt and rarely or never flying. as Mr. no less than twenty-three have all their species in this condition! Several facts. elsewhere excessively numerous.

It is well known that several animals. with respect to the insects alone. who had often caught them. and if so. This is certainly not the case if we look at the two whole faunas. and as eyes are certainly not necessary to animals having subterranean habits. and I was assured by a Spaniard. that they were frequently blind. having in most cases ordinary powers of vision. a burrowing rodent. In one of the blind animals." On my view we must suppose that American animals. the stand for the telescope is there. the tuco-tuco. belonging to the most different classes. though useless. but aided perhaps by natural selection. so that. with the adhesion of the eyelids and growth of fur over them. As it is difficult to imagine that eyes. in accordance with the old view of the blind animals having been separately created for the American and European caverns. In some of the crabs the foot-stalk for the eye remains. which inhabit the caves of Carniola and Kentucky. two of which were captured by Professor Silliman at above half a mile distance from the mouth of the cave. Schiodte has remarked: "We are accordingly prevented from considering the entire phenomenon in any other light than something purely local. namely. the eyes were lustrous and of large size. otherwise than as a very plain expression of that analogy which subsists generally between the fauna of Europe and of North America. as did 139 . having been inflammation of the nictitating membrane. very close similarity in their organisation and affinities might have been expected. and these animals. is even more subterranean in its habits than the mole. as appeared on dissection. their loss may be attributed to disuse. though the eye is gone. and therefore not in the profoundest depths. One which I kept alive was certainly in this condition. the cave-rat (Neotoma). and the similarity which is exhibited in a few forms between the Mammoth Cave (in Kentucky) and the caves in Carniola. In South America. or Ctenomys. the cause. acquired a dim perception of objects. are blind. though the telescope with its glasses has been lost. might in such case be an advantage. a reduction in their size. natural selection would aid the effects of disuse.the eyes is probably due to gradual reduction from disuse. As frequent inflammation of the eyes must be injurious to any animal. It is difficult to imagine conditions of life more similar than deep limestone caverns under a nearly similar climate. slowly migrated by successive generations from the outer world into the deeper and deeper recesses of the Kentucky caves. after having been exposed for about a month to a graduated light. as I am informed by Professor Silliman. could be in any way injurious to animals living in darkness.

with reference to the reptiles of 140 . Next follow those that are constructed for twilight. and as is the case with the blind Proteus. By the time that an animal had reached. we might expect from the well-known relationship of most of their other productions. Animals not far remote from ordinary forms. those destined for total darkness. for it is natural that an insect already deprived of vision should readily become adapted to dark caverns. Notwithstanding such modifications. as I hear from Professor Dana. after numberless generations.European animals into the caves of Europe. the Amblyopsis. but to distinct species. affinities to the other inhabitants of that continent. for. and. And this is the case with some of the American cave-animals. have not as yet been found anywhere except in caves. prepare the transition from light to darkness." These remarks of Schiodte's it should be understood. yet those which inhabit the several caves of Europe and America are distinct. 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. as Agassiz has remarked in regard to the blind fish. last of all. Another blind genus (Anophthalmus) offers this remarkable peculiarity. That several of the inhabitants of the caves of the Old and New Worlds should be closely related. Murray observes. the loss of vision in the cave species of this one genus has probably had no relation to its dark habitation. but it is possible that the progenitors of these several species. and then have become extinct. as a compensation for blindness. that the species. may formerly have ranged over both continents. while they were furnished with eyes. we might expect still to see in the caveanimals of America. and some of the European cave-insects are very closely allied to those of the surrounding country. and natural selection will often have effected other changes. have been accommodated to surrounding circumstances. apply not to the same. and which. as they extended themselves into darkness. the deepest recesses. such as an increase in the length of the antennae or palpi. Far from feeling surprise that some of the cave-animals should be very anomalous. and in those of Europe to the inhabitants of the European continent. 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. We have some evidence of this gradation of habit. As a blind species of Bathyscia is found in abundance on shady rocks far from caves. as Mr. and whose formation is quite peculiar. excepting in their present secluded abodes. disuse will on this view have more or less perfectly obliterated its eyes.

I am only surprised that more wrecks of ancient life have not been preserved. owing to the less severe competition to which the scanty inhabitants of these dark abodes will have been exposed. 141 .Europe.

Watson on European species of plants brought from the Azores to England. and not because they were subsequently found capable of far-extended transportation. several authentic instances could be adduced of species having largely extended. naturally habituated to different temperatures. as in the period of flowering. But whether or not this adaptation is in most cases very close. 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. we have evidence with some few plants. by adaptation to particular climates.. they become acclimatised: thus the pines and rhododendrons.3. 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. or more than. In regard to animals. As we may infer that our domestic animals were originally chosen by uncivilised man because they were useful. and from the number of plants and animals brought from different countries which are here perfectly healthy. so as to be better fitted for them than they were at first. H. in the time of sleep. the common and extraordinary 142 . and conversely. Thwaites informs me that he has observed similar facts in Ceylon. within historical times. Mr. were found to possess in this country different constitutional powers of resisting cold. but we do not positively know that these animals were strictly adapted to their native climate. raised from seed collected by Dr. and this leads me to say a few words on acclimatisation. We may infer this from our frequent inability to predict whether or not an imported plant will endure our climate. though in all ordinary cases we assume such to be the case. of their becoming. that is. acclimatisation must be readily effected during a long course of descent. analogous observations have been made by Mr. many succulent plants cannot endure a damp climate. etc.C. if it be true that all the species of the same genus are descended from a single parent-form. Acclimatisation Habit is hereditary with plants. in the amount of rain requisite for seeds to germinate. But the degree of adaptation of species to the climates under which they live is often overrated. to a certain extent. Hooker from the same species growing at different heights on the Himalayas. or conversely. As it is extremely common for distinct species belonging to the same genus to inhabit hot and cold countries. their range from warmer to colder latitudes. and because they bred readily under confinement. nor do we know that they have subsequently become specially acclimatised to their new homes. and I could give other cases. So again.

for they live under the cold climate of Faroe in the north and of the Falklands in the south. in which certain varieties are habitually recommended for the northern and others for the southern states. the result must. and now have a far wider range than any other rodent.capacity in our domestic animals of not only withstanding the most different climates. The case of the Jerusalem artichoke. even in the ancient Encyclopaedias of China. common to most animals. they cannot owe their constitutional differences to habit. and how much to both means combined. On this view. but they have been transported by man to many parts of the world. the capacity of enduring the most different climates by man himself and by his domestic animals. That habit or custom has some influence. In treatises on many kinds of cultivated plants. And as it is not likely that man should have succeeded in selecting so many breeds and sub-breeds with constitutions specially fitted for their own districts. and how much to the natural selection of varieties having different innate constitutions. for instance. but as examples of a very common flexibility of constitution. certain varieties are said to withstand certain climates better than others. push the foregoing argument too far. and the fact of the extinct elephant and rhinoceros having formerly endured a glacial climate. How much of the acclimatisation of species to any peculiar climate is due to mere habit. ought not to be looked at as anomalies. brought. whereas the living species are now all tropical or subtropical in their habits. on account of the probable origin of some of our domestic animals from several wild stocks: the blood. both from analogy and from the incessant advice given in agricultural works. and on many an island in the torrid zones. natural selection would inevitably tend to preserve those individuals which were born with constitutions best adapted to any country which they inhabited. and 143 . be due to habit. I think. this is strikingly shown in works on fruit-trees published in the United States. I must believe. We must not. however. is an obscure question. to be very cautious in transporting animals from one district to another. into action. Hence adaptation to any special climate may be looked at as a quality readily grafted on an innate wide flexibility of constitution. and as most of these varieties are of recent origin. The rat and mouse cannot be considered as domestic animals. but of being perfectly fertile (a far severer test) under them. of a tropical and arctic wolf may perhaps be mingled in our domestic breeds. On the other hand. 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. under peculiar circumstances. which is never propagated in England by seed.

and of this fact I have myself observed striking instances. Nor let it be supposed that differences in the constitution of seedling kidneybeans never appear. his kidneybeans so early that a very large proportion are destroyed by frost. as proving that acclimatisation cannot be effected. played a considerable part in the modification of the constitution and structure. consequently. have. has even been advanced. in some cases. and then again get seed from these seedlings. but that the effects have often been largely combined with. for it is now as tender as ever it was! The case. 144 . and then collect seed from the few survivors.of which. and with much greater weight. On the whole. the natural selection of innate variations. or use and disuse. during a score of generations. but until some one will sow. of the kidney-bean has been often cited for a similar purpose. for an account has been published how much more hardy some seedlings are than others. and sometimes overmastered by. new varieties have not been produced. with the same precautions. the experiment cannot be said to have been even tried. with care to prevent accidental crosses. we may conclude that habit. also.

most imperfectly understood. We shall presently see that simple inheritance often gives the false appearance of correlation. 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.4. that when slight variations in any one part occur and are accumulated through natural selection. Hard parts seem to affect the form of adjoining soft parts. as in the union of the petals into a tube. tend to cohere. in the front and hind legs. 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. One of the most obvious real cases is. coexist without our being able to assign any reason. during its growth and development. may be mastered more or less completely by natural selection: thus a family of stags once existed with an antler only on one side. The several parts which are homologous. this is often seen in monstrous plants: and nothing is more common than the union of homologous parts in normal structures. and which. it might probably have been rendered permanent by natural selection. are identical in structure. Geoffroy St. Is. What can be more singular than the relation in cats between complete whiteness and blue eyes with deafness. Others believe that the shape of the pelvis in the human mother influences by pressure the shape of the head of the child. Correlated Variation I mean by this expression that the whole organisation is so tied together. I do not doubt. and even in the jaws and limbs. These tendencies. This is a very important subject. In snakes. and that others rarely. between their feathered feet and skin betwixt the outer toes. varying together. at an early embryonic period. M. and which are necessarily exposed to similar conditions. Hilaire has forcibly remarked that certain malconformations frequently. that variations of structure arising in the young or larvae naturally tend to affect the structure of the mature animal. The nature of the bond is frequently quite obscure. or between the presence of more or 145 . or between the tortoiseshell colour and the female sex. or in pigeons. Homologous parts. the shape of the body and the manner of swallowing determine the position and form of several of the most important viscera. as has been remarked by some authors. and no doubt wholly different classes of facts may be here easily confounded together. other parts become modified. for the lower jaw is believed by some anatomists to be homologous with the limbs. and if this had been of any great use to the breed. according to Schlegel.

Everyone is familiar with the difference between the ray and central florets of. for in some Compositae the seeds of the outer and inner florets differ. Sprengel's idea that the ray-florets serve to attract insects. the daisy. and the shape of the seeds in the rayflorets of some Compositae countenances this idea. With respect to the development of the corolla. but this can hardly be the sole case. or to their mutual pressure. etc. and this difference is often accompanied with the partial or complete abortion of the reproductive organs. with the future colour of its plumage. than that of the difference between the outer and inner flowers in some Compositous and Umbelliferous plants. the nectary is not quite aborted but is much shortened. the central flower thus becoming peloric or regular. is highly probable. that it has little value. Hooker informs me. but there are so many exceptions to this rule. are likewise on the whole the most abnormal in their teeth.less down on the young pigeon when first hatched. viz. or necessary for the fertilisation of these plants. When the colour is absent from only one of the two upper petals. though here no doubt homology comes into play? With respect to this latter case of correlation. and therefore of natural selection. again. Cetacea (whales) and Edentata (armadilloes. by drawing nourishment from the reproductive organs causes their abortion. but with the Umbelliferae it is by no means. without any difference in the corolla. as Dr. Possibly these several differences may be connected with the different flow of nutriment towards the central and external flowers. or. as an instance of this fact. the adherent nectary is quite aborted. independently of utility. that is to become abnormally symmetrical. the species with the densest heads which most frequently differ in their inner and outer flowers. I think it can hardly be accidental that the two orders of mammals which are most abnormal in their dermal covering. and as a striking case of correlation. and if so. It might have been thought that the development of the raypetals. at least. the relation between the hair and the teeth in the naked Turkish dog. whose agency is highly advantageous. natural selection may have come into play. These differences have sometimes been attributed to the pressure of the involucra on the florets. I may add. as Mr.). that with irregular flowers those nearest to the axis are most subject to peloria.. But with respect to the 146 . and when this occurs. that in many pelargoniums the two upper petals in the central flower of the truss often lose their patches of darker colour. But in some of these plants the seeds also differ in shape and sculpture. scaly ant-eaters. for instance. I know of no case better adapted to show the importance of the laws of correlation and variation. Mivart has remarked. We know.

without being. would naturally be thought to be in some necessary manner correlated. and. after thousands of generations. For instance. Hence modifications of structure. unless the capsules were open. I should explain this rule by the impossibility of seeds gradually becoming winged through natural selection. and these two modifications. 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. having been transmitted to a whole group of descendants with diverse habits. 147 . of the slightest service to the species. and which in truth are simply due to inheritance. which are not always correlated with any difference in the corolla. for an ancient progenitor may have acquired through natural selection some one modification in structure. viewed by systematists as of high value. Some other correlations are apparently due to the manner in which natural selection can alone act. some other and independent modification. Alph. gain an advantage over others less well fitted for wide dispersal. it seems impossible that their differences in shape. for in this case alone could the seeds. We may often falsely attribute to correlated variation structures which are common to whole groups of species.seeds. may be wholly due to the laws of variation and correlation. as far as we can judge. De Candolle has remarked that winged seeds are never found in fruits which do not open. can be in any way beneficial.

but in the parasitic and protected Proteolepas. it rarely flows. on the other hand. that when a cirripede is parasitic within another cirripede and is thus protected. would 148 . at about the same time. In our poultry. a large tuft of feathers on the head is generally accompanied by a diminished comb. believe in its truth. 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. to another part. here give any instances. that natural selection is continually trying to economise in every part of the organisation. the whole anterior part of the head is reduced to the merest rudiment attached to the bases of the prehensile antennae. "in order to spend on one side. I suspect.5. This is the case with the male Ibla. or. before useful. for it will profit the individual not to have its nutriment wasted in building up a useless structure. more especially botanists. and a large beard by diminished wattles. and. Compensation And Economy Of Growth The elder Geoffroy and Goethe propounded. I will not. its diminution will be favoured. but many good observers. namely. the actual withdrawal of nutriment from one part owing to the excess of growth in another and adjoining part. that some of the cases of compensation which have been advanced. however. as Goethe expressed it. thus it is difficult to get a cow to give much milk and to fatten readily." I think this holds true to a certain extent with our domestic productions: if nourishment flows to one part or organ in excess. it loses more or less completely its own shell or carapace. the fruit itself gains largely in size and quality. When the seeds in our fruits become atrophied. at least in excess. of a part being largely developed through natural selection and another and adjoining part being reduced by the same process or by disuse. Now the saving of a large and complex structure. I can thus only understand a fact with which I was much struck when examining cirripedes. and likewise some other facts. becomes less useful. for I see hardly any way of distinguishing between the effects. their law of compensation or balancement of growth. If under changed conditions of life a structure. also. when rendered superfluous. and furnished with great nerves and muscles. on the one hand. The same varieties of the cabbage do not yield abundant and nutritious foliage and a copious supply of oil-bearing seeds. nature is forced to economise on the other side. and of which many other instances could be given: namely. may be merged under a more general principle. With species in a state of nature it can hardly be maintained that the law is of universal application.

each would have a better chance of supporting itself. natural selection will tend in the long run to reduce any part of the organisation. that natural selection may perfectly well succeed in largely developing an organ without requiring as a necessary compensation the reduction of some adjoining part. superfluous. as soon as it becomes. without by any means causing some other part to be largely developed in a corresponding degree. by less nutriment being wasted. Thus. for in the struggle for life to which every animal is exposed.be a decided advantage to each successive individual of the species. And conversely. 149 . as I believe. through changed habits.

that beings which stand low in the scale of nature are more variable than those which are higher. Natural selection. both with varieties and species. that when any part or organ is repeated many times in the same individual (as the vertebrae in snakes. and the stamens in polyandrous flowers) the number is variable. whilst a tool for some particular purpose must be of some particular shape. Geoffroy St. 150 . and as long as the same part has to perform diversified work. Hilaire. As "vegetative repetition. We shall have to recur to this subject. Multiple. when it occurs in lesser numbers. as is generally admitted. is constant. and consequently from natural selection having had no power to check deviations in their structure." to use Professor Owen's expression. Rudimentary parts. it should never be forgotten. Rudimentary. I presume that lowness here means that the several parts of the organisation have been but little specialised for particular functions. The same author as well as some botanists. are apt to be highly variable. can act solely through and for the advantage of each being. the foregoing statements accord with the common opinion of naturalists. have further remarked that multiple parts are extremely liable to vary in structure. 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. that is. In the same way that a knife which has to cut all sorts of things may be of almost any shape. we can perhaps see why it should remain variable. and I will here only add that their variability seems to result from their uselessness.6. as remarked by Is. is a sign of low organisation. And Lowly-Organised Structures Are Variable It seems to be a rule. whereas the number of the same part or organ.

but are not directly connected with the act of reproduction. I shall. as they seldom offer remarkable secondary sexual characters. and which cannot possibly be here introduced. 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. and the amount of variation in the individuals of the same species is so great that 151 . in a future work. A Part Developed In Any Species In An Extraordinary Degree Or Manner. and I am fully convinced that the rule almost always holds good. very important structures. but I hope that I have made due allowances for them. when displayed in any unusual manner. whether or not displayed in any unusual manner—of which fact I think there can be little doubt. The term. The opercular valves of sessile cirripedes (rock barnacles) are. Thus. The rule applies to males and females.7. I will here give only one. the homologous valves in the different species being sometimes wholly unlike in shape. the wing of the bat is a most abnormal structure in the class of mammals. Professor Owen. unless it be unusually developed in one species or in a few species in comparison with the same part in many closely allied species. seems to have come to a nearly similar conclusion. relates to characters which are attached to one sex. I am aware of several causes of error. The rule being so plainly applicable in the case of secondary sexual characters. in every sense of the word. these valves present a marvellous amount of diversification. It should be understood that the rule by no means applies to any part. but more rarely to females. also. 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. as it illustrates the rule in its largest application. The rule applies very strongly in the case of secondary sexual characters. Tends To Be Highly Variable Several years ago I was much struck by a remark to the above effect made by Mr. I can only state my conviction that it is a rule of high generality. Waterhouse. secondary sexual characters. and they differ extremely little even in distinct genera. whilst investigating this order. but the rule would not apply here. used by Hunter. but in the several species of one genus. I particularly attended to Mr. But that our rule is not confined to secondary sexual characters is clearly shown in the case of hermaphrodite cirripedes. may be due to the great variability of these characters. Waterhouse's remark. In Comparison With The Same Part In Allied Species. because the whole group of bats possesses wings. Pyrgoma. give a list of all the more remarkable cases.

we see a nearly parallel case. the fair presumption is that it is of high importance to that species: nevertheless it is in this case eminently liable to variation. and no selection be applied. Why should this be so? On the view that each species has been independently created. Look at the individuals of the same breed of the pigeon.it is no exaggeration to state that the varieties of the same species differ more from each other in the characters derived from these important organs. and have been modified through natural selection. that those points in our domestic animals. the tendency to reversion to a less perfect state. and this would have seriously shaken my belief in its truth. 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- 152 . with all its parts as we now see them. vary extremely little. If. these being the points now mainly attended to by English fanciers. in the beak and wattle of carriers. and the rule certainly seems to hold good in this class. as well as an innate tendency to new variations. are also eminently liable to variation. In the long run selection gains the day. etc. that part (for instance. I can see no explanation. for in such cases natural selection either has not or cannot come into full play. any part or the whole animal be neglected. When we see any part or organ developed in a remarkable degree or manner in a species. But what here more particularly concerns us is. and. As with birds the individuals of the same species. in our domestic animals.. had not the great variability in plants made it particularly difficult to compare their relative degrees of variability. I cannot make out that it applies to plants. I think we can obtain some light. on the one hand. many departing widely from the standard. There may truly be said to be a constant struggle going on between. First let me make some preliminary remarks. 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. and thus the organisation is left in a fluctuating condition. which at the present time are undergoing rapid change by continued selection. and in those which have been but little specialised for any particular purpose. and perhaps in polymorphic groups. I have particularly attended to them. the power of steady selection to keep the breed true. than do the species belonging to other distinct genera. But on the view that groups of species are descended from some other species. Even in the same sub-breed. as in that of the short-faced tumbler. and see what a prodigious amount of difference there is in the beak of tumblers. In rudimentary organs. in the carriage and tail of fantails. inhabiting the same country. it is notoriously difficult to breed nearly perfect birds. on the other hand.

An extraordinary amount of modification implies an unusually large and long-continued amount of variability. When a part has been developed in an extraordinary manner in any one species. is the case. as species rarely endure for more than one geological period. That the struggle between natural selection on the one hand. This period will seldom be remote in any extreme degree. Now let us turn to nature. will in the course of time cease. and that the most abnormally developed organs may be made constant.faced strain. it must have existed. But as long as selection is rapidly going on. But as the variability of the extraordinarily developed part or organ has been so great and long-continued within a period not excessively remote. according to our theory. as a general rule. we might. as it may be called. 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. however abnormal it may be. and thus it has come not to be more variable than any other structure. 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. which has continually been accumulated by natural selection for the benefit of the species. has been transmitted in approximately the same condition to many modified descendants. compared with the other species of the same genus. when an organ. and by the continued rejection of those tending to revert to a former and less modified condition. 153 . 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. for an immense period in nearly the same state. Hence. still present in a high degree. And this. and the tendency to reversion and variability on the other hand. 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. much variability in the parts undergoing modification may always be expected. I am convinced. I see no reason to doubt. as in the case of the wing of the bat.

have to return to this point in the chapter on Classification. yet only indirectly. 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. because they are taken from parts of less physiological importance than those commonly used for classing genera. Or to state the case in another manner: the points in which all the species of a genus resemble each other. often becomes variable. Geoffroy St. which differs from the same part in other independently created species of the same genus. 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. though its physiological importance may remain the same. DIFFERS considerably in closely-allied species. Something of the same kind applies to monstrosities: at least Is. that the more an organ normally differs in the different species of the same group. On the ordinary view of each species having been independently created. it is often VARIABLE in the individuals of the same species. the colour would be only a specific character. but if all the species had blue flowers. I have chosen this example because the explanation which most naturalists would advance is not here applicable. and no one would be surprised at one of the blue species varying into red. or conversely. It would be almost superfluous to adduce evidence in support of the statement. the more subject it is to anomalies in the individuals. Hilaire apparently entertains no doubt. I shall. and in which they 154 . and which have thus come to differ. and its variation would be a more unusual circumstance. true. the colour would become a generic character. that when an author remarks with surprise that some important organ or part. And this fact shows that a character. however. be more variable than those parts which are closely alike in the several species? I do not see that any explanation can be given. why should that part of the structure. I have repeatedly noticed in works on natural history. It is notorious that specific characters are more variable than generic. when it sinks in value and becomes only of specific value. But on the view that species are only strongly marked and fixed varieties. which is generally of generic value. that ordinary specific characters are more variable than generic.8. namely. which is generally very constant throughout a large group of species. but with respect to important characters. I believe this explanation is partly. Specific Characters More Variable Than Generic Characters The principle discussed under the last heading may be applied to our present subject.

or only in a slight degree. 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. and these characters may be attributed to inheritance from a common progenitor. the points in which species differ from other species of the same genus are called specific characters. On the other hand. fitted to more or less widely different habits. and subsequently have not varied or come to differ in any degree.differ from allied genera. are called generic characters. for it can rarely have happened that natural selection will have modified several distinct species. it is not probable that they should vary at the present day. it is probable that they should still often be in some degree variable—at least more variable than those parts of the organisation which have for a very long period remained constant. and as these specific characters have varied and come to differ since the period when the species branched off from a common progenitor. 155 .

than in other parts of their organisation. without my entering on details. the amount of difference between the males of gallinaceous birds. and as the differences in these cases are of a very unusual nature. which is less rigid in its action than ordinary selection. that secondary sexual characters are highly variable. became variable. but in certain genera the neuration differs in the different species. because common to large groups. Lubbock has recently remarked. for instance. sexual selection will have had a wide scope for action. 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. the number varies greatly and the number likewise differs in the two sexes of the same species. Again in the fossorial hymenoptera. 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. Secondary Sexual Characters Variable I think it will be admitted by naturalists. but in the Engidae. 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. it is highly probable. Of this fact I will give in illustration the first two instances which happen to stand on my list. as have the two sexes of any one species. The same number of joints in the tarsi is a character common to very large groups of beetles. be 156 . "In Pontella. the relation can hardly be accidental. for they are accumulated by sexual selection. It will also be admitted that species of the same group differ from each other more widely in their secondary sexual characters. Sir J. as Westwood has remarked. and likewise in the two sexes of the same species. The cause of the original variability of these characters is not manifest.9. 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. in which secondary sexual characters are strongly displayed. with the amount of difference between the females. for instance. compare. that several minute crustaceans offer excellent illustrations of this law. but we can see why they should not have been rendered as constant and uniform as others. or of its early descendants. Whatever the cause may be of the variability of secondary sexual characters. the neuration of the wings is a character of the highest importance. It is a remarkable fact. variations of this part would. Consequently. as it does not entail death. but only gives fewer offspring to the less favoured males. whatever part of the structure of the common progenitor.

if it be common to a whole group of species. are all principles closely connected together. 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. from whom they have inherited much in common. and for ordinary purposes. I conclude that the greater variability of specific characters. and the slight degree of variability in a part. according to the lapse of time. that secondary sexual and ordinary specific differences are generally displayed in the same parts of the organisation. and likewise to fit the two sexes of the same species to each other. that the great variability of secondary sexual characters and their great difference in closely allied species. than of generic characters. All being mainly due to the species of the same group being the descendants of a common progenitor. or to fit the males to struggle with other males for the possession of the females. or those which distinguish species from species.taken advantage of by natural and sexual selection. to natural selection having more or less completely. and thus having been adapted for secondary sexual. then. and to variations in the same parts having been accumulated by natural and sexual selection. 157 . or those which are possessed by all the species. Finally. in order to fit the several places in the economy of nature. however extraordinarily it may be developed. overmastered the tendency to reversion and to further variability. to sexual selection being less rigid than ordinary selection. 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.

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. and a consequent tendency to vary in a like manner. So That A Variety Of One Species Often Assumes A Character Proper To An Allied Species. in the enlarged stems. I think. As all these marks are characteristic of the parent rock-pigeon. of the Swedish turnip and ruta-baga. the common turnip. these then are analogous variations in two or more distinct races. We may. we have another case. with the outer feathers externally edged near their bases with white. characters not possessed by the aboriginal rock-pigeon. or as commonly called roots. because.10. Similar cases occurring with insects under natural conditions have lately been discussed with much ability by Mr. and by various authors in our cereals. of slaty-blue birds with two black bars on the wings. however. not to the vera causa of community of descent. white loins. and to these a third may be added. namely. present sub-varieties with reversed feathers on the head. confidently come to this conclusion. who has grouped them under his law of equable variability. In the vegetable kingdom we have a case of analogous variation. but to three separate yet closely related acts of creation. these coloured marks are eminently liable to appear in the crossed offspring of two distinct and 158 . as we have seen. and not of a new yet analogous variation appearing in the several breeds. The most distinct breeds of the pigeon. the occasional appearance in all the breeds. plants which several botanists rank as varieties produced by cultivation from a common parent: if this be not so. 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. and with feathers on the feet. in countries widely apart. Walsh. a bar at the end of the tail. we should have to attribute this similarity in the enlarged stems of these three plants. Distinct Species Present Analogous Variations. Or Reverts To Some Of The Characters Of An Early Progenitor These propositions will be most readily understood by looking to our domestic races. According to the ordinary view of each species having been independently created. Many similar cases of analogous variation have been observed by Naudin in the great gourd family. the fantail. With pigeons. the case will then be one of analogous variation in two so-called distinct species. when acted on by similar unknown influences. I presume that no one will doubt that this is a case of reversion. namely.

is not greater than that of quite useless or rudimentary organs being similarly transmitted. with the several marks. But characters exclusively due to analogous variation would probably be of an unimportant nature. for all that we can see to the contrary. and at last. this other species being. reappears after a great number of generations. to use a common expression. which very rarely produces a blue bird. beyond the influence of the mere act of crossing on the laws of inheritance. But when a breed has been crossed only once by some other breed. When a character which has been lost in a breed. from one ancestor. the most probable hypothesis is. A mere tendency to produce a rudiment is indeed sometimes thus inherited. and yet. It might further be expected that the species of the same genus would 159 . whether strong or weak.differently coloured breeds. under unknown favourable conditions. and in this case there is nothing in the external conditions of life to cause the reappearance of the slaty-blue. probably for hundreds of generations. it is probable that there is a latent tendency in each generation to produce blue plumage. it might be expected that they would occasionally vary in an analogous manner. In a breed which has not been crossed. but in which BOTH parents have lost some character which their progenitor possessed. it is generally believed that a tendency to reversion is retained by this remnant of foreign blood. to reproduce the lost character might. as was formerly remarked. in accordance with the different habits of the species. After twelve generations. With the barb-pigeon. for the preservation of all functionally important characters will have been determined through natural selection. or that a variety of one species would resemble in certain characters another and distinct species. as we see. the tendency. the offspring occasionally show for many generations a tendency to revert in character to the foreign breed—some say. As all the species of the same genus are supposed to be descended from a common progenitor. is only 1 in 2048. for instance. only a well-marked and permanent variety. is developed. but that in each successive generation the character in question has been lying latent. not that one individual suddenly takes after an ancestor removed by some hundred generations. according to our view. The abstract improbability of such a tendency being transmitted through a vast number of generations. so that the varieties of two or more species would resemble each other. No doubt it is a very surprising fact that characters should reappear after having been lost for many. be transmitted for almost any number of generations. for a dozen or even a score of generations. the proportion of blood.

A white ass. what cases are reversions to formerly existing characters. The difficulty in distinguishing variable species is largely due to the varieties mocking. The stripe on the shoulder is sometimes double. we do not know the common ancestor of any natural group. yet we ought. on our theory. The ass sometimes has very distinct transverse bars on its legs. partly under domestication and partly under nature. but here. but we might have inferred that the blue colour was a case of reversion from the number of the markings. and is very variable in length and outline. as it were. like those on the legs of a zebra. I can only repeat that such cases certainly occur. and what are new but analogous variations. we could not have told. A considerable catalogue. that they have in varying assumed some of the characters of the others. or actually quite 160 . however. which themselves can only doubtfully be ranked as species. Hence. not indeed as affecting any important character. But the best evidence of analogous variations is afforded by parts or organs which are generally constant in character. As. and from inquiries which I have made. the same part or organ in an allied species. whether such characters in our domestic breeds were reversions or only analogous variations. I believe this to be true. however. could be given of forms intermediate between two other forms. we did not know that the parent rock-pigeon was not feather-footed or turn-crowned. we cannot distinguish between reversionary and analogous characters. has been described without either spinal or shoulder stripe. in some degree. also. but from occurring in several species of the same genus. which are correlated with this tint. and this shows. I will. unless all these closely allied forms be considered as independently created species. and which would not probably have all appeared together from simple variation. More especially we might have inferred this from the blue colour and the several marks so often appearing when differently coloured breeds are crossed. for instance.occasionally exhibit reversions to long-lost characters. It has been asserted that these are plainest in the foal. If. It is a case almost certainly of reversion. although under nature it must generally be left doubtful. and seem to me very remarkable. but which occasionally vary so as to resemble. and these stripes are sometimes very obscure. sometimes to find the varying offspring of a species assuming characters which are already present in other members of the same group. I have collected a long list of such cases. And this undoubtedly is the case. but NOT an albino. as before. I lie under the great disadvantage of not being able to give them. other species of the same genus. give one curious and complex case.

in dark-coloured asses. I have also reason to suspect. 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. 161 . The spine is always striped. and a small dun Welsh pony has been carefully described to me. mouse-duns. Edwards. Blyth has seen a specimen of the hemionus with a distinct shoulder-stripe. and the shoulder-stripe.W. who examined this breed for the Indian Government. and from Norway in the north to the Malay Archipelago in the south. as I hear from Colonel Poole. but Dr. is sometimes striped. W. by the term dun a large range of colour is included. and sometimes quite disappear in old horses. 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. In the northwest part of India the Kattywar breed of horses is so generally striped. from information given me by Mr. from one between brown and black to a close approach to cream colour. 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. is common. both with THREE parallel stripes on each shoulder. Mr. The stripes are often plainest in the foal. the legs are generally barred. transverse bars on the legs are not rare in duns. and I have been informed by Colonel Poole that foals of this species are generally striped on the legs and faintly on the shoulder. that. In all parts of the world these stripes occur far oftenest in duns and mouse-duns. moreover.lost. With respect to the horse. and in one instance in a chestnut. is without bars on the legs. The quagga. This foal. though so plainly barred like a zebra over the body. I have myself seen a dun Devonshire pony. I have collected cases in England of the spinal stripe in horses of the most distinct breeds. which is sometimes double and sometimes treble. when a week old. and I have seen a trace in a bay horse. that with the English race-horse the spinal stripe is much commoner in the foal than in the full-grown animal. The koulan of Pallas is said to have been seen with a double shoulder-stripe. dark. Gray has figured one specimen with very distinct zebralike bars on the hocks. a horse without stripes is not considered as purely bred. and its legs were feebly striped. was marked on its hinder quarters and on its forehead with numerous very narrow. zebra-like bars. and of ALL colours. the side of the face. Colonel Poole has seen both gray and bay Kattywar horses striped when first foaled. though it properly has none. a faint shoulder-stripe may sometimes be seen in duns. All the stripes soon disappeared completely.

But this view may be safely rejected. who has written on this subject. and this hybrid. In Lord Morton's famous hybrid. The appearance of the stripes is not accompanied by any change 162 . What now are we to say to these several facts? We see several distinct species of the horse genus becoming. one of which. striped on the legs like a zebra. the lanky Kattywar race. nevertheless had all four legs barred. inhabiting the most distant parts of the world. like those on the dun Devonshire and Welsh ponies. for it is highly improbable that the heavy Belgian cart-horse. Norwegian cobs. Martin. etc. in certain parts of the United States. which I have seen. Gray (and he informs me that he knows of a second case) from the ass and the hemionus. Welsh ponies. Rollin asserts that the common mule from the ass and horse is particularly apt to have bars on its legs. according to Mr. or striped on the shoulders like an ass. and even had some zebra-like stripes on the sides of its face. the hybrid and even the pure offspring subsequently produced from the same mare by a black Arabian sire. should have all have been crossed with one supposed aboriginal stock.I am aware that Colonel Hamilton Smith. Gosse.. were much more plainly barred across the legs than is even the pure quagga. the legs were much more plainly barred than the rest of the body. and had three short shoulder-stripes. and Mr. about nine out of ten mules have striped legs. and was. was striped. the dun. though the ass only occasionally has stripes on his legs and the hemionus has none and has not even a shoulder-stripe. I was so convinced that not even a stripe of colour appears from what is commonly called chance. 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. in his excellent treatise on the horse. answered in the affirmative. In four coloured drawings. believes that the several breeds of the horse are descended from several aboriginal species. as we have seen. Lastly. by simple variation.C. Now let us turn to the effects of crossing the several species of the horse genus. a hybrid has been figured by Dr. of hybrids between the ass and zebra. has given a figure of a similar mule. from a chestnut mare and male 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. and that the above-described appearances are all due to ancient crosses with the dun stock. and in one of them there was a double shoulder-stripe. and this is another most remarkable case. With respect to this last fact. I once saw a mule with its legs so much striped that any one might have thought that it was a hybrid zebra. W.

sometimes prevails. and when any breed assumes by simple variation a bluish tint. we see a strong tendency for the blue tint and bars and marks to reappear in the mongrels. He who believes that each equine species was independently created. Call the breeds of pigeons. quagga. is—that there is a TENDENCY in the young of each successive generation to produce the long-lost character. so as often to become striped like the other species of the genus. and that each has been created with a strong tendency. I have stated that the most probable hypothesis to account for the reappearance of very ancient characters. 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. assert that each species has been created with a tendency to vary. but had been created in stone so as to mock the shells now living on the sea-shore. and zebra. the hemionus. It makes the works of God a mere mockery and deception. I venture confidently to look back thousands on thousands of generations. as it seems to me. but other species of the genus. or by any other new character. with certain bars and other marks. to produce hybrids resembling in their stripes. We see this tendency to become striped most strongly displayed in hybrids from between several of the most distinct species. and that this tendency. species. 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. to reject a real for an unreal. and I see an animal striped like a zebra. 163 . from unknown causes. that fossil shells had never lived. I would almost as soon believe with the old and ignorant cosmogonists. the common parent of our domestic horse (whether or not it be descended from one or more wild stocks) of the ass. not their own parents. To admit this view is. and how exactly parallel is the case with that of the species of the horse genus! For myself.of form. I presume. some of which have bred true for centuries. When the oldest and truest breeds of various colours are crossed. these bars and other marks invariably reappear. or at least for an unknown cause. will. in this particular manner. both under nature and under domestication. when crossed with species inhabiting distant quarters of the world. but perhaps otherwise very differently constructed.

Changes of structure at an early age may affect parts subsequently developed. that organic beings low in the scale are more variable than those standing higher in the scale. appear in many cases to have been potent in their effects. but we have also seen in the second chapter that the same principle applies to the whole individual. on an average. we now find. and homologous parts tend to cohere. undoubtedly occur. the nature of which we are unable to understand. so that their modifications have not been closely checked by natural selection. But whenever we have the means of instituting a comparison. Changed conditions generally induce mere fluctuating variability. Specific characters—that is. and many cases of correlated variation. perhaps arising from such parts not having been closely specialised for any particular function. and which have their whole organisation more specialised. 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. for in a district where many species of a genus are found—that is. and these may become strongly marked in the course of time. are not regulated by natural selection. but sometimes they cause direct and definite effects. and use in strengthening. Homologous parts tend to vary in the same manner. though we have not sufficient evidence on this head. or those which have long been inherited. In these remarks we have referred to special parts or organs being still variable. and have not differed within this same period. Not in one case out of a hundred can we pretend to assign any reason why this or that part has varied. the same laws appear to have acted in producing the lesser differences between varieties of the same species. most 164 . because they have recently varied and thus come to differ. It follows probably from this same cause. and the greater differences between species of the same genus. and disuse in weakening and diminishing organs. Multiple parts are variable in number and in structure. Modifications in hard parts and in external parts sometimes affect softer and internal parts.11. perhaps it tends to draw nourishment from the adjoining parts. Rudimentary organs. Summary Our ignorance of the laws of variation is profound. and every part of the structure which can be saved without detriment will be saved. and hence are variable. Habit in producing constitutional peculiarities. When one part is largely developed. where there has been much former variation and differentiation. or where the manufactory of new specific forms has been actively at work—in that district and among these species.

in comparison with the same part or organ in the allied species. must have gone through an extraordinary amount of modification since the genus arose. Secondary sexual characters are highly variable. requiring a long lapse of time—in this case. for variation is a longcontinued and slow process. Although new and important modifications may not arise from reversion and analogous variation. Any part or organ developed to an extraordinary size or in an extraordinary manner. and thus we can understand why it should often still be variable in a much higher degree than other parts. or these same species may occasionally revert to some of the characters of their ancient progenitors. 165 . and specific differences to the several species of the same genus. in however extraordinary a manner it may have been developed. 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. naturally tend to present analogous variations. 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. Species inheriting nearly the same constitution from a common parent. 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.varieties. and exposed to similar influences. natural selection has succeeded in giving a fixed character to the organ. such modifications will add to the beautiful and harmonious diversity of nature. 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. and such characters differ much in the species of the same group.

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. 166 .

the greater number are only apparent. can instincts be acquired and modified through natural selection? What shall we say to the instinct which leads the bee to make cells. is it possible that an animal having. which serves as a flyflapper. but. do we not everywhere see innumerable transitional forms? Why is not all nature in confusion. being sterile and producing sterile offspring. well defined? Secondly. why. a crowd of difficulties will have occurred to him. if species have descended from other species by fine gradations. and those that are real are not. 167 . an organ so wonderful as the eye? Thirdly. I think. Instinct and Hybridism in the two succeeding chapters. such as the tail of a giraffe. and which has practically anticipated the discoveries of profound mathematicians? Fourthly. Long before the reader has arrived at this part of my work. for instance.1. how can we account for species. the structure and habits of a bat. when crossed. 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. on the other hand. when varieties are crossed. and. instead of the species being. on the one hand. an organ of trifling importance. to the best of my judgment. as we see them. their fertility is unimpaired? The two first heads will be here discussed. Some of them are so serious that to this day I can hardly reflect on them without being in some degree staggered. fatal to the theory. These difficulties and objections may be classed under the following heads: First. some miscellaneous objections in the following chapter. whereas.

Hence. and has supplanted and exterminated its original parentform and all the transitional varieties between its past and present states. These representative species often meet and interlock. till the one replaces the other. and I will here only state that I believe the answer mainly lies in the record being incomparably less perfect than is generally supposed. 168 . Hence we ought not to expect at the present time to meet with numerous transitional varieties in each region. and may be embedded there in a fossil condition. we generally meet at successive intervals with closely allied or representative species. But I think it can be in large part explained. though they must have existed there. and only at long intervals of time. each has become adapted to the conditions of life of its own region. The crust of the earth is a vast museum. But it may be urged that when several closely allied species inhabit the same territory. But if we compare these species where they intermingle. as by this theory innumerable transitional forms must have existed. why do we not now find closely-linking intermediate varieties? This difficulty for a long time quite confounded me. But. 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. if we look at each species as descended from some unknown form. evidently filling nearly the same place in the natural economy of the land. and finally to exterminate. 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. each new form will tend in a fully-stocked country to take the place of. we surely ought to find at the present time many transitional forms. On The Absence Or Rarity Of Transitional Varieties As natural selection acts solely by the preservation of profitable modifications. they are generally as absolutely distinct from each other in every detail of structure as are specimens taken from the metropolis inhabited by each. Let us take a simple case: in travelling from north to south over a continent. and during the process of modification. its own less improved parent-form and other less-favoured forms with which it comes into competition.2. and as the one becomes rarer and rarer. but the natural collections have been imperfectly made. By my theory these allied species are descended from a common parent. Thus extinction and natural selection go hand in hand. having intermediate conditions of life. the other becomes more and more frequent. But in the intermediate region.

Forbes in sounding the depths of the sea with the dredge. but in large part on the presence of other species. we generally find them tolerably numerous over a large territory. were it not for other competing species. the range of any one species. would increase immensely in numbers. will. Geology would lead us to believe that most continents have been broken up into islands even during the later tertiary periods. 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. In looking at species as they are now distributed over a wide area. The same fact has been noticed by E. be extremely liable 169 . not blending one into another by insensible gradations. But I will pass over this way of escaping from the difficulty.In the first place we should be extremely cautious in inferring. then becoming somewhat abruptly rarer and rarer on the confines. We see the same fact in ascending mountains. during fluctuations in the number of its enemies or of its prey. or in the nature of the seasons. in short. 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. will tend to be sharply defined. Hence the neutral territory between two representative species is generally narrow in comparison with the territory proper to each. where it exists in lessened numbers. for I believe that many perfectly defined species have been formed on strictly continuous areas. or with which it comes into competition. depending as it does on the range of others. even in its metropolis. these facts ought to cause surprise. as Alph. each species on the confines of its range. that nearly all either prey on or serve as prey for others. and sometimes it is quite remarkable how abruptly. as climate and height or depth graduate away insensibly. has played an important part in the formation of new species. Moreover. on which it lives. or by which it is destroyed. De Candolle has observed. To those who look at climate and the physical conditions of life as the all-important elements of distribution. But when we bear in mind that almost every species. and in such islands distinct species might have been separately formed without the possibility of intermediate varieties existing in the intermediate zones. By changes in the form of the land and of climate. because an area is now continuous. and finally disappearing. though I do not doubt that the formerly broken condition of areas now continuous. that it has been continuous during a long period. a common alpine species disappears. and as these species are already defined objects.

from inhabiting larger areas. run a greater chance of being exterminated than one existing in large numbers. will have a great advantage over the intermediate variety. sooner than the forms which they originally linked together. we shall have to adapt two varieties to two large areas. It is the same principle which. they should be exterminated and disappear. than will the rarer forms which exist in lesser numbers. For forms existing in larger numbers will have a better chance. accounts for the common species in each 170 . Asa Gray. And it would appear from information given me by Mr. by which two varieties are supposed to be converted and perfected into two distinct species. will tend to beat and supplant the less common forms. Hence. in which they become rather suddenly rarer and rarer. will exist in lesser numbers from inhabiting a narrow and lesser area. of presenting further favourable variations for natural selection to seize on. I have met with striking instances of the rule in the case of varieties intermediate between well-marked varieties in the genus Balanus. as a general rule. if we may trust these facts and inferences. and Mr. the two which exist in larger numbers. as varieties do not essentially differ from species. within any given period. Dr. then. which exists in smaller numbers in a narrow and intermediate zone. as I believe. for these will be more slowly modified and improved. and practically. Wollaston. with a comparatively narrow neutral territory between them. when inhabiting a continuous area. when varieties intermediate between two other forms occur. and a third variety to a narrow intermediate zone. As allied or representative species. then we can understand why intermediate varieties should not endure for very long periods: why. and if we take a varying species inhabiting a very large area. the same rule will probably apply to both. Now. The intermediate variety. they are much rarer numerically than the forms which they connect. are generally distributed in such a manner that each has a wide range. as far as I can make out. in the race for life. this rule holds good with varieties in a state of nature. consequently. and conclude that varieties linking two other varieties together generally have existed in lesser numbers than the forms which they connect. and thus its geographical range will come to be still more sharply defined. 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 already remarked. that generally. But it is a far more important consideration. Watson. For any form existing in lesser numbers would. that during the process of further modification. the more common forms.to utter extermination.

or on the occasional immigration of new inhabitants. in any one region and at any one time. 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. To sum up. presenting on an average a greater number of well-marked varieties than do the rarer species. so that they will no longer be found in a living state. must formerly have existed within each isolated portion of the land. with the new forms thus produced and the old ones acting and reacting on each other. And such new places will depend on slow changes of climate. we ought to see only a few species presenting slight modifications of structure in some degree permanent. areas now continuous must often have existed within the recent period as isolated portions. but these links during the process of natural selection will have been supplanted and exterminated. hilly tract. it is probable. Thirdly. a second to a comparatively narrow. but they 171 . I may illustrate what I mean by supposing three varieties of sheep to be kept.country. when two or more varieties have been formed in different portions of a strictly continuous area. one adapted to an extensive mountainous region. in which many forms. and consequently the improved mountain or plain breed will soon take the place of the less improved hill breed. will come into close contact with each other. as shown in the second chapter. 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. hilly tract. and thus the two breeds. intermediate varieties will. and natural selection can do nothing until favourable individual differences or variations occur. without the interposition of the supplanted. probably. may have separately been rendered sufficiently distinct to rank as representative species. because new varieties are very slowly formed. on some of the old inhabitants becoming slowly modified. So that. more especially among the classes which unite for each birth and wander much. intermediate varieties between the several representative species and their common parent. and this assuredly we do see. and a third to the wide plains at the base. for variation is a slow process. I believe that species come to be tolerably well-defined objects. and. intermediate hill variety. and that the inhabitants are all trying with equal steadiness and skill to improve their stocks by selection. In this case. Secondly. and do not at any one period present an inextricable chaos of varying and intermediate links: first. at first have been formed in the intermediate zones. in a still more important degree. which originally existed in greater numbers.

for these. to exterminate the parent forms and the intermediate links. they will almost certainly be beaten and supplanted by the forms which they connect. which are preserved. and thus be further improved through natural selection and gain further advantages. From this cause alone the intermediate varieties will be liable to accidental extermination. from reasons already assigned (namely from what we know of the actual distribution of closely allied or representative species. linking closely together all the species of the same group. 172 . Lastly. as has been so often remarked. but at all time. For these intermediate varieties will. must assuredly have existed. but the very process of natural selection constantly tends. present more varieties. and during the process of further modification through natural selection. and likewise of acknowledged varieties). in an extremely imperfect and intermittent record.will generally have had a short duration. looking not to any one time. as we shall attempt to show in a future chapter. in the aggregate. Consequently evidence of their former existence could be found only among fossil remains. numberless intermediate varieties. if my theory be true. exist in the intermediate zones in lesser numbers than the varieties which they tend to connect. from existing in greater numbers will.

during summer this animal dives for and preys on fish. to lessen the danger from occasional falls. Here. could a land carnivorous animal have been converted into one with aquatic habits. for. Richardson has remarked. or. by enabling it to escape birds or beasts of prey. and all analogy would lead us to believe that some. and it had been asked how an insectivorous quadruped could possibly have been converted into a flying 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. short legs. 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. We cannot doubt that each structure is of use to each kind of squirrel in its own country. either constant or occasional. or to collect food more quickly. If a different case had been taken. for instance.3. at least. and preys. as there is reason to believe. like other polecats on mice and land animals. let other competing rodents or new beasts of prey immigrate. I can give only one or two instances of transitional habits and structures in allied species. Look at the family of squirrels. 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. Let the climate and vegetation change. as Sir J. how. and from others. which has webbed feet. to the so-called flying squirrels. of the squirrels would decrease in numbers or become 173 . Yet I think such difficulties have little weight. out of the many striking cases which I have collected. it is clear that each must be well adapted to its place in nature. and of diversified habits. but during the long winter it leaves the frozen waters. here we have the finest gradation from animals with their tails only slightly flattened. I lie under a heavy disadvantage. as on other occasions. and form of tail. and flying squirrels have their limbs and even the base of the tail united by a broad expanse of skin. with the posterior part of their bodies rather wide and with the skin on their flanks rather full. or old ones become modified. and which resembles an otter in its fur. the question would have been far more difficult to answer. 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 as each exists by a struggle for life. Look at the Mustela vison of North America. which serves as a parachute and allows them to glide through the air to an astonishing distance from tree to tree. in the same species.

each modification being useful. each grade of structure having been useful to its possessor. as sails. as far as the organs of flight are concerned. like the logger headed duck (Micropterus of Eyton). I can see no difficulty. Now look at the Galeopithecus or so-called flying lemur. we perhaps see traces of an apparatus originally fitted for gliding through the air rather than for flight. Therefore. under the conditions of life to which it is exposed. Although no graduated links of structure. unless they also become modified and improved in structure in a corresponding manner. which was formerly ranked among bats. now connect the Galeopithecus with the other Insectivora. a perfect so-called flying squirrel was produced. but is now believed to belong to the Insectivora. which perhaps may all be the result of disuse. like the apteryx? Yet the structure of each of these birds is good for it. An extremely wide flank-membrane stretches from the corners of the jaw to the tail. This flank. indicate the steps by which birds actually acquired their perfect power of flight. and this. would have converted the animal into a bat. like the penguin. for each has to live by a struggle: but it is not necessarily the best possible under all possible conditions. and that each was developed in the same manner as with the less perfectly gliding squirrels. each being propagated. and seeing that 174 . fitted for gliding through the air. in the continued preservation of individuals with fuller and fuller flank-membranes. like the ostrich. and includes the limbs with the elongated fingers. If about a dozen genera of birds were to become extinct.exterminated. yet there is no difficulty in supposing that such links formerly existed.membrane is furnished with an extensor muscle. but they serve to show what diversified means of transition are at least possible. as fins in the water and as front legs on the land. Seeing that a few members of such water-breathing classes as the Crustacea and Mollusca are adapted to live on the land. until. 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. who would have ventured to surmise that birds might have existed which used their wings solely as flappers. more especially under changing conditions of life. and functionally for no purpose. by the accumulated effects of this process of natural selection. In certain bats in which the wing-membrane extends from the top of the shoulder to the tail and includes the hind-legs.

difficult to decide and immaterial for us. slightly rising and turning by the aid of their fluttering fins. which were gradually rendered more perfect through natural selection. in the individuals of the same species. In either case it would be easy for natural selection to adapt the structure of the animal to its changed habits. In our own country the larger titmouse (Parus major) 175 . both probably often occurring almost simultaneously. Hence the chance of discovering species with transitional grades of structure in a fossil condition will always be less.we have flying birds and mammals. to return to our imaginary illustration of the flying-fish. until their organs of flight had come to a high stage of perfection. and formerly had flying reptiles. or exclusively on artificial substances. flying insects of the most diversified types. which now glide far through the air. who would have ever imagined that in an early transitional state they had been inhabitants of the open ocean. so as to have given them a decided advantage over other animals in the battle for life. we should bear in mind that animals displaying early transitional grades of the structure will seldom have survived to the present day. so far as we know. for they will have been supplanted by their successors. or whether slight modifications of structure lead to changed habits. Furthermore. 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. might have been modified into perfectly winged animals. than in the case of species with fully developed structures. whether habits generally change first and structure afterwards. from their having existed in lesser numbers. It is. hovering over one spot and then proceeding to another. for taking prey of many kinds in many ways. however. and at other times standing stationary on the margin of water. both of diversified and of changed habits. or exclusively to one of its several habits. to escape being devoured by other fish? When we see any structure highly perfected for any particular habit. and then dashing into it like a kingfisher at a fish. it does not seem probable that fishes capable of true flight would have been developed under many subordinate forms. If this had been effected. like a kestrel. Of cases of changed habits it will suffice merely to allude to that of the many British insects which now feed on exotic plants. it is conceivable that flying-fish. on the land and in the water. Thus. and had used their incipient organs of flight exclusively. I will now give two or three instances. Of diversified habits innumerable instances could be given: I have often watched a tyrant flycatcher (Saurophagus sulphuratus) in South America. as the wings of a bird for flight.

however. Petrels are the most aerial and oceanic of birds. in its general habits. and thus breaking them like a nuthatch. Hudson states. the Puffinuria berardi. and a straight. we might expect that such individuals would occasionally give rise to new species. as Mr. On the plains of La Plata. insects in the water. and others with elongated wings which chase insects on the wing. and it makes its nest in holes in banks! In certain other districts. strong beak. would be mistaken by any one for an auk or a grebe. In North America the black bear was seen by Hearne swimming for hours with widely open mouth. almost like a whale. 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. but not so stiff as in the typical wood-peckers. as I can assert. whereas the woodpecker of La Plata has had its structure only slightly 176 . almost like a creeper. where hardly a tree grows. not only from my own observations. this same woodpecker. in all the essential parts of its structure. but with many parts of its organisation profoundly modified in relation to its new habits of life. and undulatory flight. and bores holes in the trunk for its nest. in the quiet sounds of Tierra del Fuego. however. and with their structure either slightly or considerably modified from that of their type. pointed tail-feathers. kills small birds by blows on the head. As we sometimes see individuals following habits different from those proper to their species and to the other species of the same genus.may be seen climbing branches. in certain large districts it does not climb trees. but from those of the accurate Azara. in its astonishing power of diving. frequents trees. The beak. and I have many times seen and heard it hammering the seeds of the yew on a branch. it is essentially a petrel. a long. but. Hence this Colaptes. there is a woodpecker (Colaptes campestris) which has two toes before and two behind. nevertheless. the harsh tone of the voice. thus catching. in its manner of swimming and of flying when made to take flight. And such instances occur in nature. is a woodpecker. its close blood-relationship to our common woodpecker is plainly declared. I may mention as another illustration of the varied habits of this genus.pointed tongue. is not so straight or so strong as in the typical woodpeckers but it is strong enough to bore into wood. yet. sufficiently stiff to support the bird in a vertical position on a post. it sometimes. Even in such trifling characters as the colouring. like a shrike. 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. having anomalous habits.

—using its wings under water and grasping stones with its feet. yet this bird. In the case of the water-ouzel. yet the first is nearly as aquatic as the coot. either in habits or structure. the deeply scooped membrane between the toes shows that structure has begun to change. The water-hen and landrail are members of this order.modified. however different that may be from its own place. has seen the frigate-bird. by examining its dead body. the acutest observer. excepting the genus Proctotrupes. On the other hand. All the members of the great order of Hymenopterous insects are terrestrial. habits have changed without a corresponding change of structure. and the second is nearly as terrestrial as the quail or partridge. subsists by diving. and no one except Audubon. it often enters the water and dives about by the use not of its legs but of its wings. He who believes that each being has been created as we now see it. and many others could be given. which is allied to the thrush family. In such cases. and thus gains an advantage over some other inhabitant of the same country. grebes and coots are eminently aquatic. He who believes in separate and innumerable acts of creation may say. and that if any one being varies ever so little. 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. Hence it will cause him no surprise that there should be geese and frigate-birds with webbed feet. though not in structure. living on the dry land and rarely alighting on the water. it will seize on the place of that inhabitant. will acknowledge that every organic being is constantly endeavouring to increase in numbers. In the frigate-bird. but this seems to me only restating the fact in dignified language. which Sir John Lubbock has discovered to be aquatic in its habits. which has all its four toes webbed. of the Grallatores. 177 . must occasionally have felt surprise when he has met with an animal having habits and structure not in agreement. 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. are formed for walking over swamps and floating plants. yet it exhibits no modification in structure in accordance with its abnormal habits. He who believes in the struggle for existence and in the principle of natural selection. What seems plainer than that the long toes. would never have suspected its sub-aquatic habits. although their toes are only bordered by membrane. not furnished with membrane. and remains as long as four hours beneath the surface. alight on the surface of the ocean. The webbed feet of the upland goose may be said to have become almost rudimentary in function.

living in meadows instead of in swamps. and petrels with the habits of auks.that there should be long-toed corncrakes. that there should be diving thrushes and diving Hymenoptera. 178 . that there should be woodpeckers where hardly a tree grows.

but without any lens or other refractive body. and resting merely on sarcodic tissue. cannot be trusted in science. as some of the lowest organisms in which nerves cannot be detected. and we are forced to look to other species and genera of the same group. endowed with this special sensibility. Reason tells me. but the old saying of Vox populi. hardly concerns us more than how life itself originated. seems. Eyes of the above simple nature are not capable of distinct vision. if further. as is likewise certainly the case. for admitting different amounts of light. Jourdain. and for the correction of spherical and chromatic aberration.4. How a nerve comes to be sensitive to light. When it was first said that the sun stood still and the world turned round. according to M. 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. that is to the collateral descendants from the same parent-form. Organs Of Extreme Perfection And Complication To suppose that the eye with all its inimitable contrivances for adjusting the focus to different distances. the common sense of mankind declared the doctrine false. apparently serving as organs of vision. in order to see what gradations are possible. but I may remark that. are capable of perceiving light. vox Dei. that if numerous gradations from a simple and imperfect eye to one complex and perfect can be shown to exist. absurd in the highest degree. we ought to look exclusively to its lineal progenitors. though insuperable by our imagination. but this is scarcely ever possible. however. and serve only to distinguish 179 . and if such variations should be useful to any animal under changing conditions of life. should not be considered as subversive of the theory. We may. as every philosopher knows. 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. But the state of the same organ in distinct classes may incidentally throw light on the steps by which it has been perfected. The simplest organ which can be called an eye consists of an optic nerve. as is certainly the case. I freely confess. In searching for the gradations through which an organ in any species has been perfected. could have been formed by natural selection. surrounded by pigment-cells and covered by translucent skin. it does not seem impossible that certain sensitive elements in their sarcode should become aggregated and developed into nerves. without any nerves. each grade being useful to its possessor.

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


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


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?


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


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.


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


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.


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


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


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


of the hind legs. it is improbable in the highest degree that their common progenitor should have been adapted for breathing air. and in the several groups they are formed by the modification of the fore legs. These organs must have been independently developed. the nature of the organism and that of the surrounding conditions. belonging to distinct sub-families and families. as they could not have been inherited from a common progenitor. of the maxillae or lips. viz. But as the vast majority of the species in the above two families. Consequently natural selection would have had different materials or variations to work on. from belonging to distinct families. which are furnished with hair-claspers. closely similar. their variability assuredly would not have been exactly the same. by the most diversified means. has argued in the same manner. He shows that there are parasitic mites (Acaridae). and he found it to differ in each in several important points. For these species. it is a common rule throughout nature that the same end should be gained. are aquatic in their habits.for by inheritance from a common progenitor. and might even have been expected. we see the same end gained and the same function performed. Muller was thus led carefully to examine the apparatus in the air-breathing species. even sometimes in the case of closely related beings. How differently constructed is the feathered wing of a bird and the membrane-covered wing of a bat. the late Professor Claparede. as well as most other crustaceans. In the foregoing cases. and the structures thus acquired would almost necessarily have differed. in the manner in which they are opened and closed. on the supposition that species belonging to distinct families had slowly become adapted to live more and more out of water. On the other hand. Now such differences are intelligible. as in the position of the orifices. and to breathe the air. by organs in appearance. but on what 190 . the two wings of a fly.. This line of argument seems to have had great weight in leading Fritz Muller to accept the views maintained by me in this volume. would have differed to a certain extent. Another distinguished zoologist. and the two wings with the elytra of a beetle. and in some accessory details. in order to arrive at the same functional result. Bivalve shells are made to open and shut. and of appendages on the under side of the hind part of the body. On the hypothesis of separate acts of creation the whole case remains unintelligible. and still more so the four wings of a butterfly. though not in development. and in accordance with the principle that the nature of each variation depends on two factors. in beings not at all or only remotely allied. and has arrived at the same result.

though hermaphrodites. With several kinds this is effected by the pollen-grains. all for the same purpose and effected in essentially the same manner. Cruger saw crowds of large 191 . and these carry the pollen from the anthers to the stigma. and rendered nutritious. and sometimes capable of neatly adapted movements through irritability or elasticity. but such a view of nature is incredible. The basal part of the labellum stands over the bucket. sometimes forming trap-like contrivances. Cruger in the Coryanthes. This orchid has part of its labellum or lower lip hollowed out into a great bucket. With plants having separated sexes. and is consequently visited by insects. An almost equally simple. by being embedded in pulp or flesh. the pollen does not spontaneously fall on the stigma. and with those in which. and by being furnished with wings and plumes.a number of patterns is the hinge constructed. The nectar may be stored in variously shaped receptacles. from the long row of neatly interlocking teeth in a Nucula to the simple ligament of a Mussel! Seeds are disseminated by their minuteness. but entailing changes in every part of the flower. From such structures we may advance till we come to such a case of extraordinary adaptation as that lately described by Dr. But Dr. as different in shape as they are elegant in structure. if he had not witnessed what takes place. into which drops of almost pure water continually fall from two secreting horns which stand above it. so as to attract and be devoured by birds. by their capsule being converted into a light balloon-like envelope. formed of the most diverse parts. which are light and incoherent. From this simple stage we may pass through an inexhaustible number of contrivances. within this chamber there are curious fleshy ridges. as well as conspicuously coloured. The most ingenious man. almost like toys in a shop. and this is the simplest plan which can well be conceived. Some authors maintain that organic beings have been formed in many ways for the sake of mere variety. the water overflows by a spout on one side. so as to adhere to the fur of quadrupeds. could never have imagined what purpose all these parts serve. and when the bucket is half-full. and is itself hollowed out into a sort of chamber with two lateral entrances. so as to be wafted by every breeze. being blown by the wind through mere chance on to the stigma. some aid is necessary for their fertilisation. by having hooks and grapnels of many kinds and serrated awns. with the stamens and pistils modified in many ways. though very different plan occurs in many plants in which a symmetrical flower secretes a few drops of nectar. I will give one other instance: for this subject of the same end being gained by the most diversified means well deserves attention.

transmits a sensation or vibration to a certain membrane which is instantly ruptured. When the bee. The construction of the flower in another closely allied orchid. in doing this they inevitably touch a long. first rubs its back against the viscid stigma and then against the viscid glands of the pollenmasses. flies to another flower. with a pollen-mass still fastened to its back. and forces them to crawl out through the spout. Cruger saw a "continual procession" of bees thus crawling out of their involuntary bath. Dr. as I have called it. of the water-secreting horns of the bucket half-full of water. the pollen-mass necessarily comes first into contact with the viscid stigma. and is roofed over by the column. like an arrow. but to gnaw off the ridges within the chamber above the bucket. like those of the Coryanthes. is widely different. in order to gnaw the labellum. Dr. as already remarked. or. namely. Bees visit these flowers. and their wings being thus wetted they could not fly away. can we understand the graduated scale of complexity and the multifarious means for gaining the same end. the Catasetum. it may be asked. This antenna. and are thus carried away. the variability will not be of the same 192 . thus provided. How. The passage is narrow. in the right direction. Cruger sent me a flower in spirits of wine. and the flower is fertilised. but were compelled to crawl out through the passage formed by the spout or overflow. which prevents the bees from flying away. The answer no doubt is. which already differ from each other in some slight degree. so that a bee.humble-bees visiting the gigantic flowers of this orchid. or to the same flower a second time. Now at last we see the full use of every part of the flower. 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 antenna. in doing this they frequently pushed each other into the bucket. with a bee which he had killed before it had quite crawled out. in forcing its way out. though serving the same end. thus effecting fertilisation. sensitive projection. tapering. and rub against the properly placed viscid pollen-masses and the viscid stigma. and adheres to it. not in order to suck nectar. and adheres by its viscid extremity to the back of the bee. that when two forms vary. 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. this sets free a spring by which the pollen-mass is shot forth. where it is brought into contact with the stigma. and retain the pollen. in the foregoing and in innumerable other instances. and is equally curious. and is pushed by its comrades into the bucket and then crawls out by the passage. when touched. which is viscid enough to break certain elastic threads.

as indeed is shown by that old. we can clearly understand why she should not. towards which no transitional grade is known to lead. that new organs appearing as if created for some special purpose rarely or never appear in any being. Finally. she can never take a great and sudden leap. then. "Nature is prodigal in variety. We should also bear in mind that every highly developed organism has passed through many changes. considering how small the proportion of living and known forms is to the extinct and unknown. 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. but somewhat exaggerated. on the theory of Creation. the structure of each part of each species. as Milne Edwards has well expressed it.exact nature. but may be again and again further altered. 193 . Hence. each supposed to have been separately created for its own proper place in nature. I have been astonished how rarely an organ can be named. through which the species has passed during its successive adaptations to changed habits and conditions of life." Why. and that each modified structure tends to be inherited. should there be so much variety and so little real novelty? Why should all the parts and organs of many independent beings. although in many cases it is most difficult even to conjecture by what transitions organs could have arrived at their present state. and consequently the results obtained through natural selection for the same general purpose will not be the same. but must advance by the short and sure." We meet with this admission in the writings of almost every experienced naturalist. for whatever purpose it may serve. is the sum of many inherited changes. It is certainly true. but niggard in innovation. yet. though slow steps. or. for natural selection acts only by taking advantage of slight successive variations. so that each modification will not readily be quite lost. canon in natural history of "Natura non facit saltum.

yet we should pause before being too positive even in this case. and it seems at first incredible that this could have been adapted for its present purpose by successive slight modifications. The tail of the giraffe looks like an artificially constructed fly-flapper. its general presence and use for many purposes in so many land animals. It is not that the larger quadrupeds are actually destroyed (except in some rare cases) by flies. though of a very different kind. and by the destruction of the less well-fitted individuals. for so trifling an object as to drive away flies. from being correlated with constitutional differences. would be able to range into new pastures and thus gain a great advantage. by the survival of the fittest. each better and better fitted. but any actually injurious deviations in their structure would of course have been checked by natural selection. may perhaps be thus accounted for. A well-developed tail having been formed in an aquatic animal. which. and. In the first place. Organs now of trifling importance have probably in some cases been of high importance to an early progenitor. after having been slowly perfected at a former period. 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. As Affected By Natural Selection As natural selection acts by life and death. so that they are more subject to disease. it might subsequently come to be worked in for all sorts 194 . might assuredly be acted on by natural selection.7. or to escape from beasts of prey. although now of very slight use. In a former chapter I have given instances of very trifling characters. which in their lungs or modified swim-bladders betray their aquatic origin. or not so well enabled in a coming dearth to search for food. but they are incessantly harassed and their strength reduced. the colour of the skin and hair of quadrupeds. Organs Of Little Apparent Importance. or from determining the attacks of insects. 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. such as the down on fruit and the colour of its flesh. I have sometimes felt great difficulty in understanding the origin or formation of parts of little importance. have been transmitted to existing species in nearly the same state. Seeing how important an organ of locomotion the tail is in most aquatic animals. almost as great. as in the case of the most perfect and complex organs.

In the second place. comprehension..of purposes. 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. of the tendency to reversion to long-lost characters. as it is. and finally of sexual selection. but as sutures occur in the skulls of young birds and reptiles. The naked skin on the head of a vulture is generally considered as a direct adaptation for wallowing in putridity. may subsequently have been taken advantage of by its modified descendants. and had been acquired through natural selection. although at first of no advantage to a species. as a fly-flapper. so the spikes on the palm may at first have been developed for this object. an organ of prehension. or may be indispensable for this act. If green woodpeckers alone had existed. and we did not know that there were many black and pied kinds. and which. but we should be very cautious in drawing any such inference. for the hare. by which characters of use to one sex are often gained and then transmitted more or less perfectly to the other sex. though the aid in this latter respect must be slight. serve as a defence against browsing quadrupeds. as there is reason to believe from the distribution of the thorn. the colour is probably in chief part due to sexual selection. no doubt.bearing species in Africa and South America. and this contrivance. and no doubt they facilitate. we may easily err in attributing importance to characters. But structures thus indirectly gained. The sutures in the skulls of young mammals have been advanced as a beautiful adaptation for aiding parturition. and so it may be. with hardly any tail. or it may possibly be due to the direct action of putrid matter. is of the highest service to the plant. and consequently that it was a character of importance. such as of correlation. which have only to 195 . when we see that the skin on the head of the clean-feeding male turkey is likewise naked. We must by no means overlook the effects of the definite action of changed conditions of life. and in believing that they have been developed through natural selection. as it underwent further modification and became a climber. of the complex laws of growth. under new conditions of life and newly acquired habits. but as we see nearly similar hooks on many trees which are not climbers. etc. as in the case of the dog. can double still more quickly. of the pressure of one part on another. of so-called spontaneous variations. which seem to depend in a quite subordinate degree on the nature of the conditions. 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. or as an aid in turning. and subsequently have been improved and taken advantage of by the plant. though of no use to the sex.

together with abundant food. and are exposed to a certain extent to natural selection. With cattle susceptibility to the attacks of flies is correlated with colour. as we have good reason to believe. is apparently one chief cause of the great modification which the breeds of swine have undergone. which nevertheless are generally admitted to have arisen through ordinary generation from one or a few parent-stocks. we ought not to lay too much stress on our ignorance of the precise cause of the slight analogous differences between true species. if we are unable to account for the characteristic differences of our several domestic breeds. and we are immediately made conscious of this by reflecting on the differences between the breeds of our domesticated animals in different countries. and then by the law of homologous variation. to increase the size of the chest. more especially in the less civilized countries. on the whole organisation is probably still more important. and individuals with slightly different constitutions would succeed best under different climates. and possibly even the form of the pelvis. Animals kept by savages in different countries often have to struggle for their own subsistence. Some observers are convinced that a damp climate affects the growth of the hair. and this. and that with the hair the horns are correlated. The shape. as is the liability to be poisoned by certain plants. and again correlation would come into play. where there has been but little methodical selection. as H. also.escape from a broken egg. so that even colour would be thus subjected to the action of natural selection. The laborious breathing necessary in high regions tends. the front limbs and the head would probably be affected. and I have made these remarks only to show that. and has been taken advantage of in the parturition of the higher animals. of the pelvis might affect by pressure the shape of certain parts of the young in the womb. and a mountainous country would probably affect the hind limbs from exercising them more. we may infer that this structure has arisen from the laws of growth. We are profoundly ignorant of the cause of each slight variation or individual difference. 196 . von Nathusius has lately shown in his excellent Treatise. The effects of lessened exercise. Mountain breeds always differ from lowland breeds. But we are far too ignorant to speculate on the relative importance of the several known and unknown causes of variation.

Thus. But a still more important consideration is that the chief part of the organisation of every living creature is due to inheritance. It is scarcely possible to decide how much allowance ought to be made for such causes of change. were originally developed. though each being assuredly is well fitted for its place in nature. horse and bat. as the definite action of external conditions. independently of any advantage thus gained.8. so-called spontaneous variations. or for the sake of mere variety. or was formerly. to delight man or the Creator (but this latter point is beyond the scope of scientific discussion). and the complex laws of growth. we may conclude that the structure of every living creature either now is. we can hardly believe that the webbed feet of the upland goose. and we may further venture to believe that the several bones in the limbs of the monkey. many structures have now no very close and direct relation to present habits of life. No doubt the definite action of changed conditions. They believe that many structures have been created for the sake of beauty. 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. as they now are to the most aquatic of living birds. a view already discussed. on the principle of utility. would be absolutely fatal to my theory. and the various causes of modifications. are of special use to these animals. or of the frigate. have all produced an effect. and in the flipper of the seal. So we may believe that the progenitor of the seal did not possess a flipper. but with these important exceptions. Utilitarian Doctrine.bird. but a foot with five toes fitted for walking or grasping. and consequently. How Far True: Beauty. 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 197 . probably a great effect. Such doctrines. in the fore leg of the horse. in the wing of the bat. we cannot believe that the similar bones in the arm of the monkey. lately specified. I fully admit that many structures are now of no direct use to their possessors. but this does not prove that they were formed solely for beauty or variety. probably through the reduction of more numerous bones in the fin of some ancient fish-like progenitor of the whole class. if true. are of special use to these birds. We may safely attribute these structures to inheritance. But webbed feet no doubt were as useful to the progenitor of the upland goose and of the frigate-bird. and may never have been of any use to their progenitors.

our plants would not have been decked with beautiful flowers. is not innate or unalterable. and never visited by insects. not coloured. A similar line of argument holds good with fruits. Flowers rank among the most beautiful productions of nature. I willingly admit that a great number of male animals. but would have produced only such poor flowers as we see on our fir. and that the idea of what is beautiful. Were the beautiful volute and cone shells of the Eocene epoch.subversive of my whole theory—I may first remark that the sense of beauty obviously depends on the nature of the mind. docks and nettles. in order that the fruit may be devoured and the matured seeds disseminated. On the other hand. or rendered conspicuous by being white or black. oak. on grasses. but they have been rendered conspicuous in contrast with the green leaves. which are all fertilised through the agency of the wind. 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. if it be coloured of any brilliant tint. we may conclude that. But this beauty serves merely as a guide to birds and beasts. 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. nut and ash trees. in the men of different races admiring an entirely different standard of beauty in their women. reptiles. and in many others. Several plants habitually produce two kinds of flowers. and the gracefully sculptured ammonites of the Secondary period. If beautiful objects had been created solely for man's gratification. spinach. for instance. as all our most gorgeous birds. irrespective of any real quality in the admired object. and in consequence at the same time beautiful. some fishes. 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. 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). so that they may be easily observed by insects. 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. destitute of nectar. one kind open and coloured so as to attract insects. 198 . is apparently wholly due to symmetry of growth. and mammals. Hence. if insects had not been developed on the face of the earth. the other closed. We see this.

it would annihilate my theory. have been rendered beautiful for beauty's sake. the reception of a peculiar kind of pleasure from certain colours. forms and sounds—was first developed in the mind of man and of the lower animals. the cause apparently lies in the colours acquired through sexual selection having been transmitted to both sexes. but some authors suppose that at the same time it is furnished with a rattle for its own injury. and in the ovipositor of the ichneumon. But natural selection can and does often produce structures for the direct injury of other animals. that is. How the sense of beauty in its simplest form—that is. by which its eggs are deposited in the living bodies of other insects. instead of to the males alone. in order to alarm the many birds and beasts which are known to attack even the most venomous species. in order to warn the doomed mouse. by the more beautiful males having been continually preferred by the females. It is a much more probable view that the rattlesnake uses its rattle. Habit in all these cases appears to have come to a certain extent into play. for such could not have been produced through natural selection. is a very obscure subject. Although many statements may be found in works on natural history to this effect. though throughout nature one species incessantly takes advantage of. and not for the delight of man. Snakes act on the same principle which makes the hen ruffle her feathers and expand her wings when a dog 199 . So it is with the music of birds. and others displeasure. but there must be some fundamental cause in the constitution of the nervous system in each species. I cannot find even one which seems to me of any weight. namely. as we see in the fang of the adder. Natural selection cannot possibly produce any modification in a species exclusively for the good of another species. It is admitted that the rattlesnake has a poison-fang for its own defence and for the destruction of its prey. 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 would almost as soon believe that the cat curls the end of its tail when preparing to spring. When the female is as beautifully coloured as the male. The same sort of difficulty is presented if we enquire how it is that certain flavours and odours give pleasure. But this has been effected through sexual selection. 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. which is not rarely the case with birds and butterflies.and a host of magnificently coloured butterflies. and profits by the structures of others. to warn its prey. the cobra expands its frill and the puff-adder swells while hissing so loudly and harshly.

as a boring and serrated instrument. owing to the backward serratures. for natural selection acts solely by and for the good of each. it will be modified. The correction for the aberration of light is said by Muller not to be perfect even in that most perfect organ. Helmholtz. nor do we always meet. whose judgment no one will dispute. the human eye. as having existed in a remote progenitor. with this high standard under nature. Natural selection tends only to make each organic being as perfect as." If our reason leads us to admire with enthusiasm a multitude of inimitable contrivances in nature. Natural selection will not produce absolute perfection. after describing in the strongest terms the wonderful powers of the human eye. or if it be not so. cannot be withdrawn. No organ will be formed. If a fair balance be struck between the good and evil caused by each part. The endemic productions of New Zealand. as far as we can judge. And we see that this is the standard of perfection attained under nature. is as nothing in comparison with the incongruities which we have just come across in the domain of the sensations. but they are now rapidly yielding before the advancing legions of plants and animals introduced from Europe. if any part comes to be injurious. as myriads have become extinct. this same reason tells us. for the purpose of causing pain or for doing an injury to its possessor. 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. for instance. under changing conditions of life. which. or slightly more perfect than the other inhabitants of the same country with which it comes into competition. and that it has since been modified but not perfected for its present purpose. are perfect. when used against many kinds of enemies. each will be found on the whole advantageous. But I have not space here to enlarge on the many ways by which animals endeavour to frighten away their enemies. Can we consider the sting of the bee as perfect. Natural selection will never produce in a being any structure more injurious than beneficial to that being. that some other contrivances are less perfect. though we may easily err on both sides. as Paley has remarked. one compared with another. with the poison originally adapted for 200 . like that in so many members of the same great order. After the lapse of time.approaches her chickens. and thus inevitably causes the death of the insect by tearing out its viscera? If we look at the sting of the bee. the being will become extinct. 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.

since intensified. can we consider as equally perfect the elaboration of dense clouds of pollen by our fir-trees. but we ought to admire the savage instinctive hatred of the queen-bee. which are utterly useless to the community for any other purpose. as soon as they are born. 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. so that a few granules may be wafted by chance on to the ovules? 201 . her daughters. If we admire the truly wonderful power of scent by which the males of many insects find their females. 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. it will fulfil all the requirements of natural selection. can we admire the production for this single purpose of thousands of drones. and which are ultimately slaughtered by their industrious and sterile sisters? It may be difficult.some other object. though it may cause the death of some few members. though the latter fortunately is most rare. such as to produce galls. or to perish herself in the combat. and maternal love or maternal hatred.

bearing in mind that each organic being is trying to live wherever it can live. if we know of a long series of gradations in complexity. the intermediate variety will usually exist in lesser numbers than the two forms which it connects. Hence we can understand. Closely allied species. fitted for an intermediate zone. will have a great advantage over the less numerous intermediate variety. and when the conditions of life did not insensibly graduate away from one part to another. and at any one time acts only on a few forms. 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. partly because the process of natural selection is always very slow. but from reasons assigned.9. and partly because the very process of natural selection implies the continual supplanting and extinction of preceding and intermediate gradations. and will thus generally succeed in supplanting and exterminating it. each good for its possessor. from existing in greater numbers. during the course of further modification. and are not linked together by a multitude of intermediate gradations. which on the belief of independent acts of creation are utterly obscure. Many of them are serious. We have seen that species at any one period are not indefinitely variable. then under changing conditions of life. how it has arisen that there are upland geese with webbed feet. that a bat. is enough to stagger any one. there is no logical impossibility in the acquirement of any conceivable degree 202 . When two varieties are formed in two districts of a continuous area. diving thrushes. We have seen that a species under new conditions of life may change its habits. now living on a continuous area. but I think that in the discussion light has been thrown on several facts. Although the belief that an organ so perfect as the eye could have been formed by natural selection. or it may have diversified habits. 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. consequently the two latter. yet in the case of any organ. with some very unlike those of its nearest congeners. must often have been formed when the area was not continuous. ground woodpeckers. for instance. could not have been formed by natural selection from an animal which at first only glided through the air. and petrels with the habits of auks. an intermediate variety will often be formed.

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.of perfection through natural selection. been subsequently taken advantage of. independently of any good having been thus gained. a swim-bladder has apparently been converted into an air-breathing lung. have been acquired by means of natural selection. often yield to 203 . though it has become of such small importance that it could not. organs. In the cases in which we know of no intermediate or transitional states. that modifications in its structure could not have been slowly accumulated by means of natural selection. Natural selection can produce nothing in one species for the exclusive good or injury of another. In many other cases. but in all cases at the same time useful to the possessor. Hence the inhabitants of one country. for the metamorphoses of many organs show what wonderful changes in function are at least possible. The same organ having performed simultaneously very different functions. But even such structures have often. also. generally the smaller one. and two distinct organs having performed at the same time the same function. In each well-stocked country natural selection acts through the competition of the inhabitants and consequently leads to success in the battle for life. For instance. and excretions highly useful or even indispensable. the common rule throughout nature is infinite diversity of structure for gaining the same end. and then having been in part or in whole specialised for one function. On the other hand. We may. and still further modified. must often have largely facilitated transitions. in its present state. or highly injurious to another species. we should be extremely cautious in concluding that none can have existed. modifications are probably the direct result of the laws of variation or of growth. essential differences in their structure can almost always be detected. 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 may well produce parts. the one having been perfected whilst aided by the other. and this naturally follows from the principle of natural selection. only in accordance with the standard of that particular country. and this again naturally follows from the same great principle. as we may feel assured. We have seen that in two beings widely remote from each other in the natural scale. organs serving for the same purpose and in external appearance closely similar may have been separately and independently formed. for the good of species under new conditions of life. but when such organs are closely examined.

is fully embraced by the principle of natural selection. as it includes.the inhabitants of another and generally the larger country. through the inheritance of former variations and adaptations. "Natura non facit saltum. unity of type is explained by unity of descent. nor. that of Unity of Type. and thus the standard of perfection will have been rendered higher. Hence. is not strictly correct. as far as we can judge by our limited faculties. The expression of conditions of existence. On the theory of natural selection we can clearly understand the full meaning of that old canon in natural history. and subjected in all cases to the several laws of growth and variation. if we look to the present inhabitants alone of the world. and more diversified forms. the law of the Conditions of Existence is the higher law. Natural selection will not necessarily lead to absolute perfection. 204 . It is generally acknowledged that all organic beings have been formed on two great laws—Unity of Type. in fact. and the competition will have been severer." This canon. and the Conditions of Existence. For in the larger country there will have existed more individuals. so often insisted on by the illustrious Cuvier. can absolute perfection be everywhere predicated. it must on this theory be strictly true. By unity of type is meant that fundamental agreement in structure which we see in organic beings of the same class. being affected by the direct action of external conditions of life. On my theory. whether known or unknown. For natural selection acts by either now adapting the varying parts of each being to its organic and inorganic conditions of life. but if we include all those of past times. or by having adapted them during past periods of time: the adaptations being aided in many cases by the increased use or disuse of parts. and which is quite independent of their habits of life.

Chapter 7 Miscellaneous Objections To The Theory Of Natural Selection 205 .

E.Longevity — Modifications not necessarily simultaneous — Modifications apparently of no direct service — Progressive development — Characters of small functional importance. And these conditions have. have remained so. 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. 206 . Thus a distinguished German naturalist has asserted that the weakest part of my theory is. when their conditions changed. Ray Lankester has recently discussed this subject. so that he who believes in natural selection "must arrange his genealogical tree" in such a manner that all the descendants have longer lives than their progenitors! Cannot our critics conceive that a biennial plant or one of the lower animals might range into a cold climate and perish there every winter. have undergone many mutations. as some of the previous discussions may thus be made clearer. as well as to the amount of expenditure in reproduction and in general activity. 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. with some parade of mathematical accuracy. developed from one and the same source — Reasons for disbelieving in great and abrupt modifications. as many have been made by writers who have not taken the trouble to understand the subject. I will devote this chapter to the consideration of various miscellaneous objections which have been advanced against my views. 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. unless they themselves likewise changed. as far as its extreme complexity allows him to form a judgment. as well as the number and kinds of its inhabitants. that longevity is a great advantage to all species. that longevity is generally related to the standard of each species in the scale of organisation. A critic has lately insisted. and he concludes. owing to advantages gained through natural selection. it is probable. that all are not as perfect as they might have been in relation to their conditions. even if they were at any one time perfectly adapted to their conditions of life. and yet. and no one will dispute that the physical conditions of each country. but it would be useless to discuss all of them. been largely determined through natural selection. Nor can organic beings.

at the close of his German translation of this work. as far as I can discover. dry or moist districts. the conditions of life. would have been of some avail against those who believe in an innate and necessary law of development. during the last several thousand years. have remained absolutely uniform. their varieties seem to be generally confined to distinct regions. The many animals which have remained unchanged since the commencement of the glacial period. but in many parts. if slight. as was formerly remarked. first in one part and then in another. but this will be effected only under certain favourable circumstances. would have been an incomparably stronger case. The fact of little or no modification having been effected since the glacial period. for the ancient domestic races figured on the Egyptian monuments. might. albinism. inhabiting distinct stations. be acquired by successive variations. G. these will be preserved. they would appear to us as if 207 . so probably have none in any part of the world. The celebrated palaeontologist. as far as we know. this line of argument proves too much. Bronn also insists that distinct species never differ from each other in single characters. excellently adapted for some purpose. Lewes has remarked. in the case of animals which wander much about and cross freely. have changed during the last three or four thousand years. they might live together. such as size. asks how. Moreover. in Egypt. in which the variability seems to be of a peculiar nature. or embalmed. and all mere temporary variations. whereas.H.. as none of the animals and plants of Egypt. of which we know anything. 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. such as high land or low land. the more permanent varieties are generally found. are closely similar or even identical with those now living. and as they would be transmitted all together. Bronn.It has been argued that. as Mr. on the principle of natural selection. yet all naturalists admit that such races have been produced through the modification of their original types. etc. The most striking modifications. for these have been exposed to great changes of climate and have migrated over great distances. But. which implies that when variations or individual differences of a beneficial nature happen to arise. and he asks. but is powerless against the doctrine of natural selection or the survival of the fittest. can a variety live side by side with the parent species? If both have become fitted for slightly different habits of life or conditions. and if we lay on one side polymorphic species.

and so forth —as well as through other causes which lead to the many mysterious cases of correlation. fruit. this subject has been discussed by Nageli in an admirable essay. so will be other parts. has been greatly changed. and therefore cannot have been influenced through natural selection. when not of any use for dissemination. Nevertheless. it should always be borne in mind that when one part is modified. 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. There is much force in the above objection. the position of the ovules. 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. which appear to be quite unimportant for the welfare of the species. almost all the other parts have been slightly modified. whether it be the flower. Look at the race and dray-horse. but he insists that the families of plants differ chiefly from each other in morphological characters. which we do not in the least understand. or have formerly been. in the first place. 208 . but first one part and then another slightly modified and improved. 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. of use to each species. such as an increased or diminished flow of nutriment to a part. through certain dimly seen causes. chiefly through man's power of selection. Their whole frames. With respect to plants. or at the greyhound and mastiff. He admits that natural selection has effected much. for some special purpose. however. or leaves. and of the leaves on the axis. have been modified. we ought. to be extremely cautious in pretending to decide what structures now are. to the above objection is afforded by those domestic races which have been modified. as cases in which natural selection could not have acted.they had been simultaneously developed. He specifies the arrangement of the cells in the tissues. and a multitude of analogous cases. He consequently believes in an innate tendency towards progressive and more perfect development. and even their mental characteristics. namely. In the second place. etc. This may be attributed partly to the principle of correlated growth. and recently by Broca. To these may be added the numerical divisions in the parts of the flower. that many characters appear to be of no service whatever to their possessors. mutual pressure. A much more serious objection has been urged by Bronn. an early developed part affecting one subsequently developed. The best answer. the shape of the seed. and partly to so-called spontaneous variation.

also.These agencies may be all grouped together. In the third place. There must be some efficient cause for each slight individual difference. which a 209 . offer good instances of spontaneous variations. in which the nature of the conditions apparently plays a quite subordinate part. and its use would be much influence by its length. presently see that the tail is a highly useful prehensile organ to some of the species. With respect to plants. we have to allow for the direct and definite action of changed conditions of life. it is almost certain that all the individuals of the species would be similarly modified. excited so much surprise in the minds of the older naturalists. hence the length of the ears can hardly be quite unimportant. and if the unknown cause were to act persistently. it is hardly necessary to observe that even in the higher and best-known animals many structures exist. or of a nectarine on a peach-tree. 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. due to some change in the conditions. according to Dr. I can no more believe in this than that the well-adapted form of a racehorse or greyhound. under the expression of the laws of growth. for the sake of brevity. of differences in structure which can be of no special use. Bud-variations. It may be worth while to illustrate some of the foregoing remarks. Schobl. so that they no doubt serve as tactile organs. which are so highly developed that no one doubts that they are of importance. As Bronn gives the length of the ears and tail in the several species of mice as instances. In the earlier editions of this work I underrated. the external ears of the common mouse are supplied in an extraordinary manner with nerves. or has only recently been. ascertained. to which on account of Nageli's essay I shall confine myself in the following remarks. can thus be explained. We shall. With respect to the assumed inutility of various parts and organs. which before the principle of selection by man was well understood. and for so-called spontaneous variations. though trifling ones. such as the appearance of a moss-rose on a common rose. it will be admitted that the flowers of the orchids present a multitude of curious structures. yet their use has not been. the frequency and importance of modifications due to spontaneous variability. as it now seems probable. 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. as well as for more strongly marked variations which occasionally arise. but even in these cases. I may mention that. if we bear in mind the power of a minute drop of poison in producing complex galls.

one ovule holds the former and a second ovule the latter position. The ordinary and open flowers can be intercrossed. and the benefits which certainly are derived from this process are thus secured. In six out of thirty of the closed flowers in an Indian violet (name unknown.few years ago would have been considered as mere morphological differences without any special function. with the expenditure of wonderfully little pollen. as just stated. The closed and imperfect flowers are. and in others they are suspended. and have probably been gained through natural selection. the other closed and imperfect. are still further modified. for the plants have never produced with me perfect flowers). and this stamen is not present in the 210 . de Jussieu. two retaining their proper function. for the five stamens which stand opposite to the sepals are all aborted. and their arrangement. according to A. and in some species of Viola three stamens are in this state. but they are now known to be of the highest importance for the fertilisation of the species through the aid of insects. In Ononis columnae five of the alternate stamens are rudimentary. but now we know this to be the case. the sepals are reduced from the normal number of five to three. No one until lately would have imagined that in dimorphic and trimorphic plants the different lengths of the stamens and pistils. and the production of seed. These positions seem at first purely morphological. but Dr. and in others the lower ones alone are fertilised. of the ordinary structure. These two kinds of flowers sometimes differ wonderfully in structure. Several plants belonging to distinct orders habitually produce flowers of two kinds—the one open. and the pollen-grains are reduced in diameter. and within the same ovarium of some few plants. or of no physiological signification. Hooker informs me that within the same ovarium the upper ovules alone in some cases. even when one is erect and the other suspended within the same ovarium. would follow the selection of any slight deviations in position which favoured their fertilisation. a sixth stamen standing opposite to a petal being alone developed. If so. In one section of the Malpighiaceae the closed flowers. as they yield with the utmost safety a large stock of seed. The petals in the imperfect flowers almost always consist of mere rudiments. but being of very small size. could have been of any service. manifestly of high importance. In certain whole groups of plants the ovules stand erect. in structure. and he suggests that this probably depends on the direction in which the pollen-tubes enter the ovarium. yet may be seen to graduate into each other on the same plant. the position of the ovules. The two kinds of flowers often differ much. however.

and in certain fir. as far as we can judge. and the central one coelospermous. while the surrounding flowers generally have three calyx-lobes with the other organs pentamerous. In the British Adoxa the uppermost flower generally has two calyx-lobes with the other organs tetramerous. In the common rue and some other plants. yet hardly any of the above special modifications can have been thus determined. opens first. It is so necessary to appreciate the important effects of the laws of growth. that the achenes or seeds of the circumference and centre sometimes differ greatly in form. with the exception of that of the well-developed ray. that I will give some additional cases of another kind. but must have followed from the laws of growth.florets. and the ovaria are reduced from three to two. colour and other characters. In certain Umbelliferae the exterior seeds. and in Hyoseris the same head yields achenes of three different forms.trees. and to reduce the amount of pollen. due to differences in relative position on the same plant. ribbed. previously referred to. are orthospermous. natural selection cannot. and five divisions to the ovarium. have come into play. during the progress of the reduction of the pollen and the closure of the flowers. and has five sepals and petals. lanceolate. Professor Braun mentions a Fumariaceous genus. In many Compositae and Umbelliferae (and in some other plants) the circumferential flowers have their corollas much more developed than those of the centre. according to Schacht. and it can hardly be doubted that if 211 . according to Tausch. two-valved and two-seeded siliques. usually the central or terminal one. and this seems often connected with the abortion of the reproductive organs. in the nearly horizontal and in the upright branches. which are of service in making the flowers conspicuous to insects. Now although natural selection may well have had the power to prevent some of the flowers from expanding. and this is a character which was considered by De Candolle to be in other species of the highest systematic importance. the angles of divergence of the leaves differ. In Carthamus and some other Compositae the central achenes alone are furnished with a pappus. the style is aborted. All these modifications follow from the relative position and inter-action of the parts. or only in a quite subordinate manner. in which the flowers in the lower part of the spike bear oval. when rendered by the closure of the flowers superfluous. one flower. In these several cases. while all the other flowers on the plant are tetramerous.ordinary flowers of this species. It is a more curious fact. namely of differences in the same part or organ. including the functional inactivity of parts. and in the upper part of the spike. one-seeded nutlets. In the Spanish chestnut.

that the plants have been caught in the act of progressing towards a higher state of 212 . flowers indifferently tetramerous. s'etend entre le pericarpe et le placenta. I may mention that. I will give only a few instances. mutabile. Of their cause we are quite ignorant. to any proximate agency. they cannot have been influenced by natural selection. But with respect to Nageli's doctrine of an innate tendency towards perfection or progressive development. to which latter tribe the genus belongs. that I need not give examples. with either one or two ovaries. St.. can it be said in the case of these strongly pronounced variations. all would have been modified in the same manner. etc. "Voila donc dans un meme individu des loges et un style qui se rattachent tantot a un axe verticale et tantot a un gynobase. such as relative position. independently of natural selection." We thus see that with plants many morphological changes may be attributed to the laws of growth and the inter-action of parts. or occurring on distinct plants. but in some species flowers may be found on the same plant. 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. as are the flowers and leaves in certain positions. but Professor Asa Gray states that with some species of Mimulus. Hilaire found towards the southern extreme of the range of Gomphia oleaeformis two forms which he did not at first doubt were distinct species. and he then adds. Hilaire gives the following cases: the genus Zanthoxylon belongs to a division of the Rutaceae with a single ovary. Masters has observed instances of both marginal and free central placentation. which are considered by botanists to be generally of a highly important nature. the flowers of Papaver bracteatum offer either two sepals with four petals (which is the common type with poppies).all the flowers and leaves on the same plant had been subjected to the same external and internal condition. It is so common to observe on the same plant. according to De Candolle. as in the last class of cases. we cannot even attribute them. In numerous other cases we find modifications of structure. affecting only some of the flowers on the same plant. St. or three sepals with six petals. pentamerous. In Helianthemum the capsule has been described as unilocular or tri-locular. and even in the same panicle. but he subsequently saw them growing on the same bush. "Une lame PLUS OU MOINS LARGE. Lastly. and in H. which grow close together under the same conditions. As these variations seem of no special use to the plants." In the flowers of Saponaria officinalis Dr. Aug. The manner in which the petals are folded in the bud is in most groups a very constant morphological character.

the divisions of the flower or of the ovarium. for it will no longer be regulated by this same power of selection. transmitted in nearly the same state to numerous. as I am inclined to believe. generally becomes variable. as well as through the intercrossing of distinct individuals. which we consider as important—such as the arrangement of the leaves. otherwise modified. and in the case of the imperfect. descendants. if any new principle has to be invoked. But when. I should infer from the mere fact of the parts in question differing or varying greatly on the same plant. any slight deviations in them could not have been governed or accumulated through this latter agency. but not through natural selection. feathers or scales. From the fact of the above characters being unimportant for the welfare of the species.development? On the contrary. it must be one of retrogression rather than of progression. we may infer that the result would be almost uniform. they may be. namely. first appeared in many cases as fluctuating variations. from the nature of the organism and of the conditions. A structure which has been developed through long-continued selection. It cannot have been of much importance to the greater number of mammals. and scales to all true reptiles. when it ceases to be of service to a species. that characters of slight 213 . closed flowers. above described. and apparently often have been. yet hair has been transmitted to almost all mammals. We are ignorant of the exciting cause of the above specified modifications. Thus. 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. whatever it may be. modifications have been induced which are unimportant for the welfare of the species. whether they were clothed with hair. and consequently is often assumed to be of high vital importance to the species. the position of the ovules. which sooner or later became constant through the nature of the organism and of the surrounding conditions. birds. any slight variations which occurred in them would not have been accumulated and augmented through natural selection. etc. for as these morphological characters do not affect the welfare of the species. morphological differences. of whatever importance they may generally be to us for our classifications. It is a strange result which we thus arrive at. as we see with rudimentary organs. or reptiles. feathers to all birds. and in this case all the individuals of the species would be modified in the same manner. but if the unknown cause were to act almost uniformly for a length of time. which is common to many allied forms. A structure. is ranked by us as of high systematic importance. that such modifications were of extremely small importance to the plants themselves..

yet this necessarily follows. Mr. I will here consider in some detail several of the cases 214 . this is by no means so paradoxical as it may at first appear. through the continued action of natural selection." This subject is intimately connected with that of the gradation of the characters. as I have attempted to show in the fourth chapter. I never before felt so strongly convinced of the general truth of the conclusions here arrived at. independently of natural selection. considered in the present volume. but. the conversion of a swim-bladder into lungs. and have treated in my "Variation under Domestication" at greater length than. subject. Mivart's book.vital importance to the species. Mr. who may wish to weigh the evidence on both sides. for instance. or have been. Mivart's objections will be. Nevertheless. The one new point which appears to have struck many readers is. Wallace and myself. When thus marshalled. and natural selection tends towards this end. as propounded by Mr. "That natural selection is incompetent to account for the incipient stages of useful structures. 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. My judgment may not be trustworthy. St. All Mr. as we shall hereafter see when we treat of the genetic principle of classification. Mivart passes over the effects of the increased use and disuse of parts. A distinguished zoologist. points which were discussed in the last chapter under two headings. inasmuch as the parts are thus enabled to perform their functions more efficiently. Although we have no good evidence of the existence in organic beings of an innate tendency towards progressive development. they make a formidable array. any other writer. but after reading with care Mr. has recently collected all the objections which have ever been advanced by myself and others against the theory of natural selection. and comparing each section with what I have said on the same head. For the best definition which has ever been given of a high standard of organisation. is the degree to which the parts have been specialised or differentiated. When discussing special cases. and has illustrated them with admirable art and force. He likewise often assumes that I attribute nothing to variation. of course. to much partial error. often accompanied by a change of function. Mivart's plan to give the various facts and considerations opposed to his conclusions. no slight effort of reason and memory is left to the reader. as I believe. which I have always maintained to be highly important. George Mivart. and as it forms no part of Mr. are the most important to the systematist. in so intricate a subject.

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. By this process long-continued. a great difference in preserving an animal's life.. browse on the twigs of trees. Mivart's objections. will often have been preserved. head and tongue. for those individuals which had some one part or several parts of their bodies rather more elongated than usual. has its whole frame beautifully adapted for browsing on the higher branches of trees. as with the race-horse and greyhound. Before coming to Mr. by breeding from the victorious birds. It can thus obtain food beyond the reach of the other Ungulata or hoofed animals inhabiting the same country. when he methodically improves a breed: natural selection will preserve and thus separate all the superior individuals. considering its probable habits of life. as man does. either inheriting the same bodily peculiarities. in which careful measurements are given. or with a tendency to vary again in the same manner. would generally have survived. fore legs. allowing them freely to intercross. to which food the common cattle and horses are then driven. etc. but from the projection of the lower jaw they cannot.advanced by Mr. reeds. while the individuals less favoured in the same respects will have been the most liable to perish. These will have intercrossed and left offspring. which exactly corresponds with what I have called 215 . during such periods. and will destroy all the inferior individuals. due to the laws of growth and variation. These cattle can browse as well as others on grass. The Niata cattle in South America show us how small a difference in structure may make. Man has modified some of his animals. without necessarily having attended to special points of structure. The giraffe. the individuals which were the highest browsers and were able during dearths to reach even an inch or two above the others. are not of the slightest use or importance to most species. by simply preserving and breeding from the fleetest individuals. These slight proportional differences. for they will have roamed over the whole country in search of food. during the often recurrent droughts. and this must be a great advantage to it during dearths. it may be well to explain once again how natural selection will act in all ordinary cases. much elongated neck. if not fed by their owners. as want of space prevents me from considering all. So under nature with the nascent giraffe. or as with the game-cock. selecting those which are the most illustrative. so that at these times the Niatas perish. But it will have been otherwise with the nascent giraffe. Mivart. by its lofty stature. We here see that there is no need to separate single pairs.

why has not any other hoofed quadruped acquired a long neck and lofty stature. that no animal is more difficult to stalk than the giraffe. to a supply of food. left untouched by the other hoofed quadrupeds of the country. no doubt. It is from this cause. The preservation of each species can rarely be determined by any one advantage. Chauncey Wright has remarked. combined. by violently swinging its head armed with stump-like horns. to acquire slightly longer necks? In every district some one kind of animal will almost certainly be able to browse higher than the others." But as the giraffe does actually exist in large numbers in Africa. more than counterbalance the advantages. and against this animal. again. great and small. the answer is not difficult. in which trees grow. abound there. if natural selection be so potent. Mivart brings forward two objections. through natural selection and the effects of increased use. besides the giraffe. and it is almost equally certain that this one kind alone could have its neck elongated for this purpose. we see the lower branches trimmed or planed to an exact level by the browsing of the horses or cattle. and as some of the largest antelopes in the world. why has not any member of the group acquired a long proboscis? With respect to South Africa. taller than an ox. Mr. that increased bulk would act as a protection against almost all beasts of prey excepting the lion. and if high browsing be so great an advantage. if kept there. 216 . in a most important manner with the inherited effects of the increased use of parts. why should we doubt that. Nor must we overlook the fact. which was formerly inhabited by numerous herds of the giraffe. as Sir S. for instance. to sheep. In every meadow in England. Mivart then asks (and this is his second objection).unconscious selection by man. it seems to me almost certain that an ordinary hoofed quadruped might be converted into a giraffe. subjected as now to severe dearths. but by the union of all. at each stage of increased size. intermediate gradations could formerly have existed there. and what advantage would it be. Baker remarks. would have been of some advantage to the nascent giraffe. This animal also uses its long neck as a means of offence or defence. its tall neck—and the taller the better— would. in times of scarcity. as far as size is concerned. the camel. in a lesser degree. serve as a watch-tower. and. guanaco and macrauchenia? Or. and can best be given by an illustration. One is that the increased size of the body would obviously require an increased supply of food. and he considers it as "very problematical whether the disadvantages thence arising would not. as Mr. To this conclusion Mr. Assuredly the being able to reach.

while South Africa abounds with them to an unparalleled degree. for instance South America.In South Africa the competition for browsing on the higher branches of the acacias and other trees must be between giraffe and giraffe. then natural selection will be able to do little. Except by assigning such 217 . however. Whatever the causes may have been. Lastly. Why this should be so we do not know.—as seems often to be the case. Even if the fitting variations did arise. in other quarters of the world. cannot be distinctly answered. Why. etc. To reach the foliage at a considerable height (without climbing. In order that an animal should acquire some structure specially and largely developed. it is almost indispensable that several other parts should be modified and coadapted. natural selection is a slow process. if the number of individuals existing in a country is determined chiefly through destruction by beasts of prey—by external or internal parasites. see in a general manner that various causes might have interfered with the development of a long neck or proboscis. in modifying any particular structure for obtaining food. 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. 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. and we know that some areas support singularly few large quadrupeds. 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. We can. and the same favourable conditions must long endure in order that any marked effect should thus be produced. and that some species are much more variable than others. We are ignorant with respect to the conditions which determine the numbers and range of each species. it does not follow that the necessary parts should always vary in the right direction and to the right degree. for which hoofed animals are singularly ill-constructed) implies greatly increased bulk of body. nor why the later tertiary periods should have been much more favourable for their existence than the present time. and we cannot even conjecture what changes of structure would be favourable to its increase in some new country. Although every part of the body varies slightly. For instance. or will be greatly retarded. With the different species of our domesticated animals we know that the parts vary in a different manner and degree. though it is so luxuriant. various animals belonging to this same order have not acquired either an elongated neck or a proboscis. and not with the other ungulate animals.

like the so-called flying squirrels. which it is thought would be beneficial to certain species. but these would already be largely preyed on by the reptiles or birds. Therefore Sir C. it would never be reconverted back. A strictly terrestrial animal. by occasionally hunting for food in shallow water. into the less efficient power of gliding through the air. 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. Gradations of structure. Bats. hoofed quadrupeds have not acquired much elongated necks or other means for browsing on the higher branches of trees. in many quarters of the world. might. like many birds. indeed. yet as some of these bats are peculiar species. 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. for the former there would be no prey. as formerly shown. but by no terrestrial mammals. One writer asks. through disuse. besides the general ones just indicated. for the sake of escaping from their enemies. and bats into terrestrial insectivorous animals. so as to compete with birds or other ground animals. probably acquired their wings by at first gliding through the air from tree to tree. by the aid of their hind legs alone. or completely lost. But seals would not find on oceanic islands the conditions favourable to their gradual reconversion into a terrestrial form. with each stage beneficial to a changing species. and for such a change a bat seems singularly ill-fitted. then in streams or lakes. for the bats groundinsects would serve as food. with each step 218 . which first colonise and abound on most oceanic islands. Bats. In each case various causes. and assigns certain reasons in answer. but in this case it would be necessary that they should first have acquired the power of running quickly on the ground. they must have long inhabited their present homes. Objections of the same nature as the foregoing have been advanced by many writers. at least for the above purposes. will be favoured only under certain peculiar conditions. have probably interfered with the acquisition through natural selection of structures. have had their wings greatly reduced in size. might at last be converted into an animal so thoroughly aquatic as to brave the open ocean. These conjectural remarks have been made merely to show that a transition of structure. but when the power of true flight had once been acquired. we cannot explain why. Oceanic islands are inhabited by bats and seals.general and vague reasons. Lyell asks.

The resemblance is often wonderfully close. for natural selection to seize upon and perpetuate. Insects often resemble for the sake of protection various objects. considering the almost infinite number of surrounding objects and the diversity in form and colour of the hosts of insects which exist. Darwin's theory. and even to the manner in which the insects hold themselves. as such development would be advantageous to all? Why have not apes acquired the intellectual powers of man? Various causes could be assigned. but only the surface which commonly surrounds them. On this head. but extends to form. bamboo. such as green or decayed leaves. of one fish) resemble for the sake of protection special objects. and their relative probability cannot be weighed. "As. excrement of birds. we can understand how it is that the larger and higher animals do not (with the exception. to see how such indefinite oscillations of infinitesimal beginnings can ever build up a sufficiently appreciable resemblance to a leaf. Mr. of two races of savages.beneficial. are rare and exceptional." But in all the foregoing cases the insects in their original state no doubt presented some rude and accidental resemblance to an object commonly found in the stations frequented by them. if not impossible. is a highly complex affair. As some rude resemblance is necessary for the first start. but to this latter point I shall hereafter recur. A definite answer to the latter question ought not to be expected. there is a constant tendency to indefinite variation. and this chiefly in colour. as far as I know. We will return to Mr. more than one writer has asked why have some animals had their mental powers more highly developed than others. they must tend to neutralize each other. The cases of the imitation of such objects as the excrement of birds. and as the minute incipient variations will be in ALL DIRECTIONS. according to Mr. Lastly. offer an excellent instance of a resemblance of this kind. one has risen higher in the scale of civilisation than the other. and living insects. dead twigs. Mivart remarks. flowers. seeing that no one can solve the simpler problem. Nor is this at all improbable. and at first to form such unstable modifications that it is difficult. and this apparently implies increased brain power. bits of lichen. why. and is not confined to colour. spines. and that it varied slightly in 219 . and that there is nothing strange in a transition not having occurred in any particular case. Assuming that an insect originally happened to resemble in some degree a dead twig or a decayed leaf. The caterpillars which project motionless like dead twigs from the bushes on which they feed. Mivart's other objections. or other object. but as they are conjectural. it would be useless to give them.

if they rendered the insect at all less like the imitated object. of about 300 plates or laminae. then all the variations which rendered the insect at all more like any such object. With respect to the baleen. independently of natural selection. the middle lamina being in one species. would tend towards its preservation. for in every group the characters which differ in the several species are the most apt to vary. or those common to all the species. while the generic characters. The extremities and inner margins of all the plates are frayed into stiff bristles. of a walking-stick insect (Ceroxylus laceratus). and in these having become more or less green-coloured. and the baleen. Insects are preyed on by birds and other enemies whose sight is probably sharper than ours. Mivart's difficulty with respect to "the last touches of perfection in the mimicry. one of its greatest peculiarities. then its 220 . or even fifteen feet in length. and thus favoured its escape. that a native Dyak maintained that the foliaceous excrescences were really moss. and every grade in resemblance which aided an insect to escape notice or detection. but in the different species of Cetaceans there are gradations in length. The middle and longest lamina in the Greenland whale is ten. Mr. which clothe the whole gigantic palate. Considering the nature of the differences between the species in the group which includes the above Ceroxylus. on each side of the upper jaw. which resembles "a stick grown over by a creeping moss or jungermannia. would be preserved. Wallace. The Greenland whale is one of the most wonderful animals in the world. and in the Balaenoptera rostrata only about nine inches in length. which stand close together transversely to the longer axis of the mouth. but as the case stands there is none. and serve to strain or sift the water. There would indeed be force in Mr. or whalebone. there is nothing improbable in this insect having varied in the irregularities on its surface. The baleen consists of a row. Mivart remarks that if it "had once attained such a size and development as to be at all useful. through mere fluctuating variability. four feet. or. Within the main row there are some subsidiary rows. according to Scoresby. and thus to secure the minute prey on which these great animals subsist. while other variations would be neglected and ultimately lost. if we were to attempt to account for the above resemblances. twelve. are the most constant." as in the case given by Mr. in another three. Nor can I see any force in Mr." So close was this resemblance. they would be eliminated. and the more perfect the resemblance so much the better for the insect. Mivart's objection. The quality of the whalebone also differs in the different species. in another eighteen inches.many ways.

and that the immense plates of baleen in the Greenland whale might have been developed from such lamellae by finely graduated steps. the lamellae would be six inches in length. and in being thus furnished it differs conspicuously from the lower jaw of a whale. like whales. or sifters. but finer. in which species the baleen is only nine inches long. instead of straight downward. I wish only to show that this is not incredible. 221 . and the family has sometimes been called Criblatores. being about one-third of an inch in length. In the genus Prion. two-thirds of the length of the baleen in this species of whale. But how to obtain the beginning of such useful development?" In answer. 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. so that they thus curiously resemble the plates of baleen. On the other hand. At their bases there is a short subsidiary row of obliquely transverse lamellae. as they project inward. They arise from the palate. elastic lamellae. 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. though incomparably less bulky. But towards the extremity of the beak they differ much. and they project fourteen onehundredths of an inch beneath the edge. The beak of a shoveller-duck (Spatula clypeata) is a more beautiful and complex structure than the mouth of a whale. In these several respects they resemble the plates of baleen in the mouth of a whale.preservation and augmentation within serviceable limits would be promoted by natural selection alone. and placed transversely to the longer axis of the mouth. The upper mandible is furnished on each side (in the specimen examined by me) with a row or comb formed of 188 thin. so that if we were to make the head of the shoveller as long as that of the Balaenoptera. obliquely bevelled so as to be pointed. it may be asked. a member of the distinct family of the Petrels. Those standing towards the middle are the longest. The entire head of the shoveller. each of service to its possessor. which are well developed and project beneath the margin. subsist by sifting the mud and water. the extremities of these lower lamellae are frayed into fine bristly points. is about one-eighteenth of the length of the head of a moderately large Balaenoptera rostrata. so that the beak of this bird resembles in this respect the mouth of a whale. which is destitute of baleen. The lower mandible of the shoveller-duck is furnished with lamellae of equal length with these above. and are attached by flexible membrane to the sides of the mandible. the upper mandible alone is furnished with lamellae. that is.

and are edged with translucent. The palate is also covered with hard rounded knobs. In this latter bird the lamellae of the upper mandible are much coarser than in the common duck. they are only about fifty in number on each side. in which the lamellae are less developed than in the common goose. for seizing or tearing solid food. constantly uses it for this purpose. 222 . which it crops like the common goose. In the Egyptian goose (Chenalopex) the beak closely resembles that of the common duck. nor so distinct from each other. as I hear from Mr. and are firmly attached to the sides of the mandible. lastly. Bartlett. Salvin. which project very little. but I do not know whether they use their beaks for sifting the water. and terminating upward in teeth-like knobs. We thus see that a member of the duck family. into one like the shoveller. There are other species of geese. nor do they project so much inward. for this bird could hardly use any part of its beak. The beak of a goose. and in some respects through that of the Aix sponsa. however. almost confluent. or even a member with a beak having less well-developed lamellae. without any great break. Turning to another group of the same family. yet this goose. provided with a beak almost exclusively adapted for sifting the water. They are square-topped. for which purpose it is so well fitted that it can crop grass closer than almost any other animal. as I hear from Mr. as if for crushing food. through the beak of the Merganetta armata. hardish tissue. coarser and sharper than in the duck. and do not project at all beneath the margin. There are other species. is grass. as every one knows." Its chief food. except the hooked tip. might be converted by small changes into a species like the Egyptian goose—this into one like the common duck—and. E. about twentyseven in number on each side.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. as I may add. but the lamellae are not so numerous. yet this bird. Salvin). In this latter species the lamellae are much coarser than in the shoveller. The common goose does not sift the water. as far as fitness for sifting is concerned. The edges of the lower mandible are crossed by numerous fine ridges. to the beak of the common duck. as I am informed by Mr. with a beak constructed like that of a common goose and adapted solely for grazing. "uses its bill like a duck by throwing the water out at the corners. but uses its beak exclusively for tearing or cutting herbage. Although the beak is thus very inferior as a sifter to that of a shoveller. in which the lamellae are considerably less developed than in the common duck. Bartlett. The edges of the lower mandible are serrated with teeth much more prominent.

like the knobs on the beak of the goose. There is. and that the structure of every part of its frame must be well adapted to its conditions of life. or resting-surface. however. serving for the widely different purpose of securing live fish. but rather more regularly placed.might also be converted by small changes into one provided with prominent. with the lateral fins often of smaller size. lead us onward to the enormous plates of baleen in the Greenland whale. with both sides equally coloured. are remarkable for their asymmetrical bodies. If so. but does not pass right through the skull. recurved teeth. 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. It is obvious that unless the lower eye did thus travel round. resembles at first sight the ventral surface of an ordinary fish. hard points of horn. but in some on the right side. The Pleuronectidae. with the functions of the parts slowly changing during the progress of development. which may be observed in still-existing Cetaceans. it could not be 223 . The Hyperoodon bidens is destitute of true teeth in an efficient condition. The lower. nothing improbable in supposing that some early Cetacean form was provided with similar points of horn on the palate. gradations. Nor is there the least reason to doubt that each step in this scale might have been as serviceable to certain ancient Cetaceans. But the eyes offer the most remarkable peculiarity. less developed in many ways than the upper side. they stand opposite to each other. in which the lamellae would be two-thirds of the length of the plates of baleen in the Balaenoptera rostrata. but its palate is roughened. until they became as well constructed as those of the shoveller. or Flat-fish. During early youth. for they are both placed on the upper side of the head. in which case they would have been used both for seizing objects and for sifting the water. We should bear in mind that each species of duck is subjected to a severe struggle for existence. it is of a white colour. according to Lacepede. like those of the Merganser (a member of the same family). and occasionally reversed adult specimens occur. aided it in seizing or tearing its food. then into lamellae like those of the domestic duck. and which. From this stage. as was formerly thought to be the case. therefore. and the whole body is then symmetrical. They rest on one side—in the greater number of species on the left. Soon the eye proper to the lower side begins to glide slowly round the head to the upper side. as are the gradations in the beaks of the different existing members of the duck-family. with small unequal. and so onward. in which case they would have served exclusively as a sifting apparatus. Returning to the whales.

the small size of their lateral fins.used by the fish while lying in its habitual position on one side. etc. flounders." Mr. and they do this so vigorously that the eye is pressed hard against the upper part of the orbit. have the habit of occasionally resting on one side at the bottom. The forehead between the eyes consequently becomes. while very young and still symmetrical. "a long series of forms exhibiting a gradual transition from Hippoglossus pinguis. and he has observed that they often then strain their lower 224 . On one occasion Malm saw a young fish raise and depress the lower eye through an angular distance of about seventy degrees. temporarily contracted in breadth. It is also known with the higher animals. in which I quite agree with him. of which I have given a figure. and to their being destitute of a swim-bladder. and facility for feeding on the ground. which are entirely thrown to one side. also. and remarks that a sudden spontaneous transformation in the position of the eyes is hardly conceivable. the lower eye upward. far from clear. The lower eye would. they fall to the bottom on one side. soon growing tired. That the Pleuronectidae are admirably adapted by their flattened and asymmetrical structure for their habits of life. which does not in any considerable degree alter the shape in which it leaves the ovum. with their eyes standing on opposite sides of the head. however. being extremely common. of the family present. We should remember that the skull at this early age is cartilaginous and flexible. cannot long retain a vertical position. so that it readily yields to muscular action. as Schiodte remarks. The chief advantages thus gained seem to be protection from their enemies. to the soles. to see above them.. its weight drags forward all the bones of the skull on the same side. even. that the skull yields and is altered in shape. is manifest from several species. indeed. He then adds: "If the transit was gradual. The Pleuronectidae. owing to the excessive depth of their bodies. even after early youth. as could be plainly seen. While thus at rest they often twist. if one ear flops forward and downward. salmon. that such an incipient transformation must rather have been injurious. 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. as Malm observed. Mivart has taken up this case. such as soles. The different members." But he might have found an answer to this objection in the excellent observations published in 1867 by Malm. It seems. Hence. if the skin or muscles be permanently contracted through disease or some accident. have been liable to be abraded by the sandy bottom. and several other symmetrical fishes. With long-eared rabbits. Malm states that the newly-hatched young of perches.

may likewise be accounted for by disuse. concludes his abstract of Malm's paper. and if this be so. on the other hand." Perhaps the lesser number of teeth in the proportion of four to seven in the upper halves of the two jaws of the plaice. including the lateral fins. by remarking that "the author gives a very simple explanation of the abnormal condition of the Pleuronectoids. and other species on the right side. Disuse. so like the sandy bed 225 .most. Mivart considers would be injurious. Gunther. Schiodte believes. Dr. the eyeless side of the head. while resting on one side at the bottom. on the other hand. and a permanent effect is thus produced on the form of the head. of feeding with ease on the ground. of endeavouring to look upward with both eyes. we could understand how it is that certain species. though Yarrel thinks that the reduced size of these fins is advantageous to the fish. with the jaw bones stronger and more effective on this. Traquair supposes. With the Pleuronectidae. and in this fish. while young." We thus see that the first stages of the transit of the eye from one side of the head to the other. which Mr. we may reasonably suppose that the absence of colour in flatfish on the side. the older they grow the more habitually they rest on one side. These fishes. 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. and no permanent effect is thus produced. the tendency to distortion would no doubt be increased through the principle of inheritance. Our great authority on Fishes. as Dr. habitually fall over and rest on the left side. to twenty-five to thirty in the lower halves. are soon able to hold themselves in a vertical position. and swims diagonally through the water. rests on its left side at the bottom. will account for the less developed condition of the whole inferior half of the body. owing to the increasing flatness of their bodies. But it cannot be supposed that the peculiar speckled appearance of the upper side of the sole. in confirmation of the above view. whether it be the right or left. Judging from analogy. no doubt beneficial to the individual and to the species. Malm adds. which is under.eyes so as to look upward. which is not a member of the Pleuronectidae. in opposition to some other naturalists. may be attributed to the habit. is due to the exclusion of light. that the adult Trachypterus arcticus. for the sake. however. than on the other. as "there is so much less room for their action than with the larger fins above. From the colourless state of the ventral surface of most fishes and of many other animals. the two sides of the head are said to be somewhat dissimilar. that the Pleuronectidae are not quite symmetrical even in the embryo. and their skulls are thus rendered rather crooked. and on the position of the eyes.

Gunther. as I have before insisted.of the sea. Habit. considering its habits while young. and thus aided themselves in climbing. If the harvest mouse had been more strictly arboreal. and how much to natural selection. Brehm saw the young of an African monkey (Cercopithecus) clinging to the under surface of their mother by their hands. or the power in some species. and perhaps of their disuse. or favour their chance of having and of rearing offspring. therefore." But there is no necessity for any such belief. Professor Henslow kept in confinement some harvest mice (Mus messorius) which do not possess a structurally prehensive tail. and at the same time they hooked their little tails round that of their mother. and this almost implies that some benefit great or small is thus derived. but he frequently observed that they curled their tails round the branches of a bush placed in the cage. it would be difficult to say. and many other peculiarities. and serves as a fifth hand. For all spontaneous variations in the right direction will thus be preserved. as will those individuals which inherit in the highest degree the effects of the increased and beneficial use of any part. they must. Why Cercopithecus. 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. I have received an analogous account from Dr. How much to attribute in each particular case to the effects of use. We should keep in mind. and are indispensable for their existence. it seems impossible to decide. would in all probability suffice for the work. or the presence of bony tubercles on the upper side of the turbot. Here natural selection has probably come into play. of changing their colour in accordance with the surrounding surface. The extremity of the tail in some American monkeys has been converted into a wonderfully perfect prehensile organ. as well as in adapting the general shape of the body of these fishes. will be strengthened by natural selection. The mammary glands are common to the whole class of mammals. as is the case with some members of the same order. who has seen a mouse thus suspend itself. are due to the action of the light. has not become thus provided. It is. A reviewer. possible that the long tail of this monkey may be of more service to it as a balancing organ in making its prodigious leaps. Mivart in every detail. I may give another instance of a structure which apparently owes its origin exclusively to use or habit. have been 226 . it would perhaps have had its tail rendered structurally prehensile. however. than as a prehensile organ. to their habits of life. who agrees with Mr. that the inherited effects of the increased use of parts. as recently shown by Pouchet.

developed at an extremely remote period, and we can know nothing positively about their manner of development. Mr. 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, what chance was there of the perpetuation of such a variation?" But the case is not here put fairly. It is admitted by most evolutionists that mammals are descended from a marsupial form; and if so, the mammary glands will have been at first developed within the marsupial sack. In the case of the fish (Hippocampus) the eggs are hatched, and the young are reared for a time, within a sack of this nature; and an American naturalist, Mr. Lockwood, believes from what he has seen of the development of the young, that they are nourished by a secretion from the cutaneous glands of the sack. Now, with the early progenitors of mammals, almost before they deserved to be thus designated, is it not at least possible that the young might have been similarly nourished? And in this case, the individuals which secreted a fluid, in some degree or manner the most nutritious, so as to partake of the nature of milk, would in the long run have reared a larger number of well-nourished offspring, than would the individuals which secreted a poorer fluid; and thus the cutaneous glands, which are the homologues of the mammary glands, would have been improved or rendered more effective. It accords with the widely extended principle of specialisation, that the glands over a certain space of the sack should have become more highly developed than the remainder; and they would then have formed a breast, but at first without a nipple, as we see in the Ornithorhyncus, at the base of the mammalian series. Through what agency the glands over a certain space became more highly specialised than the others, I will not pretend to decide, whether in part through compensation of growth, the effects of use, or of natural selection. The development of the mammary glands would have been of no service, and could not have been affected through natural selection, unless the young at the same time were able to partake of the secretion. There is no greater difficulty in understanding how young mammals have instinctively learned to suck the breast, than in understanding how unhatched chickens have learned to break the egg-shell by tapping against it with their specially adapted beaks; or how a few hours after leaving the shell they have learned to pick up grains of food. In such cases the most probable solution seems to be, that the habit was at first acquired by practice at a more advanced age, and afterwards transmitted to the


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


many co-ordinated modifications are almost indispensable. and with the vibracula we can understand how the successive gradations might have been of service. as the struggle for life did not depend on such structures. became attached to the free ends of the caudicles—all these gradations being of manifest benefit to the plants in question. would have been beneficial to a species. I need not repeat what has been so lately said. In most cases only general reasons. which we imagine would have been gained through natural selection from appearing to us advantageous to the species. and still subserving nearly but not quite the same purpose. are necessary for the development of a structure. the threads which originally served to tie together the pollen-grains. considering our ignorance of the past history of each species. serviceable modifications in the ultimate and penultimate segments of a limb. It has often been asked. such as that secreted by the stigmas of ordinary flowers. and concentrated into a confined area. is opposed to what we can 237 . can be traced cohering into caudicles. often erroneously. became developed through natural selection into tridactyle pedicellariae. and the requisite conditions may seldom have concurred. if natural selection be so potent. The belief that any given structure. to which it would apparently have been advantageous? But it is unreasonable to expect a precise answer to such questions. With respect to climbing plants. which was at first used solely for locomotion. but in some few cases special reasons. through slight. There is no more difficulty in understanding how the branched spines of some ancient Echinoderm. which we think.these glands were improved in function through natural selection. In the avicularia and vibracula of the Polyzoa we have organs widely different in appearance developed from the same source. With the pollinia of orchids. 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 destructive agencies. In this case. which served as a defence. they could not have been acquired through natural selection. can be assigned. which stood in no relation to certain structures. in which case they would have formed a mamma. and the steps can likewise be followed by which viscid matter. than in understanding the development of the pincers of crustaceans. and of the conditions which at the present day determine its numbers and range. In many cases complex and long-enduring conditions. would have been gained under all circumstances through natural selection. Thus to adapt a species to new habits of life. why has not this or that structure been gained by certain species. often of a peculiar nature.

or even under domestication. by man has given rise to many well-adapted domestic races. That species have a capacity for change will be admitted by all evolutionists." about which it is not pretended that anything is known. aided by the other agencies often specified." For instance. Mivart is further inclined to believe. it is not probable that such great and abrupt variations have often occurred under nature. Mivart does not deny that natural selection has effected something. 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. I am bound to add." At the present day almost all naturalists admit evolution under some form. and the others will hereafter be considered. which implies great breaks or discontinuity in the series. that new species manifest themselves "with suddenness and by modifications appearing at once. Mr. But as species are more variable when domesticated or cultivated than under their natural conditions. 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. he supposes that the differences between the extinct three-toed Hipparion and the horse arose suddenly. and the characters which thus reappear were. which through the aid of selection. but he considers it as "demonstrably insufficient" to account for the phenomena which I explain by its agency. have been advanced for the same purpose in an able article lately published in the "Medico-Chirurgical Review. as already explained. would equally well give rise by graduated steps to natural races or species. in many cases at first gained in a 238 . Of these latter variations several may be attributed to reversion. Everyone who believes in slow and gradual evolution.understand of its manner of action. Mivart believes that species change through "an internal force or tendency. through the aid of natural selection. an advance. appears to me improbable in the highest degree. as it seems to me. His chief arguments have now been considered. in organisation. but in some few cases a retrogression. it is probable. to invoke any internal force beyond the tendency to ordinary variability." and apparently he would extend the same view to the wings of bats and pterodactyles. This conclusion. Mr. The final result will generally have been. but there is no need. and some naturalists agree with him. as are known occasionally to arise under domestication. and which. Mr. 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. that some of the facts and arguments here used by me.

in proceeding from north to south. which varied more or less in any favourable direction. it is almost necessary to believe. in order that a new species should suddenly appear in the manner supposed by Mr. and of the destruction of a large number which varied in an opposite manner. If such occurred under nature. they throw very little light on our subject. porcupine men. and compare the species which have just passed away with those still living within the same areas. or if we compare the fossil species embedded in the sub-stages of the same geological formation. Excluding such cases of abrupt variations. abrupt and strongly marked variations occur in our domesticated productions. in opposition to all analogy. we meet with a host of closely related or representative species. Ancon sheep. etc. 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. are as follows. On every continent. Mivart. etc. unless abrupt variations of this kind are specially preserved and separated by the care of man. to be lost by accidental causes of destruction and by subsequent intercrossing. That many species have been evolved in an extremely gradual manner. the few which remain would at best constitute. or have lately existed. According to our experience. which we have reason to believe were formerly connected. and see how many of their inhabitants can be raised only to the rank of doubtful species. Hence. and so it is known to be under domestication. from lowland to upland. and as they are widely different in character from natural species. Look at the many outlying islands round a continent. and for entirely disbelieving that they have changed in the wonderful manner indicated by Mr. they would be liable. closely related to their parental types... My reasons for doubting whether natural species have changed as abruptly as have occasionally domestic races. through the preservation of a large number of individuals. I am compelled to allude to subjects hereafter to be discussed. if found in a state of nature.gradual manner. as formerly explained. Niata cattle. such as six-fingered men. But in making these and the following remarks. doubtful species. as in the case of unconscious selection by man. So it is if we look to past times. as we likewise do on certain distinct continents. that several wonderfully changed individuals appeared simultaneously within the same district. It is indeed manifest that multitudes of species are related in the closest manner to other species that still exist. A still greater number must be called monstrosities. This difficulty. is avoided on the theory of gradual evolution. Mivart. singly and at rather long intervals of time. and it will hardly be 239 . there can hardly be a doubt.

or the falling of a facetted spheroid from one facet to another. Many analogous facts. ought not to be made without strong evidence being assigned. and present a greater number of varieties than do the species in the smaller genera. instead of to distinct species. are undistinguishable at an early embryonic period. there is nothing strange in new forms appearing as if suddenly developed. Such an admission. and how the parts which are developed in an extraordinary degree or manner are more variable than other parts of the same species. that numerous and wonderfully fine gradations can be traced. or the sudden conversion of a Hipparion into a horse. such as the sudden crystallisation of inorganic substances. hardly any light is thrown by the belief in abrupt modifications on the deficiency of connecting links in our geological formations. connecting together widely different structures. Many large groups of facts are intelligible only on the principle that species have been evolved by very small steps. Nor should it be forgotten. It is notorious that the wings of birds and bats. and they present other analogies with varieties. Unless we admit transformations as prodigious as those advocated by Mr. yet it may be maintained that some have been developed in a different and abrupt manner. hardly deserve consideration. One class of facts. as was shown in our second chapter. Mivart. the sudden appearance of new and distinct forms of life in our geological formations supports at first sight the belief in abrupt development. in relation to periods remote in the history of the world. If the record is as fragmentary as many geologists strenuously assert. the fact that the species included in the larger genera are more closely related to each other. and that they 240 . On this same principle we can understand how it is that specific characters are more variable than generic characters. could be added.maintained that such species have been developed in an abrupt or sudden manner. such as the sudden development of the wings of birds or bats. But against the belief in such abrupt changes. however. when we look to the special parts of allied species. as they have been shown to be by Mr. however. For instance. The vague and in some respects false analogies. embryology enters a strong protest. The former are also grouped in little clusters. Although very many species have almost certainly been produced by steps not greater than those separating fine varieties. all pointing in the same direction. Chauncey Wright. like varieties round species. namely. and the legs of horses or other quadrupeds. But the value of this evidence depends entirely on the perfection of the geological record. which have been advanced in favour of this view.

On this view of the meaning of embryological resemblances. in opposition to all analogy. one furnished with wings. Embryological resemblances of all kinds can be accounted for.become differentiated by insensibly fine steps. 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. have been suddenly produced. for instance. every detail in its structure being developed by insensibly fine steps. and yet should not bear even a trace in its embryonic condition of any sudden modification. as we shall hereafter see. will be almost compelled to assume. and to leave those of science. by the progenitors of our existing species having varied after early youth. The embryo is thus left almost unaffected. It cannot be denied that such abrupt and great changes of structure are widely different from those which most species apparently have undergone. that many individuals varied simultaneously. as it seems to me. and serves as a record of the past condition of the species. and having transmitted their newly-acquired characters to their offspring. it is incredible that an animal should have undergone such momentous and abrupt transformations as those above indicated. at a corresponding age. and indeed on any view. and of such complex and wonderful co-adaptations. He who believes that some ancient form was transformed suddenly through an internal force or tendency into. to enter into the realms of miracle. To admit all this is. 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. 241 . He will be forced to admit that these great and sudden transformations have left no trace of their action on the embryo. he will not be able to assign a shadow of an explanation.

their origin — Natural instincts of the cuckoo. and parasitic bees — Slave-making ants — Hive-bee. but different in their origin — Instincts graduated — Aphides and ants — Instincts variable — Domestic instincts. 242 . molothrus.Chapter Instinct 8 Instincts comparable with habits. 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.

that I have nothing to do with the origin of the mental powers. often comes into play. or in repeating anything by rote. an accurate notion of the frame of mind under which an instinctive action is performed. the sixth stage of construction. is usually said to be instinctive. which makes a very complicated hammock. Huber found it was with a caterpillar. Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. When once acquired. If. one action follows another by a sort of rhythm. as Pierre Huber expresses it. without their knowing for what purpose it is performed. so in instincts. they often remain constant throughout life. but every one understands what is meant. he is generally forced to go back to recover the habitual train of thought: so P. fifth. even with animals low in the scale of nature. say. if a person be interrupted in a song. when performed by an animal. A little dose of judgment or reason. with certain periods of time and states of the body. for if he took a caterpillar which had completed its hammock up to. when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds' nests. Habits easily become associated with other habits. but not necessarily of its origin. and sixth stages of construction. any more than I have with that of life itself. without experience. An action. and were put into one finished up to the sixth stage. which we ourselves require experience to enable us to perform. I may here premise. a caterpillar were taken out of a hammock made up. It would be easy to show that several distinct mental actions are commonly embraced by this term. for instance. so that much of its 243 . I think. and when performed by many individuals in the same way. We are concerned only with the diversities of instinct and of the other mental faculties in animals of the same class. As in repeating a well-known song. more especially by a very young one. But I could show that none of these characters are universal.1. I will not attempt any definition of instinct. to the third stage. This comparison gives. Several other points of resemblance between instincts and habits could be pointed out. and put it into a hammock completed up only to the third stage. How unconsciously many habitual actions are performed. the caterpillar simply re-performed the fourth. Many instincts are so wonderful that their development will probably appear to the reader a difficulty sufficient to overthrow my whole theory. however. indeed not rarely in direct opposition to our conscious will! yet they may be modified by the will or reason.

seemed forced to start from the third stage. it is at least possible that slight modifications of instinct might be profitable to a species. I have been surprised to find.work was already done for it. and then transmitted by inheritance to succeeding generations. we ought to find in nature. be might truly be said to have done so instinctively. where it had left off. so I do not doubt it has been with instincts. it was much embarrassed. making 244 . or we ought at least to be able to show that gradations of some kind are possible. as I believe. under its present conditions of life. those of the hive-bee and of many ants. Hence. far from deriving any benefit from this. 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. and if it can be shown that instincts do vary ever so little. then I can see no difficulty in natural selection preserving and continually accumulating variations of instinct to any extent that was profitable. and are increased by.—that is of variations produced by the same unknown causes which produce slight deviations of bodily structure. Under changed conditions of life. As modifications of corporeal structure arise from. had played a tune with no practice at all. It can be clearly shown that the most wonderful instincts with which we are acquainted. slight. as in the case of corporeal structures. and this we certainly can do. It is thus. No complex instinct can possibly be produced through natural selection. namely. 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. But it would be a serious error to suppose that the greater number of instincts have been acquired by habit in one generation. and. It will be universally admitted that instincts are as important as corporeal structures for the welfare of each species. use or habit. that all the most complex and wonderful instincts have originated. variations. yet profitable. except by the slow and gradual accumulation of numerous. If we suppose any habitual action to become inherited—and it can be shown that this does sometimes happen—then the resemblance between what originally was a habit and an instinct becomes so close as not to be distinguished. and thus tried to complete the already finished work. could not possibly have been acquired by habit. If Mozart. instead of playing the pianoforte at three years old with wonderfully little practice. in order to complete its hammock. and are diminished or lost by disuse.

except in Europe and North America. as the ants do with their antennae. I felt sure that the aphides would want to excrete. I allowed an ant to visit them. therefore probably they do not excrete solely for the good of the ants. but has never. or at different seasons of the year. I removed all the ants from a group of about a dozen aphides on a dockplant. showing that the action was instinctive. how very generally gradations. as was first observed by Huber. or when placed under different circumstances. Afterwards. Changes of instinct may sometimes be facilitated by the same species having different instincts at different periods of life. their sweet excretion to ants: that they do so voluntarily. Although there is no evidence that any animal performs an action for the exclusive good of another species. But as the excretion is extremely viscid. is that of aphides voluntarily yielding. the following facts show. I then tickled and stroked them with a hair in the same manner. it is no doubt a convenience to the aphides to have it removed. It is certain. can be discovered. leading to the most complex instincts. After this interval. from the observations of Huber. it then began to play with its antennae on the abdomen first of one aphis and then of another. Even the quite young aphides behaved in this manner. by its eager way of running about to be well aware what a rich flock it had discovered. and each. but not one excreted. which was eagerly devoured by the ant. and not the result of experience. as in the case of corporeal structure. been produced for the exclusive good of others. yet each tries to take advantage of the instincts of others. and for no instinct being known among extinct species.allowance for the instincts of animals having been but little observed. One of the strongest instances of an animal apparently performing an action for the sole good of another. and prevented their attendance during several hours. the instinct of each species is good for itself. And such instances of diversity of instinct in the same species can be shown to occur in nature. as each takes advantage of the weaker bodily structure of other species. that the aphides show no dislike to the ants: if the latter be not present they are at last compelled to eject their excretion. and conformably to my theory. as far as we can judge. in which case either the one or the other instinct might be preserved by natural selection. immediately lifted up its abdomen and excreted a limpid drop of sweet juice. with which I am acquainted. I watched them for some time through a lens. as soon as it felt the antennae. Again.. and it immediately seemed. So again certain instincts cannot be considered as 245 . etc. as well as I could. but not one excreted.

The fear of man is slowly acquired. but want of space prevents me. for the large birds have been most persecuted by man. so wary in England. As some degree of variation in instincts under a state of nature. has it not granted to the bee "the ability to use some other material when wax was deficient?" But what other natural material could bees use? They will work. it has been asked. used a cement of wax and turpentine. as may be seen in nestling birds. both in extent and direction. Fear of any particular enemy is certainly an instinctive quality. for in uninhabited islands large birds are not more fearful than small. which vary partly in dependence on the situations chosen. though it is strengthened by experience. will gladly use a very different substance. in the greater wildness of all our large birds in comparison with our small birds. the migratory instinct. is tame in Norway. I can only assert that instincts certainly do vary—for instance. So it is with the nests of birds. are indispensable for the action of natural selection. can produce but a feeble effect on the reader's mind. with wax hardened with vermilion or softened with lard. without the facts in detail. that I do not speak without good evidence. by the various animals which inhabit desert islands. if advantageous to the species.absolutely perfect. even in England. with which he had covered decorticated trees. as many instances as possible ought to be given. instead of laboriously collecting propolis. 246 . instead of searching for pollen. but often from causes wholly unknown to us. to new instincts. oatmeal. and the magpie. as is the hooded crow in Egypt. and in its total loss. as I have seen. and we see an instance of this. if instinct be variable. and the inheritance of such variations. and on the nature and temperature of the country inhabited. We may safely attribute the greater wildness of our large birds to this cause. born in a state of nature. might have given rise. It has lately been shown that bees. which. namely. That the mental qualities of animals of the same kind. through natural selection. But I am well aware that these general statements. could be shown by many facts. Several cases could also be adduced of occasional and strange habits in wild animals. but as details on this and other such points are not indispensable. Andrew Knight observed that his bees. vary much. Why. Audubon has given several remarkable cases of differences in the nests of the same species in the northern and southern United States. and by the sight of fear of the same enemy in other animals. they may be here passed over. as I have elsewhere shown. I can only repeat my assurance.

Domestic instincts. habits. 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. and a tendency to run round. and have been transmitted for an incomparably shorter period. when young and without any training. With cats. of inherited variations of instinct in a state of nature will be strengthened by briefly considering a few cases under domestication. are certainly far less fixed than natural instincts. and dispositions are inherited. 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. a herd of deer. a flock of sheep. one naturally takes to catching rats. and likewise of the oddest tricks. for instance. Thus it is known that a cross with a bull-dog has affected for many generations the courage and obstinacy of 247 . and then slowly crawl forward with a peculiar gait. I cannot see that these actions. according to Mr. and these tendencies are known to be inherited. and another kind of wolf rushing round. It is notorious how much domestic animals vary in their mental qualities. and driving them to a distant point. and another hunted on marshy ground and almost nightly caught woodcocks or snipes. A number of curious and authentic instances could be given of various shades of disposition and taste. performed with eager delight by each breed. instead of at. as soon as it scented its prey.2. Inherited Changes Of Habit Or Instinct In Domesticated Animals The possibility. If we were to behold one kind of wolf. and another mice. always brought home game birds. performed without experience by the young. stand motionless like a statue. and in nearly the same manner by each individual. and how curiously they become mingled. is well shown when different breeds of dogs are crossed. another hares or rabbits. instead of at. St. How strongly these domestic instincts. associated with certain frames of mind or periods of time. as they may be called. by shepherd-dogs. but they have been acted on by far less rigorous selection. 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. or even probability. under less fixed conditions of life. One cat. we should assuredly call these actions instinctive. and without the end being known—for the young pointer can no more know that he points to aid his master. retrieving is certainly in some degree inherited by retrievers. John.

but I can hardly suppose that domestic rabbits have often been selected for tameness alone. and that the long-continued selection of the best individuals in successive generations made tumblers what they now are. Le Roy describes a dog. to habit and long-continued close confinement. When the first tendency to point was once displayed. when called. habit alone in some cases has sufficed. without intending to improve the breed. as I once saw. when thus tested by crossing.greyhounds. Brent. which in a like manner become curiously blended together. as each man tries to procure. These domestic instincts." that is. so that we must attribute at least the greater part of the inherited change from extreme wildness to extreme tameness. by not coming in a straight line to his master. is performed by young birds. the tumbler-pigeon to tumble—an action which. and unconscious selection is still in progress. Domestic instincts are sometimes spoken of as actions which have become inherited solely from long-continued and compulsory habit. that have never seen a pigeon tumble. Familiarity alone prevents our seeing how largely and how permanently the minds of our domestic animals have been modified. or probably could have taught. whose great-grandfather was a wolf. resemble natural instincts. in a pure terrier: the act of pointing is probably. but this is not true. had not some one dog naturally shown a tendency in this line. We may believe that some one pigeon showed a slight tendency to this strange habit. which cannot fly eighteen inches high without going head over heels. 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. only the exaggerated pause of an animal preparing to spring on its prey. It may be doubted whether any one would have thought of training a dog to point. It is scarcely possible to doubt that 248 . as I hear from Mr. never wish to sit on their eggs. methodical selection and the inherited effects of compulsory training in each successive generation would soon complete the work. dogs which stand and hunt best. scarcely any animal is tamer than the young of the tame rabbit. and this dog showed a trace of its wild parentage only in one way. as many have thought. and a cross with a greyhound has given to a whole family of shepherd-dogs a tendency to hunt hares. hardly any animal is more difficult to tame than the young of the wild rabbit. No one would ever have thought of teaching. and near Glasgow there are house-tumblers. and this is known occasionally to happen. and for a long period exhibit traces of the instincts of either parent: for example. On the other hand. as I have witnessed.

on the other hand. pursued both methodically and unconsciously. they are destroyed. when reared in India under a hen. 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. Hence. in other cases compulsory habit has done nothing. 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. but fear only of dogs and cats. for I am informed by Captain Hutton that the young chickens of the parent stock. as we see in wild ground-birds. when kept tame. their mother to fly away. during successive generations. How rarely. It is not that chickens have lost all fear.the love of man has become instinctive in the dog. But this instinct retained by our chickens has become useless under domestication. even when quite young. so that habit and some degree of selection have probably concurred in civilising by inheritance our dogs. but in most cases habit and selection have probably concurred. the Gallus bankiva. we may conclude that under domestication instincts have been acquired and natural instincts have been lost. do our civilised dogs. In some cases compulsory habit alone has sufficed to produce inherited mental changes. and this is evidently done for the instinctive purpose of allowing. All wolves. that fear of the dog and cat which no doubt was originally instinctive in them. and are then beaten. which at first appeared from what we must in our ignorance call an accident. jackals and species of the cat genus. sheep. where the savages do not keep these domestic animals. foxes. young chickens have lost wholly by habit. 249 . and all has been the result of selection. So it is with young pheasants reared in England under a hen. sheep and pigs. peculiar mental habits and actions. and if not cured. are most eager to attack poultry. require to be taught not to attack poultry. partly by habit and partly by man selecting and accumulating. and pigs! No doubt they occasionally do make an attack. On the other hand. for the mother-hen has almost lost by disuse the power of flight. are at first excessively wild.

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. the slave-making instinct of certain ants.3. namely. 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. I will select only three. 250 . perhaps.

aboriginal instinct of nidification. It has been both asserted and denied that the American cuckoo occasionally lays her eggs in other birds' nests. It has. that the cuckoo occasionally lays her eggs on the bare ground. and in their turn would be apt to lay their eggs in other birds' nests. not daily. that he once found in Illinois a young cuckoo. I could also give several instances of various birds which have been known occasionally to lay their eggs in other birds' nests. 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. of Iowa. If the old bird profited by this occasional habit through being enabled to emigrate earlier or through any other cause. But the American cuckoo is in this predicament. there could be no mistake in their identification. Now let us suppose that the ancient progenitor of our European cuckoo had the habits of the American cuckoo. By a continued process of this nature. If this were the case the process of laying and hatching might be inconveniently long. 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. for she makes her own nest and has eggs and young successively hatched. This rare event is probably a case of reversion to the long-lost. 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. and that she occasionally laid an egg in another bird's nest. but at intervals of two or three days. and the first hatched young would probably have to be fed by the male alone. together with a young jay in the nest of a blue jay (Garrulus cristatus). I believe that the strange instinct of our cuckoo has been generated. but I have lately heard from Dr. But in all cases. by Adolf Muller.4. speculation on an instinct 251 . so that. 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 thus be more successful in rearing their young. then the old birds or the fostered young would gain an advantage. and as both were nearly full feathered. all at the same time. Merrill. if she were to make her own nest and sit on her own eggs. It has been objected that I have not noticed other related instincts and adaptations of structure in the cuckoo. more especially as she migrates at a very early period. also recently been ascertained on sufficient evidence. sits on them and feeds her young. encumbered as she could hardly fail to be by having eggs and young of different ages at the same time. which are spoken of as necessarily co-ordinated.

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. from eight to ten lines in length. 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. as is more probable. 252 . or.parasitic American cuckoo alone were known. 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. is useless. when they lay their eggs in an open nest. it would assuredly have been added to those which it is assumed must all have been acquired together. the strength and a properly shaped back for ejecting its foster. we have learned something about three Australian species. which lay their eggs in other birds' nests. not exceeding those of the skylark—a bird about one-fourth as large as the cuckoo. with rare exceptions. The eggs of the Australian bronze cuckoo vary. as well as in size. but not rarely departs from it. has the instinct. Ramsay remarks that two of the Australian cuckoos. natural selection might have secured and fixed any advantageous variation.brothers. so as to have deceived certain foster-parents. that the young cuckoo. for these would have been more safely hatched and reared. soon after birth. for we have hitherto had no facts to guide us. Ramsay. Secondly.known to us only in a single species. This has been boldly called a beneficent arrangement. The chief points to be referred to are three: first. so that in this respect. Had our cuckoo invariably displayed the above instinct. according to Mr. lays only one egg in a nest. Now. that the common cuckoo. to an extraordinary degree in colour. as is shown by her laying her dull and pale-coloured eggs in the nest of the hedgewarbler with bright greenish-blue eggs. now. manifest a decided preference for nests containing eggs similar in colour to their own. so that the large and voracious young bird receives ample food. 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). owing to Mr. Ramsay's observations. in order that the young cuckoo may get sufficient food. that the eggs are remarkably small. Thirdly. it is not rare to find two and even three eggs in the same nest. which then perish from cold and hunger. The European species apparently manifests some tendency towards a similar instinct. In the bronze cuckoo the eggs vary greatly in size. Until recently the instincts of the European and of the non. if it had been of an advantage to this species to have laid eggs even smaller than those now laid.

in the act of ejecting its foster-brothers. during successive generations. as Owen has remarked. Mr. the habit having been afterwards improved. They either lay their eggs in the nest thus appropriated. They usually sit on their own eggs and rear their own young. Mr. or oddly enough build one for themselves on the top of it. Hudson. and transmitted to an earlier age. The sexes of Molothrus badius are stated by an excellent observer. and structure necessary for the work of ejection. With respect to the means by which this strange and odious instinct was acquired. the blind desire. Gould was formerly inclined to believe that the act of ejection was performed by the foster-parents themselves. sometimes to live promiscuously together in flocks. to receive as much food as possible soon after birth. 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. 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. if it were of great importance for the young cuckoo. I can see no special difficulty in its having gradually acquired. For if each part is liable to individual variations at all ages. as is probably the case. have parasitic habits like those of the cuckoo. while still blind and not able even to hold up its own head. 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. and both cases must stand or fall together with the whole theory of natural selection. Some species of Molothrus. for he has seen the young of 253 . The first step towards the acquisition of the proper instinct might have been mere unintentional restlessness on the part of the young bird. They either build a nest of their own or seize on one belonging to some other bird. the offspring of the foster-parents are commonly ejected from the nest within three days after the cuckoo is hatched. but Mr. and sometimes to pair. a widely distinct genus of American birds. or than in young snakes acquiring in their upper jaws. allied to our starlings. a transitory sharp tooth for cutting through the tough egg-shell. Hudson says it is probable that they are occasionally parasitic. and was again thrown out.In the case of the European cuckoo. occasionally throwing out the nestlings of the stranger. the strength. One of these was replaced in the nest by the observer. and the species present an interesting gradation in the perfection of their instincts. when somewhat advanced in age and strength. and as the latter at this age is in a most helpless condition.

as has already been remarked. are much more highly developed than those of the last. but as small consequences of one general law. has acquired instincts as perfect as those of the cuckoo. complete a nest for themselves. In this family several hen birds unite and lay first a few eggs in one nest and then in another. which are thus wasted. so that the young bird is securely reared. This habit is not very uncommon with the Gallinaceae. which they find in the appropriated nests. and throws some light on the singular instinct of the ostrich.this species following old birds of a distinct kind and clamouring to be fed by them. bonariensis. This instinct may probably be accounted for by the fact of the hens laying a large number of eggs. at intervals of two or three days. but it is remarkable that several together sometimes commence to build an irregular untidy nest of their own. and regularly lay their eggs in the nests of other kinds of bees. transition?" Various birds. Hudson has ascertained. moreover. namely. for they do not possess the pollen-collecting apparatus which would have been indispensable if they had stored up food for their own young. This bird. They have. whether of their own species or of their foster parents. as far as Mr. pecoris of North America. for a surprising number of eggs lie strewed over the plains. of the American ostrich. however. however. Hudson is a strong disbeliever in evolution. occasionally lay their eggs in the nests of other birds. Some species of Sphegidae (wasp-like insects) are likewise parasitic. has not as yet been perfected. Many bees are parasitic. Mr. but. The parasitic habits of another species of Molothrus. the extraordinary habit of pecking holes in the eggs. invariably lays its eggs in the nests of strangers. for it never lays more than one egg in a foster-nest. They never. This case is more remarkable than that of the cuckoo. and M. A third species. as in the case of the Molothrus bonariensis. Fabre has lately shown good reason for believing that. but he appears to have been so much struck by the imperfect instincts of the Molothrus bonariensis that he quotes my words. They often lay so many eggs—from fifteen to twenty—in the same foster-nest. "Must we consider these habits. but are still far from perfect. although the Tachytes nigra generally 254 . The instinct. as on the leaves of a large thistle. so that in one day's hunting I picked up no less than twenty lost and wasted eggs. and these are hatched by the males. that few or none can possibly be hatched. and asks. not as especially endowed or created instincts. They drop also many eggs on the bare ground. the M. as with the cuckoo. placed in singular ill-adapted situations. as far as it is known. the M. for these bees have not only had their instincts but their structure modified in accordance with their parasitic habits.

be not thus exterminated. 255 .makes its own burrow and stores it with paralysed prey for its own larvae. and if the insect whose nest and stored food are feloniously appropriated. it takes advantage of the prize. as with that of the Molothrus or cuckoo. when this insect finds a burrow already made and stored by another sphex. if of advantage to the species. and becomes for the occasion parasitic. In this case. I can see no difficulty in natural selection making an occasional habit permanent. yet that.

When the old nest is found inconvenient. a better observer even than his celebrated father. and found a few slaves in all. they could not even feed themselves. and the larvae and pupae are exposed. and put all to rights. and the workers or sterile females. sanguinea. Huber to be a slave-making ant. Huber then introduced a single slave (F. Another species. the slaves work energetically together with their masters in carrying them away to a place of safety. Slave-Making Instinct This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber. fusca). Hence. without their aid. F. that when Huber shut up thirty of them without a slave. but with plenty of the food which they like best. of the British Museum. and actually carry their masters in their jaws. do no other work. fusca) are found only in their own proper communities. and she instantly set to work. the species would certainly become extinct in a single year. The slaves are black and not above half the size of their red masters. When the nest is slightly disturbed. it is the slaves which determine the migration. They are incapable of making their own nests. and many perished of hunger. I opened fourteen nests of F. as any one may well be excused for doubting the existence of so extraordinary an instinct as that of making slaves. and with their larvae and pupae to stimulate them to work.5. Hence. though most energetic and courageous in capturing slaves. so that the contrast in their appearance is great. So utterly helpless are the masters. Although fully trusting to the statements of Huber and Mr. fed and saved the survivors. and they have to migrate. they did nothing. and have never been observed in the nests of F. Males and fertile females of the slave-species (F. it is clear that the slaves feel quite at home. and like their masters are much agitated and defend the nest: when the nest is much disturbed. to whom I am much indebted for information on this and other subjects. Smith. This ant is absolutely dependent on its slaves. the slaves occasionally come out. sanguinea. and its habits have been attended to by Mr. made some cells and tended the larvae. Formica sanguinea. The males and fertile females do no work of any kind. What can be more extraordinary than these well-ascertained facts? If we had not known of any other slave-making ant. was likewise first discovered by P. or of feeding their own larvae. it would have been hopeless to speculate how so wonderful an instinct could have been perfected. Smith. I tried to approach the subject in a sceptical frame of mind. This species is found in the southern parts of England. During the months of June and 256 . I will give the observations which I made in some little detail.

after all. This species is sometimes. sanguinea. Smith informs me that he has watched the nests at various hours during May. During the year 1860. June and August. the slaves are very few in number. may be constantly seen bringing in materials for the nest. probably depends merely on the slaves being captured in greater numbers in Switzerland than in England. with a few of these little yellow ants still clinging to the fragments of their nest. he considers them as strictly household slaves. probably in search of aphides or cocci. The latter ruthlessly killed their small opponents and carried their dead bodies as food to their nest. Hence. as Huber expressly states. during these months. fusca). Another day my attention was struck by about a score of the slave-makers haunting the same spot. and put them down on a bare spot near the place of combat. and. they had been victorious in their late combat. though present in large numbers in August. both in Surrey and Hampshire. At the same time I laid on the same place a small parcel of the pupae of another species. This difference in the usual habits of the masters and slaves in the two countries. either leave or enter the nest. and they alone open and close the doors in the morning and evening. rufescens. As. their principal office is to search for aphides. however. and I observed a few slaves mingled with their masters leaving the nest. they were eagerly seized and carried off by the tyrants. in the month of July.July. as in the case of F. they approached and were vigorously repulsed by an independent community of the slave species (F. which they ascended together. who had ample opportunities for observation. and food of all kinds. but Mr. F. and evidently not in search of food. I then dug up a small parcel of the pupae of F. and has never seen the slaves. I watched for many hours several nests in Surrey and Sussex. The masters. flava. the slaves in Switzerland habitually work with their masters in making the nest. One day I fortunately witnessed a migration of F. who perhaps fancied that. and marching along the same road to a tall Scotch-fir tree. but they were prevented from getting any pupae to rear as slaves. on the other hand. I came across a community with an unusually large stock of slaves. According to Huber. sometimes as many as three of these ants clinging to the legs of the slave-making F. on three successive years. twenty-nine yards distant. I thought that they might behave differently when more numerous. twenty-five yards distant. 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 never saw a slave either leave or enter a nest. though 257 . sanguinea from one nest to another. fusca from another nest.

but chiefly the slaves. The nest.rarely. and one was perched motionless with its own pupa in its mouth on the top of a spray of heath. in regard to the wonderful instinct of making slaves. as has been described by Mr. Now I was curious to ascertain whether F. and I have seen it ferociously attack other ants. fusca (showing that it was not a migration) and numerous pupae. and when they migrate. fusca. One evening I visited another community of F. In Switzerland the slaves and masters work together. but in about a quarter of an hour. does not determine its own migrations. though they did not need confirmation by me. must have been close at hand. sanguinea. the little ants attacked their big neighbours with surprising courage. Such are the facts. flava. and in the early part of the summer extremely few. the masters carry the slaves. whereas they were much terrified when they came across the pupae. possesses much fewer slaves. shortly after all the little yellow ants had crawled away. of F. whence I saw the last individual of F. for we have seen that they eagerly and instantly seized the pupae of F. and quickly ran away. sanguinea emerge. I traced a long file of ants burdened with booty. tend and milk as it may be called. Let it be observed what a contrast the instinctive habits of F. and cannot even feed itself: it is absolutely dependent on its numerous slaves. their 258 . and found a number of these ants returning home and entering their nests. sanguinea could distinguish the pupae of F. and the masters alone go on slave-making expeditions. which they rarely capture. Both in Switzerland and England the slaves seem to have the exclusive care of the larvae. fusca. and when I had accidentally disturbed both nests. sanguinea present with those of the continental F. Although so small a species. making and bringing materials for the nest: both. however. The latter does not build its own nest. for about forty yards back. In one instance I found to my surprise an independent community of F. rufescens. and it was evident that they did at once distinguish them. flava under a stone beneath a nest of the slave-making F. they took heart and carried off the pupae. sanguinea. it is very courageous. made into slaves. Formica sanguinea. or even the earth from the nest. but I was not able to find the desolated nest in the thick heath. carrying the dead bodies of F. flava. does not collect food for itself or its young. carrying a pupa. from those of the little and furious F. The masters determine when and where a new nest shall be formed. which they habitually make into slaves. Smith. fusca were rushing about in the greatest agitation. on the other hand. for two or three individuals of F. to a very thick clump of heath. an image of despair over its ravaged home.

aphides. is less aided by its slaves than the same species in Switzerland. When the instinct was once acquired. By what steps the instinct of F. which. So that the masters in this country receive much less service from their slaves than they do in Switzerland. and the foreign ants thus unintentionally reared would then follow their proper instincts. if scattered near their nests. it is possible that such pupae originally stored as food might become developed. 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. carry off pupae of other species. as we have seen. But as ants which are not slave-makers. to capture workers than to procreate them—the habit of collecting pupae. and do what work they could. and thus both collect food for the community. sanguinea. will. if carried out to a much less extent even than in our British F. as I have seen. originally for food. might by natural selection be strengthened and rendered permanent for the very different purpose of raising slaves. sanguinea originated I will not pretend to conjecture. If their presence proved useful to the species which had seized them—if it were more advantageous to this species. 259 . their slaves and larvae. In England the masters alone usually leave the nest to collect building materials and food for themselves.

some large cells of wax for holding honey. with the basal edges of its six sides bevelled so as to join an inverted pyramid. perhaps. But the important point to notice is. though this is effected by a crowd of bees working in a dark hive. and the following view may. who has shown that the form of the cell stands in close relation to the presence of adjoining cells. and have made their cells of the proper shape to hold the greatest possible amount of honey. Granting whatever instincts you please. to follow from a few simple instincts. The Melipona itself is intermediate in structure between the hive and humble bee. but will merely give an outline of the conclusions at which I have arrived. enter into the composition of the bases of three adjoining cells on the opposite side. sometimes adding to them short tubes of wax. He must be a dull man who can examine the exquisite structure of a comb. Cell-Making Instinct Of The Hive-Bee I will not here enter on minute details on this subject. is an hexagonal prism. These rhombs have certain angles. as is well known. carefully described and figured by Pierre Huber. We hear from mathematicians that bees have practically solved a recondite problem. Waterhouse. At the other end of the series we have the cells of the hive-bee. These latter cells are nearly spherical and of nearly equal sizes. with the least possible consumption of precious wax in their construction. so beautifully adapted to its end. without enthusiastic admiration. with fitting tools and measures. and. in addition. of three rhombs. In the series between the extreme perfection of the cells of the hivebee and the simplicity of those of the humble-bee. I think. and are aggregated into an irregular mass. I was led to investigate this subject by Mr. or even perceive when they are correctly made. and likewise making separate and very irregular rounded cells of wax. would find it very difficult to make cells of wax of the true form. we have the cells of the Mexican Melipona domestica. It has been remarked that a skilful workman. be considered only as a modification of his theory. in which the young are hatched. But the difficulty is not nearly so great as at first appears: all this beautiful work can be shown. which use their old cocoons to hold honey. but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells. it seems at first quite inconceivable how they can make all the necessary angles and planes. placed in a double layer: each cell.6. 260 . and see whether Nature does not reveal to us her method of work. Let us look to the great principle of gradation. At one end of a short series we have humble-bees. and the three which form the pyramidal base of a single cell on one side of the comb.

each cell consists of an outer spherical portion. and tells me that it is strictly correct:If a number of equal spheres be described with their centres placed in two parallel layers. if planes of intersection between the several spheres in both layers be formed. according as the cell adjoins two. but are of the same thickness as the outer spherical portions. three. is manifestly a gross imitation of the three-sided pyramidal base of the cell of the hive-bee. or more flat surfaces. and what is more important. it occurred to me that if the Melipona had made its spheres at some given distance from each other. It is obvious that the Melipona saves wax. if ever. Hence. drawn up from his information. there will result a double layer of hexagonal prisms united together by pyramidal bases formed of three rhombs. As in the cells of the hive-bee. and of two. then. but this is never permitted. Hence we may safely conclude that. and had made them of equal sizes and had arranged them symmetrically in a double layer. and in themselves not very 261 . so here. the resulting structure would have been as perfect as the comb of the hive-bee. Accordingly I wrote to Professor Miller. realised. from the centres of the six surrounding spheres in the same layer. as Huber has remarked.41421 (or at some lesser distance). and this pyramid. is very frequently and necessarily the case. the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. with the centre of each sphere at the distance of radius x sqrt(2) or radius x 1. if we could slightly modify the instincts already possessed by the Melipona. But I hear from Professor Wyman. When one cell rests on three other cells. Reflecting on this case. so much so. that whatever the typical form of the cell may be. who has made numerous careful measurements. it is rarely. and the rhombs and the sides of the hexagonal prisms will have every angle identically the same with the best measurements which have been made of the cells of the hive-bee. the three flat surfaces are united into a pyramid. for the flat walls between the adjoining cells are not double. from the spheres being nearly of the same size. and yet each flat portion forms a part of two cells. which. the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. and this geometer has kindly read over the following statement.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. that the accuracy of the workmanship of the bee has been greatly exaggerated. labour. and at the same distance from the centres of the adjoining spheres in the other and parallel layer. by this manner of building. three or more other cells. of Cambridge.

knife-edged ridge. As soon as this occurred. rectangular piece of wax. if they had been excavated to the same depth as in the former experiment. and began to build up flat walls of wax on the lines of intersection between the basins. as she already does her cylindrical cells. and put between them a long. but she is already so far enabled to judge of distance. the rims of the basins intersected or broke into each other. It was most interesting to observe that. seeing that she already does so to a certain extent. instead of a thick. and seeing what perfectly cylindrical burrows many insects make in wood. in the same way as before. coloured with vermilion. wherever several bees had begun to excavate these basins near together. appearing to the eye perfectly true or parts of a sphere. rectangular strip of wax: the bees instantly began to excavate minute circular pits in it. 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. through natural selection. apparently by turning round on a fixed point. and were in depth about one sixth of the diameter of the sphere of which they formed a part. But this theory can be tested by experiment. would have broken into each other from the opposite 262 . and this is the greatest difficulty. and as they deepened these little pits. so that each hexagonal prism was built upon the scalloped edge of a smooth basin. Tegetmeier. The bees instantly began on both sides to excavate little basins near to each other. thick. about the width of an ordinary cell). this bee would make a structure as wonderfully perfect as that of the hive-bee. 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. Following the example of Mr. and of equal sizes. they made them wider and wider until they were converted into shallow basins. and this would not be very surprising. instead of on the straight edges of a three-sided pyramid as in the case of ordinary cells. that she always describes her spheres so as to intersect to a certain extent. but the ridge of wax was so thin. the bees ceased to excavate. We must suppose the Melipona to arrange her cells in level layers. a thin and narrow. and of about the diameter of a cell.e. I separated two combs. that the bottoms of the basins. and we must further suppose. and then she unites the points of intersection by perfectly flat surfaces. I then put into the hive. We must suppose the Melipona to have the power of forming her cells truly spherical. her inimitable architectural powers.wonderful. that she can somehow judge accurately at what distance to stand from her fellow-labourers when several are making their spheres.

where I suppose that the bees had excavated too quickly. In some parts. that they could have effected this by gnawing away the convex side. circumferential wall or rim all round the comb.sides. and these flat bases. In one wellmarked instance. only small portions. and again examined the cell. They do not make the 263 . and they gnaw this away from the opposite sides. if the bees were to build for themselves a thin wall of wax. and by endeavouring to make equal spherical hollows. so that the basins. and they stopped their excavations in due time. had not been neatly performed. in order to have thus succeeded in leaving flat plates between the basins. and convex on the opposed side where the bees had worked less quickly. Now bees. and then stopping their work. and I suspect that the bees in such cases stand in the opposed cells and push and bend the ductile and warm wax (which as I have tried is easily done) into its proper intermediate plane. and had become PERFECTLY FLAT: it was absolutely impossible. do make a rough. they could make their cells of the proper shape. which were slightly concave on one side. by stopping work at the planes of intersection. in other parts. and allowed the bees to go on working for a short time. from the unnatural state of things. came to have flat bases. exactly along the planes of imaginary intersection between the basins on the opposite side of the ridge of wax. and I found that the rhombic plate had been completed. always working circularly as they deepen each cell. but never allowing the spheres to break into each other. formed by thin little plates of the vermilion wax left ungnawed. I put the comb back into the hive. perceiving when they have gnawed the wax away to the proper thinness. for I have noticed half-completed rhombs at the base of a just-commenced cell. however. were situated. from the extreme thinness of the little plate. From the experiment of the ridge of vermilion wax we can see that. I do not see that there is any difficulty in the bees. but the work. Considering how flexible thin wax is. 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. In ordinary combs it has appeared to me that the bees do not always succeed in working at exactly the same rate from the opposite sides. and thus flatten it. by excavating at the same rate. The bees. by standing at the proper distance from each other. large portions of a rhombic plate were thus left between the opposed basins. did not suffer this to happen. as far as the eye could judge. whilst at work on the two sides of a strip of wax. as soon as they had been a little deepened. as may be clearly seen by examining the edge of a growing comb.

that a multitude of bees all work together. flexures may sometimes be observed. 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. nearly in the proportion of three to two. Huber's statement. It seems at first to add to the difficulty of understanding how the cells are made. until the hexagonal walls are commenced. From all the cells. and they never complete the upper edges of the rhombic plates. with the utmost ultimate economy of wax. as far as I have seen. as the case may be. and if I had space. having a mean thickness. We see how important a part excavation plays in the construction of the cells. By the above singular manner of building. till a smooth. and then to begin cutting it away equally on both sides near the ground. on an average of twelve measurements made near the border of the comb. that the very first cell is excavated out of a little parallel-sided wall of wax. from twenty-one measurements. along the plane of intersection between two adjoining spheres. which will ultimately be left. the first commencement having always been a little hood of wax. very thin wall is left in the middle. We shall understand how they work. they always make the first rough wall from ten to twenty times thicker than the excessively thin finished wall of the cell. vary greatly in thickness. but I will not here enter on details. strength is continually given to the comb. but I am convinced of their accuracy. These walls. or the two plates. The manner in which the bees build is curious. being thus crowned by a strong coping of wax. as Professor Miller has kindly ascertained for me. Some of these statements differ from those made by the justly celebrated elder Huber. of 1/229 of an inch. and adding fresh cement on the summit of the ridge. corresponding in position to the planes of the rhombic basal plates of future cells. But the rough wall of wax has in every case to be finished off. the masons always piling up the cut-away cement. I could show that they are conformable with my theory. Even in the rude circumferential rim or wall of wax round a growing comb. is not. by supposing masons first to pile up a broad ridge of cement. I have several specimens showing clearly that they can do this. but only that one rhombic plate which stands on the extreme growing margin. one bee after 264 . being. 1/352 of an inch in thickness. whereas the basal rhomboidal plates are thicker. strictly correct. the bees can cluster and crawl over the comb without injuring the delicate hexagonal walls. by being largely gnawed away on both sides. both those just commenced and those completed. We shall thus have a thin wall steadily growing upward but always crowned by a gigantic coping.whole three-sided pyramidal base of any one cell at the same time.

placed directly under the middle of a comb growing downwards. sometimes recurring to a shape which they had at first rejected. 265 . Nor does there seem to me any great difficulty in a single insect (as in the case of a queen-wasp) making hexagonal cells. on a slip of wood. by striking imaginary spheres. by covering the edges of the hexagonal walls of a single cell. sweeping spheres or cylinders. all trying to sweep equal spheres. and then building up. all instinctively standing at the same relative distance from each other. as when two pieces of comb met at an angle. projecting beyond the other completed cells. 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. and worked into the growing edges of the cells all round. as far as I have seen. 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 it bears on a fact. how often the bees would pull down and rebuild in different ways the same cell. which seems at first subversive of the foregoing theory. This capacity in bees of laying down under certain circumstances a rough wall in its proper place between two justcommenced cells. 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.working a short time at one cell going to another. or leaving ungnawed. namely. in its strictly proper place. that the cells on the extreme margin of wasp-combs are sometimes strictly hexagonal. I was able practically to show this fact. but I have not space here to enter on this subject. is important. the planes of intersection between these spheres. a score of individuals work even at the commencement of the first cell. so that. always standing at the proper relative distance from the parts of the cells just begun. It was really curious to note in cases of difficulty. 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. with an extremely thin layer of melted vermilion wax. When bees have a place on which they can stand in their proper positions for working—for instance. The work of construction seems to be a sort of balance struck between many bees. and then. or the extreme margin of the circumferential rim of a growing comb. and building up intermediate planes. as Huber has stated. if she were to work alternately on the inside and outside of two or three cells commenced at the same time. they can build up a wall intermediate between two adjoining spheres.

and I am informed by Mr. whether a bee allied to our humble-bees could exist in large numbers in any country. many bees have to remain idle for many days during the process of secretion. Again. But let us suppose that this latter circumstance determined. and save much in labour and space. nearer together. if they were to make their cells more and more regular. Moreover. as it probably often has determined. all tending towards the present perfect plan of construction. A large store of honey is indispensable to support a large stock of bees during the winter.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. it would continually be more and more advantageous to our humble-bees. the 266 . or parasites. as we have seen. and in the materials of which they are constructed. it would be advantageous to the Melipona. Hence. or on quite distinct causes. and aggregated into a mass. if she were to make her cells closer together. and so be altogether independent of the quantity of honey which the bees can collect. Hence the saving of wax by largely saving honey. and more regular in every way than at present. and the security of the hive is known mainly to depend on a large number of bees being supported. it may reasonably be asked. for in this case a large part of the bounding surface of each cell would serve to bound the adjoining cells. for then. must be an important element of success any family of bees. With respect to the formation of wax. and much labour and wax would be saved. 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. 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. how a long and graduated succession of modified architectural instincts. from the same cause. and let us further suppose that the community lived through the winter. 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. it is known that bees are often hard pressed to get sufficient nectar. and the time consumed in collecting the honey. each profitable to the individual under its conditions of life. like the cells of the Melipona. Of course the success of the species may be dependent on the number of its enemies. so as to intersect a little. 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.

as I believe. this being effected with the greatest possible economy of labour and wax. and to build up and excavate the wax along the planes of intersection. and least waste of honey in the secretion of wax. the most wonderful of all known instincts. more and more perfectly led the bees to sweep equal spheres at a given distance from each other in a double layer. 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. and the Melipona would make a comb as perfect as that of the hive-bee.spherical surfaces would wholly disappear and be replaced by plane surfaces. no more knowing that they swept their spheres at one particular distance from each other. The bees. is absolutely perfect in economising labour and wax. Beyond this stage of perfection in architecture. for the comb of the hive-bee. that of the hive-bee. 267 . by slow degrees. natural selection could not lead. that individual swarm which thus made the best cells with least labour. than they know what are the several angles of the hexagonal prisms and of the basal rhombic plates. which in their turn will have had the best chance of succeeding in the struggle for existence. having succeeded best. slight modifications of simpler instincts. natural selection having. and having transmitted their newly-acquired economical instincts to new swarms. Thus. can be explained by natural selection having taken advantage of numerous. of course. successive. as far as we can see.

To take as an illustration the case of the larger titmouse. 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. It must. but in accordance with. (Parus major) alluded to in a previous chapter. subsequently to. cases. at the same time that habit. and even with flakes of this substance. which secreted more and more saliva. but let the feet of the titmouse vary and grow larger from correlation with the beak. 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. in which we cannot see how an instinct could have originated. believed to be moistened with saliva. 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. however. cases of instincts of such trifling 268 . and hammers with its beak till it gets at the kernel. in which no intermediate gradations are known to exist. or compulsion. 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. No doubt many instincts of very difficult explanation could be opposed to the theory of natural selection—cases. as well constructed for this purpose as that of the nuthatch. 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." The force of this objection rests entirely on the assumption that the changes in the instincts and structure are abrupt. slowly changing habits or taste. or from any other unknown cause. be admitted that in many instances we cannot conjecture whether it was instinct or structure which first varied. Some birds build their nests of mud. Is it then very improbable that the natural selection of individual swifts. In this case a gradual change of structure is supposed to lead to changed instinctive habits. until a beak was formed. 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. this bird often holds the seeds of the yew between its feet on a branch. Now what special difficulty would there be in natural selection preserving all the slight individual variations in the shape of the beak.7. and one of the swifts of North America makes its nest (as I have seen) of sticks agglutinated with saliva. or spontaneous variations of taste.

It may well be asked how it is possible to reconcile this case with the theory of natural selection? First. If a working ant or other neuter insect had been an ordinary animal. of all sorts of differences of inherited structure which are correlated with certain ages and with either sex. and yet. that they could hardly have been acted on by natural selection. but will confine myself to one special difficulty. they cannot propagate their kind. But with the working ant we have an insect differing greatly from its parents. let it be remembered that we have innumerable instances. which at first appeared to me insuperable. How the workers have been rendered sterile is a difficulty. yet absolutely sterile. for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile. which were inherited by the offspring. but incapable of procreation. We have differences correlated not only with one 269 .importance. I will not here enter on these several cases. that of working or sterile ants. and so onwards. by individuals having been born with slight profitable modifications. so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny. and actually fatal to the whole theory. 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. 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. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure. I should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection. and it had been profitable to the community that a number should have been annually born capable of work. namely. and in instinct. but I will here take only a single case. but not much greater than that of any other striking modification of structure. the wonderful difference in this respect between the workers and the perfect females would have been better exemplified by the hivebee. as in the shape of the thorax. and that these again varied and again were selected. and if such insects had been social. But I must pass over this preliminary difficulty. and consequently must believe that they were independently acquired through natural selection. and in being destitute of wings and sometimes of eyes. both in our domestic productions and in those in a state of nature. As far as instinct alone is concerned. from being sterile. The subject well deserves to be discussed at great length. I can see no especial difficulty in this having been effected through natural selection.

relatively to the length of the horns in both the bulls and cows of these same breeds. is lessened. Hence. Here is a better and real illustration: According to M. These latter. though appearing insuperable. always produce a large proportion of seedlings bearing double and quite sterile flowers. and transmitted to their fertile offspring a tendency to produce sterile members with the same modifications. as in the nuptial plumage of many birds. 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. could. and the double sterile plants with the neuters of the same community. from having been long and carefully selected to the right degree. be formed by carefully watching which individual bulls and cows. 270 . We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male sex. for the sake of gaining a serviceable end. This difficulty. always yielding oxen with extraordinarily long horns. and not to the individual. An animal thus characterized has been slaughtered. we may conclude that slight modifications of structure or of instinct. disappears. Hence. I can see no great difficulty in any character becoming correlated with the sterile condition of certain members of insect communities. but with that short period when the reproductive system is active. but the breeder has gone with confidence to the same stock and has succeeded. selection has been applied to the family. as I believe.sex. or. This process must have been repeated many times. by which alone the variety can be propagated. Breeders of cattle wish the flesh and fat to be well marbled together. as well as to the individual. the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection. correlated with the sterile condition of certain members of the community. some varieties of the double annual stock. consequently the fertile males and females have flourished. have proved advantageous. produced oxen with the longest horns. so with social insects. may be compared with the fertile male and female ants. when matched. and yet no one ox would ever have propagated its kind. As with the varieties of the stock. Such faith may be placed in the power of selection that a breed of cattle. Verlot. but they likewise yield some single and fertile plants. it is probable. for oxen of certain breeds have longer horns than the oxen of other breeds. when it is remembered that selection may be applied to the family. and may thus gain the desired end. and in the hooked jaws of the male salmon.

with jaws and instincts extraordinarily different: in Cryptocerus. while those of an intermediate size are scanty in numbers. or rather as any two genera of the same family. the use of which is quite unknown: in the Mexican Myrmecocystus. there are working and soldier neuters. Smith has shown that the neuters of several British ants differ surprisingly from each other in size and sometimes in colour. all the neuters ultimately came to be thus characterized. the workers of one caste alone carry a wonderful sort of shield on their heads. F. do not generally graduate into each other. Formica flava has larger and smaller workers. or the domestic cattle as they may be called. Thus. when I do not admit that such wonderful and well-established facts at once annihilate the theory. and are thus divided into two or even three castes. in this species. and that by the survival of the communities with females which produced most neuters having the advantageous modification. being as distinct from each other as are any two species of the same genus. and this we do find. with some few of intermediate size. supplying the place of that excreted by the aphides. even not rarely. whereas the smaller workers have 271 . and they have an enormously developed abdomen which secretes a sort of honey. profitable modifications did not first arise in all the neuters in the same nest. the larger workers have simple eyes (ocelli). According to this view we ought occasionally to find in the same nest neuter-insects. that the successive.But we have not as yet touched on the acme of the difficulty. which. and. but from each other. sometimes to an almost incredible degree. Mr. moreover. the workers of one caste never leave the nest. It sometimes happens that the larger or the smaller sized workers are the most numerous. but are perfectly well defined. in Eciton. namely. not only from the fertile females and males. the fact that the neuters of several ants differ. F. presenting gradations of structure. It will indeed be thought that I have an overweening confidence in the principle of natural selection. Smith has observed. In the simpler case of neuter insects all of one caste. which. which our European ants guard and imprison. can be plainly distinguished. considering how few neuter-insects out of Europe have been carefully examined. as Mr. slight. though small. 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. as I believe. we may conclude from the analogy of ordinary variations. have been rendered different from the fertile males and females through natural selection. but in some few alone. The castes. they are fed by the workers of another caste. or that both large and small are numerous.

though the male and female ants of this genus have well-developed ocelli. yet connected by some few members in an intermediate condition. or lastly. as Sir J. of the working ants of the several sizes differed wonderfully in shape. The jaws. I believe that natural selection. with the camera lucida. but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house. I speak confidently on this latter point. though I dare not assert so positively. Smith's offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa. So that here we have two bodies of sterile workers in the same nest. Bates. The reader will perhaps best appreciate the amount of difference in these workers by my giving. differing not only in size. 272 . F. by acting on the fertile ants or parents. that the workers of intermediate size have their ocelli in an exactly intermediate condition. and this is the greatest difficulty. and in the form and number of the teeth. which produced more and more of the smaller workers. could form a species which should regularly produce neuters. we should then have had a species of ant with neuters in nearly the same condition as those of Myrmica. all of large size with one form of jaw. and jaws nearly five times as big. 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. With these facts before me. but in their organs of vision. moreover. yet they graduate insensibly into each other." has described analogous cases. and I fully believe. that if the smaller workers had been the most useful to the community. Mr. of whom many were five feet four inches high. though the workers can be grouped into castes of different sizes. I may digress by adding. of the jaws which I dissected from the workers of the several sizes. For the workers of Myrmica have not even rudiments of ocelli. that I gladly availed myself of Mr. not the actual measurements. 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. But the important fact for us is that. 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. until all the workers were in this condition. as does the widely-different structure of their jaws. Having carefully dissected several specimens of these workers. and those males and females had been continually selected.their ocelli rudimentary. or all of small size with widely different jaws. Lubbock made drawings for me. in his interesting "Naturalist on the Amazons. and many sixteen feet high.

discussed this case. also. 273 . I believe. however. of certain Malayan butterflies regularly appearing under two or even three distinct female forms. For peculiar habits. and likewise because this is by far the most serious special difficulty which my theory has encountered. in order to show the power of natural selection. with all my faith in natural selection. as advanced by Lamarck. Wallace. An analogous explanation has been given by Mr.one set of workers of one size and structure. is very interesting. of the equally complex case. at some little but wholly insufficient length. as it proves that with animals. against the well-known doctrine of inherited habit. could not possibly affect the males and fertile females. slight. as in the case of the driver ant. without exercise or habit having been brought into play. confined to the workers of sterile females. which alone leave descendants. The case. through the survival of the parents which generated them. a graduated series having first been formed. as with plants. and then the extreme forms having been produced in greater and greater numbers. I have. But I must confess. work by inherited instincts and by inherited organs or tools. We can see how useful their production may have been to a social community of ants. has originated. until none with an intermediate structure were produced. however long they might be followed. I have now explained how. the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest. while man works by acquired knowledge and manufactured instruments. I should never have anticipated that this principle could have been efficient in so high a degree. and simultaneously another set of workers of a different size and structure. which are in any way profitable. that. any amount of modification may be effected by the accumulation of numerous. of certain Brazilian crustaceans likewise appearing under two widely distinct male forms. But this subject need not here be discussed. and by Fritz Muller. I am surprised that no one has advanced this demonstrative case of neuter insects. both widely different from each other and from their parents. had not the case of these neuter insects led me to this conclusion. on the same principle that the division of labour is useful to civilised man. spontaneous variations. therefore. Ants.

multiply.—a habit wholly unlike that of any other known bird. annihilate it. that no instinct can be shown to have been produced for the good of other animals. 274 . how it is that the thrush of tropical South America lines its nest with mud. Finally. vary. in natural selection accumulating to any extent slight modifications of instinct which are in any way useful. we can understand. like the males of our Kittywrens. but distinct. For instance. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. under changing conditions of life. as by that common case of closely allied. not as specially endowed or created instincts. and is plainly explicable on the foregoing views. that the canon in natural history. In many cases habit or use and disuse have probably come into play. in the same peculiar manner as does our British thrush. but is otherwise inexplicable—all tend to corroborate the theory of natural selection." to roost in. Therefore. on the principle of inheritance. when inhabiting distant parts of the world and living under considerably different conditions of life. though animals take advantage of the instincts of others. No one will dispute that instincts are of the highest importance to each animal. I do not pretend that the facts given in this chapter strengthen in any great degree my theory. This theory is also strengthened by some few other facts in regard to instincts. there is no real difficulty. build "cock-nests. let the strongest live and the weakest die. 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.8. Summary I have endeavoured in this chapter briefly to show that the mental qualities of our domestic animals vary. how it is that the male wrens (Troglodytes) of North America." is applicable to instincts as well as to corporeal structure. with only a small hole left in the plaster through which the males feed them and their young when hatched. the fact that instincts are not always absolutely perfect and are liable to mistakes. yet often retaining nearly the same instincts. but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its fosterbrothers. On the other hand. and that the variations are inherited. to the best of my judgment. of "Natura non facit saltum. species. but none of the cases of difficulty. it may not be a logical deduction. the larvae of ichneumonidae feeding within the live bodies of caterpillars. ants making slaves. but as small consequences of one general law leading to the advancement of all organic beings—namely.

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

In treating this subject. in the second case they are either not at all developed. more especially as the sterility of species when first crossed. that is of the forms known or believed to be descended from common parents. It is an incidental result of differences in the reproductive systems of the parentspecies. in order to prevent their confusion. Pure species have of course their organs of reproduction in a perfect condition. on the other hand. The distinction probably has been slurred over. of equal importance with the sterility of species. which is common to the two cases. and the sterility of the hybrids produced from them. when crossed. to a large extent fundamentally different. namely.1. have been specially endowed with sterility. In the first case the two sexual elements which go to form the embryo are perfect. This distinction is important. when the cause of the sterility. with reference to my theory. 276 . The subject is in many ways important for us. The fertility of varieties. by the preservation of successive profitable degrees of sterility. have their reproductive organs functionally impotent. when intercrossed. the sterility of species when first crossed. beyond the province of our reasoning powers. as far as the microscope reveals. have generally been confounded. or are imperfectly developed. is. and that of their hybrid offspring. as may be clearly seen in the state of the male element in both plants and animals. Hybrids. for species living together could hardly have been kept distinct had they been capable of freely crossing. and likewise the fertility of their mongrel offspring. for it seems to make a broad and clear distinction between varieties and species. two classes of facts. yet when intercrossed they produce either few or no offspring. owing to the sterility in both cases being looked on as a special endowment. cannot have been acquired. though the formative organs themselves are perfect in structure. The view commonly entertained by naturalists is that species. This view certainly seems at first highly probable. has to be considered. as I shall show.

also. that the fertility of pure species is so easily affected by various circumstances. that the sterility of various species when crossed is so different in degree and graduates away so insensibly. when intercrossed. and were kept in a chamber in his house. But causes of serious error here intervene: a plant. namely Kolreuter and Gartner. and artificially fertilised with their own pollen. Degrees Of Sterility First. as Gartner repeatedly crossed some forms. for Gartner gives in his table about a score of cases of plants which he castrated. Kolreuter makes the rule universal. must be castrated. It is certain. he unhesitatingly ranks them as varieties. for the sterility of species when crossed and of their hybrid offspring. quite fertile together. which the best botanists rank as varieties. and. Moreover. Gartner is obliged carefully to count the seeds. Kolreuter and Gartner. But in these and in many other cases. for in ten cases in which he found two forms. and. and (excluding all cases such as the Leguminosae. and the maximum produced by their hybrid offspring. It is impossible to study the several memoirs and works of those two conscientious and admirable observers. I think no better evidence of this can be required than that the two most experienced observers who have ever lived. what is often more important. such as the common red and blue pimpernels (Anagallis arvensis and coerulea). He always compares the maximum number of seeds produced by two species when first crossed. to be hybridised. as he believed. must be secluded in order to prevent pollen being brought to it by insects from other plants. and found them absolutely sterile. that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins. in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. but then he cuts the knot. on the other hand. in order to show that there is any degree of sterility. Nearly all the plants experimented on by Gartner were potted. considered by most authors as distinct species.2. That these processes are often injurious to the fertility of a plant cannot be doubted. and he disputes the entire fertility of Kolreuter's ten cases. we may doubt whether many species are really so sterile. who almost devoted their lives to this subject. without being deeply impressed with the high generality of some degree of sterility. with the average number produced by both pure parent-species in a state of nature. on the one hand. makes the rule equally universal. arrived at diametrically opposite conclusions in regard to some of the very same 277 . Gartner.

I have made so many experiments and collected so many facts. would be thus effected. for six or seven. and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences. I am strengthened in this conviction by a remarkable statement repeatedly made by Gartner. that I cannot doubt the correctness of this conclusion.forms. and on the other hand that very close interbreeding lessens their vigour and fertility. With respect to this decrease. and this would probably be injurious to their fertility. and as the parent-species. In regard to the sterility of hybrids in successive generations. by too close interbreeding. the visits of insects must be carefully prevented during the flowering season: hence hybrids. with the evidence from fertility adduced by different hybridisers. whenever complicated 278 . if left to themselves. but generally decreases greatly and suddenly. generally grow in the same garden. It can thus be shown that neither sterility nor fertility affords any certain distinction between species and varieties. already lessened by their hybrid origin. showing on the one hand that an occasional cross with a distinct individual or variety increases the vigour and fertility of the offspring. it may first be noticed that when any deviation in structure or constitution is common to both parents. sometimes decidedly increases. 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. pollen is as often taken by chance (as I know from my own experience) from the anthers of another flower. namely. Hybrids are seldom raised by experimentalists in great numbers. Moreover. But I believe that their fertility has been diminished in nearly all these cases by an independent cause. namely. and both sexual elements in hybrid plants are already affected in some degree. or other allied hybrids. though probably often on the same plant. so that a cross between two flowers. carefully guarding them from a cross with either pure parent. in the process of artificial fertilisation. and in one case for ten generations. The evidence from this source graduates away. notwithstanding the frequent ill effects from manipulation. that if even the less fertile hybrids be artificially fertilised with hybrid pollen of the same kind. and goes on increasing. their fertility. yet he asserts positively that their fertility never increases. Now. though Gartner was enabled to rear some hybrids. or by the same observer from experiments made during different years. as from the anthers of the flower itself which is to be fertilised. will generally be fertilised during each generation by pollen from the same flower. this is often transmitted in an augmented degree to the offspring.

" Mr. So that with some species. which vegetated freely. certain abnormal individuals. though this pollen can be proved to be perfectly sound by fertilising other plants or species. which I never saw to occur in a case of its natural fecundation. the strange fact of an increase of fertility in the successive generations of ARTIFICIALLY FERTILISED hybrids. a bulb of Hippeastrum aulicum produced four flowers. Herbert tried similar 279 . may. I think. Verbascum and Passiflora. in various orchids as shown by Mr. W. and Rev. namely. 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. whereas the pod impregnated by the pollen of the hybrid made vigorous growth and rapid progress to maturity. in a first cross between two distinct species. all the individuals are in this peculiar condition.experiments are in progress. and after a few days perished entirely. namely. Herbert. and this would have insured in each generation a cross with pollen from a distinct flower. that "every ovule in a pod of Crinum capense fertilised by C. He is as emphatic in his conclusion that some hybrids are perfectly fertile—as fertile as the pure parent-species—as are Kolreuter and Gartner that some degree of sterility between distinct species is a universal law of nature. In the genus Hippeastrum." So that here we have perfect. in contrast with those spontaneously self.fertilised. can easily be fertilised by the pollen from a distinct species. in Corydalis as shown by Professor Hildebrand. be in part accounted for by Herbert's great horticultural skill. so careful an observer as Gartner would have castrated his hybrids. Of his many important statements I will here give only a single one as an example. as I believe. or even more than commonly perfect fertility. Now let us turn to the results arrived at by a third most experienced hybridiser. three were fertilised by Herbert with their own pollen. be accounted for by too close interbreeding having been avoided. And thus. The difference in their results may. and in other species all the individuals. the Hon. can actually be hybridised much more readily than they can be fertilised by pollen from the same individual plant! To give one instance. but not by pollen from the same plant. and bore good seed. that individual plants of certain species of Lobelia. He experimented on some of the very same species as did Gartner. This case of the Crinum leads me to refer to a singular fact. namely. Scott and Fritz Muller. revolutum produced a plant. and by his having hot-houses at his command. either from the same plant or from another plant of the same hybrid nature.

Petunia. with respect to the fertility in successive generations of the more fertile hybrid animals. and the injurious influence of close interbreeding is thus prevented. if the genera of animals are as distinct from each other as are the genera of plants. the canary-bird has been crossed with nine distinct species of finches. always gone on decreasing in fertility in each successive generation. have been crossed. and always with the same result. more sterile. Mr. owing to few animals breeding freely under confinement. for he will find on their stigmas plenty of pollen brought from other flowers. and I am assured that many of them are perfectly fertile. so as to avoid the ill effects of close 280 .. 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. Herbert asserts that a hybrid from Calceolaria integrifolia and plantaginea. be borne in mind that. few experiments have been fairly tried: for instance. informs me that he raises stocks for grafting from a hybrid between Rhod. In regard to animals. deserve some notice. The practical experiments of horticulturists. yet many of these hybrids seed freely. and such alone are fairly treated. as Gartner believed to be the case. Any one may readily convince himself of the efficiency of insect agency by examining the flowers of the more sterile kinds of hybrid Rhododendrons. but. Calceolaria. It is notorious in how complicated a manner the species of Pelargonium. ponticum and catawbiense. much fewer experiments have been carefully tried than with plants. Fuchsia. for instance. which produce no pollen. for by insect agency the several individuals are allowed to cross freely with each other. or that their hybrids. that is." I have taken some pains to ascertain the degree of fertility of some of the complex crosses of Rhododendrons. Noble. though not made with scientific precision. should be perfectly fertile. I hardly know of an instance in which two families of the same hybrid have been raised at the same time from different parents. species most widely dissimilar in general habit. For instance. If our systematic arrangements can be trusted. as not one of these breeds freely in confinement. when fairly treated. C. "reproduces itself as perfectly as if it had been a natural species from the mountains of Chile. but the hybrids themselves are. These cases serve to show on what slight and mysterious causes the lesser or greater fertility of a species sometimes depends." Had hybrids. the fact would have been notorious to nurserymen.experiments during many years. Again. Horticulturists raise large beds of the same hybrid. however. and that this hybrid "seeds as freely as it is possible to imagine. I think. we have no right to expect that the first crosses between them and the canary. It should. Rhododendron. etc.

whether the several aboriginal species would at first have freely bred together and have produced quite fertile hybrids. On the contrary. but from different hatches. but analogy makes me greatly doubt. yet in many cases they are descended from two or more wild species. Quatrefages states that the hybrids from two moths (Bombyx cynthia and arrindia) were proved in Paris to be fertile inter se for eight generations. With our domesticated animals. where neither pure parent-species exists. with perhaps the exception of certain indigenous domestic dogs of South America. it is not at all surprising that the inherent sterility in the hybrids should have gone on increasing. all are quite fertile together. The hybrids from the common and Chinese geese (A. which was first propounded by Pallas. This was effected by Mr. Blyth and Captain Hutton. And in this case. M. Eyton. cygnoides). in opposition to the constantly repeated admonition of every breeder. species which are so different that they are generally ranked in distinct genera. From this fact we must conclude either that the aboriginal parent-species at first produced perfectly fertile hybrids. and can. and from Phasianus colchicus with P. hardly be doubted. Although I know of hardly any thoroughly well-authenticated cases of perfectly fertile hybrid animals. however. produce offspring. and in one single instance they have bred inter se. torquatus. This latter alternative. the various races when crossed together are quite fertile.interbreeding. indeed. In India. that whole flocks of these crossed geese are kept in various parts of the country. brothers and sisters have usually been crossed in each successive generation. they must certainly be highly or perfectly fertile. It is. yet. almost certain that our dogs are descended from several wild stocks. or that the hybrids subsequently reared under domestication became quite fertile. I have reason to believe that the hybrids from Cervulus vaginalis and Reevesii. when they can be got to breed together. who raised two hybrids from the same parents. are perfectly fertile. which are highly fertile when crossed with one of the parent-species. and from these two birds he raised no less than eight hybrids (grandchildren of the pure geese) from one nest. for instance. these cross-bred geese must be far more fertile. namely Mr. seems by far the most probable. have often bred in this country with either pure parent. and as they are kept for profit. for I am assured by two eminently capable judges. It has lately been asserted that two such distinct species as the hare and rabbit. So again I have lately acquired decisive evidence that the crossed offspring from the Indian humped and common cattle are inter se perfectly 281 .

considering all the ascertained facts on the intercrossing of plants and animals. The same remarks may be extended to the two chief races of the pig. Blyth on their differences in habits. it may be concluded that some degree of sterility. Finally. constitution. under our present state of knowledge. either give up the belief of the universal sterility of species when crossed. is an extremely general result. be considered as absolutely universal. and from the observations by Rutimeyer on their important osteological differences. these two forms must be regarded as good and distinct species. therefore.fertile. etc. or we must look at this sterility in animals. but as one capable of being removed by domestication. both in first crosses and in hybrids. voice.. 282 . not as an indelible characteristic. as well as from those by Mr. but that it cannot. We must.

as we have seen. There are many 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. beyond that which the plant's own pollen produces. considering how scanty our knowledge is in regard to hybrid animals. but only the barest outline of the facts can here be given. can be united with unusual facility. are generally very sterile. It is surprising in how many curious ways this gradation can be shown. The following conclusions are drawn up chiefly from Gartner's admirable work on the hybridisation of plants. yet these hybrids are remarkably sterile. it exerts no more influence than so much inorganic dust. Our chief object will be to see whether or not these laws indicate that species have been specially endowed with this quality. graduates from zero to perfect fertility. The hybrids raised from two species which are very difficult to cross. the pollen of different species applied to the stigma of some one species of the same genus. So in hybrids themselves. as in the genus Verbascum.3. up to nearly complete or even quite complete fertility. yields a perfect gradation in the number of seeds produced. It has been already remarked. and. by the pollen of one of the pure parent-species causing the flower of the hybrid to wither earlier than it otherwise would have done. and which rarely produce any offspring. a single fertile seed: but in some of these cases a first trace of fertility may be detected. but the parallelism between the difficulty of making a first cross. that the degree of fertility. 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. there are some which never have produced. On the other hand. and produce numerous hybrid offspring. and the early withering of the flower is well known to be a sign of incipient fertilisation. I have taken much pains to ascertain how far they apply to animals. and probably never would produce. When pollen from a plant of one family is placed on the stigma of a plant of a distinct family. From this extreme degree of sterility we have self-fertilised hybrids producing a greater and greater number of seeds up to perfect fertility. I have been surprised to find how generally the same rules apply to both kingdoms. in certain abnormal cases. both of first crosses and of hybrids. even with the pollen of the pure parents. even to an excess of fertility. there are 283 . and. in which two pure species. in order to prevent their crossing and blending together in utter confusion.

and having strongly marked differences in every part of the flower.species which can be crossed very rarely. 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. by no less than eight other species of Nicotiana. it depends in part upon the constitution of the individuals which happen to have been chosen for the experiment. Even within the limits of the same genus. Now the fertility of first crosses. Many analogous facts could be given. is largely governed by their systematic affinity. A multitude of cases could be given of very closely allied species which will not unite. and another genus. obstinately failed to fertilise. or to be fertilised. in any recognisable character. is sufficient to prevent two species crossing. and on the other hand. the general resemblance between species in structure and constitution. as Silene. for instance in Dianthus. the many species of Nicotiana have been more largely crossed than the species of almost any other genus. and of the hybrids produced from them. as Dianthus. are very fertile. for instance. This is clearly shown by hybrids never having been raised between species ranked by systematists in distinct families. The fertility. But the correspondence between systematic affinity and the facility of crossing is by no means strict. Even within the limits of the same genus. than is that of pure species. these two opposite cases occur. It can be shown that plants most widely different in habit and general appearance. No one has been able to point out what kind or what amount of difference. But the fertility of first crosses is likewise innately variable. So it is with hybrids. In the same family there may be a genus. acuminata. we meet with this same difference. in which very many species can most readily be crossed. both of first crosses and of hybrids. by very closely allied species generally uniting with facility. Annual and perennial plants. and in the cotyledons. but the hybrids. is more easily affected by unfavourable conditions. but Gartner found that N. even in the pollen. which is not a particularly distinct species. for it is not always the same in degree when the same two species are crossed under the same circumstances. when at last produced. and on the other hand of very distinct species which unite with the utmost facility. in the fruit. in which the most persevering efforts have failed to produce between extremely close species a single hybrid. can be crossed. or only with extreme difficulty. deciduous and evergreen 284 . or with extreme difficulty. By the term systematic affinity is meant.

moreover. longiflora. some species have a remarkable power of crossing with other species. to fertilise reciprocally M. yet generally differ in fertility in a small. and occasionally in a high degree. longiflora with the pollen of M. but Kolreuter tried more than two hundred times. Such cases are highly important. and then a mare by a male-ass: these two species may then be said to have been reciprocally crossed. and such hybrids. that is of any difference in their structure or constitution. are with rare exceptions extremely sterile.trees. though externally so like one of their pure parent-species. So again among hybrids which are usually intermediate in structure between their parents. I mean the case. of a female-ass being first crossed by a stallion. for they prove that the capacity in any two species to cross is often completely independent of their systematic affinity. which closely resemble one of their pure parents. the one species having first been used as the father and then as the mother. always closely resemble one of them. though of course compounded of the very same two species. can often be crossed with ease. during eight following years. Several other singular rules could be given from Gartner: for instance. for instance. though they rarely differ in external characters. By a reciprocal cross between two species. and utterly failed. excepting in their reproductive systems. but these two powers do not at all necessarily go together. and these hybrids are almost always utterly sterile. Thuret has observed the same fact with certain sea-weeds or Fuci. as is usual. instead of having. It is also a remarkable fact that hybrids raised from reciprocal crosses. other species of the same genus have a remarkable power of impressing their likeness on their hybrid offspring. exceptional and abnormal individuals sometimes are born. and the hybrids thus produced are sufficiently fertile. even when the other hybrids raised from seed from the same capsule have a considerable 285 . To give an instance: Mirabilis jalapa can easily be fertilised by the pollen of M. plants inhabiting different stations and fitted for extremely different climates. Several other equally striking cases could be given. jalapa. There are certain hybrids which. There is often the widest possible difference in the facility of making reciprocal crosses. found that this difference of facility in making reciprocal crosses is extremely common in a lesser degree. The diversity of the result in reciprocal crosses between the same two species was long ago observed by Kolreuter. Gartner. an intermediate character between their two parents. He has observed it even between closely related forms (as Matthiola annua and glabra) which many botanists rank only as varieties.

according as the one species or the other is used as the father or the mother. or even to fertility under certain conditions in excess. produced from reciprocal crosses often differ in fertility. These facts show how completely the fertility of a hybrid may be independent of its external resemblance to either pure parent. not strictly related to the facility of the first union between their parents. that. in the facility of effecting an union. seems a strange arrangement. that their fertility. has the production of hybrids been permitted? To grant to species the special power of producing hybrids. their fertility graduates from zero to perfect fertility. the differences being of so peculiar and limited a nature. that the fertility of hybrids is not related to the degree in which they resemble in external appearance either parent. on the other hand. and lastly. and occasionally the widest possible difference. This latter statement is clearly proved by the difference in the result of reciprocal crosses between the same two species. and other species cross with extreme difficulty. the male sexual 286 . which govern the fertility of first crosses and of hybrids. we see that when forms. that it is by no means always the same in degree in the first cross and in the hybrids produced from this cross. 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. both of first crosses and of hybrids.degree of fertility. when various species are crossed. for. appear to me clearly to indicate that the sterility. 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. is simply incidental or dependent on unknown differences in their reproductive systems. moreover. is innately variable. there is generally some difference. The foregoing rules and facts. and then to stop their further propagation by different degrees of sterility. besides being eminently susceptible to favourable and unfavourable conditions. which must be considered as good and distinct species. it may even be asked. Considering the several rules now given. are united. For why should the sterility be so extremely different in degree. The hybrids. in reciprocal crosses between the same two species. 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. 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.

though with difficulty. We have seen that the sterility of hybrids which have their reproductive organs in an imperfect condition. and not a specially endowed quality. but not in a reversed direction. for no one has been able to graft together trees belonging to quite distinct families. but will admit that it is incidental on differences in the laws of growth of the two plants. yet these two distinct classes of cases run to a large extent parallel. can usually. Even different varieties of the pear take with different degrees of facility on the quince. in the period of the flow or nature of their sap. The pear can be grafted far more readily on the quince. so it sometimes is in grafting. in the hardness of their wood. for Thouin found that three 287 . on the gooseberry.. is a different case from the difficulty of uniting two pure species. what I mean by sterility being incidental on other differences. be grafted with ease. cannot be grafted on the currant. It will be advisable to explain a little more fully. which is a member of the same genus.element of the one will often freely act on the female sexual element of the other. etc. whereas the currant will take. Although many distinct genera within the same family have been grafted together. does not always prevent the two grafting together. is by no means absolutely governed by systematic affinity. As in hybridisation. which is ranked as a distinct genus. the facility of effecting an union is often very far from equal. We can sometimes see the reason why one tree will not take on another from differences in their rate of growth. which have their reproductive organs perfect. As Gartner found that there was sometimes an innate difference in different INDIVIDUALS of the same two species in crossing. one being evergreen and the other deciduous. and. in other cases species of the same genus will not take on each other. so do different varieties of the apricot and peach on certain varieties of the plum. one being woody and the other herbaceous. and adaptation to widely different climates. But this capacity. As in reciprocal crosses. but not invariably. than on the apple. closely allied species and varieties of the same species. so Sagaret believes this to be the case with different individuals of the same two species in being grafted together. As the capacity of one plant to be grafted or budded on another is unimportant for their welfare in a state of nature. on the other hand. by an example. Something analogous occurs in grafting. as in hybridisation. but in a multitude of cases we can assign no reason whatever. the capacity is limited by systematic affinity. so with grafting. Great diversity in the size of two plants. The common gooseberry. for instance. I presume that no one will suppose that this capacity is a SPECIALLY endowed quality.

as might have been expected. We are reminded by this latter fact of the extraordinary cases of Hippeastrum. These differences in both cases follow. yet that there is a rude degree of parallelism in the results of grafting and of crossing distinct species. to a certain extent.species of Robinia. 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. which seed much more freely when fertilised with the pollen of a distinct species than when fertilised with pollen from the same plant. On the other hand. 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. when grafted on other species. certain species of Sorbus. etc. although in the case of crossing. Passiflora. 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. so I believe that the still more complex laws governing the facility of first crosses are incidental on unknown differences in their reproductive systems. which seeded freely on their own roots.. The facts by no means seem to indicate that the greater or lesser difficulty of either grafting or crossing various species has been a special endowment. when thus grafted were rendered barren. We thus see that. and which could be grafted with no great difficulty on a fourth species. yielded twice as much fruit as when on their own roots. by which term every kind of resemblance and dissimilarity between organic beings is attempted to be expressed. 288 . systematic affinity.

produced few and sterile offspring. In considering the probability of natural selection having come into action. while at the same time the male element of this second form is enabled freely to fertilise the first form. which. For it would clearly be advantageous to two varieties or incipient species if they could be kept from blending. if a species was rendered sterile with some one compatriot. it seems to me that 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. and consequently this could not have been effected through natural selection. that in reciprocal crosses the male element of one form should have been rendered utterly impotent on a second form. it is almost as much opposed to the theory of natural selection as to that of special creation. but it may perhaps be argued. what is there which could favour the 289 . for thus fewer bastardised and deteriorated offspring would be produced to commingle their blood with the new species in process of formation. it may be remarked that species inhabiting distinct regions are often sterile when crossed. when man is selecting at the same time two varieties. when crossed. now. Take the case of any two species which. sterility with other species would follow as a necessary contingency. from slightly lessened fertility to absolute sterility. in rendering species mutually sterile. It may be admitted that it would profit an incipient species. After mature reflection. Origin And Causes Of The Sterility Of First Crosses And Of Hybrids At one time it appeared to me probable. In the first place. that the sterility of first crosses and of hybrids might have been slowly acquired through the natural selection of slightly lessened degrees of fertility. will find the subject extraordinarily complex. the greatest difficulty will be found to lie in the existence of many graduated steps. and which is universal with species which have been differentiated to a generic or family rank. 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. In the second place. spontaneously appeared in certain individuals of one variety when crossed with those of another variety. on the same principle that. it is necessary that he should keep them separate. as it has to others. that. if it were rendered in some slight degree sterile when crossed with its parent form or with some other variety. now it could clearly have been of no advantage to such separated species to have been rendered mutually sterile. like any other variation.4.

if rendered slightly sterile when crossed with some other variety. which have already ceased to yield seeds. would not thus itself gain any advantage or indirectly give any advantage to the other individuals of the same variety. as seems to have been the case with some of Thuret's 290 . the greater or less difficulty in effecting a union and in obtaining offspring apparently depends on several distinct causes. It is here manifestly impossible to select the more sterile individuals. is the same or nearly the same in all cases. Again.survival of those individuals which happened to be endowed in a slightly higher degree with mutual infertility. from an advantage having been thus indirectly given to the community to which they belonged over other communities of the same species. though the pollen-tubes protrude. cannot have been gained through selection. Both Gartner and Kolreuter have proved that in genera including numerous species. It has also been observed that when the pollen of one species is placed on the stigma of a distantly allied species. a series can be formed from species which when crossed yield fewer and fewer seeds. In the case of first crosses. we may infer that the cause. when the germen alone is effected. thus leading to their preservation. for the germen swells. and from the laws governing the various grades of sterility being so uniform throughout the animal and vegetable kingdoms. whatever it may be. the male element may reach the female element. There must sometimes be a physical impossibility in the male element reaching the ovule. but be incapable of causing an embryo to be developed. but an individual animal not belonging to a social community. 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. as would be the case with a plant having a pistil too long for the pollen-tubes to reach the ovarium. for a multitude are mutually quite barren. they do not penetrate the stigmatic surface. 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. With sterile neuter insects we have reason to believe that modifications in their structure and fertility have been slowly accumulated by natural selection. but yet are affected by the pollen of certain other species. to species which never produce a single seed. and which thus approached by one small step towards absolute sterility? Yet an advance of this kind. so that this acme of sterility. quite independent of natural selection. if the theory of natural selection be brought to bear. must have incessantly occurred with many species.

or within the egg or seed produced by the mother. rather than in the conditions to which it is subsequently exposed. 291 . more especially as all very young beings are eminently sensitive to injurious or unnatural conditions of life. are differently circumstanced before and after birth: when born and living in a country where their two parents live. the majority of these eggs had been fertilised. and in the majority of the fertilised eggs. the cause more probably lies in some imperfection in the original act of impregnation. who has had great experience in hybridising pheasants and fowls. Hewitt. and then perish at an early period.experiments on Fuci. as we see in the case of the common mule. any more than why certain trees cannot be grafted on others. it may therefore. No explanation can be given of these facts. more than four-fifths died within the first few days. Of the chickens which were born. that the early death of the embryo is a very frequent cause of sterility in first crosses. pass through their early stages of development and then perish like the embryos produced by a cross between distinct species. or at latest weeks. Hybrids. "without any obvious cause. Lastly. and perish at an early age. hybridized embryos probably often perish in a like manner. be exposed to conditions in some degree unsuitable. Until becoming acquainted with these facts. however. are generally healthy and long-lived. but the young chickens had been unable to break through the shell. for hybrids. Mr. as long as it is nourished within its mother's womb." so that from the 500 eggs only twelve chickens were reared. causing the embryo to be imperfectly developed. But after all. This latter alternative has not been sufficiently attended to. or had become nearly mature. With plants. when once born. before birth. the embryos had either been partially developed and had then perished. It may be here worth noticing that in some cases of parthenogenesis. at least it is known that hybrids raised from very distinct species are sometimes weak and dwarfed. apparently from mere inability to live. I was unwilling to believe in the frequent early death of hybrid embryos. But a hybrid partakes of only half of the nature and constitution of its mother. of which fact Max Wichura has recently given some striking cases with hybrid willows. but I believe. they are generally placed under suitable conditions of life. the embryos within the eggs of silk moths which had not been fertilised. and consequently be liable to perish at an early period. from observations communicated to me by Mr. an embryo may be developed. 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.

On the other hand. When hybrids are able to breed inter se. the tendency goes to a certain extent with systematic affinity. in fact. and certain species in a group will produce unusually fertile hybrids. when animals and plants are removed from their natural conditions. or periodical action. Lastly. Between the sterility thus superinduced and that of hybrids. without some disturbance occurring in the development. So it is with hybrids. In the one case. or that of hybrids. or any exotic plant seed freely under culture. though in a lesser degree than when sterility ensues. though often in so slight a degree as to be inappreciable by us. In both cases the sterility occurs in various degrees. but sometimes the female more than the male. No one can tell till he tries. For it is scarcely possible that two organisations should be compounded into one. when organic beings are placed during several generations under conditions not natural to them. as every experimentalist has observed. the external conditions have remained the same. but the organisation has been disturbed by two distinct structures and constitutions. they are extremely liable to have their reproductive systems seriously affected. the reproductive system. in the other case. This. for whole groups of animals and plants are rendered impotent by the same unnatural conditions. or mutual relations of the different parts and organs one to another or to the conditions of life. the conditions of life have been disturbed. is affected in a very similar manner. whether any two species of a genus will produce more or less sterile hybrids. in both. In both cases the sterility is independent of general health. and when hybrids are produced by the unnatural crossing of two species. one species in a group will sometimes resist great changes of conditions with unimpaired fertility. independently of the general state of health. which seems to be partly due to their reproductive systems having been specially affected. having been blended into one. the male element is the most liable to be affected. they are extremely liable to vary. for their offspring in successive generations are eminently liable to vary. and is often accompanied by excess of size or great luxuriance. in which the sexual elements are imperfectly developed. nor can he tell till he tries.In regard to the sterility of hybrids. and whole groups of species tend to produce sterile hybrids. In both. whether any particular animal will breed under confinement. including of course the reproductive systems. is the great bar to the domestication of animals. they transmit to their offspring from generation to generation the same 292 . there are many points of similarity. Thus we see that when organic beings are placed under new and unnatural conditions. the case is somewhat different. I have more than once alluded to a large body of facts showing that.

A similar parallelism holds good with an allied yet very different class of facts. and hence we need not be surprised that their sterility. if these be kept under the same conditions of life. tubers.compounded organisation. which have been subjected to slightly different conditions. does not diminish. of too close interbreeding. there is the clearest evidence that a cross between individuals of the same species. very frequently are rendered more 293 . on the above or any other view. weakness. It must. be owned that we cannot understand. as we have seen. crosses between the males and females of the same species. etc.. to a considerable change in their conditions. that slight changes in the conditions of life are beneficial to all living things. and back again. from one soil or climate to another. But. We see this acted on by farmers and gardeners in their frequent exchanges of seed. which I have elsewhere given. Hence it seems that. when placed under unnatural conditions. sterility is the common result—in the one case from the conditions of life having been disturbed. and that close interbreeding continued during several generations between the nearest relations. the unequal fertility of hybrids produced from reciprocal crosses. During the convalescence of animals. several facts with respect to the sterility of hybrids. great benefit is derived from almost any change in their habits of life. The above view of the sterility of hybrids being caused by two constitutions being compounded into one has been strongly maintained by Max Wichura. Again. give vigour and fertility to the offspring. is rendered sterile. or which have slightly varied. in the other case from the organisation having been disturbed by two organisations being compounded into one. Nor do I pretend that the foregoing remarks go to the root of the matter: no explanation is offered why an organism. All that I have attempted to show is. as under confinement. or sterility. and on the other hand. gives vigour and fertility to the offspring. almost always leads to decreased size. which differ to a certain extent. this being generally the result. for instance. as before explained. it is even apt to increase. when subjected. organic beings long habituated to certain uniform conditions under a state of nature. that slight crosses. though in some degree variable. on the one hand. slight changes in the conditions of life benefit all organic beings. both with plants and animals. It is an old and almost universal belief. however. or the increased sterility in those hybrids which occasionally and exceptionally resemble closely either pure parent. in some respects allied. that in two cases. that is. founded on a considerable body of evidence.

which. the incessant action and reaction of various forces. as throughout nature. and a multitude of other animals. 294 . 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. Herbert Spencer. and when this tendency is slightly disturbed by any change.or less sterile. He will at the same time be able to explain how it is that the races of some of our domesticated animals. are always tending towards an equilibrium. are incapable of breeding when kept under only partial confinement in their native country. produce hybrids which are almost always in some degree sterile. although they are descended from distinct species. He who is able to explain why the elephant. which is essentially related to the principle of life. being that life depends on. will be able to explain the primary cause of hybrids being so generally sterile. are quite fertile together. or consists in. according to Mr. this principle. The above two parallel series of facts seem to be connected together by some common but unknown bond. the vital forces gain in power. which would probably have been sterile if aboriginally crossed. I am fully persuaded that this double parallelism is by no means an accident or an illusion.

just in the same manner as occurs in crossing distinct species. Now I have shown. had wholly destroyed 295 . its action is so strongly prepotent that it generally annihilates the effect of the foreign pollen. As the degree of sterility in the latter case depends in an eminent degree on the conditions of life being more or less favourable. even after a considerable interval of time. and all the seedlings were similarly coloured. first illegitimately. placed on the same stigma. when both are placed on the same stigma. and two. With trimorphic plants there are three forms likewise differing in the lengths of their pistils and stamens. in the size and colour of the pollen-grains. up to absolute and utter sterility. and twenty-four hours afterwards legitimately. with pollen taken from a peculiarly coloured variety. are fully fertile. I ascertained this by fertilising several flowers. the other a short pistil with long stamens. and the result has been confirmed by other observers. So that with dimorphic species two unions. Reciprocal Dimorphism And Trimorphism This subject may be here briefly discussed. and as in each of the three forms there are two sets of stamens. that in order to obtain full fertility with these plants. and in some other respects. one form having a long pistil with short stamens. for legitimate pollen is strongly prepotent over illegitimate pollen. though applied twenty-four hours subsequently. differs much in degree. 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. With trimorphic species six unions are legitimate. and twelve are illegitimate. Several plants belonging to distinct orders present two forms. and its own pollen be afterwards. which may be called legitimate. which may be called illegitimate. which exist in about equal numbers and which differ in no respect except in their reproductive organs. 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. the two having differently sized pollen-grains. or more or less infertile.5. this shows that the legitimate pollen. so I have found it with illegitimate unions. when they are illegitimately fertilised. or fully fertile. the three forms possess altogether six sets of stamens and three kinds of pistils. and will be found to throw some light on hybridism. so it is with the pollen of the several forms of the same species. It is well known that if pollen of a distinct species be placed on the stigma of a flower. The infertility which may be observed in various dimorphic and trimorphic plants. that is by pollen taken from stamens not corresponding in height with the pistil. are more or less infertile.

many hybrids are profuse and persistent flowerers. It is possible to raise from dimorphic species. 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. It is hardly an exaggeration to maintain that illegitimate plants are hybrids. some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even seed-capsule. but the latter form did not yield a single seed when fertilised by the longer stamens of the mid-styled form. are not fully fertile. When this is done. exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants. produced within the limits of the same species by the improper union of certain forms. both long-styled and short-styled illegitimate plants. Altogether there is the closest identity in character and behaviour between illegitimate plants and hybrids. and yielded many seeds. so that sterility of certain illegitimate plants was unusually great. miserable dwarfs. With hybrids raised from the same seed. there is occasionally a great difference in the result. when illegitimately united. the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. The chief result is that these illegitimate plants. Again. and from trimorphic plants all three illegitimate forms. there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. behave in exactly the same manner as do two distinct species when crossed. They are all infertile. and in others which might be added. In all these respects. raised from several illegitimate unions. 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. But such is not the case. the forms of the same undoubted species. so the same thing occurs with trimorphic plants. may be strictly compared with that of hybrids when crossed inter se. as they may be called. on the other hand. for instance.capsule the degree of sterility is innately variable. as in making reciprocal crosses between the same two species. This led me carefully to observe during four years many seedlings. while other and more sterile hybrids produce few flowers. Lastly. while the sterility of the union from which they were derived was by no means great. while 296 . These can then be properly united in a legitimate manner. so it is in a marked manner with illegitimate plants. If.or prevented the action of the previously applied illegitimate pollen. when united with each other in a legitimate manner. and are weak. The sterility of these illegitimate plants. a hybrid is crossed with either pure parent-species.

styled forms. either in structure or in constitution. in the ordinary unions of the individuals of the same species and with crosses between distinct species. and this seems to me of especial importance. of two long. The facts now given on dimorphic and trimorphic plants are important. relatively to external conditions. He would find that they yielded only about one-fifth of the proper number of seed. however. infer as probable from the consideration of dimorphic and trimorphic plants. doubtful whether this is really so. first. and that they behaved in all the other above specified respects as if they had been two distinct species.ordinary hybrids are produced from an improper union between socalled distinct species. and yet be sterile when united in certain ways. but I will not enlarge on this obscure subject. and we are led to extend the same view to first crosses and hybrids. depends exclusively on the nature of their sexual elements. he would raise plants from his supposed hybridised seed. thirdly. that the sterility of distinct species when crossed and of their hybrid progeny. and not on any difference in their structure or general constitution. 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. while it is the union of the sexual elements proper to two distinct forms which is fertile. that the physiological test of lessened fertility. This will perhaps be made more fully apparent by an illustration. We have also already seen that there is the closest similarity in all respects between first illegitimate unions and first crosses between distinct species. But to make the case sure. He might then maintain that he had actually proved. is no safe criterion of specific distinction. and he would find that the seedlings were miserably dwarfed and utterly sterile. which results in sterility. We may. It is. because they show us. We are also led to this same conclusion by 297 . Hence the case appears at first sight exactly the reverse of what occurs. we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lythrum salicaria. because we may conclude that there is some unknown bond which connects the infertility of illegitimate unions with that of their illegitimate offspring. because we find. for instance. and that they behaved in all other respects like ordinary hybrids. both in first crosses and in hybrids. however. that two or three forms of the same species may exist and may differ in no respect whatever. and that he determined to try by crossing whether they were specifically distinct. that his two varieties were as good and as distinct species as any in the world. secondly. but he would be completely mistaken.

Gartner. That excellent observer. in which the male of one species cannot be united. while the converse cross can be effected with perfect facility. or can be united with great difficulty.considering reciprocal crosses. 298 . with the female of a second species. likewise concluded that species when crossed are sterile owing to differences confined to their reproductive systems.

so far is 299 . Several considerations. And Of Their Mongrel Offspring. those of the pigeon. have had so little tendency towards modifying the reproductive system in a manner leading to mutual sterility. however much they may differ from each other in external appearance. the fertility of all varieties produced under nature will assuredly have to be granted. if two forms hitherto reputed to be varieties be found in any degree sterile together. render the fertility of domestic varieties less remarkable. It is certain that with species the cause lies exclusively in differences in their sexual constitution. which. which are considered by most botanists as varieties. so that similar differences in the case of varieties would be no sure guide. become perfectly fertile together. under domestication. for. are utterly sterile when intercrossed. are said by Gartner to be quite sterile when crossed. we are still involved in some doubt. produced. the blue and red pimpernel. is a remarkable fact. and yield perfectly fertile offspring. namely. and probably the true one. With plants. the explanation which will occur to everyone. looking to varieties produced under nature. for instance.6. and he consequently ranks them as undoubted species. presently to be given. In the first place. which in their natural state probably would have been in some degree sterile when crossed. or of the cabbage. so that the domesticated descendants of species. that such conditions generally eliminate this tendency. But the subject is surrounded by difficulties. or supposed to have been produced. For instance. Now the varying conditions to which domesticated animals and cultivated plants have been subjected. For when it is stated. 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. more especially when we reflect how many species there are. though resembling each other most closely. cross with perfect facility. that certain South American indigenous domestic dogs do not readily unite with European dogs. With some exceptions. they are at once ranked by most naturalists as species. Fertility Of Varieties When Crossed. is that they are descended from aboriginally distinct species. for instance. that we have good grounds for admitting the directly opposite doctrine of Pallas. however. If we turn to varieties. I fully admit that this is the rule. it may be observed that the amount of external difference between two species is no sure guide to their degree of mutual sterility. differing widely from each other in appearance. Nevertheless the perfect fertility of so many domestic races. If we thus argue in a circle.

are rendered sterile. it becomes in the highest degree improbable that similar conditions long. and which can now generally resist with undiminished fertility repeated changes of conditions. as shown by the mere fact of their domestication. but why this has so generally occurred with natural varieties. 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.authenticated cases already alluded to. The evidence 300 . while still retaining the capacity of fertilising. sterility might occasionally be thus caused. But it is impossible to resist the evidence of the existence of a certain amount of sterility in the few following cases. and this may well make a wide difference in the result. For we know how commonly wild animals and plants. and why with plants only a few such cases. other species. with domesticated animals.cultivation from giving a tendency towards sterility between distinct species. with species having a peculiar constitution. that in several well. seeing how profoundly ignorant we are in regard to the normal and abnormal action of the reproductive system. why domestic varieties have not become mutually infertile when crossed. immediately to be given. certain plants have been affected in an opposite manner. 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. on the other hand. for they have become self-impotent. Thus. varieties have not been produced which are mutually sterile. We are far from precisely knowing the cause. though in certain cases. as soon as they have been permanently modified in a sufficient degree to take rank as species.continued should likewise induce this tendency. which I will briefly abstract. have been observed. I have as yet spoken as if the varieties of the same species were invariably fertile when intercrossed. than have domestic varieties. and it can hardly be rejected. The real difficulty in our present subject is not. If the Pallasian doctrine of the elimination of sterility through long-continued domestication be admitted. were not originally highly sensitive to changes in their conditions of life. owing to their struggle for existence with numerous competitors. Domesticated productions. and being fertilised by. we can understand why. when taken from their natural conditions and subjected to captivity. nor is this surprising. will have been exposed during long periods of time to more uniform conditions. But we can see that species. as it appears to me. might be expected to produce varieties. which. as I believe.

when crossed with a widely distinct species. a dwarf kind of maize with yellow seeds. whose accuracy has been confirmed by every subsequent observer. except in the colour of their flowers. No one. he finds that the dissimilarly coloured varieties of the same species yield fewer seeds. that the yellow and white varieties when crossed produce less seed than the similarly coloured varieties of the same species. has proved the remarkable fact that one particular variety of the common tobacco was more fertile than the other varieties. as the plants have separated sexes. and seems at first incredible. Mr. than between those which are differently coloured. when yellow and white varieties of one species are crossed with yellow and white varieties of a DISTINCT species. Scott also has experimented on the species and varieties of Verbascum. Kolreuter.is also derived from hostile witnesses. he asserts that. during several years. He then fertilised thirteen flowers of the one kind with pollen of the other. He experimented on five forms which are commonly reputed to be varieties. I know not. and one variety can sometimes be raised from the seed of another. as varieties. and although unable to confirm Gartner's results on the crossing of the distinct species. but only a single head produced any seed. which like the maize has separated sexes. who in all other cases consider fertility and sterility as safe criterions of specific distinction. and it is important to notice that the hybrid plants thus raised were themselves PERFECTLY fertile. they never naturally crossed. I believe. than the similarly coloured varieties. and which he tested by the 301 . who mainly founds his classification by the test of infertility. How far these experiments may be trusted. more seed is produced by the crosses between the similarly coloured flowers. The following case is far more remarkable. and Naudin has come to the same conclusion. Girou de Buzareingues crossed three varieties of gourd. by so good an observer and so hostile a witness as Gartner: namely. Gartner kept. has suspected that these varieties of maize are distinct species. and he asserts that their mutual fertilisation is by so much the less easy as their differences are greater. but the forms experimented on are ranked by Sagaret. Manipulation in this case could not have been injurious. in the proportion of eighty-six to 100. Yet these varieties differ in no respect. but it is the result of an astonishing number of experiments made during many years on nine species of Verbascum. and this one head produced only five grains. and a tall variety with red seeds growing near each other in his garden. Moreover. so that even Gartner did not venture to consider the two varieties as specifically distinct. and although these plants have separated sexes.

by reciprocal crosses. But one of these five varieties. but as incidental on changes of an unknown nature in their sexual elements. 302 . not as a special acquirement or endowment. From the great difficulty of ascertaining the infertility of varieties in a state of nature. From these facts it can no longer be maintained that varieties when crossed are invariably quite fertile. and he found their mongrel offspring perfectly fertile. and crossed with the Nicotiana glutinosa. always yielded hybrids not so sterile as those which were produced from the four other varieties when crossed with N. when used either as the father or mother.severest trial. for a supposed variety. The general sterility of crossed species may safely be looked at. namely. from these several considerations we may conclude that fertility does not constitute a fundamental distinction between varieties and species when crossed. would almost universally be ranked as a species. if proved to be infertile in any degree. and from such varieties not having been exposed for very long periods to uniform conditions of life. Hence the reproductive system of this one variety must have been in some manner and in some degree modified. from man attending only to external characters in his domestic varieties. glutinosa.

and I have myself seen striking instances of this fact. is a curious fact and deserves attention. and they are not variable. but Gartner admits that hybrids from species which have long been cultivated are often variable in the first generation. from being eminently sensitive to changed conditions of life. but some few instances of both hybrids and mongrels long retaining a uniform character could be given. and mostly domestic varieties (very few experiments having been tried on natural varieties). and this implies that there has been recent variability. that in the first generation mongrels are more variable than hybrids. Independently Of Their Fertility Independently of the question of fertility. the offspring of species and of varieties when crossed may be compared in several other respects. whose strong wish it was to draw a distinct line between species and varieties. namely. fails under these circumstances to perform its proper function of producing offspring closely similar in all respects to the parent-form. Gartner. and their descendants are highly variable. For it bears on the view which I have taken of one of the causes of ordinary variability. The variability. And. Now. that the reproductive system. This greater variability in mongrels than in hybrids does not seem at all surprising. The slight variability of hybrids in the first generation. which would often continue and would augment that arising from the act of crossing. and. on the other hand. in contrast with that in the succeeding generations. they agree most closely in many important respects. For the parents of mongrels are varieties. hybrids in the first generation are descended from species (excluding those long cultivated) which have not had their reproductive systems in any way affected. Hybrids And Mongrels Compared. 303 . I shall here discuss this subject with extreme brevity.7. Gartner further admits that hybrids between very closely allied species are more variable than those from very distinct species. When mongrels and the more fertile hybrids are propagated for several generations. perhaps. but hybrids themselves have their reproductive systems seriously affected. as it seems to me. The most important distinction is. could find very few. and the socalled mongrel offspring of varieties. in the successive generations of mongrels is. an extreme amount of variability in the offspring in both cases is notorious. however. and this shows that the difference in the degree of variability graduates away. quite unimportant differences between the so-called hybrid offspring of species. greater than in hybrids.

is founded on a single experiment. On the other hand. more especially in hybrids produced from nearly related species. Gartner further states that when any two species. follow. and with animals. partly owing to the existence of secondary sexual characters. one has sometimes a prepotent power of impressing its likeness on the hybrid. and seems directly opposed to the results of several experiments made by Kolreuter. but this. the hybrids are widely different from each other. although most closely allied to each other. but more especially owing to prepotency in transmitting likeness running more strongly in one sex than in the other. But this conclusion. Both hybrids and mongrels can be reduced to either pure parent form. by repeated crosses in successive generations with either parent. When two species are crossed. if it be true. Thus Max Wichura doubts whether hybrids ever revert to their parent forms. are crossed with a third species. Such alone are the unimportant differences which Gartner is able to point out between hybrid and mongrel plants. and he experimented on uncultivated species of willows.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. as far as I can make out. and he experimented chiefly on cultivated plants. So I believe it to be with varieties of plants. while Naudin. I think those authors are right who maintain that the ass has a prepotent power over the horse. the degrees and kinds of resemblance in mongrels and in hybrids to their respective parents. insists in the strongest terms on the almost universal tendency to reversion in hybrids. Hybrid plants produced from a reciprocal cross generally resemble each other closely. whereas if two very distinct varieties of one species are crossed with another species. Moreover. is certainly only a difference in degree. These several remarks are apparently applicable to animals. so that both the mule and the hinny resemble more closely the ass than the horse. and this probably explains the singular difference in the results arrived at by different observers. one variety certainly often has this prepotent power over another variety. and so it is with mongrel plants from a reciprocal cross. on the other hand. both when one species is crossed with another and when one variety is crossed with another variety. For instance. but the subject is here much complicated. the hybrids do not differ much. Gartner expressly states that the hybrids from long cultivated plants are more subject to reversion than hybrids from species in their natural state. but that the prepotency runs more strongly in the male than in the 304 . according to Gartner the same laws.

which is the offspring of the female-ass and stallion. be much more likely to occur with mongrels. and of crossed varieties. whether the two parents differ little or much from each other. On the whole. also. Independently of the question of fertility and sterility. If we look at species as having been specially created. which are descended from species slowly and naturally produced. but this does sometimes occur with hybrids. in all other respects there seems to be a general and close similarity in the offspring of crossed species. 305 . comes to the conclusion that the laws of resemblance of the child to its parents are the same.female ass. who. that it is only with mongrels that the offspring are not intermediate in character. which are descended from varieties often suddenly produced and semi-monstrous in character. after arranging an enormous body of facts with respect to animals. melanism. the resemblances seem chiefly confined to characters almost monstrous in their nature. Much stress has been laid by some authors on the supposed fact. I entirely agree with Dr. in the union of individuals of the same variety. this similarity would be an astonishing fact. and which have suddenly appeared—such as albinism. yet I grant much less frequently than with mongrels. and do not relate to characters which have been slowly acquired through selection. than with hybrids. deficiency of tail or horns. or of different varieties. or of distinct species. Looking to the cases which I have collected of cross-bred animals closely resembling one parent. so that the mule. namely. or additional fingers and toes. which is an offspring of the male ass and mare. but closely resemble one of their parents. But it harmonises perfectly with the view that there is no essential distinction between species and varieties. A tendency to sudden reversions to the perfect character of either parent would. and at varieties as having been produced by secondary laws. Prosper Lucas. is more like an ass than is the hinny.

and is eminently susceptible to action of favourable and unfavourable conditions. It is not always equal in degree in a first cross and in the hybrids produced from this cross. the capacity in one species or variety to take on another. and sometimes widely different in reciprocal crosses between the same two species. the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. and secondly. perhaps render it probable that some unknown bond in all cases connects the degree of fertility of first unions with that 306 . that. In the same manner as in grafting trees. so in crossing. He who will explain these latter cases will be able to explain the sterility of hybrids. in some instances in chief part on the early death of the embryo. or which have varied. 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. sterile. In the case of first crosses it seems to depend on several circumstances. it apparently depends on their whole organisation having been disturbed by being compounded from two distinct forms. the sterility being closely allied to that which so frequently affects pure species. firstly. but not universally. favours the size. and is often so slight that the most careful experimentalists have arrived at diametrically opposite conclusions in ranking forms by this test. which have been exposed to slightly different conditions of life. The sterility of first crosses and of their hybrid progeny has not been acquired through natural selection. slight changes in the conditions of life add to the vigour and fertility of all organic beings. The sterility is of all degrees. 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. generally of an unknown nature. The facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny. The degree of sterility does not strictly follow systematic affinity. sufficiently distinct to be ranked as species. and their hybrids. Summary Of Chapter First crosses between forms. that the crossing of forms. The sterility is innately variable in individuals of the same species. This view is strongly supported by a parallelism of another kind: namely. but is governed by several curious and complex laws. when exposed to new and unnatural conditions of life. are very generally. is incidental on differences. In the case of hybrids. in their vegetative systems.8. vigour and fertility of their offspring. It is generally different.

But why. then. yet the facts given in this chapter do not seem to me opposed to the belief that species aboriginally existed as varieties. in the case of distinct species. and the sterility of their hybrid offspring. The consideration of these facts on dimorphism. First crosses between forms known to be varieties. in all other respects there is the closest general resemblance between hybrids and mongrels. Nor is this almost universal and perfect fertility surprising. parallel with the systematic affinity of the forms subjected to experiment. but not. It is not surprising that the difficulty in crossing any two species. the sexual elements should so generally have become more or less modified. and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences. even if due to distinct causes: for both depend on the amount of difference between the species which are crossed. and is therefore little likely to induce this same quality. for systematic affinity includes resemblances of all kinds. and their mongrel offspring. Nor is it surprising that the facility of effecting a first cross.of their offspring. and in their inheritance of characters from both parent-forms. 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 the fertility of the hybrids thus produced. when it is remembered how liable we are to argue in a circle with respect to varieties in a state of nature. It should also be especially kept in mind. are very generally. leading to their mutual infertility. should in most cases correspond. Independently of the question of fertility. we do not know. invariably fertile. that longcontinued domestication tends to eliminate sterility. and that they have not been long exposed to uniform conditions of life. to a certain extent. as is so often stated. or sufficiently alike to be considered as varieties. 307 . in their variability. as well as of the results of reciprocal crosses. clearly leads to the conclusion that the primary cause of the sterility of crossed species is confined to differences in their sexual elements. in their power of absorbing each other by repeated crosses. 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. and the capacity of being grafted together—though this latter capacity evidently depends on widely different circumstances—should all run. Finally.

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 their number — On the lapse of time. 308 . 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.

from existing in lesser numbers than the forms which they connect. I endeavoured to show. on the theory. Most of them have now been discussed. when looking at any two species. namely. The explanation lies. is the most obvious and serious objection which can be urged against my theory. of innumerable intermediate links not now occurring everywhere throughout nature depends. will generally be beaten out and exterminated during the course of further modification and improvement. if we possessed all the intermediate varieties which have ever existed. To give a simple illustration: the fantail and pouter pigeons are both descended from the rock-pigeon. In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. none. which have formerly existed. that the life of each species depends in a more important manner on the presence of other already defined organic forms.1. perhaps. namely. One. be truly enormous. therefore. to show that intermediate varieties. also. we should always look for forms intermediate between each species and a common but unknown progenitor. than on climate. we should have an extremely close series between both and the rock-pigeon. on an extensive and continuous area with graduated physical conditions. is a very obvious difficulty. it should always be borne in mind what sort of intermediate forms must. and the progenitor will generally have differed in some respects from all its modified descendants. so must the number of intermediate varieties. and. I assigned reasons why such links do not commonly occur at the present day under the circumstances apparently most favourable for their presence. in the extreme imperfection of the geological record. The main cause. the distinctness of specific forms and their not being blended together by innumerable transitional links. But just in proportion as this process of extermination has acted on an enormous scale. as I believe. through which new varieties continually take the places of and supplant their parent-forms. however. on the very process of natural selection. In the first place. I endeavoured. But this is a wholly false view. and this. for 309 . that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. but we should have no varieties directly intermediate between the fantail and pouter. to avoid picturing to myself forms DIRECTLY intermediate between them. 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. have formerly existed. I have found it difficult.

that. and the principle of competition between organism and organism. oenas. have become so much modified.instance. but between each and an unknown common parent. unless at the same time we had a nearly perfect chain of the intermediate links. we have no reason to suppose that links directly intermediate between them ever existed. So that the number of intermediate and transitional links. such as C. whether they had descended from this species or from some other allied species. But assuredly. will render this a very rare event. in all such cases. combining a tail somewhat expanded with a crop somewhat enlarged. for in all cases the new and improved forms of life tend to supplant the old and unimproved forms. 310 . if we had no historical or indirect evidence regarding their origin. Hence. and these parent-species. it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon. if we look to forms very distinct. even perhaps more than they differ from each other. such have lived upon the earth. and so on backwards. livia. a horse from a tapir. moreover. So with natural species. C. for instance. must have been inconceivably great. between child and parent. have in their turn been similarly connected with more ancient forms. even if we closely compared the structure of the parent with that of its modified descendants. But such a case would imply that one form had remained for a very long period unaltered. by differences not greater than we see between the natural and domestic varieties of the same species at the present day. By the theory of natural selection all living species have been connected with the parent-species of each genus. always converging to the common ancestor of each great class. whilst its descendants had undergone a vast amount of change. if this theory be true. for instance to the horse and tapir. These two breeds. by the theory. we should be unable to recognise the parent-form of any two or more species. the characteristic features of these two breeds. now generally extinct. The common parent will have had in its whole organisation much general resemblance to the tapir and to the horse. and in this case DIRECT intermediate links will have existed between them. but in some points of structure may have differed considerably from both. that one of two living forms might have descended from the other. between all living and extinct species. It is just possible.

it may be objected that time cannot have sufficed for so great an amount of organic change. or even of each stratum. But how often do we see along the bases of retreating cliffs rounded boulders. until after being reduced in size they can be rolled about by the waves. It is good to wander along the coast. As Inferred From The Rate Of Deposition And Extent Of Denudation Independently of our not finding fossil remains of such infinitely numerous connecting links.2. and the waves eat into them only when they are charged with sand or pebbles. sand. all thickly clothed by marine productions. which the future historian will recognise as having produced a revolution in natural science. Not that it suffices to study the Principles of Geology. Therefore a man should examine for himself the great piles of superimposed strata. On The Lapse Of Time. have to be worn away atom by atom. As Lyell has well remarked. and mark the process of degradation. showing how little they are abraded and how seldom they are rolled about! Moreover. may at once close this volume. and yet does not admit how vast have been the past periods of time. which is undergoing degradation. for there is good evidence that pure water effects nothing in wearing away rock. We can best gain some idea of past time by knowing the agencies at work. He who can read Sir Charles Lyell's grand work on the Principles of Geology. and then they are more quickly ground into pebbles. and watch the rivulets bringing down mud. 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. along a short length or round a promontory. and these remaining fixed. or mud. The appearance of the surface and 311 . and to mark how each author attempts to give an inadequate idea of the duration of each formation. or to read special treatises by different observers on separate formations. in order to comprehend something about the duration of past time. and how much sediment has been deposited. the facts leading the mind feebly to comprehend the lapse of time. we find that it is only here and there. when formed of moderately hard rocks. the monuments of which we see all around us. that the cliffs are at the present time suffering. huge fragments fall down. At last the base of the cliff is undermined. The tides in most cases reach the cliffs only for a short time twice a day. It is hardly possible for me to recall to the reader who is not a practical geologist. all changes having been effected slowly. and learning how deeply the surface of the land has been denuded. if we follow for a few miles any line of rocky cliff. and the waves wearing away the sea-cliffs.

Messrs. have produced great results. the disintegrated matter is carried down even gentle slopes during heavy rain. The same story is told still more plainly by faults—those great cracks along which the strata have been upheaved on one side. and the observation is a most striking one. more forcibly than the conviction thus gained that subaerial agencies. for each line is composed of one and the same formation. in order to appreciate the past duration of time. it is then transported by the streams and rivers. or thrown down on the 312 . we see the effects of subaerial degradation in the muddy rills which flow down every slope. and in colder countries to frost. we are compelled to admit that the escarpments owe their origin in chief part to the rocks of which they are composed. on the one hand. which formerly were looked at as ancient seacoasts. which have been worn by the waves and pared all round into perpendicular cliffs of one or two thousand feet in height. and to a greater extent than might be supposed. due to their formerly liquid state. and which seem to work so slowly. it is good. for the gentle slope of the lavastreams. showed at a glance how far the hard. with the lines of harder rock left projecting. by the wind. with its dissolved carbonic acid. Croll and others. or the power of the waves. which. however. This being the case. having resisted subaerial denudation better than the surrounding surface. that the great lines of escarpment in the Wealden district and those ranging across England. this surface consequently has been gradually lowered. which apparently have so little power. rocky beds had once extended into the open ocean. When thus impressed with the slow rate at which the land is worn away through subaerial and littoral action. especially in arid districts. Geikie. while our sea-cliffs are everywhere formed by the intersection of various formations.the vegetation show that elsewhere years have elapsed since the waters washed their base. Ramsay and Whitaker have shown. and on the other hand the thickness of our sedimentary formations. The whole surface of the land is exposed to the chemical action of the air and of the rainwater. and triturate the fragments. On a rainy day. the masses of rock which have been removed over many extensive areas. that subaerial degradation is a much more important agency than coast-action. recently learned from the observations of Ramsay. according to our ideas of time. to consider. when rapid deepen their channels. Nothing impresses the mind with the vast duration of time. We have. cannot have been thus formed. I remember having been much struck when viewing volcanic islands. even in a gently undulating country. in the van of many excellent observers—of Jukes.

. Mr.154 Secondary strata… … … … … … … … … … . Croll shows. was slow and effected by many starts. the surface of the land has been so completely planed down that no trace of these vast dislocations is externally visible. between each successive formation we have. which are represented in England by thin beds. Mr.13. and then at the figures representing several million years. In regard to subaerial denudation. for instance. by calculating the 313 .other. On the other hand. extends for upward of thirty miles. from actual measurement in most cases. and the figures are at once pronounced too small. blank periods of enormous length. are thousands of feet in thickness on the Continent. in all parts of the world the piles of sedimentary strata are of wonderful thickness. to the height or depth of thousands of feet. and he informs me that he fully believes that there is one in Merionethshire of 12. they are good to show how slowly the mass must have been heaped together. In the Cordillera. in an interesting paper.000 feet. remarks that we do not err "in forming too great a conception of the length of geological periods.. Professor Ramsay has given me the maximum thickness. and this is the result: Feet Palaeozoic strata (not including igneous beds). Moreover. and along this line the vertical displacement of the strata varies from 600 to 3. The Craven fault. very nearly thirteen and threequarters British miles. When geologists look at large and complicated phenomena.57. Professor Ramsay has published an account of a downthrow in Anglesea of 2.584 feet. or. Croll.000 feet. in the opinion of most geologists. of the successive formations in DIFFERENT parts of Great Britain.2. as most geologists now believe." but in estimating them by years. and it makes no great difference whether the upheaval was sudden. for since the crust cracked.300 feet.240 —making altogether 72. 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. and although conglomerates have probably been accumulated at a quicker rate than finer sediments. I estimated one mass of conglomerate at ten thousand feet. Some of these formations.190 Tertiary strata… … … … … … … … … … … . Nevertheless this impression is partly false. yet from being formed of worn and rounded pebbles. that is. the two produce a totally different effect on the mind. the pile of rocks on either side of the crack having been smoothly swept away. yet in these cases there is nothing on the surface of the land to show such prodigious movements. So that the lofty pile of sedimentary rocks in Britain gives but an inadequate idea of the time which has elapsed during their accumulation. each of which bears the stamp of time.

during a single lifetime. by which some of the inhabitants might be better fitted to their new places under the altered circumstance. so that a hundred years represents the work of two breeders in succession. various breeds have been sensibly changed in the course of two or three centuries. Species. how long a period it takes to modify a species. Few of us. but by this process of unconscious selection. would thus be removed from the mean level of the whole area in the course of six million years. or through the immigration of new forms. Few men have attended with due care to any one strain for more than half a century. that is.known amount of sediment annually brought down by certain rivers. Several eminent breeders. represented as it is by a measure utterly insignificant in a hall of the above dimensions. have so largely modified some of the higher animals. then mark off at one end the tenth of an inch. variations or individual differences of the right nature. would not always occur at once. which propagate their kind much more slowly than most of the lower animals. due to the occurrence of physical changes of some kind. but to the subject of time we must return. Moreover. 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. This seems an astonishing result. but if halved or quartered it is still very surprising. that new places in the polity of nature do not occur until after long intervals. with no intention of modifying the breed.000 feet of solid rock. in relation to the subject of this work. what a hundred years implies. relatively to their areas of drainage. 314 . and within the same country only a few change at the same time. The comparison would be in every way fairer with the effects which follow from unconscious selection. probably change much more slowly. Croll gives the following illustration: Take a narrow strip of paper. Unfortunately we have no means of determining. eighty-three feet four inches in length. the preservation of the most useful or beautiful animals. that they have formed what well deserves to be called a new sub-breed. and stretch it along the wall of a large hall. as it became gradually disintegrated. and the entire strip a million years. that 1. know what a million really means: Mr. 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. This tenth of an inch will represent one hundred years. according to the standard of years. however. however. But let it be borne in mind.

and although this great mass has been most carefully searched. if in sand or gravel. at a rate sufficiently quick to embed and preserve fossil remains. seem explicable only on the view of the bottom of the sea not rarely lying for ages in an unaltered condition. or from a few specimens collected on some one spot. that very many fossil species are known and named from single and often broken specimens. except a few vegetable remains. without the underlying bed having suffered in the interval any wear and tear. the bright blue tint of the water bespeaks its purity. Only a small portion of the surface of the earth has been geologically explored. when the beds are upraised. many great deposits. Throughout an enormously large proportion of the ocean. We probably take a quite erroneous view. should never be forgotten. 315 . and no part with sufficient care. where sediment is not accumulating. are entirely destitute of organic remains. will. and extending for at least 300 miles from Vienna to Switzerland.3. requiring a vast length of time for their accumulation. occasionally even six thousand feet in thickness. by another and later formation. generally be dissolved by the percolation of rain water charged with carbonic acid. 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. which consists of shale and sandstone. The remains which do become embedded. Edward Forbes. without our being able to assign any reason: one of the most striking instances is that of the Flysch formation. No organism wholly soft can be preserved. The many cases on record of a formation conformably covered. For instance. have been found. On The Poorness Of Palaeontological Collections Now let us turn to our richest museums. several thousand. Some of the many kinds of animals which live on the beach between high and low water mark seem to be rarely preserved. which inhabits deep water and this has been found fossil in Sicily. after an immense interval of time. namely. with the exception of a single Mediterranean species. 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. Shells and bones decay and disappear when left on the bottom of the sea. when we assume that sediment is being deposited over nearly the whole bed of the sea. and what a paltry display we behold! That our collections are imperfect is admitted by every one. Lastly. no fossils. The remark of that admirable palaeontologist. as the important discoveries made every year in Europe prove.

so it is in North America. we may infer that this could nowhere be ascertained. But we know. whence the sediment was derived. Nor is their rarity surprising. for instance. entirely disbelieve in the change of species. from the several formations being separated from each other by wide intervals of time. But the imperfection in the geological record largely results from another and more important cause than any of the foregoing. namely. We can. that is. accord with the belief of vast intervals of time having elapsed between each formation. Dawson in the carboniferous strata of North America. The most skilful geologist. charged with new and peculiar forms of life. in every separate territory. it is superfluous to state that our evidence is fragmentary in an extreme degree. had elsewhere been accumulated. When we see the formations tabulated in written works. from Sir R. with the exception of one species discovered by Sir C. would never have suspected that during the periods which were blank and barren in his own country. And if. Murchison's great work on Russia. Scarcely any fact struck me more when examining many hundred miles of the South American coasts. great piles of sediment. and in many other parts of the world. hardly any idea can be formed of the length of time which has elapsed between the consecutive formations. This doctrine has been emphatically admitted by many geologists and palaeontologists. far better than pages of detail. For instance.With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods. how accidental and rare is their preservation. which have been upraised 316 . or when we follow them in nature. I think. until recently not a land-shell was known belonging to either of these vast periods. when we remember how large a proportion of the bones of tertiary mammals have been discovered either in caves or in lacustrine deposits. like E. and that not a cave or true lacustrine bed is known belonging to the age of our secondary or palaeozoic formations. what wide gaps there are in that country between the superimposed formations. Forbes. see why the geological formations of each region are almost invariably intermittent. but now land-shells have been found in the lias. who. will bring home the truth. a glance at the historical table published in Lyell's Manual. generally implying great changes in the geography of the surrounding lands. it is difficult to avoid believing that they are closely consecutive. In regard to mammiferous remains. Lyell and Dr. have not followed each other in close sequence. The frequent and great changes in the mineralogical composition of consecutive formations. if his attention had been confined exclusively to these large territories.

conclude that sediment must be accumulated in extremely thick. no extensive formations with recent or tertiary remains can anywhere be found. have thus been formed during subsidence. when upraised. and have been surprised to note how author after author. if it continue slowly to subside.several hundred feet within the recent period. no doubt. has come to the conclusion that it was accumulated during subsidence. to resist a large amount of denudation. which are throughout the greater part of their thickness RICH IN FOSSILS. that the only ancient tertiary formation on the west coast of South America. along the rising coast of the western side of South America. from the enormous degradation of the coast rocks and from the muddy streams entering the sea. as well as the subsequent subaerial degradation. or extensive masses. tertiary beds are so poorly developed that no record of several successive and peculiar marine faunas will probably be preserved to a distant age. when first upraised and during subsequent oscillations of level. though the supply of sediment must for ages have been great. I have watched the progress of geology. Along the whole west coast. Such thick and extensive accumulations of sediment may be formed in two ways. either in profound depths of the sea. I am convinced that nearly all our ancient formations. which is inhabited by a peculiar marine fauna. In this latter case. in order to withstand the incessant action of the waves. and thus a rich fossiliferous formation. I may add. as long as the rate of subsidence and supply of sediment nearly balance each other. 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. thick enough. than the absence of any recent deposits sufficiently extensive to last for even a short geological period. A little reflection will explain why. Or sediment may be deposited to any thickness and extent over a shallow bottom. and the mass when upraised will give an imperfect record of the organisms which existed in the neighbourhood during the period of its accumulation. is that the littoral and sub-littoral deposits are continually worn away. The explanation. as soon as they are brought up by the slow and gradual rising of the land within the grinding action of the coast-waves. may be formed. I think. but which will hardly last to a 317 . We may. in treating of this or that great formation. solid. the sea will remain shallow and favourable for many and varied forms. which has been bulky enough to resist such degradation as it has as yet suffered.

however. nor be capped by overlying formations.distant geological age.. the former protecting mantle of rock may not have been very thick. Admitting then that gneiss. were once 318 . or. Still less can this have happened during the alternate periods of elevation. the deposit formed during the rising movement. granite. the beds which were then accumulated will generally have been destroyed by being upraised and brought within the limits of the coast-action. that if one part of the area. might afterwards become protected by fresh accumulations. Hopkins. as long as the bed of the sea remained stationary. a widely extended formation might be formed during a period of elevation. In the case of an extensive and shallow sea. and thus be preserved for a long period. which are the most favourable to life. such as that within a large part of the Malay Archipelago. It has. These remarks apply chiefly to littoral and sublittoral deposits. after rising and before being denuded. nor would the deposit be much consolidated. etc. But all geologists. Consequently. formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation. mica-schist. where the depth varies from thirty or forty to sixty fathoms. For it is scarcely possible that such rocks could have been solidified and crystallised while uncovered. Mr. was deposited during a downward oscillation of level. subsided. excepting the few who believe that our present metamorphic schists and plutonic rocks once formed the primordial nucleus of the globe. will have been formed over wide spaces during periods of subsidence. though not thick. but the thickness of the formation could not be great. 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. diorite. been suggested by Mr. THICK deposits cannot have been accumulated in the shallow parts. and thus gained considerable thickness. All geological facts tell us plainly that each area has undergone numerous slow oscillations of level. and apparently these oscillations have affected wide spaces. and yet not suffer excessively from denudation during its slow upheaval. Hopkins also expresses his belief that sedimentary beds of considerable horizontal extent have rarely been completely destroyed. but if the metamorphic action occurred at profound depths of the ocean. for owing to the elevatory movement it would be less than the depth in which it was formed. On the other hand. will admit that these latter rocks have been stripped of their covering to an enormous extent. to speak more accurately. 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.

Inland. if all the sedimentary beds were removed which rest unconformably on them. Numerous specimens. I have estimated the areas by cutting out and weighing the paper. 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. during subsidence. along the whole northern bank of the Plata. and saw nothing but granitic rocks. the whole of the newer Palaeozoic formations. with not a wreck left behind. how can we account for the naked and extensive areas of such rocks in many parts of the world.necessarily covered up. the granitic area is very large: thus Von Eschwege gives a detailed section of these rocks. Italy. all conjoined. Rogers' beautiful map. Hence. which alone could have formed a part of the original capping of the granitic series. for the formation of new varieties and species. One remark is here worth a passing notice. 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.5. collected along the whole coast. I saw. part of Germany. stretching from Rio de Janeiro for 260 geographical miles inland in a straight line. Boue colours an area composed of rocks of this nature as equal to that of Spain. in the proportion of 19 to 12. and consequently during subsidence. On the other hand. and it is during these very 319 . as shown in Professor H. and they all belonged to this class. as previously explained. namely. it is probable that in some parts of the world whole formations have been completely denuded. but from the concurrent testimony of travellers. Turning to a wellknown region. few new varieties or species will be formed.100 geographical miles.D. besides modern tertiary beds. but during such periods there will generally be a blank in the geological record. were examined by me. and the British Islands. from near Rio de Janeiro to the mouth of the Plata. and which could not have formed part of the original mantle under which they were crystallised. though there will be much extinction. France. and I find that the metamorphic (excluding the "semi-metamorphic") and granite rocks exceed. South of the Amazon. This region has not been carefully explored. the inhabited area and number of inhabitants will decrease (excepting on the shores of a continent when first broken up into an archipelago). and I travelled for 150 miles in another direction. only one small patch of slightly metamorphosed rock. a distance of 1. In many regions the metamorphic and granite rocks would be found much more widely extended than they appear to be. to the United States and Canada.

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

when we find a species disappearing before the last layers have been deposited. the probability is that it only then first immigrated into that area. Several cases are on record of the same species presenting varieties in the upper and lower parts of the same formation. as it seems to me. time having apparently been required for their migration from the American to the European 321 . Although each formation has indisputably required a vast number of years for its deposition. prevent us from coming to any just conclusion on this head. But insuperable difficulties. nor have the several stages of the same formation throughout Europe been correlated with perfect accuracy. viewed as a whole. We forget how small the area of Europe is compared with the rest of the world. I am aware that two palaeontologists. is extremely imperfect. Although each formation may mark a very long lapse of years. that several species appear somewhat earlier in the palaeozoic beds of North America than in those of Europe. have concluded that the average duration of each formation is twice or thrice as long as the average duration of specific forms.4. but if we confine our attention to any one formation. for instance. 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. 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. 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. On The Absence Of Numerous Intermediate Varieties In Any Single Formation From these several considerations it cannot be doubted that the geological record. each probably is short compared with the period requisite to change one species into another. it would be equally rash to suppose that it then became extinct. and when we see a species first appearing in any 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. When we see a species first appearing in the middle of any formation. Thus Trautschold gives a number of instances with Ammonites. but I cannot assign due proportional weight to the following considerations. It is well known. So again. it would be rash in the extreme to infer that it had not elsewhere previously existed. whose opinions are worthy of much deference. namely Bronn and Woodward.

for instance.seas. whilst the downward movement continues. the deposit must have gone on continuously accumulating during a long period. hence. But we have seen that a thick formation. Yet it may be doubted whether. But this same movement of subsidence will tend to submerge the area whence the sediment is derived. instead of having been really far greater. And in the distant future. examining these beds. INCLUDING FOSSIL REMAINS. for it has been observed by more 322 . 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. sufficient for the slow process of modification. owing to the migrations of species and to geographical changes. that is. the deposit must be a very thick one. In fact. organic remains will probably first appear and disappear at different levels. and likewise to reflect on the changes of level. which forms only a part of one whole geological period. on the extreme change of climate. probable that sediment was deposited during the whole of the glacial period near the mouth of the Mississippi. this nearly exact balancing between the supply of sediment and the amount of subsidence is probably a rare contingency. it has everywhere been noted. in any quarter of the world. would be tempted to conclude that the average duration of life of the embedded fossils had been less than that of the glacial period. and on the great lapse of time. fossiliferous throughout its entire thickness. and the species undergoing change must have lived in the same district throughout the whole time. It is an excellent lesson to reflect on the ascertained amount of migration of the inhabitants of Europe during the glacial epoch. In order to get a perfect gradation between two forms in the upper and lower parts of the same formation. conversely. sedimentary deposits. that some few still existing species are common in the deposit. a geologist. that some are now abundant in the neighbouring sea. have gone on accumulating within the same area during the whole of this period. It is not. and to keep the depth approximately the same. or. can accumulate only during a period of subsidence. 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. extending from before the glacial epoch to the present day. and thus diminish the supply. but have become extinct in the immediately surrounding sea. In examining the latest deposits. which is necessary that the same marine species may live on the same space. but are rare or absent in this particular deposit. in various quarters of the world. the supply of sediment must nearly counterbalance the amount of subsidence. all included within this same glacial period.

though perhaps slight. a formation composed of beds of widely different mineralogical composition. still standing upright as they grew. Many instances could be given of beds. even if 323 . at no less than sixty-eight different levels. Supposing B and C to be two species. we may reasonably suspect that the process of deposition has been more or less interrupted. Consequently if it were to undergo a considerable amount of modification during the deposition of any one geological formation.than one palaeontologist that very thick deposits are usually barren of organic remains. they grant some little variability to each species. which would not have been suspected. Lyell and Dr. Hence. yet no one ignorant of this fact would have even suspected the vast lapse of time represented by the thinner formation. they rank both as species. A. when the same species occurs at the bottom. and this. to be found in an older and underlying bed. and which must have required an enormous period for their accumulation. as is so often the case. In other cases we have the plainest evidence in great fossilised trees. like the whole pile of formations in any country. but when they meet with a somewhat greater amount of difference between any two forms. middle. one above the other. When we see. the probability is that it has not lived on the same spot during the whole period of deposition. but abrupt. yet easily overlooked. and then re-covered by the upper beds of the same formation—facts. had not the trees been preserved: thus Sir C. with ancient root-bearing strata. and a third. Dawson found carboniferous beds 1.400 feet thick in Nova Scotia. It would seem that each separate formation. denuded. submerged. perhaps many times. has generally been intermittent in its accumulation. from the reasons just assigned. It is all-important to remember that naturalists have no golden rule by which to distinguish species and varieties. except near their upper or lower limits. only a few feet in thickness. showing what wide. intervals have occurred in its accumulation. and top of a formation. we can seldom hope to effect in any one geological section. unless they are enabled to connect them together by the closest intermediate gradations. changes of form. Nor will the closest inspection of a formation give us any idea of the length of time which its deposition may have consumed. during the same geological period. of many long intervals of time and changes of level during the process of deposition. representing formations which are elsewhere thousands of feet in thickness. but has disappeared and reappeared. a section would not include all the fine intermediate gradations which must on our theory have existed. Many cases could be given of the lower beds of a formation having been upraised.

that their varieties are generally at first local. that A might be the actual progenitor of B and C. maintain that all these tertiary species are specifically distinct. we have evidence of the frequent occurrence of slight modifications of the kind required. So that we might obtain the parentspecies and its several modified descendants from the lower and upper beds of the same formation. It is notorious on what excessively slight differences many palaeontologists have founded their species. to distinct but consecutive stages of the same great formation. and yet would not necessarily be strictly intermediate between them in all respects. and on this view we do find the kind of evidence of change which on the theory we ought to find. unless we believe that these eminent naturalists have been misled by their imaginations. it would simply be ranked as a third and distinct species. but some excellent naturalists. If we look to rather wider intervals of time. and that these late tertiary species really present no difference whatever from their living representatives. so that here again we have undoubted evidence of change in the direction required by the theory. we find that the embedded fossils. and should consequently rank them as distinct species. which include many shells believed by the majority of naturalists to be identical with existing species. Nor should it be forgotten. With animals and plants that propagate rapidly and do not wander much. we should not recognise their blood-relationship. unless at the same time it could be closely connected by intermediate varieties with either one or both forms. Look again at the later tertiary deposits. as Agassiz and Pictet. Some experienced conchologists are now sinking many of the very fine species of D'Orbigny and others into the rank of varieties. as before explained. though universally ranked as specifically different. though the distinction is admitted to be very slight. namely. as we have formerly seen. yet are far more closely related to each other than are the species found in more widely separated formations. there is reason to suspect. so that here. that these tertiary species are all truly distinct from the recent. in opposition to the judgment of most naturalists. but to this latter subject I shall return in the following chapter. 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. the chance of discovering in a formation in any one country all the early 324 . and unless we obtained numerous transitional gradations. or unless we admit.A were strictly intermediate between B and C. According to this view. and they do this the more readily if the specimens come from different sub-stages of the same formation.

and dogs are descended from a single stock or from several aboriginal stocks. by asking ourselves whether. perhaps. and thus proved to be the same species. that geology yields no linking forms. that the period during which each species underwent modification. are really varieties. that at the present day. recent and extinct. This could be effected by the future geologist only by his discovering in a fossil state numerous intermediate gradations. namely. fine. that oftenest present varieties. have oftenest given rise. for the successive changes are supposed to have been local or confined to some one spot." If we take a genus having a score of species. and by other conchologists as only varieties.stages of transition between any two forms. as it is called. no one doubts that the remainder will stand much more distinct from each other. with shells and other marine animals. It should not be forgotten. for instance. first to local varieties and ultimately to new species. As Sir J. and with fossil species this can rarely be done. Lubbock has remarked. until many specimens are collected from many places. and destroy four-fifths of them. it is probable that those which had the widest range. intermediate. "Every species is a link between other allied forms. and we have seen that with plants it is those which have the widest range. with perfect specimens for examination. Falconer. and this again would greatly lessen the chance of our being able to trace the stages of transition in any one geological formation. best perceive the improbability of our being enabled to connect species by numerous. far exceeding the limits of the known geological formations in Europe. was probably short in comparison with that during which it remained without undergoing any change. It is a more important consideration. or are. We shall. as we shall see in the next chapter. the genus itself will stand more distinct from other allied genera. If the extreme forms in the genus happen to have been thus destroyed. two forms can seldom be connected by intermediate varieties. Most marine animals have a wide range. or. is small. leading to the same result. This assertion. What geological research has not 325 . by writers who believe in the immutability of species. specifically distinct. again. geologists at some future period will be able to prove that our different breeds of cattle. horses. sheep. as lately insisted on by Dr. and such success is improbable in the highest degree. It has been asserted over and over again. is certainly erroneous. fossil links. which are ranked by some conchologists as distinct species from their European representatives. though long as measured by years. whether certain sea-shells inhabiting the shores of North America. so that.

revealed. excepting those of the United States of America. But this ought not to be expected. as we now see on the shores of South America. the fossiliferous formations on the steeper shores would be destroyed. there would probably 326 . Formations rich in fossils of many kinds. sedimentary beds could hardly be accumulated of great thickness during the periods of elevation. or of those which lived on naked submarine rocks. Not many of the strictly littoral animals. 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. during which the area would be either stationary or rising. Even throughout the extensive and shallow seas within the archipelago. The Malay Archipelago is about the size of Europe from the North Cape to the Mediterranean. These periods of subsidence would be separated from each other by immense intervals of time. Godwin. During the periods of subsidence. is the former existence of infinitely numerous gradations. that the present condition of the Malay Archipelago.Austen. so as to have a good chance of enduring to a very distant future. with its numerous large islands separated by wide and shallow seas. no remains could be preserved. and therefore equals all the geological formations which have been examined with any accuracy. or where it did not accumulate at a sufficient rate to protect organic bodies from decay. I fully agree with Mr. Wherever sediment did not accumulate on the bed of the sea. would generally be formed in the archipelago only during periods of subsidence. 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 of thickness sufficient to last to an age as distant in futurity as the secondary formations lie in the past. The Malay Archipelago is one of the richest regions in organic beings. yet if all the species were to be collected which have ever lived there. probably represents the former state of Europe. while most of our formations were accumulating. and from Britain to Russia. by the incessant coast-action. whilst rising. yet this has been repeatedly advanced as a most serious objection against my views. almost as soon as accumulated. and those embedded in gravel or sand would not endure to a distant epoch. as fine as existing varieties. or become capped and protected by subsequent deposits. connecting together nearly all existing and extinct species. would be embedded.

be much extinction of life. be ranked as distinct species. pressed so hardly on my theory. 327 . and such assuredly we do find—some more distantly. and these links. have connected all the past and present species of the same group into one long and branching chain of life. an infinite number of those fine transitional forms. though perhaps extremely slight degree. they would slowly spread and supplant their parent-forms. together with a contemporaneous accumulation of sediment. related to each other. and that slight climatical changes would intervene during such lengthy periods. If then there be some degree of truth in these remarks. by many palaeontologists. on our theory. we have no right to expect to find. Very many of the marine inhabitants of the archipelago now range thousands of miles beyond its confines. though closely allied species. and analogy plainly leads to the belief that it would be chiefly these far-ranging species. and the varieties would at first be local or confined to one place. some more closely. as they would differ from their former state in a nearly uniform. and no closely consecutive record of their modifications could be preserved in any one formation. But I do not pretend that I should ever have suspected how poor was the record in the best preserved geological sections. if found in different stages of the same formation. When such varieties returned to their ancient homes. which. and these contingencies are indispensable for the preservation of all the transitional gradations between any two or more species. though only some of them. but if possessed of any decided advantage. let them be ever so close. or when further modified and improved. and in these cases the inhabitants of the archipelago would migrate. there would be much variation. It is also probable that each great period of subsidence would be interrupted by oscillations of level. If such gradations were not all fully preserved. We ought only to look for a few links. they would. but the geological record would then be less perfect. would. which would oftenest produce new varieties. in our geological formations. transitional varieties would merely appear as so many new. during the periods of elevation. according to the principles followed by many palaeontologists. would EXCEED the average duration of the same specific forms. be ranked as new and distinct species. It may be doubted whether the duration of any one great period of subsidence over the whole or part of the archipelago. had not the absence of innumerable transitional links between the species which lived at the commencement and close of each formation. and as they would be found embedded in slightly different substages of the same formation.

before they invaded the ancient archipelagoes of Europe and the United States. and consequently that the transitional forms would often long remain confined to some one region. that they did not exist before that stage. and the progenitors must have lived long before their modified descendants. I may here recall a remark formerly made. when this adaptation had once been effected. We do not make due allowance for the enormous intervals of time which have elapsed between our consecutive formations. Professor Pictet. cannot see how the successive modifications of the anterior limbs of a supposed prototype could possibly have been of any advantage. and falsely infer. in commenting on early transitional forms. On The Sudden Appearance Of Whole Groups Of Allied Species The abrupt manner in which whole groups of species suddenly appear in certain formations. longer perhaps in many cases than the time required for the accumulation of each formation. we forget that groups of species may elsewhere have long existed.5. the fact would be fatal to the theory of evolution through natural selection. to fly through the air. by Agassiz. Pictet. but that. But look at the 328 . These intervals will have given time for the multiplication of species from some one parent-form: and in the succeeding formation. and Sedgwick. in his excellent Review of this work. which would spread rapidly and widely throughout the world. as a fatal objection to the belief in the transmutation of species. have really started into life at once. for instance. as experience has so often shown. and have slowly multiplied. negative evidence is worthless. because certain genera or families have not been found beneath a certain stage. that it might require a long succession of ages to adapt an organism to some new and peculiar line of life. has been urged by several palaeontologists—for instance. namely. and a few species had thus acquired a great advantage over other organisms. compared with the area over which our geological formations have been carefully examined. For the development by this means of a group of forms. and taking birds as an illustration. a comparatively short time would be necessary to produce many divergent forms. all of which are descended from some one progenitor. We continually forget how large the world is. must have been an extremely slow process. If numerous species. But we continually overrate the perfection of the geological record. such groups or species will appear as if suddenly created. belonging to the same genera or families. In all cases positive palaeontological evidence may be implicitly trusted.

and with its wings furnished with two free claws. existed during that period? Not a fragment of bone has been discovered in these beds. and ultimately to rise from its surface and glide through the air? I will now give a few examples to illustrate the foregoing remarks. Cuvier used to urge that no monkey occurred in any tertiary stratum. published in 1844-46 and in 1853-57. Not long ago. that a bird certainly lived during the deposition of the upper greensand. I may recall the well-known fact that in geological treatises. bearing a pair of feathers on each joint. that strange bird. and to show how liable we are to error in supposing that whole groups of species have suddenly been produced. Had it not been for the rare accident of the preservation of footsteps in the new red sandstone of the United States. for they exist in infinite numbers and of many kinds. And now one of the richest known accumulations of fossil mammals belongs to the middle of the secondary series. the conclusions on the first appearance and disappearance of several groups of animals have been considerably modified. the Archeopteryx. South America and in Europe. In a memoir on Fossil Sessile Cirripedes. from having passed under my own eyes has much struck me. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world. and a third edition would require still further changes. with a long lizard-like tail.penguins of the Southern Ocean. I may give another instance. Even in so short an interval as that between the first and second editions of Pictet's great work on Palaeontology. but what special difficulty is there in believing that it might profit the modified descendants of the penguin. I 329 . first to become enabled to flap along the surface of the sea like the logger-headed duck. who would have ventured to suppose that no less than at least thirty different bird-like animals. published not many years ago. some of gigantic size. 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. which. has been discovered in the oolitic slates of Solenhofen. and still more recently. on the authority of Professor Owen. mammals were always spoken of as having abruptly come in at the commencement of the tertiary series. but now extinct species have been discovered in India. I do not suppose that we here see the real transitional grades through which the wings of birds have passed. and true mammals have been discovered in the new red sandstone at nearly the commencement of this great series. as far back as the miocene stage. but now we know. palaeontologists maintained that the whole class of birds came suddenly into existence during the eocene period.

from all these circumstances. But certain Jurassic and Triassic forms are now commonly admitted to be teleostean. 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. It is almost superfluous to remark that hardly any fossil-fish are known from south of the equator. inhabiting various zones of depths. and as not one species had then been discovered in beds of this age. Some few families of fish now have a confined range. low down. they would certainly have been preserved and discovered. I inferred that. from the large number of existing and extinct tertiary species. this cirripede was a Chthamalus. is that of the teleostean fishes. sent me a drawing of a perfect specimen of an unmistakable sessile cirripede. and ubiquitous genus. and after having been largely developed in 330 . a member of a distinct subfamily of sessile cirripedes. I concluded that this great group had been suddenly developed at the commencement of the tertiary series. M. Still more recently. 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. the fact would have been highly remarkable. as if to make the case as striking as possible. If the teleosteans had really appeared suddenly in the northern hemisphere at the commencement of the chalk formation. large. of which not one species has as yet been found even in any tertiary stratum. according to Agassiz. Woodward in the upper chalk. from the Arctic regions to the equator. has been discovered by Mr. a Pyrgoma. in the Chalk period. as I then thought. from the perfect manner in which specimens are preserved in the oldest tertiary beds. But my work had hardly been published. had sessile cirripedes existed during the secondary periods. from the ease with which even a fragment of a valve can be recognised. a very common. from the extraordinary abundance of the individuals of many species all over the world. and even some palaeozoic forms have thus been classed by one high authority. one more instance of the abrupt appearance of a great group of species. Bosquet. but it would not have formed an insuperable difficulty. This group includes the large majority of existing species. 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. This was a sore trouble to me. the teleostean fishes might formerly have had a similarly confined range.stated that. which he had himself extracted from the chalk of Belgium. And. from the upper tidal limits to fifty fathoms. adding.

and then to discuss the number and range of its productions. Even at this day. 331 . have spread widely. as it would be for a naturalist to land for five minutes on a barren point in Australia. and were enabled to double the southern capes of Africa or Australia.some one sea. 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. it seems to me to be about as rash to dogmatize on the succession of organic forms throughout the world. in which any great group of marine animals might be multiplied. and from the revolution in our palaeontological knowledge effected by the discoveries of the last dozen years. and here they would remain confined. from our ignorance of the geology of other countries beyond the confines of Europe and the United States. until some of the species became adapted to a cooler climate. and thus reach other and distant seas. From these considerations. the tropical parts of the Indian Ocean would form a large and perfectly enclosed basin. if the Malay Archipelago were converted into land.

Lingula. which have certainly occurred since the Cambrian formation. and it cannot on our theory be supposed. These very wide limits show how doubtful the data are. Mr. the whole interval from the Cambrian age to the present day. and other elements may have hereafter to be introduced into the problem.. Croll estimates that about sixty million years have elapsed since the Cambrian period. however. but probably not less than ninety-eight or more than two hundred million years. and which probably differed greatly from any known animal. but this. do not differ much from living species. as long as. as Sir William Thompson insists. etc. if the theory be true. it cannot be doubted that all the Cambrian and Silurian trilobites are descended from some one crustacean. as the Nautilus. Most of the arguments which have convinced me that all the existing species of the same group are descended from a single progenitor. and that during these vast periods the world swarmed with living creatures. Here we encounter a formidable objection. Some of the most ancient animals. that the world at a very early period was subjected to more rapid and violent changes in its physical conditions than those now occurring. and such 332 . that these old species were the progenitors of all the species belonging to the same groups which have subsequently appeared. for it seems doubtful whether the earth. On The Sudden Appearance Of Groups Of Allied Species In The Lowest Known Fossiliferous Strata There is another and allied difficulty. which must have lived long before the Cambrian age. has lasted long enough. It is. Consequently. 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. Sir W.6. apply with equal force to the earliest known species. judging from the small amount of organic change since the commencement of the Glacial epoch. which is much more serious. For instance. 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. for they are not in any degree intermediate in character. probable. appears a very short time for the many and great mutations of life. Thompson concludes that the consolidation of the crust can hardly have occurred less than twenty or more than four hundred million years ago. it is indisputable that before the lowest Cambrian stratum was deposited long periods elapsed. in a fit state for the habitation of living creatures. or probably far longer than.

Logan states that their "united thickness may possibly far surpass that of all the succeeding rocks. Logan. We are thus carried back to a period so remote. To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system. Other highly competent judges. were until recently convinced that we beheld in the organic remains of the lowest Silurian stratum the first dawn of life. it existed in countless numbers. Thus the words. and the existence of the Eozoon in the Laurentian formation of Canada is generally admitted. Forbes. and which are almost the same with those since used by Sir W. But the descriptions which we possess of the Silurian deposits over immense territories in Russia and in North 333 . and now." The Eozoon belongs to the most lowly organised of all classes of animals. Barrande added another and lower stage. abounding with new and peculiar species.changes would have tended to induce changes at a corresponding rate in the organisms which then existed. Mr Hicks has found South Wales beds rich in trilobites. Nevertheless. Sir W. There are three great series of strata beneath the Silurian system in Canada. probably indicates life at these periods. and containing various molluscs and annelids. Not very long ago M. with Sir R. Murchison at their head. 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. that the appearance of the so-called primordial fauna (of Barrande) may by some be considered as a comparatively modern event. beneath the then known Silurian system. as Dr. which I wrote in 1859. from the base of the palaeozoic series to the present time. have disputed this conclusion. I can give no satisfactory answer. It does not seem probable that the most ancient beds have been quite worn away by denudation. in the lowest of which the Eozoon is found. still lower down in the Lower Cambrian formation. or that their fossils have been wholly obliterated by metamorphic action. for if this had been the case we should have found only small remnants of the formations next succeeding them in age. even in some of the lowest azotic rocks. about the existence of living beings long before the Cambrian period. Several eminent geologists. and. have proved true. certainly preyed on other minute organic beings. as Lyell and E. We should not forget that only a small portion of the world is known with accuracy. Dawson has remarked. The presence of phosphatic nodules and bituminous matter. but is highly organised for its class. which must have lived in great numbers. and these would always have existed in a partially metamorphosed condition.

that during the palaeozoic and secondary periods. whence the sediment was derived. oceans have extended from the remotest period of which we have any record. led me to conclude that the great oceans are still mainly areas of subsidence. Looking to the existing oceans. and the continents areas of elevation. and may be truly urged as a valid argument against the views here entertained. which must have intervened during these enormously long periods. But we do not know what was the state of things in the intervals between the several successive formations. To show that it may hereafter receive some explanation. that where continents now exist. This same view has since been maintained by Agassiz and others. we see them studded with many islands. palaeozoic and secondary formations would in all probability have been accumulated from sediment derived from their wear and tear. of which the formations are composed. since the Cambrian period. large tracts of land have existed. But we have no reason to assume that things have thus remained from the beginning of the world. miles in thickness. From the nature of the organic remains which do not appear to have inhabited profound depths. then. and on the other hand. where our oceans now extend. whether Europe and the United States during these intervals existed as dry land. Our continents seem to have been formed by a preponderance. but hardly one truly oceanic island (with the exception of New Zealand. and would have been at least partially upheaved by the oscillations of level. But may not the areas of preponderant movement have changed in 334 . and from the amount of sediment. on which sediment was not deposited. If. Hence. we may perhaps infer. I will give the following hypothesis. in the several formations of Europe and of the United States.America. which are thrice as extensive as the land. no doubt. we may infer anything from these facts. The coloured map appended to my volume on Coral Reefs. of the force of elevation. we may infer that. we may infer that from first to last large islands or tracts of land. The case at present must remain inexplicable. occurred in the neighbourhood of the now existing continents of Europe and North America. subjected. or as the bed of an open and unfathomable sea. during many oscillations of level. if this can be called a truly oceanic island) is as yet known to afford even a remnant of any palaeozoic or secondary formation. or as a submarine surface near land. neither continents nor continental islands existed where our oceans now extend. for had they existed. do not support the view that the older a formation is the more invariably it has suffered extreme denudation and metamorphism. the great archipelagoes still areas of oscillations of level. to great oscillations of level.

and which had been pressed on by an enormous weight of superincumbent water. in recognisable condition. though we find in our geological formations many links between the species which now exist and which formerly existed. may represent the forms of life. The immense areas in some parts of the world. the almost entire absence. The sudden manner in which several groups of species first appear in our European formations. Agassiz. for instance. Pictet. and clear and open oceans may have existed where our continents now stand. Of this history we possess the last volume alone. and all our greatest geologists. supposing such to have been formerly deposited. for it might well happen that strata which had subsided some miles nearer to the centre of the earth. Sedgwick. for instance in South America. we do not find infinitely numerous fine transitional forms closely joining them all together. The several difficulties here discussed.. relating only to two or three countries. namely. Falconer. and of each page. which must have been heated under great pressure. of naked metamorphic rocks. the bed of the Pacific Ocean were now converted into a continent we should there find sedimentary formations. which are entombed in our consecutive formations. Nor should we be justified in assuming that if. Those who believe that the geological record is in any degree perfect. following out Lyell's metaphor. maintained the immutability of species. as at present known. Murchison. namely. Cuvier. of formations rich in fossils beneath the Cambrian strata. as Lyell. E. often vehemently. and which falsely appear to have been 335 .the lapse of ages? At a period long antecedent to the Cambrian epoch. 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. that. and most geologists and palaeontologists are much shaken in their former belief. older than the Cambrian strata.. more or less different in the successive chapters. have always seemed to me to require some special explanation. Each word of the slowly-changing language. For my part. have unanimously. continents may have existed where oceans are now spread out. etc. We see this in the fact that the most eminent palaeontologists. only here and there a few lines. Of this volume. only here and there a short chapter has been preserved. Barrande. etc. might have undergone far more metamorphic action than strata which have always remained nearer to the surface. will undoubtedly at once reject my theory. But Sir Charles Lyell now gives the support of his high authority to the opposite side. are all undoubtedly of the most serious nature. I look at the geological record as a history of the world imperfectly kept and written in a changing dialect. Forbes.

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



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.


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


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.


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


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.


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


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


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


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.


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


and find how slight is the relation between the physical conditions of various countries and the nature of their inhabitants. MM. their extinction. de Verneuil and d'Archiac. and if this be so. look to some special law. So. and Australia. but depend on general laws which govern the whole animal kingdom. Nevertheless. indeed. It is. This great fact of the parallel succession of the forms of life throughout the world. "If struck by this strange sequence. under the most different climates. it will appear certain that all these modifications of species. We must. and all those that lived in Europe during the pleistocene period (a very remote period as measured by years. and the introduction of new ones. the pleistocene and strictly modern beds of Europe. they add. as the cause of these great mutations in the forms of life throughout the world. from containing fossil remains in some degree allied. 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. which are already dominant. again. looking to a remotely future epoch. New species are formed by having some advantage over older forms. North and South America. The fact of the forms of life changing simultaneously in the above large sense. climate. After referring to the parallelism of the palaeozoic forms of life in various parts of Europe. would be correctly ranked as simultaneous in a geological sense. there can be little doubt that all the more modern MARINE formations.living in Europe. has greatly struck those admirable observers. including the whole glacial epoch) were compared with those now existing in South America or in Australia. namely. and the forms. Barrande has made forcible remarks to precisely the same effect. and from not including those forms which are found only in the older underlying deposits. cannot be owing to mere changes in marine currents or other causes more or less local and temporary. and there discover a series of analogous phenomena. as Barrande has remarked. We shall see this more clearly when we treat of the present distribution of organic beings. the upper pliocene. at distant parts of the world. or other physical conditions. 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. 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." M. quite futile to look to changes of currents. or have some advantage over the other 347 . is explicable on the theory of natural selection. than to the present inhabitants of Europe. we turn our attention to North America.

in the plants which are dominant. accords well with the principle of new species having been formed by dominant species spreading widely and varying. to a certain extent. and again spreading. We might therefore expect to find. and that there was 348 . and producing new forms. from inheriting some inferiority in common. and. a less strict degree of parallelism in the succession of the productions of the land than with those of the sea. give birth to the greatest number of new varieties or incipient species.forms in their own country. taken in a large sense. varying. rich in fossils. and. occurred during the periods when the bed of the sea was either stationary or rising. It is also natural that the dominant. therefore. as it seems to me. it is probable. which are commonest and most widely diffused. that is. There is one other remark connected with this subject worth making. as well as over other species. The process of diffusion would often be very slow. The old forms which are beaten and which yield their places to the new and victorious forms. owing to their having had some advantage over their already dominant parents. depending on climatal and geographical changes. the parallel. and that blank intervals of vast duration. the territories of other species. old groups disappear from the world. and of giving rise in new countries to other new varieties and species. and likewise when sediment was not thrown down quickly enough to embed and preserve organic remains. as new and improved groups spread throughout the world. were deposited during periods of subsidence. varying and far-spreading species. which have already invaded. be slower with the terrestrial inhabitants of distinct continents than with the marine inhabitants of the continuous sea. The diffusion would. simultaneous. the new species thus produced being themselves dominant. will generally be allied in groups. and the succession of forms everywhere tends to correspond both in their first appearance and final disappearance. We have distinct evidence on this head. I have given my reasons for believing that most of our great formations. as far as fossils are concerned. producing the greatest number of new varieties. should be those which would have the best chance of spreading still further. Thus. succession of the same forms of life throughout the world. and on the gradual acclimatization of new species to the various climates through which they might have to pass. on strange accidents. as we do find. but in the course of time the dominant forms would generally succeed in spreading and would ultimately prevail. During these long and blank intervals I suppose that the inhabitants of each region underwent a considerable amount of modification and extinction.

When two formations have been deposited in two regions during nearly. the same period. also. from the causes explained in the foregoing paragraphs. but we are very far from having any right to conclude that this has invariably been the case. indeed. in his admirable Memoirs on the eocene deposits of England and France. As we have reason to believe that large areas are affected by the same movement. 349 . but not exactly. but the species would not exactly correspond. but when he compares certain stages in England with those in France. it be assumed that an isthmus separated two seas inhabited by distinct. and immigration. the same general succession in the forms of life. Mr. in accordance with the general succession of the forms of life. and the order would falsely appear to be strictly parallel. unless. although he finds in both a curious accordance in the numbers of the species belonging to the same genera. for there will have been a little more time in the one region than in the other for modification. and that large areas have invariably been affected by the same movements. we should find in both. is able to draw a close general parallelism between the successive stages in the two countries. it is probable that strictly contemporaneous formations have often been accumulated over very wide spaces in the same quarter of the world. yet the species themselves differ in a manner very difficult to account for considering the proximity of the two areas. extinction. Barrande. Prestwich. 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. nevertheless he finds a surprising amount of difference in the species. in this case the several formations in the two regions could be arranged in the same order. I suspect that cases of this nature occur in Europe. nevertheless the species would not all be the same in the apparently corresponding stages in the two regions. shows that there is a striking general parallelism in the successive Silurian deposits of Bohemia and Scandinavia. but contemporaneous faunas. Lyell has made similar observations on some of the later tertiary formations.much migration from other parts of the world.

of prophetic or synthetic types. has shown in the most striking manner that many of the fossil mammals discovered by him in Attica serve to break down the intervals between existing genera. as the name given to it by Professor Gervais expresses. in fact. families. but the Macrauchenia of South America connects to a certain extent these two grand divisions. If we confine our attention either to the living or to the extinct species of the same class. Gaudry. is certainly true. but the extinct Halitherium had. No one will deny that the Hipparion is intermediate between the existing horse and certain other ungulate forms. M. it may be well to make some remarks on this subject. as applied to extinct animals. intermediate or connecting links. and these terms imply that such forms are." and it thus makes some approach to ordinary hoofed quadrupeds. and has placed certain Pachyderms in the same sub-order with ruminants. Another distinguished palaeontologist.4. and in the writings of Agassiz. On The Affinities Of Extinct Species To Each Other. All fall into a few grand classes. What a wonderful connecting link in the chain of mammals is the Typotherium from South America. That the extinct forms of life help to fill up the intervals between existing genera. But. it differs from living forms. as Buckland long ago remarked. and which cannot be placed in any existing order. or between them. but as this statement has often been ignored or even denied. The more ancient any form is. and this fact is at once explained on the principle of descent. according to Professor Flower. And To Living Forms Let us now look to the mutual affinities of extinct and living species. to which the Sirenia are in other respects allied. The Sirenia form a very distinct group of the mammals. and orders. without even a rudiment being left. he dissolves by gradations the apparently wide interval between the pig and the camel. Cuvier ranked the Ruminants and Pachyderms as two of the most distinct orders of mammals. an ossified thigh-bone "articulated to a well-defined acetabulum in the pelvis. In the writings of Professor Owen we continually meet with the expression of generalised forms. The Ungulata or hoofed quadrupeds are now divided into the even-toed or odd-toed divisions. extinct species can all be classed either in still existing groups. for example. the series is far less perfect than if we combine both into one general system. as a general rule. the more. but so many fossil links have been disentombed that Owen has had to alter the whole classification. The 350 . and to give some instances. and one of the most remarkable peculiarities in existing dugong and lamentin is the entire absence of hind limbs.

But in a natural classification many fossil species certainly stand between living species. by the ostrich and extinct Archeopteryx. "and to constitute connecting links with the aquatic carnivora. If by this term it is meant that an extinct form is directly intermediate in all its characters between two living forms or groups. such as fish and reptiles. we must admit that there is truth in the remark. and it would be difficult to prove the truth of the proposition. so that the two groups formerly made a somewhat nearer approach to each other than they now do. and some extinct genera between living genera. seems to be that. The most common case. and the eocene Mammals. the older Fish. Turning to the Invertebrata. the older Cephalopods. on the one hand. one of the Dinosaurians—that group which includes the most gigantic of all terrestrial reptiles. as the Lepidosiren. a higher authority could not be named. which have been placed by some naturalists in an order by themselves. or group of species." 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. Barrande asserts. are considered by Professor Huxley to be undoubtedly cetaceans. or groups of species. is discovered having affinities directed towards very distinct groups. for every now and then even a living animal. This remark no doubt must be restricted to those groups which have undergone much change in the course of geological ages. with the recent members of the same classes. It is a common belief that the more ancient a form is. Yet if we compare the older Reptiles and Batrachians. 351 . by so much the more it tends to connect by some of its characters groups now widely separated from each other. Some writers have objected to any extinct species. the ancient members are separated by a somewhat lesser number of characters. yet that at this ancient period the groups were not so distinctly separated from each other as they now are. and on the other hand by the Compsognathus. even between genera belonging to distinct families. but the tertiary Zeuglodon and Squalodon. the objection is probably valid. being considered as intermediate between any two living species. although palaeozoic animals can certainly be classed under existing groups. supposing them to be distinguished at the present day by a score of characters. that he is every day taught that. especially with respect to very distinct groups.cetaceans or whales are widely different from all other mammals.

On the principle of the continued tendency to divergence of character. the three existing families on the uppermost line would be rendered less distinct from each other. and some to have endured to the present day. a third family. If. As the subject is somewhat complex. m3. a14. i14. were disinterred. which was formerly illustrated by this diagram. Therefore it is quite possible. extinct and recent. f8. The three existing genera. for all will have inherited something in common from their ancient progenitor. q14. p14. and all the forms beneath the uppermost line may be considered as extinct. The diagram is much too simple. it depends solely on the descendants from a species being thus enabled to seize on many and different places in the economy of nature. make. We may suppose that the numbered letters in italics represent genera. assume that divergence of character is a necessary contingency. the genera a1. We must not. descended from (A). Yet he who objected to consider as intermediate the extinct genera. m9. for instance. The horizontal lines may represent successive geological formations. one order. which thus link 352 . however. that a species might go on being slightly modified in relation to its slightly altered conditions of life. All the many forms. and o14. These three families. from the continued effects of extinction and divergence of character. I must request the reader to turn to the diagram in the fourth chapter. the more recent any form is the more it will generally differ from its ancient progenitor. m6. and the dotted lines diverging from them the species in each genus. we can understand the rule that the most ancient fossils differ most from existing forms. in nearly the same manner as has occurred with ruminants and certain pachyderms. some of which are supposed to have perished at different periods. and yet retain throughout a vast period the same general characteristics. By looking at the diagram we can see that if many of the extinct forms supposed to be embedded in the successive formations. b14 and f14. were discovered at several points low down in the series. Hence. m14. will form a small family. a10.Let us see how far these several facts and inferences accord with the theory of descent with modification. as before remarked. and this order. as we have seen in the case of some Silurian forms. a closely allied family or subfamily. a5. This is represented in the diagram by the letter F14. has become divided into several sub-families and families. these three families would be so closely linked together that they probably would have to be united into one great family. but this is unimportant for us. too few genera and too few species being given. together with the many extinct genera on the several lines of descent diverging from the parent form (A) will form an order.

etc. for they would at this early stage of descent have diverged in a less degree from their common progenitor. and this by the concurrent evidence of our best palaeontologists is frequently the case. On this same theory. we have no right to expect. And they are wholly inexplicable on any other view. and will have been modified in various degrees. undergone much modification. All that we have a right to expect is.) would have to be united into one. are explained in a satisfactory manner. 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.together the living genera of three families. and thus to unite distinct families or orders. be supposed to differ from each other by half-a-dozen important characters.. and are the parents of those which became still more modified at the seventh stage. on the theory of descent with modification. for they are intermediate. then only two of the families (those on the left hand a14. except in rare cases. etc. or between their collateral relations. on the uppermost line.—but none from beneath this line. Thus. but only by a long and circuitous course through many widely different forms. for the groups will have been more numerous. then the families which existed at a period marked VI would certainly have differed from each other by a less number of characters. Thus it comes that ancient and extinct genera are often in a greater or less degree intermediate in character between their modified descendants. As we possess only the last volume of the geological record. and b14. would be partly justified. not directly. above No. 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. the main facts with respect to the mutual affinities of the extinct forms of life to each other and to living forms. to fill up the wide intervals in the natural system. and that in a very broken condition. within known geological periods. 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. if the three families formed of eight genera (a14 to m14). 353 . VI. Under nature the process will be far more complicated than is represented in the diagram. So again. they will have endured for extremely unequal lengths of time. that those groups which have. should in the older formations make some slight approach to each other. If many extinct forms were to be discovered above one of the middle horizontal lines or geological formations—for instance. and there would remain two families which would be less distinct from each other than they were before the discovery of the fossils.

for the parent rock-pigeon still lives. especially in the case of terrestrial productions inhabiting separated districts. if the principal living and extinct races of the domestic pigeon were arranged in serial affinity. and many varieties between the rock-pigeon and the carrier have become extinct. namely. when arranged by Dr. were at once recognised by palaeontologists as intermediate in character between those of the overlying carboniferous and underlying Silurian systems. this arrangement would not closely accord with the order in time of their production. when this system was first discovered. and in any one region for the immigration of new forms from other regions. Subject to these allowances. Closely connected with the statement. 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. allow for the entire extinction of some preceding forms. 354 . But supposing for an instant. But each fauna is not necessarily exactly intermediate. that the organic remains from an intermediate formation are in some degree intermediate in character. between the preceding and succeeding faunas. the species of mastodons and elephants. the manner in which the fossils of the Devonian system. in this and other such cases. nor are those which are intermediate in character. the fauna of each geological period undoubtedly is intermediate in character. Falconer in two series—in the first place according to their mutual affinities. as unequal intervals of time have elapsed between consecutive formations. intermediate in age. and even less with the order of their disappearance. that certain genera offer exceptions to the rule. that the record of the first appearance and disappearance of the species was complete. which is far from the case. and for a large amount of modification during the long and blank intervals between the successive formations. I need give only one instance. and carriers which are extreme in the important character of length of beak originated earlier than short-beaked tumblers. For instance. The species extreme in character are not the oldest or the most recent. we have no reason to believe that forms successively produced necessarily endure for corresponding lengths of time. To compare small things with great. which are at the opposite end of the series in this respect. however. A very ancient form may occasionally have lasted much longer than a form elsewhere subsequently produced. We must.hence they could hardly fail to be nearly intermediate in character between the forms of life above and below. and in the second place according to their periods of existence—do not accord in arrangement.

seems to have shaken Professor Pictet in his belief in the immutability of species. will not attempt to account for the close resemblance of distinct species in closely consecutive formations. such evidence of the slow and scarcely sensible mutations of specific forms. as they have been called by some authors. On the theory of descent. all the intermediate varieties between the species which appeared at the commencement and close of these periods: but we ought to find after intervals. though the species are distinct in each stage. as I attempted to show in the last chapter. in short. the full meaning of the fossil remains from closely consecutive formations. that fossils from two consecutive formations are far more closely related to each other. and note how little the specific forms of the inhabitants of the sea have been affected. have changed almost simultaneously throughout the world. we ought not to expect to find. and these assuredly we do find. or. very long as measured by years. Let it be remembered that the forms of life. This fact alone. closely allied forms. though ranked as distinct species. He who is acquainted with the distribution of existing species over the globe. is obvious. As the accumulation of each formation has often been interrupted. the general resemblance of the organic remains from the several stages of the Chalk formation. than are the fossils from two remote formations. by the physical conditions of the ancient areas having remained nearly the same. We find. from its generality. representative species. being closely related. and therefore under the most different climates and conditions. but only moderately long as measured geologically. and as long blank intervals have intervened between successive formations. which includes the whole glacial epoch. as we have the right to expect. at least those inhabiting the sea. 355 . Consider the prodigious vicissitudes of climate during the pleistocene period. insisted on by all palaeontologists.is the fact. in any one or in any two formations. Pictet gives as a well-known instance.

They would likewise be fatal. if it included advance in organisation as a necessary contingent. as well as by the standard of the specialisation of organs. as yet suggested. as would the secondary by the eocene. as far as is known. the former would be beaten and exterminated by the latter. 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. or the above Brachiopods during the Cambrian formation. for some organisms would have to remain fitted for simple conditions of life. they first appeared. and in this sense higher. and the palaeozoic by the secondary forms. on the theory of natural selection. So that by this fundamental test of victory in the battle for life. It is not an insuperable difficulty that Foraminifera have not. for they have to beat in the struggle for life all the older forms. progressed in organisation since even the Laurentian epoch. new species become superior to their predecessors. yet leaving such degraded beings better fitted for their new walks of life. to stand higher than ancient forms. could be proved to have first come into existence during the Laurentian epoch. We have also seen that. and in some cases will even degrade or simplify the organisation. and it seems that this answer must be admitted as true.5. and that certain land and fresh-water shells have remained nearly the same. for instance. if the above Foraminifera. 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. is the best standard. with which they come into close competition. there would not have been time sufficient for the development 356 . from the time when. It is no valid objection to this conclusion. so natural selection will tend to render the organisation of each being more specialised and perfect. not but that it may leave many creatures with simple and unimproved structures fitted for simple conditions of life. In another and more general manner. though difficult of proof. Is this the case? A large majority of palaeontologists would answer in the affirmative. for in this case. of their degree of perfection or highness. as the specialisation of parts is an advantage to each being. as insisted on by Dr. that certain Brachiopods have been but slightly modified from an extremely remote geological epoch. when arrived at maturity. 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. modern forms ought. Carpenter.

of these organisms up to the standard which they had then reached. 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. some look at the selaceans or sharks. at two periods. for their further continued process. from their approach in some important points of structure to reptiles. does not extend far enough back to show with unmistakable clearness that within the known history of the world organisation has largely advanced. the latter at the present day are largely preponderant in number. as the highest fish. When advanced up to any given point. Even at the present day. cephalopods and brachiopods. naturalists are not unanimous which forms ought to be ranked as highest: thus. 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. Beside these inherent difficulties in deciding which forms are the most advanced in organisation. The geological record. and in this case. not very high in their own class. according to the standard of highness chosen. The problem whether organisation on the whole has advanced is in many ways excessively intricate. though not highly developed. would stand very high in the scale of invertebrate animals. on the theory of natural selection. and such crustaceans. others look at the teleosteans as the highest. and at what period the various forms of life first appeared. although upon another type?" In the complex struggle for life it is quite credible that crustaceans. so as to hold their places in relation to slight changes in their conditions. consequently some naturalists maintain that molluscs were formerly more highly developed than at present. there is no necessity. namely. looking to members of the same class. might beat cephalopods. high and low. The ganoids stand intermediate between the selaceans and teleosteans. At an ancient epoch the highest and lowest molluscoidal animals. if judged by the most decisive of all trials—the law of battle. though they will. intermediate in organisation. during each successive age. so will it be said that fishes have advanced or retrograded in organisation. swarmed in numbers. while others. at all times imperfect. at the present time both groups are greatly reduced. but a stronger case can 357 . have largely increased. To attempt to compare members of distinct types in the scale of highness seems hopeless. the highest molluscs. have to be slightly modified. but formerly selaceans and ganoids alone existed. The foregoing objections hinge on the question whether we really know how old the world is. and this may well be disputed.

we may well doubt whether. and having been inherited at a corresponding age. we must believe. We ought also to compare the relative proportional numbers. any considerable number would be enabled to seize on places now occupied by our native plants and animals. From the extraordinary manner in which European productions have recently spread over New Zealand. for instance. are more highly organised than their ancient representatives. at any two periods. preserved by nature. This view accords admirably well with our theory. owing to variations having supervened at a not early age. if all the productions of New Zealand were set free in Great Britain. in the course of successive generations. and would exterminate many of the natives. of the former 358 . Thus the embryo comes to be left as a sort of picture. and the fact that our existing cephalopods. continually adds. that if all the animals and plants of Great Britain were set free in New Zealand. This process. from an examination of the species of the two countries. Under this point of view. more and more difference to the adult. under such extremely complex relations. We thus see how hopelessly difficult it is to compare with perfect fairness. We shall appreciate this difficulty more clearly by looking to certain existing faunas and floras. as a decided advance in the organisation of the world. and have seized on places which must have been previously occupied by the indigenes. we ought to look at this increase in number in the highest class. and if we knew that at some former period only ten thousand kinds existed. On the other hand. though few in number. could not have foreseen this result. which implies a great displacement of lower forms.be made out on the opposite side. In a future chapter I shall attempt to show that the adult differs from its embryo. at the present day fifty thousand kinds of vertebrate animals exist. whilst it leaves the embryo almost unaltered. by considering the vast reduction of brachiopods. from the fact that hardly a single inhabitant of the southern hemisphere has become wild in any part of Europe. of the high and low classes throughout the world: if. a multitude of British forms would in the course of time become thoroughly naturalized there. the standard of organisation of the imperfectly-known faunas of successive periods. the productions of Great Britain stand much higher in the scale than those of New Zealand. 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. Yet the most skilful naturalist. and that the geological succession of extinct forms is nearly parallel with the embryological development of existing forms.

that the oldest known mammals. This view may be true. and fishes strictly belong to their proper classes. and yet may never be capable of proof. for instance. reptiles.and less modified condition of the species. Seeing. until beds rich in fossils are discovered far beneath the lowest Cambrian strata—a discovery of which the chance is small. it would be vain to look for animals having the common embryological character of the Vertebrata. 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. 359 .

Now. after comparing the present climate of Australia and of parts of South America. would attempt to account. in 1839 and 1845. for the dissimilarity of the inhabitants of these two continents." Professor Owen has subsequently extended the same generalisation to the mammals of the Old World. in the gigantic pieces of armour. found in several parts of La Plata. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent. on this "law of the succession of types. are related to South American types. on the other hand through similarity of conditions. or that Edentata and other American types should have been solely produced in South America. what does this remarkable law of the succession of the same types within the same areas mean? He would be a bold man who. North America formerly partook strongly of the present character of the southern half of the continent. We see it also in the birds of the caves of Brazil. and the southern half was formerly more 360 . Woodward has shown that the same law holds good with sea-shells. and between the extinct and living brackish water-shells of the Aralo-Caspian Sea. like those of the armadillo. even to an uneducated eye."—on "this wonderful relationship in the same continent between the dead and the living.6. it is not well displayed by them. I was so much impressed with these facts that I strongly insisted. and I have shown in the publications above alluded to. under the same latitude. Nor can it be pretended that it is an immutable law that marsupials should have been chiefly or solely produced in Australia. and Professor Owen has shown in the most striking manner that most of the fossil mammals. but. on the one hand through dissimilar physical conditions. that in America the law of distribution of terrestrial mammals was formerly different from what it now is. Lund and Clausen in the caves of Brazil. buried there in such numbers. In South America. from the wide distribution of most molluscs. On The Succession Of The Same Types Within The Same Areas. During The Later Tertiary Periods Mr. a similar relationship is manifest. For we know that Europe in ancient times was peopled by numerous marsupials. We see the same law in this author's restorations of the extinct and gigantic birds of New Zealand. Mr. as the relation between the extinct and living land-shells of Madeira. and. Other cases could be added. This relationship is even more clearly seen in the wonderful collection of fossil bones made by MM. for the uniformity of the same types in each continent during the later tertiary periods.

This cannot for an instant be admitted. on our theory. It must not be forgotten that. It may be asked in ridicule whether I suppose that the megatherium and other allied huge monsters. be found in one geological formation. than it is at present. the great law of the long enduring. and anteater. succession of the same types within the same areas. and some of these fossils may have been the actual progenitors of the living species. then we may conclude that generally only one species of each of the older genera has left modified descendants. but not immutable. each having eight species. the other seven species of each old genus having died out and left no progeny. which formerly lived in South America. each with the same number of species. the feebler will yield to the more dominant forms. so will their modified descendants still differ in nearly the same manner and degree. On the theory of descent with modification. closely allied though in some degree modified descendants. If the inhabitants of one continent formerly differed greatly from those of another continent. 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. as their degenerate descendants. with the genera and species decreasing in numbers as is the case with the Edentata of South America. Analogous facts could be given in relation to the distribution of marine animals. 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. for the inhabitants of each quarter of the world will obviously tend to leave in that quarter. is at once explained. These huge animals have become wholly extinct. In failing orders. permitting much intermigration. armadillo. and in a succeeding formation there be six other allied or representative genera. so that. and after great geographical changes. that Northern India was formerly more closely related in its mammals to Africa than it is at the present time. and there will be nothing immutable in the distribution of organic beings. to the northern half. which constitute the new genera containing the several species. and have left no progeny. Or. still fewer genera and species will leave modified blood-descendants. during the next succeeding period of time. But after very long intervals of time. In a similar manner we know.closely allied. from Falconer and Cautley's discoveries. 361 . all the species of the same genus are the descendants of some one species. if six genera. have left behind them the sloth. and this will be a far commoner case.

although each species must have passed through numerous transitional stages. for it is not pretended that we have any sure criterion by which species and varieties can be discriminated. is absolutely as nothing compared with the number of generations which must have passed away even during a single formation. would be ranked. owing to subsidence being almost necessary for the accumulation of deposits rich in fossil species of many kinds.7. preserved in our museums. connecting together all extinct and existing forms by the finest graduated steps. It should also be constantly borne in mind that any linking variety between two forms. when the 362 . and lastly. and have oftenest given rise to new species. have been short in comparison with the periods during which each remained in an unchanged condition. that only a small portion of the globe has been geologically explored with care. that migration has played an important part in the first appearance of new forms in any one area and formation. that varieties have at first been local. though many and long as measured by years. that widely ranging species are those which have varied most frequently. and during the latter the record will have been least perfectly kept. taken conjointly. that the number both of specimens and of species. that only certain classes of organic beings have been largely preserved in a fossil state. which might be found. For he may ask in vain where are the numberless transitional links which must formerly have connected the closely allied or representative species. will to a large extent explain why—though we do find many links—we do not find interminable varieties. that. as a new and distinct species. He who rejects this view of the imperfection of the geological record. 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. These causes. it is probable that the periods. that there has probably been more extinction during the periods of subsidence. and thick enough to outlast future degradation. and more variation during the periods of elevation. during which each underwent modification. that each single formation has not been continuously deposited. will rightly reject the whole theory. unless the whole chain could be perfectly restored. Summary Of The Preceding And Present Chapters I have attempted to show that the geological record is extremely imperfect. great intervals of time must have elapsed between most of our successive formations. he may overlook how important a part migration has played. that the duration of each formation is probably short compared with the average duration of specific forms.

Hence. lingering in protected and isolated situations. but that. the productions of the world appear to have changed simultaneously. the world presented a widely different aspect. and to leave no modified offspring on the face of the earth. 363 . We can thus understand how it is that new species come in slowly and successively. are considered. Groups of species increase in numbers slowly. or lie still buried under the ocean. 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. long before that epoch. which form new sub-groups and groups. formed of formations older than any known to us. but modified. and endure for unequal periods of time. how species of different classes do not necessarily change together. from their inferiority inherited from a common progenitor. exist now only as remnants in a metamorphosed condition. The dominant species belonging to large and dominant groups tend to leave many modified descendants. or at the same rate. and where our oscillating continents now stand they have stood since the commencement of the Cambrian system. or in the same degree. We can understand why. tend to become extinct together. the other great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and natural selection. as those of Europe. and depends on many complex contingencies. and that the older continents. he may urge the apparent.formations of any one great region. yet in the long run that all undergo modification to some extent. As these are formed. for the link of generation has been broken. after long intervals of time. when a species has once disappeared. it never reappears. descendants. and these will generally succeed in displacing the groups which are their inferiors in the struggle for existence. but often falsely apparent. the species of the less vigorous groups. But the utter extinction of a whole group of species has sometimes been a slow process. but I can answer this last question only by supposing that where our oceans now extend they have extended for an enormous period. 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. The extinction of old forms is the almost inevitable consequence of the production of new forms. When a group has once wholly disappeared. Passing from these difficulties. sudden coming in of whole groups of species. it does not reappear. for the process of modification is necessarily slow. from the survival of a few descendants.

We can clearly see why the remains of an intermediate formation are intermediate in character. the more often it stands in some degree intermediate between groups now distinct. for they are closely linked together by generation. the more it generally differs from those now living. 364 . in so far. and are. and it may at least be asserted that the record cannot be proved to be much more perfect. but more commonly bringing them only a little closer together. the more nearly it will be related to. from the continued tendency to divergence of character. that species have been produced by ordinary generation: old forms having been supplanted by new and improved forms of life. the main objections to the theory of natural selection are greatly diminished or disappear. On the other hand. and this may account for the common belief held by so many palaeontologists. and their structure has generally become more specialised. all the chief laws of palaeontology plainly proclaim. but are intermediate only by a long and circuitous course through other extinct and different forms. The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious. and this wonderful fact receives a simple explanation according to our views. sometimes blending two groups. then. the common progenitor of groups. Extinct and ancient animals resemble to a certain extent the embryos of the more recent animals belonging to the same classes.We can understand how it is that all the forms of life. the products of variation and the survival of the fittest. and consequently resemble. since become widely divergent. If. the geological record be as imperfect as many believe. previously classed as distinct into one. 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. why the more ancient a form is. The more ancient a form is. as it seems to me. Why ancient and extinct forms often tend to fill up gaps between existing forms. We can clearly see why the organic remains of closely consecutive formations are closely allied. higher in the scale. for the more ancient a form is. We can understand. Extinct forms are seldom directly intermediate between existing forms. ancient and recent. make together a few grand classes. that organisation on the whole has progressed.

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. 365 . and by occasional means — Dispersal during the Glacial period — Alternate Glacial periods in the North and South.

We see this in the great difference in nearly all the terrestrial productions of the New and Old Worlds. from the central parts of the United States to its extreme southern point. grassy plains. and western South America. all authors agree that one of the most fundamental divisions in geographical distribution is that between the New and Old Worlds. In considering the distribution of organic beings over the face of the globe. South Africa. marshes. 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. The case of America alone would almost suffice to prove its truth. forests. how widely different are their living productions! In the southern hemisphere. arid deserts. Or. are related in a close and important manner to the differences between the productions of various regions. for it is rare to find a group of organisms confined to a small area. but these are not inhabited by a fauna different from that of the surrounding districts. the first great fact which strikes us is. we may compare the productions of South America south of latitude 35 degrees with those north of 25 degrees.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. that barriers of any kind. No doubt small areas can be pointed out in the Old World hotter than any in the New World. which consequently are separated by a space of ten degrees of latitude. lakes and great rivers. if we compare large tracts of land in Australia. yet it would not be possible to point out three faunas and floras more utterly dissimilar. of which the conditions are peculiar in only a slight degree. under almost every temperature. yet if we travel over the vast American continent. again. A second great fact which strikes us in our general review is. humid districts. almost every author who has studied the subject has come to this conclusion. 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. 366 . between latitudes 25 and 35 degrees. we meet with the most diversified conditions. Notwithstanding this general parallelism in the conditions of Old and New Worlds. for if we exclude the arctic and northern temperate parts. or obstacles to free migration. lofty mountains. Analogous facts could be given with respect to the inhabitants of the sea. Of late. excepting in the northern parts. we shall find parts extremely similar in all their conditions.

A third great fact. though the species themselves are distinct at different points and stations. and over this vast space we meet with no well-defined and distinct marine faunas. partly included in the foregoing statement. 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. Gunther has recently shown that about thirty per cent of the fishes are the same on the opposite sides of the isthmus of Panama. etc. though as mountain chains. On the other hand. until. or likely to have endured so long. we encounter no impassable barriers. they are almost wholly distinct. Westward of the shores of America. we come to the shores of Africa. but from being separated from each other by impassable barriers. and South America under the same latitude. and every continent offers innumerable 367 . for these countries are almost as much isolated from each other as is possible. there might have been free migration for the northern temperate forms. we find different productions. and where. as the oceans separating continents. we find the same law. or continuous coasts. after travelling over a hemisphere. and this fact has led naturalists to believe that the isthmus was formerly open. with extremely few shells. here we have a barrier of another kind. It is a law of the widest generality. we see the same fact. is the affinity of the productions of the same continent or of the same sea. under corresponding climate. with not an island as a halting-place for emigrants. under a slightly different climate. and as soon as this is passed we meet in the eastern islands of the Pacific with another and totally distinct fauna. The marine inhabitants of the eastern and western shores of South America are very distinct.. either of land or open sea. yet many fishes range from the Pacific into the Indian Ocean. for on the opposite sides of lofty and continuous mountain-ranges. Although so few marine animals are common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands. as there now is for the strictly arctic productions. proceeding still further westward from the eastern islands of the tropical parts of the Pacific. but Dr.where the land almost joins. a wide space of open ocean extends. also. and of great deserts and even of large rivers. and we have innumerable islands as halting-places. So that three marine faunas range northward and southward in parallel lines not far from each other. deserts. We see the same fact in the great difference between the inhabitants of Australia. Africa. On each continent. or echinodermata in common. the differences are very inferior in degree to those characteristic of distinct continents. Turning to the sea. are not as impassable. crustacea.

the most important of all relations. however much they may differ in geological structure. as we see in the case of varieties. animals having nearly the same habits as our hares and rabbits. On these same plains of La Plata we see the agouti and bizcacha. but not quite alike. or. in travelling. but they plainly display an American type of structure. Nevertheless. He hears from closely allied. Innumerable other instances could be given. independently of physical conditions. replace each other. yet distinct kinds of birds. the relation of organism to organism in the struggle for life being. notes nearly similar. like those inhabiting Africa and Australia under the same latitude. though nearly related. with eggs coloured in nearly the same manner. though they may be all peculiar species. Thus the high importance of barriers comes into play by checking migration. the naturalist. and northward the plains of La Plata by another species of the same genus. We ascend the lofty peaks of the Cordillera. The plains near the Straits of Magellan are inhabited by one species of Rhea (American ostrich). as I have already often remarked. and probably in a subordinate degree to the definite influence of different physical conditions. throughout space and time. nearly alike.instances. from north to south. for instance. and belonging to the same order of Rodents. The degrees of dissimilarity will depend on the migration of the more dominant forms of life from one region into another having been more or less effectually prevented. The bond is simply inheritance. The naturalist must be dull who is not led to inquire what this bond is. we look to the waters. never fails to be struck by the manner in which successive groups of beings. and we find an alpine species of bizcacha. and we find American types then prevailing on the American continent and in the American seas. the inhabitants are essentially American. and sees their nests similarly constructed. and we do not find the beaver or muskrat. Widely-ranging 368 . We see in these facts some deep organic bond. We may look back to past ages. The dissimilarity of the inhabitants of different regions may be attributed to modification through variation and natural selection. over the same areas of land and water. specifically distinct. 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. rodents of the South American type. as shown in the last chapter. as far as we positively know. and not by a true ostrich or emu. If we look to the islands off the American shore. that cause which alone. as does time for the slow process of modification through natural selection. produces organisms quite like each other. but the coypu and capybara.

must originally have proceeded from the same source. as is so commonly and notoriously the case. and thus they will become still further victorious. But in many other cases. were to migrate in a body into a new and afterwards isolated country. and will frequently undergo further modification and improvement. though inhabiting the most distant quarters of the world. of the existence of any law of necessary development. they would be little liable to modification. whole genera. In their new homes they will be exposed to new conditions. will have the best chance of seizing on new places. only so far as it profits each individual in its complex struggle for life. As the variability of each species is an independent property. According to these views.species. and even families. and have not become greatly or at all modified. during whole geological periods. If a number of species. though now inhabiting distant and isolated regions. must have proceeded from one spot. there is not much difficulty in believing that they have migrated from the same region. where their parents were first produced: for. There is no evidence. and will be taken advantage of by natural selection. almost any amount of migration is possible. 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. after having long competed with each other in their old home. and will produce groups of modified descendants. are confined to the same areas. In the case of those species which have undergone. for neither migration nor isolation in themselves effect anything. so certain species have migrated over vast spaces. it is obvious that the several species of the same genus. It is also obvious that the individuals of the same species. when they spread out into new countries. which have already triumphed over many competitors in their own widely-extended homes. as they are descended from the same progenitor. On this principle of inheritance with modification we can understand how it is that sections of genera. so the amount of modification in different species will be no uniform quantity. for during the vast geographical and climatical changes which have supervened since ancient times. as was remarked in the last chapter. it is incredible that 369 . As we have seen in the last chapter that some forms have retained nearly the same character from an enormously remote geological period. abounding in individuals. there is great difficulty on this head. as has been explained. in which we have reason to believe that the species of a genus have been produced within comparatively recent times. little modification.

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

Single Centres Of Supposed Creation We are thus brought to the question which has been largely discussed by naturalists. No geologist feels any difficulty in Great Britain possessing the same quadrupeds with the rest of Europe. What a strange anomaly it would be if a directly opposite rule were to prevail when we go down one step lower in the 371 . we find no inexplicable instances of the same mammals inhabiting distant points of the world. and it has been observed by several naturalists that the most natural genera. and have not been able to migrate to the opposite side. accordingly. where now found. and calls in the agency of a miracle. He who rejects it. so that a multitude of European animals and plants have become naturalised in America and Australia. as I believe. or if they have a wide range that their range is continuous. that in most cases the area inhabited by a species is continuous. Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. namely. the fact is given as something remarkable and exceptional. But if the same species can be produced at two separate points. is. are generally confined to the same country. are confined to a single region. have migrated across the wide and broken interspaces. is intelligible only on the view that the great majority of species have been produced on one side. that the space could not have been easily passed over by migration. Some few families. or with an interval of such a nature. and that when a plant or animal inhabits two points so distant from each other. It is universally admitted. and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres? The answer. 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. rejects the vera causa of ordinary generation with subsequent migration. whereas some plants. or those genera in which the species are most closely related to each other. whether species have been created at one or more points of the earth's surface. that mammals have not been able to migrate. for they were no doubt once united. The incapacity of migrating across a wide sea is more clear in the case of terrestrial mammals than perhaps with any other organic beings. The great and striking influence of barriers of all kinds.2. a still greater number of sections of genera. and. many subfamilies. from their varied means of dispersal. why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same. very many genera.

whether the several species of a genus which must on our theory all be descended from a common progenitor. that we ought to give up the belief. and these had not been. the wide distribution of fresh water productions. In discussing this subject we shall be enabled at the same time to consider a point equally important for us. It would be hopelessly tedious to discuss all the exceptional cases of the same species. nor do I for a moment pretend that any explanation could be offered of many instances. and secondly (in the following chapter). the occurrence of the same terrestrial species on islands and on the nearest mainland. it can be shown that migration from the one region to the other has probably occurred at some former period. that the view of each species having been produced in one area alone. namely. the existence of the same species on the summits of distant mountain ranges. and at distant points in the Arctic and Antarctic regions. and to the various occasional means of transport. is the most probable. rendered probable by general considerations. undergoing modification during their migration from some one area. But the geographical and climatical changes which have certainly occurred within recent geological times. 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. that each species has been produced within one area. though closely allied to them. our general view will be much strengthened. now living at distant and separated points. for the explanation is obvious on the 372 . So that we are reduced to consider whether the exceptions to continuity of range are so numerous. can have migrated. I will discuss a few of the most striking classes of facts. and thirdly. when most of the species inhabiting one region are different from those of another region. If. Undoubtedly many cases occur in which we cannot explain how the same species could have passed from one point to the other. and has migrated thence as far as it could. must have rendered discontinuous the formerly continuous range of many species. But. after some preliminary remarks. confined to some one region! Hence. and having subsequently migrated from that area as far as its powers of migration and subsistence under past and present conditions permitted. the belief that a single birthplace is the law seems to me incomparably the safest. it seems to me. considering our ignorance with respect to former climatical and geographical changes. and of so grave a nature. though separated by hundreds of miles of open sea. then. namely to the individuals of the same species. namely. at least at first. as it has to many other naturalists.series.

upheaved and formed at the distance of a few hundreds of miles from a continent. and the whole amount of modification at each stage will not be due to descent from a single parent. whether all the individuals of the same species are descended from a single pair. Cases of this nature are common. Wallace." And it is now well known that he attributes this coincidence to descent with modification. with all organisms which habitually unite for each birth. or whether. does not differ much from that advanced by Mr. from many individuals simultaneously created.principle of descent with modification. as we shall hereafter see. would still be related by inheritance to the inhabitants of that continent. or single hermaphrodite. inexplicable on the theory of independent creation. A volcanic island. that have supplanted each other. at each successive stage of modification. would probably receive from it in the course of time a few colonists. With organic beings which never intercross. But in the great majority of cases. for instance. This view of the relation of the species of one region to those of another. or which occasionally intercross. namely. To illustrate what I mean: our English race-horses differ from the horses of every other breed. but to continued care in the selecting and training of many individuals during each generation. if such exist. the individuals of the same species inhabiting the same area will be kept nearly uniform by intercrossing. but have never blended with other individuals or varieties of the same species. must be descended from a succession of modified varieties. and their descendants. each species. and are. so that many individuals will go on simultaneously changing. all the individuals of the same form will be descended from a single parent. The question of single or multiple centres of creation differs from another though allied question. though modified. 373 ." I must say a few words on the means of dispersal. as some authors suppose. which I have selected as presenting the greatest amount of difficulty on the theory of "single centres of creation. Before discussing the three classes of facts. but they do not owe their difference and superiority to descent from any single pair. who concludes that "every species has come into existence coincident both in space and time with a pre-existing closely allied species. so that. namely.

and the two faunas will now blend together. as it will some day be. I can give here only the briefest abstract of the more important facts. have been 374 . Edward Forbes insisted that all the islands in the Atlantic must have been recently connected with Europe or Africa. and thus have allowed terrestrial productions to pass from one to the other. It seems to me that we have abundant evidence of great oscillations in the level of the land or sea. and Europe likewise with America. we shall be enabled to speculate with security on the former extension of the land.3. when the climate was different. which may have served as halting places for plants and for many animals during their migration. now buried beneath the sea. indeed. that each species has proceeded from a single birthplace. however. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas. or may formerly have blended. and removes many a difficulty. Where the sea now extends. Change of climate must have had a powerful influence on migration. presently have to discuss this branch of the subject in some detail. If. Means Of Dispersal Sir C. A region now impassable to certain organisms from the nature of its climate. but not of such vast changes in the position and extension of our continents. In the coral-producing oceans such sunken islands are now marked by rings of coral or atolls standing over them. land may at a former period have connected islands or possibly even continents together. I freely admit the former existence of many islands. 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. I shall. No geologist disputes that great mutations of level have occurred within the period of existing organisms. Lyell and other authors have ably treated this subject. and united almost every island with some mainland. Whenever it is fully admitted. the arguments used by Forbes are to be trusted. it must be admitted that scarcely a single island exists which has not recently been united to some continent. Other authors have thus hypothetically bridged over every ocean. submerge it. might have been a high road for migration. This view cuts the Gordian knot of the dispersal of the same species to the most distant points. or let it formerly have been submerged. and when in the course of time we know something definite about the means of distribution. but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. as to have united them within the recent period to each other and to the several intervening oceanic islands.

but which more properly should be called occasional means of distribution. For convenience sake I chiefly tried small seeds without the capsules or fruit. etc. Nor does the almost universally volcanic composition of such islands favour the admission that they are the wrecks of sunken continents. if they had originally existed as continental mountain ranges. The nature and relative proportions of the inhabitants of oceanic islands are likewise opposed to the belief of their former continuity of continents. metamorphic schists. 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. or almost continuously united with each other. sixtyfour germinated after an immersion of twenty-eight days. were all killed by a month's immersion. of granite. seven species of the allied orders. To my surprise I found that out of eighty-seven kinds. and as all of these sank in a few days. some at least of the islands would have been formed. Hydrophyllaceae and Polemoniaceae. Until I tried. whether or not they were injured by salt water. a few experiments. It is well known what a difference there is in the buoyancy of green and seasoned timber. they could not have been floated across wide spaces of the sea. with Mr. I shall here confine myself to plants. like other mountain summits. Hence I was led to dry the stems and branches of ninety-four plants with ripe 375 . and some of these floated for a long time. It deserves notice that certain orders were far more injured than others: nine Leguminosae were tried. capsules. and. they resisted the salt-water badly. 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. and a few survived an immersion of 137 days. I must now say a few words on what are called accidental means. and with the many existing oceanic islands.continuously. Berkeley's aid. but the greater or less facilities for transport across the sea may be said to be almost wholly unknown. 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. Afterwards I tried some larger fruits. this or that plant is often stated to be ill adapted for wide dissemination. In botanical works. with one exception. instead of consisting of mere piles of volcanic matter. it was not even known how far seeds could resist the injurious action of sea-water. old fossiliferous and other rocks.. as are necessary on the view advanced by Forbes and admitted by his followers.

and some of the eighteen floated for a very much longer period. ripe hazel-nuts sank immediately. when dried floated much longer. out of the ninety-four dried plants. floated for a very short time. and this. and afterwards germinated. after having been dried. would have caused some of them to have floated much longer. on this average. but in a much better manner. as we have seen. we may conclude. So that as 64/87 kinds of seeds germinated after an immersion of twenty-eight days. M. so that they were alternately wet and exposed to the air like really floating plants. for above twenty-eight days. and would then germinate. 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). from plants which live near the sea. for he placed the seeds in a box in the actual sea. when dried they floated for above ninety days. Altogether. could be floated across a space of sea 900 miles in width. The fact of the larger fruits often floating longer than the small. Subsequently to my experiments. that the seeds of 14/ 100 kinds of plants of any country might be floated by sea-currents during twenty-eight days. and would retain their power of germination. but he chose many large fruits. 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. and to place them on sea-water. On the other hand. but when dried they floated for ninety days. the seeds of 14/100 plants belonging to one country might be floated across 924 miles of sea to another country. The majority sank quickly. when dried it floated for eighty-five days. after being dried. and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days. mostly different from mine. and this would have favoured both the average length of their flotation and their resistance to the injurious action of the salt-water. He tried ninety-eight seeds. eighteen floated for above twenty-eight days. Therefore. and were then capable of germination. he did not previously dry the plants or branches with the fruit. and as 18/94 distinct species with ripe fruit (but not all the same species as in the foregoing experiment) floated. is 376 . but some which. an asparagus plant with ripe berries floated for twenty-three days. would germinate. for instance. as far as anything can be inferred from these scanty facts. and when stranded. and afterwards when planted germinated. The result was that 18/98 of his seeds of different kinds floated for forty-two days.fruit. Martens tried similar ones. it would perhaps be safer to assume that the seeds of about 10/100 plants of a flora. In Johnston's Physical Atlas. whilst green. if blown by an inland gale to a favourable spot. and likewise seeds.

for instance. I picked up in my garden twelve kinds of seeds. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the ocean. Again. Seeds may be occasionally transported in another manner. I have never seen an instance of nutritious seeds passing through the intestines of a bird. are killed by even a few days' immersion in sea-water. Drift timber is thrown up on most islands. which were tried. and hawks are known to look out for tired birds. solely from the roots of drifted trees. and the natives of the coral islands in the Pacific procure stones for their tools. sometimes escape being immediately devoured. and the contents of their torn crops might thus readily get scattered. could hardly be transported by any other means. which had floated on artificial sea-water for thirty days. In the course of two months. Some hawks and owls bolt their prey whole. and after an interval of from twelve to twenty hours. generally have restricted ranges.interesting. now. germinated. and do not. when floating on the sea. disgorge pellets. and some authors have given a far higher estimate. as I know from experiments made 377 . 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. A bird in this interval might easily be blown to the distance of five hundred miles. and these seemed perfect. to my surprise nearly all germinated. de Candolle has shown. as Alph. and many kinds of seeds in the crops of floating birds long retain their vitality: peas and vetches. but hard seeds of fruit pass uninjured through even the digestive organs of a turkey. after a bird has found and devoured a large supply of food. injure in the least the germination of seeds. it is positively asserted that all the grains do not pass into the gizzard for twelve or even eighteen hours. Living birds can hardly fail to be highly effective agents in the transportation of seeds. out of the excrement of small birds. But the following fact is more important: the crops of birds do not secrete gastric juice. as plants with large seeds or fruit which. even on those in the midst of the widest oceans. which. these stones being a valuable royal tax. small parcels of earth are very frequently enclosed in their interstices and behind them. We may safely assume that under such circumstances their rate of flight would often be thirty-five miles an hour. I find that when irregularly shaped stones are embedded in the roots of trees. but some taken out of the crop of a pigeon. three dicotyledonous plants germinated: I am certain of the accuracy of this observation. as I know by trial. I can show that the carcasses of birds. and some of them.

Certain seeds. Some seeds of the oat. out of which I extracted under the microscope several seeds. I forced many kinds of seeds into the stomachs of dead fish. They then disappeared over the sea. with a little cake of dry earth attached to the shank. as thick as the flakes of snow in the heaviest snowstorm. and in another case twenty-two grains of dry argillaceous earth from the foot of a partridge. R. I find. and several of these seeds retained the power of germination. such as that which visited Madeira. as suddenly as they had appeared. though on insufficient evidence. at least five or six miles in diameter. of Brighton. They were in countless numbers. were always killed by this process. and at night alighted on the taller trees. In consequence of this belief Mr. Swaysland. these birds. 378 . and then gave their bodies to fishing-eagles. clover.in the Zoological Gardens. Hence a swarm of locusts. wheat. fish are frequently devoured by birds. millet. Fresh-water fish. might readily be the means of introducing several kinds of plants into an island lying far from the mainland. Lowe informed Sir C. hemp. swarms of locusts visited the island of Madeira. which were completely coated with them. and thus the seeds might be transported from place to place. and beet germinated after having been from twelve to twenty-one hours in the stomachs of different birds of prey.T. in parts of Natal it is believed by some farmers. and have heard of others caught at greater distances. 1844. and extended upward as far as could be seen with a telescope. earth sometimes adheres to them: in one case I removed sixty-one grains. and this contained a seed of the toad-rush (Juncus bufonius) which germinated and flowered. Weale sent me in a letter a small packet of the dried pellets. Lyell that in November. canary. Mr. weighing only nine grains. and raised from them seven grass plants. I myself caught one 370 miles from the coast of Africa. include seeds capable of germination. and two seeds of beet grew after having been thus retained for two days and fourteen hours. Here is a better case: the leg of a woodcock was sent to me by a friend. that injurious seeds are introduced into their grass-land in the dung left by the great flights of locusts which often visit that country. Locusts are sometimes blown to great distances from the land. and in the earth there was a pebble as large as the seed of a vetch. eat seeds of many land and water plants. and have not since visited the island. either rejected the seeds in pellets or passed them in their excrement. storks. however. after an interval of many hours. Although the beaks and feet of birds are generally clean. and pelicans. Now. belonging to two species. During two or three days they slowly careered round and round in an immense ellipse. The Rev. of two genera.

which do not occur in the archipelago. I think. I suspected that these islands had been partly stocked by ice-borne seeds during the Glacial epoch. In the Azores. and it is at least possible that they may have brought thither the seeds of northern plants. With such facts before us. and which annually migrate—for instance. H. and he has several times noticed little cakes of earth attached to their feet.C. in comparison with the latitude. and at least one kind of grass. Hence we may safely infer that icebergs formerly landed their rocky burdens on the shores of these mid-ocean islands. have been in action year after year for tens of thousands of years. For instance. Hartung to inquire whether he had observed erratic boulders on these islands. judging from the young leaves. Considering that these several means of transport. which without doubt remain to be discovered. Lyell wrote to M. watered and placed under a bell glass. and the nest of a land-bird. on their first arrival on our shores. wheatears. bones. Many facts could be given showing how generally soil is charged with seeds. and during the Glacial period from one part of the now temperate regions to another. and that other means. but when broken. Watson) from their somewhat northern character. The earth had been kept for three years. which stand nearer to the mainland. as suggested by Lyell. and weighing six and a half ounces. and he answered that he had found large fragments of granite and other rocks. and of seventy dicotyledons. from the large number of plants common to Europe. have transported seeds from one part to another of the arctic and antarctic regions. As icebergs are known to be sometimes loaded with earth and stones. informs me that he has often shot wagtails (Motacillae). no less than eighty-two plants sprung from it: these consisted of twelve monocotyledons. At my request Sir C.who during the last forty years has paid close attention to our migratory birds. can we doubt that the many birds which are annually blown by gales across great spaces of ocean. before they had alighted. Professor Newton sent me the leg of a red-legged partridge (Caccabis rufa) which had been wounded and could not fly. it would. of at least three distinct species. with a ball of hard earth adhering to it. be a 379 . and whinchats (Saxicolae). including the common oat. and have even carried brushwood. which consisted. it can hardly be doubted that they must occasionally. 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. in comparison with the species on the other islands of the Atlantic. and (as remarked by Mr.

during the long lapse of geological time. would germinate and survive. but would remain as distinct as they now are. one or two land-birds are blown across the whole Atlantic Ocean. for seeds do not retain their vitality when exposed for a great length of time to the action of sea water. 380 . they could not endure our climate. though standing more remote from the mainland. and coming to maturity! But it would be a great error to argue that because a well-stocked island. that a poorlystocked island. though they might and do bring seeds from the West Indies to our western shores. but this is not strictly correct: the currents of the sea are not accidental. Almost every year. On almost bare land. however. would suffice for occasional transport across tracts of sea some hundred miles in breadth. would never bring seeds from North America to Britain. like Great Britain. but not from one distant continent to another. where. or from a continent to a neighbouring island. how small would be the chance of a seed falling on favourable soil. received within the last few centuries. nor is the direction of prevalent gales of wind. as far as is known (and it would be very difficult to prove this). through occasional means of transport. by dirt adhering to their feet or beaks. if not killed by their very long immersion in salt water. and before it had become fully stocked with inhabitants. namely. as to become naturalised. even if far less well-stocked than Britain. or from island to island. from North America to the western shores of Ireland and England. immigrants from Europe or any other continent. nor could they be long carried in the crops or intestines of birds. These means of transport are sometimes called accidental. It should be observed that scarcely any means of transport would carry seeds for very great distances. whilst the island was being upheaved.marvellous fact if many plants had not thus become widely transported. if fitted for the climate. has not. The currents. would not receive colonists by similar means. with few or no destructive insects or birds living there. Even in this case. which is in itself a rare accident. These means. perhaps not more than one would be so well fitted to its new home. Out of a hundred kinds of seeds or animals transported to an island. But this is no valid argument against what would be effected by occasional means of transport. from their course. nearly every seed which chanced to arrive. but seeds could be transported by these rare wanderers only by one means. The floras of distant continents would not by such means become mingled.

As the cold came on. unless they were stopped by barriers.4. and then pass away. We have evidence of almost every conceivable kind. within a very recent geological period. The former influence of the glacial climate on the distribution of the inhabitants of Europe. and perched boulders. on mountain-summits. with their scored flanks. 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. had not Agassiz and others called vivid attention to the Glacial period. By the time that the cold had reached its maximum. central Europe and North America suffered under an Arctic climate. gigantic moraines. these would take the places of the former inhabitants of the temperate regions. The ruins of a house burnt by fire do not tell their tale more plainly than do the mountains of Scotland and Wales. and in the extreme northern parts of Europe. as formerly occurred. but it is far more remarkable. that the plants on the White Mountains. with those on the loftiest mountains of Europe. left by old glaciers. and we might have remained in this same belief. Throughout a large part of the United States. erratic boulders and scored rocks plainly reveal a former cold period. The mountains would become covered with snow and ice. and as each more southern zone became fitted for the inhabitants of the north. is one of the most striking cases known of the same species living at distant points. at the same time would travel further and further southward. So greatly has the climate of Europe changed. affords a simple explanation of these facts. we should have an arctic fauna and flora. are now clothed by the vine and maize. in the United States of America. Even as long ago as 1747. and their former Alpine inhabitants would descend to the plains. are all the same with those of Labrador. by supposing a new glacial period slowly to come on. separated from each other by hundreds of miles of lowlands. The latter. But we shall follow the changes more readily. is substantially as follows. as we hear from Asa Gray. without the apparent possibility of their having migrated from one point to the other. such facts led Gmelin to conclude that the same species must have been independently created at many distinct points. in which case they would perish. where Alpine species could not possibly exist. and nearly all the same. of the icy streams with which their valleys were lately filled. Dispersal During The Glacial Period The identity of many plants and animals. that. which. that in Northern Italy. covering the central 381 . as we shall immediately see. organic and inorganic. polished surfaces. as explained by Edward Forbes.

for example. those of the mountains of Siberia to the arctic regions of that country. H. Watson. the arctic forms would seize on the cleared and thawed ground. always ascending. as remarked by Mr. 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. of Scotland. and the re-migration on the returning warmth. closely followed up in their retreat by the productions of the more temperate regions. whilst their brethren were pursuing their northern journey. for the present circumpolar inhabitants. and even stretching into Spain. those of the United States to Labrador. are remarkably uniform round the world. 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. As the warmth returned. And as the snow melted from the bases of the mountains. in unison with the changing climate. Hence. the same species. These views. and as they all migrated in a body together. As the arctic forms moved first southward and afterwards backward to the north. they will not have been exposed during their long migrations to any great diversity of temperature. which had lately lived together on the European and North American lowlands. when the warmth had fully returned. would again be found in the arctic regions of the Old and New Worlds. and those of the Pyrenees. without other evidence. as the warmth increased and the snow still further disappeared. The Alpine plants. 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. their mutual relations 382 . the arctic forms would retreat northward. higher and higher. that when in other regions we find the same species on distant mountain-summits. 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. grounded as they are on the perfectly well-ascertained occurrence of a former Glacial period. are more especially allied to the plants of northern Scandinavia. would generally have been due south and north. as remarked by Ramond. which we suppose to have everywhere travelled southward.parts of Europe. now become too warm for their existence. and on many isolated mountain-summits far distant from each other.C. as far south as the Alps and Pyrenees. that a colder climate formerly permitted their migration across the intervening lowlands.

At the present day. the arctic productions were as uniform round the polar regions as they are at the present day. they will. 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.will not have been much disturbed. and have survived there ever since. I believe. some exist as varieties. for it is not likely that all the same arctic species will have been left on mountain ranges far distant from each other. in accordance with the principles inculcated in this volume. The explanation. have become mingled with ancient Alpine species. the majority of the inhabitants of the world were specifically the same as now. Hence. which must have existed on the mountains before the commencement of the Glacial epoch. so that it may well be asked how the same species could then or previously have entered the two continents. left isolated from the moment of the returning warmth. I have assumed that at the commencement of our imaginary Glacial period. During the Glacial period. they will also. and which during the coldest period will have been temporarily driven down to the plains. lies in the nature of the climate before the commencement of the Glacial period. these forms will not have been liable to much modification. in all probability. at the commencement of the real Glacial period. Their mutual relations will thus have been in some degree disturbed. for if we compare the present Alpine plants and animals of the several great European mountain ranges. and we have good reason to believe that the climate was 383 . But it is also necessary to assume that many sub-arctic and some few temperate forms were the same round the world. consequently they will have been liable to modification. But with the Alpine productions. some as doubtful forms or sub-species and some as distinct yet closely allied species representing each other on the several ranges. and it may be asked how I account for this degree of uniformity of the sub-arctic and temperate forms round the world. In the foregoing illustration. first at the bases and ultimately on the summits of the mountains. though many of the species remain identically the same. the newer Pliocene period. also. At this. and they have been modified. when the inhabitants of the Old and New Worlds lived further southwards than they do at present. the case will have been somewhat different. have been subsequently exposed to somewhat different climatical influences. 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. they must have been still more completely separated from each other by wider spaces of ocean. one with another.

to eastern America. both in the Old and New Worlds. must have taken place long ages ago. which inhabited the New and Old Worlds. under the Polar Circle. serving as a bridge. and that these plants and animals. Believing. and to infer that during some earlier and still warmer period. and in the other great region. with those of 384 . Hence. their descendants. migrated south of the Polar Circle. Now. at a period anterior to the Glacial epoch. will account for the supposed uniformity of the sub-arctic and temperate productions of the Old and New Worlds. between the productions of North America and Europe—a relationship which is highly remarkable.warmer than at the present day. that our continents have long remained in nearly the same relative position. began slowly to migrate southwards as the climate became less warm. lived during the Pliocene period further north. mostly in a modified condition. we may suppose that the organisms which now live under latitude 60 degrees. though subjected to great oscillations of level. long before the commencement of the Glacial period. they will have become mingled in the one great region with the native American productions. During the slowly decreasing warmth of the Pliocene period. from reasons before alluded to. for during these warmer periods the northern parts of the Old and New Worlds will have been almost continuously united by land. with the consequent freedom under a more favourable climate for intermigration. and their separation by the whole Atlantic Ocean. in latitude 66-67 degrees. they will have been completely cut off from each other. considering the distance of the two areas. as soon as the species in common. and would have had to compete with them. such as the older Pliocene period. as far as the more temperate productions are concerned. We now see. as I believe. This separation. and that the present arctic productions then lived on the broken land still nearer to the pole. 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. As the plants and animals migrated southward. On this view we can understand the relationship with very little identity. for the intermigration of their inhabitants. we see under the Polar Circle that there is almost continuous land from western Europe through Siberia. if we look at a terrestrial globe. 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. since rendered impassable by cold. And this continuity of the circumpolar land. I am strongly inclined to extend the above view.

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

at points 900 miles apart. glaciers have left the marks of their former low descent. W.B. from the western shores of Britain to the Ural range. I am convinced that Forbes's view may be largely extended. including grooved pebbles. as far south as latitude 36 and 37 degrees. Dr. from latitude 41 degrees to the southernmost extremity. where the climate is now so different. Hector. we have the clearest evidence of former glacial action. 386 . From facts communicated to me by the Rev. also. Clarke. nearly under the equator. Further south. perpetual snow formerly covered the central axis. according to Dr. Haast and Dr. Along the Himalaya. Hooker. at about the height of 12. crossing the Portillo valley. glaciers once extended far below their present level. and in Sikkim. as far south as latitude 46 degrees. it appears also that there are traces of former glacial action on the mountains of the south.000 feet down the valleys. Along this whole space of the Cordillera true glaciers do not now exist even at much more considerable heights. In central Chile I examined a vast mound of detritus with great boulders. ice-borne fragments of rock have been observed on the eastern side of the continent. In the Lebanon. Alternate Glacial Periods In The North And South But we must return to our more immediate subject. Southward of the Asiatic continent. we know. deeply-furrowed rocks. from the excellent researches of Dr. J. been noticed on the Rocky Mountains. and the same plants. which. Forbes informs me that he found in various parts of the Cordillera. in numerous immense boulders transported far from their parent source. resembling those with which he was familiar in Norway. found by Dr. Hooker on widely separated mountains in this island tell the same story of a former cold period. and fed glaciers which rolled 4.eastern corner of Australia. and Mr. that Siberia was similarly affected. on both sides of the continent.5. from latitude 13 to 30 degrees south. Looking to America: in the northern half. on the opposite side of the equator. We may infer from the frozen mammals and nature of the mountain vegetation. D. and likewise great masses of detritus. that in New Zealand immense glaciers formerly descended to a low level. there can hardly be a doubt. Hooker saw maize growing on ancient and gigantic moraines. Erratic boulders have. once formed a huge moraine. In Europe we meet with the plainest evidence of the Glacial period. and on the shores of the Pacific.000 feet. In the Cordillera of South America. and southward to the Pyrenees. The same observer has recently found great moraines at a low level on the Atlas range in North Africa.

Mr. On the Organ Mountains of Brazil some few temperate European. cold periods regularly recur every ten or fifteen thousand years. All these causes tend towards the same end. are common to North America and Europe. Dr. is the relative position of the land and water. brought into operation by an increase in the eccentricity of the earth's orbit. while the southern passes through a glacial period. not to mention still more ancient formations. from glaciers having recently descended to a low level along the whole line of the Cordillera. with slight alterations of climate. Croll. In South America. arrived at by Mr. 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. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur.000 years ago. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it. Croll believes that the last great glacial period occurred about 240. Lyell has shown.From these several facts. Hooker has shown that besides many closely allied species. With respect to more ancient glacial periods. forming no inconsiderable part of its scanty flora. 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. and these at long intervals are extremely severe. namely. So conversely it will be with the northern hemisphere. Mr. from direct evidence. According to Mr. with the winters rendered much milder. chiefly through changes in the direction of the ocean currents. that such occurred during the miocene and eocene formations. owing to certain contingencies. But now. Croll. enormously remote as these areas in opposite hemispheres are from each other. for about 160.000 years. but I will first give the facts which demand an explanation. but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. and endured. as Sir C. Croll. has attempted to show that a glacial condition of climate is the result of various physical causes. is that whenever the northern hemisphere passes through a cold period the temperature of the southern hemisphere is actually raised. But the most important result for us. between forty and fifty of the flowering plants of Tierra del Fuego. some Antarctic and some Andean genera were found by Gardner which do not exist in the low intervening hot countries. On the Silla of Caraccas 387 . as may be inferred from the amount of work effected—and lastly. in a series of admirable memoirs. several geologists are convinced. of which the most important.

on the heights of Ceylon. believed not to have been introduced by man." by Dr. In the admirable "Introduction to the Flora of New Zealand. This extension of the same temperate forms. On the Himalaya. Some of these Australian forms. occur on the mountains of Abyssinia. and on the other hand as far north as Japan. as I hear from Dr. other species. occur on the lowlands. but not in the intermediate torrid regions. however. Hooker has also lately shown that several of the plants living on the upper parts of the lofty island of Fernando Po. and likewise to those of temperate Europe. In Africa. Hooker. On the southern mountains of Australia. many plants occur either identically the same or representing each other. It now also appears. and some few representatives of the flora of the Cape of Good Hope. F. Lowe on the mountains of the Cape Verde Islands. almost under the equator. across the whole continent of Africa and to the mountains of the Cape Verde archipelago. is one of the most astonishing facts ever recorded in the distribution of plants.T. extend along the heights of the peninsula of Malacca. that some of these same temperate plants have been discovered by the Rev. as I hear from Dr. of European genera. Hooker. and are thinly scattered on the one hand over India. Hooker. we see that certain plants growing on the more lofty mountains of the tropics in all parts of the world. Hence. not introduced by man. Hooker. Still more striking is the fact that peculiar Australian forms are represented by certain plants growing on the summits of the mountains of Borneo. analogous and striking facts are given in regard to the plants of that large island. and at the same time representing plants of Europe not found in the intervening hot lowlands. several forms characteristic of Europe. be observed that 388 . Muller has discovered several European species. and on the temperate plains of the north and south. and on the mountains several representative European forms are found which have not been discovered in the intertropical parts of Africa. It should. and on the volcanic cones of Java. and a long list can be given. in the Gulf of Guinea. as I am informed by Dr. Dr. are closely related to those on the mountains of Abyssinia. R. found in Australia. are either the same species or varieties of the same species. At the Cape of Good Hope a very few European species. raises a picture of a collection made on a hillock in Europe. A list of the genera of plants collected on the loftier peaks of Java.the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera. and on the neighbouring Cameroon Mountains. and on the isolated mountain ranges of the peninsula of India. Dr.

of northern forms of fish.. In marine productions. as an example. Dana. speaks of the reappearance on the shores of New Zealand.C. During this. Croll's conclusion that when the northern hemisphere suffered from the extreme cold of the great Glacial period. I may quote a statement by the highest authority. Prof. So again in the mountainous island of Fernando Po. These brief remarks apply to plants alone. its antipode. the lowlands under the equator must have been clothed with a mingled tropical and temperate vegetation. the southern hemisphere was actually warmer. etc." Besides these identical and closely allied forms. to Ceylon and the Malay Archipelago. that "it is certainly a wonderful fact that New Zealand should have a closer resemblance in its crustacea to Great Britain. or perhaps even rather cooler. Dr." Sir J.these plants are not strictly arctic forms. for. the lowlands of these great continents were everywhere tenanted under the equator by a considerable number of temperate forms. "in receding from polar toward equatorial latitudes. Seemann found the vegetation like that of Mexico. Richardson. it appears almost certain that at some former period." Now let us see whether Mr. On the mountains of Panama. and along the Peninsula of India. at the height of only two thousand feet. "with forms of the torrid zone harmoniously blended with those of the temperate. but with perhaps a still greater preponderance of temperate forms. the Alpine or mountain flora really become less and less Arctic. 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. in the Gulf of Guinea. and in a less wellmarked manner across the wide expanse of tropical South America. also. as Mr. but have not been found in the intermediate tropical seas. the presence of temperate forms on the highlands across the whole of equatorial Africa. similar cases likewise occur. 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. Dr. Mann found temperate European forms beginning to appear at the height of about five thousand feet. belong to genera not now found in the intermediate tropical lowlands. H. namely. throws any clear light on 389 . no doubt during the most severe part of a Glacial period. Watson has remarked. many species inhabiting the same widely sundered areas. than to any other part of the world. but some few analogous facts could be given in regard to terrestrial animals. From the foregoing facts. Hooker informs me that twenty-five species of Algae are common to New Zealand and to Europe. the coldest period. Tasmania. Mr.

They might have survived. These latter. 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. But the species left during a long time on these mountains. would return to their former homes. for the change of temperature must have been very slow. the northern temperate forms living on the lowlands under the equator. as measured by years. The Glacial period. if sufficiently lofty. with the northern hemisphere rendered warmer. The northern forms which had before been left on the mountains would now descend and mingle with the southern forms. Thus. hence. The inhabitants of these hot lowlands would at the same time have migrated to the tropical and subtropical regions of the south. this period will have been ample for any amount of migration. would have to compete with many new forms and would be exposed to somewhat different physical conditions. for the southern hemisphere was at this period warmer. In the regular course of events the southern hemisphere would in its turn be subjected to a severe Glacial period. they would have long survived like the Arctic forms on the mountains of Europe. as both hemispheres gradually recovered their former temperature. dominant and widest-spreading temperate forms invaded the equatorial lowlands. and would generally 390 . and then the southern temperate forms would invade the equatorial lowlands. and from the facts just given. leaving some few species on the mountains. we know that Arctic forms invaded the temperate regions. and plants undoubtedly possess a certain capacity for acclimatisation. there can hardly be a doubt that some of the more vigorous. of the northern temperate forms would almost certainly have ascended any adjoining high land. and on the mountains of the tropics. however. when the warmth returned. As the cold became more and more intense. would have been driven to their former homes or have been destroyed. even if the climate was not perfectly fitted for them. being replaced by the equatorial forms returning from the south. and when we remember over what vast spaces some naturalised plants and animals have spread within a few centuries. they would be eminently liable to modification. as shown by their transmitting to their offspring different constitutional powers of resisting heat and cold.the present apparently inexplicable distribution of various organisms in the temperate parts of both hemispheres. and carrying southward with them some of the northern temperate forms which had descended from their mountain fastnesses. or in opposite hemispheres. On the decline of the Glacial period. Some. where. must have been very long.

wool. or dominating power. for the many quite distinct species inhabiting the same widely separated areas. but these have almost everywhere yielded to the more dominant forms generated in the larger areas and more efficient workshops of the north. or even outnumbered. In many islands the native productions are nearly equalled. than the southern forms. also. at the present day. Hooker. offer a partial exception. but not so the southern in regard to the northern forms. It is a remarkable fact. that many more identical or slightly modified species have migrated from the north to the south. no doubt the intertropical mountains were stocked with endemic Alpine forms. just in the same way as the inhabitants of real islands have everywhere 391 . New Zealand. And thus. and by Alph. We must. however. and having consequently been advanced through natural selection and competition to a higher stage of perfection. as I hear from Dr. and belonging to genera not now found in the intermediate torrid zones. and this is the first stage towards their extinction. Before the last great Glacial period. strongly insisted on by Hooker in regard to America. during the alternations of the Glacial periods. de Candolle in regard to Australia. a few southern forms on the mountains of Borneo and Abyssinia. I suspect that this preponderant migration from the north to the south is due to the greater extent of land in the north. for here. by those which have become naturalised. when the two sets became commingled in the equatorial regions. though hides. we see that very many European productions cover the ground in La Plata.now exist as varieties or as representative species. Mountains are islands on the land. 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. for these will account. and this is the case. than in a reversed direction. the northern forms were the more powerful and were able to hold their places on the mountains. Australian forms are rapidly sowing themselves and becoming naturalised. in accordance with the same principles. The Neilgherrie Mountains in India. whereas extremely few southern forms have become naturalised in any part of the northern hemisphere. and to a lesser degree in Australia. and have beaten the natives. We see. and to the northern forms having existed in their own homes in greater numbers. and afterwards migrate southward with the southern forms. however. In the same manner. bear in mind the occurrence in both hemispheres of former Glacial periods. and their inhabitants have yielded to those produced within the larger areas of the north.

We cannot hope to explain such facts. according to Forbes. now covered with ice. before the commencement of the last Glacial period. in nearly the same manner as. are removed on the views above given. some of the inhabitants of the temperate seas might have reached the equator. Various special difficulties also remain to be solved. and on the intertropical mountains. 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. but icebergs. while others have remained unaltered. isolated spaces inhabited by Arctic productions exist to the present day in the deeper parts of the northern temperate seas. which. by keeping to the cooler currents. and is twice or thrice as common. New Zealand. The facts seem to indicate that distinct species belonging to the same genera have migrated in radiating lines from a common centre. The exact lines of migration cannot be indicated. during the height of the Glacial period. until we can say why one species and not another becomes naturalised by man's agency in a foreign land. as in the northern hemisphere. is a more remarkable case. of the same plants at points so enormously remote as Kerguelen Land. The existence at these and other distant points of the southern hemisphere. We cannot say why certain species and not others have migrated. Some of these species are so distinct. When. why one species ranges twice or thrice as far. may have been concerned in their dispersal. to a former and warmer period. though distinct. and sometimes on the intermediate mountain ranges. I am far from supposing that all the difficulties in regard to the distribution and affinities of the identical and allied species. as another species within their own homes. which now live so widely separated in the north and south. when the Antarctic lands. and I am inclined to look in the southern. and Fuegia. why certain species have been modified and have given rise to new forms. as suggested by Lyell. of these a few would perhaps at once be able to migrate southwards. of species. the ocean-currents were widely different to what they now are. belong to genera exclusively confined to the south. the occurrence. The same principles apply to the distribution of terrestrial animals and of marine productions. supported a highly peculiar and isolated 392 . for instance. in the northern and southern temperate zones. as shown by Dr. Hooker. 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.yielded and are still yielding to continental forms naturalised through man's agency.

so have the living waters left their living drift on our mountain summits. driven up and surviving in the mountain fastnesses of almost every land. on the effects of great alternations of climate throughout the world on geographical distribution. in a line gently rising from the Arctic lowlands to a great latitude under the equator. Sir C. as halting-places. a few forms had been already widely dispersed to various points of the southern hemisphere by occasional means of transport. As the tide leaves its drift in horizontal lines. It may be suspected that before this flora was exterminated during the last Glacial epoch. of the former inhabitants of the surrounding lowlands. Australia. rising higher on the shores where the tide rises highest. and in both cases have reached the equator. And we have now seen that Mr. of now sunken islands. and New Zealand may have become slightly tinted by the same peculiar forms of life. in language almost identical with mine. but the stream of life has flowed with greater force from the north than in the opposite direction. Thus the southern shores of America. The various beings thus left stranded may be compared with savage races of man. The living waters have flowed during one period from the north and during another from the south. full of interest to us. Croll's conclusion that successive Glacial periods in the one hemisphere coincide with warmer periods in the opposite hemisphere. together with the admission of the slow modification of species. and by the aid. Lyell in a striking passage has speculated. and has consequently more freely inundated the south. which serves as a record. explains a multitude of facts in the distribution of the same and of the allied forms of life in all parts of the globe. 393 .flora.

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. 394 .

compared with those of Britain. though separated by a distance of about 230 miles. could be given of this having occurred during 395 . Thus fishes still alive are not very rarely dropped at distant points by whirlwinds. for in two adjoining river systems some of the species may be the same and some wholly different. 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. This case. Instances. in most cases be explained by their having become fitted. On the same continent fresh-water fish often range widely.1. that they would never have extended to distant countries. I think. belonging to different classes. be mainly attributed to changes in the level of the land within the recent period. some of the most difficult to explain are presented by fish. Gunther has lately shown that the Galaxias attenuatus inhabits Tasmania. We can here consider only a few cases. and at the dissimilarity of the surrounding terrestrial beings. and as if capriciously. and liability to wide dispersal would follow from this capacity as an almost necessary consequence. Not only have many fresh-water species. however. it might have been thought that fresh-water productions would not have ranged widely within the same country.. also. of these. shells. It was formerly believed that the same fresh-water species never existed on two continents distant from each other. an enormous range. But Dr. It is probable that they are occasionally transported by what may be called accidental means. I well remember feeling much surprise at the similarity of the fresh-water insects. in a manner highly useful to them. New Zealand. and it is known that the ova retain their vitality for a considerable time after removal from the water. within their own countries. etc. causing rivers to flow into each other. the Falkland Islands and the mainland of South America. But the wide ranging power of fresh-water productions can. and as the sea is apparently a still more formidable barrier. Fresh-Water Productions As lakes and river-systems are separated from each other by barriers of land. or from stream to stream. But the case is exactly the reverse. This is a wonderful case. Their dispersal may. for short and frequent migrations from pond to pond. When first collecting in the fresh waters of Brazil. and probably indicates dispersal from an Antarctic centre during a former warm period. but allied species prevail in a remarkable manner throughout the world. however.

These just-hatched molluscs. would be sure to alight on a pool or rivulet. and in this length of time a duck or heron might fly at least six or seven hundred miles. though at a somewhat more advanced age they would voluntarily drop off. and allied species which. as well as the adults. and if blown across the sea to an oceanic island. and could. in removing a little duck-weed from one aquarium to another. and the ova. Moreover. Salt-water fish can with care be slowly accustomed to live in fresh water. are descended from a common parent. which are continuous and consequently must. it is probable. which I have observed—and many others no doubt will be discovered—throw some light on this subject. Their distribution at first perplexed me much. from an early period. and. without any change of level. or to any other distant point.floods. I have twice seen these little plants adhering to their backs. The wide differences of the fish on the opposite sides of most mountain-ranges. survived on the duck's feet. When ducks suddenly emerge from a pond covered with duck-weed. as their ova are not likely to be transported by birds. But two facts. Gunther has recently been led by several considerations to infer that with fishes the same forms have a long endurance. Dr. from twelve to twenty hours. and clung to them so firmly that when taken out of the water they could not be jarred off. so that a marine species belonging to a fresh-water group might travel far along the shores of the sea. and in such cases there will have been ample time for great geographical changes. where many ova of fresh-water shells were hatching. become adapted without much difficulty to the fresh waters of a distant land. and consequently time and means for much migration. have completely prevented the inosculation of the river systems on the two sides. that I have unintentionally stocked the one with fresh-water shells from the other. leads to the same conclusion. and I found that numbers of the extremely minute and just-hatched shells crawled on the feet. there is hardly a single group of which all the members are confined to fresh water. on our theory. are immediately killed by sea-water. according to Valenciennes. in damp air. Some fresh-water fish belong to very ancient forms. Some species of fresh-water shells have very wide ranges. though aquatic in their nature. I could not even understand how some naturalised species have spread rapidly throughout the same country. prevail throughout the world. and it has happened to me. and must have proceeded from a single source. But another agency is perhaps more effectual: I suspended the feet of a duck in an aquarium. Sir Charles Lyell informs me that a Dyticus has been caught with an Ancylus (a fresh-water shell like a 396 .

according to Alph. With respect to plants. on the edge of a little pond. in those large groups of terrestrial plants. and even marsh-species. pulling up and counting each plant as it grew. I do not believe that botanists are aware how charged the mud of ponds is with seeds: I have tried several little experiments. they would not be likely to alight on the surface of the sea. when rejected many hours afterwards in pellets or in the excrement. century after century. they then take flight and go to other waters. they would be sure to fly to their natural fresh-water haunts. or are blown across the sea. and are occasionally found on the most remote and barren islands of the open ocean. but will here give only the most striking case: I took in February three tablespoonfuls of mud from three different points. as if in consequence. a wide range. a Colymbetes. The same agency may have come into play with the eggs of some of the smaller fresh-water animals. both over continents and to the most remote oceanic islands.limpet) firmly adhering to it." 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. the 397 . this mud when dry weighed only 6 and 3/4 ounces. have gone on daily devouring fish. beneath water. I have stated that fresh-water fish eat some kinds of seeds. and were altogether 537 in number. it has long been known what enormous ranges many fresh-water. once flew on board the "Beagle. 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. would be the most likely to have muddy feet. situated at very distant points. Other and unknown agencies probably have also played a part. Herons and other birds. though they reject many other kinds after having swallowed them. so that any dirt on their feet would not be washed off. if suddenly flushed. and we have seen that seeds retain their power of germination. When I saw the great size of the seeds of that fine water-lily. which have very few aquatic members. which frequent the muddy edges of ponds. as of the yellow water-lily and Potamogeton. and yet the viscid mud was all contained in a breakfast cup! Considering these facts. the plants were of many kinds. and a water-beetle of the same family. I have before mentioned that earth occasionally adheres in some quantity to the feet and beaks of birds. even small fish swallow seeds of moderate size. de Candolle. I think favourable means of dispersal explain this fact. Birds of this order wander more than those of any other. This is strikingly illustrated. I kept it covered up in my study for six months. have. and when gaining the land. for the latter seem immediately to acquire. Wading birds.

Hooker. however few in kind. the Nelumbium luteum) in a heron's stomach. We should not forget the probability of many fresh-water forms having formerly ranged continuously over immense areas. more especially by fresh-water birds. But the wide distribution of fresh-water plants. for instance on a rising islet. Although there will always be a struggle for life between the inhabitants of the same pond. and. it will be unoccupied. de Candolle's remarks on the distribution of this plant. but Audubon states that he found the seeds of the great southern water-lily (probably according to Dr. 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. 398 . then getting a hearty meal of fish. apparently depends in main part on the wide dispersal of their seeds and eggs by animals. whether retaining the same identical form.Nelumbium. We should also remember that many fresh-water productions are low in the scale of nature. I thought that the means of its dispersal must remain inexplicable. and a single seed or egg will have a good chance of succeeding. and this will give time for the migration of aquatic species. and naturally travel from one piece of water to another. In considering these several means of distribution. analogy makes me believe that it would have rejected the seeds in the pellet in a fit state for germination. than in the case of terrestrial colonists. and then having become extinct at intermediate points. Now this bird must often have flown with its stomach thus well stocked to distant ponds. the competition between them will probably be less severe than between terrestrial species. and remembered Alph. which have great powers of flight. and of the lower animals. it should be remembered that when a pond or stream is first formed. and we have reason to believe that such beings become modified more slowly than the high. or in some degree modified. consequently an intruder from the waters of a foreign country would have a better chance of seizing on a new place.

and the comparison in some other respects is not quite fair. This view removes many difficulties. for man has unintentionally stocked them far more fully and perfectly than did nature.2. but a few ferns and a few introduced plants are included in these numbers. but shall consider some other cases bearing on the truth of the two theories of independent creation and of descent with modification. together with the outlying islands of Auckland. with its lofty mountains and diversified stations. extending over 780 miles of latitude. yet many species have now become naturalised on it. the birthplace of their early progenitors. New Zealand. but it does not accord with all the facts in regard to the productions of islands. On The Inhabitants Of Oceanic Islands We now come to the last of the three classes of facts. 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. although now inhabiting the most distant points. contain altogether only 960 kinds of flowering plants. We have evidence that the barren island of Ascension aboriginally possessed less than half-a-dozen flowering plants. for instance. In the following remarks I shall not confine myself to the mere question of dispersal. which I have selected as presenting the greatest amount of difficulty with respect to distribution. independently of different physical conditions. The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. but that allied species. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions. we must admit that some cause. 399 . de Candolle admits this for plants. if we compare this moderate number with the species which swarm over equal areas in Southwestern Australia or at the Cape of Good Hope. has given rise to so great a difference in number. on the view that not only all the individuals of the same species have migrated from some one area. have proceeded from a single area. Campbell and Chatham. as they have in New Zealand and on every other oceanic island which can be named. and the little island of Anglesea 764. In St. and Wollaston for insects. will have to admit that a sufficient number of the best adapted plants and animals were not created for oceanic islands. He who admits the doctrine of the creation of each separate species. Even the uniform county of Cambridge has 847 plants.

Bermuda. for instance. or of another section of the same class. after long intervals of time in the new and isolated district. though very closely related to a European form. does not possess a single endemic land bird. when settled in their new homes.e. so that their mutual relations have not been much disturbed. If we compare. are peculiar. this island is inhabited by ninetynine kinds. whereas of the eleven marine birds only two are peculiar. species occasionally arriving. and it is obvious that marine birds could arrive at these islands much more easily and frequently than land-birds. I will give a few illustrations of the foregoing remarks: in the Galapagos Islands there are twenty-six land birds. which for long ages have there struggled together. Any tendency to 400 .Although in oceanic islands the species are few in number. It should be borne in mind that the offspring of such crosses would certainly gain in vigour. as already explained. with the number found on any continent. each kind will have been kept by the others to its proper place and habits. Jones's admirable account of Bermuda. J. would be eminently liable to modification. and which has a very peculiar soil. or of endemic birds in the Galapagos Archipelago. and this difference seems to depend partly on the species which are not modified having immigrated in a body. which lies at about the same distance from North America as the Galapagos Islands do from South America. and having to compete with new associates.V. as I am informed by Mr. we shall see that this is true. the number of endemic land-shells in Madeira. Almost every year. those found nowhere else in the world) is often extremely large. Hence. Harcourt. of which one alone is peculiar. So that the islands of Bermuda and Madeira have been stocked from the neighbouring continents with birds. so that even an occasional cross would produce more effect than might have been anticipated. But it by no means follows that. and we know from Mr. and then compare the area of the island with that of the continent. E. and would often produce groups of modified descendants. those of another class. because in an island nearly all the species of one class are peculiar. and partly on the frequent arrival of unmodified immigrants from the mother-country. of these twenty-one (or perhaps twentythree) are peculiar.M. on the other hand. many European and African birds are blown to Madeira. for. that very many North American birds occasionally or even frequently visit this island. with which the insular forms have intercrossed. the proportion of endemic kinds (i. and three or four other species are confined to this island and to the Canaries. and have become mutually co-adapted. and will consequently have been but little liable to modification. This fact might have been theoretically expected.

yet we can see that their eggs or larvae. the place of mammals. Hence trees would be little likely to reach distant oceanic islands. as Alph. Turning to plants. but this explanation is not a little doubtful. and their places are occupied by other classes. The different orders of insects inhabiting Madeira present nearly parallel cases. Madeira again is inhabited by a wonderful number of peculiar land-shells. Although New Zealand is here spoken of as an oceanic island. from its geological character and the direction of its mountain ranges. in certain islands not tenanted by a single mammal. should be considered as appurtenances of Australia. as well as New Caledonia. confined ranges. But a hooked seed might be carried to an island by other means. often arriving from the mother-country. islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species.modification will also have been checked by intercrossing with the unmodified immigrants. or recently took. and is not separated from Australia by a profoundly deep sea. whereas not one species of sea-shell is peculiar to its shores: now. it is in some degree doubtful whether it should be so ranked. de Candolle has shown. which had no chance of successfully competing with the many 401 . W. it is of large size. Oceanic islands are sometimes deficient in animals of certain whole classes. All such differences in number.B. whatever the cause may be. Many remarkable little facts could be given with respect to the inhabitants of oceanic islands. and an herbaceous plant. Again. For instance. are generally accounted for by supposed differences in the physical conditions of the islands. now trees. and in New Zealand gigantic wingless birds. yet few relations are more manifest than that hooks serve for the transportal of seeds in the wool or fur of quadrupeds. though we do not know how sea-shells are dispersed. Clarke has lately maintained that this island. some of the endemic plants have beautifully hooked seeds. perhaps attached to seaweed or floating timber. Facility of immigration seems to have been fully as important as the nature of the conditions. and the absence of certain whole groups of animals and plants. might be transported across three or four hundred miles of open sea far more easily than land-shells. take. thus in the Galapagos Islands reptiles. and the plant then becoming modified would form an endemic species. still retaining its hooks. which would form a useless appendage. Hooker has shown that in the Galapagos Islands the proportional numbers of the different orders are very different from what they are elsewhere. or to the feet of wading birds. generally have. like the shrivelled wings under the soldered wing-covers of many insular beetles. Dr. the Rev.

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


3. Absence Of Batrachians And Terrestrial Mammals On Oceanic Islands
With respect to the absence of whole orders of animals on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts) are never found on any of the many islands with which the great oceans are studded. I have taken pains to verify this assertion, and have found it true, with the exception of New Zealand, New Caledonia, the Andaman Islands, and perhaps the Solomon Islands and the Seychelles. But I have already remarked that it is doubtful whether New Zealand and New Caledonia ought to be classed as oceanic islands; and this is still more doubtful with respect to the Andaman and Solomon groups and the Seychelles. This general absence of frogs, toads and newts on so many true oceanic islands cannot be accounted for by their physical conditions; indeed it seems that islands are peculiarly fitted for these animals; for frogs have been introduced into Madeira, the Azores, and Mauritius, and have multiplied so as to become a nuisance. But as these animals and their spawn are immediately killed (with the exception, as far as known, of one Indian species) by sea-water, there would be great difficulty in their transportal across the sea, and therefore we can see why they do not exist on strictly oceanic islands. But why, on the theory of creation, they should not have been created there, it would be very difficult to explain. Mammals offer another and similar case. I have carefully searched the oldest voyages, and have not found a single instance, free from doubt, of a terrestrial mammal (excluding domesticated animals kept by the natives) inhabiting an island situated above 300 miles from a continent or great continental island; and many islands situated at a much less distance are equally barren. The Falkland Islands, which are inhabited by a wolf-like fox, come nearest to an exception; but this group cannot be considered as oceanic, as it lies on a bank in connection with the mainland at a distance of about 280 miles; moreover, icebergs formerly brought boulders to its western shores, and they may have formerly transported foxes, as now frequently happens in the arctic regions. Yet it cannot be said that small islands will not support at least small mammals, for they occur in many parts of the world on very small islands, when lying close to a continent; and hardly an island can be named on which our smaller quadrupeds have not become naturalised and greatly multiplied. It cannot be said, on the ordinary view of creation, that there has not been time for the creation of mammals; many volcanic islands are sufficiently


ancient, as shown by the stupendous degradation which they have suffered, and by their tertiary strata: there has also been time for the production of endemic species belonging to other classes; and on continents it is known that new species of mammals appear and disappear at a quicker rate than other and lower animals. Although terrestrial mammals do not occur on oceanic islands, aerial mammals do occur on almost every island. New Zealand possesses two bats found nowhere else in the world: Norfolk Island, the Viti Archipelago, the Bonin Islands, the Caroline and Marianne Archipelagoes, and Mauritius, all possess their peculiar bats. Why, it may be asked, has the supposed creative force produced bats and no other mammals on remote islands? On my view this question can easily be answered; for no terrestrial mammal can be transported across a wide space of sea, but bats can fly across. Bats have been seen wandering by day far over the Atlantic Ocean; and two North American species, either regularly or occasionally, visit Bermuda, at the distance of 600 miles from the mainland. I hear from Mr. Tomes, who has specially studied this family, that many species have enormous ranges, and are found on continents and on far distant islands. Hence, we have only to suppose that such wandering species have been modified in their new homes in relation to their new position, and we can understand the presence of endemic bats on oceanic islands, with the absence of all other terrestrial mammals. Another interesting relation exists, namely, between the depth of the sea separating islands from each other, or from the nearest continent, and the degree of affinity of their mammalian inhabitants. Mr. Windsor Earl has made some striking observations on this head, since greatly extended by Mr. Wallace's admirable researches, in regard to the great Malay Archipelago, which is traversed near Celebes by a space of deep ocean, and this separates two widely distinct mammalian faunas. On either side, the islands stand on a moderately shallow submarine bank, and these islands are inhabited by the same or by closely allied quadrupeds. I have not as yet had time to follow up this subject in all quarters of the world; but as far as I have gone, the relation holds good. For instance, Britain is separated by a shallow channel from Europe, and the mammals are the same on both sides; and so it is with all the islands near the shores of Australia. The West Indian Islands, on the other hand, stand on a deeply submerged bank, nearly one thousand fathoms in depth, and here we find American forms, but the species and even the genera are quite distinct. As the amount of modification which animals of all kinds undergo partly depends on the lapse of time, and as the islands which


are separated from each other, or from the mainland, by shallow channels, are more likely to have been continuously united within a recent period than the islands separated by deeper channels, we can understand how it is that a relation exists between the depth of the sea separating two mammalian faunas, and the degree of their affinity, a relation which is quite inexplicable on the theory of independent acts of creation. The foregoing statements in regard to the inhabitants of oceanic islands, namely, the fewness of the species, with a large proportion consisting of endemic forms—the members of certain groups, but not those of other groups in the same class, having been modified—the absence of certain whole orders, as of batrachians and of terrestrial mammals, notwithstanding the presence of aerial bats, the singular proportions of certain orders of plants, herbaceous forms having been developed into trees, etc., seem to me to accord better with the belief in the efficiency of occasional means of transport, carried on during a long course of time, than with the belief in the former connection of all oceanic islands with the nearest continent; for on this latter view it is probable that the various classes would have immigrated more uniformly, and from the species having entered in a body, their mutual relations would not have been much disturbed, and consequently, they would either have not been modified, or all the species in a more equable manner. I do not deny that there are many and serious difficulties in understanding how many of the inhabitants of the more remote islands, whether still retaining the same specific form or subsequently modified, have reached their present homes. But the probability of other islands having once existed as halting-places, of which not a wreck now remains, must not be overlooked. I will specify one difficult case. Almost all oceanic islands, even the most isolated and smallest, are inhabited by land-shells, generally by endemic species, but sometimes by species found elsewhere striking instances of which have been given by Dr. A.A. Gould in relation to the Pacific. Now it is notorious that land-shells are easily killed by sea-water; their eggs, at least such as I have tried, sink in it and are killed. Yet there must be some unknown, but occasionally efficient means for their transportal. Would the just-hatched young sometimes adhere to the feet of birds roosting on the ground and thus get transported? It occurred to me that land-shells, when hybernating and having a membranous diaphragm over the mouth of the shell, might be floated in chinks of drifted timber across moderately wide arms of the sea. And I find that several species in this state withstand uninjured an immersion in sea-water during seven days. One shell, the Helix pomatia,


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


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


nearest continent, or of the nearest large island. The exceptions are few, and most of them can be explained. Thus, although Kerguelen Land stands nearer to Africa than to America, the plants are related, and that very closely, as we know from Dr. Hooker's account, to those of America: but on the view that this island has been mainly stocked by seeds brought with earth and stones on icebergs, drifted by the prevailing currents, this anomaly disappears. New Zealand in its endemic plants is much more closely related to Australia, the nearest mainland, than to any other region: and this is what might have been expected; but it is also plainly related to South America, which, although the next nearest continent, is so enormously remote, that the fact becomes an anomaly. But this difficulty partially disappears on the view that New Zealand, South America, and the other southern lands, have been stocked in part from a nearly intermediate though distant point, namely, from the antarctic islands, when they were clothed with vegetation, during a warmer tertiary period, before the commencement of the last Glacial period. The affinity, which, though feeble, I am assured by Dr. Hooker is real, between the flora of the south-western corner of Australia and of the Cape of Good Hope, is a far more remarkable case; but this affinity is confined to the plants, and will, no doubt, some day be explained. The same law which has determined the relationship between the inhabitants of islands and the nearest mainland, is sometimes displayed on a small scale, but in a most interesting manner, within the limits of the same archipelago. Thus each separate island of the Galapagos Archipelago is tenanted, and the fact is a marvellous one, by many distinct species; but these species are related to each other in a very much closer manner than to the inhabitants of the American continent, or of any other quarter of the world. This is what might have been expected, for islands situated so near to each other would almost necessarily receive immigrants from the same original source, and from each other. But how is it that many of the immigrants have been differently modified, though only in a small degree, in islands situated within sight of each other, having the same geological nature, the same height, climate, etc? This long appeared to me a great difficulty: but it arises in chief part from the deeply-seated error of considering the physical conditions of a country as the most important; whereas it cannot be disputed that the nature of the other species with which each has to compete, is at least as important, and generally a far more important element of success. Now if we look to the species which inhabit the Galapagos Archipelago, and are likewise found in other parts of the world, we find that they differ


considerably in the several islands. This difference might indeed have been expected if the islands have been stocked by occasional means of transport—a seed, for instance, of one plant having been brought to one island, and that of another plant to another island, though all proceeding from the same general source. Hence, when in former times an immigrant first settled on one of the islands, or when it subsequently spread from one to another, it would undoubtedly be exposed to different conditions in the different islands, for it would have to compete with a different set of organisms; a plant, for instance, would find the ground 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


large proportion of cases, as shown by Alph. de Candolle, to distinct genera. In the Galapagos Archipelago, many even of the birds, though so well adapted for flying from island to island, differ on the different islands; thus there are three closely allied species of mocking-thrush, each confined to its own island. Now let us suppose the mocking-thrush of Chatham Island to be blown to Charles Island, which has its own mocking-thrush; why should it succeed in establishing itself there? We may safely infer that Charles Island is well stocked with its own species, for annually more eggs are laid and young birds hatched than can possibly be reared; and we may infer that the mocking-thrush peculiar to Charles Island is at least as well fitted for its home as is the species peculiar to Chatham Island. Sir C. Lyell and Mr. Wollaston have communicated to me a remarkable fact bearing on this subject; namely, that Madeira and the adjoining islet of Porto Santo possess many distinct but representative species of land-shells, some of which live in crevices of stone; and although large quantities of stone are annually transported from Porto Santo to Madeira, yet this latter island has not become colonised by the Porto Santo species: nevertheless, both islands have been colonised by some European land-shells, which no doubt had some advantage over the indigenous species. From these considerations I think we need not greatly marvel at the endemic species which inhabit the several islands of the Galapagos Archipelago not having all spread from island to island. On the same continent, also, pre-occupation has probably played an important part in checking the commingling of the species which inhabit different districts with nearly the same physical conditions. Thus, the south-east and south-west corners of Australia have nearly the same physical conditions, and are united by continuous land, yet they are inhabited by a vast number of distinct mammals, birds, and plants; so it is, according to Mr. Bates, with the butterflies and other animals inhabiting the great, open, and continuous valley of the Amazons. The same principle which governs the general character of the inhabitants of oceanic islands, namely, the relation to the source whence colonists could have been most easily derived, together with their subsequent modification, is of the widest application throughout nature. We see this on every mountain-summit, in every lake and marsh. For Alpine species, excepting in as far as the same species have become widely spread during the Glacial epoch, are related to those of the surrounding lowlands; thus we have in South America, Alpine humming-birds, Alpine rodents, Alpine plants, etc., all strictly belonging to American forms; and it is obvious that a mountain, as it became slowly upheaved, would be


colonised from the surrounding lowlands. So it is with the inhabitants of lakes and marshes, excepting in so far as great facility of transport has allowed the same forms to prevail throughout large portions of the world. We see the same principle in the character of most of the blind animals inhabiting the caves of America and of Europe. Other analogous facts could be given. It will, I believe, be found universally true, that wherever in two regions, let them be ever so distant, many closely allied or representative species occur, there will likewise be found some identical species; and wherever many closely-allied species occur, there will be found many forms which some naturalists rank as distinct species, and others as mere varieties; these doubtful forms showing us the steps in the process of modification. The relation between the power and extent of migration in certain species, either at the present or at some former period, and the existence at remote points of the world of closely allied species, is shown in another and more general way. Mr. Gould remarked to me long ago, that in those genera of birds which range over the world, many of the species have very wide ranges. I can hardly doubt that this rule is generally true, though difficult of proof. Among mammals, we see it strikingly displayed in Bats, and in a lesser degree in the Felidae and Canidae. We see the same rule in the distribution of butterflies and beetles. So it is with most of the inhabitants of fresh water, for many of the genera in the most distinct classes range over the world, and many of the species have enormous ranges. It is not meant that all, but that some of the species have very wide ranges in the genera which range very widely. Nor is it meant that the species in such genera have, on an average, a very wide range; for this will largely depend on how far the process of modification has gone; for instance, two varieties of the same species inhabit America and Europe, and thus the species has an immense range; but, if variation were to be carried a little further, the two varieties would be ranked as distinct species, and their range would be greatly reduced. Still less is it meant, that species which have the capacity of crossing barriers and ranging widely, as in the case of certain powerfully-winged birds, will necessarily range widely; for we should never forget that to range widely implies not only the power of crossing barriers, but the more important power of being victorious in distant lands in the struggle for life with foreign associates. But according to the view that all the species of a genus, though distributed to the most remote points of the world, are descended from a single progenitor, we ought to find, and I believe as a


that some at least of the species range very widely. but are explicable if we admit colonisation from the nearest or readiest source. consequently they will have had a better chance of ranging widely and of still retaining the same specific character. as alpine. probably accounts for a law which has long been observed. lacustrine. lower organisms ranging more widely than the higher—some of the species of widely-ranging genera themselves ranging widely—such facts.general rule we do find. from geological evidence. together with the subsequent adaptation of the colonists to their new homes. namely. 412 . together with that of the seeds and eggs of most lowly organised forms being very minute and better fitted for distant transportal. and the species in this case will have had ample time for dispersal and subsequent modification. There is also reason to believe. This fact. de Candolle in regard to plants. We should bear in mind that many genera in all classes are of ancient origin. 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. and which has lately been discussed by Alph. that the lower any group of organisms stands the more widely it ranges. The relations just discussed—namely. that within each great class the lower organisms change at a slower rate than the higher.

together with subsequent modification and the multiplication of new forms. though in this case. which has been arrived at by many naturalists under the designation of single centres of creation. and likewise of the several species belonging to the same genus. wherever found. more especially from the importance of barriers of all kinds. as in that of the individuals of the same species. enormous periods of time having been thus granted for their migration.5. whether of land or water. the difficulties are far from insuperable. As showing how diversified are the means of occasional transport. and left some of them stranded on the mountain-summits in all parts of the world. and remember that some forms of life have changed very slowly. and this is a very important consideration. and then have become extinct in the intermediate tracts—the difficulty is not insuperable in believing that all the individuals of the same species. If the difficulties be not insuperable in admitting that in the long course of time all the individuals of the same species. and which. in not only separating but in apparently forming the several zoological and 413 . by various general considerations. We can thus understand the high importance of barriers. With respect to distinct species belonging to the same genus. I have discussed at some little length the means of dispersal of fresh-water productions. And we are led to this conclusion. if we make the same allowances as before for our ignorance. which affected even the equatorial regions. 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. 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. they are often great. As exemplifying the effects of climatical changes on distribution. how often a species may have ranged continuously over a wide area. then all the grand leading facts of geographical distribution are explicable on the theory of migration. genera. which have certainly occurred within the recent period. and from the analogical distribution of subgenera. have proceeded from some one source. during the alternations of the cold in the north and the south. I have attempted to show how important a part the last Glacial period has played. allowed the productions of opposite hemispheres to mingle. and families. are descended from common parents. which on our theory have spread from one parent-source.

whilst the most isolated islands should possess their own peculiar species of aerial mammals or bats. and the depth of the sea between such islands and the mainland. We can thus understand the concentration of related species within the same areas. though specifically distinct on the several islets. according to the nature of the communication which allowed certain forms and not others to enter. one group of beings should have all its species peculiar. are linked together in so mysterious a manner. of the forests. there should be some relation between the presence of mammals. there would ensue in the to or more regions. there would be an almost endless amount of organic action and reaction. We can see why. should be absent from oceanic islands.botanical provinces. and we should find some groups of beings greatly. should have all its species the same with those in an adjoining quarter of the world. some developed in great force. as I have endeavoured to show. for according to the length of time which has elapsed since the colonists entered one of the regions. and should likewise be related. We can see why. or other source whence immigrants might have been derived. On these same principles we can understand. as batrachians and terrestrial mammals. but less closely. and why. We can clearly see why all the inhabitants of an archipelago. in a more or less modified condition. should be closely related to each other. or both. independently of their physical conditions. either in greater or lesser numbers. even within the same class. in relation to the means of migration. a large proportion should be endemic or peculiar. if there exist very closely allied or 414 . but that of these. some existing in scanty numbers—and this we do find in the several great geographical provinces of the world. the inhabitants of the plains and mountains. We can see why whole groups of organisms. infinitely diversified conditions of life. for instance. why oceanic islands should have few inhabitants. and another group. and some only slightly modified. should often be inhabited by very different forms of life. according or not as those which entered happened to come into more or less direct competition with each other and with the aborigines. we can see why two areas. to those of the nearest continent. having nearly the same physical conditions. and how it is that under different latitudes. and deserts. in islands. and according as the immigrants were capable of varying more or less rapidly. Bearing in mind that the mutual relation of organism to organism is of the highest importance. and are likewise linked to the extinct beings which formerly inhabited the same continent. marshes. in South America.

but there are in both cases marked exceptions to the rule. living during the same period of time. as of sculpture or colour. We see this in many facts. there is a striking parallelism in the laws of life throughout time and space. as I have attempted to show. in both cases the laws of variation have been the same. 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. for whether we look to the allied forms of life which have changed during successive ages. some identical species will almost always there be found. or only in a different section of the same order. it certainly is the general rule that the area inhabited by a single species. and modifications have been accumulated by the same means of natural selection. or by a group of species. As the late Edward Forbes often insisted. The endurance of each species and group of species is continuous in time. According to our theory. differ greatly from each other. is continuous. however distant from each other. as in looking to distant provinces throughout the world. or by the species having become extinct in the intermediate tracts. Both in time and space species and groups of species have their points of maximum development. but which occur above and below: so in space. which are not rare. or through occasional means of transport. are often characterised by trifling features in common. whilst those in another class. and the exceptions. Groups of species. 415 . 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. may. In looking to the long succession of past ages. in both cases they are connected by the same bond of ordinary generation. we find that species in certain classes differ little from each other.representative species in two areas. or to those which have changed after having migrated into distant quarters. In both time and space the lowly organised members of each class generally change less than the highly organised. or living within the same area. these several relations throughout time and space are intelligible. be accounted for by former migrations under different circumstances.

between parts of the same individual — Embryology.Chapter 14 Mutual Affinities Of Organic Beings: Morphology -Embryology -. laws of. and being inherited at a corresponding age — Rudimentary organs. their origin explained — Summary.Rudimentary Organs Classification. general. groups subordinate to groups — Natural system — Rules and difficulties in classification. explained on the theory of descent with modification — Classification of varieties — Descent always used in classification — Analogical or adaptive characters — Affinities. 416 . explained by variations not supervening at an early age. complex and radiating — Extinction separates and defines groups — Morphology. between members of the same class.

belonging to the larger genera in each class. one to feed on flesh. 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. to supplant and exterminate the preceding. distinct from that containing the next two genera on the right hand. and. This classification is not arbitrary like the grouping of the stars in constellations. much in common. and form a subfamily. and so on. Consequently the groups which are now large. In the second and fourth chapters.1. and these. or incipient species. and which generally include many dominant species. I have attempted to show that within each country it is the widely ranging. come into the closest competition. and he will see that the inevitable result is. I request the reader to turn to the diagram illustrating the action. that is the dominant species. which. tend to produce other new and dominant species. on the principle of inheritance. ultimately become converted into new and distinct species. as formerly explained. for all are descended from one ancient parent. which vary most. though 417 . and another the water. In the diagram each letter on the uppermost line may represent a genus including several species. consequently. and the whole of the genera along this upper line form together one class. But the three genera on the left hand have. which diverged from a common parent at the fifth stage of descent. but the case is widely different. they constantly tend to diverge in character. in any small area. another on vegetable matter. The varieties. The existence of groups would have been of simple significance. thus produced. have inherited something in common. and by certain facts in naturalisation. This latter conclusion is supported by observing the great diversity of forms. for it is notorious how commonly members of even the same subgroup have different habits. so that they can be classed in groups under groups. that the modified descendants proceeding from one progenitor become broken up into groups subordinate to groups. if one group had been exclusively fitted to inhabit the land. on this same principle. tend to go on increasing in size. of these several principles. the much diffused and common. These five genera have also much in common. on Variation and on Natural Selection. 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. I attempted also to show that there is a steady tendency in the forms which are increasing in number and diverging in character.

for instance. descended from (A). on what is called the Natural System. and for separating those which are most unlike. or as an artificial method of enunciating. it seems to me that nothing is thus added to our knowledge. But many naturalists think that something more is meant by the Natural System. that the characters do not make the genus. which diverged at an earlier period. and the genera into subfamilies. genera and families in each class. is in my judgment thus explained. or what else is meant by the plan of the Creator. Expressions such as that famous one by Linnaeus.less than when grouped in subfamilies. or more naturally by a number of characters. We know. or both. from its familiarity. is partially revealed to us by our classifications. like all other objects. try to arrange the species. either artificially by single characters. and that community of descent—the one known cause of close similarity in organic beings—is the bond. by one sentence to give the characters common. but unless it be specified whether order in time or space. which. does not always sufficiently strike us. can be classed in many ways. and then. and no cause can at present be assigned for their falling into groups. but that the genus gives the characters. namely. which. a full description is given of each kind of dog. Naturalists. 418 . and the view above given accords with their natural arrangement in group under group. which we often meet with in a more or less concealed form. as we have seen. No doubt organic beings. And all these genera. all under one great class. So that we here have many species descended from a single progenitor grouped into genera. general propositions—that is. families and orders. and no other explanation has ever been attempted. In this case there is of course no relation to genealogical succession. that minerals and the elemental substances can be thus arranged. But with organic beings the case is different. The ingenuity and utility of this system are indisputable. they believe that it reveals the plan of the Creator. seem to imply that some deeper bond is included in our classifications than mere resemblance. 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. for instance. I believe that this is the case. form an order distinct from the genera descended from (I). to all mammals. though observed by various degrees of modification. and they form a family distinct from that containing the three genera still further to the right hand. by another those common to all carnivora. The grand fact of the natural subordination of organic beings in groups under groups. as briefly as possible. by adding a single sentence. by another those common to the doggenus.

As an instance: Owen. its classificatory value is widely different. being those which are most remotely related to the habits and food of an animal. and it has been fully acknowledged in the writings of almost every author. It may even be given as a general rule. I have always regarded as affording very clear indications of its true affinities. 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. that in allied groups." With plants how remarkable it is that the organs of vegetation. That the mere physiological importance of an organ does not determine its classificatory value. "The generative organs. Robert Brown. is very unequal." Again. and in some cases seems to be entirely lost. not only in this. This depends on their constancy throughout many allied groups. are of little signification. in speaking of certain organs in the Proteaceae. as I apprehend in every natural family. the 419 . No naturalist can have worked at any group without being struck with this fact.Let us now consider the rules followed in classification. are ranked as merely "adaptive or analogical characters. It might have been thought (and was in ancient times thought) that those parts of the structure which determined the habits of life. or is simply a scheme for enunciating general propositions and of placing together the forms most like each other. in which the same organ. No one regards the external similarity of a mouse to a shrew. which acts only on serviceable characters. These resemblances. are of paramount importance! So again. is almost proved by the fact. as of any importance. "like that of all their parts. in speaking of the dugong. It will suffice to quote the highest authority. says. of a dugong to a whale. the more important it becomes for classification. and the general place of each being in the economy of nature. as we have every reason to suppose. but. in another work he says. whereas the organs of reproduction. with their product the seed and embryo. and their constancy chiefly depends on any slight deviations not having been preserved and accumulated by natural selection. has nearly the same physiological value. we have seen that they are often of the highest service in classification. of a whale to a fish. says their generic importance. and the difficulties which are encountered on the view that classification either gives some unknown plan of creation. on which their nutrition and life depend. would be of very high importance in classification. Nothing can be more false. who." but to the consideration of these resemblances we shall recur. that the less any part of the organisation is concerned with special habits. though so intimately connected with the whole life of the being.

" To give an example among insects: in one great division of the Hymenoptera. The importance. as hair or feathers. it has been found that a 420 . Robert Brown has strongly insisted on the fact that the position of the rudimentary florets is of the highest importance in the classification of the Grasses. organs in this condition are often of much value in classification. For instance. but which are universally admitted as highly serviceable in the definition of whole groups.genera of the Connaraceae "differ in having one or more ovaria. no one will say that rudimentary or atrophied organs are of high physiological or vital importance. of trifling characters. and certain rudimentary bones of the leg. whether or not there is an open passage from the nostrils to the mouth. Again. in the existence or absence of albumen. a species may depart from its allies in several characters. Numerous instances could be given of characters derived from parts which must be considered of very trifling physiological importance. Hence. yet. If the Ornithorhynchus had been covered with feathers instead of hair. as Westwood has remarked. Any one of these characters singly is frequently of more than generic importance. when all taken together. also. mainly depends on their being correlated with many other characters of more or less importan