On the Origin of Species, 6th Edition

Darwin, Charles

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


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"But with regard to the material world, we can at least go so far as this— we can perceive that events are brought about not by insulated interpositions of Divine power, exerted in each particular case, but by the establishment of general laws."—Whewell: "Bridgewater Treatise". "The only distinct meaning of the word 'natural' is STATED, FIXED or SETTLED; since what is natural as much requires and presupposes an intelligent agent to render it so, i.e., to effect it continually or at stated times, as what is supernatural or miraculous does to effect it for once."—Butler: "Analogy of Revealed Religion". "To conclude, therefore, let no man out of a weak conceit of sobriety, or an ill-applied moderation, think or maintain, that a man can search too far or be too well studied in the book of God's word, or in the book of God's works; divinity or philosophy; but rather let men endeavour an endless progress or proficience in both."—Bacon: "Advancement of Learning".


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 "Untersuchungen uber die Entwickelungs-Gesetze", it appears that the


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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Chapter 2 Variation Under Nature Variability — Individual differences — Doubtful species — Wide ranging. and common species. vary most — Species of the larger genera in each country vary more frequently than the species of the smaller genera — Many of the species of the larger genera resemble varieties in being very closely. but unequally. much diffused. related to each other. and in having restricted ranges. 49 .

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

68 .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.

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

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

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

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

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

It would suffice to keep up the full number of a tree. a small number may be produced. in all cases.number. 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. and the number of the species will almost instantaneously increase to any amount. the average number of any animal or plant depends only indirectly on the number of its eggs or seeds. Lighten any check. If an animal can in any way protect its own eggs or young. but if many eggs or young are destroyed. so that. 74 . many must be produced or the species will become extinct. which lived on an average for a thousand years. it is most necessary to keep the foregoing considerations always in mind—never to forget that every single organic being may be said to be striving to the utmost to increase in numbers. if 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. In looking at Nature. and yet the average stock be fully kept up. that each lives by a struggle at some period of its life. that heavy destruction inevitably falls either on the young or old during each generation or at recurrent intervals. mitigate the destruction ever so little.

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

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

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

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

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

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

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

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

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

Or The Survival Of The Fittest Natural Selection — its power compared with man's selection — its power on characters of trifling importance — its power at all ages and on both sexes — Sexual Selection — On the generality of intercrosses between individuals of the same species — Circumstances favourable and unfavourable to the results of Natural Selection. intercrossing. isolation. namely.Chapter 4 Natural Selection. on the descendants from a common parent — Explains the Grouping of all organic beings — Advance in organisation — Low forms preserved — Convergence of character — Indefinite multiplication of species — Summary. related to the diversity of inhabitants of any small area and to naturalisation — Action of Natural Selection. through Divergence of Character and Extinction. number of individuals — Slow action — Extinction caused by Natural Selection — Divergence of Character. 84 .

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

forming sub-families. If species had been independently created. and so on in almost endless cycles. when the tree was young. and these into lesser and lesser branches. the nature of the affinities. budding twigs. and those produced during former years may represent the long succession of extinct species. and widely ranging species. and these round other points. in the manner which we everywhere behold—namely. the more diversified the descendants become. The limbs divided into great branches. as we have seen illustrated in the diagram. leads to divergence of character and to much extinction of the less improved and intermediate forms of life.struggle of all species to increase in numbers. species of the same genus less closely and unequally related. and genera related in different degrees. The green and budding twigs may represent existing species. but it is explained through inheritance and the complex action of natural selection. varieties of the same species most closely related. 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. but seem clustered round points. and these tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. Thus the small differences distinguishing varieties of the same species. orders. entailing extinction and divergence of character. or even of distinct genera. sub-classes. On these principles. Natural selection. the widely diffused. in the same manner as species and groups of species have at all times overmastered other species in the great battle for life. and classes. steadily tend to increase. I believe this simile largely speaks the truth. At each period of growth all the growing twigs have tried to branch out on all sides. forming sections and sub-genera. as has just been remarked. and the generally well defined distinctions between the innumerable organic beings in each class throughout the world. families. which vary most. belonging to the larger genera within each class. were themselves once. and to overtop and kill the surrounding twigs and branches. subordinate to groups. 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. the better will be their chance of success in the battle for life. no explanation would have been possible of this kind of classification. till they equal the greater differences between species of the same genus. species of distinct genera much less closely related. We have seen that it is the common. may be explained. and this connexion of the former and present buds by ramifying 131 .

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

organs of flight and of vision — Acclimatisation — Correlated variation — Compensation and economy of growth — False correlations — Multiple. 133 . and lowly organised structures variable — Parts developed in an unusual manner are highly variable: specific characters more variable than generic: secondary sexual characters variable — Species of the same genus vary in an analogous manner — Reversions to long-lost characters — Summary. rudimentary.Chapter 5 Laws Of Variation Effects of changed conditions — Use and disuse. combined with natural selection.

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

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

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

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

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

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

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

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

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

but until some one will sow. or use and disuse. with the same precautions. played a considerable part in the modification of the constitution and structure. new varieties have not been produced. of the kidney-bean has been often cited for a similar purpose. has even been advanced. 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. the natural selection of innate variations. during a score of generations. and of this fact I have myself observed striking instances. 143 . and sometimes overmastered by. in some cases. and then collect seed from the few survivors. the experiment cannot be said to have been even tried. for it is now as tender as ever it was! The case. and with much greater weight. consequently. for an account has been published how much more hardy some seedlings are than others. have. as proving that acclimatisation cannot be effected. also. and then again get seed from these seedlings. with care to prevent accidental crosses. we may conclude that habit. On the whole. but that the effects have often been largely combined with.of which.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Secondary sexual characters are highly variable. such modifications will add to the beautiful and harmonious diversity of nature. 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. must have gone through an extraordinary amount of modification since the genus arose. 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. or these same species may occasionally revert to some of the characters of their ancient progenitors. natural selection has succeeded in giving a fixed character to the organ. for variation is a longcontinued and slow process. 164 . and exposed to similar influences. 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. in however extraordinary a manner it may have been developed. requiring a long lapse of time—in this case. and specific differences to the several species of the same genus. Species inheriting nearly the same constitution from a common parent. 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. and thus we can understand why it should often still be variable in a much higher degree than other parts. Variability in the same parts of the organisation has generally been taken advantage of in giving secondary sexual differences to the two sexes of the same species. in comparison with the same part or organ in the allied species. and such characters differ much in the species of the same group.varieties.

165 .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.

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

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

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

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

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

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

I can give only one or two instances of transitional habits and structures in allied species. it is clear that each must be well adapted to its place in nature. how. which serves as a parachute and allows them to glide through the air to an astonishing distance from tree to tree. 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. at least. short legs. and it had been asked how an insectivorous quadruped could possibly have been converted into a flying bat. of the squirrels would decrease in numbers or become 172 . 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. 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. Look at the Mustela vison of North America. either constant or occasional. here we have the finest gradation from animals with their tails only slightly flattened. and all analogy would lead us to believe that some.3. Look at the family of squirrels. for. And it seems to me that nothing less than a long list of such cases is sufficient to lessen the difficulty in any particular case like that of the bat. out of the many striking cases which I have collected. or. We cannot doubt that each structure is of use to each kind of squirrel in its own country. Let the climate and vegetation change. as Sir J. as on other occasions. to lessen the danger from occasional falls. by enabling it to escape birds or beasts of prey. the question would have been far more difficult to answer. and preys. Yet I think such difficulties have little weight. for instance. in the same species. or to collect food more quickly. Richardson has remarked. like other polecats on mice and land animals. and as each exists by a struggle for life. and which resembles an otter in its fur. to the so-called flying squirrels. and from others. If a different case had been taken. or old ones become modified. during summer this animal dives for and preys on fish. which has webbed feet. but during the long winter it leaves the frozen waters. and form of tail. Here. I lie under a heavy disadvantage. as there is reason to believe. and flying squirrels have their limbs and even the base of the tail united by a broad expanse of skin. could a land carnivorous animal have been converted into one with aquatic habits. let other competing rodents or new beasts of prey immigrate. with the posterior part of their bodies rather wide and with the skin on their flanks rather full. and of diversified habits.

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

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

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

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

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

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

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

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


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


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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

whereas. would have been of some avail against those who believe in an innate and necessary law of development. Bronn. been largely determined through natural selection. but this will be effected only under certain favourable circumstances. The fact of little or no modification having been effected since the glacial period.. survive from year to year by means of its seeds or ova? Mr. they might live together. on the principle of natural selection. such as high land or low land. The many animals which have remained unchanged since the commencement of the glacial period. the conditions of life. But. G. as well as to the amount of expenditure in reproduction and in general activity. of which we know anything. inhabiting distinct stations. the more permanent varieties are generally found. have changed during the last three or four thousand years. Moreover. E. and all mere temporary variations. owing to advantages gained through natural selection. in which the variability seems to be of a peculiar nature. it is probable.Cannot our critics conceive that a biennial plant or one of the lower animals might range into a cold climate and perish there every winter. and yet.H. as none of the animals and plants of Egypt. dry or moist districts. during the last several thousand years. such as size. as far as we know. but is powerless against the doctrine of natural selection or the survival of the fittest. which implies that when variations or individual differences of a beneficial nature happen to arise. can a variety live side by side with the parent species? If both have become fitted for slightly different habits of life or conditions. as far as its extreme complexity allows him to form a judgment. and if we lay on one side polymorphic species. or embalmed. and he concludes. for the ancient domestic races figured on the Egyptian monuments. Lewes has remarked. Ray Lankester has recently discussed this subject. in the case of 205 . at the close of his German translation of this work. yet all naturalists admit that such races have been produced through the modification of their original types. so probably have none in any part of the world. asks how. as Mr. this line of argument proves too much. as far as I can discover. would have been an incomparably stronger case. have remained absolutely uniform. in Egypt. And these conditions have. etc. that longevity is generally related to the standard of each species in the scale of organisation. It has been argued that. for these have been exposed to great changes of climate and have migrated over great distances. The celebrated palaeontologist. albinism. these will be preserved. are closely similar or even identical with those now living.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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


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


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

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

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

ostrich. their origin — Natural instincts of the cuckoo. 240 . its cell-making instinct — Changes of instinct and structure not necessarily simultaneous — Difficulties of the theory of the Natural Selection of instincts — Neuter or sterile insects — Summary.Chapter Instinct 8 Instincts comparable with habits. and parasitic bees — Slave-making ants — Hive-bee. but different in their origin — Instincts graduated — Aphides and ants — Instincts variable — Domestic instincts. molothrus.

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

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

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

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

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

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

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

perhaps.3. I will select only three. 248 . 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. the slave-making instinct of certain ants. namely. 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. Special Instincts We shall.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

as it bears on a fact. so that. how often the bees would pull down and rebuild in different ways the same cell. 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. but. I was able practically to show this fact. by covering the edges of the hexagonal walls of a single cell. 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. or the extreme margin of the circumferential rim of a growing comb. or leaving ungnawed. all trying to sweep equal spheres. and building up intermediate planes. sometimes recurring to a shape which they had at first rejected. projecting beyond the other completed cells. as when two pieces of comb met at an angle. Nor does there seem to me any great difficulty in a single insect (as in the case of a queen-wasp) making hexagonal cells. and worked into the growing edges of the cells all round. they can build up a wall intermediate between two adjoining spheres. It suffices that the bees should be enabled to stand at their proper relative distances from each other and from the walls of the last completed cells. as Huber has stated. placed directly under the middle of a comb growing downwards. and then. in its strictly proper place. This capacity in bees of laying down under certain circumstances a rough wall in its proper place between two justcommenced cells. It was really curious to note in cases of difficulty. 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. When bees have a place on which they can stand in their proper positions for working—for instance. a score of individuals work even at the commencement of the first cell. namely. is important. with an extremely thin layer of melted vermilion wax. as far as I have seen. on a slip of wood. which seems at first subversive of the foregoing theory. 263 . and then building up. by striking imaginary spheres. sweeping spheres or cylinders.working a short time at one cell going to another. that the cells on the extreme margin of wasp-combs are sometimes strictly hexagonal. but I have not space here to enter on this subject. the planes of intersection between these spheres. The work of construction seems to be a sort of balance struck between many bees. all instinctively standing at the same relative distance from each other. always standing at the proper relative distance from the parts of the cells just begun. if she were to work alternately on the inside and outside of two or three cells commenced at the same time.

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

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

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

and it had been profitable to the community that a number should have been annually born capable of work. The subject well deserves to be discussed at great length.importance. I can see no especial difficulty in this having been effected through natural selection. It may well be asked how it is possible to reconcile this case with the theory of natural selection? First. We have differences correlated not only with one 267 . that they could hardly have been acted on by natural selection. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure. namely. and yet. and in being destitute of wings and sometimes of eyes. but will confine myself to one special difficulty. How the workers have been rendered sterile is a difficulty. as in the shape of the thorax. But with the working ant we have an insect differing greatly from its parents. by individuals having been born with slight profitable modifications. from being sterile. let it be remembered that we have innumerable instances. and if such insects had been social. As far as instinct alone is concerned. which were inherited by the offspring. the wonderful difference in this respect between the workers and the perfect females would have been better exemplified by the hivebee. of all sorts of differences of inherited structure which are correlated with certain ages and with either sex. and that these again varied and again were selected. But I must pass over this preliminary difficulty. and in instinct. so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny. but not much greater than that of any other striking modification of structure. they cannot propagate their kind. for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile. and so onwards. that of working or sterile ants. 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. which at first appeared to me insuperable. but incapable of procreation. and actually fatal to the whole theory. I should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection. and consequently must believe that they were independently acquired through natural selection. yet absolutely sterile. but I will here take only a single case. If a working ant or other neuter insect had been an ordinary animal. 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. both in our domestic productions and in those in a state of nature. I will not here enter on these several cases.

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

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

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

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

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

removed by domestication — Laws governing the sterility of hybrids — Sterility not a special endowment. not universal. not accumulated by natural selection — Causes of the sterility of first crosses and of hybrids — Parallelism between the effects of changed conditions of life and of crossing — Dimorphism and trimorphism — Fertility of varieties when crossed and of their mongrel offspring not universal — Hybrids and mongrels compared independently of their fertility — Summary. affected by close interbreeding. 273 .Chapter 9 Hybridism Distinction between the sterility of first crosses and of hybrids — Sterility various in degree. but incidental on other differences.

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

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

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

so careful an observer as Gartner would have castrated his hybrids. and Rev. all the individuals are in this peculiar condition.experiments are in progress. certain abnormal individuals. He is as emphatic in his conclusion that some hybrids are perfectly fertile—as fertile as the pure parent-species—as are Kolreuter and Gartner that some degree of sterility between distinct species is a universal law of nature. He experimented on some of the very same species as did Gartner. that "every ovule in a pod of Crinum capense fertilised by C. in a first cross between two distinct species. revolutum produced a plant. or even more than commonly perfect fertility. The difference in their results may." Mr. Of his many important statements I will here give only a single one as an example. can easily be fertilised by the pollen from a distinct species. and by his having hot-houses at his command. though this pollen can be proved to be perfectly sound by fertilising other plants or species. which vegetated freely. a bulb of Hippeastrum aulicum produced four flowers. the strange fact of an increase of fertility in the successive generations of ARTIFICIALLY FERTILISED hybrids. but not by pollen from the same plant. be accounted for by too close interbreeding having been avoided. Herbert.fertilised. namely. three were fertilised by Herbert with their own pollen. And thus. I think. and in other species all the individuals. W. 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. as I believe. which I never saw to occur in a case of its natural fecundation. in Corydalis as shown by Professor Hildebrand." So that here we have perfect. the Hon. namely. In the genus Hippeastrum. This case of the Crinum leads me to refer to a singular fact. Verbascum and Passiflora. can actually be hybridised much more readily than they can be fertilised by pollen from the same individual plant! To give one instance. and after a few days perished entirely. either from the same plant or from another plant of the same hybrid nature. in contrast with those spontaneously self. in various orchids as shown by Mr. and bore good seed. and this would have insured in each generation a cross with pollen from a distinct flower. Herbert tried similar 277 . So that with some species. Now let us turn to the results arrived at by a third most experienced hybridiser. namely. be in part accounted for by Herbert's great horticultural skill. that individual plants of certain species of Lobelia. Scott and Fritz Muller. whereas the pod impregnated by the pollen of the hybrid made vigorous growth and rapid progress to maturity. may.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

I am fully persuaded that this double parallelism is by no means an accident or an illusion. He will at the same time be able to explain how it is that the races of some of our domesticated animals. the incessant action and reaction of various forces. will be able to explain the primary cause of hybrids being so generally sterile. which would probably have been sterile if aboriginally crossed. The above two parallel series of facts seem to be connected together by some common but unknown bond. Herbert Spencer. or consists in. 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. according to Mr. are incapable of breeding when kept under only partial confinement in their native country. are quite fertile together. and a multitude of other animals. produce hybrids which are almost always in some degree sterile. the vital forces gain in power. and when this tendency is slightly disturbed by any change. as throughout nature. which. He who is able to explain why the elephant. are always tending towards an equilibrium. although they are descended from distinct species. which is essentially related to the principle of life.or less sterile. being that life depends on. this principle. 292 .

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

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

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

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

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

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

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

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

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

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

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

the capacity in one species or variety to take on another. vigour and fertility of their offspring. so in crossing. that. firstly. In the case of first crosses it seems to depend on several circumstances. generally of an unknown nature. is incidental on differences. slight changes in the conditions of life add to the vigour and fertility of all organic beings. He who will explain these latter cases will be able to explain the sterility of hybrids. There is no more reason to think that species have been specially endowed with various degrees of sterility to prevent their crossing and blending in nature. the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. It is not always equal in degree in a first cross and in the hybrids produced from this cross. but is governed by several curious and complex laws. are very generally. 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. and is often so slight that the most careful experimentalists have arrived at diametrically opposite conclusions in ranking forms by this test. In the case of hybrids. and sometimes widely different in reciprocal crosses between the same two species. The sterility is innately variable in individuals of the same species. the sterility being closely allied to that which so frequently affects pure species. that the crossing of forms. Summary Of Chapter First crosses between forms. It is generally different. The sterility is of all degrees. 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. or which have varied. In the same manner as in grafting trees. favours the size. and secondly. perhaps render it probable that some unknown bond in all cases connects the degree of fertility of first unions with that 304 . in their vegetative systems. it apparently depends on their whole organisation having been disturbed by being compounded from two distinct forms. in some instances in chief part on the early death of the embryo. This view is strongly supported by a parallelism of another kind: namely. The degree of sterility does not strictly follow systematic affinity.8. The facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny. sterile. and their hybrids. and is eminently susceptible to action of favourable and unfavourable conditions. sufficiently distinct to be ranked as species. when exposed to new and unnatural conditions of life. but not universally.

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

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. 306 . on their number — On the lapse of time.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.

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

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

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

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

Nevertheless this impression is partly false. 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. are thousands of feet in thickness on the Continent. blank periods of enormous length.300 feet. Croll shows.154 Secondary strata… … … … … … … … … … . to the height or depth of thousands of feet.other. Professor Ramsay has published an account of a downthrow in Anglesea of 2. by calculating the 311 . and then at the figures representing several million years. and it makes no great difference whether the upheaval was sudden. very nearly thirteen and threequarters British miles.000 feet. that is. the two produce a totally different effect on the mind.240 —making altogether 72.190 Tertiary strata… … … … … … … … … … … . as most geologists now believe." but in estimating them by years.. between each successive formation we have. Mr. In regard to subaerial denudation..57. When geologists look at large and complicated phenomena. Professor Ramsay has given me the maximum thickness.13. and this is the result: Feet Palaeozoic strata (not including igneous beds). Some of these formations. and the figures are at once pronounced too small. yet in these cases there is nothing on the surface of the land to show such prodigious movements. Croll. the pile of rocks on either side of the crack having been smoothly swept away. each of which bears the stamp of time. So that the lofty pile of sedimentary rocks in Britain gives but an inadequate idea of the time which has elapsed during their accumulation. the surface of the land has been so completely planed down that no trace of these vast dislocations is externally visible. On the other hand.000 feet. and along this line the vertical displacement of the strata varies from 600 to 3. and he informs me that he fully believes that there is one in Merionethshire of 12. The Craven fault. for since the crust cracked. from actual measurement in most cases. In the Cordillera. they are good to show how slowly the mass must have been heaped together. in an interesting paper.584 feet. remarks that we do not err "in forming too great a conception of the length of geological periods. I estimated one mass of conglomerate at ten thousand feet. was slow and effected by many starts. yet from being formed of worn and rounded pebbles. in all parts of the world the piles of sedimentary strata are of wonderful thickness. Moreover. extends for upward of thirty miles. or. in the opinion of most geologists. and although conglomerates have probably been accumulated at a quicker rate than finer sediments. which are represented in England by thin beds. for instance. Mr. of the successive formations in DIFFERENT parts of Great Britain.2.

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

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

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

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

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

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

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

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

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

to be found in an older and underlying bed. and then re-covered by the upper beds of the same formation—facts. during the same geological period. It would seem that each separate formation. but has disappeared and reappeared. even if 321 . perhaps many times. has generally been intermittent in its accumulation. the probability is that it has not lived on the same spot during the whole period of deposition. representing formations which are elsewhere thousands of feet in thickness. Many instances could be given of beds. unless they are enabled to connect them together by the closest intermediate gradations. Consequently if it were to undergo a considerable amount of modification during the deposition of any one geological formation. denuded. which would not have been suspected. and which must have required an enormous period for their accumulation. when the same species occurs at the bottom. yet easily overlooked. they rank both as species. submerged. changes of form. of many long intervals of time and changes of level during the process of deposition. yet no one ignorant of this fact would have even suspected the vast lapse of time represented by the thinner formation.400 feet thick in Nova Scotia. though perhaps slight. still standing upright as they grew. Nor will the closest inspection of a formation give us any idea of the length of time which its deposition may have consumed. middle. In other cases we have the plainest evidence in great fossilised trees. It is all-important to remember that naturalists have no golden rule by which to distinguish species and varieties. and a third. Dawson found carboniferous beds 1. but abrupt. intervals have occurred in its accumulation. they grant some little variability to each species. but when they meet with a somewhat greater amount of difference between any two forms. had not the trees been preserved: thus Sir C. showing what wide. a formation composed of beds of widely different mineralogical composition. Hence. Supposing B and C to be two species. as is so often the case. from the reasons just assigned. at no less than sixty-eight different levels. and this. and top of a formation. we may reasonably suspect that the process of deposition has been more or less interrupted.than one palaeontologist that very thick deposits are usually barren of organic remains. like the whole pile of formations in any country. When we see. we can seldom hope to effect in any one geological section. except near their upper or lower limits. A. Lyell and Dr. Many cases could be given of the lower beds of a formation having been upraised. only a few feet in thickness. with ancient root-bearing strata. a section would not include all the fine intermediate gradations which must on our theory have existed. one above the other.

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

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

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

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

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

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

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

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

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

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

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

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

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

335 .Chapter 11 On The Geological Succession Of Organic Beings On the slow and successive appearance of new species — On their different rates of change — Species once lost do not reappear — Groups of species follow the same general rules in their appearance and disappearance as do single species — On extinction — On simultaneous changes in the forms of life throughout the world — On the affinities of extinct species to each other and to living species — On the state of development of ancient forms — On the succession of the same types within the same areas — Summary of preceding and present chapters.

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

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


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


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


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

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

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

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

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

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

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

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

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

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

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

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

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. the feebler will yield to the more dominant forms. On the theory of descent with modification. be found in one geological formation. succession of the same types within the same areas. armadillo. 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. In a similar manner we know. then we may conclude that generally only one species of each of the older genera has left modified descendants. during the next succeeding period of time. and have left no progeny. each having eight species. In failing orders. for the inhabitants of each quarter of the world will obviously tend to leave in that quarter. Analogous facts could be given in relation to the distribution of marine animals. each with the same number of species. 359 . which constitute the new genera containing the several species. if six genera. have left behind them the sloth. and anteater. the other seven species of each old genus having died out and left no progeny. on our theory. This cannot for an instant be admitted. still fewer genera and species will leave modified blood-descendants. But after very long intervals of time.closely allied. and after great geographical changes. than it is at present. permitting much intermigration. and some of these fossils may have been the actual progenitors of the living species. with the genera and species decreasing in numbers as is the case with the Edentata of South America. as their degenerate descendants. which formerly lived in South America. closely allied though in some degree modified descendants. so will their modified descendants still differ in nearly the same manner and degree. If the inhabitants of one continent formerly differed greatly from those of another continent. Or. These huge animals have become wholly extinct. from Falconer and Cautley's discoveries. is at once explained. It may be asked in ridicule whether I suppose that the megatherium and other allied huge monsters. and in a succeeding formation there be six other allied or representative genera. so that. It must not be forgotten that. and this will be a far commoner case. but not immutable. the great law of the long enduring. 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. all the species of the same genus are the descendants of some one species.

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

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

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

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

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

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

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

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

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

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

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

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

the arguments used by Forbes are to be trusted.3. have been 372 . and united almost every island with some mainland. It seems to me that we have abundant evidence of great oscillations in the level of the land or sea. now buried beneath the sea. Means Of Dispersal Sir C. or let it formerly have been submerged. might have been a high road for migration. when the climate was different. submerge it. In the coral-producing oceans such sunken islands are now marked by rings of coral or atolls standing over them. however. 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. presently have to discuss this branch of the subject in some detail. 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. and when in the course of time we know something definite about the means of distribution. If. Lyell and other authors have ably treated this subject. Whenever it is fully admitted. which may have served as halting places for plants and for many animals during their migration. it must be admitted that scarcely a single island exists which has not recently been united to some continent. but not of such vast changes in the position and extension of our continents. I can give here only the briefest abstract of the more important facts. A region now impassable to certain organisms from the nature of its climate. 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. and Europe likewise with America. and the two faunas will now blend together. This view cuts the Gordian knot of the dispersal of the same species to the most distant points. land may at a former period have connected islands or possibly even continents together. Other authors have thus hypothetically bridged over every ocean. I freely admit the former existence of many islands. that each species has proceeded from a single birthplace. indeed. or may formerly have blended. I shall. as it will some day be. Where the sea now extends. and removes many a difficulty. Change of climate must have had a powerful influence on migration. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas. No geologist disputes that great mutations of level have occurred within the period of existing organisms. we shall be enabled to speculate with security on the former extension of the land.

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

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

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

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

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

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

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

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

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

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

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

and fed glaciers which rolled 4. and Mr. in numerous immense boulders transported far from their parent source. Southward of the Asiatic continent. glaciers once extended far below their present level. from latitude 13 to 30 degrees south. and the same plants. J. In the Cordillera of South America. perpetual snow formerly covered the central axis. In Europe we meet with the plainest evidence of the Glacial period. D.B. according to Dr. Along this whole space of the Cordillera true glaciers do not now exist even at much more considerable heights. Hooker saw maize growing on ancient and gigantic moraines. Hector. from the excellent researches of Dr. at about the height of 12. we have the clearest evidence of former glacial action. We may infer from the frozen mammals and nature of the mountain vegetation.000 feet down the valleys.000 feet. we know. glaciers have left the marks of their former low descent. where the climate is now so different. Alternate Glacial Periods In The North And South But we must return to our more immediate subject. from latitude 41 degrees to the southernmost extremity. once formed a huge moraine. on the opposite side of the equator. In central Chile I examined a vast mound of detritus with great boulders.5. resembling those with which he was familiar in Norway. I am convinced that Forbes's view may be largely extended. there can hardly be a doubt. and southward to the Pyrenees. at points 900 miles apart. Further south. The same observer has recently found great moraines at a low level on the Atlas range in North Africa. and likewise great masses of detritus. Hooker. ice-borne fragments of rock have been observed on the eastern side of the continent. and in Sikkim. Erratic boulders have. which. From facts communicated to me by the Rev. W. that Siberia was similarly affected. as far south as latitude 36 and 37 degrees. Hooker on widely separated mountains in this island tell the same story of a former cold period. and on the shores of the Pacific. also. as far south as latitude 46 degrees.eastern corner of Australia. found by Dr. Forbes informs me that he found in various parts of the Cordillera. Along the Himalaya. Haast and Dr. nearly under the equator. 384 . Clarke. on both sides of the continent. deeply-furrowed rocks. been noticed on the Rocky Mountains. crossing the Portillo valley. In the Lebanon. it appears also that there are traces of former glacial action on the mountains of the south. including grooved pebbles. from the western shores of Britain to the Ural range. Dr. that in New Zealand immense glaciers formerly descended to a low level. Looking to America: in the northern half.

has attempted to show that a glacial condition of climate is the result of various physical causes. from the glacial action having extended all round the northern and southern hemispheres—from the period having been in a geological sense recent in both hemispheres—from its having lasted in both during a great length of time. that such occurred during the miocene and eocene formations. owing to certain contingencies. Mr. with the winters rendered much milder. All these causes tend towards the same end. is the relative position of the land and water. are common to North America and Europe. several geologists are convinced. between forty and fifty of the flowering plants of Tierra del Fuego. 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. is that whenever the northern hemisphere passes through a cold period the temperature of the southern hemisphere is actually raised. from direct evidence. But now. with slight alterations of climate. But the most important result for us. So conversely it will be with the northern hemisphere. brought into operation by an increase in the eccentricity of the earth's orbit. cold periods regularly recur every ten or fifteen thousand years. In South America. Croll believes that the last great glacial period occurred about 240.000 years. from glaciers having recently descended to a low level along the whole line of the Cordillera. Croll. for about 160. but I will first give the facts which demand an explanation. With respect to more ancient glacial periods. Mr. Croll.From these several facts. while the southern passes through a glacial period. as Sir C. enormously remote as these areas in opposite hemispheres are from each other. chiefly through changes in the direction of the ocean currents. Dr. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it. forming no inconsiderable part of its scanty flora. On the Organ Mountains of Brazil some few temperate European. arrived at by Mr. According to Mr. Croll. as may be inferred from the amount of work effected—and lastly. in a series of admirable memoirs. and these at long intervals are extremely severe.000 years ago. some Antarctic and some Andean genera were found by Gardner which do not exist in the low intervening hot countries. and endured. but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. On the Silla of Caraccas 385 . of which the most important. Lyell has shown. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur. namely. Hooker has shown that besides many closely allied species. not to mention still more ancient formations.

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

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

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

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

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

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

392 .Chapter 13 Geographical Distribution--continued Distribution of fresh-water productions — On the inhabitants of oceanic islands — Absence of Batrachians and of terrestrial Mammals — On the relation of the inhabitants of islands to those of the nearest mainland — On colonisation from the nearest source with subsequent modification — Summary of the last and present chapters.

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

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

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

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

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

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

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

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

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

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


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


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


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

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

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

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

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

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

organs in this condition are often of much value in classification. and the differences are of quite subordinate value in classification. 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. If the Ornithorhynchus had been covered with feathers instead of hair. when all taken together. and yet leave us in no doubt where it should be ranked. though here even. for classification. whether or not there is an open passage from the nostrils to the mouth. also. yet no one will say that the antennae in these two divisions of the same order are of unequal physiological importance. both of high physiological importance. For instance. in another division they differ much. they appear insufficient to separate Cnestis from Connarus. the antennae. the only character. No one will dispute that the rudimentary teeth in the upper jaws of young ruminants. yet. are most constant in structure." To give an example among insects: in one great division of the Hymenoptera. Again. Any number of instances could be given of the varying importance for classification of the same important organ within the same group of beings. a species may depart from its allies in several characters. and of almost universal prevalence. as has often been remarked. Numerous instances could be given of characters derived from parts which must be consi