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The Origin of Species by means of Natural Selection;


Preservation of Favoured Races in the Struggle for Life.

By Charles DarwinM.A.F.R.S.
Author of "The Descent of Man etc., etc.

Sixth London Edition, with all Additions and Corrections.
The 6th Edition is often considered the definititive edition.
Also see Project Gutenberg Etext #1228 for an earlier edition.

But with regard to the material worldwe can at least go so far as this--
we can perceive that events are brought about not by insulated
interpositions of Divine powerexerted in each particular casebut 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".




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 productionsand had been separately created. This
view has been ably maintained by many authors. Some few naturalistson
the other handhave believed that species undergo modificationand 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 (Aristotlein his "Physicae Auscultationes" (lib.2cap.8s.2)
after remarking that rain does not fall in order to make the corn growany
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 Grecewho 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 forthbut how little Aristotle
fully comprehended the principleis 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 periodsand as he does not enter on the causes or means of the
transformation of speciesI 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
subsequently1815in the Introduction to his "Hist. Nat. des Animaux sans
Vertebres". In these works he up holds the doctrine that all species
including manare descended from other species. He first did the eminent
service of arousing attention to the probability of all change in the
organicas well as in the inorganic worldbeing the result of lawand
not of miraculous interposition. Lamarck seems to have been chiefly led to
his conclusion on the gradual change of speciesby the difficulty of
distinguishing species and varietiesby the almost perfect gradation of
forms in certain groupsand by the analogy of domestic productions. With
respect to the means of modificationhe attributed something to the direct
action of the physical conditions of lifesomething to the crossing of
already existing formsand much to use and disusethat isto 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
developmentand as all the forms of life thus tend to progressin order
to account for the existence at the present day of simple productionshe
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 4051859) 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
grandfatherDr. Erasmus Darwinanticipated 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 viewsas shown in the
introduction to a work written in 1794 and 1795but not published till
long afterward; he has pointedly remarked ("Goethe als Naturforscher"von
Dr. Karl Medings. 34) that the future question for naturalists will be
howfor instancecattle 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 timethat Goethe in GermanyDr. Darwin in Englandand
Geoffroy Saint-Hilaire (as we shall immediately see) in Francecame to the
same conclusion on the origin of speciesin the years 1794-5.)

Geoffroy Saint-Hilaireas is stated in his "Life"written by his son
suspectedas early as 1795that 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 lifeor the "monde ambiant" as the cause of change. He was
cautious in drawing conclusionsand did not believe that existing species
are now undergoing modification; andas his son addsC'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
Femalepart 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 selectionand this is the first recognition which has been
indicated; but he applies it only to the races of manand to certain
characters alone. After remarking that negroes and mulattoes enjoy an
immunity from certain tropical diseaseshe observesfirstlythat all
animals tend to vary in some degreeandsecondlythat agriculturists
improve their domesticated animals by selection; and thenhe addsbut
what is done in this latter case "by artseems to be done with equal
efficacythough more slowlyby naturein the formation of varieties of
mankindfitted for the country which they inhabit. Of the accidental
varieties of manwhich would occur among the first few and scattered
inhabitants of the middle regions of Africasome one would be better
fitted than others to bear the diseases of the country. This race would
consequently multiplywhile the others would decrease; not only from their
in ability to sustain the attacks of diseasebut from their incapacity of
contending with their more vigorous neighbours. The colour of this
vigorous race I take for grantedfrom what has been already saidwould be
dark. But the same disposition to form varieties still existinga darker
and a darker race would in the course of time occur: and as the darkest
would be the best fitted for the climatethis would at length become the
most prevalentif not the only racein 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. Rowleyof the United Statesfor
having called my attentionthrough Mr. Braceto the above passage of Dr.
Wells' work.

The Hon. and Rev. W. Herbertafterward Dean of Manchesterin the fourth
volume of the "Horticultural Transactions"1822and in his work on the
Amaryllidaceae(1837pages 19339)declares that "horticultural
experiments have establishedbeyond the possibility of refutationthat
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 conditionand
that these have producedchiefly by inter-crossingbut likewise by
variationall our existing species.

In 1826 Professor Grantin the concluding paragraph in his well-known
paper ("Edinburgh Philosophical Journal"vol. XIVpage 283) on the
Spongillaclearly declares his belief that species are descended from
other speciesand that they become improved in the course of modification.
This same view was given in his Fifty-fifth Lecturepublished in the
Lancetin 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 subjectso that
it remained unnoticed until Mr. Matthew himself drew attention to it in the
Gardeners' Chronicleon April 71860. 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 periodsand then restocked; and
he gives as an alternativethat 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 sawhoweverthe
full force of the principle of natural selection.

The celebrated geologist and naturalistVon Buchin his excellent
Description Physique des Isles Canaries(1836page 147)clearly
expresses his belief that varieties slowly become changed into permanent
specieswhich are no longer capable of intercrossing.

Rafinesquein his "New Flora of North America"published in 1836wrote

(page 6) as follows: "All species might have been varieties onceand many
varieties are gradually becoming species by assuming constant and peculiar
characters;" but further on (page 18) he addsexcept the original types
or ancestors of the genus.

In 1843-44 Professor Haldeman ("Boston Journal of Nat. Hist. U. States"
vol. ivpage 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 isthat the several
series of animated beingsfrom the simplest and oldest up to the highest
and most recentareunder the providence of Godthe resultsFIRSTof
an impulse which has been imparted to the forms of lifeadvancing themin
definite timesby generationthrough grades of organisation terminating
in the highest dicotyledons and vertebratathese grades being few in
numberand generally marked by intervals of organic characterwhich we
find to be a practical difficulty in ascertaining affinities; SECONDof
another impulse connected with the vital forcestendingin the course of
generationsto modify organic structures in accordance with external
circumstancesas foodthe nature of the habitatand the meteoric
agenciesthese 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 howfor
instancea woodpecker has become adapted to its peculiar habits of life.
The workfrom its powerful and brilliant stylethough displaying in the
early editions little accurate knowledge and a great want of scientific
cautionimmediately had a very wide circulation. In my opinion it has
done excellent service in this country in calling attention to the subject
in removing prejudiceand 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.
xiiipage 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 Owenin 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 Associationin 1858he speaks (page li) of "the
axiom of the continuous operation of creative poweror of the ordained
becoming of living things." Further on (page xc)after referring to
geographical distributionhe addsThese 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
islandshe chiefly expresses that he knows not how the Red Grouse came to
be thereand there exclusively; signifying alsoby this mode of
expressing such ignorancehis 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 addressone by the otherit 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 Speciespresently to be referred tohad been read before
the Linnean Society. When the first edition of this work was publishedI
was so completely deceivedas were many othersby 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 inferredand the inference still seems to me
perfectly justfrom a passage beginning with the words "no doubt the typeform
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 myselfthat 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 otheras I do. As far as the mere enunciation of the
principle of natural selection is concernedit is quite immaterial whether
or not Professor Owen preceded mefor both of usas shown in this
historical sketchwere long ago preceded by Dr. Wells and Mr. Matthews.

M. Isidore Geoffroy Saint-Hilairein 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
fixespour chaque especetant qu'elle se perpetue au milieu des memes
circonstances: ils se modifientsi les circonstances ambiantes viennent a
changer. En resumeL'OBSERVATION des animaux sauvages demontre deja la
variabilite LIMITEE des especes. Les EXPERIENCES sur les animaux sauvages
devenus domestiqueset sur les animaux domestiques redevenus sauvagesla
demontrent plus clairment encore. Ces memes experiences prouventde plus
que les differences produites peuvent etre de VALEUR GENERIQUE." In his
Hist. Nat. Generale(tom. iipage 4301859) he amplifies analogous
>From a circular lately issued it appears that Dr. Frekein 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 Spencerin an Essay (originally published in the "Leader"
March1852and 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
productionsfrom the changes which the embryos of many species undergo
from the difficulty of distinguishing species and varietiesand from the
principle of general gradationthat 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. Naudina distinguished botanistexpressly statedin 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 believeslike Dean Herbertthat specieswhen nascentwere
more plastic than at present. He lays weight on what he calls the
principle of finalitypuissance mysterieuse, indeterminee; fatalite pour
les uns; pour les autres volonte providentielle, dont l'action incessante
sur les etres vivantes determine, a toutes les epoques de l'existence du
monde, la forme, le volume, et la duree de chacun d'eux, en raison de sa
destinee dans l'ordre de choses dont il fait partie. C'est cette puissance
qui harmonise chaque membre a l'ensemble, en l'appropriant a la fonction
qu'il doit remplir dans l'organisme general de la nature, fonction qui est
pour lui sa raison d'etre.(From references in Bronn's "Untersuchungen
uber die Entwickelungs-Gesetze"it appears that the celebrated botanist
and palaeontologist Unger publishedin 1852his belief that species
undergo development and modification. Daltonlikewisein Pander and
Dalton's work on Fossil Slothsexpressedin 1821a similar belief.
Similar views haveas is well knownbeen maintained by Oken in his
mystical "Natur-Philosophie". From other references in Godron's work "Sur
l'Espece"it seems that Bory St. VincentBurdachPoiret and Frieshave
all admitted that new species are continually being produced. I may add
that of the thirty-four authors named in this Historical Sketchwho
believe in the modification of speciesor at least disbelieve in separate
acts of creationtwenty-seven have written on special branches of natural
history or geology.)

In 1853 a celebrated geologistCount Keyserling ("Bulletin de la Soc.
Geolog."2nd Ser.tom. xpage 357)suggested that as new diseases
supposed to have been caused by some miasma have arisen and spread over the
worldso at certain periods the germs of existing species may have been
chemically affected by circumambient molecules of a particular natureand
thus have given rise to new forms.

In this same year1853Dr. Schaaffhausen published an excellent pamphlet
("Verhand. des Naturhist. Vereins der Preuss. Rheinlands"etc.)in which
he maintains the development of organic forms on the earth. He infers that
many species have kept true for long periodswhereas a few have become
modified. The distinction of species he explains by the destruction of
intermediate graduated forms. "Thus living plants and animals are not
separated from the extinct by new creationsbut are to be regarded as
their descendants through continued reproduction."

A well-known French botanistM. Lecoqwrites in 1854 ("Etudes sur
Geograph. Bot. tom. ipage 250)On voit que nos recherches sur la fixite
ou la variation de l'espece, nous conduisent directement aux idees emises
par deux hommes justement celebres, Geoffroy Saint-Hilaire et Goethe.
Some other passages scattered through M. Lecoq's large work make it a
little doubtful how far he extends his views on the modification of

The "Philosophy of Creation" has been treated in a masterly manner by the
Rev. Baden Powellin his "Essays on the Unity of Worlds"1855. Nothing
can be more striking than the manner in which he shows that the
introduction of new species is "a regularnot a casual phenomenon or, as
Sir John Herschel expresses it, a natural in contradistinction to a
miraculous process."

The third volume of the "Journal of the Linnean Society" contains papers
read July 11858by Mr. Wallace and myselfin whichas stated in the
introductory remarks to this volumethe theory of Natural Selection is
promulgated by Mr. Wallace with admirable force and clearness.

Von Baertoward whom all zoologists feel so profound a respectexpressed
about the year 1859 (see Prof. Rudolph WagnerZoologisch-Anthropologische
Untersuchungen1861s. 51) his convictionchiefly grounded on the laws
of geographical distributionthat forms now perfectly distinct have
descended from a single parent-form.

In June1859Professor Huxley gave a lecture before the Royal Institution
on the "Persistent Types of Animal Life". Referring to such caseshe
remarksIt is difficult to comprehend the meaning of such facts as these,
if we suppose that each species of animal and plant, or each great type of
organisation, was formed and placed upon the surface of the globe at long
intervals by a distinct act of creative power; 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. If, on the other hand, 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 speciesa hypothesis whichthough unprovenand sadly
damaged by some of its supportersis yet the only one to which physiology
lends any countenance; 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."

In December1859Dr. Hooker published his "Introduction to the Australian
Flora". In the first part of this great work he admits the truth of the
descent and modification of speciesand supports this doctrine by many
original observations.

The first edition of this work was published on November 241859and the
second edition on January 71860.





Causes of Variability -- Effects of Habit and the use or disuse of Parts --
Correlated Variation -- Inheritance -- Character of Domestic Varieties --
Difficulty of distinguishing between Varieties and Species -- Origin of
Domestic Varieties from one or more Species -- Domestic Pigeonstheir
Differences and Origin -- Principles of Selectionanciently followed
their Effects -- Methodical and Unconscious Selection -- Unknown Origin of
our Domestic Productions -- Circumstances favourable to Man's power of



Variability -- Individual Differences -- Doubtful species -- Wide ranging
much diffusedand common speciesvary 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 closelybut unequallyrelated to each otherand in having
restricted ranges.



Its bearing on natural selection -- The term used in a wide sense --
Geometrical ratio of increase -- Rapid increase of naturalised animals and
plants -- Nature of the checks to increase -- Competition universal --
Effects of climate -- Protection from the number of individuals -- Complex
relations of all animals and plants throughout nature -- Struggle for life
most severe between individuals and varieties of the same species; often
severe between species of the same genus -- The relation of organism to
organism the most important of all relations.



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 Selectionnamelyintercrossing
isolationnumber of individuals -- Slow action -- Extinction caused by
Natural Selection -- Divergence of Characterrelated to the diversity of
inhabitants of any small area and to naturalisation -- Action of Natural
Selectionthrough Divergence of Character and Extinctionon 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.



Effects of changed conditions -- Use and disusecombined with natural
selection; organs of flight and of vision -- Acclimatisation -- Correlated
variation -- Compensation and economy of growth -- False correlations -Multiple
rudimentaryand 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.



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.



Longevity -- Modifications not necessarily simultaneous -- Modifications
apparently of no direct service -- Progressive development -- Characters of
small functional importancethe 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 classdeveloped from one
and the same source -- Reasons for disbelieving in great and abrupt



Instincts comparable with habitsbut different in their origin --
Instincts graduated -- Aphides and ants -- Instincts variable -- Domestic
instinctstheir origin -- Natural instincts of the cuckoomolothrus
ostrichand parasitic bees -- Slave-making ants -- Hive-beeits cellmaking
instinct -- Changes of instinct and structure not necessarily
simultaneous -- Difficulties on the theory of the Natural Selection of
instincts -- Neuter or sterile insects -- Summary.



Distinction between the sterility of first crosses and of hybrids --
Sterility various in degreenot universalaffected by close
interbreedingremoved by domestication -- Laws governing the sterility of
hybrids -- Sterility not a special endowmentbut incidental on other
differencesnot 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.



On the absence of intermediate varieties at the present day -- On the
nature of extinct intermediate varieties; on their number -- On the lapse
of timeas inferred from the rate of denudation and of deposition -- On
the lapse of time as estimated in years -- On the poorness of our
palaeontological collections -- On the intermittence of geological
formations -- On the denudation of granitic areas -- On the absence of
intermediate varieties in any one formation -- On the sudden appearance of
groups of species -- On their sudden appearance in the lowest known
fossiliferous strata -- Antiquity of the habitable earth.



On the slow and successive appearance of new species -- On their different
rates of change -- Species once lost do not reappear -- Groups of species
follow the same general rules in their appearance and disappearance as do

single species -- On extinction -- On simultaneous changes in the forms of
life throughout the world -- On the affinities of extinct species to each
other and to living species -- On the state of development of ancient forms
-- On the succession of the same types within the same areas -- Summary of
preceding and present chapter.



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 landand by occasional means -- Dispersal
during the Glacial period -- Alternate Glacial periods in the north and



Distribution of fresh-water productions -- On the inhabitants of oceanic
islands -- Absence of Batrachians and of terrestrial Mammals -- On the
relation of the inhabitants of islands to those of the nearest mainland --
On colonisation from the nearest source with subsequent modification --
Summary of the last and present chapter.



Classificationgroups subordinate to groups -- Natural system -- Rules and
difficulties in classificationexplained on the theory of descent with
modification -- Classification of varieties -- Descent always used in
classification -- Analogical or adaptive characters -- Affinitiesgeneral
complex and radiating -- Extinction separates and defines groups --
Morphologybetween members of the same classbetween parts of the same
individual -- Embryologylaws ofexplained by variations not supervening
at an early ageand being inherited at a corresponding age -- Rudimentary
Organs; their origin explained -- Summary.



Recapitulation of the objections to the theory of Natural Selection --
Recapitulation of the general and special circumstances in its favour --
Causes of the general belief in the immutability of species -- How far the
theory of Natural Selection may be extended -- Effects of its adoption on
the study of Natural history -- Concluding remarks.





When on board H.M.S. Beagleas naturalistI was much struck with certain
facts in the distribution of the organic beings inhabiting South America
and in the geological relations of the present to the past inhabitants of
that continent. These factsas will be seen in the latter chapters of
this volumeseemed to throw some light on the origin of species--that
mystery of mysteriesas it has been called by one of our greatest
philosophers. On my return homeit occurred to mein 1837that
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. After five years' work I allowed myself to speculate on
the subjectand drew up some short notes; these I enlarged in 1844 into a
sketch of the conclusionswhich then seemed to me probable: from that
period to the present day I have steadily pursued the same object. I hope
that I may be excused for entering on these personal detailsas I give
them to show that I have not been hasty in coming to a decision.

My work is now (1859) nearly finished; but as it will take me many more
years to complete itand as my health is far from strongI have been
urged to publish this abstract. I have more especially been induced to do
thisas Mr. Wallacewho is now studying the natural history of the Malay
Archipelagohas arrived at almost exactly the same general conclusions
that I have on the origin of species. In 1858 he sent me a memoir on this
subjectwith a request that I would forward it to Sir Charles Lyellwho
sent it to the Linnean Societyand it is published in the third volume of
the Journal of that Society. Sir C. Lyell and Dr. Hookerwho both knew of
my work--the latter having read my sketch of 1844--honoured me by thinking
it advisable to publishwith Mr. Wallace's excellent memoirsome brief
extracts from my manuscripts.

This abstractwhich I now publishmust necessarily be imperfect. I
cannot here give references and authorities for my several statements; and
I must trust to the reader reposing some confidence in my accuracy. No
doubt errors may have crept inthough I hope I have always been cautious
in trusting to good authorities alone. I can here give only the general
conclusions at which I have arrivedwith a few facts in illustrationbut
whichI hopein most cases will suffice. No one can feel more sensible
than I do of the necessity of hereafter publishing in detail all the facts
with referenceson which my conclusions have been grounded; and I hope in
a future work to do this. For I am well aware that scarcely a single point
is discussed in this volume on which facts cannot be adducedoften
apparently leading to conclusions directly opposite to those at which I
have arrived. A fair result can be obtained only by fully stating and
balancing the facts and arguments on both sides of each question; and this
is here impossible.

I much regret that want of space prevents my having the satisfaction of
acknowledging the generous assistance which I have received from very many
naturalistssome of them personally unknown to me. I cannothoweverlet
this opportunity pass without expressing my deep obligations to Dr. Hooker
whofor the last fifteen yearshas aided me in every possible way by his
large stores of knowledge and his excellent judgment.

In considering the origin of speciesit is quite conceivable that a
naturalistreflecting on the mutual affinities of organic beingson their
embryological relationstheir geographical distributiongeological
successionand other such factsmight come to the conclusion that species
had not been independently createdbut had descendedlike varietiesfrom
other species. Neverthelesssuch a conclusioneven if well founded

would be unsatisfactoryuntil it could be shown how the innumerable
speciesinhabiting this world have been modifiedso as to acquire that
perfection of structure and coadaptation which justly excites our
admiration. Naturalists continually refer to external conditionssuch as the only possible cause of variation. In one
limited senseas we shall hereafter seethis may be true; but it is
preposterous to attribute to mere external conditionsthe structurefor
instanceof the woodpeckerwith its feettailbeakand tongueso
admirably adapted to catch insects under the bark of trees. In the case of
the mistletoewhich draws its nourishment from certain treeswhich has
seeds that must be transported by certain birdsand which has flowers with
separate sexes absolutely requiring the agency of certain insects to bring
pollen from one flower to the otherit is equally preposterous to account
for the structure of this parasitewith its relations to several distinct
organic beingsby the effects of external conditionsor of habitor of
the volition of the plant itself.

It isthereforeof the highest importance to gain a clear insight into
the means of modification and coadaptation. 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. Nor have I been disappointed; in this and in all
other perplexing cases I have invariably found that our knowledge
imperfect though it beof variation under domesticationafforded the best
and safest clue. I may venture to express my conviction of the high value
of such studiesalthough they have been very commonly neglected by

>From these considerationsI shall devote the first chapter of this
abstract to variation under domestication. We shall thus see that a large
amount of hereditary modification is at least possible; andwhat is
equally or more importantwe shall see how great is the power of man in
accumulating by his selection successive slight variations. I will then
pass on to the variability of species in a state of nature; but I shall
unfortunatelybe compelled to treat this subject far too brieflyas it
can be treated properly only by giving long catalogues of facts. We shall
howeverbe enabled to discuss what circumstances are most favourable to
variation. In the next chapter the struggle for existence among all
organic beings throughout the worldwhich inevitably follows from the high
geometrical ratio of their increasewill be considered. This is the
doctrine of Malthusapplied to the whole animal and vegetable kingdoms.
As many more individuals of each species are born than can possibly
survive; and asconsequentlythere is a frequently recurring struggle for
existenceit follows that any beingif it vary however slightly in any
manner profitable to itselfunder the complex and sometimes varying
conditions of lifewill have a better chance of survivingand thus be
NATURALLY SELECTED. From the strong principle of inheritanceany selected
variety will tend to propagate its new and modified form.

This fundamental subject of natural selection will be treated at some
length in the fourth chapter; and we shall then see how natural selection
almost inevitably causes much extinction of the less improved forms of
lifeand leads to what I have called divergence of character. In the next
chapter I shall discuss the complex and little known laws of variation. In
the five succeeding chaptersthe most apparent and gravest difficulties in
accepting the theory will be given: namelyfirstthe difficulties of
transitionsor how a simple being or a simple organ can be changed and
perfected into a highly developed being or into an elaborately constructed
organ; secondly the subject of instinctor the mental powers of animals;
thirdlyhybridismor the infertility of species and the fertility of
varieties when intercrossed; and fourthlythe imperfection of the
geological record. In the next chapter I shall consider the geological
succession of organic beings throughout time; in the twelfth and
thirteenththeir geographical distribution throughout space; in the

fourteenththeir classification or mutual affinitiesboth when mature and
in an embryonic condition. In the last chapter I shall give a brief
recapitulation of the whole workand a few concluding remarks.

No one ought to feel surprise at much remaining as yet unexplained in
regard to the origin of species and varietiesif he make due allowance for
our profound ignorance in regard to the mutual relations of the many beings
which live around us. Who can explain why one species ranges widely and is
very numerousand why another allied species has a narrow range and is
rare? Yet these relations are of the highest importancefor they
determine the present welfare andas I believethe future success and
modification of every inhabitant of this world. 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. Although much remains obscure
and will long remain obscureI can entertain no doubtafter the most
deliberate study and dispassionate judgment of which I am capablethat the
view which most naturalists until recently entertainedand which I
formerly entertained--namelythat each species has been independently
created--is erroneous. I am fully convinced that species are not
immutable; but that those belonging to what are called the same genera are
lineal descendants of some other and generally extinct speciesin the same
manner as the acknowledged varieties of any one species are the descendants
of that species. FurthermoreI am convinced that natural selection has
been the most importantbut not the exclusivemeans of modification.



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 Pigeonstheir
Differences and Origin -- Principles of Selectionanciently followed
their Effects -- Methodical and Unconscious Selection -- Unknown Origin of
our Domestic Productions -- Circumstances favourable to Man's power of


When we compare the individuals of the same variety or sub-variety of our
older cultivated plants and animalsone of the first points which strikes
us isthat they generally differ more from each other than do the
individuals of any one species or variety in a state of nature. And if we
reflect on the vast diversity of the plants and animals which have been
cultivatedand which have varied during all ages under the most different
climates and treatmentwe are driven to conclude that this great
variability is due to our domestic productions having been raised under
conditions of life not so uniform asand somewhat different fromthose to
which the parent species had been exposed under nature. There isalso
some probability in the view propounded by Andrew Knightthat this
variability may be partly connected with excess of food. It seems clear
that organic beings must be exposed during several generations to new
conditions to cause any great amount of variation; and thatwhen the
organisation has once begun to varyit generally continues varying for
many generations. No case is on record of a variable organism ceasing to
vary under cultivation. Our oldest cultivated plantssuch as wheatstill
yield new varieties: our oldest domesticated animals are still capable of
rapid improvement or modification.

As far as I am able to judgeafter long attending to the subjectthe
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. With respect to the direct actionwe must bear in
mind that in every caseas Professor Weismann has lately insistedand as
I have incidently shown in my work on "Variation under Domestication
there are two factors: namely, the nature of the organism and the nature
of the conditions. The former seems to be much the more important; for
nearly similar variations sometimes arise under, as far as we can judge,
dissimilar conditions; and, on the other hand, dissimilar variations arise
under conditions which appear to be nearly uniform. The effects on the
offspring are either definite or in definite. 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. It is extremely difficult to come to any conclusion in regard to
the extent of the changes which have been thus definitely induced. There
can, however, be little doubt about many slight changes, such as size from
the amount of food, colour from the nature of the food, thickness of the
skin and hair from climate, etc. Each of the endless variations which we
see in the plumage of our fowls must have had some efficient cause; and if
the same cause were to act uniformly during a long series of generations on
many individuals, all probably would be modified in the same manner. 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,
shows us what singular modifications might result in the case of plants
from a chemical change in the nature of the sap.

In definite variability is a much more common result of changed conditions
than definite variability, and has probably played a more important part in
the formation of our domestic races. We see in definite variability in the
endless slight peculiarities which distinguish the individuals of the same
species, and which cannot be accounted for by inheritance from either
parent or from some more remote ancestor. Even strongly-marked differences
occasionally appear in the young of the same litter, and in seedlings from
the same seed-capsule. At long intervals of time, out of millions of
individuals reared in the same country and fed on nearly the same food,
deviations of structure so strongly pronounced as to deserve to be called
monstrosities arise; but monstrosities cannot be separated by any distinct
line from slighter variations. All such changes of structure, whether
extremely slight or strongly marked, which appear among many individuals
living together, may be considered as the in definite effects of the
conditions of life on each individual organism, in nearly the same manner
as the chill effects different men in an in definite manner, according to
their state of body or constitution, causing coughs or colds, rheumatism,
or inflammation of various organs.

With respect to what I have called the in direct action of changed
conditions, namely, through the reproductive system of being affected, we
may infer that variability is thus induced, partly from the fact of this
system being extremely sensitive to any change in the conditions, and
partly from the similarity, as Kolreuter and others have remarked, between
the variability which follows from the crossing of distinct species, and
that which may be observed with plants and animals when reared under new or
unnatural conditions. Many facts clearly show how eminently susceptible
the reproductive system is to very slight changes in the surrounding
conditions. Nothing is more easy than to tame an animal, and few things
more difficult than to get it to breed freely under confinement, even when
the male and female unite. How many animals there are which will not
breed, though kept in an almost free state in their native country! This
is generally, but erroneously attributed to vitiated instincts. Many
cultivated plants display the utmost vigour, and yet rarely or never seed!
In some few cases it has been discovered that a very trifling change, such
as a little more or less water at some particular period of growth, will
determine whether or not a plant will produce seeds. I cannot here give
the details which I have collected and elsewhere published on this curious
subject; but to show how singular the laws are which determine the
reproduction of animals under confinement, I may mention that carnivorous

animals, even from the tropics, breed in this country pretty freely under
confinement, with the exception of the plantigrades or bear family, which
seldom produce young; whereas, carnivorous birds, with the rarest
exception, hardly ever lay fertile eggs. Many exotic plants have pollen
utterly worthless, in the same condition as in the most sterile hybrids.
When, on the one hand, we see domesticated animals and plants, though often
weak and sickly, breeding freely under confinement; and when, on the other
hand, we see individuals, though taken young from a state of nature
perfectly tamed, long-lived, and healthy (of which I could give numerous
instances), yet having their reproductive system so seriously affected by
unperceived causes as to fail to act, we need not be surprised at this
system, when it does act under confinement, acting irregularly, and
producing offspring somewhat unlike their parents. I may add that as some
organisms breed freely under the most unnatural conditions--for instance,
rabbits and ferrets kept in hutches--showing that their reproductive organs
are not easily affected; so will some animals and plants withstand
domestication or cultivation, and vary very slightly--perhaps hardly more
than in a state of nature.

Some naturalists have maintained that all variations are connected with the
act of sexual reproduction; but this is certainly an error; for I have
given in another work a long list of sporting plants;" as they are called
by gardeners; that isof 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. These bud variationsas they may be namedcan be
propagated by graftsoffsetsetc.and sometimes by seed. They occur
rarely under naturebut are far from rare under culture. As a single bud
out of many thousands produced year after year on the same tree under
uniform conditionshas been known suddenly to assume a new character; and
as buds on distinct treesgrowing under different conditionshave
sometimes yielded nearly the same variety--for instancebuds on peachtrees
producing nectarinesand buds on common roses producing moss-roses-we
clearly see that the nature of the conditions is of subordinate
importance in comparison with the nature of the organism in determining
each particular form of variation; perhaps of not more importance than the
nature of the sparkby which a mass of combustible matter is ignitedhas
in determining the nature of the flames.


Changed habits produce an inherited effect as in the period of the
flowering of plants when transported from one climate to another. With
animals the increased use or disuse of parts has had a more marked
influence; thus I find in the domestic duck that the bones of the wing
weigh less and the bones of the leg morein proportion to the whole
skeletonthan do the same bones in the wild duck; and this change may be
safely attributed to the domestic duck flying much lessand walking more
than its wild parents. The great and inherited development of the udders
in cows and goats in countries where they are habitually milkedin
comparison with these organs in other countriesis probably another
instance of the effects of use. Not one of our domestic animals can be
named which has not in some country drooping ears; and the view which has
been suggested that the drooping is due to disuse of the muscles of the
earfrom the animals being seldom much alarmedseems probable.

Many laws regulate variationsome few of which can be dimly seenand will
hereafter be briefly discussed. I will here only allude to what may be
called correlated variation. Important changes in the embryo or larva will
probably entail changes in the mature animal. In monstrositiesthe
correlations between quite distinct parts are very curious; and many
instances are given in Isidore Geoffroy St. Hilaire's great work on this
subject. Breeders believe that long limbs are almost always accompanied by
an elongated head. Some instances of correlation are quite whimsical; thus

cats which are entirely white and have blue eyes are generally deaf; but it
has been lately stated by Mr. Tait that this is confined to the males.
Colour and constitutional peculiarities go togetherof which many
remarkable cases could be given among animals and plants. From facts
collected by Heusingerit appears that white sheep and pigs are injured by
certain plantswhile dark-coloured individuals escape: Professor Wyman
has recently communicated to me a good illustration of this fact; on asking
some farmers in Virginia how it was that all their pigs were blackthey
informed him that the pigs ate the paint-root (Lachnanthes)which coloured
their bones pinkand which caused the hoofs of all but the black varieties
to drop off; and one of the "crackers" (i.e. Virginia squatters) addedwe
select the black members of a litter for raising, as they alone have a good
chance of living.Hairless dogs have imperfect teeth; long-haired and
coarse-haired animals are apt to haveas is assertedlong or many horns;
pigeons with feathered feet have skin between their outer toes; pigeons
with short beaks have small feetand those with long beaks large feet.
Hence if man goes on selectingand thus augmentingany peculiarityhe
will almost certainly modify unintentionally other parts of the structure
owing to the mysterious laws of correlation.

The results of the variousunknownor but dimly understood laws of
variation are infinitely complex and diversified. It is well worth while
carefully to study the several treatises on some of our old cultivated
plantsas on the hyacinthpotatoeven the dahliaetc.; 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. The
whole organisation seems to have become plasticand departs in a slight
degree from that of the parental type.

Any variation which is not inherited is unimportant for us. But the number
and diversity of inheritable deviations of structureboth those of slight
and those of considerable physiological importanceare endless. Dr.
Prosper Lucas' treatisein two large volumesis the fullest and the best
on this subject. No breeder doubts how strong is the tendency to
inheritance; that like produces like is his fundamental belief: doubts
have been thrown on this principle only by theoretical writers. When any
deviation of structure often appearsand we see it in the father and
childwe cannot tell whether it may not be due to the same cause having
acted on both; but when among individualsapparently exposed to the same
conditionsany very rare deviationdue to some extraordinary combination
of circumstancesappears in the parent--sayonce among several million
individuals--and it reappears in the childthe mere doctrine of chances
almost compels us to attribute its reappearance to inheritance. Every one
must have heard of cases of albinismprickly skinhairy bodiesetc.
appearing in several members of the same family. If strange and rare
deviations of structure are truly inheritedless strange and commoner
deviations may be freely admitted to be inheritable. Perhaps the correct
way of viewing the whole subject would beto look at the inheritance of
every character whatever as the ruleand non-inheritance as the anomaly.

The laws governing inheritance are for the most part unknown; no one can
say why the same peculiarity in different individuals of the same species
or in different speciesis sometimes inherited and sometimes not so; why
the child often reverts in certain characteristics to its grandfather or
grandmother or more remote ancestor; why a peculiarity is often transmitted
from one sex to both sexesor to one sex alonemore commonly but not
exclusively to the like sex. It is a fact of some importance to usthat
peculiarities appearing in the males of our domestic breeds are often
transmittedeither exclusively or in a much greater degreeto the males
alone. A much more important rulewhich I think may be trustedis that
at whatever period of life a peculiarity first appearsit tends to
reappear in the offspring at a corresponding agethough sometimes earlier.
In many cases this could not be otherwise; thus the inherited peculiarities
in the horns of cattle could appear only in the offspring when nearly

mature; peculiarities in the silk-worm are known to appear at the
corresponding caterpillar or cocoon stage. But hereditary diseases and
some other facts make me believe that the rule has a wider extensionand
thatwhen there is no apparent reason why a peculiarity should appear at
any particular ageyet that it does tend to appear in the offspring at the
same period at which it first appeared in the parent. I believe this rule
to be of the highest importance in explaining the laws of embryology.
These remarks are of course confined to the first APPEARANCE of the
peculiarityand not to the primary cause which may have acted on the
ovules or on the male element; in nearly the same manner as the increased
length of the horns in the offspring from a short-horned cow by a
long-horned bullthough appearing late in lifeis clearly due to the male

Having alluded to the subject of reversionI may here refer to a statement
often made by naturalists--namelythat our domestic varietieswhen run
wildgradually but invariably revert in character to their aboriginal
stocks. Hence it has been argued that no deductions can be drawn from
domestic races to species in a state of nature. I have in vain endeavoured
to discover on what decisive facts the above statement has so often and so
boldly been made. 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. In many cases we do not
know what the aboriginal stock wasand so could not tell whether or not
nearly perfect reversion had ensued. It would be necessaryin order to
prevent the effects of intercrossingthat only a single variety should be
turned loose in its new home. Neverthelessas our varieties certainly do
occasionally revert in some of their characters to ancestral formsit
seems to me not improbable that if we could succeed in naturalisingor
were to cultivateduring many generationsthe several racesfor
instanceof the cabbagein very poor soil--in which casehoweversome
effect would have to be attributed to the DEFINITE action of the poor soil
--that they wouldto a large extentor even whollyrevert to the wild
aboriginal stock. Whether or not the experiment would succeed is not of
great importance for our line of argument; for by the experiment itself the
conditions of life are changed. If it could be shown that our domestic
varieties manifested a strong tendency to reversion--that isto lose their
acquired characterswhile kept under the same conditions and while kept in
a considerable bodyso that free intercrossing might checkby blending
togetherany slight deviations in their structurein such caseI grant
that we could deduce nothing from domestic varieties in regard to species.
But there is not a shadow of evidence in favour of this view: to assert
that we could not breed our cart and race-horseslong and short-horned
cattleand poultry of various breedsand esculent vegetablesfor an
unlimited number of generationswould be opposed to all experience.


When we look to the hereditary varieties or races of our domestic animals
and plantsand compare them with closely allied specieswe generally
perceive in each domestic raceas already remarkedless uniformity of
character than in true species. Domestic races often have a somewhat
monstrous character; by which I meanthatalthough differing from each
other and from other species of the same genusin several trifling
respectsthey often differ in an extreme degree in some one partboth
when compared one with anotherand more especially when compared with the
species under nature to which they are nearest allied. With these
exceptions (and with that of the perfect fertility of varieties when
crossed--a subject hereafter to be discussed)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 naturebut the differences in most
cases are less in degree. This must be admitted as truefor the domestic

races of many animals and plants have been ranked by some competent judges
as the descendants of aboriginally distinct speciesand by other competent
judges as mere varieties. If any well marked distinction existed between a
domestic race and a speciesthis source of doubt would not so perpetually
recur. It has often been stated that domestic races do not differ from
each other in characters of generic value. It can be shown that this
statement is not correct; but naturalists differ much in determining what
characters are of generic value; all such valuations being at present
empirical. When it is explained how genera originate under natureit will
be seen that we have no right to expect often to find a generic amount of
difference in our domesticated races.

In attempting to estimate the amount of structural difference between
allied domestic raceswe are soon involved in doubtfrom not knowing
whether they are descended from one or several parent species. This point
if it could be cleared upwould be interesting; iffor instanceit could
be shown that the greyhoundbloodhoundterrierspaniel and bull-dog
which we all know propagate their kind trulywere the offspring of any
single speciesthen such facts would have great weight in making us doubt
about the immutability of the many closely allied natural species--for
instanceof the many foxes--inhabiting the different quarters of the
world. I do not believeas we shall presently seethat the whole amount
of difference between the several breeds of the dog has been produced under
domestication; I believe that a small part of the difference is due to
their being descended from distinct species. In the case of strongly
marked races of some other domesticated speciesthere is presumptive or
even strong evidence that all are descended from a single wild stock.

It has often been assumed that man has chosen for domestication animals and
plants having an extraordinary inherent tendency to varyand likewise to
withstand diverse climates. I do not dispute that these capacities have
added largely to the value of most of our domesticated productions; but how
could a savage possibly knowwhen he first tamed an animalwhether it
would vary in succeeding generationsand whether it would endure other
climates? Has the little variability of the ass and gooseor the small
power of endurance of warmth by the reindeeror of cold by the common
camelprevented their domestication? I cannot doubt that if other animals
and plantsequal in number to our domesticated productionsand belonging
to equally diverse classes and countrieswere taken from a state of
natureand could be made to breed for an equal number of generations under
domesticationthey would on an average vary as largely as the parent
species of our existing domesticated productions have varied.

In the case of most of our anciently domesticated animals and plantsit is
not possible to come to any definite conclusionwhether they are descended
from one or several wild species. The argument mainly relied on by those
who believe in the multiple origin of our domestic animals isthat we find
in the most ancient timeson the monuments of Egyptand in the lakehabitations
of Switzerlandmuch diversity in the breeds; and that some of
these ancient breeds closely resembleor are even identical withthose
still existing. But this only throws far backward the history of
civilisationand shows that animals were domesticated at a much earlier
period than has hitherto been supposed. The lake-inhabitants of
Switzerland cultivated several kinds of wheat and barleythe peathe
poppy for oil and flax; and they possessed several domesticated animals.
They also carried on commerce with other nations. All this clearly shows
as Heer has remarkedthat they had at this early age progressed
considerably in civilisation; and this again implies a long continued
previous period of less advanced civilisationduring which the
domesticated animalskept by different tribes in different districts
might have varied and given rise to distinct races. Since the discovery of
flint tools in the superficial formations of many parts of the worldall
geologists believe that barbarian men existed at an enormously remote
period; and we know that at the present day there is hardly a tribe so

barbarous as not to have domesticated at least the dog.

The origin of most of our domestic animals will probably forever remain
vague. But I may here state thatlooking to the domestic dogs of the
whole worldI haveafter a laborious collection of all known factscome
to the conclusion that several wild species of Canidae have been tamedand
that their bloodin some cases mingled togetherflows in the veins of our
domestic breeds. In regard to sheep and goats I can form no decided
opinion. From facts communicated to me by Mr. Blython the habitsvoice
constitution and structure of the humped Indian cattleit is almost
certain that they are descended from a different aboriginal stock from our
European cattle; and some competent judges believe that these latter have
had two or three wild progenitorswhether or not these deserve to be
called species. This conclusionas well as that of the specific
distinction between the humped and common cattlemayindeedbe looked
upon as established by the admirable researches of Professor Rutimeyer.
With respect to horsesfrom reasons which I cannot here giveI am
doubtfully inclined to believein opposition to several authorsthat all
the races belong to the same species. Having kept nearly all the English
breeds of the fowl alivehaving bred and crossed themand examined their
skeletonsit appears to me almost certain that all are the descendants of
the wild Indian fowlGallus bankiva; and this is the conclusion of Mr.
Blythand of others who have studied this bird in India. In regard to
ducks and rabbitssome breeds of which differ much from each otherthe
evidence is clear that they are all descended from the common duck and wild

The doctrine of the origin of our several domestic races from several
aboriginal stockshas been carried to an absurd extreme by some authors.
They believe that every race which breeds truelet the distinctive
characters be ever so slighthas had its wild prototype. At this rate
there must have existed at least a score of species of wild cattleas many
sheepand several goatsin Europe aloneand several even within Great
Britain. 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 mammaland France but few distinct from
those of Germanyand so with HungarySpainetc.but that each of these
kingdoms possesses several peculiar breeds of cattlesheepetc.we must
admit that many domestic breeds must have originated in Europe; for whence
otherwise could they have been derived? So it is in India. Even in the
case of the breeds of the domestic dog throughout the worldwhich I admit
are descended from several wild speciesit cannot be doubted that there
has been an immense amount of inherited variation; for who will believe
that animals closely resembling the Italian greyhoundthe bloodhoundthe
bull-dogpug-dogor Blenheim spanieletc.--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; but by crossing we can only get forms in some degree
intermediate between their parents; and if we account for our several
domestic races by this processwe must admit the former existence of the
most extreme formsas the Italian greyhoundbloodhoundbull-dogetc.
in the wild state. Moreoverthe possibility of making distinct races by
crossing has been greatly exaggerated. 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; but to
obtain a race intermediate between two quite distinct races would be very
difficult. Sir J. Sebright expressly experimented with this object and
failed. The offspring from the first cross between two pure breeds is
tolerably and sometimes (as I have found with pigeons) quite uniform in
characterand every thing seems simple enough; but when these mongrels are
crossed one with another for several generationshardly two of them are
alikeand then the difficulty of the task becomes manifest.


Believing that it is always best to study some special groupI haveafter
deliberationtaken up domestic pigeons. I have kept every breed which I
could purchase or obtainand have been most kindly favoured with skins
from several quarters of the worldmore especially by the Hon. W. Elliot
from Indiaand by the Hon. C. Murray from Persia. Many treatises in
different languages have been published on pigeonsand some of them are
very importantas being of considerable antiquity. I have associated with
several eminent fanciersand have been permitted to join two of the London
Pigeon Clubs. The diversity of the breeds is something astonishing.
Compare the English carrier and the short-faced tumblerand see the
wonderful difference in their beaksentailing corresponding differences in
their skulls. The carriermore especially the male birdis also
remarkable from the wonderful development of the carunculated skin about
the headand this is accompanied by greatly elongated eyelidsvery large
external orifices to the nostrilsand a wide gape of mouth. The
short-faced tumbler has a beak in outline almost like that of a finch; and
the common tumbler has the singular inherited habit of flying at a great
height in a compact flockand tumbling in the air head over heels. The
runt is a bird of great sizewith longmassive beak and large feet; some
of the sub-breeds of runts have very long necksothers very long wings and
tailsothers singularly short tails. The barb is allied to the carrier
butinstead of a long beakhas a very short and broad one. The pouter
has a much elongated bodywingsand legs; and its enormously developed
cropwhich it glories in inflatingmay well excite astonishment and even
laughter. The turbit has a short and conical beakwith a line of reversed
feathers down the breast; and it has the habit of continually expanding
slightlythe upper part of the oesophagus. The Jacobin has the feathers
so much reversed along the back of the neck that they form a hoodand it
hasproportionally to its sizeelongated wing and tail feathers. The
trumpeter and laugheras their names expressutter a very different coo
from the other breeds. The fantail has thirty or even forty tail-feathers
instead of twelve or fourteenthe 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. Several other less distinct breeds might be specified.

In the skeletons of the several breedsthe development of the bones of the
facein length and breadth and curvaturediffers enormously. The shape
as well as the breadth and length of the ramus of the lower jawvaries in
a highly remarkable manner. The caudal and sacral vertebrae vary in
number; as does the number of the ribstogether with their relative
breadth and the presence of processes. The size and shape of the apertures
in the sternum are highly variable; so is the degree of divergence and
relative size of the two arms of the furcula. The proportional width of
the gape of mouththe proportional length of the eyelidsof the orifice
of the nostrilsof the tongue (not always in strict correlation with the
length of beak)the size of the crop and of the upper part of the
oesophagus; the development and abortion of the oil-gland; the number of
the primary wing and caudal feathers; the relative length of the wing and
tail to each other and to the body; the relative length of the leg and
foot; the number of scutellae on the toesthe development of skin between
the toesare all points of structure which are variable. The period at
which the perfect plumage is acquired variesas does the state of the down
with which the nestling birds are clothed when hatched. The shape and size
of the eggs vary. The manner of flightand in some breeds the voice and
dispositiondiffer remarkably. Lastlyin certain breedsthe males and
females have come to differ in a slight degree from each other.

Altogether at least a score of pigeons might be chosenwhichif shown to
an ornithologistand he were told that they were wild birdswould
certainly be ranked by him as well-defined species. MoreoverI do not
believe that any ornithologist would in this case place the English
carrierthe short-faced tumblerthe runtthe barbpouterand fantail

in the same genus; more especially as in each of these breeds several
truly-inherited sub-breedsor speciesas he would call themcould be
shown him.

Great as are the differences between the breeds of the pigeonI am fully
convinced that the common opinion of naturalists is correctnamelythat
all are descended from the rock-pigeon (Columba livia)including under
this term several geographical races or sub-specieswhich differ from each
other in the most trifling respects. As several of the reasons which have
led me to this belief are in some degree applicable in other casesI will
here briefly give them. If the several breeds are not varietiesand have
not proceeded from the rock-pigeonthey must have descended from at least
seven or eight aboriginal stocks; for it is impossible to make the present
domestic breeds by the crossing of any lesser number: howfor instance
could a pouter be produced by crossing two breeds unless one of the
parent-stocks possessed the characteristic enormous crop? The supposed
aboriginal stocks must all have been rock-pigeonsthat isthey did not
breed or willingly perch on trees. But besides C. liviawith its
geographical sub-speciesonly two or three other species of rock-pigeons
are known; and these have not any of the characters of the domestic breeds.
Hence the supposed aboriginal stocks must either still exist in the
countries where they were originally domesticatedand yet be unknown to
ornithologists; and thisconsidering their sizehabits and remarkable
charactersseems improbable; or they must have become extinct in the wild
state. But birds breeding on precipicesand good flyersare unlikely to
be exterminated; and the common rock-pigeonwhich has the same habits with
the domestic breedshas not been exterminated even on several of the
smaller British isletsor on the shores of the Mediterranean. Hence the
supposed extermination of so many species having similar habits with the
rock-pigeon seems a very rash assumption. Moreoverthe several
above-named domesticated breeds have been transported to all parts of the
worldandthereforesome of them must have been carried back again into
their native country; but not one has become wild or feralthough the
dovecot-pigeonwhich is the rock-pigeon in a very slightly altered state
has become feral in several places. Againall recent experience shows
that it is difficult to get wild animals to breed freely under
domestication; yet on the hypothesis of the multiple origin of our pigeons
it must be assumed that at least seven or eight species were so thoroughly
domesticated in ancient times by half-civilized manas to be quite
prolific under confinement.

An argument of great weightand applicable in several other casesis
that the above-specified breedsthough agreeing generally with the wild
rock-pigeon in constitutionhabitsvoicecolouringand in most parts of
their structureyet are certainly highly abnormal in other parts; we may
look in vain through the whole great family of Columbidae for a beak like
that of the English carrieror that of the short-faced tumbleror barb;
for reversed feathers like those of the Jacobin; for a crop like that of
the pouter; for tail-feathers like those of the fantail. Hence it must be
assumednot only that half-civilized man succeeded in thoroughly
domesticating several speciesbut that he intentionally or by chance
picked out extraordinarily abnormal species; and furtherthat these very
species have since all become extinct or unknown. So many strange
contingencies are improbable in the highest degree.

Some facts in regard to the colouring of pigeons well deserve
consideration. The rock-pigeon is of a slaty-bluewith white loins; but
the Indian sub-speciesC. intermedia of Stricklandhas this part bluish.
The tail has a terminal dark barwith the outer feathers externally edged
at the base with white. The wings have two black bars. Some semi-domestic
breedsand some truly wild breedshavebesides the two black barsthe
wings chequered with black. These several marks do not occur together in
any other species of the whole family. Nowin every one of the domestic
breedstaking thoroughly well-bred birdsall the above markseven to the

white edging of the outer tail-featherssometimes concur perfectly
developed. Moreoverwhen birds belonging to two or more distinct breeds
are crossednone of which are blue or have any of the above-specified
marksthe mongrel offspring are very apt suddenly to acquire these
characters. To give one instance out of several which I have observed:
crossed some white fantailswhich breed very truewith some black barbs-and
it so happens that blue varieties of barbs are so rare that I never
heard of an instance in England; and the mongrels were blackbrown and
mottled. I also crossed a barb with a spotwhich is a white bird with a
red tail and red spot on the foreheadand which notoriously breeds very
true; the mongrels were dusky and mottled. I then crossed one of the
mongrel barb-fantails with a mongrel barb-spotand they produced a bird of
as beautiful a blue colourwith the white loinsdouble black wing-bar
and barred and white-edged tail-feathersas any wild rock-pigeon! We can
understand these factson the well-known principle of reversion to
ancestral charactersif all the domestic breeds are descended from the
rock-pigeon. But if we deny thiswe must make one of the two following
highly improbable suppositions. Eitherfirstthat all the several
imagined aboriginal stocks were coloured and marked like the rock-pigeon
although no other existing species is thus coloured and markedso that in
each separate breed there might be a tendency to revert to the very same
colours and markings. Orsecondlythat each breedeven the puresthas
within a dozenor at most within a scoreof generationsbeen crossed by
the rock-pigeon: I say within a dozen or twenty generationsfor no
instance is known of crossed descendants reverting to an ancestor of
foreign bloodremoved by a greater number of generations. 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 lessas in each
succeeding generation there will be less of the foreign blood; but when
there has been no crossand there is a tendency in the breed to revert to
a character which was lost during some former generationthis tendency
for all that we can see to the contrarymay be transmitted undiminished
for an indefinite number of generations. These two distinct cases of
reversion are often confounded together by those who have written on

Lastlythe hybrids or mongrels from between all the breeds of the pigeon
are perfectly fertileas I can state from my own observationspurposely
madeon the most distinct breeds. Nowhardly any cases have been
ascertained with certainty of hybrids from two quite distinct species of
animals being perfectly fertile. Some authors believe that long-continued
domestication eliminates this strong tendency to sterility in species.
>From the history of the dogand of some other domestic animalsthis
conclusion is probably quite correctif applied to species closely related
to each other. But to extend it so far as to suppose that species
aboriginally as distinct as carrierstumblerspoutersand fantails now
areshould yield offspring perfectly fertileinter seseems to me rash
in the extreme.

>From these several reasonsnamelythe 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
stateand their not having become anywhere feral--these species presenting
certain very abnormal charactersas compared with all other Columbidae
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
both when kept pure and when crossed--and lastlythe mongrel offspring
being perfectly fertile--from these several reasonstaken togetherwe may
safely conclude that all our domestic breeds are descended from the rockpigeon
or Columba livia with its geographical sub-species.

In favour of this viewI may addfirstlythat the wild C. livia has been
found capable of domestication in Europe and in India; and that it agrees
in habits and in a great number of points of structure with all the

domestic breeds. Secondlythat although an English carrier or a
short-faced tumbler differs immensely in certain characters from the
rock-pigeonyet that by comparing the several sub-breeds of these two
racesmore especially those brought from distant countrieswe can make
between them and the rock-pigeonan almost perfect series; so we can in
some other casesbut not with all the breeds. Thirdlythose characters
which are mainly distinctive of each breed are in each eminently variable
for instancethe wattle and length of beak of the carrierthe shortness
of that of the tumblerand the number of tail-feathers in the fantail; and
the explanation of this fact will be obvious when we treat of selection.
Fourthlypigeons have been watched and tended with the utmost careand
loved by many people. They have been domesticated for thousands of years
in several quarters of the world; the earliest known record of pigeons is
in the fifth Aegyptian dynastyabout 3000 was pointed out to me
by Professor Lepsius; but Mr. Birch informs me that pigeons are given in a
bill of fare in the previous dynasty. In the time of the Romansas we
hear from Plinyimmense prices were given for pigeons; "naythey are come
to this passthat they can reckon up their pedigree and race." Pigeons
were much valued by Akber Khan in Indiaabout the year 1600; never less
than 20000 pigeons were taken with the court. "The monarchs of Iran and
Turan sent him some very rare birds;" andcontinues the courtly historian
His Majesty, by crossing the breeds, which method was never practised
before, has improved them astonishingly.About this same period the Dutch
were as eager about pigeons as were the old Romans. The paramount
importance of these considerations in explaining the immense amount of
variation which pigeons have undergonewill likewise be obvious when we
treat of selection. We shall thenalsosee how it is that the several
breeds so often have a somewhat monstrous character. It is also a most
favourable circumstance for the production of distinct breedsthat male
and female pigeons can be easily mated for life; and thus different breeds
can be kept together in the same aviary.

I have discussed the probable origin of domestic pigeons at someyet quite
insufficientlength; because when I first kept pigeons and watched the
several kindswell knowing how truly they breedI felt fully as much
difficulty in believing that since they had been domesticated they had all
proceeded from a common parentas any naturalist could in coming to a
similar conclusion in regard to the many species of finchesor other
groups of birdsin nature. One circumstance has struck me much; namely
that nearly all the breeders of the various domestic animals and the
cultivators of plantswith whom I have conversedor whose treatises I
have readare firmly convinced that the several breeds to which each has
attendedare descended from so many aboriginally distinct species. Ask
as I have askeda celebrated raiser of Hereford cattlewhether his cattle
might not have descended from Long-hornsor both from a common parentstock
and he will laugh you to scorn. I have never met a pigeonor
poultryor duckor rabbit fancierwho was not fully convinced that each
main breed was descended from a distinct species. Van Monsin his
treatise on pears and applesshows how utterly he disbelieves that the
several sortsfor instance a Ribston-pippin or Codlin-applecould ever
have proceeded from the seeds of the same tree. Innumerable other examples
could be given. The explanationI thinkis 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 slightlyfor they
win their prizes by selecting such slight differencesyet they ignore all
general argumentsand refuse to sum up in their minds slight differences
accumulated during many successive generations. May not those naturalists
whoknowing far less of the laws of inheritance than does the breederand
knowing no more than he does of the intermediate links in the long lines of
descentyet admit that many of our domestic races are descended from the
same parents--may they not learn a lesson of cautionwhen they deride the
idea of species in a state of nature being lineal descendants of other


Let us now briefly consider the steps by which domestic races have been
producedeither from one or from several allied species. Some effect may
be attributed to the direct and definite action of the external conditions
of lifeand some to habit; but he would be a bold man who would account by
such agencies for the differences between a dray and race-horsea
greyhound and bloodhounda carrier and tumbler pigeon. One of the most
remarkable features in our domesticated races is that we see in them
adaptationnot indeed to the animal's or plant's own goodbut to man's
use or fancy. Some variations useful to him have probably arisen suddenly
or by one step; many botanistsfor instancebelieve that the fuller's
teaselwith its hookswhich can not be rivalled by any mechanical
contrivanceis only a variety of the wild Dipsacus; and this amount of
change may have suddenly arisen in a seedling. So it has probably been
with the turnspit dog; and this is known to have been the case with the
ancon sheep. But when we compare the dray-horse and race-horsethe
dromedary and camelthe various breeds of sheep fitted either for
cultivated land or mountain pasturewith the wool of one breed good for
one purposeand that of another breed for another purpose; when we compare
the many breeds of dogseach good for man in different ways; when we
compare the game-cockso pertinacious in battlewith other breeds so
little quarrelsomewith "everlasting layers" which never desire to sit
and with the bantam so small and elegant; when we compare the host of
agriculturalculinaryorchardand flower-garden races of plantsmost
useful to man at different seasons and for different purposesor so
beautiful in his eyeswe mustI thinklook further than to mere
variability. We can not suppose that all the breeds were suddenly produced
as perfect and as useful as we now see them; indeedin many caseswe know
that this has not been their history. The key is man's power of
accumulative selection: nature gives successive variations; man adds them
up in certain directions useful to him. In this sense he may be said to
have made for himself useful breeds.

The great power of this principle of selection is not hypothetical. It is
certain that several of our eminent breeders haveeven within a single
lifetimemodified to a large extent their breeds of cattle and sheep. In
order fully to realise what they have done it is almost necessary to read
several of the many treatises devoted to this subjectand to inspect the
animals. Breeders habitually speak of an animal's organisation as
something plasticwhich they can model almost as they please. If I had
space I could quote numerous passages to this effect from highly competent
authorities. Youattwho was probably better acquainted with the works of
agriculturalists than almost any other individualand who was himself a
very good judge of animalsspeaks of the principle of selection as "that
which enables the agriculturistnot only to modify the character of his
flockbut to change it altogether. It is the magician's wandby means of
which he may summon into life whatever form and mould he pleases." Lord
Somervillespeaking of what breeders have done for sheepsays: "It would
seem as if they had chalked out upon a wall a form perfect in itselfand
then had given it existence." In Saxony the importance of the principle of
selection in regard to merino sheep is so fully recognisedthat men follow
it as a trade: the sheep are placed on a table and are studiedlike a
picture by a connoisseur; this is done three times at intervals of months
and the sheep are each time marked and classedso that the very best may
ultimately be selected for breeding.

What English breeders have actually effected is proved by the enormous
prices given for animals with a good pedigree; and these have been exported
to almost every quarter of the world. The improvement is by no means
generally due to crossing different breeds; all the best breeders are
strongly opposed to this practiceexcept sometimes among closely allied
sub-breeds. And when a cross has been madethe closest selection is far
more indispensable even than in ordinary cases. If selection consisted

merely in separating some very distinct variety and breeding from itthe
principle would be so obvious as hardly to be worth notice; but its
importance consists in the great effect produced by the accumulation in one
directionduring successive generationsof differences absolutely
inappreciable by an uneducated eye--differences which I for one have vainly
attempted to appreciate. Not one man in a thousand has accuracy of eye and
judgment sufficient to become an eminent breeder. If gifted with these
qualitiesand he studies his subject for yearsand devotes his lifetime
to it with indomitable perseverancehe will succeedand may make great
improvements; if he wants any of these qualitieshe will assuredly fail.
Few would readily believe in the natural capacity and years of practice
requisite to become even a skilful pigeon-fancier.

The same principles are followed by horticulturists; but the variations are
here often more abrupt. No one supposes that our choicest productions have
been produced by a single variation from the aboriginal stock. We have
proofs that this is not so in several cases in which exact records have
been kept; thusto give a very trifling instancethe steadily increasing
size of the common gooseberry may be quoted. We see an astonishing
improvement in many florists' flowerswhen the flowers of the present day
are compared with drawings made only twenty or thirty years ago. When a
race of plants is once pretty well establishedthe seed-raisers do not
pick out the best plantsbut merely go over their seed-bedsand pull up
the "rogues as they call the plants that deviate from the proper
standard. With animals this kind of selection is, in fact, likewise
followed; for hardly any one is so careless as to breed from his worst

In regard to plants, there is another means of observing the accumulated
effects of selection--namely, by comparing the diversity of flowers in the
different varieties of the same species in the flower-garden; the diversity
of leaves, pods, or tubers, or whatever part is valued, in the
kitchen-garden, in comparison with the flowers of the same varieties; and
the diversity of fruit of the same species in the orchard, in comparison
with the leaves and flowers of the same set of varieties. See how
different the leaves of the cabbage are, and how extremely alike the
flowers; how unlike the flowers of the heartsease are, and how alike the
leaves; how much the fruit of the different kinds of gooseberries differ in
size, colour, shape, and hairiness, and yet the flowers present very slight
differences. It is not that the varieties which differ largely in some one
point do not differ at all in other points; this is hardly ever--I speak
after careful observation--perhaps never, the case. The law of correlated
variation, the importance of which should never be overlooked, will ensure
some differences; but, as a general rule, it cannot be doubted that the
continued selection of slight variations, either in the leaves, the
flowers, or the fruit, will produce races differing from each other chiefly
in these characters.

It may be objected that the principle of selection has been reduced to
methodical practice for scarcely more than three-quarters of a century; it
has certainly been more attended to of late years, and many treatises have
been published on the subject; and the result has been, in a corresponding
degree, rapid and important. But it is very far from true that the
principle is a modern discovery. I could give several references to works
of high antiquity, in which the full importance of the principle is
acknowledged. In rude and barbarous periods of English history choice
animals were often imported, and laws were passed to prevent their
exportation: the destruction of horses under a certain size was ordered,
and this may be compared to the roguing" of plants by nurserymen. The
principle of selection I find distinctly given in an ancient Chinese
encyclopaedia. Explicit rules are laid down by some of the Roman classical
writers. From passages in Genesisit is clear that the colour of domestic
animals was at that early period attended to. Savages now sometimes cross
their dogs with wild canine animalsto improve the breedand they

formerly did soas is attested by passages in Pliny. The savages in South
Africa match their draught cattle by colouras do some of the Esquimaux
their teams of dogs. Livingstone states that good domestic breeds are
highly valued by the negroes in the interior of Africa who have not
associated with Europeans. Some of these facts do not show actual
selectionbut they show that the breeding of domestic animals was
carefully attended to in ancient timesand is now attended to by the
lowest savages. It wouldindeedhave been a strange facthad attention
not been paid to breedingfor the inheritance of good and bad qualities is
so obvious.


At the present timeeminent breeders try by methodical selectionwith a
distinct object in viewto make a new strain or sub-breedsuperior to
anything of the kind in the country. Butfor our purposea form of
selectionwhich may be called unconsciousand which results from every
one trying to possess and breed from the best individual animalsis more
important. Thusa man who intends keeping pointers naturally tries to get
as good dogs as he canand afterwards breeds from his own best dogsbut
he has no wish or expectation of permanently altering the breed.
Nevertheless we may infer that this processcontinued during centuries
would improve and modify any breedin the same way as BakewellCollins this very same processonly carried on more methodicallydid
greatly modifyeven during their lifetimesthe forms and qualities of
their cattle. 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 agowhich may serve for comparison. In some
caseshoweverunchangedor but little changedindividuals of the same
breed exist in less civilised districtswhere the breed has been less
improved. There is reason to believe that King Charles' spaniel has been
unconsciously modified to a large extent since the time of that monarch.
Some highly competent authorities are convinced that the setter is directly
derived from the spanieland has probably been slowly altered from it. It
is known that the English pointer has been greatly changed within the last
centuryand in this case the change hasit is believedbeen chiefly
effected by crosses with the foxhound; but what concerns us isthat the
change has been effected unconsciously and graduallyand yet so
effectually thatthough the old Spanish pointer certainly came from Spain
Mr. Borrow has not seenas I am informed by himany native dog in Spain
like our pointer.

By a similar process of selectionand by careful trainingEnglish racehorses
have come to surpass in fleetness and size the parent Arabsso that
the latterby the regulations for the Goodwood Racesare favoured in the
weights which they carry. Lord Spencer and others have shown how the
cattle of England have increased in weight and in early maturitycompared
with the stock formerly kept in this country. By comparing the accounts
given in various old treatises of the former and present state of carrier
and tumbler pigeons in BritainIndiaand Persiawe can trace the stages
through which they have insensibly passedand come to differ so greatly
from the rock-pigeon.

Youatt gives an excellent illustration of the effects of a course of
selection which may be considered as unconsciousin so far that the
breeders could never have expectedor even wishedto produce the result
which ensued--namelythe production of the distinct strains. The two
flocks of Leicester sheep kept by Mr. Buckley and Mr. Burgessas Mr.
Youatt remarksHave been purely bred from the original stock of Mr.
Bakewell for upwards of fifty years. 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.
Bakewell's flock, 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.

If there exist savages so barbarous as never to think of the inherited
character of the offspring of their domestic animalsyet any one animal
particularly useful to themfor any special purposewould be carefully
preserved during famines and other accidentsto which savages are so
liableand such choice animals would thus generally leave more offspring
than the inferior ones; so that in this case there would be a kind of
unconscious selection going on. We see the value set on animals even by
the barbarians of Tierra del Fuegoby their killing and devouring their
old womenin times of dearthas of less value than their dogs.

In plants the same gradual process of improvement through the occasional
preservation of the best individualswhether or not sufficiently distinct
to be ranked at their first appearance as distinct varietiesand whether
or not two or more species or races have become blended together by
crossingmay plainly be recognised in the increased size and beauty which
we now see in the varieties of the heartseaserosepelargoniumdahlia
and other plantswhen compared with the older varieties or with their
parent-stocks. No one would ever expect to get a first-rate heartsease or
dahlia from the seed of a wild plant. No one would expect to raise a
first-rate melting pear from the seed of a wild pearthough he might
succeed from a poor seedling growing wildif it had come from a
garden-stock. The pearthough cultivated in classical timesappears
from Pliny's descriptionto have been a fruit of very inferior quality. 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; but the art has been simpleandas far as the final result is
concernedhas been followed almost unconsciously. It has consisted in
always cultivating the best known varietysowing its seedsandwhen a
slightly better variety chanced to appearselecting itand so onwards.
But the gardeners of the classical periodwho cultivated the best pears
which they could procurenever thought what splendid fruit we should eat;
though we owe our excellent fruit in some small degree to their having
naturally chosen and preserved the best varieties they could anywhere find.

A large amount of changethus slowly and unconsciously accumulated
explainsas I believethe well-known factthat in a number of cases we
cannot recogniseand therefore do not knowthe wild parent-stocks of the
plants which have been longest cultivated in our flower and kitchen
gardens. 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
manwe can understand how it is that neither Australiathe Cape of Good
Hopenor any other region inhabited by quite uncivilised manhas afforded
us a single plant worth culture. It is not that these countriesso rich
in speciesdo not by a strange chance possess the aboriginal stocks of any
useful plantsbut 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.

In regard to the domestic animals kept by uncivilised manit should not be
overlooked that they almost always have to struggle for their own foodat
least during certain seasons. And in two countries very differently
circumstancedindividuals of the same specieshaving slightly different
constitutions or structurewould often succeed better in the one country
than in the otherand thus by a process of "natural selection as will
hereafter be more fully explained, two sub-breeds might be formed. This,
perhaps, partly explains why the varieties kept by savages, as has been
remarked by some authors, have more of the character of true species than
the varieties kept in civilised countries.

On the view here given of the important part which selection by man has
played, it becomes at once obvious, how it is that our domestic races show
adaptation in their structure or in their habits to man's wants or fancies.

We can, I think, further understand the frequently abnormal character of
our domestic races, and likewise their differences being so great in
external characters, and relatively so slight in internal parts or organs.
Man can hardly select, or only with much difficulty, any deviation of
structure excepting such as is externally visible; and indeed he rarely
cares for what is internal. He can never act by selection, excepting on
variations which are first given to him in some slight degree by nature.
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, or a pouter till he
saw a pigeon with a crop of somewhat unusual size; and the more abnormal or
unusual any character was when it first appeared, the more likely it would
be to catch his attention. But to use such an expression as trying to make
a fantail is, I have no doubt, in most cases, utterly incorrect. The man
who first selected a pigeon with a slightly larger tail, never dreamed what
the descendants of that pigeon would become through long-continued, partly
unconscious and partly methodical, selection. Perhaps the parent bird of
all fantails had only fourteen tail-feathers somewhat expanded, like the
present Java fantail, or like individuals of other and distinct breeds, in
which as many as seventeen tail-feathers have been counted. 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, as it is not one of the points of the breed.

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, and it is in human nature to value any novelty, however
slight, in one's own possession. Nor must the value which would formerly
have been set on any slight differences in the individuals of the same
species, be judged of by the value which is now set on them, after several
breeds have fairly been established. It is known that with pigeons many
slight variations now occasionally appear, but these are rejected as faults
or deviations from the standard of perfection in each breed. The common
goose has not given rise to any marked varieties; hence the Toulouse and
the common breed, which differ only in colour, that most fleeting of
characters, have lately been exhibited as distinct at our poultry-shows.

These views appear to explain what has sometimes been noticed, namely, that
we know hardly anything about the origin or history of any of our domestic
breeds. But, in fact, a breed, like a dialect of a language, can hardly be
said to have a distinct origin. A man preserves and breeds from an
individual with some slight deviation of structure, or takes more care than
usual in matching his best animals, and thus improves them, and the
improved animals slowly spread in the immediate neighbourhood. But they
will as yet hardly have a distinct name, and from being only slightly
valued, their history will have been disregarded. When further improved by
the same slow and gradual process, they will spread more widely, and will
be recognised as something distinct and valuable, and will then probably
first receive a provincial name. In semi-civilised countries, with little
free communication, the spreading of a new sub-breed will be a slow
process. As soon as the points of value are once acknowledged, the
principle, as I have called it, of unconscious selection will always
tend--perhaps more at one period than at another, as the breed rises or
falls in fashion--perhaps more in one district than in another, according
to the state of civilisation of the inhabitants--slowly to add to the
characteristic features of the breed, whatever they may be. But the chance
will be infinitely small of any record having been preserved of such slow,
varying, and insensible changes.


I will now say a few words on the circumstances, favourable or the reverse,
to man's power of selection. A high degree of variability is obviously
favourable, as freely giving the materials for selection to work on; not
that mere individual differences are not amply sufficient, with extreme

care, to allow of the accumulation of a large amount of modification in
almost any desired direction. But as variations manifestly useful or
pleasing to man appear only occasionally, the chance of their appearance
will be much increased by a large number of individuals being kept. Hence
number is of the highest importance for success. On this principle
Marshall formerly remarked, with respect to the sheep of part of Yorkshire,
As they generally belong to poor peopleand are mostly IN SMALL LOTS
they never can be improved." On the other handnurserymenfrom keeping
large stocks of the same plantare generally far more successful than
amateurs in raising new and valuable varieties. A large number of
individuals of an animal or plant can be reared only where the conditions
for its propagation are favourable. When the individuals are scanty all
will be allowed to breedwhatever their quality may beand this will
effectually prevent selection. But probably the most important element is
that the animal or plant should be so highly valued by manthat the
closest attention is paid to even the slightest deviations in its qualities
or structure. Unless such attention be paid nothing can be effected. I
have seen it gravely remarkedthat it was most fortunate that the
strawberry began to vary just when gardeners began to attend to this plant.
No doubt the strawberry had always varied since it was cultivatedbut the
slight varieties had been neglected. As soonhoweveras gardeners picked
out individual plants with slightly largerearlieror better fruitand
raised seedlings from themand again picked out the best seedlings and
bred from themthen (with some aid by crossing distinct species) those
many admirable varieties of the strawberry were raised which have appeared
during the last half-century.

With animalsfacility in preventing crosses is an important element in the
formation of new races--at leastin a country which is already stocked
with other races. In this respect enclosure of the land plays a part.
Wandering savages or the inhabitants of open plains rarely possess more
than one breed of the same species. Pigeons can be mated for lifeand
this is a great convenience to the fancierfor thus many races may be
improved and kept truethough mingled in the same aviary; and this
circumstance must have largely favoured the formation of new breeds.
PigeonsI may addcan be propagated in great numbers and at a very quick
rateand inferior birds may be freely rejectedas when killed they serve
for food. On the other handcatsfrom their nocturnal rambling habits
can not be easily matchedandalthough so much valued by women and
childrenwe rarely see a distinct breed long kept up; such breeds as we do
sometimes see are almost always imported from some other country. Although
I do not doubt that some domestic animals vary less than othersyet the
rarity or absence of distinct breeds of the catthe donkeypeacock
gooseetc.may be attributed in main part to selection not having been
brought into play: in catsfrom the difficulty in pairing them; in
donkeysfrom only a few being kept by poor peopleand little attention
paid to their breeding; for recently in certain parts of Spain and of the
United States this animal has been surprisingly modified and improved by
careful selection; in peacocksfrom not being very easily reared and a
large stock not kept; in geesefrom being valuable only for two purposes
food and feathersand more especially from no pleasure having been felt in
the display of distinct breeds; but the gooseunder the conditions to
which it is exposed when domesticatedseems to have a singularly
inflexible organisationthough it has varied to a slight extentas I have
elsewhere described.

Some authors have maintained that the amount of variation in our domestic
productions is soon reachedand can never afterward be exceeded. It would
be somewhat rash to assert that the limit has been attained in any one
case; for almost all our animals and plants have been greatly improved in
many ways within a recent period; and this implies variation. It would be
equally rash to assert that characters now increased to their utmost limit
could notafter remaining fixed for many centuriesagain vary under new
conditions of life. No doubtas Mr. Wallace has remarked with much truth

a limit will be at last reached. For instancethere must be a limit to
the fleetness of any terrestrial animalas this will be determined by the
friction to be overcomethe weight of the body to be carriedand the
power of contraction in the muscular fibres. But what concerns us is that
the domestic varieties of the same species differ from each other in almost
every characterwhich man has attended to and selectedmore than do the
distinct species of the same genera. Isidore Geoffroy St. Hilaire has
proved this in regard to sizeand so it is with colourand probably with
the length of hair. With respect to fleetnesswhich depends on many
bodily charactersEclipse was far fleeterand a dray-horse is comparably
strongerthan any two natural species belonging to the same genus. So
with plantsthe seeds of the different varieties of the bean or maize
probably differ more in size than do the seeds of the distinct species in
any one genus in the same two families. The same remark holds good in
regard to the fruit of the several varieties of the plumand still more
strongly with the melonas well as in many other analogous cases.

To sum up on the origin of our domestic races of animals and plants.
Changed conditions of life are of the highest importance in causing
variabilityboth by acting directly on the organisationand indirectly by
affecting the reproductive system. It is not probable that variability is
an inherent and necessary contingentunder all circumstances. The greater
or less force of inheritance and reversion determine whether variations
shall endure. Variability is governed by many unknown lawsof which
correlated growth is probably the most important. Somethingbut how much
we do not knowmay be attributed to the definite action of the conditions
of life. Someperhaps a greateffect may be attributed to the increased
use or disuse of parts. The final result is thus rendered infinitely
complex. In some cases the intercrossing of aboriginally distinct species
appears to have played an important part in the origin of our breeds. When
several breeds have once been formed in any countrytheir occasional
intercrossingwith the aid of selectionhasno doubtlargely aided in
the formation of new sub-breeds; but the importance of crossing has been
much exaggeratedboth in regard to animals and to those plants which are
propagated by seed. With plants which are temporarily propagated by
cuttingsbudsetc.the importance of crossing is immense; for the
cultivator may here disregard the extreme variability both of hybrids and
of mongrelsand the sterility of hybrids; but plants not propagated by
seed are of little importance to usfor their endurance is only temporary.
Over all these causes of changethe accumulative action of selection
whether applied methodically and quicklyor unconsciously and slowlybut
more efficientlyseems to have been the predominant power.



Variability -- Individual differences -- Doubtful species -- Wide ranging
much diffusedand common speciesvary 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 closelybut unequallyrelated to each otherand in having
restricted ranges.

Before applying the principles arrived at in the last chapter to organic
beings in a state of naturewe must briefly discuss whether these latter
are subject to any variation. To treat this subject properlya long
catalogue of dry facts ought to be given; but these I shall reserve for a
future work. Nor shall I here discuss the various definitions which have
been given of the term species. No one definition has satisfied all
naturalists; yet every naturalist knows vaguely what he means when he
speaks of a species. Generally the term includes the unknown element of a
distinct act of creation. The term "variety" is almost equally difficult

to define; but here community of descent is almost universally implied
though it can rarely be proved. We have also what are called
monstrosities; but they graduate into varieties. By a monstrosity I
presume is meant some considerable deviation of structuregenerally
injuriousor not useful to the species. Some authors use the term
variationin a technical senseas implying a modification directly due
to the physical conditions of life; and "variations" in this sense are
supposed not to be inherited; but who can say that the dwarfed condition of
shells in the brackish waters of the Balticor dwarfed plants on Alpine
summitsor the thicker fur of an animal from far northwardswould 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.

It may be doubted whether sudden and considerable deviations of structure
such as we occasionally see in our domestic productionsmore especially
with plantsare ever permanently propagated in a state of nature. 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 perfectas that a complex machine should have been
invented by man in a perfect state. Under domestication monstrosities
sometimes occur which resemble normal structures in widely different
animals. Thus pigs have occasionally been born with a sort of proboscis
and if any wild species of the same genus had naturally possessed a
proboscisit might have been argued that this had appeared as a
monstrosity; but I have as yet failed to findafter diligent searchcases
of monstrosities resembling normal structures in nearly allied formsand
these alone bear on the question. 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)as they occur rarely and singlytheir preservation would
depend on unusually favourable circumstances. They wouldalsoduring the
first and succeeding generations cross with the ordinary formand thus
their abnormal character would almost inevitably be lost. But I shall have
to return in a future chapter to the preservation and perpetuation of
single or occasional variations.


The many slight differences which appear in the offspring from the same
parentsor which it may be presumed have thus arisenfrom being observed
in the individuals of the same species inhabiting the same confined
localitymay be called individual differences. No one supposes that all
the individuals of the same species are cast in the same actual mould.
These individual differences are of the highest importance for usfor they
are often inheritedas must be familiar to every one; and they thus afford
materials for natural selection to act on and accumulatein the same
manner as man accumulates in any given direction individual differences in
his domesticated productions. These individual differences generally
affect what naturalists consider unimportant parts; but I could showby a
long catalogue of factsthat parts which must be called importantwhether
viewed under a physiological or classificatory point of viewsometimes
vary in the individuals of the same species. I am convinced that the most
experienced naturalist would be surprised at the number of the cases of
variabilityeven in important parts of structurewhich he could collect
on good authorityas I have collectedduring a course of years. It
should be remembered that systematists are far from being pleased at
finding variability in important charactersand that there are not many
men who will laboriously examine internal and important organsand compare
them in many specimens of the same species. It would never have been
expected that the branching of the main nerves close to the great central
ganglion of an insect would have been variable in the same species; it
might have been thought that changes of this nature could have been
effected only by slow degrees; yet Sir J. Lubbock has shown a degree of
variability in these main nerves in Coccuswhich may almost be compared to
the irregular branching of the stem of a tree. This philosophical

naturalistI may addhas also shown that the muscles in the larvae of
certain insects are far from uniform. Authors sometimes argue in a circle
when they state that important organs never vary; for these same authors
practically rank those parts as important (as some few naturalists have
honestly confessed) which do not vary; andunder this point of viewno
instance will ever be found of an important part varying; but under any
other point of view many instances assuredly can be given.

There is one point connected with individual differences which is extremely
perplexing: I refer to those genera which have been called "protean" or
polymorphic,in which species present an inordinate amount of variation.
With respect to many of these formshardly two naturalists agree whether
to rank them as species or as varieties. We may instance RubusRosaand
Hieracium among plantsseveral genera of insectsand of Brachiopod
shells. In most polymorphic genera some of the species have fixed and
definite characters. Genera which are polymorphic in one country seem to
bewith a few exceptionspolymorphic in other countriesand likewise
judging from Brachiopod shellsat former periods of time. These facts are
very perplexingfor they seem to show that this kind of variability is
independent of the conditions of life. I am inclined to suspect that we
seeat least in some of these polymorphic generavariations which are of
no service or disservice to the speciesand which consequently have not
been seized on and rendered definite by natural selectionas hereafter to
be explained.

Individuals of the same species often presentas is known to every one
great differences of structureindependently of variationas in the two
sexes of various animalsin the two or three castes of sterile females or
workers among insectsand in the immature and larval states of many of the
lower animals. There arealsocases of dimorphism and trimorphismboth
with animals and plants. ThusMr. Wallacewho has lately called
attention to the subjecthas shown that the females of certain species of
butterfliesin the Malayan Archipelagoregularly appear under two or even
three conspicuously distinct formsnot connected by intermediate
varieties. Fritz Muller has described analogous but more extraordinary
cases with the males of certain Brazilian Crustaceans: thusthe male of a
Tanais regularly occurs under two distinct forms; one of these has strong
and differently shaped pincersand the other has antennae much more
abundantly furnished with smelling-hairs. Although in most of these cases
the two or three formsboth with animals and plantsare not now connected
by intermediate gradationsit is possible that they were once thus
connected. Mr. Wallacefor instancedescribes a certain butterfly which
presents in the same island a great range of varieties connected by
intermediate linksand the extreme links of the chain closely resemble the
two forms of an allied dimorphic species inhabiting another part of the
Malay Archipelago. Thus also with antsthe several worker-castes are
generally quite distinct; but in some casesas we shall hereafter seethe
castes are connected together by finely graduated varieties. So it isas
I have myself observedwith some dimorphic plants. 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; and that an hermaphrodite plant should produce from the same seedcapsule
three distinct hermaphrodite formsbearing 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.


The forms which possess in some considerable degree the character of
speciesbut which are so closely similar to other formsor are so closely
linked to them by intermediate gradationsthat naturalists do not like to
rank them as distinct speciesare in several respects the most important

for us. We have every reason to believe that many of these doubtful and
closely allied forms have permanently retained their characters for a long
time; for as longas far as we knowas have good and true species.
Practicallywhen a naturalist can unite by means of intermediate links any
two formshe treats the one as a variety of the otherranking the most
commonbut sometimes the one first described as the speciesand the other
as the variety. But cases of great difficultywhich I will not here
enumeratesometimes arise in deciding whether or not to rank one form as a
variety of anothereven when they are closely connected by intermediate
links; nor will the commonly assumed hybrid nature of the intermediate
forms always remove the difficulty. In very many caseshoweverone form
is ranked as a variety of anothernot because the intermediate links have
actually been foundbut because analogy leads the observer to suppose
either that they do now somewhere existor may formerly have existed; and
here a wide door for the entry of doubt and conjecture is opened.

Hencein determining whether a form should be ranked as a species or a
varietythe opinion of naturalists having sound judgment and wide
experience seems the only guide to follow. We musthoweverin many
casesdecide by a majority of naturalistsfor few well-marked and
well-known varieties can be named which have not been ranked as species by
at least some competent judges.

That varieties of this doubtful nature are far from uncommon cannot be
disputed. Compare the several floras of Great Britainof Franceor of
the United Statesdrawn up by different botanistsand see what a
surprising number of forms have been ranked by one botanist as good
speciesand by another as mere varieties. Mr. H.C. Watsonto whom I lie
under deep obligation for assistance of all kindshas marked for me 182
British plantswhich are generally considered as varietiesbut which have
all been ranked by botanists as species; and in making this list he has
omitted many trifling varietiesbut which nevertheless have been ranked by
some botanists as speciesand he has entirely omitted several highly
polymorphic genera. Under generaincluding the most polymorphic forms
Mr. Babington gives 251 specieswhereas Mr. Bentham gives only 112--a
difference of 139 doubtful forms! Among animals which unite for each
birthand which are highly locomotivedoubtful formsranked by one
zoologist as a species and by another as a varietycan rarely be found
within the same countrybut are common in separated areas. How many of
the birds and insects in North America and Europewhich differ very
slightly from each otherhave been ranked by one eminent naturalist as
undoubted speciesand by another as varietiesoras they are often
calledgeographical races! Mr. Wallacein several valuable papers on the
various animalsespecially on the Lepidopterainhabiting the islands of
the great Malayan Archipelagoshows that they may be classed under four
headsnamelyas variable formsas local formsas geographical races or
sub-speciesand as true representative species. 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; but when all from
the several islands are compared togetherthe differences are seen to be
so slight and graduated that it is impossible to define or describe them
though at the same time the extreme forms are sufficiently distinct. The
geographical races or sub-species are local forms completely fixed and
isolated; but as they do not differ from each other by strongly marked and
important charactersThere is no possible test but individual opinion to
determine which of them shall be considered as species and which as
varieties.Lastlyrepresentative species fill the same place in the
natural economy of each island as do the local forms and sub-species; but
as they are distinguished from each other by a greater amount of difference
than that between the local forms and sub-speciesthey are almost
universally ranked by naturalists as true species. Neverthelessno
certain criterion can possibly be given by which variable formslocal
formssub species and representative species can be recognised.

Many years agowhen comparingand seeing others comparethe birds from
the closely neighbouring islands of the Galapagos Archipelagoone with
anotherand with those from the American mainlandI was much struck how
entirely vague and arbitrary is the distinction between species and
varieties. On the islets of the little Madeira group there are many
insects which are characterized as varieties in Mr. Wollaston's admirable
workbut which would certainly be ranked as distinct species by many
entomologists. Even Ireland has a few animalsnow generally regarded as
varietiesbut which have been ranked as species by some zoologists.
Several experienced ornithologists consider our British red grouse as only
a strongly marked race of a Norwegian specieswhereas the greater number
rank it as an undoubted species peculiar to Great Britain. A wide distance
between the homes of two doubtful forms leads many naturalists to rank them
as distinct species; but what distanceit has been well askedwill
suffice if that between America and Europe is amplewill that between
Europe and the Azoresor Madeiraor the Canariesor between the several
islets of these small archipelagosbe sufficient?

Mr. B.D. Walsha distinguished entomologist of the United Stateshas
described what he calls Phytophagic varieties and Phytophagic species.
Most vegetable-feeding insects live on one kind of plant or on one group of
plants; some feed indiscriminately on many kindsbut do not in consequence
vary. In several caseshoweverinsects found living on different plants
have been observed by Mr. Walsh to present in their larval or mature state
or in both statesslightthough constant differences in coloursizeor
in the nature of their secretions. In some instances the males alonein
other instancesboth males and femaleshave been observed thus to differ
in a slight degree. When the differences are rather more strongly marked
and when both sexes and all ages are affectedthe forms are ranked by all
entomologists as good species. But no observer can determine for another
even if he can do so for himselfwhich of these Phytophagic forms ought to
be called species and which varieties. Mr. Walsh ranks the forms which it
may be supposed would freely intercrossas varieties; and those which
appear to have lost this poweras species. As the differences depend on
the insects having long fed on distinct plantsit cannot be expected that
intermediate links connecting the several forms should now be found. The
naturalist thus loses his best guide in determining whether to rank
doubtful forms as varieties or species. This likewise necessarily occurs
with closely allied organismswhich inhabit distinct continents or
islands. Whenon the other handan animal or plant ranges over the same
continentor inhabits many islands in the same archipelagoand presents
different forms in the different areasthere is always a good chance that
intermediate forms will be discovered which will link together the extreme
states; and these are then degraded to the rank of varieties.

Some few naturalists maintain that animals never present varieties; but
then these same naturalists rank the slightest difference as of specific
value; and when the same identical form is met with in two distant
countriesor in two geological formationsthey believe that two distinct
species are hidden under the same dress. The term species thus comes to be
a mere useless abstractionimplying and assuming a separate act of
creation. It is certain that many formsconsidered by highly competent
judges to be varietiesresemble species so completely in character that
they have been thus ranked by other highly competent judges. But to
discuss whether they ought to be called species or varietiesbefore any
definition of these terms has been generally acceptedis vainly to beat
the air.

Many of the cases of strongly marked varieties or doubtful species well
deserve consideration; for several interesting lines of argumentfrom
geographical distributionanalogical variationhybridismetc.have been
brought to bear in the attempt to determine their rank; but space does not
here permit me to discuss them. Close investigationin many caseswill
no doubt bring naturalists to agree how to rank doubtful forms. Yet it

must be confessed that it is in the best known countries that we find the
greatest number of them. I have been struck with the fact that if any
animal or plant in a state of nature be highly useful to manor from any
cause closely attracts his attentionvarieties of it will almost
universally be found recorded. These varietiesmoreoverwill often be
ranked by some authors as species. Look at the common oakhow closely it
has been studied; yet a German author makes more than a dozen species out
of formswhich are almost universally considered by other botanists to be
varieties; 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.

I may here allude to a remarkable memoir lately published by A. de
Candolleon the oaks of the whole world. No one ever had more ample
materials for the discrimination of the speciesor could have worked on
them with more zeal and sagacity. He first gives in detail all the many
points of structure which vary in the several speciesand estimates
numerically the relative frequency of the variations. He specifies above a
dozen characters which may be found varying even on the same branch
sometimes according to age or developmentsometimes without any assignable
reason. Such characters are not of course of specific valuebut they are
as Asa Gray has remarked in commenting on this memoirsuch as generally
enter into specific definitions. 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 treeand never found connected by intermediate states.
After this discussionthe result of so much labourhe emphatically
remarks: "They are mistakenwho repeat that the greater part of our
species are clearly limitedand that the doubtful species are in a feeble
minority. This seemed to be trueso long as a genus was imperfectly
knownand its species were founded upon a few specimensthat is to say
were provisional. Just as we come to know them betterintermediate forms
flow inand 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. Thus Quercus robur has twenty-eight
varietiesall of whichexcepting sixare clustered round three subspecies
namely Q. pedunculatasessiliflora and pubescens. The forms
which connect these three sub-species are comparatively rare; andas Asa
Gray again remarksif 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. FinallyDe
Candolle admits that out of the 300 specieswhich will be enumerated in
his Prodromus as belonging to the oak familyat least two-thirds are
provisional speciesthat isare not known strictly to fulfil the
definition above given of a true species. It should be added that De
Candolle no longer believes that species are immutable creationsbut
concludes that the derivative theory is the most natural oneand the most
accordant with the known facts in palaeontology, geographical botany and
zoology, of anatomical structure and classification.

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; for he knows
nothing of the amount and kind of variation to which the group is subject;
and this showsat leasthow very generally there is some variation. 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. His general
tendency will be to make many speciesfor he will become impressedjust
like the pigeon or poultry fancier before alluded towith the amount of
difference in the forms which he is continually studying; and he has little
general knowledge of analogical variation in other groups and in other
countries by which to correct his first impressions. As he extends the
range of his observations he will meet with more cases of difficulty; for
he will encounter a greater number of closely-allied forms. But if his

observations be widely extended he will in the end generally be able to
make up his own mind; but he will succeed in this at the expense of
admitting much variationand the truth of this admission will often be
disputed by other naturalists. When he comes to study allied forms brought
from countries not now continuousin which case he cannot hope to find
intermediate linkshe will be compelled to trust almost entirely to
analogyand his difficulties will rise to a climax.

Certainly no clear line of demarcation has as yet been drawn between
species and sub-species--that isthe forms which in the opinion of some
naturalists come very near tobut do not quite arrive atthe rank of
species; oragainbetween sub-species and well-marked varietiesor
between lesser varieties and individual differences. These differences
blend into each other by an insensible series; and a series impresses the
mind with the idea of an actual passage.

Hence I look at individual differencesthough of small interest to the
systematistas of the highest importance for usas being the first step
towards such slight varieties as are barely thought worth recording in
works on natural history. And I look at varieties which are in any degree
more distinct and permanentas steps toward more strongly marked and
permanent varieties; and at the latteras leading to sub-speciesand then
to species. The passage from one stage of difference to another mayin
many casesbe the simple result of the nature of the organism and of the
different physical conditions to which it has long been exposed; but with
respect to the more important and adaptive charactersthe passage from one
stage of difference to another may be safely attributed to the cumulative
action of natural selectionhereafter to be explainedand to the effects
of the increased use or disuse of parts. A well-marked variety may
therefore be called an incipient species; but whether this belief is
justifiable must be judged by the weight of the various facts and
considerations to be given throughout this work.

It need not be supposed that all varieties or incipient species attain the
rank of species. They may become extinctor they may endure as varieties
for very long periodsas has been shown to be the case by Mr. Wollaston
with the varieties of certain fossil land-shells in Madeiraand with
plants by Gaston de Saporta. If a variety were to flourish so as to exceed
in numbers the parent speciesit would then rank as the speciesand the
species as the variety; or it might come to supplant and exterminate the
parent species; or both might co-existand both rank as independent
species. But we shall hereafter return to this subject.

>From these remarks it will be seen that I look at the term species as one
arbitrarily givenfor the sake of convenienceto a set of individuals
closely resembling each otherand that it does not essentially differ from
the term varietywhich is given to less distinct and more fluctuating
forms. The term varietyagainin comparison with mere individual
differencesis also applied arbitrarilyfor convenience sake.


Guided by theoretical considerationsI thought that some interesting
results might be obtained in regard to the nature and relations of the
species which vary mostby tabulating all the varieties in several
well-worked floras. At first this seemed a simple task; but Mr. H.C.
Watsonto whom I am much indebted for valuable advice and assistance on
this subjectsoon convinced me that there were many difficultiesas did
subsequently Dr. Hookereven in stronger terms. I shall reserve for a
future work the discussion of these difficultiesand the tables of the
proportional numbers of the varying species. Dr. Hooker permits me to add
that after having carefully read my manuscriptand examined the tableshe
thinks that the following statements are fairly well established. The
whole subjecthowevertreated as it necessarily here is with much

brevityis rather perplexingand allusions cannot be avoided to the
struggle for existence,divergence of character,and other questions
hereafter to be discussed.

Alphonse de Candolle and others have shown that plants which have very wide
ranges generally present varieties; and this might have been expectedas
they are exposed to diverse physical conditionsand as they come into
competition (whichas we shall hereafter seeis a far more important
circumstance) with different sets of organic beings. But my tables further
show thatin any limited countrythe species which are the most common
that is abound most in individualsand the species which are most widely
diffused within their own country (and this is a different consideration
from wide rangeand to a certain extent from commonness)oftenest give
rise to varieties sufficiently well-marked to have been recorded in
botanical works. Hence it is the most flourishingoras they may be
calledthe dominant species--those which range widelyare the most
diffused in their own countryand are the most numerous in
individuals--which oftenest produce well-marked varietiesoras I
consider themincipient species. And thisperhapsmight have been
anticipated; foras varietiesin order to become in any degree permanent
necessarily have to struggle with the other inhabitants of the countrythe
species which are already dominant will be the most likely to yield
offspringwhichthough in some slight degree modifiedstill inherit
those advantages that enabled their parents to become dominant over their
compatriots. In these remarks on predominenceit should be understood
that reference is made only to the forms which come into competition with
each otherand more especially to the members of the same genus or class
having nearly similar habits of life. With respect to the number of
individuals or commonness of speciesthe comparison of course relates only
to the members of the same group. 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 countrywhich live under nearly the same
conditions. 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 individualsand more widely diffused. But if the conferva or
parasitic fungus exceeds its allies in the above respectsit will then be
dominant within its own class.


If the plants inhabiting a country as described in any Florabe divided
into two equal massesall those in the larger genera (i.e.those
including many species) being placed on one sideand all those in the
smaller genera on the other sidethe former will be found to include a
somewhat larger number of the very common and much diffused or dominant
species. This might have been anticipatedfor the mere fact of many
species of the same genus inhabiting any countryshows that there is
something in the organic or inorganic conditions of that country favourable
to the genus; andconsequentlywe might have expected to have found in
the larger generaor those including many speciesa larger proportional
number of dominant species. But so many causes tend to obscure this
resultthat I am surprised that my tables show even a small majority on
the side of the larger genera. I will here allude to only two causes of
obscurity. Fresh water and salt-loving plants generally have very wide
ranges and are much diffusedbut this seems to be connected with the
nature of the stations inhabited by themand has little or no relation to
the size of the genera to which the species belong. Againplants low in
the scale of organisation are generally much more widely diffused than
plants higher in the scale; and here again there is no close relation to
the size of the genera. The cause of lowly-organised plants ranging widely
will be discussed in our chapter on Geographical Distribution.

>From looking at species as only strongly marked and well-defined varieties

I was led to anticipate that the species of the larger genera in each
country would oftener present varietiesthan the species of the smaller
genera; for wherever many closely related species (i.e.species of the
same genus) have been formedmany varieties or incipient species oughtas
a general ruleto be now forming. Where many large trees growwe expect
to find saplings. Where many species of a genus have been formed through
variationcircumstances have been favourable for variation; and hence we
might expect that the circumstances would generally still be favourable to
variation. On the other handif we look at each species as a special act
of creationthere is no apparent reason why more varieties should occur in
a group having many speciesthan in one having few.

To test the truth of this anticipation I have arranged the plants of twelve
countriesand the coleopterous insects of two districtsinto two nearly
equal massesthe species of the larger genera on one sideand those of
the smaller genera on the other sideand it has invariably proved to be
the case that a larger proportion of the species on the side of the larger
genera presented varietiesthan on the side of the smaller genera.
Moreoverthe species of the large genera which present any varieties
invariably present a larger average number of varieties than do the species
of the small genera. Both these results follow when another division is
madeand when all the least generawith from only one to four species
are altogether excluded from the tables. These facts are of plain
signification on the view that species are only strongly marked and
permanent varieties; for wherever many species of the same genus have been
formedor whereif we may use the expressionthe manufactory of species
has been activewe ought generally to find the manufactory still in
actionmore 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; for my tables clearly show
as a general rulethatwherever many species of a genus have been formed
the species of that genus present a number of varietiesthat isof
incipient speciesbeyond the average. It is not that all large genera are
now varying muchand are thus increasing in the number of their species
or that no small genera are now varying and increasing; for if this had
been soit would have been fatal to my theory; inasmuch as geology plainly
tells us that small genera have in the lapse of time often increased
greatly in size; and that large genera have often come to their maxima
declinedand disappeared. All that we want to show isthat where many
species of a genus have been formedon an average many are still forming;
and this certainly holds good.


There are other relations between the species of large genera and their
recorded varieties which deserve notice. We have seen that there is no
infallible criterion by which to distinguish species and well-marked
varieties; and when intermediate links have not been found between doubtful
formsnaturalists are compelled to come to a determination by the amount
of difference between themjudging by analogy whether or not the amount
suffices to raise one or both to the rank of species. Hence the amount of
difference is one very important criterion in settling whether two forms
should be ranked as species or varieties. Now Fries has remarked in regard
to plantsand Westwood in regard to insectsthat in large genera the
amount of difference between the species is often exceedingly small. I
have endeavoured to test this numerically by averagesandas far as my
imperfect results gothey confirm the view. I have also consulted some
sagacious and experienced observersandafter deliberationthey concur
in this view. In this respectthereforethe species of the larger genera
resemble varietiesmore than do the species of the smaller genera. Or the
case may be put in another wayand it may be saidthat in the larger
generain which a number of varieties or incipient species greater than

the average are now manufacturingmany of the species already manufactured
still to a certain extent resemble varietiesfor they differ from each
other by a less than the usual amount of difference.

Moreoverthe species of the larger genera are related to each otherin
the same manner as the varieties of any one species are related to each
other. No naturalist pretends that all the species of a genus are equally
distinct from each other; they may generally be divided into sub-generaor
sectionsor lesser groups. As Fries has well remarkedlittle groups of
species are generally clustered like satellites around other species. And
what are varieties but groups of formsunequally related to each other
and clustered round certain forms--that isround their parent-species.
Undoubtedly there is one most important point of difference between
varieties and speciesnamelythat the amount of difference between
varietieswhen compared with each other or with their parent-speciesis
much less than that between the species of the same genus. But when we
come to discuss the principleas I call itof divergence of characterwe
shall see how this may be explainedand how the lesser differences between
varieties tend to increase into the greater differences between species.

There is one other point which is worth notice. Varieties generally have
much restricted ranges. This statement is indeed scarcely more than a
truismfor if a variety were found to have a wider range than that of its
supposed parent-speciestheir denominations would be reversed. But there
is reason to believe that the species which are very closely allied to
other speciesand in so far resemble varietiesoften have much restricted
ranges. For instanceMr. H.C. Watson has marked for me in the well-sifted
London catalogue of Plants (4th edition) sixty-three plants which are
therein ranked as speciesbut 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 6.9 of the provinces into which Mr. Watson
has divided Great Britain. Nowin this same cataloguefifty-three
acknowledged varieties are recordedand these range over 7.7 provinces;
whereasthe species to which these varieties belong range over 14.3
provinces. So that the acknowledged varieties have very nearly the same
restricted average rangeas have the closely allied formsmarked for me
by Mr. Watson as doubtful speciesbut which are almost universally ranked
by British botanists as good and true species.


Finallyvarieties cannot be distinguished from species--exceptfirstby
the discovery of intermediate linking forms; andsecondlyby a certain
indefinite amount of difference between them; for two formsif differing
very littleare generally ranked as varietiesnotwithstanding that they
cannot be closely connected; but the amount of difference considered
necessary to give to any two forms the rank of species cannot be defined.
In genera having more than the average number of species in any country
the species of these genera have more than the average number of varieties.
In large genera the species are apt to be closely but unequally allied
togetherforming little clusters round other species. Species very
closely allied to other species apparently have restricted ranges. In all
these respects the species of large genera present a strong analogy with
varieties. And we can clearly understand these analogiesif species once
existed as varietiesand thus originated; whereasthese analogies are
utterly inexplicable if species are independent creations.

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 varietiesand varietiesas we shall hereafter seetend to
become converted into new and distinct species. Thus the larger genera
tend to become larger; and throughout nature the forms of life which are
now dominant tend to become still more dominant by leaving many modified
and dominant descendants. Butby steps hereafter to be explainedthe

larger genera also tend to break up into smaller genera. And thusthe
forms of life throughout the universe become divided into groups
subordinate to groups.



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.

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. 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. It is immaterial for us whether a
multitude of doubtful forms be called species or sub-species or varieties;
what rankfor instancethe two or three hundred doubtful forms of British
plants are entitled to holdif the existence of any well-marked varieties
be admitted. But the mere existence of individual variability and of some
few well-marked varietiesthough necessary as the foundation for the work
helps us but little in understanding how species arise in nature. How have
all those exquisite adaptations of one part of the organisation to another
partand to the conditions of life and of one organic being to another
beingbeen perfected? We see these beautiful co-adaptations most plainly
in the woodpecker and the mistletoe; and only a little less plainly in the
humblest parasite which clings to the hairs of a quadruped or feathers of a
bird; in the structure of the beetle which dives through the water; in the
plumed seed which is wafted by the gentlest breeze; in shortwe see
beautiful adaptations everywhere and in every part of the organic world.

Againit may be askedhow is it that varietieswhich I have called
incipient speciesbecome ultimately converted into good and distinct
specieswhich in most cases obviously differ from each other far more than
do the varieties of the same species? How do those groups of species
which constitute what are called distinct genera and which differ from each
other more than do the species of the same genusarise? All these
resultsas we shall more fully see in the next chapterfollow from the
struggle for life. Owing to this strugglevariationshowever slight and
from whatever cause proceedingif they be in any degree profitable to the
individuals of a speciesin their infinitely complex relations to other
organic beings and to their physical conditions of lifewill tend to the
preservation of such individualsand will generally be inherited by the
offspring. The offspringalsowill thus have a better chance of
survivingforof the many individuals of any species which are
periodically bornbut a small number can survive. I have called this
principleby which each slight variationif usefulis preservedby the
term natural selectionin order to mark its relation to man's power of
selection. But the expression often used by Mr. Herbert Spencerof the
Survival of the Fittestis more accurateand is sometimes equally
convenient. We have seen that man by selection can certainly produce great
resultsand can adapt organic beings to his own usesthrough the
accumulation of slight but useful variationsgiven to him by the hand of
Nature. But Natural Selectionwe shall hereafter seeis a power
incessantly ready for actionand is as immeasurably superior to man's
feeble effortsas the works of Nature are to those of Art.

We will now discuss in a little more detail the struggle for existence. In
my future work this subject will be treatedas it well deservesat
greater length. The elder De Candolle and Lyell have largely and
philosophically shown that all organic beings are exposed to severe
competition. In regard to plantsno one has treated this subject with
more spirit and ability than W. HerbertDean of Manchesterevidently the
result of his great horticultural knowledge. Nothing is easier than to
admit in words the truth of the universal struggle for lifeor more
difficult--at least I found it so--than constantly to bear this conclusion
in mind. Yet unless it be thoroughly engrained in the mindthe whole
economy of naturewith every fact on distributionrarityabundance
extinctionand variationwill be dimly seen or quite misunderstood. We
behold the face of nature bright with gladnesswe often see superabundance
of food; we do not see or we forget that the birds which are idly singing
round us mostly live on insects or seedsand are thus constantly
destroying life; or we forget how largely these songstersor their eggs
or their nestlingsare destroyed by birds and beasts of prey; we do not
always bear in mindthatthough food may be now superabundantit is not
so at all seasons of each recurring year.


I should premise that I use this term in a large and metaphorical sense
including dependence of one being on anotherand including (which is more
important) not only the life of the individualbut success in leaving
progeny. Two canine animalsin a time of dearthmay 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 droughtthough
more properly it should be said to be dependent on the moisture. A plant
which annually produces a thousand seedsof which only one of an average
comes to maturitymay be more truly said to struggle with the plants of
the same and other kinds which already clothe the ground. The mistletoe is
dependent on the apple and a few other treesbut can only in a far-fetched
sense be said to struggle with these treesforif too many of these
parasites grow on the same treeit languishes and dies. But several
seedling mistletoesgrowing close together on the same branchmay more
truly be said to struggle with each other. As the mistletoe is
disseminated by birdsits existence depends on them; and it may
metaphorically be said to struggle with other fruit-bearing plantsin
tempting the birds to devour and thus disseminate its seeds. In these
several senseswhich pass into each otherI use for convenience sake the
general term of Struggle for Existence.


A struggle for existence inevitably follows from the high rate at which all
organic beings tend to increase. Every beingwhich during its natural
lifetime produces several eggs or seedsmust suffer destruction during
some period of its lifeand during some season or occasional year
otherwiseon the principle of geometrical increaseits numbers would
quickly become so inordinately great that no country could support the
product. Henceas more individuals are produced than can possibly
survivethere must in every case be a struggle for existenceeither one
individual with another of the same speciesor with the individuals of
distinct speciesor with the physical conditions of life. It is the
doctrine of Malthus applied with manifold force to the whole animal and
vegetable kingdoms; for in this case there can be no artificial increase of
foodand no prudential restraint from marriage. Although some species may
be now increasingmore or less rapidlyin numbersall cannot do sofor
the world would not hold them.

There is no exception to the rule that every organic being naturally
increases at so high a ratethatif not destroyedthe earth would soon
be covered by the progeny of a single pair. Even slow-breeding man has

doubled in twenty-five yearsand at this ratein less than a thousand
yearsthere would literally not be standing room for his progeny.
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 twoand so onthen in twenty years there would be a
million plants. The elephant is reckoned the slowest breeder of all known
animalsand I have taken some pains to estimate its probable minimum rate
of natural increase; it will be safest to assume that it begins breeding
when thirty years oldand goes on breeding till ninety years oldbringing
forth six young in the intervaland surviving till one hundred years old;
if this be soafter a period of from 740 to 750 years there would be
nearly nineteen million elephants alive descended from the first pair.

But we have better evidence on this subject than mere theoretical
calculationsnamelythe numerous recorded cases of the astonishingly
rapid increase of various animals in a state of naturewhen circumstances
have been favourable to them during two or three following seasons. Still
more striking is the evidence from our domestic animals of many kinds which
have run wild in several parts of the world; if the statements of the rate
of increase of slow-breeding cattle and horses in South Americaand
latterly in Australiahad not been well authenticatedthey would have
been incredible. So it is with plants; cases could be given of introduced
plants which have become common throughout whole islands in a period of
less than ten years. Several of the plantssuch as the cardoon and a tall
thistlewhich are now the commonest over the wide plains of La Plata
clothing square leagues of surface almost to the exclusion of every other
planthave been introduced from Europe; and there are plants which now
range in Indiaas I hear from Dr. Falconerfrom Cape Comorin to the
Himalayawhich have been imported from America since its discovery. In
such casesand endless others could be givenno one supposes that the
fertility of the animals or plants has been suddenly and temporarily
increased in any sensible degree. The obvious explanation is that the
conditions of life have been highly favourableand that there has
consequently been less destruction of the old and young and that nearly all
the young have been enabled to breed. Their geometrical ratio of increase
the result of which never fails to be surprisingsimply explains their
extraordinarily rapid increase and wide diffusion in their new homes.

In a state of nature almost every full-grown plant annually produces seed
and among animals there are very few which do not annually pair. 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 existand that this geometrical tendency to increase
must be checked by destruction at some period of life. Our familiarity
with the larger domestic animals tendsI thinkto mislead us; we see no
great destruction falling on themand we do not keep in mind that
thousands are annually slaughtered for foodand that in a state of nature
an equal number would have somehow to be disposed of.

The only difference between organisms which annually produce eggs or seeds
by the thousandand those which produce extremely fewisthat the slow
breeders would require a few more years to peopleunder favourable
conditionsa whole districtlet it be ever so large. The condor lays a
couple of eggs and the ostrich a scoreand yet in the same country the
condor may be the more numerous of the two. The Fulmar petrel lays but one
eggyet it is believed to be the most numerous bird in the world. One fly
deposits hundreds of eggsand anotherlike the hippoboscaa single one.
But this difference does not determine how many individuals of the two
species can be supported in a district. A large number of eggs is of some
importance to those species which depend on a fluctuating amount of food
for it allows them rapidly to increase in number. But the real importance
of a large number of eggs or seeds is to make up for much destruction at
some period of life; and this period in the great majority of cases is an
early one. If an animal can in any way protect its own eggs or younga

small number may be producedand yet the average stock be fully kept up;
but if many eggs or young are destroyedmany must be produced or the
species will become extinct. It would suffice to keep up the full number
of a treewhich lived on an average for a thousand yearsif a single seed
were produced once in a thousand yearssupposing that this seed were never
destroyed and could be ensured to germinate in a fitting place; so thatin
all casesthe average number of any animal or plant depends only
indirectly on the number of its eggs or seeds.

In looking at Natureit 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; 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. Lighten any checkmitigate the destruction ever so
littleand the number of the species will almost instantaneously increase
to any amount.


The causes which check the natural tendency of each species to increase are
most obscure. Look at the most vigorous species; by as much as it swarms
in numbersby so much will it tend to increase still further. We know not
exactly what the checks are even in a single instance. Nor will this
surprise any one who reflects how ignorant we are on this headeven in
regard to mankindalthough so incomparably better known than any other
animal. This subject of the checks to increase has been ably treated by
several authorsand I hope in a future work to discuss it at considerable
lengthmore especially in regard to the feral animals of South America.
Here I will make only a few remarksjust to recall to the reader's mind
some of the chief points. Eggs or very young animals seem generally to
suffer mostbut this is not invariably the case. With plants there is a
vast destruction of seedsbut from some observations which I have made it
appears that the seedlings suffer most from germinating in ground already
thickly stocked with other plants. Seedlingsalsoare destroyed in vast
numbers by various enemies; for instanceon a piece of ground three feet
long and two widedug and clearedand where there could be no choking
from other plantsI marked all the seedlings of our native weeds as they
came upand out of 357 no less than 295 were destroyedchiefly by slugs
and insects. If turf which has long been mownand the case would be the
same with turf closely browsed by quadrupedsbe let to growthe more
vigorous plants gradually kill the less vigorousthough fully grown
plants; thus out of twenty species grown on a little plot of mown turf
(three feet by four) nine species perishedfrom the other species being
allowed to grow up freely.

The amount of food for each speciesof coursegives the extreme limit to
which each can increase; but very frequently it is not the obtaining food
but the serving as prey to other animalswhich determines the average
number of a species. Thusthere seems to be little doubt that the stock
of partridgesgrouseand hares on any large estate depends chiefly on the
destruction of vermin. If not one head of game were shot during the next
twenty years in Englandandat the same timeif no vermin were
destroyedthere wouldin all probabilitybe less game than at present
although hundreds of thousands of game animals are now annually shot. On
the other handin some casesas with the elephantnone are destroyed by
beasts of prey; for even the tiger in India most rarely dares to attack a
young elephant protected by its dam.

Climate plays an important part in determining the average numbers of a
speciesand periodical seasons of extreme cold or drought seem to be the
most effective of all checks. I estimated (chiefly from the greatly
reduced numbers of nests in the spring) that the winter of 1854-5 destroyed
four-fifths of the birds in my own grounds; and this is a tremendous

destructionwhen we remember that ten per cent. is an extraordinarily
severe mortality from epidemics with man. The action of climate seems at
first sight to be quite independent of the struggle for existence; but in
so far as climate chiefly acts in reducing foodit brings on the most
severe struggle between the individualswhether of the same or of distinct
specieswhich subsist on the same kind of food. Even when climatefor
instanceextreme coldacts directlyit will be the least vigorous
individualsor those which have got least food through the advancing
winterwhich will suffer the most. When we travel from south to northor
from a damp region to a drywe invariably see some species gradually
getting rarer and rarerand finally disappearing; and the change of
climate being conspicuouswe are tempted to attribute the whole effect to
its direct action. But this is a false view; we forget that each species
even where it most aboundsis constantly suffering enormous destruction at
some period of its lifefrom enemies or from competitors for the same
place and food; and if these enemies or competitors be in the least degree
favoured by any slight change of climatethey will increase in numbers;
and as each area is already fully stocked with inhabitantsthe other
species must decrease. When we travel southward and see a species
decreasing in numberswe may feel sure that the cause lies quite as much
in other species being favouredas in this one being hurt. So it is when
we travel northwardbut in a somewhat lesser degreefor the number of
species of all kindsand therefore of competitorsdecreases northward;
hence in going northwardor in ascending a mountainwe far oftener meet
with stunted formsdue to the DIRECTLY injurious action of climatethan
we do in proceeding southward or in descending a mountain. When we reach
the Arctic regionsor snow-capped summitsor absolute desertsthe
struggle for life is almost exclusively with the elements.

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 climatebut which never become naturalisedfor
they cannot compete with our native plants nor resist destruction by our
native animals.

When a speciesowing to highly favourable circumstancesincreases
inordinately in numbers in a small tractepidemics--at leastthis seems
generally to occur with our game animals--often ensue; and here we have a
limiting check independent of the struggle for life. But even some of
these so-called epidemics appear to be due to parasitic wormswhich have
from some causepossibly in part through facility of diffusion among the
crowded animalsbeen disproportionally favoured: and here comes in a sort
of struggle between the parasite and its prey.

On the other handin many casesa large stock of individuals of the same
speciesrelatively to the numbers of its enemiesis absolutely necessary
for its preservation. Thus we can easily raise plenty of corn and our fieldsbecause the seeds are in great excess
compared with the number of birds which feed on them; nor can the birds
though having a superabundance of food at this one seasonincrease in
number proportionally to the supply of seedas their numbers are checked
during the winter; 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; I have in this
case lost every single seed. This view of the necessity of a large stock
of the same species for its preservationexplainsI believesome
singular facts in nature such as that of very rare plants being sometimes
extremely abundantin the few spots where they do exist; and that of some
social plants being socialthat is abounding in individualseven on the
extreme verge of their range. For in such caseswe may believethat a
plant could exist only where the conditions of its life were so favourable
that many could exist togetherand thus save the species from utter
destruction. I should add that the good effects of intercrossingand the
ill effects of close interbreedingno doubt come into play in many of
these cases; but I will not here enlarge on this subject.


Many cases are on record showing how complex and unexpected are the checks
and relations between organic beingswhich have to struggle together in
the same country. I will give only a single instancewhichthough a
simple oneinterested me. In Staffordshireon the estate of a relation
where I had ample means of investigationthere was a large and extremely
barren heathwhich had never been touched by the hand of man; but several
hundred acres of exactly the same nature had been enclosed twenty-five
years previously and planted with Scotch fir. The change in the native
vegetation of the planted part of the heath was most remarkablemore than
is generally seen in passing from one quite different soil to another: not
only the proportional numbers of the heath-plants were wholly changedbut
twelve species of plants (not counting grasses and carices) flourished in
the plantationswhich could not be found on the heath. The effect on the
insects must have been still greaterfor six insectivorous birds were very
common in the plantationswhich were not to be seen on the heath; and the
heath was frequented by two or three distinct insectivorous birds. Here we
see how potent has been the effect of the introduction of a single tree
nothing whatever else having been donewith the exception of the land
having been enclosedso that cattle could not enter. But how important an
element enclosure isI plainly saw near Farnhamin Surrey. Here there
are extensive heathswith a few clumps of old Scotch firs on the distant
hill-tops: within the last ten years large spaces have been enclosedand
self-sown firs are now springing up in multitudesso close together that
all cannot live. 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 viewwhence I could examine hundreds of acres of the
unenclosed heathand literally I could not see a single Scotch firexcept
the old planted clumps. But on looking closely between the stems of the
heathI found a multitude of seedlings and little treeswhich had been
perpetually browsed down by the cattle. In one square yardat a point
some hundred yards distant from one of the old clumpsI counted thirty-two
little trees; and one of themwith twenty-six rings of growthhadduring
many years tried to raise its head above the stems of the heathand had
failed. No wonder thatas soon as the land was enclosedit became
thickly clothed with vigorously growing young firs. Yet the heath was so
extremely barren and so extensive that no one would ever have imagined that
cattle would have so closely and effectually searched it for food.

Here we see that cattle absolutely determine the existence of the Scotch
fir; but in several parts of the world insects determine the existence of
cattle. Perhaps Paraguay offers the most curious instance of this; for
here neither cattle nor horses nor dogs have ever run wildthough they
swarm southward and northward in a feral state; and Azara and Rengger have
shown that this is caused by the greater number in Paraguay of a certain
flywhich lays its eggs in the navels of these animals when first born.
The increase of these fliesnumerous as they aremust be habitually
checked by some meansprobably by other parasitic insects. Henceif
certain insectivorous birds were to decrease in Paraguaythe parasitic
insects would probably increase; and this would lessen the number of the
navel-frequenting flies--then cattle and horses would become feraland
this would certainly greatly alter (as indeed I have observed in parts of
South America) the vegetation: this again would largely affect the
insects; and thisas we have just seen in Staffordshirethe insectivorous
birdsand so onwards in ever-increasing circles of complexity. Not that
under nature the relations will ever be as simple as this. Battle within
battle must be continually recurring with varying success; and yet in the
long-run the forces are so nicely balanced that the face of nature remains
for long periods of time uniformthough assuredly the merest trifle would
give the victory to one organic being over another. Neverthelessso
profound is our ignoranceand so high our presumptionthat we marvel when

we hear of the extinction of an organic being; and as we do not see the
causewe invoke cataclysms to desolate the worldor invent laws on the
duration of the forms of life!

I am tempted to give one more instance showing how plants and animals
remote in the scale of natureare bound together by a web of complex
relations. I shall hereafter have occasion to show that the exotic Lobelia
fulgens is never visited in my garden by insectsand consequentlyfrom
its peculiar structurenever sets a seed. Nearly all our orchidaceous
plants absolutely require the visits of insects to remove their
pollen-masses and thus to fertilise them. I find from experiments that
humble-bees are almost indispensable to the fertilisation of the heartsease
(Viola tricolor)for other bees do not visit this flower. I have also
found that the visits of bees are necessary for the fertilisation of some
kinds of clover; for instance twenty heads of Dutch clover (Trifolium
repens) yielded 2290 seedsbut twenty other headsprotected from bees
produced not one. Again100 heads of red clover (T. pratense) produced
2700 seedsbut the same number of protected heads produced not a single
seed. Humble bees alone visit red cloveras other bees cannot reach the
nectar. It has been suggested that moths may fertilise the clovers; but I
doubt whether they could do so in the case of the red cloverfrom their
weight not being sufficient to depress the wing petals. Hence we may infer
as highly probable thatif the whole genus of humble-bees became extinct
or very rare in Englandthe heartsease and red clover would become very
rareor wholly disappear. The number of humble-bees in any district
depends in a great measure upon the number of field-micewhich destroy
their combs and nests; and Colonel Newmanwho has long attended to the
habits of humble-beesbelieves that "more than two-thirds of them are thus
destroyed all over England." Now the number of mice is largely dependent
as every one knowson the number of cats; and Colonel Newman saysNear
villages and small towns I have found the nests of humble-bees more
numerous than elsewhere, which I attribute to the number of cats that
destroy the mice.Hence it is quite credible that the presence of a
feline animal in large numbers in a district might determinethrough the
intervention first of mice and then of beesthe frequency of certain
flowers in that district!

In the case of every speciesmany different checksacting at different
periods of lifeand during different seasons or yearsprobably come into
play; some one check or some few being generally the most potentbut all
will concur in determining the average numberor even the existence of the
species. In some cases it can be shown that widely-different checks act on
the same species in different districts. When we look at the plants and
bushes clothing an entangled bankwe are tempted to attribute their
proportional numbers and kinds to what we call chance. But how false a
view is this! Every one has heard that when an American forest is cut
downa very different vegetation springs up; but it has been observed that
ancient Indian ruins in the Southern United Stateswhich must formerly
have been cleared of treesnow display the same beautiful diversity and
proportion of kinds as in the surrounding virgin forests. What a struggle
must have gone on during long centuries between the several kinds of trees
each annually scattering its seeds by the thousand; what war between insect
and insect--between insectssnailsand other animals with birds and
beasts of prey--all striving to increaseall feeding on each otheror on
the treestheir seeds and seedlingsor on the other plants which first
clothed the ground and thus checked the growth of the trees. Throw up a
handful of feathersand all fall to the ground according to definite laws;
but how simple is the problem where each shall fall compared to that of the
action and reaction of the innumerable plants and animals which have
determinedin the course of centuriesthe proportional numbers and kinds
of trees now growing on the old Indian ruins!

The dependency of one organic being on anotheras of a parasite on its
preylies generally between beings remote in the scale of nature. This is

likewise sometimes the case with those which may strictly be said to
struggle with each other for existenceas in the case of locusts and
grass-feeding quadrupeds. But the struggle will almost invariably be most
severe between the individuals of the same speciesfor they frequent the
same districtsrequire the same foodand are exposed to the same dangers.
In the case of varieties of the same speciesthe struggle will generally
be almost equally severeand we sometimes see the contest soon decided:
for instanceif several varieties of wheat be sown togetherand the mixed
seed be resownsome of the varieties which best suit the soil or climate
or are naturally the most fertilewill beat the others and so yield more
seedand will consequently in a few years supplant the other varieties.
To keep up a mixed stock of even such extremely close varieties as the
variously coloured sweet-peasthey must be each year harvested separately
and the seed then mixed in due proportionotherwise the weaker kinds will
steadily decrease in number and disappear. So again with the varieties of
sheep: it has been asserted that certain mountain-varieties will starve
out other mountain-varietiesso that they cannot be kept together. The
same result has followed from keeping together different varieties of the
medicinal leech. It may even be doubted whether the varieties of any of
our domestic plants or animals have so exactly the same strengthhabits
and constitutionthat the original proportions of a mixed stock (crossing
being prevented) could be kept up for half-a-dozen generationsif they
were allowed to struggle togetherin the same manner as beings in a state
of natureand if the seed or young were not annually preserved in due


As the species of the same genus usually havethough by no means
invariablymuch similarity in habits and constitutionand always in
structurethe struggle will generally be more severe between themif they
come into competition with each otherthan between the species of distinct
genera. 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. The recent increase of the missel-thrush in parts of Scotland has
caused the decrease of the song-thrush. 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. In Australia the imported hive-bee is
rapidly exterminating the smallstingless native bee. One species of
charlock has been known to supplant another species; and so in other cases.
We can dimly see why the competition should be most severe between allied
formswhich fill nearly the same place in the economy of nature; but
probably in no one case could we precisely say why one species has been
victorious over another in the great battle of life.

A corollary of the highest importance may be deduced from the foregoing
remarksnamelythat the structure of every organic being is relatedin
the most essential yet often hidden mannerto that of all other organic
beingswith which it comes into competition for food or residenceor from
which it has to escapeor on which it preys. This is obvious in the
structure of the teeth and talons of the tiger; and in that of the legs and
claws of the parasite which clings to the hair on the tiger's body. But in
the beautifully plumed seed of the dandelionand in the flattened and
fringed legs of the water-beetlethe relation seems at first confined to
the elements of air and water. Yet the advantage of the plumed seeds no
doubt stands in the closest relation to the land being already thickly
clothed with other plants; so that the seeds may be widely distributed and
fall on unoccupied ground. In the water-beetlethe structure of its legs
so well adapted for divingallows it to compete with other aquatic
insectsto hunt for its own preyand to escape serving as prey to other

The store of nutriment laid up within the seeds of many plants seems at
first sight to have no sort of relation to other plants. But from the
strong growth of young plants produced from such seedsas peas and beans
when sown in the midst of long grassit may be suspected that the chief
use of the nutriment in the seed is to favour the growth of the seedlings
whilst struggling with other plants growing vigorously all around.

Look at a plant in the midst of its range! Why does it not double or
quadruple its numbers? We know that it can perfectly well withstand a
little more heat or colddampness or drynessfor elsewhere it ranges into
slightly hotter or colderdamper or drier districts. In this case we can
clearly see that if we wish in imagination to give the plant the power of
increasing in numberswe should have to give it some advantage over its
competitorsor over the animals which prey on it. On the confines of its
geographical rangea change of constitution with respect to climate would
clearly be an advantage to our plant; but we have reason to believe that
only a few plants or animals range so farthat they are destroyed
exclusively by the rigour of the climate. Not until we reach the extreme
confines of lifein the Arctic regions or on the borders of an utter
desertwill competition cease. The land may be extremely cold or dryyet
there will be competition between some few speciesor between the
individuals of the same speciesfor the warmest or dampest spots.

Hence we can see that when a plant or animal is placed in a new country
among new competitorsthe conditions of its life will generally be changed
in an essential manneralthough the climate may be exactly the same as in
its former home. If its average numbers are to increase in its new home
we should have to modify it in a different way to what we should have had
to do in its native country; for we should have to give it some advantage
over a different set of competitors or enemies.

It is good thus to try in imagination to give any one species an advantage
over another. Probably in no single instance should we know what to do.
This ought to convince us of our ignorance on the mutual relations of all
organic beings; a conviction as necessaryas it is difficult to acquire.
All that we can do is to keep steadily in mind that each organic being is
striving to increase in a geometrical ratio; that eachat some period of
its lifeduring some season of the yearduring each generationor at
intervalshas to struggle for life and to suffer great destruction. When
we reflect on this struggle we may console ourselves with the full belief
that the war of nature is not incessantthat no fear is feltthat death
is generally promptand that the vigorousthe healthyand the happy
survive and multiply.



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 Selectionnamelyintercrossing
isolationnumber of individuals -- Slow action -- Extinction caused by
Natural Selection -- Divergence of Characterrelated to the diversity of
inhabitants of any small area and to naturalisation -- Action of Natural
Selectionthrough Divergence of Character and Extinctionon 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.

How will the struggle for existencebriefly discussed in the last chapter
act in regard to variation? Can the principle of selectionwhich we have

seen is so potent in the hands of manapply 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
productionsandin a lesser degreein those under naturebe borne in
mind; as well as the strength of the hereditary tendency. Under
domesticationit may truly be said that the whole organisation becomes in
some degree plastic. But the variabilitywhich we almost universally meet
with in our domestic productions is not directly producedas Hooker and
Asa Gray have well remarkedby man; he can neither originate varieties nor
prevent their occurrence; he can only preserve and accumulate such as do
occur. Unintentionally he exposes organic beings to new and changing
conditions of lifeand variability ensues; but similar changes of
conditions might and do occur under nature. 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; and
consequently what infinitely varied diversities of structure might be of
use to each being under changing conditions of life. Can it then be
thought improbableseeing that variations useful to man have undoubtedly
occurredthat other variations useful in some way to each being in the
great and complex battle of lifeshould occur in the course of many
successive generations? If such do occurcan we doubt (remembering that
many more individuals are born than can possibly survive) that individuals
having any advantagehowever slightover otherswould have the best
chance of surviving and procreating their kind? On the other handwe may
feel sure that any variation in the least degree injurious would be rigidly
destroyed. This preservation of favourable individual differences and
variationsand the destruction of those which are injuriousI have called
Natural Selectionor the Survival of the Fittest. Variations neither
useful nor injurious would not be affected by natural selectionand would
be left either a fluctuating elementas perhaps we see in certain
polymorphic speciesor would ultimately become fixedowing to the nature
of the organism and the nature of the conditions.

Several writers have misapprehended or objected to the term Natural
Selection. Some have even imagined that natural selection induces
variabilitywhereas it implies only the preservation of such variations as
arise and are beneficial to the being under its conditions of life. No one
objects to agriculturists speaking of the potent effects of man's
selection; and in this case the individual differences given by nature
which man for some object selectsmust of necessity first occur. Others
have objected that the term selection implies conscious choice in the
animals which become modified; and it has even been urged thatas plants
have no volitionnatural selection is not applicable to them! In the
literal sense of the wordno doubtnatural selection is a false term; but
who ever objected to chemists speaking of the elective affinities of the
various elements?--and yet an acid cannot strictly be said to elect the
base with which it in preference combines. It has been said that I speak
of natural selection as an active power or Deity; but who objects to an
author speaking of the attraction of gravity as ruling the movements of the
planets? Every one knows what is meant and is implied by such metaphorical
expressions; and they are almost necessary for brevity. So again it is
difficult to avoid personifying the word Nature; but I mean by natureonly
the aggregate action and product of many natural lawsand by laws the
sequence of events as ascertained by us. With a little familiarity such
superficial objections will be forgotten.

We shall best understand the probable course of natural selection by taking
the case of a country undergoing some slight physical changefor instance
of climate. The proportional numbers of its inhabitants will almost
immediately undergo a changeand some species will probably become
extinct. We may concludefrom what we have seen of the intimate and
complex manner in which the inhabitants of each country are bound together
that any change in the numerical proportions of the inhabitants
independently of the change of climate itselfwould seriously affect the

others. If the country were open on its bordersnew forms would certainly
immigrateand this would likewise seriously disturb the relations of some
of the former inhabitants. Let it be remembered how powerful the influence
of a single introduced tree or mammal has been shown to be. But in the
case of an islandor of a country partly surrounded by barriersinto
which new and better adapted forms could not freely enterwe 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; for
had the area been open to immigrationthese same places would have been
seized on by intruders. In such casesslight modificationswhich in any
way favoured the individuals of any speciesby better adapting them to
their altered conditionswould tend to be preserved; and natural selection
would have free scope for the work of improvement.

We have good reason to believeas shown in the first chapterthat changes
in the conditions of life give a tendency to increased variability; and in
the foregoing cases the conditions the changedand this would manifestly
be favourable to natural selectionby affording a better chance of the
occurrence of profitable variations. Unless such occurnatural selection
can do nothing. Under the term of "variations it must never be forgotten
that mere individual differences are included. As man can produce a great
result with his domestic animals and plants by adding up in any given
direction individual differences, so could natural selection, but far more
easily from having incomparably longer time for action. Nor do I believe
that any great physical change, as of climate, or any unusual degree of
isolation, to check immigration, 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. For as all the inhabitants of
each country are struggling together with nicely balanced forces, extremely
slight modifications in the structure or habits of one species would often
give it an advantage over others; and still further modifications of the
same kind would often still further increase the advantage, as long as the
species continued under the same conditions of life and profited by similar
means of subsistence and defence. 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, that none of them could be still
better adapted or improved; for in all countries, the natives have been so
far conquered by naturalised productions that they have allowed some
foreigners to take firm possession of the land. And as foreigners have
thus in every country beaten some of the natives, we may safely conclude
that the natives might have been modified with advantage, so as to have
better resisted the intruders.

As man can produce, and certainly has produced, a great result by his
methodical and unconscious means of selection, what may not natural
selection effect? Man can act only on external and visible characters:
Nature, if I may be allowed to personify the natural preservation or
survival of the fittest, cares nothing for appearances, except in so far as
they are useful to any being. She can act on every internal organ, on
every shade of constitutional difference, on the whole machinery of life.
Man selects only for his own good; Nature only for that of the being which
she tends. Every selected character is fully exercised by her, as is
implied by the fact of their selection. Man keeps the natives of many
climates in the same country. He seldom exercises each selected character
in some peculiar and fitting manner; he feeds a long and a short-beaked
pigeon on the same food; he does not exercise a long-backed or long-legged
quadruped in any peculiar manner; he exposes sheep with long and short wool
to the same climate; does not allow the most vigorous males to struggle for
the females; he does not rigidly destroy all inferior animals, but protects
during each varying season, as far as lies in his power, all his
productions. He often begins his selection by some half-monstrous form, or
at least by some modification prominent enough to catch the eye or to be
plainly useful to him. Under nature, the slightest differences of
structure or constitution may well turn the nicely-balanced scale in the

struggle for life, and so be preserved. How fleeting are the wishes and
efforts of man! How short his time, and consequently how poor will be his
results, compared with those accumulated by Nature during whole geological
periods! Can we wonder, then, that Nature's productions should be far
truer" in character than man's productions; that they should be infinitely
better adapted to the most complex conditions of lifeand should plainly
bear the stamp of far higher workmanship?

It may metaphorically be said that natural selection is daily and hourly
scrutinisingthroughout the worldthe slightest variations; rejecting
those that are badpreserving and adding up all that are good; silently
improvement of each organic being in relation to its organic and inorganic
conditions of life. We see nothing of these slow changes in progress
until the hand of time has marked the long lapse of agesand 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
speciesa varietywhen once formed must againperhaps after a long
interval of timevary or present individual differences of the same
favourable nature as before; and these must again be preservedand so
onwardstep by step. Seeing that individual differences of the same kind
perpetually recurthis can hardly be considered as an unwarrantable
assumption. But whether it is truewe can judge only by seeing how far
the hypothesis accords with and explains the general phenomena of nature.
On the other handthe ordinary belief that the amount of possible
variation is a strictly limited quantityis likewise a simple assumption.

Although natural selection can act only through and for the good of each
beingyet characters and structureswhich we are apt to consider as of
very trifling importancemay thus be acted on. When we see leaf-eating
insects greenand bark-feeders mottled-grey; the alpine ptarmigan white in
winterthe red-grouse the colour of heatherwe must believe that these
tints are of service to these birds and insects in preserving them from
danger. Grouseif not destroyed at some period of their liveswould
increase in countless numbers; they are known to suffer largely from birds
of prey; and hawks are guided by eyesight to their prey--so much so that
on parts of the continent persons are warned not to keep white pigeonsas
being the most liable to destruction. Hence natural selection might be
effective in giving the proper colour to each kind of grouseand in
keeping that colourwhen once acquiredtrue and constant. Nor ought we
to think that the occasional destruction of an animal of any particular
colour would produce little effect; we should remember how essential it is
in a flock of white sheep to destroy a lamb with the faintest trace of
black. We have seen how the colour of hogswhich feed on the "paint-root"
in Virginiadetermines whether they shall live or die. In plantsthe
down on the fruit and the colour of the flesh are considered by botanists
as characters of the most trifling importance; yet we hear from an
excellent horticulturistDowningthat in the United States smooth-skinned
fruits suffer far more from a beetlea Curculiothan those with down;
that purple plums suffer far more from a certain disease than yellow plums;
whereas another disease attacks yellow-fleshed peaches far more than those
with other coloured flesh. Ifwith all the aids of artthese slight
differences make a great difference in cultivating the several varieties
assuredlyin a state of naturewhere the trees would have to struggle
with other trees and with a host of enemiessuch differences would
effectually settle which varietywhether a smooth or downya yellow or a
purple-fleshed fruitshould succeed.

In looking at many small points of difference between specieswhichas
far as our ignorance permits us to judgeseem quite unimportantwe must
not forget that climatefoodetc.have no doubt produced some direct
effect. It is also necessary to bear in mind thatowing to the law of

correlationwhen one part varies and the variations are accumulated
through natural selectionother modificationsoften of the most
unexpected naturewill ensue.

As we see that those variations whichunder domesticationappear at any
particular period of lifetend to reappear in the offspring at the same
period; for instancein the shapesize and flavour of the seeds of the
many varieties of our culinary and agricultural plants; in the caterpillar
and cocoon stages of the varieties of the silkworm; in the eggs of poultry
and in the colour of the down of their chickens; in the horns of our sheep
and cattle when nearly adult; so in a state of nature natural selection
will be enabled to act on and modify organic beings at any ageby the
accumulation of variations profitable at that ageand by their inheritance
at a corresponding age. If it profit a plant to have its seeds more and
more widely disseminated by the windI can see no greater difficulty in
this being effected through natural selectionthan in the cotton-planter
increasing and improving by selection the down in the pods on his
cotton-trees. Natural selection may modify and adapt the larva of an
insect to a score of contingencieswholly different from those which
concern the mature insect; and these modifications may affectthrough
correlationthe structure of the adult. Soconverselymodifications in
the adult may affect the structure of the larva; but in all cases natural
selection will ensure that they shall not be injurious: for if they were
sothe species would become extinct.

Natural selection will modify the structure of the young in relation to the
parent and of the parent in relation to the young. In social animals it
will adapt the structure of each individual for the benefit of the whole
community; if the community profits by the selected change. What natural
selection cannot dois to modify the structure of one specieswithout
giving it any advantagefor the good of another species; and though
statements to this effect may be found in works of natural historyI
cannot find one case which will bear investigation. A structure used only
once in an animal's lifeif of high importance to itmight be modified to
any extent by natural selection; for instancethe great jaws possessed by
certain insectsused exclusively for opening the cocoon--or the hard tip
to the beak of unhatched birdsused for breaking the eggs. It has been
assertedthat of the best short-beaked tumbler-pigeons a greater number
perish in the egg than are able to get out of it; so that fanciers assist
in the act of hatching. Nowif nature had to make the beak of a
full-grown pigeon very short for the bird's own advantagethe process of
modification would be very slowand there would be simultaneously the most
rigorous selection of all the young birds within the eggwhich had the
most powerful and hardest beaksfor all with weak beaks would inevitably
perish: ormore delicate and more easily broken shells might be selected
the thickness of the shell being known to vary like every other structure.

It may be well here to remark that with all beings there must be much
fortuitous destructionwhich can have little or no influence on the course
of natural selection. For instancea vast number of eggs or seeds are
annually devouredand these could be modified through natural selection
only if they varied in some manner which protected them from their enemies.
Yet many of these eggs or seeds would perhapsif not destroyedhave
yielded individuals better adapted to their conditions of life than any of
those which happened to survive. So again a vast number of mature animals
and plantswhether or not they be the best adapted to their conditions
must be annually destroyed by accidental causeswhich 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. But let the
destruction of the adults be ever so heavyif 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 survivethe best
adapted individualssupposing that there is any variability in a

favourable directionwill tend to propagate their kind in larger numbers
than the less well adapted. If the numbers be wholly kept down by the
causes just indicatedas will often have been the casenatural selection
will be powerless in certain beneficial directions; but this is no valid
objection to its efficiency at other times and in other ways; 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.


Inasmuch as peculiarities often appear under domestication in one sex and
become hereditarily attached to that sexso no doubt it will be under
nature. Thus it is rendered possible for the two sexes to be modified
through natural selection in relation to different habits of lifeas is
sometimes the case; or for one sex to be modified in relation to the other
sexas commonly occurs. This leads me to say a few words on what I have
called sexual selection. This form of selection dependsnot on a struggle
for existence in relation to other organic beings or to external
conditionsbut on a struggle between the individuals of one sexgenerally
the malesfor the possession of the other sex. The result is not death to
the unsuccessful competitorbut few or no offspring. Sexual selection is
thereforeless rigorous than natural selection. Generallythe most
vigorous malesthose which are best fitted for their places in nature
will leave most progeny. But in many cases victory depends not so much on
general vigourbut on having special weaponsconfined to the male sex. A
hornless stag or spurless cock would have a poor chance of leaving numerous
offspring. Sexual selectionby always allowing the victor to breedmight
surely give indomitable couragelength of spurand strength to the wing
to strike in the spurred legin nearly the same manner as does the brutal
cockfighter by the careful selection of his best cocks. How low in the
scale of nature the law of battle descends I know not; male alligators have
been described as fightingbellowingand whirling roundlike Indians in
a war-dancefor the possession of the females; male salmons have been
observed fighting all day long; male stag-beetles sometimes bear wounds
from the huge mandibles of other males; the males of certain hymenopterous
insects have been frequently seen by that inimitable observer M. Fabre
fighting for a particular female who sits byan apparently unconcerned
beholder of the struggleand then retires with the conqueror. The war is
perhapsseverest between the males of polygamous animalsand these seem
oftenest provided with special weapons. The males of carnivorous animals
are already well armed; though to them and to othersspecial means of
defence may be given through means of sexual selectionas the mane of the
lionand the hooked jaw to the male salmon; for the shield may be as
important for victory as the sword or spear.

Among birdsthe contest is often of a more peaceful character. All those
who have attended to the subjectbelieve that there is the severest
rivalry between the males of many species to attractby singingthe
females. The rock-thrush of Guianabirds of paradiseand some others
congregateand successive males display with the most elaborate careand
show off in the best mannertheir gorgeous plumage; they likewise perform
strange antics before the femaleswhichstanding by as spectatorsat
last choose the most attractive partner. Those who have closely attended
to birds in confinement well know that they often take individual
preferences and dislikes: thus Sir R. Heron has described how a pied
peacock was eminently attractive to all his hen birds. I cannot here enter
on the necessary details; but if man can in a short time give beauty and an
elegant carriage to his bantamsaccording to his standard of beautyI can
see no good reason to doubt that female birdsby selectingduring
thousands of generationsthe most melodious or beautiful malesaccording
to their standard of beautymight produce a marked effect. Some
well-known lawswith respect to the plumage of male and female birdsin
comparison with the plumage of the youngcan partly be explained through
the action of sexual selection on variations occurring at different ages

and transmitted to the males alone or to both sexes at corresponding ages;
but I have not space here to enter on this subject.

Thus it isas I believethat when the males and females of any animal
have the same general habits of lifebut differ in structurecolouror
ornamentsuch differences have been mainly caused by sexual selection:
that isby individual males having hadin successive generationssome
slight advantage over other malesin their weaponsmeans of defenceor
charms; which they have transmitted to their male offspring alone. YetI
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 sexwhich apparently have not been augmented through selection by
man. The tuft of hair on the breast of the wild turkey-cock cannot be of
any useand it is doubtful whether it can be ornamental in the eyes of the
female bird; indeedhad the tuft appeared under domestication it would
have been called a monstrosity.


In order to make it clear howas I believenatural selection actsI must
beg permission to give one or two imaginary illustrations. Let us take the
case of a wolfwhich preys on various animalssecuring some by craft
some by strengthand some by fleetness; and let us suppose that the
fleetest preya deer for instancehad from any change in the country
increased in numbersor that other prey had decreased in numbersduring
that season of the year when the wolf was hardest pressed for food. Under
such circumstances the swiftest and slimmest wolves have the best chance of
survivingand so be preserved or selectedprovided always that they
retained strength to master their prey at this or some other period of the
yearwhen they were compelled to prey on other animals. I can see no more
reason to doubt that this would be the resultthan that man should be able
to improve the fleetness of his greyhounds by careful and methodical
selectionor by that kind of unconscious selection which follows from each
man trying to keep the best dogs without any thought of modifying the
breed. I may add thataccording to Mr. Piercethere are two varieties of
the wolf inhabiting the Catskill Mountainsin the United Statesone with
a light greyhound-like formwhich pursues deerand the other more bulky
with shorter legswhich more frequently attacks the shepherd's flocks.

Even without any change in the proportional numbers of the animals on which
our wolf preyeda cub might be born with an innate tendency to pursue
certain kinds of prey. Nor can this be thought very improbable; for we
often observe great differences in the natural tendencies of our domestic
animals; one catfor instancetaking to catch ratsanother mice; one
cataccording to Mr. St. Johnbringing home winged gameanother hares or
rabbitsand another hunting on marshy ground and almost nightly catching
woodcocks or snipes. The tendency to catch rats rather than mice is known
to be inherited. Nowif any slight innate change of habit or of structure
benefited an individual wolfit would have the best chance of surviving
and of leaving offspring. Some of its young would probably inherit the
same habits or structureand by the repetition of this processa new
variety might be formed which would either supplant or coexist with the
parent-form of wolf. Oragainthe wolves inhabiting a mountainous
districtand those frequenting the lowlandswould naturally be forced to
hunt different prey; and from the continued preservation of the individuals
best fitted for the two sitestwo varieties might slowly be formed. These
varieties would cross and blend where they met; but to this subject of
intercrossing we shall soon have to return. I may addthataccording to
Mr. Piercethere are two varieties of the wolf inhabiting the Catskill
Mountains in the United Statesone with a light greyhound-like formwhich
pursues deerand the other more bulkywith shorter legswhich more
frequently attacks the shepherd's flocks.

It should be observed that in the above illustrationI speak of the
slimmest individual wolvesand not of any single strongly marked variation
having been preserved. In former editions of this work I sometimes spoke
as if this latter alternative had frequently occurred. I saw the great
importance of individual differencesand this led me fully to discuss the
results of unconscious selection by manwhich depends on the preservation
of all the more or less valuable individualsand on the destruction of the
worst. I sawalsothat the preservation in a state of nature of any
occasional deviation of structuresuch as a monstrositywould be a rare
event; and thatif at first preservedit would generally be lost by
subsequent intercrossing with ordinary individuals. Neverthelessuntil
reading an able and valuable article in the "North British Review" (1867)
I did not appreciate how rarely single variationswhether slight or
strongly markedcould be perpetuated. The author takes the case of a pair
of animalsproducing during their lifetime two hundred offspringof
whichfrom various causes of destructiononly two on an average survive
to pro-create their kind. This is rather an extreme estimate for most of
the higher animalsbut by no means so for many of the lower organisms. He
then shows that if a single individual were bornwhich varied in some
mannergiving it twice as good a chance of life as that of the other
individualsyet the chances would be strongly against its survival.
Supposing it to survive and to breedand that half its young inherited the
favourable variation; stillas the Reviewer goes onto showthe young
would have only a slightly better chance of surviving and breeding; and
this chance would go on decreasing in the succeeding generations. The
justice of these remarks cannotI thinkbe disputed. Iffor instancea
bird of some kind could procure its food more easily by having its beak
curvedand if one were born with its beak strongly curvedand which
consequently flourishednevertheless there would be a very poor chance of
this one individual perpetuating its kind to the exclusion of the common
form; but there can hardly be a doubtjudging by what we see taking place
under domesticationthat this result would follow from the preservation
during many generations of a large number of individuals with more or less
strongly curved beaksand from the destruction of a still larger number
with the straightest beaks.

It should nothoweverbe overlooked that certain rather strongly marked
variationswhich no one would rank as mere individual differences
frequently recur owing to a similar organisation being similarly acted on-of
which fact numerous instances could be given with our domestic
productions. In such casesif the varying individual did not actually
transmit to its offspring its newly-acquired characterit would
undoubtedly transmit to themas long as the existing conditions remained
the samea still stronger tendency to vary in the same manner. 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. Or only a
thirdfifthor tenth part of the individuals may have been thus affected
of which fact several instances could be given. Thus Graba estimates that
about one-fifth of the guillemots in the Faroe Islands consist of a variety
so well markedthat it was formerly ranked as a distinct species under the
name of Uria lacrymans. In cases of this kindif the variation were of a
beneficial naturethe original form would soon be supplanted by the
modified formthrough the survival of the fittest.

To the effects of intercrossing in eliminating variations of all kindsI
shall have to recur; but it may be here remarked that most animals and
plants keep to their proper homesand do not needlessly wander about; we
see this even with migratory birdswhich almost always return to the same
spot. Consequently each newly-formed variety would generally be at first
localas seems to be the common rule with varieties in a state of nature;
so that similarly modified individuals would soon exist in a small body
togetherand would often breed together. If the new variety were
successful in its battle for lifeit would slowly spread from a central

districtcompeting with and conquering the unchanged individuals on the
margins of an ever-increasing circle.

It may be worth while to give another and more complex illustration of the
action of natural selection. Certain plants excrete sweet juice
apparently for the sake of eliminating something injurious from the sap:
this is effectedfor instanceby glands at the base of the stipules in
some Leguminosaeand at the backs of the leaves of the common laurel.
This juicethough small in quantityis greedily sought by insects; but
their visits do not in any way benefit the plant. Nowlet 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 dusted with pollenand would often transport it from one flower
to another. The flowers of two distinct individuals of the same species
would thus get crossed; and the act of crossingas can be fully proved
gives rise to vigorous seedlingswhich consequently would have the best
chance of flourishing and surviving. The plants which produced flowers
with the largest glands or nectariesexcreting most nectarwould oftenest
be visited by insectsand would oftenest be crossed; and so in the
long-run would gain the upper hand and form a local variety. The flowers
alsowhich had their stamens and pistils placedin relation to the size
and habits of the particular insect which visited themso as to favour in
any degree the transportal of the pollenwould likewise be favoured. We
might have taken the case of insects visiting flowers for the sake of
collecting pollen instead of nectar; and as pollen is formed for the sole
purpose of fertilisationits destruction appears to be a simple loss to
the plant; yet if a little pollen were carriedat first occasionally and
then habituallyby the pollen-devouring insects from flower to flowerand
a cross thus effectedalthough nine-tenths of the pollen were destroyed it
might still be a great gain to the plant to be thus robbed; and the
individuals which produced more and more pollenand had larger anthers
would be selected.

When our plantby the above process long continuedhad been rendered
highly attractive to insectsthey wouldunintentionally on their part
regularly carry pollen from flower to flower; and that they do this
effectually I could easily show by many striking facts. I will give only
oneas likewise illustrating one step in the separation of the sexes of
plants. Some holly-trees bear only male flowerswhich have four stamens
producing a rather small quantity of pollenand a rudimentary pistil;
other holly-trees bear only female flowers; these have a full-sized pistil
and four stamens with shrivelled anthersin which not a grain of pollen
can be detected. Having found a female tree exactly sixty yards from a
male treeI put the stigmas of twenty flowerstaken from different
branchesunder the microscopeand on allwithout exceptionthere were a
few pollen-grainsand on some a profusion. As the wind had set for
several days from the female to the male treethe pollen could not thus
have been carried. The weather had been cold and boisterous and therefore
not favourable to beesnevertheless every female flower which I examined
had been effectually fertilised by the beeswhich had flown from tree to
tree in search of nectar. But to return to our imaginary case; as soon as
the plant had been rendered so highly attractive to insects that pollen was
regularly carried from flower to floweranother process might commence.
No naturalist doubts the advantage of what has been called the
physiological division of labour;hence we may believe that it would be
advantageous to a plant to produce stamens alone in one flower or on one
whole plantand pistils alone in another flower or on another plant. In
plants under culture and placed under new conditions of lifesometimes the
male organs and sometimes the female organs become more or less impotent;
now if we suppose this to occur in ever so slight a degree under nature
thenas pollen is already carried regularly from flower to flowerand as
a more complete separation of the sexes of our plant would be advantageous
on the principle of the division of labourindividuals with this tendency
more and more increasedwould be continually favoured or selecteduntil

at last a complete separation of the sexes might be effected. It would
take up too much space to show the various stepsthrough dimorphism and
other meansby which the separation of the sexes in plants of various
kinds is apparently now in progress; but I may add that some of the species
of holly in North America areaccording to Asa Grayin an exactly
intermediate conditionoras he expresses itare more or less
dioeciously polygamous.

Let us now turn to the nectar-feeding insects; we may suppose the plant of
which we have been slowly increasing the nectar by continued selectionto
be a common plant; and that certain insects depended in main part on its
nectar for food. I could give many facts showing how anxious bees are to
save time: for instancetheir habit of cutting holes and sucking the
nectar at the bases of certain flowerswhich with a very little more
trouble they can enter by the mouth. Bearing such facts in mindit may be
believed that under certain circumstances individual differences in the
curvature or length of the proboscisetc.too slight to be appreciated by
usmight profit a bee or other insectso that certain individuals would
be able to obtain their food more quickly than others; and thus the
communities to which they belonged would flourish and throw off many swarms
inheriting the same peculiarities. 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; yet the hive-bee can easily suck
the nectar out of the incarnate cloverbut not out of the common red
cloverwhich is visited by humble-bees alone; so that whole fields of the
red clover offer in vain an abundant supply of precious nectar to the
hive-bee. That this nectar is much liked by the hive-bee is certain; for I
have repeatedly seenbut only in the autumnmany hive-bees sucking the
flowers through holes bitten in the base of the tube by humble bees. The
difference in the length of the corolla in the two kinds of cloverwhich
determines the visits of the hive-beemust be very trifling; for I have
been assured that when red clover has been mownthe flowers of the second
crop are somewhat smallerand that these are visited by many hive-bees. I
do not know whether this statement is accurate; nor whether another
published statement can be trustednamelythat the Ligurian beewhich is
generally considered a mere variety of the common hive-beeand which
freely crosses with itis able to reach and suck the nectar of the red
clover. Thusin a country where this kind of clover aboundedit might be
a great advantage to the hive-bee to have a slightly longer or differently
constructed proboscis. On the other handas the fertility of this clover
absolutely depends on bees visiting the flowersif humble-bees were to
become rare in any countryit might be a great advantage to the plant to
have a shorter or more deeply divided corollaso that the hive-bees should
be enabled to suck its flowers. Thus I can understand how a flower and a
bee might slowly becomeeither simultaneously or one after the other
modified and adapted to each other in the most perfect mannerby the
continued preservation of all the individuals which presented slight
deviations of structure mutually favourable to each other.

I am well aware that this doctrine of natural selectionexemplified in the
above imaginary instancesis open to the same objections which were first
urged against Sir Charles Lyell's noble views on "the modern changes of the
earthas illustrative of geology;" but we now seldom hear the agencies
which we see still at workspoken of as trifling and insignificantwhen
used in explaining the excavation of the deepest valleys or the formation
of long lines of inland cliffs. Natural selection acts only by the
preservation and accumulation of small inherited modificationseach
profitable to the preserved being; and as modern geology has almost
banished such views as the excavation of a great valley by a single
diluvial waveso will natural selection banish the belief of the continued
creation of new organic beingsor of any great and sudden modification in
their structure.


I must here introduce a short digression. In the case of animals and
plants with separated sexesit 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. Nevertheless there is reason to
believe that with all hermaphrodites two individualseither occasionally
or habituallyconcur for the reproduction of their kind. This view was
long ago doubtfully suggested by SprengelKnight and Kolreuter. We shall
presently see its importance; but I must here treat the subject with
extreme brevitythough I have the materials prepared for an ample
discussion. All vertebrate animalsall insects and some other large
groups of animalspair for each birth. Modern research has much
diminished the number of supposed hermaphrodites and of real hermaphrodites
a large number pair; that istwo individuals regularly unite for
reproductionwhich is all that concerns us. But still there are many
hermaphrodite animals which certainly do not habitually pairand a vast
majority of plants are hermaphrodites. What reasonit may be askedis
there for supposing in these cases that two individuals ever concur in
reproduction? As it is impossible here to enter on detailsI must trust
to some general considerations alone.

In the first placeI have collected so large a body of factsand made so
many experimentsshowingin accordance with the almost universal belief
of breedersthat with animals and plants a cross between different
varietiesor between individuals of the same variety but of another
straingives vigour and fertility to the offspring; and on the other hand
that CLOSE interbreeding diminishes vigour and fertility; 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; but that a
cross with another individual is occasionally--perhaps at long intervals of

On the belief that this is a law of naturewe canI thinkunderstand
several large classes of factssuch as the followingwhich on any other
view are inexplicable. Every hybridizer knows how unfavourable exposure to
wet is to the fertilisation of a floweryet what a multitude of flowers
have their anthers and stigmas fully exposed to the weather! If an
occasional cross be indispensablenotwithstanding that the plant's own
anthers and pistil stand so near each other as almost to ensure selffertilisation
the fullest freedom for the entrance of pollen from another
individual will explain the above state of exposure of the organs. Many
flowerson the other handhave their organs of fructification closely
enclosedas in the great papilionaceous or pea-family; but these almost
invariably present beautiful and curious adaptations in relation to the
visits of insects. So necessary are the visits of bees to many
papilionaceous flowersthat their fertility is greatly diminished if these
visits be prevented. Nowit is scarcely possible for insects to fly from
flower to flowerand not to carry pollen from one to the otherto the
great good of the plant. Insects act like a camel-hair penciland it is
sufficientto ensure fertilisationjust to touch with the same brush the
anthers of one flower and then the stigma of another; but it must not be
supposed that bees would thus produce a multitude of hybrids between
distinct species; for if a plant's own pollen and that from another species
are placed on the same stigmathe former is so prepotent that it
invariably and completely destroysas has been shown by Gartnerthe
influence of the foreign pollen.

When the stamens of a flower suddenly spring towards the pistilor slowly
move one after the other towards itthe contrivance seems adapted solely
to ensure self-fertilisation; and no doubt it is useful for this end: but
the agency of insects is often required to cause the stamens to spring
forwardas Kolreuter has shown to be the case with the barberry; and in
this very genuswhich seems to have a special contrivance for

self-fertilisationit is well known thatif closely-allied forms or
varieties are planted near each otherit is hardly possible to raise pure
seedlingsso largely do they naturally cross. In numerous other cases
far from self-fertilisation being favouredthere are special contrivances
which effectually prevent the stigma receiving pollen from its own flower
as I could show from the works of Sprengel and othersas well as from my
own observations: for instancein Lobelia fulgensthere is a really
beautiful and elaborate contrivance by which all the infinitely numerous
pollen-granules are swept out of the conjoined anthers of each flower
before the stigma of that individual flower is ready to receive them; and
as this flower is never visitedat least in my gardenby insectsit
never sets a seedthough by placing pollen from one flower on the stigma
of anotherI raise plenty of seedlings. Another species of Lobeliawhich
is visited by beesseeds freely in my garden. In very many other cases
though there is no special mechanical contrivance to prevent the stigma
receiving pollen from the same floweryetas Sprengeland more recently
Hildebrand and others have shownand as I can confirmeither the anthers
burst before the stigma is ready for fertilisationor the stigma is ready
before the pollen of that flower is readyso that these so-named
dichogamous plants have in fact separated sexesand must habitually be
crossed. So it is with the reciprocally dimorphic and trimorphic plants
previously alluded to. How strange are these facts! How strange that the
pollen and stigmatic surface of the same flowerthough placed so close
togetheras if for the very purpose of self-fertilisationshould 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 cabbageradishonionand of some other
plantsbe allowed to seed near each othera large majority of the
seedlings thus raised turn outas I foundmongrels: for instanceI
raised 233 seedling cabbages from some plants of different varieties
growing near each otherand of these only 78 were true to their kindand
some even of these were not perfectly true. Yet the pistil of each
cabbage-flower is surrounded not only by its own six stamens but by those
of the many other flowers on the same plant; and the pollen of each flower
readily gets on its stigma without insect agency; for I have found that
plants carefully protected from insects produce the full number of pods.
Howthencomes 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; and that this is part of the
general law of good being derived from the intercrossing of distinct
individuals of the same species. When distinct SPECIES are crossed the
case is reversedfor a plant's own pollen is always prepotent over foreign
pollen; but to this subject we shall return in a future chapter.

In the case of a large tree covered with innumerable flowersit may be
objected that pollen could seldom be carried from tree to treeand at most
only from flower to flower on the same tree; and flowers on the same tree
can be considered as distinct individuals only in a limited sense. I
believe this objection to be validbut that nature has largely provided
against it by giving to trees a strong tendency to bear flowers with
separated sexes. When the sexes are separatedalthough the male and
female flowers may be produced on the same treepollen must be regularly
carried from flower to flower; and this will give a better chance of pollen
being occasionally carried from tree to tree. That trees belonging to all
orders have their sexes more often separated than other plantsI find to
be the case in this country; and at my request Dr. Hooker tabulated the
trees of New Zealandand Dr. Asa Gray those of the United Statesand the
result was as I anticipated. On the other handDr. Hooker informs me that
the rule does not hold good in Australia: but if most of the Australian
trees are dichogamousthe same result would follow as if they bore flowers
with separated sexes. I have made these few remarks on trees simply to
call attention to the subject.

Turning for a brief space to animals: various terrestrial species are
hermaphroditessuch as the land-mollusca and earth-worms; but these all
pair. As yet I have not found a single terrestrial animal which can
fertilise itself. This remarkable factwhich offers so strong a contrast
with terrestrial plantsis intelligible on the view of an occasional cross
being indispensable; for owing to the nature of the fertilising element
there are no meansanalogous to the action of insects and of the wind with
plantsby which an occasional cross could be effected with terrestrial
animals without the concurrence of two individuals. Of aquatic animals
there are many self-fertilising hermaphrodites; but here the currents of
water offer an obvious means for an occasional cross. As in the case of
flowersI have as yet failedafter consultation with one of the highest
authoritiesnamelyProfessor Huxleyto discover a single hermaphrodite
animal with the organs of reproduction so perfectly enclosed that access
from withoutand the occasional influence of a distinct individualcan be
shown to be physically impossible. Cirripedes long appeared to me to
presentunder this point of viewa case of great difficulty; but I have
been enabledby a fortunate chanceto prove that two individualsthough
both are self-fertilising hermaphroditesdo sometimes cross.

It must have struck most naturalists as a strange anomaly thatboth with
animals and plantssome species of the same family and even of the same
genusthough agreeing closely with each other in their whole organisation
are hermaphroditesand some unisexual. But ifin factall
hermaphrodites do occasionally intercrossthe difference between them and
unisexual species isas far as function is concernedvery small.

>From these several considerations and from the many special facts which I
have collectedbut which I am unable here to giveit appears that with
animals and plants an occasional intercross between distinct individuals is
a very generalif not universallaw of nature.


This is an extremely intricate subject. A great amount of variability
under which term individual differences are always includedwill evidently
be favourable. A large number of individualsby giving a better chance
within any given period for the appearance of profitable variationswill
compensate for a lesser amount of variability in each individualand isI
believea highly important element of success. Though nature grants long
periods of time for the work of natural selectionshe does not grant an
indefinite period; for as all organic beings are striving to seize on each
place in the economy of natureif any one species does not become modified
and improved in a corresponding degree with its competitors it will be
exterminated. Unless favourable variations be inherited by some at least
of the offspringnothing can be effected by natural selection. The
tendency to reversion may often check or prevent the work; but as this
tendency has not prevented man from forming by selection numerous domestic
raceswhy should it prevail against natural selection?

In the case of methodical selectiona breeder selects for some definite
objectand if the individuals be allowed freely to intercrosshis work
will completely fail. But when many menwithout intending to alter the
breedhave a nearly common standard of perfectionand all try to procure
and breed from the best animalsimprovement surely but slowly follows from
this unconscious process of selectionnotwithstanding that there is no
separation of selected individuals. Thus it will be under nature; for
within a confined areawith some place in the natural polity not perfectly
occupiedall the individuals varying in the right directionthough in
different degreeswill tend to be preserved. But if the area be large
its several districts will almost certainly present different conditions of
life; and thenif the same species undergoes modification in different

districtsthe newly formed varieties will intercross on the confines of
each. But we shall see in the sixth chapter that intermediate varieties
inhabiting intermediate districtswill in the long run generally be
supplanted by one of the adjoining varieties. Intercrossing will chiefly
affect those animals which unite for each birth and wander muchand which
do not breed at a very quick rate. Hence with animals of this naturefor
instance birdsvarieties will generally be confined to separated
countries; and this I find to be the case. With hermaphrodite organisms
which cross only occasionallyand likewise with animals which unite for
each birthbut which wander little and can increase at a rapid ratea new
and improved variety might be quickly formed on any one spotand might
there maintain itself in a body and afterward spreadso that the
individuals of the new variety would chiefly cross together. On this
principle nurserymen always prefer saving seed from a large body of plants
as the chance of intercrossing is thus lessened.

Even with animals which unite for each birthand which do not propagate
rapidlywe must not assume that free intercrossing would always eliminate
the effects of natural selection; 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 distinctfrom haunting different stationsfrom
breeding at slightly different seasonsor from the individuals of each
variety preferring to pair together.

Intercrossing plays a very important part in nature by keeping the
individuals of the same speciesor of the same varietytrue and uniform
in character. It will obviously thus act far more efficiently with those
animals which unite for each birth; butas already statedwe have reason
to believe that occasional intercrosses take place with all animals and
plants. Even if these take place only at long intervals of timethe young
thus produced will gain so much in vigour and fertility over the offspring
from long-continued self-fertilisationthat they will have a better chance
of surviving and propagating their kind; and thus in the long run the
influence of crosseseven at rare intervalswill be great. With respect
to organic beings extremely low in the scalewhich do not propagate
sexuallynor conjugateand which cannot possibly intercrossuniformity
of character can be retained by them under the same conditions of life
only through the principle of inheritanceand through natural selection
which will destroy any individuals departing from the proper type. If the
conditions of life change and the form undergoes modificationuniformity
of character can be given to the modified offspringsolely by natural
selection preserving similar favourable variations.

Isolation also is an important element in the modification of species
through natural selection. In a confined or isolated areaif not very
largethe organic and inorganic conditions of life will generally be
almost uniform; so that natural selection will tend to modify all the
varying individuals of the same species in the same manner. Intercrossing
with the inhabitants of the surrounding districtswill also be thus
prevented. Moritz Wagner has lately published an interesting essay on this
subjectand has shown that the service rendered by isolation in preventing
crosses between newly-formed varieties is probably greater even than I
supposed. But from reasons already assigned I can by no means agree with
this naturalistthat migration and isolation are necessary elements for
the formation of new species. The importance of isolation is likewise
great in preventingafter any physical change in the conditionssuch as
of climateelevation of the landetc.the immigration of better adapted
organisms; 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.
Lastlyisolation will give time for a new variety to be improved at a slow
rate; and this may sometimes be of much importance. Ifhoweveran
isolated area be very smalleither from being surrounded by barriersor
from having very peculiar physical conditionsthe total number of the
inhabitants will be small; and this will retard the production of new

species through natural selectionby decreasing the chances of favourable
variations arising.

The mere lapse of time by itself does nothingeither for or against
natural selection. 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 speciesas if all the forms of life were necessarily
undergoing change through some innate law. Lapse of time is only so far
importantand its importance in this respect is greatthat it gives a
better chance of beneficial variations arising and of their being selected
accumulatedand fixed. It likewise tends to increase the direct action of
the physical conditions of lifein relation to the constitution of each

If we turn to nature to test the truth of these remarksand look at any
small isolated areasuch as an oceanic islandalthough the number of the
species inhabiting it is smallas we shall see in our chapter on
Geographical Distribution; yet of these species a very large proportion are
endemic--that ishave been produced there and nowhere else in the world.
Hence an oceanic island at first sight seems to have been highly favourable
for the production of new species. But we may thus deceive ourselvesfor
to ascertain whether a small isolated areaor a large open area like a
continenthas been most favourable for the production of new organic
formswe ought to make the comparison within equal times; and this we are
incapable of doing.

Although isolation is of great importance in the production of new species
on the whole I am inclined to believe that largeness of area is still more
importantespecially for the production of species which shall prove
capable of enduring for a long periodand of spreading widely. Throughout
a great and open areanot only will there be a better chance of favourable
variationsarising from the large number of individuals of the same
species there supportedbut the conditions of life are much more complex
from the large number of already existing species; and if some of these
many species become modified and improvedothers will have to be improved
in a corresponding degreeor they will be exterminated. Each new form
alsoas soon as it has been much improvedwill be able to spread over the
open and continuous areaand will thus come into competition with many
other forms. Moreovergreat areasthough now continuouswill often
owing to former oscillations of levelhave existed in a broken condition
so that the good effects of isolation will generallyto a certain extent
have concurred. FinallyI conclude thatalthough small isolated areas
have been in some respects highly favourable for the production of new
speciesyet that the course of modification will generally have been more
rapid on large areas; and what is more importantthat the new forms
produced on large areaswhich already have been victorious over many
competitorswill be those that will spread most widelyand will give rise
to the greatest number of new varieties and species. They will thus play a
more important part in the changing history of the organic world.

In accordance with this viewwe canperhapsunderstand some facts which
will be again alluded to in our chapter on Geographical Distribution; for
instancethe fact of the productions of the smaller continent of Australia
now yielding before those of the larger Europaeo-Asiatic area. Thusalso
it is that continental productions have everywhere become so largely
naturalised on islands. On a small islandthe race for life will have
been less severeand there will have been less modification and less
extermination. Hencewe can understand how it is that the flora of
Madeiraaccording to Oswald Heerresembles to a certain extent the
extinct tertiary flora of Europe. All fresh water basinstaken together
make a small area compared with that of the sea or of the land.
Consequentlythe competition between fresh water productions will have
been less severe than elsewhere; new forms will have been more slowly
producedand old forms more slowly exterminated. And it is in fresh water

basins that we find seven genera of Ganoid fishesremnants of a once
preponderant order: and in fresh water we find some of the most anomalous
forms now known in the worldas the Ornithorhynchus and Lepidosiren
whichlike fossilsconnect to a certain extent orders at present widely
separated in the natural scale. These anomalous forms may be called living
fossils; they have endured to the present dayfrom having inhabited a
confined areaand from having been exposed to less variedand therefore
less severecompetition.

To sum upas far as the extreme intricacy of the subject permitsthe
circumstances favourable and unfavourable for the production of new species
through natural selection. I conclude that for terrestrial productions a
large continental areawhich has undergone many oscillations of level
will have been the most favourable for the production of many new forms of
lifefitted to endure for a long time and to spread widely. While the
area existed as a continent the inhabitants will have been numerous in
individuals and kindsand will have been subjected to severe competition.
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 preventedso that new places in the polity of each island will have
had to be filled up by the modification of the old inhabitants; and time
will have been allowed for the varieties in each to become well modified
and perfected. Whenby renewed elevationthe islands were reconverted
into a continental areathere will again have been very severe
competition; the most favoured or improved varieties will have been enabled
to spread; there will have been much extinction of the less improved forms
and the relative proportional numbers of the various inhabitants of the
reunited continent will again have been changed; and again there will have
been a fair field for natural selection to improve still further the
inhabitantsand thus to produce new species.

That natural selection generally act with extreme slowness I fully admit.
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
changeswhich generally take place very slowlyand on the immigration of
better adapted forms being prevented. As some few of the old inhabitants
become modified the mutual relations of others will often be disturbed; and
this will create new placesready to be filled up by better adapted forms;
but all this will take place very slowly. Although all the individuals of
the same species differ in some slight degree from each otherit would
often be long before differences of the right nature in various parts of
the organisation might occur. The result would often be greatly retarded
by free intercrossing. Many will exclaim that these several causes are
amply sufficient to neutralise the power of natural selection. I do not
believe so. But I do believe that natural selection will generally act
very slowlyonly at long intervals of timeand only on a few of the
inhabitants of the same region. I further believe that these slow
intermittent results accord well with what geology tells us of the rate and
manner at which the inhabitants of the world have changed.

Slow though the process of selection may beif feeble man can do much by
artificial selectionI can see no limit to the amount of changeto the
beauty and complexity of the coadaptations between all organic beingsone
with another and with their physical conditions of lifewhich may have
been effected in the long course of time through nature's power of
selectionthat is by the survival of the fittest.


This subject will be more fully discussed in our chapter on Geology; but it
must here be alluded to from being intimately connected with natural

selection. Natural selection acts solely through the preservation of
variations in some way advantageouswhich consequently endure. Owing to
the high geometrical rate of increase of all organic beingseach area is
already fully stocked with inhabitantsand it follows from thisthat as
the favoured forms increase in numbersogenerallywill the less
favoured decrease and become rare. Rarityas geology tells usis the
precursor to extinction. We can see that any form which is represented by
few individuals will run a good chance of utter extinctionduring great
fluctuations in the nature or the seasonsor from a temporary increase in
the number of its enemies. But we may go further than this; for as new
forms are producedunless we admit that specific forms can go on
indefinitely increasing in numbermany old forms must become extinct.
That the number of specific forms has not indefinitely increasedgeology
plainly tells us; and we shall presently attempt to show why it is that the
number of species throughout the world has not become immeasurably great.

We have seen that the species which are most numerous in individuals have
the best chance of producing favourable variations within any given period.
We have evidence of thisin the facts stated in the second chapter
showing that it is the common and diffused or dominant species which offer
the greatest number of recorded varieties. Hencerare species will be
less quickly modified or improved within any given period; they will
consequently be beaten in the race for life by the modified and improved
descendants of the commoner species.

>From these several considerations I think it inevitably followsthat as
new species in the course of time are formed through natural selection
others will become rarer and rarerand finally extinct. The forms which
stand in closest competition with those undergoing modification and
improvementwill naturally suffer most. And we have seen in the chapter
on the Struggle for Existence that it is the most closely-allied
forms--varieties of the same speciesand species of the same genus or
related genera--whichfrom having nearly the same structureconstitution
and habitsgenerally come into the severest competition with each other.
Consequentlyeach new variety or speciesduring the progress of its
formationwill generally press hardest on its nearest kindredand tend to
exterminate them. We see the same process of extermination among our
domesticated productionsthrough the selection of improved forms by man.
Many curious instances could be given showing how quickly new breeds of
cattlesheep and other animalsand varieties of flowerstake the place
of older and inferior kinds. In Yorkshireit is historically known that
the ancient black cattle were displaced by the long-hornsand that these
were swept away by the short-horns(I quote the words of an agricultural
writer) "as if by some murderous pestilence."


The principlewhich I have designated by this termis of high importance
and explainsas I believeseveral important facts. In the first place
varietieseven strongly-marked onesthough 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. Nevertheless according to my viewvarieties are
species in the process of formationor areas I have called them
incipient species. Howthendoes the lesser difference between varieties
become augmented into the greater difference between species? That this
does habitually happenwe must infer from most of the innumerable species
throughout nature presenting well-marked differences; whereas varieties
the supposed prototypes and parents of future well-marked speciespresent
slight and ill-defined differences. Mere chanceas we may call itmight
cause one variety to differ in some character from its parentsand the
offspring of this variety again to differ from its parent in the very same
character and in a greater degree; but this alone would never account for
so habitual and large a degree of difference as that between the species of

the same genus.

As has always been my practiceI have sought light on this head from our
domestic productions. We shall here find something analogous. It will be
admitted that the production of races so different as short-horn and
Hereford cattlerace and cart horsesthe several breeds of pigeonsetc.
could never have been effected by the mere chance accumulation of similar
variations during many successive generations. In practicea fancier is
for instancestruck by a pigeon having a slightly shorter beak; another
fancier is struck by a pigeon having a rather longer beak; and on the
acknowledged principle that "fanciers do not and will not admire a medium
standardbut like extremes 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. Again, we
may suppose that at an early period of history, the men of one nation or
district required swifter horses, while those of another required stronger
and bulkier horses. The early differences would be very slight; but, in
the course of time, from the continued selection of swifter horses in the
one case, and of stronger ones in the other, the differences would become
greater, and would be noted as forming two sub-breeds. Ultimately after
the lapse of centuries, these sub-breeds would become converted into two
well-established and distinct breeds. As the differences became greater,
the inferior animals with intermediate characters, being neither very swift
nor very strong, would not have been used for breeding, and will thus have
tended to disappear. Here, then, we see in man's productions the action of
what may be called the principle of divergence, causing differences, at
first barely appreciable, steadily to increase, and the breeds to diverge
in character, both from each other and from their common parent.

But how, it may be asked, 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), from the simple circumstance that the more diversified
the descendants from any one species become in structure, constitution, and
habits, by so much will they be better enabled to seize on many and widely
diversified places in the polity of nature, and so be enabled to increase
in numbers.

We can clearly discern this in the case of animals with simple habits.
Take the case of a carnivorous quadruped, of which the number that can be
supported in any country has long ago arrived at its full average. If its
natural power of increase be allowed to act, 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, for instance, being enabled to feed on new kinds of prey, either
dead or alive; some inhabiting new stations, climbing trees, frequenting
water, and some perhaps becoming less carnivorous. The more diversified in
habits and structure the descendants of our carnivorous animals become, the
more places they will be enabled to occupy. What applies to one animal
will apply throughout all time to all animals--that is, if they vary--for
otherwise natural selection can effect nothing. So it will be with plants.
It has been experimentally proved, that if a plot of ground be sown with
one species of grass, and a similar plot be sown with several distinct
genera of grasses, a greater number of plants and a greater weight of dry
herbage can be raised in the latter than in the former case. 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. Hence, if any one species
of grass were to go on varying, and the varieties were continually selected
which differed from each other in the same manner, though in a very slight
degree, as do the distinct species and genera of grasses, a greater number
of individual plants of this species, including its modified descendants,
would succeed in living on the same piece of ground. And we know that each
species and each variety of grass is annually sowing almost countless
seeds; and is thus striving, as it may be said, to the utmost to increase
in number. Consequently, in the course of many thousand generations, the

most distinct varieties of any one species of grass would have the best
chance of succeeding and of increasing in numbers, and thus of supplanting
the less distinct varieties; and varieties, when rendered very distinct
from each other, take the rank of species.

The truth of the principle that the greatest amount of life can be
supported by great diversification of structure, is seen under many natural
circumstances. In an extremely small area, especially if freely open to
immigration, and where the contest between individual and individual must
be very severe, we always find great diversity in its inhabitants. For
instance, I found that a piece of turf, three feet by four in size, which
had been exposed for many years to exactly the same conditions, supported
twenty species of plants, and these belonged to eighteen genera and to
eight orders, which shows how much these plants differed from each other.
So it is with the plants and insects on small and uniform islets: also in
small ponds of fresh water. 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. Most of the animals
and plants which live close round any small piece of ground, could live on
it (supposing its nature not to be in any way peculiar), and may be said to
be striving to the utmost to live there; but, it is seen, that where they
come into the closest competition, the advantages of diversification of
structure, with the accompanying differences of habit and constitution,
determine that the inhabitants, which thus jostle each other most closely,
shall, as a general rule, belong to what we call different genera and

The same principle is seen in the naturalisation of plants through man's
agency in foreign lands. 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; for these are commonly looked at as
specially created and adapted for their own country. It might also,
perhaps, have been expected that naturalised plants would have belonged to
a few groups more especially adapted to certain stations in their new
homes. But the case is very different; and Alph. de Candolle has well
remarked, in his great and admirable work, that floras gain by
naturalisation, proportionally with the number of the native genera and
species, far more in new genera than in new species. To give a single
instance: in the last edition of Dr. Asa Gray's Manual of the Flora of
the Northern United States 260 naturalised plants are enumerated, and
these belong to 162 genera. We thus see that these naturalised plants are
of a highly diversified nature. They differ, moreover, to a large extent,
from the indigenes, for out of the 162 naturalised genera, no less than 100
genera are not there indigenous, and thus a large proportional addition is
made to the genera now living in the United States.

By considering the nature of the plants or animals which have in any
country struggled successfully with the indigenes, and have there become
naturalised, 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; and we may at least infer that diversification of structure,
amounting to new generic differences, would be profitable to them.

The advantage of diversification of structure in the inhabitants of the
same region is, in fact, 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. No physiologist doubts that a stomach by
being adapted to digest vegetable matter alone, or flesh alone, draws most
nutriment from these substances. So in the general economy of any land,
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. A set of animals, with their
organisation but little diversified, could hardly compete with a set more
perfectly diversified in structure. It may be doubted, for instance,

whether the Australian marsupials, which are divided into groups differing
but little from each other, and feebly representing, as Mr. Waterhouse and
others have remarked, our carnivorous, ruminant, and rodent mammals, could
successfully compete with these well-developed orders. In the Australian
mammals, we see the process of diversification in an early and incomplete
stage of development.


After the foregoing discussion, which has been much compressed, we may
assume that the modified descendants of any one species will succeed so
much the better as they become more diversified in structure, and are thus
enabled to encroach on places occupied by other beings. Now let us see how
this principle of benefit being derived from divergence of character,
combined with the principles of natural selection and of extinction, tends
to act.

The accompanying diagram will aid us in understanding this rather
perplexing subject. Let A to L represent the species of a genus large in
its own country; these species are supposed to resemble each other in
unequal degrees, as is so generally the case in nature, and as is
represented in the diagram by the letters standing at unequal distances. I
have said a large genus, because as we saw in the second chapter, on an
average more species vary in large genera than in small genera; and the
varying species of the large genera present a greater number of varieties.
We have, also, seen that the species, which are the commonest and most
widely-diffused, vary more than do the rare and restricted species. Let

(A) be a common, widely-diffused, and varying species, belonging to a genus
large in its own country. The branching and diverging dotted lines of
unequal lengths proceeding from (A), may represent its varying offspring.
The variations are supposed to be extremely slight, but of the most
diversified nature; they are not supposed all to appear simultaneously, but
often after long intervals of time; nor are they all supposed to endure for
equal periods. Only those variations which are in some way profitable will
be preserved or naturally selected. And here the importance of the
principle of benefit derived from divergence of character comes in; 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. When a dotted line reaches one of the horizontal lines,
and is there marked by a small numbered letter, a sufficient amount of
variation is supposed to have been accumulated to form it into a fairly
well-marked variety, such as would be thought worthy of record in a
systematic work.
The intervals between the horizontal lines in the diagram, may represent
each a thousand or more generations. After a thousand generations, species

(A) is supposed to have produced two fairly well-marked varieties, namely
a1 and m1. These two varieties will generally still be exposed to the same
conditions which made their parents variable, and the tendency to
variability is in itself hereditary; consequently they will likewise tend
to vary, and commonly in nearly the same manner as did their parents.
Moreover, these two varieties, being only slightly modified forms, will
tend to inherit those advantages which made their parent (A) more numerous
than most of the other inhabitants of the same country; they will also
partake of those more general advantages which made the genus to which the
parent-species belonged, a large genus in its own country. And all these
circumstances are favourable to the production of new varieties.
If, then, these two varieties be variable, the most divergent of their
variations will generally be preserved during the next thousand
generations. And after this interval, variety a1 is supposed in the
diagram to have produced variety a2, which will, owing to the principle of
divergence, differ more from (A) than did variety a1. Variety m1 is

supposed to have produced two varieties, namely m2 and s2, differing from
each other, and more considerably from their common parent (A). We may
continue the process by similar steps for any length of time; some of the
varieties, after each thousand generations, producing only a single
variety, but in a more and more modified condition, some producing two or
three varieties, and some failing to produce any. Thus the varieties or
modified descendants of the common parent (A), will generally go on
increasing in number and diverging in character. In the diagram the
process is represented up to the ten-thousandth generation, and under a
condensed and simplified form up to the fourteen-thousandth generation.

But I must here remark that I do not suppose that the process ever goes on
so regularly as is represented in the diagram, though in itself made
somewhat irregular, nor that it goes on continuously; it is far more
probable that each form remains for long periods unaltered, and then again
undergoes modification. Nor do I suppose that the most divergent varieties
are invariably preserved: a medium form may often long endure, and may or
may not produce more than one modified descendant; for natural selection
will always act according to the nature of the places which are either
unoccupied or not perfectly occupied by other beings; and this will depend
on infinitely complex relations. But as a general rule, the more
diversified in structure the descendants from any one species can be
rendered, the more places they will be enabled to seize on, and the more
their modified progeny will increase. In our diagram the line of
succession is broken at regular intervals by small numbered letters marking
the successive forms which have become sufficiently distinct to be recorded
as varieties. But these breaks are imaginary, and might have been inserted
anywhere, after intervals long enough to allow the accumulation of a
considerable amount of divergent variation.

As all the modified descendants from a common and widely-diffused species,
belonging to a large genus, will tend to partake of the same advantages
which made their parent successful in life, 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). The modified offspring from the later and more highly improved
branches in the lines of descent, will, it is probable, often take the
place of, and so destroy, 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. In some cases no doubt the process of
modification will be confined to a single line of descent, and the number
of modified descendants will not be increased; although the amount of
divergent modification may have been augmented. This case would be
represented in the diagram, if all the lines proceeding from (A) were
removed, excepting that from a1 to a10. In the same way the English
racehorse and English pointer have apparently both gone on slowly diverging
in character from their original stocks, without either having given off
any fresh branches or races.

After ten thousand generations, species (A) is supposed to have produced
three forms, a10, f10, and m10, which, from having diverged in character
during the successive generations, will have come to differ largely, but
perhaps unequally, from each other and from their common parent. If we
suppose the amount of change between each horizontal line in our diagram to
be excessively small, these three forms may still be only well-marked
varieties; but we have only to suppose the steps in the process of
modification to be more numerous or greater in amount, 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.
By continuing the same process for a greater number of generations (as
shown in the diagram in a condensed and simplified manner), we get eight
species, marked by the letters between a14 and m14, all descended from (A).
Thus, as I believe, species are multiplied and genera are formed.

In a large genus it is probable that more than one species would vary. In
the diagram I have assumed that a second species (I) has produced, by
analogous steps, after ten thousand generations, either two well-marked
varieties (w10 and z10) or two species, according to the amount of change
supposed to be represented between the horizontal lines. After fourteen
thousand generations, six new species, marked by the letters n14 to z14,
are supposed to have been produced. In any genus, the species which are
already very different in character from each other, will generally tend to
produce the greatest number of modified descendants; 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), and
the nearly extreme species (I), as those which have largely varied, and
have given rise to new varieties and species. The other nine species
(marked by capital letters) of our original genus, 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.

But during the process of modification, represented in the diagram, another
of our principles, namely that of extinction, will have played an important
part. 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, 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. 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, constitution
and structure. Hence all the intermediate forms between the earlier and
later states, that is between the less and more improved states of a the
same species, as well as the original parent-species itself, will generally
tend to become extinct. So it probably will be with many whole collateral
lines of descent, which will be conquered by later and improved lines. If,
however, the modified offspring of a species get into some distinct
country, or become quickly adapted to some quite new station, in which
offspring and progenitor do not come into competition, both may continue to

If, then, our diagram be assumed to represent a considerable amount of
modification, species (A) and all the earlier varieties will have become
extinct, being replaced by eight new species (a14 to m14); and species (I)
will be replaced by six (n14 to z14) new species.

But we may go further than this. The original species of our genus were
supposed to resemble each other in unequal degrees, as is so generally the
case in nature; species (A) being more nearly related to B, C, and D than
to the other species; and species (I) more to G, H, K, L, than to the
others. These two species (A and I), were also supposed to be very common
and widely diffused species, so that they must originally have had some
advantage over most of the other species of the genus. Their modified
descendants, fourteen in number at the fourteen-thousandth generation, 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, so
as to have become adapted to many related places in the natural economy of
their country. It seems, therefore, extremely probable that they will have
taken the places of, and thus exterminated, not only their parents (A) and
(I), but likewise some of the original species which were most nearly
related to their parents. Hence very few of the original species will have
transmitted offspring to the fourteen-thousandth generation. 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, has transmitted
descendants to this late stage of descent.

The new species in our diagram, descended from the original eleven species,
will now be fifteen in number. Owing to the divergent tendency of natural

selection, 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. The new species, moreover, will be allied to each
other in a widely different manner. Of the eight descendants from (A) the
three marked a14, q14, p14, will be nearly related from having recently
branched off from a10; b14 and f14, from having diverged at an earlier
period from a5, will be in some degree distinct from the three first-named
species; and lastly, o14, e14, and m14, will be nearly related one to the
other, but, from having diverged at the first commencement of the process
of modification, will be widely different from the other five species, and
may constitute a sub-genus or a distinct genus.

The six descendants from (I) will form two sub-genera or genera. But as
the original species (I) differed largely from (A), standing nearly at the
extreme end of the original genus, the six descendants from (I) will, owing
to inheritance alone, differ considerably from the eight descendants from
(A); the two groups, moreover, are supposed to have gone on diverging in
different directions. The intermediate species, also (and this is a very
important consideration), which connected the original species (A) and (I),
have all become, except (F), extinct, and have left no descendants. Hence
the six new species descended from (I), and the eight descendants from (A),
will have to be ranked as very distinct genera, or even as distinct

Thus it is, as I believe, that two or more genera are produced by descent
with modification, from two or more species of the same genus. And the two
or more parent-species are supposed to be descended from some one species
of an earlier genus. In our diagram this is indicated by the broken lines
beneath the capital letters, converging in sub-branches downwards towards a
single point; this point represents a species, the supposed progenitor of
our several new sub-genera and genera.

It is worth while to reflect for a moment on the character of the new
species F14, which is supposed not to have diverged much in character, but
to have retained the form of (F), either unaltered or altered only in a
slight degree. In this case its affinities to the other fourteen new
species will be of a curious and circuitous nature. Being descended from a
form that stood between the parent-species (A) and (I), now supposed to be
extinct and unknown, it will be in some degree intermediate in character
between the two groups descended from these two species. But as these two
groups have gone on diverging in character from the type of their parents,
the new species (F14) will not be directly intermediate between them, but
rather between types of the two groups; and every naturalist will be able
to call such cases before his mind.

In the diagram each horizontal line has hitherto been supposed to represent
a thousand generations, but each may represent a million or more
generations; it may also represent a section of the successive strata of
the earth's crust including extinct remains. We shall, when we come to our
chapter on geology, have to refer again to this subject, and I think we
shall then see that the diagram throws light on the affinities of extinct
beings, which, though generally belonging to the same orders, families, or
genera, with those now living, yet are often, in some degree, intermediate
in character between existing groups; and we can understand this fact, for
the extinct species lived at various remote epochs when the branching lines
of descent had diverged less.

I see no reason to limit the process of modification, as now explained, to
the formation of genera alone. If, in the diagram, we suppose the amount
of change represented by each successive group of diverging dotted lines to
be great, the forms marked a14 to p14, those marked b14 and f14, and those
marked o14 to m14, will form three very distinct genera. We shall also
have two very distinct genera descended from (I), differing widely from the
descendants of (A). These two groups of genera will thus form two distinct

families, or orders, according to the amount of divergent modification
supposed to be represented in the diagram. And the two new families, or
orders, are descended from two species of the original genus; and these are
supposed to be descended from some still more ancient and unknown form.

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


Natural selection acts exclusively by the preservation and accumulation of
variations, which are beneficial under the organic and inorganic conditions
to which each creature is exposed at all periods of life. The ultimate
result is that each creature tends to become more and more improved in
relation to its conditions. This improvement inevitably leads to the
gradual advancement of the organisation of the greater number of living
beings throughout the world. But here we enter on a very intricate
subject, for naturalists have not defined to each other's satisfaction what
is meant by an advance in organisation. Among the vertebrata the degree of
intellect and an approach in structure to man clearly come into play. 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 there are cases, as with certain
parasitic crustaceans, in which several parts of the structure become less
perfect, so that the mature animal cannot be called higher than its larva.
Von Baer's standard seems the most widely applicable and the best, namely,
the amount of differentiation of the parts of the same organic being, in
the adult state, as I should be inclined to add, and their specialisation
for different functions; or, as Milne Edwards would express it, the
completeness of the division of physiological labour. But we shall see how
obscure this subject is if we look, for instance, to fishes, among which
some naturalists rank those as highest which, like the sharks, approach

nearest to amphibians; while other naturalists rank the common bony or
teleostean fishes as the highest, inasmuch as they are most strictly fishlike,
and differ most from the other vertebrate classes. We see still more
plainly the obscurity of the subject by turning to plants, among which the
standard of intellect is of course quite excluded; and here some botanists
rank those plants as highest which have every organ, as sepals, petals,
stamens and pistils, fully developed in each flower; whereas other
botanists, probably with more truth, look at the plants which have their
several organs much modified and reduced in number as the highest.

If we take as the standard of high organisation, the amount of
differentiation and specialisation of the several organs in each being when
adult (and this will include the advancement of the brain for intellectual
purposes), natural selection clearly leads towards this standard: for all
physiologists admit that the specialisation of organs, inasmuch as in this
state they perform their functions better, is an advantage to each being;
and hence the accumulation of variations tending towards specialisation is
within the scope of natural selection. On the other hand, we can see,
bearing in mind that all organic beings are striving to increase at a high
ratio and to seize on every unoccupied or less well occupied place in the
economy of nature, that it is quite possible for natural selection
gradually to fit a being to a situation in which several organs would be
superfluous or useless: in such cases there would be retrogression in the
scale of organisation. 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.

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; 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, who believed
in an innate and inevitable tendency towards perfection in all organic
beings, 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. Science has not as yet proved the truth of this
belief, whatever the future may reveal. On our theory the continued
existence of lowly organisms offers no difficulty; for natural selection,
or the survival of the fittest, does not necessarily include progressive
development--it only takes advantage of such variations as arise and are
beneficial to each creature under its complex relations of life. And it
may be asked what advantage, as far as we can see, would it be to an
infusorian animalcule--to an intestinal worm--or even to an earth-worm, to
be highly organised. If it were no advantage, these forms would be left,
by natural selection, unimproved or but little improved, and might remain
for indefinite ages in their present lowly condition. And geology tells us
that some of the lowest forms, as the infusoria and rhizopods, have
remained for an enormous period in nearly their present state. 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; for every
naturalist who has dissected some of the beings now ranked as very low in
the scale, must have been struck with their really wondrous and beautiful

Nearly the same remarks are applicable, if we look to the different grades
of organisation within the same great group; for instance, in the
vertebrata, to the co-existence of mammals and fish--among mammalia, to the
co-existence of man and the ornithorhynchus--among fishes, to the coexistence
of the shark and the lancelet (Amphioxus), which latter fish in
the extreme simplicity of its structure approaches the invertebrate
classes. But mammals and fish hardly come into competition with each
other; the advancement of the whole class of mammals, or of certain members
in this class, to the highest grade would not lead to their taking the
place of fishes. Physiologists believe that the brain must be bathed by

warm blood to be highly active, and this requires aerial respiration; so
that warm-blooded mammals when inhabiting the water lie under a
disadvantage in having to come continually to the surface to breathe. With
fishes, members of the shark family would not tend to supplant the
lancelet; for the lancelet, as I hear from Fritz Muller, has as sole
companion and competitor on the barren sandy shore of South Brazil, an
anomalous annelid. The three lowest orders of mammals, namely, marsupials,
edentata, and rodents, co-exist in South America in the same region with
numerous monkeys, and probably interfere little with each other. Although
organisation, on the whole, may have advanced and be still advancing
throughout the world, yet the scale will always present many degrees of
perfection; for the high advancement of certain whole classes, or of
certain members of each class, does not at all necessarily lead to the
extinction of those groups with which they do not enter into close
competition. In some cases, as we shall hereafter see, lowly organised
forms appear to have been preserved to the present day, from inhabiting
confined or peculiar stations, where they have been subjected to less
severe competition, and where their scanty numbers have retarded the chance
of favourable variations arising.

Finally, I believe that many lowly organised forms now exist throughout the
world, from various causes. In some cases variations or individual
differences of a favourable nature may never have arisen for natural
selection to act on and accumulate. In no case, probably, has time
sufficed for the utmost possible amount of development. In some few cases
there has been what we must call retrogression or organisation. 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, as being of a more delicate nature, and more liable to be put
out of order and injured.

Looking to the first dawn of life, when all organic beings, as we may
believe, presented the simplest structure, how, it has been asked, could
the first step in the advancement or differentiation of parts have arisen?
Mr. Herbert Spencer would probably answer that, as soon as simple
unicellular organisms came by growth or division to be compounded of
several cells, or became attached to any supporting surface, his law that
homologous units of any order become differentiated in proportion as their
relations to incident forces become different" would come into action. But
as we have no facts to guide usspeculation on the subject is almost
useless. It ishoweveran error to suppose that there would be no
struggle for existenceandconsequentlyno natural selectionuntil many
forms had been produced: variations in a single species inhabiting an
isolated station might be beneficialand thus the whole mass of
individuals might be modifiedor two distinct forms might arise. Butas
I remarked towards the close of the introductionno one ought to feel
surprise at much remaining as yet unexplained on the origin of speciesif
we make due allowance for our profound ignorance on the mutual relations of
the inhabitants of the world at the present timeand still more so during
past ages.


Mr. H.C. Watson thinks that I have overrated the importance of divergence
of character (in whichhoweverhe apparently believes)and that
convergenceas it may be calledhas likewise played a part. If two
species belonging to two distinct though allied generahad both produced a
large number of new and divergent formsit is conceivable that these might
approach each other so closely that they would have all to be classed under
the same genus; and thus the descendants of two distinct genera would
converge into one. 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. The shape of a crystal is
determined solely by the molecular forcesand it is not surprising that

dissimilar substances should sometimes assume the same form; but with
organic beings we should bear in mind that the form of each depends on an
infinitude of complex relationsnamely on the variations which have
arisenthese being due to causes far too intricate to be followed out--on
the nature of the variations which have been preserved or selectedand
this depends on the surrounding physical conditionsand in a still higher
degree on the surrounding organisms with which each being has come into
competition--and lastlyon inheritance (in itself a fluctuating element)
from innumerable progenitorsall of which have had their forms determined
through equally complex relations. It is incredible that the descendants
of two organismswhich had originally differed in a marked mannershould
ever afterwards converge so closely as to lead to a near approach to
identity throughout their whole organisation. If this had occurredwe
should meet with the same formindependently of genetic connection
recurring in widely separated geological formations; and the balance of
evidence is opposed to any such an admission.

Mr. Watson has also objected that the continued action of natural
selectiontogether with divergence of characterwould tend to make an
indefinite number of specific forms. As far as mere inorganic conditions
are concernedit seems probable that a sufficient number of species would
soon become adapted to all considerable diversities of heatmoisture
etc.; but I fully admit that the mutual relations of organic beings are
more important; and as the number of species in any country goes on
increasingthe organic conditions of life must become more and more
complex. Consequently there seems at first no limit to the amount of
profitable diversification of structureand therefore no limit to the
number of species which might be produced. We do not know that even the
most prolific area is fully stocked with specific forms: at the Cape of
Good Hope and in Australiawhich support such an astonishing number of
speciesmany European plants have become naturalised. But geology shows
usthat from an early part of the tertiary period the number of species of
shellsand that from the middle part of this same periodthe number of
mammals has not greatly or at all increased. 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
depending so largely as it does on physical conditions; thereforeif an
area be inhabited by very many specieseach or nearly each species will be
represented by few individuals; and such species will be liable to
extermination from accidental fluctuations in the nature of the seasons or
in the number of their enemies. The process of extermination in such cases
would be rapidwhereas the production of new species must always be slow.
Imagine the extreme case of as many species as individuals in Englandand
the first severe winter or very dry summer would exterminate thousands on
thousands of species. Rare speciesand each species will become rare if
the number of species in any country becomes indefinitely increasedwill
on the principal often explainedpresent within a given period few
favourable variations; consequentlythe process of giving birth to new
specific forms would thus be retarded. When any species becomes very rare
close interbreeding will help to exterminate it; authors have thought that
this comes into play in accounting for the deterioration of the aurochs in
Lithuaniaof red deer in Scotland and of bears in Norwayetc. Lastly
and this I am inclined to think is the most important elementa dominant
specieswhich has already beaten many competitors in its own homewill
tend to spread and supplant many others. Alph. de Candolle has shown that
those species which spread widely tend generally to spread VERY widely
consequently they will tend to supplant and exterminate several species in
several areasand thus check the inordinate increase of specific forms
throughout the world. Dr. Hooker has recently shown that in the southeast
corner of Australiawhereapparentlythere are many invaders from
different quarters of the globethe endemic Australian species have been
greatly reduced in number. How much weight to attribute to these several
considerations I will not pretend to say; but conjointly they must limit in
each country the tendency to an indefinite augmentation of specific forms.


If under changing conditions of life organic beings present individual
differences in almost every part of their structureand this cannot be
disputed; if there beowing to their geometrical rate of increasea
severe struggle for life at some ageseason or yearand this certainly
cannot be disputed; thenconsidering the infinite complexity of the
relations of all organic beings to each other and to their conditions of
lifecausing an infinite diversity in structureconstitutionand habits
to be advantageous to themit would be a most extraordinary fact if no
variations had ever occurred useful to each being's own welfarein the
same manner as so many variations have occurred useful to man. But if
variations useful to any organic being ever do occurassuredly individuals
thus characterised will have the best chance of being preserved in the
struggle for life; and from the strong principle of inheritancethese will
tend to produce offspring similarly characterised. This principle of
preservationor the survival of the fittestI have called natural
selection. It leads to the improvement of each creature in relation to its
organic and inorganic conditions of life; and consequentlyin most cases
to what must be regarded as an advance in organisation. Neverthelesslow
and simple forms will long endure if well fitted for their simple
conditions of life.

Natural selectionon the principle of qualities being inherited at
corresponding agescan modify the eggseedor young as easily as the
adult. Among many animals sexual selection will have given its aid to
ordinary selection by assuring to the most vigorous and best adapted males
the greatest number of offspring. Sexual selection will also give
characters useful to the males alone in their struggles or rivalry with
other males; and these characters will be transmitted to one sex or to both
sexesaccording to the form of inheritance which prevails.

Whether natural selection has really thus acted in adapting the various
forms of life to their several conditions and stationsmust be judged by
the general tenour and balance of evidence given in the following chapters.
But we have already seen how it entails extinction; and how largely
extinction has acted in the world's historygeology plainly declares.
Natural selectionalsoleads to divergence of character; for the more
organic beings diverge in structurehabits and constitutionby so much
the more can a large number be supported on the areaof which we see proof
by looking to the inhabitants of any small spotand to the productions
naturalised in foreign lands. Thereforeduring the modification of the
descendants of any one speciesand during the incessant struggle of all
species to increase in numbersthe more diversified the descendants
becomethe better will be their chance of success in the battle for life.
Thus the small differences distinguishing varieties of the same species
steadily tend to increasetill they equal the greater differences between
species of the same genusor even of distinct genera.

We have seen that it is the commonthe widely diffusedand widely ranging
speciesbelonging to the larger genera within each classwhich vary most;
and these tend to transmit to their modified offspring that superiority
which now makes them dominant in their own countries. Natural selection
as has just been remarkedleads to divergence of character and to much
extinction of the less improved and intermediate forms of life. On these
principlesthe nature of the affinitiesand the generally well defined
distinctions between the innumerable organic beings in each class
throughout the worldmay be explained. It is a truly wonderful fact--the
wonder of which we are apt to overlook from familiarity--that all animals
and all plants throughout all time and space should be related to each
other in groupssubordinate to groupsin the manner which we everywhere
behold--namelyvarieties of the same species most closely relatedspecies
of the same genus less closely and unequally relatedforming sections and

sub-generaspecies of distinct genera much less closely relatedand
genera related in different degreesforming sub-familiesfamilies
orderssub-classesand classes. The several subordinate groups in any
class cannot be ranked in a single filebut seem clustered round points
and these round other pointsand so on in almost endless cycles. If
species had been independently createdno explanation would have been
possible of this kind of classification; but it is explained through
inheritance and the complex action of natural selectionentailing
extinction and divergence of characteras we have seen illustrated in the

The affinities of all the beings of the same class have sometimes been
represented by a great tree. I believe this simile largely speaks the
truth. The green and budding twigs may represent existing species; and
those produced during former years may represent the long succession of
extinct species. At each period of growth all the growing twigs have tried
to branch out on all sidesand to overtop and kill the surrounding twigs
and branchesin the same manner as species and groups of species have at
all times overmastered other species in the great battle for life. The
limbs divided into great branchesand these into lesser and lesser
brancheswere themselves oncewhen the tree was youngbudding twigs; and
this connexion of the former and present buds by ramifying branches may
well represent the classification of all extinct and living species in
groups subordinate to groups. Of the many twigs which flourished when the
tree was a mere bushonly two or threenow grown into great branchesyet
survive and bear the other branches; so with the species which lived during
long-past geological periodsvery few have left living and modified
descendants. From the first growth of the treemany a limb and branch has
decayed and dropped off; and these fallen branches of various sizes may
represent those whole ordersfamiliesand genera which have now no living
representativesand which are known to us only in a fossil state. As we
here and there see a thinstraggling branch springing from a fork low down
in a treeand which by some chance has been favoured and is still alive on
its summitso we occasionally see an animal like the Ornithorhynchus or
Lepidosirenwhich in some small degree connects by its affinities two
large branches of lifeand which has apparently been saved from fatal
competition by having inhabited a protected station. As buds give rise by
growth to fresh budsand theseif vigorousbranch out and overtop on all
sides many a feebler branchso by generation I believe it has been with
the great Tree of Lifewhich fills with its dead and broken branches the
crust of the earthand covers the surface with its ever-branching and
beautiful ramifications.



Effects of changed conditions -- Use and disusecombined with natural
selection; organs of flight and of vision -- Acclimatisation -- Correlated
variation -- Compensation and economy of growth -- False correlations -Multiple
rudimentaryand 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.

I have hitherto sometimes spoken as if the variations--so common and
multiform with organic beings under domesticationand in a lesser degree
with those under nature--were due to chance. Thisof course is a wholly
incorrect expressionbut it serves to acknowledge plainly our ignorance of
the cause of each particular variation. Some authors believe it to be as
much the function of the reproductive system to produce individual
differencesor slight deviations of structureas to make the child like

its parents. But the fact of variations and monstrosities occurring much
more frequently under domestication than under natureand the greater
variability of species having wide ranges than of those with restricted
rangeslead to the conclusion that variability is generally related to the
conditions of life to which each species has been exposed during several
successive generations. In the first chapter I attempted to show that
changed conditions act in two waysdirectly on the whole organisation or
on certain parts aloneand indirectly through the reproductive system. In
all cases there are two factorsthe nature of the organismwhich is much
the most important of the twoand the nature of the conditions. The
direct action of changed conditions leads to definite or indefinite
results. In the latter case the organisation seems to become plasticand
we have much fluctuating variability. In the former case the nature of the
organism is such that it yields readilywhen subjected to certain
conditionsand allor nearly allthe individuals become modified in the
same way.

It is very difficult to decide how far changed conditionssuch as of
climatefoodetc.have acted in a definite manner. There is reason to
believe that in the course of time the effects have been greater than can
be proved by clear evidence. But we may safely conclude that the
innumerable complex co-adaptations of structurewhich we see throughout
nature between various organic beingscannot be attributed simply to such
action. In the following cases the conditions seem to have produced some
slight definite effect: E. Forbes asserts that shells at their southern
limitand when living in shallow waterare more brightly coloured than
those of the same species from further north or from a greater depth; but
this certainly does not always hold good. Mr. Gould believes that birds of
the same species are more brightly coloured under a clear atmospherethan
when living near the coast or on islands; and Wollaston is convinced that
residence near the sea affects the colours of insects. Moquin-Tandon gives
a list of plants whichwhen growing near the sea-shorehave their leaves
in some degree fleshythough not elsewhere fleshy. 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.

When a variation is of the slightest use to any beingwe cannot tell how
much to attribute to the accumulative action of natural selectionand how
much to the definite action of the conditions of life. Thusit is well
known to furriers that animals of the same species have thicker and better
fur the further north they live; but who can tell how much of this
difference may be due to the warmest-clad individuals having been favoured
and preserved during many generationsand 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.

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; andon the other handof dissimilar varieties being produced
under apparently the same external conditions. Againinnumerable
instances are known to every naturalistof species keeping trueor not
varying at allalthough living under the most opposite climates. Such
considerations as these incline me to lay less weight on the direct action
of the surrounding conditionsthan on a tendency to varydue to causes of
which we are quite ignorant.

In one sense the conditions of life may be saidnot only to cause
variabilityeither directly or indirectlybut likewise to include natural
selectionfor the conditions determine whether this or that variety shall
survive. But when man is the selecting agentwe clearly see that the two
elements of change are distinct; variability is in some manner excitedbut
it is the will of man which accumulates the variations in certain
direction; and it is this latter agency which answers to the survival of
the fittest under nature.


>From the facts alluded to in the first chapterI think there can be no
doubt that use in our domestic animals has strengthened and enlarged
certain partsand disuse diminished them; and that such modifications are
inherited. Under free nature we have no standard of comparison by which to
judge of the effects of long-continued use or disusefor we know not the
parent-forms; but many animals possess structures which can be best
explained by the effects of disuse. As Professor Owen has remarkedthere
is no greater anomaly in nature than a bird that cannot fly; yet there are
several in this state. The logger-headed duck of South America can only
flap along the surface of the waterand has its wings in nearly the same
condition as the domestic Aylesbury duck: it is a remarkable fact that the
young birdsaccording to Mr. Cunninghamcan flywhile the adults have
lost this power. As the larger ground-feeding birds seldom take flight
except to escape dangerit is probable that the nearly wingless condition
of several birdsnow inhabiting or which lately inhabited several oceanic
islandstenanted by no beasts of preyhas been caused by disuse. The
ostrich indeed inhabits continentsand is exposed to danger from which it
cannot escape by flightbut it can defend itselfby kicking its enemies
as efficiently as many quadrupeds. We may believe that the progenitor of
the ostrich genus had habits like those of the bustardand thatas the
size and weight of its body were increased during successive generations
its legs were used more and its wings lessuntil they became incapable of

Kirby has remarked (and I have observed the same fact) that the anterior
tarsior feetof many male dung-feeding beetles are often broken off; he
examined seventeen specimens in his own collectionand not one had even a
relic left. In the Onites apelles the tarsi are so habitually lost that
the insect has been described as not having them. In some other genera
they are presentbut in a rudimentary condition. In the Ateuchus or
sacred beetle of the Egyptiansthey are totally deficient. The evidence
that accidental mutilations can be inherited is at present not decisive;
but the remarkable cases observed by Brown-Sequard in guinea-pigsof the
inherited effects of operationsshould make us cautious in denying this
tendency. Henceit will perhaps be safest to look at the entire absence
of the anterior tarsi in Ateuchusand their rudimentary condition in some
other generanot as cases of inherited mutilationsbut as due to the
effects of long-continued disuse; for as many dung-feeding beetles are
generally found with their tarsi lostthis must happen early in life;
therefore the tarsi cannot be of much importance or be much used by these

In some cases we might easily put down to disuse modifications of structure
which are whollyor mainly due to natural selection. Mr. Wollaston has
discovered the remarkable fact that 200 beetlesout of the 550 species
(but more are now known) inhabiting Madeiraare so far deficient in wings
that they cannot fly; and thatof the twenty-nine endemic generano less
than twenty-three have all their species in this condition! Several facts
namelythat beetles in many parts of the world are very frequently blown
to sea and perish; that the beetles in Madeiraas observed by Mr.
Wollastonlie much concealeduntil the wind lulls and the sun shines;
that the proportion of wingless beetles is larger on the exposed Desertas
than in Madeira itself; and especially the extraordinary factso strongly
insisted on by Mr. Wollastonthat certain large groups of beetles
elsewhere excessively numerouswhich absolutely require the use of their
wingsare here almost entirely absent. These several considerations make
me believe that the wingless condition of so many Madeira beetles is mainly
due to the action of natural selectioncombined probably with disuse. For
during many successive generations each individual beetle which flew least
either from its wings having been ever so little less perfectly developed

or from indolent habitwill have had the best chance of surviving from not
being blown out to sea; andon the other handthose beetles which most
readily took to flight would oftenest have been blown to seaand thus

The insects in Madeira which are not ground-feedersand whichas certain
flower-feeding coleoptera and lepidopteramust habitually use their wings
to gain their subsistencehaveas Mr. Wollaston suspectstheir wings not
at all reducedbut even enlarged. This is quite compatible with the
action of natural selection. For when a new insect first arrived on the
islandthe tendency of natural selection to enlarge or to reduce the
wingswould depend on whether a greater number of individuals were saved
by successfully battling with the windsor by giving up the attempt and
rarely or never flying. As with mariners shipwrecked near a coastit
would have been better for the good swimmers if they had been able to swim
still furtherwhereas 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.

The eyes of moles and of some burrowing rodents are rudimentary in size
and in some cases are quite covered by skin and fur. This state of the
eyes is probably due to gradual reduction from disusebut aided perhaps by
natural selection. In South Americaa burrowing rodentthe tuco-tucoor
Ctenomysis even more subterranean in its habits than the mole; and I was
assured by a Spaniardwho had often caught themthat they were frequently
blind. One which I kept alive was certainly in this conditionthe cause
as appeared on dissectionhaving been inflammation of the nictitating
membrane. As frequent inflammation of the eyes must be injurious to any
animaland as eyes are certainly not necessary to animals having
subterranean habitsa reduction in their sizewith the adhesion of the
eyelids and growth of fur over themmight in such case be an advantage;
and if sonatural selection would aid the effects of disuse.

It is well known that several animalsbelonging to the most different
classeswhich inhabit the caves of Carniola and Kentuckyare blind. In
some of the crabs the foot-stalk for the eye remainsthough the eye is
gone; the stand for the telescope is therethough the telescope with its
glasses has been lost. As it is difficult to imagine that eyesthough
uselesscould be in any way injurious to animals living in darknesstheir
loss may be attributed to disuse. In one of the blind animalsnamelythe
cave-rat (Neotoma)two of which were captured by Professor Silliman at
above half a mile distance from the mouth of the caveand therefore not in
the profoundest depthsthe eyes were lustrous and of large size; and these
animalsas I am informed by Professor Sillimanafter having been exposed
for about a month to a graduated lightacquired a dim perception of

It is difficult to imagine conditions of life more similar than deep
limestone caverns under a nearly similar climate; so thatin accordance
with the old view of the blind animals having been separately created for
the American and European cavernsvery close similarity in their
organisation and affinities might have been expected. This is certainly
not the case if we look at the two whole faunas; with respect to the
insects aloneSchiodte has remarked: "We are accordingly prevented from
considering the entire phenomenon in any other light than something purely
localand the similarity which is exhibited in a few forms between the
Mammoth Cave (in Kentucky) and the caves in Carniolaotherwise than as a
very plain expression of that analogy which subsists generally between the
fauna of Europe and of North America." On my view we must suppose that
American animalshaving in most cases ordinary powers of visionslowly
migrated by successive generations from the outer world into the deeper and
deeper recesses of the Kentucky cavesas did European animals into the
caves of Europe. We have some evidence of this gradation of habit; foras
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 tractsand whichas they
extended themselves into darknesshave been accommodated to surrounding
circumstances. Animals not far remote from ordinary formsprepare the
transition from light to darkness. Next follow those that are constructed
for twilight; andlast of allthose destined for total darknessand
whose formation is quite peculiar." These remarks of Schiodte's it should
be understoodapply not to the samebut to distinct species. By the time
that an animal had reachedafter numberless generationsthe deepest
recessesdisuse will on this view have more or less perfectly obliterated
its eyesand natural selection will often have effected other changes
such as an increase in the length of the antennae or palpias a
compensation for blindness. Notwithstanding such modificationswe might
expect still to see in the cave-animals of Americaaffinities to the other
inhabitants of that continentand in those of Europe to the inhabitants of
the European continent. And this is the case with some of the American
cave-animalsas I hear from Professor Dana; and some of the European
cave-insects are very closely allied to those of the surrounding country.
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. That several of the
inhabitants of the caves of the Old and New Worlds should be closely
relatedwe might expect from the well-known relationship of most of their
other productions. As a blind species of Bathyscia is found in abundance
on shady rocks far from cavesthe loss of vision in the cave species of
this one genus has probably had no relation to its dark habitation; for it
is natural that an insect already deprived of vision should readily become
adapted to dark caverns. Another blind genus (Anophthalmus) offers this
remarkable peculiaritythat the speciesas Mr. Murray observeshave not
as yet been found anywhere except in caves; yet those which inhabit the
several caves of Europe and America are distinct; but it is possible that
the progenitors of these several specieswhile they were furnished with
eyesmay formerly have ranged over both continentsand then have become
extinctexcepting in their present secluded abodes. Far from feeling
surprise that some of the cave-animals should be very anomalousas Agassiz
has remarked in regard to the blind fishthe Amblyopsisand as is the
case with the blind Proteuswith reference to the reptiles of EuropeI am
only surprised that more wrecks of ancient life have not been preserved
owing to the less severe competition to which the scanty inhabitants of
these dark abodes will have been exposed.


Habit is hereditary with plantsas in the period of floweringin the time
of sleepin the amount of rain requisite for seeds to germinateetc.and
this leads me to say a few words on acclimatisation. As it is extremely
common for distinct species belonging to the same genus to inhabit hot and
cold countriesif it be true that all the species of the same genus are
descended from a single parent-formacclimatisation must be readily
effected during a long course of descent. 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 climateor
conversely. So againmany succulent plants cannot endure a damp climate.
But the degree of adaptation of species to the climates under which they
live is often overrated. We may infer this from our frequent inability to
predict whether or not an imported plant will endure our climateand from
the number of plants and animals brought from different countries which are
here perfectly healthy. 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 asor more thanby adaptation to particular
climates. But whether or not this adaptation is in most cases very close
we have evidence with some few plantsof their becomingto a certain
extentnaturally habituated to different temperatures; that isthey
become acclimatised: thus the pines and rhododendronsraised from seed
collected by Dr. Hooker from the same species growing at different heights

on the Himalayaswere found to possess in this country different
constitutional powers of resisting cold. Mr. Thwaites informs me that he
has observed similar facts in Ceylon; analogous observations have been made
by Mr. H.C. Watson on European species of plants brought from the Azores to
England; and I could give other cases. In regard to animalsseveral
authentic instances could be adduced of species having largely extended
within historical timestheir range from warmer to colder latitudesand
conversely; but we do not positively know that these animals were strictly
adapted to their native climatethough in all ordinary cases we assume
such to be the case; nor do we know that they have subsequently become
specially acclimatised to their new homesso as to be better fitted for
them than they were at first.

As we may infer that our domestic animals were originally chosen by
uncivilised man because they were usefuland because they bred readily
under confinementand not because they were subsequently found capable of
far-extended transportationthe common and extraordinary capacity in our
domestic animals of not only withstanding the most different climatesbut
of being perfectly fertile (a far severer test) under themmay 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. We must
nothoweverpush the foregoing argument too faron account of the
probable origin of some of our domestic animals from several wild stocks:
the bloodfor instanceof a tropical and arctic wolf may perhaps be
mingled in our domestic breeds. The rat and mouse cannot be considered as
domestic animalsbut they have been transported by man to many parts of
the worldand now have a far wider range than any other rodent; for they
live under the cold climate of Faroe in the north and of the Falklands in
the southand on many an island in the torrid zones. Hence adaptation to
any special climate may be looked at as a quality readily grafted on an
innate wide flexibility of constitutioncommon to most animals. On this
viewthe capacity of enduring the most different climates by man himself
and by his domestic animalsand the fact of the extinct elephant and
rhinoceros having formerly endured a glacial climatewhereas the living
species are now all tropical or sub-tropical in their habitsought not to
be looked at as anomaliesbut as examples of a very common flexibility of
constitutionbroughtunder peculiar circumstancesinto action.

How much of the acclimatisation of species to any peculiar climate is due
to mere habitand how much to the natural selection of varieties having
different innate constitutionsand how much to both means combinedis an
obscure question. That habit or custom has some influenceI must believe
both from analogy and from the incessant advice given in agricultural
workseven in the ancient Encyclopaedias of Chinato be very cautious in
transporting animals from one district to another. 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 districtsthe result
mustI thinkbe due to habit. On the other handnatural selection would
inevitably tend to preserve those individuals which were born with
constitutions best adapted to any country which they inhabited. In
treatises on many kinds of cultivated plantscertain varieties are said to
withstand certain climates better than others; this is strikingly shown in
works on fruit-trees published in the United Statesin which certain
varieties are habitually recommended for the northern and others for the
southern states; and as most of these varieties are of recent originthey
cannot owe their constitutional differences to habit. The case of the
Jerusalem artichokewhich is never propagated in England by seedand of
whichconsequentlynew varieties have not been producedhas even been
advancedas proving that acclimatisation cannot be effectedfor it is now
as tender as ever it was! The casealsoof the kidney-bean has been
often cited for a similar purposeand with much greater weight; but until
some one will sowduring a score of generationshis kidney-beans so early
that a very large proportion are destroyed by frostand then collect seed
from the few survivorswith care to prevent accidental crossesand then

again get seed from these seedlingswith the same precautionsthe
experiment cannot be said to have been even tried. Nor let it be supposed
that differences in the constitution of seedling kidney-beans never appear
for an account has been published how much more hardy some seedlings are
than others; and of this fact I have myself observed striking instances.

On the wholewe may conclude that habitor use and disusehavein some
casesplayed a considerable part in the modification of the constitution
and structure; but that the effects have often been largely combined with
and sometimes overmastered bythe natural selection of innate variations.


I mean by this expression that the whole organisation is so tied together
during its growth and developmentthat when slight variations in any one
part occur and are accumulated through natural selectionother parts
become modified. This is a very important subjectmost imperfectly
understoodand no doubt wholly different classes of facts may be here
easily confounded together. We shall presently see that simple inheritance
often gives the false appearance of correlation. One of the most obvious
real cases isthat variations of structure arising in the young or larvae
naturally tend to affect the structure of the mature animal. The several
parts which are homologousand whichat an early embryonic periodare
identical in structureand which are necessarily exposed to similar
conditionsseem 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; in the
front and hind legsand even in the jaws and limbsvarying togetherfor
the lower jaw is believed by some anatomists to be homologous with the
limbs. These tendenciesI do not doubtmay be mastered more or less
completely by natural selection: thus a family of stags once existed with
an antler only on one side; and if this had been of any great use to the
breedit might probably have been rendered permanent by natural selection.

Homologous partsas has been remarked by some authorstend to cohere;
this is often seen in monstrous plants: and nothing is more common than
the union of homologous parts in normal structuresas in the union of the
petals into a tube. Hard parts seem to affect the form of adjoining soft
parts; 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. Others believe that the shape of the pelvis in the human mother
influences by pressure the shape of the head of the child. In snakes
according to Schlegelthe shape of the body and the manner of swallowing
determine the position and form of several of the most important viscera.

The nature of the bond is frequently quite obscure. M. Is. Geoffroy St.
Hilaire has forcibly remarked that certain malconformations frequentlyand
that others rarelycoexist without our being able to assign any reason.
What can be more singular than the relation in cats between complete
whiteness and blue eyes with deafnessor between the tortoise-shell colour
and the female sex; or in pigeonsbetween their feathered feet and skin
betwixt the outer toesor between the presence of more or less down on the
young pigeon when first hatchedwith the future colour of its plumage; or
againthe relation between the hair and the teeth in the naked Turkish
dogthough here no doubt homology comes into play? With respect to this
latter case of correlationI think it can hardly be accidental that the
two orders of mammals which are most abnormal in their dermal covering
viz.Cetacea (whales) and Edentata (armadilloesscaly ant-eatersetc.)
are likewise on the whole the most abnormal in their teethbut there are
so many exceptions to this ruleas Mr. Mivart has remarkedthat it has
little value.

I know of no case better adapted to show the importance of the laws of
correlation and variationindependently of utilityand therefore of
natural selectionthan that of the difference between the outer and inner

flowers in some Compositous and Umbelliferous plants. Everyone is familiar
with the difference between the ray and central florets offor instance
the daisyand this difference is often accompanied with the partial or
complete abortion of the reproductive organs. But in some of these plants
the seeds also differ in shape and sculpture. These differences have
sometimes been attributed to the pressure of the involucra on the florets
or to their mutual pressureand the shape of the seeds in the ray-florets
of some Compositae countenances this idea; but with the Umbelliferae it is
by no meansas Dr. Hooker informs methe species with the densest heads
which most frequently differ in their inner and outer flowers. It might
have been thought that the development of the ray-petalsby drawing
nourishment from the reproductive organs causes their abortion; but this
can hardly be the sole casefor in some Compositae the seeds of the outer
and inner florets differwithout any difference in the corolla. Possibly
these several differences may be connected with the different flow of
nutriment towards the central and external flowers. We knowat least
that with irregular flowers those nearest to the axis are most subject to
peloriathat is to become abnormally symmetrical. I may addas an
instance of this factand as a striking case of correlationthat in many
pelargoniums the two upper petals in the central flower of the truss often
lose their patches of darker colour; and when this occursthe adherent
nectary is quite abortedthe central flower thus becoming peloric or
regular. When the colour is absent from only one of the two upper petals
the nectary is not quite aborted but is much shortened.

With respect to the development of the corollaSprengel's idea that the
ray-florets serve to attract insectswhose agency is highly advantageous
or necessary for the fertilisation of these plantsis highly probable; and
if sonatural selection may have come into play. But with respect to the
seedsit seems impossible that their differences in shapewhich are not
always correlated with any difference in the corollacan be in any way
beneficial; 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. Hence
modifications of structureviewed by systematists as of high valuemay be
wholly due to the laws of variation and correlationwithout beingas far
as we can judgeof the slightest service to the species.

We may often falsely attribute to correlated variation structures which are
common to whole groups of speciesand which in truth are simply due to
inheritance; for an ancient progenitor may have acquired through natural
selection some one modification in structureandafter thousands of
generationssome other and independent modification; and these two
modificationshaving been transmitted to a whole group of descendants with
diverse habitswould naturally be thought to be in some necessary manner
correlated. Some other correlations are apparently due to the manner in
which natural selection can alone act. For instanceAlph. De Candolle has
remarked that winged seeds are never found in fruits which do not open; I
should explain this rule by the impossibility of seeds gradually becoming
winged through natural selectionunless the capsules were open; for in
this case alone could the seedswhich were a little better adapted to be
wafted by the windgain an advantage over others less well fitted for wide


The elder Geoffroy and Goethe propoundedat about the same timetheir law
of compensation or balancement of growth; oras Goethe expressed itin
order to spend on one side, nature is forced to economise on the other
side.I think this holds true to a certain extent with our domestic
productions: if nourishment flows to one part or organ in excessit
rarely flowsat least in excessto another part; thus it is difficult to
get a cow to give much milk and to fatten readily. The same varieties of

the cabbage do not yield abundant and nutritious foliage and a copious
supply of oil-bearing seeds. When the seeds in our fruits become
atrophiedthe fruit itself gains largely in size and quality. In our
poultrya large tuft of feathers on the head is generally accompanied by a
diminished comband a large beard by diminished wattles. With species in
a state of nature it can hardly be maintained that the law is of universal
application; but many good observersmore especially botanistsbelieve in
its truth. I will nothoweverhere give any instancesfor I see hardly
any way of distinguishing between the effectson the one handof a part
being largely developed through natural selection and another and adjoining
part being reduced by the same process or by disuseandon the other
handthe actual withdrawal of nutriment from one part owing to the excess
of growth in another and adjoining part.

I suspectalsothat some of the cases of compensation which have been
advancedand likewise some other factsmay be merged under a more general
principlenamelythat natural selection is continually trying to
economise in every part of the organisation. If under changed conditions
of life a structurebefore usefulbecomes less usefulits diminution
will be favouredfor it will profit the individual not to have its
nutriment wasted in building up a useless structure. I can thus only
understand a fact with which I was much struck when examining cirripedes
and of which many other instances could be given: namelythat when a
cirripede is parasitic within another cirripede and is thus protectedit
loses more or less completely its own shell or carapace. This is the case
with the male Iblaand 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
and furnished with great nerves and muscles; but in the parasitic and
protected Proteolepasthe whole anterior part of the head is reduced to
the merest rudiment attached to the bases of the prehensile antennae. Now
the saving of a large and complex structurewhen rendered superfluous
would be a decided advantage to each successive individual of the species;
for in the struggle for life to which every animal is exposedeach would
have a better chance of supporting itselfby less nutriment being wasted.

Thusas I believenatural selection will tend in the long run to reduce
any part of the organisationas soon as it becomesthrough changed
habitssuperfluouswithout by any means causing some other part to be
largely developed in a corresponding degree. And converselythat natural
selection may perfectly well succeed in largely developing an organ without
requiring as a necessary compensation the reduction of some adjoining part.


It seems to be a ruleas remarked by Is. Geoffroy St. Hilaireboth with
varieties and speciesthat when any part or organ is repeated many times
in the same individual (as the vertebrae in snakesand the stamens in
polyandrous flowers) the number is variable; whereas the number of the same
part or organwhen it occurs in lesser numbersis constant. The same
author as well as some botanistshave further remarked that multiple parts
are extremely liable to vary in structure. As "vegetative repetition to
use Professor Owen's expression, is a sign of low organisation; the
foregoing statements accord with the common opinion of naturalists, that
beings which stand low in the scale of nature are more variable than those
which are higher. I presume that lowness here means that the several parts
of the organisation have been but little specialised for particular
functions; and as long as the same part has to perform diversified work, we
can perhaps see why it should remain variable, that is, why natural
selection should not have preserved or rejected each little deviation of
form so carefully as when the part has to serve for some one special
purpose. In the same way that a knife which has to cut all sorts of things
may be of almost any shape; whilst a tool for some particular purpose must
be of some particular shape. Natural selection, it should never be

forgotten, can act solely through and for the advantage of each being.

Rudimentary parts, as is generally admitted, are apt to be highly variable.
We shall have to recur to this subject; and I will here only add that their
variability seems to result from their uselessness, and consequently from
natural selection having had no power to check deviations in their


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

As with birds the individuals of the same species, inhabiting the same
country, vary extremely little, I have particularly attended to them; and
the rule certainly seems to hold good in this class. I cannot make out
that it applies to plants, and this would have seriously shaken my belief
in its truth, had not the great variability in plants made it particularly
difficult to compare their relative degrees of variability.

When we see any part or organ developed in a remarkable degree or manner in
a species, the fair presumption is that it is of high importance to that
species: nevertheless it is in this case eminently liable to variation.
Why should this be so? On the view that each species has been
independently created, with all its parts as we now see them, I can see no

explanation. But on the view that groups of species are descended from
some other species, and have been modified through natural selection, I
think we can obtain some light. First let me make some preliminary
remarks. If, in our domestic animals, any part or the whole animal be
neglected, and no selection be applied, that part (for instance, 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. In rudimentary
organs, and in those which have been but little specialised for any
particular purpose, and perhaps in polymorphic groups, we see a nearly
parallel case; for in such cases natural selection either has not or cannot
come into full play, and thus the organisation is left in a fluctuating
condition. But what here more particularly concerns us is, that those
points in our domestic animals, which at the present time are undergoing
rapid change by continued selection, are also eminently liable to
variation. Look at the individuals of the same breed of the pigeon; and
see what a prodigious amount of difference there is in the beak of
tumblers, in the beak and wattle of carriers, in the carriage and tail of
fantails, etc., these being the points now mainly attended to by English
fanciers. Even in the same sub-breed, as in that of the short-faced
tumbler, it is notoriously difficult to breed nearly perfect birds, many
departing widely from the standard. There may truly be said to be a
constant struggle going on between, on the one hand, the tendency to
reversion to a less perfect state, as well as an innate tendency to new
variations, and, on the other hand, the power of steady selection to keep
the breed true. In the long run selection gains the day, and we do not
expect to fail so completely as to breed a bird as coarse as a common
tumbler pigeon from a good short-faced strain. But as long as selection is
rapidly going on, much variability in the parts undergoing modification may
always be expected.

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


The principle discussed under the last heading may be applied to our
present subject. It is notorious that specific characters are more

variable than generic. 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,
the colour would be only a specific character, and no one would be
surprised at one of the blue species varying into red, or conversely; but
if all the species had blue flowers, the colour would become a generic
character, and its variation would be a more unusual circumstance. I have
chosen this example because the explanation which most naturalists would
advance is not here applicable, namely, that specific characters are more
variable than generic, because they are taken from parts of less
physiological importance than those commonly used for classing genera. I
believe this explanation is partly, yet only indirectly, true; I shall,
however, have to return to this point in the chapter on Classification. It
would be almost superfluous to adduce evidence in support of the statement,
that ordinary specific characters are more variable than generic; but with
respect to important characters, I have repeatedly noticed in works on
natural history, that when an author remarks with surprise that some
important organ or part, which is generally very constant throughout a
large group of species, DIFFERS considerably in closely-allied species, it
is often VARIABLE in the individuals of the same species. And this fact
shows that a character, which is generally of generic value, when it sinks
in value and becomes only of specific value, often becomes variable, though
its physiological importance may remain the same. Something of the same
kind applies to monstrosities: at least Is. Geoffroy St. Hilaire
apparently entertains no doubt, that the more an organ normally differs in
the different species of the same group, the more subject it is to
anomalies in the individuals.

On the ordinary view of each species having been independently created, why
should that part of the structure, which differs from the same part in
other independently created species of the same genus, be more variable
than those parts which are closely alike in the several species? I do not
see that any explanation can be given. But on the view that species are
only strongly marked and fixed varieties, 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, 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, and in which they differ from
allied genera, are called generic characters; and these characters may be
attributed to inheritance from a common progenitor, for it can rarely have
happened that natural selection will have modified several distinct
species, fitted to more or less widely different habits, in exactly the
same manner: and as these so-called generic characters have been inherited
from before the period when the several species first branched off from
their common progenitor, and subsequently have not varied or come to differ
in any degree, or only in a slight degree, it is not probable that they
should vary at the present day. On the other hand, the points in which
species differ from other species of the same genus are called specific
characters; and as these specific characters have varied and come to differ
since the period when the species branched off from a common progenitor, 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.


I think it will be admitted by naturalists, without my entering on details,
that secondary sexual characters are highly variable. It will also be
admitted that species of the same group differ from each other more widely
in their secondary sexual characters, than in other parts of their
organisation; compare, for instance, the amount of difference between the
males of gallinaceous birds, in which secondary sexual characters are
strongly displayed, with the amount of difference between the females. The
cause of the original variability of these characters is not manifest; but
we can see why they should not have been rendered as constant and uniform

as others, for they are accumulated by sexual selection, which is less
rigid in its action than ordinary selection, as it does not entail death,
but only gives fewer offspring to the less favoured males. Whatever the
cause may be of the variability of secondary sexual characters, as they are
highly variable, sexual selection will have had a wide scope for action,
and may thus have succeeded in giving to the species of the same group a
greater amount of difference in these than in other respects.

It is a remarkable fact, 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. Of this fact I will give in illustration the first two instances
which happen to stand on my list; and as the differences in these cases are
of a very unusual nature, the relation can hardly be accidental. The same
number of joints in the tarsi is a character common to very large groups of
beetles, but in the Engidae, as Westwood has remarked, the number varies
greatly and the number likewise differs in the two sexes of the same
species. Again in the fossorial hymenoptera, the neuration of the wings is
a character of the highest importance, because common to large groups; but
in certain genera the neuration differs in the different species, and
likewise in the two sexes of the same species. Sir J. Lubbock has recently
remarked, that several minute crustaceans offer excellent illustrations of
this law. In Pontellafor instancethe 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." 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 progenitoras
have the two sexes of any one species. Consequentlywhatever part of the
structure of the common progenitoror of its early descendantsbecame
variable; variations of this part wouldit is highly probablebe taken
advantage of by natural and sexual selectionin order to fit the several
places in the economy of natureand likewise to fit the two sexes of the
same species to each otheror to fit the males to struggle with other
males for the possession of the females.

FinallythenI conclude that the greater variability of specific
charactersor those which distinguish species from speciesthan of
generic charactersor those which are possessed by all the species; that
the frequent extreme variability of any part which is developed in a
species in an extraordinary manner in comparison with the same part in its
congeners; and the slight degree of variability in a parthowever
extraordinarily it may be developedif it be common to a whole group of
species; that the great variability of secondary sexual characters and
their great difference in closely allied species; that secondary sexual and
ordinary specific differences are generally displayed in the same parts of
the organisationare all principles closely connected together. All being
mainly due to the species of the same group being the descendants of a
common progenitorfrom whom they have inherited much in commonto 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 variedto
natural selection having more or less completelyaccording to the lapse of
timeovermastered the tendency to reversion and to further variabilityto
sexual selection being less rigid than ordinary selectionand to
variations in the same parts having been accumulated by natural and sexual
selectionand thus having been adapted for secondary sexualand for
ordinary purposes.


These propositions will be most readily understood by looking to our
domestic races. The most distinct breeds of the pigeonin countries
widely apartpresent sub-varieties with reversed feathers on the headand

with feathers on the feetcharacters not possessed by the aboriginal
rock-pigeon; these then are analogous variations in two or more distinct
races. 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 racethe fantail. 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 variationwhen acted on by similar unknown influences. In the
vegetable kingdom we have a case of analogous variationin the enlarged
stemsor as commonly called rootsof the Swedish turnip and ruta-baga
plants which several botanists rank as varieties produced by cultivation
from a common parent: if this be not sothe case will then be one of
analogous variation in two so-called distinct species; and to these a third
may be addednamelythe common turnip. According to the ordinary view of
each species having been independently createdwe should have to attribute
this similarity in the enlarged stems of these three plantsnot to the
vera causa of community of descentand a consequent tendency to vary in a
like mannerbut to three separate yet closely related acts of creation.
Many similar cases of analogous variation have been observed by Naudin in
the great gourd familyand by various authors in our cereals. Similar
cases occurring with insects under natural conditions have lately been
discussed with much ability by Mr. Walshwho has grouped them under his
law of equable variability.

With pigeonshoweverwe have another casenamelythe occasional
appearance in all the breedsof slaty-blue birds with two black bars on
the wingswhite loinsa bar at the end of the tailwith the outer
feathers externally edged near their bases with white. As all these marks
are characteristic of the parent rock-pigeonI presume that no one will
doubt that this is a case of reversionand not of a new yet analogous
variation appearing in the several breeds. We mayI thinkconfidently
come to this conclusionbecauseas we have seenthese coloured marks are
eminently liable to appear in the crossed offspring of two distinct and
differently coloured breeds; and in this case there is nothing in the
external conditions of life to cause the reappearance of the slaty-blue
with the several marksbeyond the influence of the mere act of crossing on
the laws of inheritance.

No doubt it is a very surprising fact that characters should reappear after
having been lost for manyprobably for hundreds of generations. But when
a breed has been crossed only once by some other breedthe offspring
occasionally show for many generations a tendency to revert in character to
the foreign breed--some sayfor a dozen or even a score of generations.
After twelve generationsthe proportion of bloodto use a common
expressionfrom one ancestoris only 1 in 2048; and yetas we seeit is
generally believed that a tendency to reversion is retained by this remnant
of foreign blood. In a breed which has not been crossedbut in which BOTH
parents have lost some character which their progenitor possessedthe
tendencywhether strong or weakto reproduce the lost character mightas
was formerly remarkedfor all that we can see to the contrarybe
transmitted for almost any number of generations. When a character which
has been lost in a breedreappears after a great number of generations
the most probable hypothesis isnot that one individual suddenly takes
after an ancestor removed by some hundred generationsbut that in each
successive generation the character in question has been lying latentand
at lastunder unknown favourable conditionsis developed. With the
barb-pigeonfor instancewhich very rarely produces a blue birdit is
probable that there is a latent tendency in each generation to produce blue
plumage. The abstract improbability of such a tendency being transmitted
through a vast number of generationsis not greater than that of quite
useless or rudimentary organs being similarly transmitted. A mere tendency
to produce a rudiment is indeed sometimes thus inherited.

As all the species of the same genus are supposed to be descended from a

common progenitorit might be expected that they would occasionally vary
in an analogous manner; so that the varieties of two or more species would
resemble each otheror that a variety of one species would resemble in
certain characters another and distinct speciesthis other species being
according to our viewonly a well-marked and permanent variety. But
characters exclusively due to analogous variation would probably be of an
unimportant naturefor the preservation of all functionally important
characters will have been determined through natural selectionin
accordance with the different habits of the species. It might further be
expected that the species of the same genus would occasionally exhibit
reversions to long-lost characters. Ashoweverwe do not know the common
ancestor of any natural groupwe cannot distinguish between reversionary
and analogous characters. Iffor instancewe did not know that the
parent rock-pigeon was not feather-footed or turn-crownedwe could not
have toldwhether such characters in our domestic breeds were reversions
or only analogous variations; but we might have inferred that the blue
colour was a case of reversion from the number of the markingswhich are
correlated with this tintand which would not probably have all appeared
together from simple variation. More especially we might have inferred
this from the blue colour and the several marks so often appearing when
differently coloured breeds are crossed. Hencealthough under nature it
must generally be left doubtfulwhat cases are reversions to formerly
existing charactersand what are new but analogous variationsyet we
oughton our theorysometimes to find the varying offspring of a species
assuming characters which are already present in other members of the same
group. And this undoubtedly is the case.

The difficulty in distinguishing variable species is largely due to the
varieties mockingas it wereother species of the same genus. A
considerable cataloguealsocould be given of forms intermediate between
two other formswhich themselves can only doubtfully be ranked as species;
and this showsunless all these closely allied forms be considered as
independently created speciesthat they have in varying assumed some of
the characters of the others. But the best evidence of analogous
variations is afforded by parts or organs which are generally constant in
characterbut which occasionally vary so as to resemblein some degree
the same part or organ in an allied species. I have collected a long list
of such cases; but hereas beforeI lie under the great disadvantage of
not being able to give them. I can only repeat that such cases certainly
occurand seem to me very remarkable.

I willhowevergive one curious and complex casenot indeed as affecting
any important characterbut from occurring in several species of the same
genuspartly under domestication and partly under nature. It is a case
almost certainly of reversion. The ass sometimes has very distinct
transverse bars on its legslike those on the legs of a zebra. It has
been asserted that these are plainest in the foaland from inquiries which
I have madeI believe this to be true. The stripe on the shoulder is
sometimes doubleand is very variable in length and outline. A white ass
but NOT an albinohas been described without either spinal or shoulder
stripe; and these stripes are sometimes very obscureor actually quite
lostin dark-coloured asses. The koulan of Pallas is said to have been
seen with a double shoulder-stripe. Mr. Blyth has seen a specimen of the
hemionus with a distinct shoulder-stripethough it properly has none; and
I have been informed by Colonel Poole that foals of this species are
generally striped on the legs and faintly on the shoulder. The quagga
though so plainly barred like a zebra over the bodyis without bars on the
legs; but Dr. Gray has figured one specimen with very distinct zebra-like
bars on the hocks.

With respect to the horseI have collected cases in England of the spinal
stripe in horses of the most distinct breedsand of ALL colours;
transverse bars on the legs are not rare in dunsmouse-dunsand in one
instance in a chestnut; a faint shoulder-stripe may sometimes be seen in

dunsand I have seen a trace in a bay horse. 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. I have myself seen a dun
Devonshire ponyand a small dun Welsh pony has been carefully described to
meboth with THREE parallel stripes on each shoulder.

In the northwest part of India the Kattywar breed of horses is so generally
stripedthatas I hear from Colonel Poolewho examined this breed for
the Indian Governmenta horse without stripes is not considered as purely
bred. The spine is always striped; the legs are generally barred; and the
shoulder-stripewhich is sometimes double and sometimes trebleis common;
the side of the facemoreoveris sometimes striped. The stripes are
often plainest in the foal; and sometimes quite disappear in old horses.
Colonel Poole has seen both gray and bay Kattywar horses striped when first
foaled. I have also reason to suspectfrom information given me by Mr.

W.W. Edwardsthat with the English race-horse the spinal stripe is much
commoner in the foal than in the full-grown animal. 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. This foalwhen a week oldwas marked
on its hinder quarters and on its forehead with numerous very narrowdark
zebra-like barsand its legs were feebly striped. All the stripes soon
disappeared completely. Without here entering on further details I may
state that I have collected cases of leg and shoulder stripes in horses of
very different breeds in various countries from Britain to Eastern China;
and from Norway in the north to the Malay Archipelago in the south. In all
parts of the world these stripes occur far oftenest in duns and mouse-duns;
by the term dun a large range of colour is includedfrom one between brown
and black to a close approach to cream colour.
I am aware that Colonel Hamilton Smithwho has written on this subject
believes that the several breeds of the horse are descended from several
aboriginal speciesone of whichthe dunwas striped; and that the
above-described appearances are all due to ancient crosses with the dun
stock. But this view may be safely rejectedfor it is highly improbable
that the heavy Belgian cart-horseWelsh poniesNorwegian cobsthe lanky
Kattywar raceetc.inhabiting the most distant parts of the worldshould
have all have been crossed with one supposed aboriginal stock.

Now let us turn to the effects of crossing the several species of the horse
genus. Rollin asserts that the common mule from the ass and horse is
particularly apt to have bars on its legs; according to Mr. Gossein
certain parts of the United Statesabout nine out of ten mules have
striped legs. I once saw a mule with its legs so much striped that any one
might have thought that it was a hybrid zebra; and Mr. W.C. Martinin his
excellent treatise on the horsehas given a figure of a similar mule. In
four coloured drawingswhich I have seenof hybrids between the ass and
zebrathe legs were much more plainly barred than the rest of the body;
and in one of them there was a double shoulder-stripe. In Lord Morton's
famous hybridfrom a chestnut mare and male quaggathe hybrid and even
the pure offspring subsequently produced from the same mare by a black
Arabian sirewere much more plainly barred across the legs than is even
the pure quagga. Lastlyand this is another most remarkable casea
hybrid has been figured by Dr. Gray (and he informs me that he knows of a
second case) from the ass and the hemionus; and this hybridthough the ass
only occasionally has stripes on his legs and the hemionus has none and has
not even a shoulder-stripenevertheless had all four legs barredand had
three short shoulder-stripeslike those on the dun Devonshire and Welsh
poniesand even had some zebra-like stripes on the sides of its face.
With respect to this last factI was so convinced that not even a stripe
of colour appears from what is commonly called chancethat 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 horsesand wasas we have
seenanswered in the affirmative.

What now are we to say to these several facts? We see several distinct
species of the horse genus becomingby simple variationstriped on the
legs like a zebraor striped on the shoulders like an ass. 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. The appearance of the stripes is not accompanied by any change of
formor by any other new character. We see this tendency to become
striped most strongly displayed in hybrids from between several of the most
distinct species. Now observe the case of the several breeds of pigeons:
they are descended from a pigeon (including two or three sub-species or
geographical races) of a bluish colourwith certain bars and other marks;
and when any breed assumes by simple variation a bluish tintthese bars
and other marks invariably reappear; but without any other change of form
or character. When the oldest and truest breeds of various colours are
crossedwe see a strong tendency for the blue tint and bars and marks to
reappear in the mongrels. I have stated that the most probable hypothesis
to account for the reappearance of very ancient charactersis--that there
is a TENDENCY in the young of each successive generation to produce the
long-lost characterand that this tendencyfrom unknown causessometimes
prevails. 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. Call the breeds of pigeonssome of which have bred true for
centuriesspecies; and how exactly parallel is the case with that of the
species of the horse genus! For myselfI venture confidently to look back
thousands on thousands of generationsand I see an animal striped like a
zebrabut perhaps otherwise very differently constructedthe common
parent of our domestic horse (whether or not it be descended from one or
more wild stocks) of the assthe hemionusquaggaand zebra.

He who believes that each equine species was independently createdwillI
presumeassert that each species has been created with a tendency to vary
both under nature and under domesticationin this particular mannerso as
often to become striped like the other species of the genus; and that each
has been created with a strong tendencywhen crossed with species
inhabiting distant quarters of the worldto produce hybrids resembling in
their stripesnot their own parentsbut other species of the genus. To
admit this view isas it seems to meto reject a real for an unrealor
at least for an unknown cause. It makes the works of God a mere mockery
and deception; I would almost as soon believe with the old and ignorant
cosmogoniststhat fossil shells had never livedbut had been created in
stone so as to mock the shells now living on the sea-shore.


Our ignorance of the laws of variation is profound. Not in one case out of
a hundred can we pretend to assign any reason why this or that part has
varied. But whenever we have the means of instituting a comparisonthe
same laws appear to have acted in producing the lesser differences between
varieties of the same speciesand the greater differences between species
of the same genus. Changed conditions generally induce mere fluctuating
variabilitybut sometimes they cause direct and definite effects; and
these may become strongly marked in the course of timethough we have not
sufficient evidence on this head. Habit in producing constitutional
peculiaritiesand use in strengtheningand disuse in weakening and
diminishing organsappear in many cases to have been potent in their
effects. Homologous parts tend to vary in the same mannerand homologous
parts tend to cohere. Modifications in hard parts and in external parts
sometimes affect softer and internal parts. When one part is largely
developedperhaps it tends to draw nourishment from the adjoining parts;
and every part of the structure which can be saved without detriment will
be saved. Changes of structure at an early age may affect parts
subsequently developed; and many cases of correlated variationthe nature
of which we are unable to understandundoubtedly occur. Multiple parts

are variable in number and in structureperhaps arising from such parts
not having been closely specialised for any particular functionso that
their modifications have not been closely checked by natural selection. It
follows probably from this same causethat organic beings low in the scale
are more variable than those standing higher in the scaleand which have
their whole organisation more specialised. Rudimentary organsfrom being
uselessare not regulated by natural selectionand hence are variable.
Specific characters--that isthe 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 charactersor those which have long
been inheritedand have not differed within this same period. In these
remarks we have referred to special parts or organs being still variable
because they have recently varied and thus come to differ; but we have also
seen in the second chapter that the same principle applies to the whole
individual; for in a district where many species of a genus are found--that
iswhere there has been much former variation and differentiationor
where the manufactory of new specific forms has been actively at work--in
that district and among these specieswe now findon an averagemost
varieties. Secondary sexual characters are highly variableand such
characters differ much in the species of the same group. Variability in
the same parts of the organisation has generally been taken advantage of in
giving secondary sexual differences to the two sexes of the same species
and specific differences to the several species of the same genus. Any
part or organ developed to an extraordinary size or in an extraordinary
mannerin comparison with the same part or organ in the allied species
must have gone through an extraordinary amount of modification since the
genus arose; and thus we can understand why it should often still be
variable in a much higher degree than other parts; for variation is a
long-continued and slow processand 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. 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 processrequiring a
long lapse of time--in this casenatural selection has succeeded in giving
a fixed character to the organin however extraordinary a manner it may
have been developed. Species inheriting nearly the same constitution from
a common parentand exposed to similar influencesnaturally tend to
present analogous variationsor these same species may occasionally revert
to some of the characters of their ancient progenitors. Although new and
important modifications may not arise from reversion and analogous
variationsuch modifications will add to the beautiful and harmonious
diversity of nature.

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.



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.

Long before the reader has arrived at this part of my worka crowd of

difficulties will have occurred to him. Some of them are so serious that
to this day I can hardly reflect on them without being in some degree
staggered; butto the best of my judgmentthe greater number are only
apparentand those that are real are notI thinkfatal to the theory.

These difficulties and objections may be classed under the following heads:
Firstwhyif species have descended from other species by fine
gradationsdo we not everywhere see innumerable transitional forms? Why
is not all nature in confusioninstead of the species beingas we see
themwell defined?

Secondlyis it possible that an animal havingfor instancethe structure
and habits of a batcould have been formed by the modification of some
other animal with widely different habits and structure? Can we believe
that natural selection could produceon the one handan organ of trifling
importancesuch as the tail of a giraffewhich serves as a fly-flapper
andon the other handan organ so wonderful as the eye?

Thirdlycan instincts be acquired and modified through natural selection?
What shall we say to the instinct which leads the bee to make cellsand
which has practically anticipated the discoveries of profound

Fourthlyhow can we account for specieswhen crossedbeing sterile and
producing sterile offspringwhereaswhen varieties are crossedtheir
fertility is unimpaired?

The two first heads will be here discussed; some miscellaneous objections
in the following chapter; Instinct and Hybridism in the two succeeding


As natural selection acts solely by the preservation of profitable
modificationseach new form will tend in a fully-stocked country to take
the place ofand finally to exterminateits own less improved parent-form
and other less-favoured forms with which it comes into competition. Thus
extinction and natural selection go hand in hand. Henceif we look at
each species as descended from some unknown formboth 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.

Butas by this theory innumerable transitional forms must have existed
why do we not find them embedded in countless numbers in the crust of the
earth? It will be more convenient to discuss this question in the chapter
on the imperfection of the geological record; and I will here only state
that I believe the answer mainly lies in the record being incomparably less
perfect than is generally supposed. The crust of the earth is a vast
museum; but the natural collections have been imperfectly madeand only at
long intervals of time.

But it may be urged that when several closely allied species inhabit the
same territorywe surely ought to find at the present time many
transitional forms. Let us take a simple case: in travelling from north
to south over a continentwe generally meet at successive intervals with
closely allied or representative speciesevidently filling nearly the same
place in the natural economy of the land. These representative species
often meet and interlock; and as the one becomes rarer and rarerthe other
becomes more and more frequenttill the one replaces the other. But if we
compare these species where they interminglethey are generally as
absolutely distinct from each other in every detail of structure as are
specimens taken from the metropolis inhabited by each. By my theory these
allied species are descended from a common parent; and during the process
of modificationeach has become adapted to the conditions of life of its

own regionand has supplanted and exterminated its original parent-form
and all the transitional varieties between its past and present states.
Hence we ought not to expect at the present time to meet with numerous
transitional varieties in each regionthough they must have existed there
and may be embedded there in a fossil condition. But in the intermediate
regionhaving intermediate conditions of lifewhy do we not now find
closely-linking intermediate varieties? This difficulty for a long time
quite confounded me. But I think it can be in large part explained.

In the first place we should be extremely cautious in inferringbecause an
area is now continuousthat it has been continuous during a long period.
Geology would lead us to believe that most continents have been broken up
into islands even during the later tertiary periods; and in such islands
distinct species might have been separately formed without the possibility
of intermediate varieties existing in the intermediate zones. By changes
in the form of the land and of climatemarine areas now continuous must
often have existed within recent times in a far less continuous and uniform
condition than at present. But I will pass over this way of escaping from
the difficulty; for I believe that many perfectly defined species have been
formed on strictly continuous areas; though I do not doubt that the
formerly broken condition of areas now continuoushas played an important
part in the formation of new speciesmore especially with freely-crossing
and wandering animals.

In looking at species as they are now distributed over a wide areawe
generally find them tolerably numerous over a large territorythen
becoming somewhat abruptly rarer and rarer on the confinesand finally
disappearing. Hence the neutral territory between two representative
species is generally narrow in comparison with the territory proper to
each. We see the same fact in ascending mountainsand sometimes it is
quite remarkable how abruptlyas Alph. De Candolle has observeda common
alpine species disappears. The same fact has been noticed by E. Forbes in
sounding the depths of the sea with the dredge. To those who look at
climate and the physical conditions of life as the all-important elements
of distributionthese facts ought to cause surpriseas climate and height
or depth graduate away insensibly. But when we bear in mind that almost
every specieseven in its metropoliswould increase immensely in numbers
were it not for other competing species; that nearly all either prey on or
serve as prey for others; in shortthat each organic being is either
directly or indirectly related in the most important manner to other
organic beings--we see that the range of the inhabitants of any country by
no means exclusively depends on insensibly changing physical conditions
but in large part on the presence of other specieson which it livesor
by which it is destroyedor with which it comes into competition; and as
these species are already defined objectsnot blending one into another by
insensible gradationsthe range of any one speciesdepending as it does
on the range of otherswill tend to be sharply defined. Moreovereach
species on the confines of its rangewhere it exists in lessened numbers
willduring fluctuations in the number of its enemies or of its preyor
in the nature of the seasonsbe extremely liable to utter extermination;
and thus its geographical range will come to be still more sharply defined.

As allied or representative specieswhen inhabiting a continuous areaare
generally distributed in such a manner that each has a wide rangewith a
comparatively narrow neutral territory between themin which they become
rather suddenly rarer and rarer; thenas varieties do not essentially
differ from speciesthe same rule will probably apply to both; and if we
take a varying species inhabiting a very large areawe shall have to adapt
two varieties to two large areasand a third variety to a narrow
intermediate zone. The intermediate varietyconsequentlywill exist in
lesser numbers from inhabiting a narrow and lesser area; and practically
as far as I can make outthis rule holds good with varieties in a state of
nature. I have met with striking instances of the rule in the case of
varieties intermediate between well-marked varieties in the genus Balanus.

And it would appear from information given me by Mr. WatsonDr. Asa Gray
and Mr. Wollastonthat generallywhen varieties intermediate between two
other forms occurthey are much rarer numerically than the forms which
they connect. Nowif we may trust these facts and inferencesand
conclude that varieties linking two other varieties together generally have
existed in lesser numbers than the forms which they connectthen we can
understand why intermediate varieties should not endure for very long
periods: whyas a general rulethey should be exterminated and
disappearsooner than the forms which they originally linked together.

For any form existing in lesser numbers wouldas already remarkedrun a
greater chance of being exterminated than one existing in large numbers;
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. But
it is a far more important considerationthat during the process of
further modificationby which two varieties are supposed to be converted
and perfected into two distinct speciesthe two which exist in larger
numbersfrom inhabiting larger areaswill have a great advantage over the
intermediate varietywhich exists in smaller numbers in a narrow and
intermediate zone. For forms existing in larger numbers will have a better
chancewithin any given periodof presenting further favourable
variations for natural selection to seize onthan will the rarer forms
which exist in lesser numbers. Hencethe more common formsin the race
for lifewill tend to beat and supplant the less common formsfor these
will be more slowly modified and improved. It is the same principle which
as I believeaccounts for the common species in each countryas shown in
the second chapterpresenting on an average a greater number of
well-marked varieties than do the rarer species. I may illustrate what I
mean by supposing three varieties of sheep to be keptone adapted to an
extensive mountainous region; a second to a comparatively narrowhilly
tract; and a third to the wide plains at the base; and that the inhabitants
are all trying with equal steadiness and skill to improve their stocks by
selection; 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 narrowhilly tract; and
consequently the improved mountain or plain breed will soon take the place
of the less improved hill breed; and thus the two breedswhich originally
existed in greater numberswill come into close contact with each other
without the interposition of the supplantedintermediate hill variety.

To sum upI believe that species come to be tolerably well-defined
objectsand do not at any one period present an inextricable chaos of
varying and intermediate links: firstbecause new varieties are very
slowly formedfor variation is a slow processand natural selection can
do nothing until favourable individual differences or variations occurand
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 such new
places will depend on slow changes of climateor on the occasional
immigration of new inhabitantsandprobablyin a still more important
degreeon some of the old inhabitants becoming slowly modifiedwith the
new forms thus produced and the old ones acting and reacting on each other.
So thatin any one region and at any one timewe ought to see only a few
species presenting slight modifications of structure in some degree
permanent; and this assuredly we do see.

Secondlyareas now continuous must often have existed within the recent
period as isolated portionsin which many formsmore especially among the
classes which unite for each birth and wander muchmay have separately
been rendered sufficiently distinct to rank as representative species. In
this caseintermediate varieties between the several representative
species and their common parentmust formerly have existed within each
isolated portion of the landbut these links during the process of natural
selection will have been supplanted and exterminatedso that they will no
longer be found in a living state.

Thirdlywhen two or more varieties have been formed in different portions
of a strictly continuous areaintermediate varieties willit is probable
at first have been formed in the intermediate zonesbut they will
generally have had a short duration. For these intermediate varieties
willfrom reasons already assigned (namely from what we know of the actual
distribution of closely allied or representative speciesand likewise of
acknowledged varieties)exist in the intermediate zones in lesser numbers
than the varieties which they tend to connect. From this cause alone the
intermediate varieties will be liable to accidental extermination; and
during the process of further modification through natural selectionthey
will almost certainly be beaten and supplanted by the forms which they
connect; for thesefrom existing in greater numbers willin the
aggregatepresent more varietiesand thus be further improved through
natural selection and gain further advantages.

Lastlylooking not to any one timebut at all timeif my theory be true
numberless intermediate varietieslinking closely together all the species
of the same groupmust assuredly have existed; but the very process of
natural selection constantly tendsas has been so often remarkedto
exterminate the parent forms and the intermediate links. Consequently
evidence of their former existence could be found only among fossil remains
which are preservedas we shall attempt to show in a future chapterin an
extremely imperfect and intermittent record.


It has been asked by the opponents of such views as I holdhowfor
instancecould a land carnivorous animal have been converted into one with
aquatic habits; for how could the animal in its transitional state have
subsisted? It would be easy to show that there now exist carnivorous
animals presenting close intermediate grades from strictly terrestrial to
aquatic habits; and as each exists by a struggle for lifeit is clear that
each must be well adapted to its place in nature. Look at the Mustela
vison of North Americawhich has webbed feetand which resembles an otter
in its furshort legsand form of tail; during summer this animal dives
for and preys on fishbut during the long winter it leaves the frozen
watersand preyslike other polecats on mice and land animals. If a
different case had been takenand it had been asked how an insectivorous
quadruped could possibly have been converted into a flying batthe
question would have been far more difficult to answer. Yet I think such
difficulties have little weight.

Hereas on other occasionsI lie under a heavy disadvantageforout of
the many striking cases which I have collectedI can give only one or two
instances of transitional habits and structures in allied species; and of
diversified habitseither constant or occasionalin the same species.
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

Look at the family of squirrels; here we have the finest gradation from
animals with their tails only slightly flattenedand from othersas Sir

J. Richardson has remarkedwith the posterior part of their bodies rather
wide and with the skin on their flanks rather fullto the so-called flying
squirrels; and flying squirrels have their limbs and even the base of the
tail united by a broad expanse of skinwhich serves as a parachute and
allows them to glide through the air to an astonishing distance from tree
to tree. We cannot doubt that each structure is of use to each kind of
squirrel in its own countryby enabling it to escape birds or beasts of
preyor to collect food more quicklyoras there is reason to believe
to lessen the danger from occasional falls. But it does not follow from
this fact that the structure of each squirrel is the best that it is

possible to conceive under all possible conditions. Let the climate and
vegetation changelet other competing rodents or new beasts of prey
immigrateor old ones become modifiedand all analogy would lead us to
believe that someat leastof the squirrels would decrease in numbers or
become exterminatedunless they also become modified and improved in
structure in a corresponding manner. ThereforeI can see no difficulty
more especially under changing conditions of lifein the continued
preservation of individuals with fuller and fuller flank-membraneseach
modification being usefuleach being propagateduntilby the accumulated
effects of this process of natural selectiona perfect so-called flying
squirrel was produced.

Now look at the Galeopithecus or so-called flying lemurwhich was formerly
ranked among batsbut is now believed to belong to the Insectivora. An
extremely wide flank-membrane stretches from the corners of the jaw to the
tailand includes the limbs with the elongated fingers. This flankmembrane
is furnished with an extensor muscle. Although no graduated links
of structurefitted for gliding through the airnow connect the
Galeopithecus with the other Insectivorayet there is no difficulty in
supposing that such links formerly existedand that each was developed in
the same manner as with the less perfectly gliding squirrels; each grade of
structure having been useful to its possessor. 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; and thisas far as the
organs of flight are concernedwould have converted the animal into a bat.
In certain bats in which the wing-membrane extends from the top of the
shoulder to the tail and includes the hind-legswe perhaps see traces of
an apparatus originally fitted for gliding through the air rather than for

If about a dozen genera of birds were to become extinctwho would have
ventured to surmise that birds might have existed which used their wings
solely as flapperslike the logger headed duck (Micropterus of Eyton); as
fins in the water and as front legs on the landlike the penguin; as
sailslike the ostrich; and functionally for no purposelike the apteryx?
Yet the structure of each of these birds is good for itunder the
conditions of life to which it is exposedfor each has to live by a
struggle: but it is not necessarily the best possible under all possible
conditions. It must not be inferred from these remarks that any of the
grades of wing-structure here alluded towhich perhaps may all be the
result of disuseindicate the steps by which birds actually acquired their
perfect power of flight; but they serve to show what diversified means of
transition are at least possible.

Seeing that a few members of such water-breathing classes as the Crustacea
and Mollusca are adapted to live on the land; and seeing that we have
flying birds and mammalsflying insects of the most diversified typesand
formerly had flying reptilesit is conceivable that flying-fishwhich now
glide far through the airslightly rising and turning by the aid of their
fluttering finsmight have been modified into perfectly winged animals.
If this had been effectedwho would have ever imagined that in an early
transitional state they had been inhabitants of the open oceanand had
used their incipient organs of flight exclusivelyso far as we knowto
escape being devoured by other fish?

When we see any structure highly perfected for any particular habitas the
wings of a bird for flightwe should bear in mind that animals displaying
early transitional grades of the structure will seldom have survived to the
present dayfor they will have been supplanted by their successorswhich
were gradually rendered more perfect through natural selection.
Furthermorewe 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. Thus

to return to our imaginary illustration of the flying-fishit does not
seem probable that fishes capable of true flight would have been developed
under many subordinate formsfor taking prey of many kinds in many ways
on the land and in the wateruntil their organs of flight had come to a
high stage of perfectionso as to have given them a decided advantage over
other animals in the battle for life. Hence the chance of discovering
species with transitional grades of structure in a fossil condition will
always be lessfrom their having existed in lesser numbersthan in the
case of species with fully developed structures.

I will now give two or three instancesboth of diversified and of changed
habitsin the individuals of the same species. In either case it would be
easy for natural selection to adapt the structure of the animal to its
changed habitsor exclusively to one of its several habits. It is
howeverdifficult to decide and immaterial for uswhether habits
generally change first and structure afterwards; or whether slight
modifications of structure lead to changed habits; both probably often
occurring almost simultaneously. Of cases of changed habits it will
suffice merely to allude to that of the many British insects which now feed
on exotic plantsor exclusively on artificial substances. Of diversified
habits innumerable instances could be given: I have often watched a tyrant
flycatcher (Saurophagus sulphuratus) in South Americahovering over one
spot and then proceeding to anotherlike a kestreland at other times
standing stationary on the margin of waterand then dashing into it like a
kingfisher at a fish. In our own country the larger titmouse (Parus major)
may be seen climbing branchesalmost like a creeper; it sometimeslike a
shrikekills small birds by blows on the head; and I have many times seen
and heard it hammering the seeds of the yew on a branchand thus breaking
them like a nuthatch. In North America the black bear was seen by Hearne
swimming for hours with widely open mouththus catchingalmost like a
whaleinsects in the water.

As we sometimes see individuals following habits different from those
proper to their species and to the other species of the same genuswe
might expect that such individuals would occasionally give rise to new
specieshaving anomalous habitsand with their structure either slightly
or considerably modified from that of their type. And such instances occur
in nature. 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
fruitand others with elongated wings which chase insects on the wing. On
the plains of La Platawhere hardly a tree growsthere is a woodpecker
(Colaptes campestris) which has two toes before and two behinda longpointed
tonguepointed tail-featherssufficiently stiff to support the
bird in a vertical position on a postbut not so stiff as in the typical
wood-peckersand a straightstrong beak. The beakhoweveris not so
straight or so strong as in the typical woodpeckers but it is strong enough
to bore into wood. Hence this Colaptesin all the essential parts of its
structureis a woodpecker. Even in such trifling characters as the
colouringthe harsh tone of the voiceand undulatory flightits close
blood-relationship to our common woodpecker is plainly declared; yetas I
can assertnot only from my own observationsbut from those of the
accurate Azarain certain large districts it does not climb treesand it
makes its nest in holes in banks! In certain other districtshowever
this same woodpeckeras Mr. Hudson statesfrequents treesand bores
holes in the trunk for its nest. I may mention as another illustration of
the varied habits of this genusthat a Mexican Colaptes has been described
by De Saussure as boring holes into hard wood in order to lay up a store of

Petrels are the most aerial and oceanic of birdsbutin the quiet sounds
of Tierra del Fuegothe Puffinuria berardiin its general habitsin its
astonishing power of divingin its manner of swimming and of flying when
made to take flightwould be mistaken by any one for an auk or a grebe;

neverthelessit is essentially a petrelbut with many parts of its
organisation profoundly modified in relation to its new habits of life;
whereas the woodpecker of La Plata has had its structure only slightly
modified. In the case of the water-ouzelthe acutest observerby
examining its dead bodywould never have suspected its sub-aquatic habits;
yet this birdwhich is allied to the thrush familysubsists by
diving--using its wings under water and grasping stones with its feet.
All the members of the great order of Hymenopterous insects are
terrestrialexcepting the genus Proctotrupeswhich Sir John Lubbock has
discovered to be aquatic in its habits; it often enters the water and dives
about by the use not of its legs but of its wingsand remains as long as
four hours beneath the surface; yet it exhibits no modification in
structure in accordance with its abnormal habits.

He who believes that each being has been created as we now see itmust
occasionally have felt surprise when he has met with an animal having
habits and structure not in agreement. What can be plainer than that the
webbed feet of ducks and geese are formed for swimming? Yet there are
upland geese with webbed feet which rarely go near the water; and no one
except Audubonhas seen the frigate-birdwhich has all its four toes
webbedalight on the surface of the ocean. On the other handgrebes and
coots are eminently aquaticalthough their toes are only bordered by
membrane. What seems plainer than that the long toesnot furnished with
membraneof the Grallatoresare formed for walking over swamps and
floating plants. The water-hen and landrail are members of this orderyet
the first is nearly as aquatic as the cootand the second is nearly as
terrestrial as the quail or partridge. In such casesand many others
could be givenhabits have changed without a corresponding change of
structure. The webbed feet of the upland goose may be said to have become
almost rudimentary in functionthough not in structure. In the
frigate-birdthe deeply scooped membrane between the toes shows that
structure has begun to change.

He who believes in separate and innumerable acts of creation may saythat
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; but this seems to me only
restating the fact in dignified language. He who believes in the struggle
for existence and in the principle of natural selectionwill acknowledge
that every organic being is constantly endeavouring to increase in numbers;
and that if any one being varies ever so littleeither in habits or
structureand thus gains an advantage over some other inhabitant of the
same countryit will seize on the place of that inhabitanthowever
different that may be from its own place. Hence it will cause him no
surprise that there should be geese and frigate-birds with webbed feet
living on the dry land and rarely alighting on the waterthat there should
be long-toed corncrakesliving in meadows instead of in swamps; that there
should be woodpeckers where hardly a tree grows; that there should be
diving thrushes and diving Hymenopteraand petrels with the habits of


To suppose that the eye with all its inimitable contrivances for adjusting
the focus to different distancesfor admitting different amounts of light
and for the correction of spherical and chromatic aberrationcould have
been formed by natural selectionseemsI freely confessabsurd in the
highest degree. When it was first said that the sun stood still and the
world turned roundthe common sense of mankind declared the doctrine
false; but the old saying of Vox populivox Deias every philosopher
knowscannot be trusted in science. Reason tells methat if numerous
gradations from a simple and imperfect eye to one complex and perfect can
be shown to existeach grade being useful to its possessoras is
certainly the case; if furtherthe eye ever varies and the variations be
inheritedas is likewise certainly the case; and if such variations should

be useful to any animal under changing conditions of lifethen the
difficulty of believing that a perfect and complex eye could be formed by
natural selectionthough insuperable by our imaginationshould not be
considered as subversive of the theory. How a nerve comes to be sensitive
to lighthardly concerns us more than how life itself originated; but I
may remark thatas some of the lowest organisms in which nerves cannot be
detectedare capable of perceiving lightit does not seem impossible that
certain sensitive elements in their sarcode should become aggregated and
developed into nervesendowed with this special sensibility.

In searching for the gradations through which an organ in any species has
been perfectedwe ought to look exclusively to its lineal progenitors; but
this is scarcely ever possibleand we are forced to look to other species
and genera of the same groupthat is to the collateral descendants from
the same parent-formin order to see what gradations are possibleand for
the chance of some gradations having been transmitted in an unaltered or
little altered condition. But the state of the same organ in distinct
classes may incidentally throw light on the steps by which it has been

The simplest organ which can be called an eye consists of an optic nerve
surrounded by pigment-cells and covered by translucent skinbut without
any lens or other refractive body. We mayhoweveraccording to M.
Jourdaindescend even a step lower and find aggregates of pigment-cells
apparently serving as organs of visionwithout any nervesand resting
merely on sarcodic tissue. Eyes of the above simple nature are not capable
of distinct visionand serve only to distinguish light from darkness. In
certain star-fishessmall depressions in the layer of pigment which
surrounds the nerve are filledas described by the author just quoted
with transparent gelatinous matterprojecting with a convex surfacelike
the cornea in the higher animals. He suggests that this serves not to form
an imagebut only to concentrate the luminous rays and render their
perception more easy. In this concentration of the rays we gain the first
and by far the most important step towards the formation of a true
picture-forming eye; for we have only to place the naked extremity of the
optic nervewhich in some of the lower animals lies deeply buried in the
bodyand in some near the surfaceat the right distance from the
concentrating apparatusand an image will be formed on it.

In the great class of the Articulatawe may start from an optic nerve
simply coated with pigmentthe latter sometimes forming a sort of pupil
but destitute of lens or other optical contrivance. With insects it is now
known that the numerous facets on the cornea of their great compound eyes
form true lensesand that the cones include curiously modified nervous
filaments. But these organs in the Articulata are so much diversified that
Muller formerly made three main classes with seven subdivisionsbesides a
fourth main class of aggregated simple eyes.

When we reflect on these factshere given much too brieflywith respect
to the widediversifiedand graduated range of structure in the eyes of
the lower animals; and when we bear in mind how small the number of all
living forms must be in comparison with those which have become extinct
the difficulty ceases to be very great in believing that natural selection
may have converted the simple apparatus of an optic nervecoated with
pigment and invested by transparent membraneinto an optical instrument as
perfect as is possessed by any member of the Articulata class.

He who will go thus farought not to hesitate to go one step furtherif
he finds on finishing this volume that large bodies of factsotherwise
inexplicablecan be explained by the theory of modification through
natural selection; he ought to admit that a structure even as perfect as an
eagle's eye might thus be formedalthough in this case he does not know
the transitional states. It has been objected that in order to modify the
eye and still preserve it as a perfect instrumentmany changes would have

to be effected simultaneouslywhichit is assumedcould not be done
through natural selection; but as I have attempted to show in my work on
the variation of domestic animalsit is not necessary to suppose that the
modifications were all simultaneousif they were extremely slight and
gradual. Different kinds of modification wouldalsoserve for the same
general purpose: as Mr. Wallace has remarkedIf 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
namelythe Vertebratawe can start from an eye so simplethat it
consistsas in the lanceletof a little sack of transparent skin
furnished with a nerve and lined with pigmentbut destitute of any other
apparatus. In fishes and reptilesas Owen has remarkedThe range of
gradation of dioptric structures is very great.It is a significant fact
that even in manaccording to the high authority of Virchowthe beautiful
crystalline lens is formed in the embryo by an accumulation of epidermic
cellslying in a sack-like fold of the skin; and the vitreous body is
formed from embryonic subcutaneous tissue. To arrivehoweverat a just
conclusion regarding the formation of the eyewith all its marvellous yet
not absolutely perfect charactersit 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 instrumentwe ought in imagination to take a thick layer of
transparent tissuewith spaces filled with fluidand with a nerve
sensitive to light beneathand then suppose every part of this layer to be
continually changing slowly in densityso as to separate into layers of
different densities and thicknessesplaced at different distances from
each otherand with the surfaces of each layer slowly changing in form.
Further we must suppose that there is a powerrepresented by natural
selection or the survival of the fittestalways intently watching each
slight alteration in the transparent layers; and carefully preserving each
whichunder varied circumstancesin any way or degreetends 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 bodiesvariation
will cause the slight alterationgeneration will multiply them almost
infinitelyand 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
glassas the works of the Creator are to those of man?


If it could be demonstrated that any complex organ existedwhich could not
possibly have been formed by numeroussuccessiveslight modificationsmy
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 speciesaround which
according to the theorythere has been much extinction. Or againif we
take an organ common to all the members of a classfor in this latter case
the organ must have been originally formed at a remote periodsince 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 formslong since become

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 respiresdigestsand excretes. In
the Hydrathe animal may be turned inside outand the exterior surface
will then digest and the stomach respire. In such cases natural selection
might specialiseif any advantage were thus gainedthe whole or part of
an organwhich had previously performed two functionsfor one function
aloneand 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 alonea great change
would be effected with comparative suddenness in the character of the
species. It ishoweverprobable 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.

Againtwo distinct organsor the same organ under two very different
formsmay 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 waterat the same time that they breathe free air in
their swim-bladdersthis 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 meansby spirally twiningby clasping a support with their
sensitive tendrilsand by the emission of aerial rootlets; these three
means are usually found in distinct groupsbut some few species exhibit
two of the meansor even all threecombined 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 workbeing aided during the progress of
modification by the other organ; and then this other organ might be
modified for some other and quite distinct purposeor be wholly

The illustration of the swim-bladder in fishes is a good onebecause it
shows us clearly the highly important fact that an organ originally
constructed for one purposenamely flotationmay be converted into one
for a widely different purposenamely respiration. The swim-bladder has
alsobeen worked in as an accessory to the auditory organs of certain
fishes. All physiologists admit that the swim-bladder is homologousor
ideally similarin 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 lungsor 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 thusas I infer from Professor Owen's interesting description of
these partsunderstand the strange fact that every particle of food and
drink which we swallow has to pass over the orifice of the tracheawith
some risk of falling into the lungsnotwithstanding 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 instanceLandois 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 organsit is so important to bear in mind
the probability of conversion from one function to anotherthat I will
give another instance. Pedunculated cirripedes have two minute folds of
skincalled by me the ovigerous frenawhich servethrough the means of a
sticky secretionto retain the eggs until they are hatched within the
sack. These cirripedes have no branchiaethe whole surface of the body
and of the sacktogether with the small frenaserving for respiration.
The Balanidae or sessile cirripedeson the other handhave no ovigerous
frenathe eggs lying loose at the bottom of the sackwithin the
well-enclosed shell; but they havein the same relative position with the
frenalargemuch-folded membraneswhich freely communicate with the
circulatory lacunae of the sack and bodyand 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; indeedthey graduate into each other.
Therefore it need not be doubted that the two little folds of skinwhich
originally served as ovigerous frenabut whichlikewisevery slightly
aided in the act of respirationhave been gradually converted by natural
selection into branchiaesimply through an increase in their size and the
obliteration of their adhesive glands. If all pedunculated cirripedes had
become extinctand they have suffered far more extinction than have
sessile cirripedeswho 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 transitionnamelythrough 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
agebefore they have acquired their perfect characters; and if this power
became thoroughly well developed in a speciesit 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 formthe character
of the species would be greatly changed and degraded. Againnot a few
animalsafter arriving at maturitygo on changing in character during
nearly their whole lives. With mammalsfor instancethe form of the
skull is often much altered with ageof which Dr. Murie has given some
striking instances with seals. Every one knows how the horns of stags
become more and more branchedand the plumes of some birds become more
finely developedas they grow older. Professor Cope states that the teeth
of certain lizards change much in shape with advancing years. With
crustaceans not only many trivialbut some important parts assume a new
characteras recorded by Fritz Mullerafter maturity. In all such cases-
and many could be given--if the age for reproduction were retardedthe
character of the speciesat least in its adult statewould 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
transitionI can form no opinion; but if this has occurredit is probable
that the differences between the young and the matureand between the
mature and the oldwere primordially acquired by graduated steps.


Although we must be extremely cautious in concluding that any organ could
not have been produced by successivesmalltransitional gradationsyet
undoubtedly serious cases of difficulty occur.

One of the most serious is that of neuter insectswhich 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
surprisingfor we do not even know of what use they are. In the gymnotus
and torpedo they no doubt serve as powerful means of defenceand perhaps
for securing prey; yet in the rayas observed by Matteuccian analogous
organ in the tail manifests but little electricityeven when the animal is
greatly irritated; so little that it can hardly be of any use for the above
purposes. Moreoverin the raybesides the organ just referred tothere
isas Dr. R. McDonnell has shownanother organ near the headnot known
to be electricalbut 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 analogyin
intimate structurein the distribution of the nervesand in the manner in
which they are acted on by various reagents. It shouldalsobe
especially observed that muscular contraction is accompanied by an
electrical discharge; andas Dr. Radcliffe insistsin 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 organsand
as we know nothing about the habits and structure of the progenitors of the
existing electric fishesit 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 fishof which several
are widely remote in their affinities. When the same organ is found in
several members of the same classespecially if in members having very
different habits of lifewe 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 thatif the electric
organs had been inherited from some one ancient progenitorwe 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 organswhich their
modified descendants have now lost. But when we look at the subject more
closelywe find in the several fishes provided with electric organsthat
these are situated in different parts of the bodythat they differ in
constructionas in the arrangement of the platesandaccording to
Paciniin the process or means by which the electricity is excited--and
lastlyin 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 organsthese cannot be
considered as homologousbut 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 organapparently
the samearising in several remotely allied speciesdisappearsleaving
only the lesser yet still great difficulty: namelyby what graduated
steps these organs have been developed in each separate group of fishes.

The luminous organs which occur in a few insectsbelonging to widely
different familiesand which are situated in different parts of the body
offerunder our present state of ignorancea difficulty almost exactly
parallel with that of the electric organs. Other similar cases could be
given; for instance in plantsthe very curious contrivance of a mass of
pollen-grainsborne on a foot-stalk with an adhesive glandis apparently
the same in Orchis and Asclepiasgenera almost as remote as is possible
among flowering plants; but here again the parts are not homologous. In
all cases of beingsfar removed from each other in the scale of

organisationwhich are furnished with similar and peculiar organsit will
be found that although the general appearance and function of the organs
may be the sameyet fundamental differences between them can always be
detected. For instancethe 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
difficultybut I am unable to see the force of his argument. An organ for
vision must be formed of transparent tissueand must include some sort of
lens for throwing an image at the back of a darkened chamber. Beyond this
superficial resemblancethere is hardly any real similarity between the
eyes of cuttle-fish and vertebratesas may be seen by consulting Hensen's
admirable memoir on these organs in the Cephalopoda. It is impossible for
me here to enter on detailsbut I may specify a few of the points of
difference. The crystalline lens in the higher cuttle-fish consists of two
partsplaced one behind the other like two lensesboth having a very
different structure and disposition to what occurs in the vertebrata. The
retina is wholly differentwith an actual inversion of the elemental
partsand 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 conceiveand 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 isof courseopen 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
anticipatedin accordance with this view of their manner of formation. As
two men have sometimes independently hit on the same inventionso in the
several foregoing cases it appears that natural selectionworking for the
good of each beingand taking advantage of all favourable variationshas
produced similar organsas far as function is concernedin distinct
organic beingswhich owe none of their structure in common to inheritance
from a common progenitor.

Fritz Mullerin 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 speciespossessing an air-breathing
apparatus and fitted to live out of the water. In two of these families
which were more especially examined by Mullerand which are nearly related
to each otherthe species agree most closely in all important characters:
namely in their sense organscirculating systemsin the position of the
tufts of hair within their complex stomachsand lastly in the whole
structure of the water-breathing branchiaeeven 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 landthe equally
important air-breathing apparatus would have been the same; for why should
this one apparatusgiven for the same purposehave been made to differ
while all the other important organs were closely similaror rather

Fritz Muller argues that this close similarity in so many points of
structure mustin accordance with the views advanced by mebe accounted
for by inheritance from a common progenitor. But as the vast majority of
the species in the above two familiesas well as most other crustaceans
are aquatic in their habitsit 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 pointsas
in the position of the orificesin the manner in which they are opened and
closedand in some accessory details. Now such differences are
intelligibleand might even have been expectedon the supposition that
species belonging to distinct families had slowly become adapted to live
more and more out of waterand to breathe the air. For these species

from belonging to distinct familieswould have differed to a certain
extentand in accordance with the principle that the nature of each
variation depends on two factorsviz.the nature of the organism and that
of the surrounding conditionstheir variability assuredly would not have
been exactly the same. Consequently natural selection would have had
different materials or variations to work onin 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 zoologistthe late Professor Claparedehas argued
in the same mannerand has arrived at the same result. He shows that
there are parasitic mites (Acaridae)belonging to distinct sub-families
and familieswhich are furnished with hair-claspers. These organs must
have been independently developedas they could not have been inherited
from a common progenitor; and in the several groups they are formed by the
modification of the fore legsof the hind legsof the maxillae or lips
and of appendages on the under side of the hind part of the body.

In the foregoing caseswe see the same end gained and the same function
performedin beings not at all or only remotely alliedby organs in
appearancethough not in developmentclosely 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 beingsby 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 butterflythe two wings of a flyand the two wings with the
elytra of a beetle. Bivalve shells are made to open and shutbut on what
a number of patterns is the hinge constructedfrom the long row of neatly
interlocking teeth in a Nucula to the simple ligament of a Mussel! Seeds
are disseminated by their minutenessby their capsule being converted into
a light balloon-like envelopeby being embedded in pulp or fleshformed
of the most diverse partsand rendered nutritiousas well as
conspicuously colouredso as to attract and be devoured by birdsby
having hooks and grapnels of many kinds and serrated awnsso as to adhere
to the fur of quadrupedsand by being furnished with wings and plumesas
different in shape as they are elegant in structureso as to be wafted by
every breeze. I will give one other instance: for this subject of the
same end being gained by the most diversified means well deserves
attention. Some authors maintain that organic beings have been formed in
many ways for the sake of mere varietyalmost like toys in a shopbut
such a view of nature is incredible. With plants having separated sexes
and with those in whichthough hermaphroditesthe pollen does not
spontaneously fall on the stigmasome aid is necessary for their
fertilisation. With several kinds this is effected by the pollen-grains
which are light and incoherentbeing blown by the wind through mere chance
on to the stigma; and this is the simplest plan which can well be
conceived. An almost equally simplethough very different plan occurs in
many plants in which a symmetrical flower secretes a few drops of nectar
and is consequently visited by insects; and these carry the pollen from the
anthers to the stigma.

>From this simple stage we may pass through an inexhaustible number of
contrivancesall for the same purpose and effected in essentially the same
mannerbut entailing changes in every part of the flower. The nectar may
be stored in variously shaped receptacleswith the stamens and pistils
modified in many wayssometimes forming trap-like contrivancesand
sometimes capable of neatly adapted movements through irritability or
elasticity. From such structures we may advance till we come to such a
case of extraordinary adaptation as that lately described by Dr. Cruger in
the Coryanthes. This orchid has part of its labellum or lower lip hollowed
out into a great bucketinto which drops of almost pure water continually

fall from two secreting horns which stand above it; and when the bucket is
half-fullthe water overflows by a spout on one side. The basal part of
the labellum stands over the bucketand is itself hollowed out into a sort
of chamber with two lateral entrances; within this chamber there are
curious fleshy ridges. The most ingenious manif he had not witnessed
what takes placecould never have imagined what purpose all these parts
serve. But Dr. Cruger saw crowds of large humble-bees visiting the
gigantic flowers of this orchidnot in order to suck nectarbut to gnaw
off the ridges within the chamber above the bucket; in doing this they
frequently pushed each other into the bucketand their wings being thus
wetted they could not fly awaybut were compelled to crawl out through the
passage formed by the spout or overflow. Dr. Cruger saw a "continual
procession" of bees thus crawling out of their involuntary bath. The
passage is narrowand is roofed over by the columnso that a beein
forcing its way outfirst rubs its back against the viscid stigma and then
against the viscid glands of the pollen-masses. 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 flowerand are thus carried away. Dr. Cruger
sent me a flower in spirits of winewith a bee which he had killed before
it had quite crawled outwith a pollen-mass still fastened to its back.
When the beethus providedflies to another floweror to the same flower
a second timeand is pushed by its comrades into the bucket and then
crawls out by the passagethe pollen-mass necessarily comes first into
contact with the viscid stigmaand adheres to itand the flower is
fertilised. Now at last we see the full use of every part of the flower
of the water-secreting horns of the bucket half-full of waterwhich
prevents the bees from flying awayand forces them to crawl out through
the spoutand rub against the properly placed viscid pollen-masses and the
viscid stigma.

The construction of the flower in another closely allied orchidnamely
the Catasetumis widely differentthough serving the same end; and is
equally curious. Bees visit these flowerslike those of the Coryanthes
in order to gnaw the labellum; in doing this they inevitably touch a long
taperingsensitive projectionoras I have called itthe antenna. This
antennawhen touchedtransmits a sensation or vibration to a certain
membrane which is instantly ruptured; this sets free a spring by which the
pollen-mass is shot forthlike an arrowin the right directionand
adheres by its viscid extremity to the back of the bee. The pollen-mass of
the male plant (for the sexes are separate in this orchid) is thus carried
to the flower of the female plantwhere it is brought into contact with
the stigmawhich is viscid enough to break certain elastic threadsand
retain the pollenthus effecting fertilisation.

Howit may be askedin the foregoing and in innumerable other instances
can we understand the graduated scale of complexity and the multifarious
means for gaining the same end. The answer no doubt isas already
remarkedthat when two forms varywhich already differ from each other in
some slight degreethe variability will not be of the same exact nature
and consequently the results obtained through natural selection for the
same general purpose will not be the same. We should also bear in mind
that every highly developed organism has passed through many changes; and
that each modified structure tends to be inheritedso that each
modification will not readily be quite lostbut may be again and again
further altered. Hencethe structure of each part of each speciesfor
whatever purpose it may serveis the sum of many inherited changes
through which the species has passed during its successive adaptations to
changed habits and conditions of life.

Finallythenalthough in many cases it is most difficult even to
conjecture by what transitions organs could have arrived at their present
state; yetconsidering how small the proportion of living and known forms
is to the extinct and unknownI have been astonished how rarely an organ
can be namedtowards which no transitional grade is known to lead. It is

certainly truethat new organs appearing as if created for some special
purpose rarely or never appear in any being; as indeed is shown by that
oldbut somewhat exaggeratedcanon in natural history of "Natura non
facit saltum." We meet with this admission in the writings of almost every
experienced naturalist; oras Milne Edwards has well expressed itNature
is prodigal in variety, but niggard in innovation.Whyon the theory of
Creationshould there be so much variety and so little real novelty? Why
should all the parts and organs of many independent beingseach supposed
to have been separately created for its own proper place in naturebe so
commonly linked together by graduated steps? Why should not Nature take a
sudden leap from structure to structure? On the theory of natural
selectionwe can clearly understand why she should not; for natural
selection acts only by taking advantage of slight successive variations;
she can never take a great and sudden leapbut must advance by the short
and surethough slow steps.


As natural selection acts by life and deathby the survival of the
fittestand by the destruction of the less well-fitted individualsI have
sometimes felt great difficulty in understanding the origin or formation of
parts of little importance; almost as greatthough of a very different
kindas in the case of the most perfect and complex organs.

In the first placewe 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. In a former chapter I have given instances of very
trifling characterssuch as the down on fruit and the colour of its flesh
the colour of the skin and hair of quadrupedswhichfrom being correlated
with constitutional differencesor from determining the attacks of
insectsmight assuredly be acted on by natural selection. The tail of the
giraffe looks like an artificially constructed fly-flapper; and it seems at
first incredible that this could have been adapted for its present purpose
by successive slight modificationseach better and better fittedfor so
trifling an object as to drive away flies; yet we should pause before being
too positive even in this casefor 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 enemieswould be
able to range into new pastures and thus gain a great advantage. It is not
that the larger quadrupeds are actually destroyed (except in some rare
cases) by fliesbut they are incessantly harassed and their strength
reducedso that they are more subject to diseaseor not so well enabled
in a coming dearth to search for foodor to escape from beasts of prey.

Organs now of trifling importance have probably in some cases been of high
importance to an early progenitorandafter having been slowly perfected
at a former periodhave been transmitted to existing species in nearly the
same statealthough now of very slight use; but any actually injurious
deviations in their structure would of course have been checked by natural
selection. Seeing how important an organ of locomotion the tail is in most
aquatic animalsits general presence and use for many purposes in so many
land animalswhich in their lungs or modified swim-bladders betray their
aquatic originmay perhaps be thus accounted for. A well-developed tail
having been formed in an aquatic animalit might subsequently come to be
worked in for all sorts of purposesas a fly-flapperan organ of
prehensionor as an aid in turningas in the case of the dogthough the
aid in this latter respect must be slightfor the harewith hardly any
tailcan double still more quickly.

In the second placewe may easily err in attributing importance to
charactersand in believing that they have been developed through natural
selection. We must by no means overlook the effects of the definite action
of changed conditions of lifeof so-called spontaneous variationswhich

seem to depend in a quite subordinate degree on the nature of the
conditionsof the tendency to reversion to long-lost charactersof the
complex laws of growthsuch as of correlationcomprehensionof the
pressure of one part on anotheretc.and finally of sexual selectionby
which characters of use to one sex are often gained and then transmitted
more or less perfectly to the other sexthough of no use to the sex. But
structures thus indirectly gainedalthough at first of no advantage to a
speciesmay subsequently have been taken advantage of by its modified
descendantsunder new conditions of life and newly acquired habits.

If green woodpeckers alone had existedand we did not know that there were
many black and pied kindsI 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 consequently that it was a character of
importanceand had been acquired through natural selection; as it isthe
colour is probably in chief part due to sexual 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
and this contrivanceno doubtis of the highest service to the plant; but
as we see nearly similar hooks on many trees which are not climbersand
whichas there is reason to believe from the distribution of the thornbearing
species in Africa and South Americaserve as a defence against
browsing quadrupedsso the spikes on the palm may at first have been
developed for this objectand subsequently have been improved and taken
advantage of by the plantas it underwent further modification and became
a climber. The naked skin on the head of a vulture is generally considered
as a direct adaptation for wallowing in putridity; and so it may beor it
may possibly be due to the direct action of putrid matter; but we should be
very cautious in drawing any such inferencewhen we see that the skin on
the head of the clean-feeding male turkey is likewise naked. The sutures
in the skulls of young mammals have been advanced as a beautiful adaptation
for aiding parturitionand no doubt they facilitateor may be
indispensable for this act; but as sutures occur in the skulls of young
birds and reptileswhich have only to escape from a broken eggwe may
infer that this structure has arisen from the laws of growthand has been
taken advantage of in the parturition of the higher animals.

We are profoundly ignorant of the cause of each slight variation or
individual difference; and we are immediately made conscious of this by
reflecting on the differences between the breeds of our domesticated
animals in different countriesmore especially in the less civilized
countrieswhere there has been but little methodical selection. Animals
kept by savages in different countries often have to struggle for their own
subsistenceand are exposed to a certain extent to natural selectionand
individuals with slightly different constitutions would succeed best under
different climates. With cattle susceptibility to the attacks of flies is
correlated with colouras is the liability to be poisoned by certain
plants; so that even colour would be thus subjected to the action of
natural selection. Some observers are convinced that a damp climate
affects the growth of the hairand that with the hair the horns are
correlated. Mountain breeds always differ from lowland breeds; and a
mountainous country would probably affect the hind limbs from exercising
them moreand possibly even the form of the pelvis; and then by the law of
homologous variationthe front limbs and the head would probably be
affected. The shapealsoof the pelvis might affect by pressure the
shape of certain parts of the young in the womb. The laborious breathing
necessary in high regions tendsas we have good reason to believeto
increase the size of the chest; and again correlation would come into play.
The effects of lessened exercisetogether with abundant foodon the whole
organisation is probably still more importantand thisas H. von
Nathusius has lately shown in his excellent Treatiseis apparently one
chief cause of the great modification which the breeds of swine have
undergone. But we are far too ignorant to speculate on the relative
importance of the several known and unknown causes of variation; and I have

made these remarks only to show thatif we are unable to account for the
characteristic differences of our several domestic breedswhich
nevertheless are generally admitted to have arisen through ordinary
generation from one or a few parent-stockswe ought not to lay too much
stress on our ignorance of the precise cause of the slight analogous
differences between true species.


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. They believe
that many structures have been created for the sake of beautyto delight
man or the Creator (but this latter point is beyond the scope of scientific
discussion)or for the sake of mere varietya view already discussed.
Such doctrinesif truewould be absolutely fatal to my theory. I fully
admit that many structures are now of no direct use to their possessors
and may never have been of any use to their progenitors; but this does not
prove that they were formed solely for beauty or variety. No doubt the
definite action of changed conditionsand the various causes of
modificationslately specifiedhave all produced an effectprobably a
great effectindependently of any advantage thus gained. But a still more
important consideration is that the chief part of the organisation of every
living creature is due to inheritance; and consequentlythough each being
assuredly is well fitted for its place in naturemany structures have now
no very close and direct relation to present habits of life. Thuswe can
hardly believe that the webbed feet of the upland gooseor of the frigatebird
are of special use to these birds; we cannot believe that the similar
bones in the arm of the monkeyin the fore leg of the horsein the wing
of the batand in the flipper of the sealare of special use to these
animals. We may safely attribute these structures to inheritance. But
webbed feet no doubt were as useful to the progenitor of the upland goose
and of the frigate-birdas they now are to the most aquatic of living
birds. So we may believe that the progenitor of the seal did not possess a
flipperbut a foot with five toes fitted for walking or grasping; and we
may further venture to believe that the several bones in the limbs of the
monkeyhorse and batwere originally developedon the principle of
utilityprobably through the reduction of more numerous bones in the fin
of some ancient fish-like progenitor of the whole class. It is scarcely
possible to decide how much allowance ought to be made for such causes of
changeas the definite action of external conditionsso-called
spontaneous variationsand the complex laws of growth; but with these
important exceptionswe may conclude that the structure of every living
creature either now isor was formerlyof some direct or indirect use to
its possessor.

With respect to the belief that organic beings have been created beautiful
for the delight of man--a belief which it has been pronounced is subversive
of my whole theory--I may first remark that the sense of beauty obviously
depends on the nature of the mindirrespective of any real quality in the
admired object; and that the idea of what is beautifulis not innate or
unalterable. We see thisfor instancein the men of different races
admiring an entirely different standard of beauty in their women. If
beautiful objects had been created solely for man's gratificationit 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 epochand the gracefully sculptured ammonites of
the Secondary periodcreated 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 caseand in many othersis apparently wholly due to symmetry of
growth. Flowers rank among the most beautiful productions of nature; but
they have been rendered conspicuous in contrast with the green leavesand

in consequence at the same time beautifulso that they may be easily
observed by insects. 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. Several plants habitually produce two kinds of
flowers; one kind open and coloured so as to attract insects; the other
closednot coloureddestitute of nectarand never visited by insects.
Hencewe may conclude thatif insects had not been developed on the face
of the earthour plants would not have been decked with beautiful flowers
but would have produced only such poor flowers as we see on our firoak
nut and ash treeson grassesspinachdocks and nettleswhich are all
fertilised through the agency of the wind. A similar line of argument
holds good with fruits; 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. But this beauty serves merely as a guide to birds
and beastsin order that the fruit may be devoured and the matured seeds
disseminated. 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)if
it be coloured of any brilliant tintor rendered conspicuous by being
white or black.

On the other handI willingly admit that a great number of male animals
as all our most gorgeous birdssome fishesreptilesand mammalsand a
host of magnificently coloured butterflieshave been rendered beautiful
for beauty's sake. But this has been effected through sexual selection
that isby the more beautiful males having been continually preferred by
the femalesand not for the delight of man. So it is with the music of
birds. 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. When the female is as beautifully coloured as the male
which is not rarely the case with birds and butterfliesthe cause
apparently lies in the colours acquired through sexual selection having
been transmitted to both sexesinstead of to the males alone. How the
sense of beauty in its simplest form--that isthe reception of a peculiar
kind of pleasure from certain coloursforms and sounds--was first
developed in the mind of man and of the lower animalsis a very obscure
subject. The same sort of difficulty is presented if we enquire how it is
that certain flavours and odours give pleasureand others displeasure.
Habit in all these cases appears to have come to a certain extent into
play; but there must be some fundamental cause in the constitution of the
nervous system in each species.

Natural selection cannot possibly produce any modification in a species
exclusively for the good of another species; though throughout nature one
species incessantly takes advantage ofand profits by the structures of
others. But natural selection can and does often produce structures for
the direct injury of other animalsas we see in the fang of the adderand
in the ovipositor of the ichneumonby which its eggs are deposited in the
living bodies of other insects. If it could be proved that any part of the
structure of any one species had been formed for the exclusive good of
another speciesit would annihilate my theoryfor such could not have
been produced through natural selection. Although many statements may be
found in works on natural history to this effectI cannot find even one
which seems to me of any weight. It is admitted that the rattlesnake has a
poison-fang for its own defence and for the destruction of its prey; but
some authors suppose that at the same time it is furnished with a rattle
for its own injurynamelyto warn its prey. I would almost as soon
believe that the cat curls the end of its tail when preparing to springin
order to warn the doomed mouse. It is a much more probable view that the
rattlesnake uses its rattlethe cobra expands its frill and the puff-adder
swells while hissing so loudly and harshlyin order to alarm the many
birds and beasts which are known to attack even the most venomous species.
Snakes act on the same principle which makes the hen ruffle her feathers

and expand her wings when a dog approaches her chickens. But I have not
space here to enlarge on the many ways by which animals endeavour to
frighten away their enemies.

Natural selection will never produce in a being any structure more
injurious than beneficial to that beingfor natural selection acts solely
by and for the good of each. No organ will be formedas Paley has
remarkedfor the purpose of causing pain or for doing an injury to its
possessor. If a fair balance be struck between the good and evil caused by
each parteach will be found on the whole advantageous. After the lapse
of timeunder changing conditions of lifeif any part comes to be
injuriousit will be modified; or if it be not sothe being will become
extinctas myriads have become extinct.

Natural selection tends only to make each organic being as perfect asor
slightly more perfect than the other inhabitants of the same country with
which it comes into competition. And we see that this is the standard of
perfection attained under nature. The endemic productions of New Zealand
for instanceare perfectone compared with another; but they are now
rapidly yielding before the advancing legions of plants and animals
introduced from Europe. Natural selection will not produce absolute
perfectionnor do we always meetas far as we can judgewith this high
standard under nature. The correction for the aberration of light is said
by Muller not to be perfect even in that most perfect organthe human eye.
Helmholtzwhose judgment no one will disputeafter describing in the
strongest terms the wonderful powers of the human eyeadds 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 retinais
as nothing in comparison with the incongruities which we have just come
across in the domain of the sensations. One might say that nature has
taken delight in accumulating contradictions in order to remove all
foundation from the theory of a pre-existing harmony between the external
and internal worlds." If our reason leads us to admire with enthusiasm a
multitude of inimitable contrivances in naturethis same reason tells us
though we may easily err on both sidesthat some other contrivances are
less perfect. Can we consider the sting of the bee as perfectwhichwhen
used against many kinds of enemiescannot be withdrawnowing to the
backward serraturesand thus inevitably causes the death of the insect by
tearing out its viscera?

If we look at the sting of the beeas having existed in a remote
progenitoras a boring and serrated instrumentlike that in so many
members of the same great orderand that it has since been modified but
not perfected for its present purposewith the poison originally adapted
for some other objectsuch as to produce gallssince intensifiedwe 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 communityit will fulfil all the requirements of
natural selectionthough it may cause the death of some few members. If
we admire the truly wonderful power of scent by which the males of many
insects find their femalescan we admire the production for this single
purpose of thousands of droneswhich are utterly useless to the community
for any other purposeand which are ultimately slaughtered by their
industrious and sterile sisters? It may be difficultbut we ought to
admire the savage instinctive hatred of the queen-beewhich urges her to
destroy the young queensher daughtersas soon as they are bornor to
perish herself in the combat; for undoubtedly this is for the good of the
community; and maternal love or maternal hatredthough the latter
fortunately is most rareis all the same to the inexorable principles of
natural selection. If we admire the several ingenious contrivances by
which orchids and many other plants are fertilised through insect agency
can we consider as equally perfect the elaboration of dense clouds of
pollen by our fir-treesso that a few granules may be wafted by chance on
to the ovules?


We have in this chapter discussed some of the difficulties and objections
which may be urged against the theory. Many of them are serious; but I
think that in the discussion light has been thrown on several factswhich
on the belief of independent acts of creation are utterly obscure. We have
seen that species at any one period are not indefinitely variableand are
not linked together by a multitude of intermediate gradationspartly
because the process of natural selection is always very slowand at any
one time acts only on a few forms; and partly because the very process of
natural selection implies the continual supplanting and extinction of
preceding and intermediate gradations. Closely allied speciesnow living
on a continuous areamust often have been formed when the area was not
continuousand when the conditions of life did not insensibly graduate
away from one part to another. When two varieties are formed in two
districts of a continuous areaan intermediate variety will often be
formedfitted for an intermediate zone; but from reasons assignedthe
intermediate variety will usually exist in lesser numbers than the two
forms which it connects; consequently the two latterduring the course of
further modificationfrom existing in greater numberswill have a great
advantage over the less numerous intermediate varietyand will thus
generally succeed in supplanting and exterminating it.

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; that
a batfor instancecould not have been formed by natural selection from
an animal which at first only glided through the air.

We have seen that a species under new conditions of life may change its
habitsor it may have diversified habitswith some very unlike those of
its nearest congeners. Hence we can understandbearing in mind that each
organic being is trying to live wherever it can livehow it has arisen
that there are upland geese with webbed feetground woodpeckersdiving
thrushesand petrels with the habits of auks.

Although the belief that an organ so perfect as the eye could have been
formed by natural selectionis enough to stagger any one; yet in the case
of any organif we know of a long series of gradations in complexityeach
good for its possessorthen under changing conditions of lifethere is no
logical impossibility in the acquirement of any conceivable degree of
perfection through natural selection. In the cases in which we know of no
intermediate or transitional stateswe should be extremely cautious in
concluding that none can have existedfor the metamorphoses of many organs
show what wonderful changes in function are at least possible. For
instancea swim-bladder has apparently been converted into an
air-breathing lung. The same organ having performed simultaneously very
different functionsand then having been in part or in whole specialised
for one function; and two distinct organs having performed at the same time
the same functionthe one having been perfected whilst aided by the other
must often have largely facilitated transitions.

We have seen that in two beings widely remote from each other in the
natural scaleorgans serving for the same purpose and in external
appearance closely similar may have been separately and independently
formed; but when such organs are closely examinedessential differences in
their structure can almost always be detected; and this naturally follows
from the principle of natural selection. On the other handthe common
rule throughout nature is infinite diversity of structure for gaining the
same end; and this again naturally follows from the same great principle.

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 speciesthat modifications

in its structure could not have been slowly accumulated by means of natural
selection. In many other casesmodifications are probably the direct
result of the laws of variation or of growthindependently of any good
having been thus gained. But even such structures have oftenas we may
feel assuredbeen subsequently taken advantage ofand still further
modifiedfor the good of species under new conditions of life. We may
alsobelieve that a part formerly of high importance has frequently been
retained (as the tail of an aquatic animal by its terrestrial descendants)
though it has become of such small importance that it could notin its
present statehave been acquired by means of natural selection.

Natural selection can produce nothing in one species for the exclusive good
or injury of another; though it may well produce partsorgansand
excretions highly useful or even indispensableor highly injurious to
another speciesbut in all cases at the same time useful to the possessor.
In each well-stocked country natural selection acts through the competition
of the inhabitants and consequently leads to success in the battle for
lifeonly in accordance with the standard of that particular country.
Hence the inhabitants of one countrygenerally the smaller oneoften
yield to the inhabitants of another and generally the larger country. For
in the larger country there will have existed more individualsand more
diversified formsand the competition will have been severerand thus the
standard of perfection will have been rendered higher. Natural selection
will not necessarily lead to absolute perfection; noras far as we can
judge by our limited facultiescan absolute perfection be everywhere

On the theory of natural selection we can clearly understand the full
meaning of that old canon in natural historyNatura non facit saltum.
This canonif we look to the present inhabitants alone of the worldis
not strictly correct; but if we include all those of past timeswhether
known or unknownit must on this theory be strictly true.

It is generally acknowledged that all organic beings have been formed on
two great laws--Unity of Typeand the Conditions of Existence. By unity
of type is meant that fundamental agreement in structure which we see in
organic beings of the same classand which is quite independent of their
habits of life. On my theoryunity of type is explained by unity of
descent. The expression of conditions of existenceso often insisted on
by the illustrious Cuvieris fully embraced by the principle of natural
selection. For natural selection acts by either now adapting the varying
parts of each being to its organic and inorganic conditions of life; or by
having adapted them during past periods of time: the adaptations being
aided in many cases by the increased use or disuse of partsbeing affected
by the direct action of external conditions of lifeand subjected in all
cases to the several laws of growth and variation. Hencein factthe law
of the Conditions of Existence is the higher law; as it includesthrough
the inheritance of former variations and adaptationsthat of Unity of



Longevity -- Modifications not necessarily simultaneous -- Modifications
apparently of no direct service -- Progressive development -- Characters of
small functional importancethe 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 classdeveloped from one
and the same source -- Reasons for disbelieving in great and abrupt

I will devote this chapter to the consideration of various miscellaneous
objections which have been advanced against my viewsas some of the
previous discussions may thus be made clearer; but it would be useless to
discuss all of themas many have been made by writers who have not taken
the trouble to understand the subject. Thus a distinguished German
naturalist has asserted that the weakest part of my theory isthat I
consider all organic beings as imperfect: what I have really said isthat
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. Nor can organic beingseven if they were at any one time
perfectly adapted to their conditions of lifehave remained sowhen their
conditions changedunless they themselves likewise changed; and no one
will dispute that the physical conditions of each countryas well as the
number and kinds of its inhabitantshave undergone many mutations.

A critic has lately insistedwith some parade of mathematical accuracy
that longevity is a great advantage to all speciesso that he who believes
in natural selection "must arrange his genealogical tree" in such a manner
that all the descendants have longer lives than their progenitors! Cannot
our critics conceive that a biennial plant or one of the lower animals
might range into a cold climate and perish there every winter; and yet
owing to advantages gained through natural selectionsurvive from year to
year by means of its seeds or ova? Mr. E. Ray Lankester has recently
discussed this subjectand he concludesas far as its extreme complexity
allows him to form a judgmentthat longevity is generally related to the
standard of each species in the scale of organisationas well as to the
amount of expenditure in reproduction and in general activity. And these
conditions haveit is probablebeen largely determined through natural

It has been argued thatas none of the animals and plants of Egyptof
which we know anythinghave changed during the last three or four thousand
yearsso probably have none in any part of the world. Butas Mr. G.H.
Lewes has remarkedthis line of argument proves too muchfor the ancient
domestic races figured on the Egyptian monumentsor embalmedare closely
similar or even identical with those now living; yet all naturalists admit
that such races have been produced through the modification of their
original types. The many animals which have remained unchanged since the
commencement of the glacial periodwould have been an incomparably
stronger casefor these have been exposed to great changes of climate and
have migrated over great distances; whereasin Egyptduring the last
several thousand yearsthe conditions of lifeas far as we knowhave
remained absolutely uniform. The fact of little or no modification having
been effected since the glacial periodwould have been of some avail
against those who believe in an innate and necessary law of development
but is powerless against the doctrine of natural selection or the survival
of the fittestwhich implies that when variations or individual
differences of a beneficial nature happen to arisethese will be
preserved; but this will be effected only under certain favourable

The celebrated palaeontologistBronnat the close of his German
translation of this workasks howon the principle of natural selection
can a variety live side by side with the parent species? If both have
become fitted for slightly different habits of life or conditionsthey
might live together; and if we lay on one side polymorphic speciesin
which the variability seems to be of a peculiar natureand all mere
temporary variationssuch as sizealbinismetc.the more permanent
varieties are generally foundas far as I can discoverinhabiting
distinct stationssuch as high land or low landdry or moist districts.
Moreoverin the case of animals which wander much about and cross freely
their varieties seem to be generally confined to distinct regions.

Bronn also insists that distinct species never differ from each other in
single charactersbut in many parts; and he askshow 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. The most striking modificationsexcellently adapted for some
purposemightas was formerly remarkedbe acquired by successive
variationsif slightfirst in one part and then in another; and as they
would be transmitted all togetherthey would appear to us as if they had
been simultaneously developed. The best answerhoweverto the above
objection is afforded by those domestic races which have been modified
chiefly through man's power of selectionfor some special purpose. Look
at the race and dray-horseor at the greyhound and mastiff. Their whole
framesand even their mental characteristicshave been modified; 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
changesbut first one part and then another slightly modified and
improved. 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 partwhether it be the
flowerfruitor leaveshas been greatly changedalmost all the other
parts have been slightly modified. This may be attributed partly to the
principle of correlated growthand partly to so-called spontaneous

A much more serious objection has been urged by Bronnand recently by
Brocanamelythat many characters appear to be of no service whatever to
their possessorsand therefore cannot have been influenced through natural
selection. 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
animalsand a multitude of analogous cases. With respect to plantsthis
subject has been discussed by Nageli in an admirable essay. He admits that
natural selection has effected muchbut he insists that the families of
plants differ chiefly from each other in morphological characterswhich
appear to be quite unimportant for the welfare of the species. He
consequently believes in an innate tendency towards progressive and more
perfect development. He specifies the arrangement of the cells in the
tissuesand of the leaves on the axisas cases in which natural selection
could not have acted. To these may be added the numerical divisions in the
parts of the flowerthe position of the ovulesthe shape of the seed
when not of any use for disseminationetc.

There is much force in the above objection. Neverthelesswe oughtin the
first placeto be extremely cautious in pretending to decide what
structures now areor have formerly beenof use to each species. In the
second placeit should always be borne in mind that when one part is
modifiedso will be other partsthrough certain dimly seen causessuch
as an increased or diminished flow of nutriment to a partmutual pressure
an early developed part affecting one subsequently developedand so forth
--as well as through other causes which lead to the many mysterious cases
of correlationwhich we do not in the least understand. These agencies
may be all grouped togetherfor the sake of brevityunder the expression
of the laws of growth. In the third placewe have to allow for the direct
and definite action of changed conditions of lifeand for so-called
spontaneous variationsin which the nature of the conditions apparently
plays a quite subordinate part. Bud-variationssuch as the appearance of
a moss-rose on a common roseor of a nectarine on a peach-treeoffer good
instances of spontaneous variations; but even in these casesif we bear in
mind the power of a minute drop of poison in producing complex gallswe
ought not to feel too sure that the above variations are not the effect of
some local change in the nature of the sapdue to some change in the
conditions. There must be some efficient cause for each slight individual
differenceas well as for more strongly marked variations which

occasionally arise; and if the unknown cause were to act persistentlyit
is almost certain that all the individuals of the species would be
similarly modified.

In the earlier editions of this work I underratedas it now seems
probablethe frequency and importance of modifications due to spontaneous
variability. 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. I can no more believe in this than that the well-adapted
form of a race-horse or greyhoundwhich before the principle of selection
by man was well understoodexcited so much surprise in the minds of the
older naturalistscan thus be explained.

It may be worth while to illustrate some of the foregoing remarks. With
respect to the assumed inutility of various parts and organsit is hardly
necessary to observe that even in the higher and best-known animals many
structures existwhich are so highly developed that no one doubts that
they are of importanceyet their use has not beenor has only recently
beenascertained. As Bronn gives the length of the ears and tail in the
several species of mice as instancesthough trifling onesof differences
in structure which can be of no special useI may mention thataccording
to Dr. Schoblthe external ears of the common mouse are supplied in an
extraordinary manner with nervesso that they no doubt serve as tactile
organs; hence the length of the ears can hardly be quite unimportant. We
shallalsopresently see that the tail is a highly useful prehensile
organ to some of the species; and its use would be much influence by its

With respect to plantsto which on account of Nageli's essay I shall
confine myself in the following remarksit will be admitted that the
flowers of the orchids present a multitude of curious structureswhich a
few years ago would have been considered as mere morphological differences
without any special function; but they are now known to be of the highest
importance for the fertilisation of the species through the aid of insects
and have probably been gained through natural selection. No one until
lately would have imagined that in dimorphic and trimorphic plants the
different lengths of the stamens and pistilsand their arrangementcould
have been of any servicebut now we know this to be the case.

In certain whole groups of plants the ovules stand erectand in others
they are suspended; and within the same ovarium of some few plantsone
ovule holds the former and a second ovule the latter position. These
positions seem at first purely morphologicalor of no physiological
signification; but Dr. Hooker informs me that within the same ovarium the
upper ovules alone in some casesand 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. If sothe position of the
ovuleseven when one is erect and the other suspended within the same
ovariumwould follow the selection of any slight deviations in position
which favoured their fertilisationand the production of seed.

Several plants belonging to distinct orders habitually produce flowers of
two kinds--the one openof the ordinary structurethe other closed and
imperfect. These two kinds of flowers sometimes differ wonderfully in
structureyet may be seen to graduate into each other on the same plant.
The ordinary and open flowers can be intercrossed; and the benefits which
certainly are derived from this process are thus secured. The closed and
imperfect flowers arehowevermanifestly of high importanceas they
yield with the utmost safety a large stock of seedwith the expenditure of
wonderfully little pollen. The two kinds of flowers often differ muchas
just statedin structure. The petals in the imperfect flowers almost
always consist of mere rudimentsand the pollen-grains are reduced in
diameter. In Ononis columnae five of the alternate stamens are
rudimentary; and in some species of Viola three stamens are in this state

two retaining their proper functionbut being of very small size. In six
out of thirty of the closed flowers in an Indian violet (name unknownfor
the plants have never produced with me perfect flowers)the sepals are
reduced from the normal number of five to three. In one section of the
Malpighiaceae the closed flowersaccording to A. de Jussieuare still
further modifiedfor the five stamens which stand opposite to the sepals
are all aborteda sixth stamen standing opposite to a petal being alone
developed; and this stamen is not present in the ordinary flowers of this
species; the style is aborted; and the ovaria are reduced from three to
two. Now although natural selection may well have had the power to prevent
some of the flowers from expandingand to reduce the amount of pollen
when rendered by the closure of the flowers superfluousyet hardly any of
the above special modifications can have been thus determinedbut must
have followed from the laws of growthincluding the functional inactivity
of partsduring the progress of the reduction of the pollen and the
closure of the flowers.

It is so necessary to appreciate the important effects of the laws of
growththat I will give some additional cases of another kindnamely of
differences in the same part or organdue to differences in relative
position on the same plant. In the Spanish chestnutand in certain firtrees
the angles of divergence of the leaves differaccording to Schacht
in the nearly horizontal and in the upright branches. In the common rue
and some other plantsone flowerusually the central or terminal one
opens firstand has five sepals and petalsand five divisions to the
ovarium; while all the other flowers on the plant are tetramerous. In the
British Adoxa the uppermost flower generally has two calyx-lobes with the
other organs tetramerouswhile the surrounding flowers generally have
three calyx-lobes with the other organs pentamerous. In many Compositae
and Umbelliferae (and in some other plants) the circumferential flowers
have their corollas much more developed than those of the centre; and this
seems often connected with the abortion of the reproductive organs. It is
a more curious factpreviously referred tothat the achenes or seeds of
the circumference and centre sometimes differ greatly in formcolour and
other characters. In Carthamus and some other Compositae the central
achenes alone are furnished with a pappus; and in Hyoseris the same head
yields achenes of three different forms. In certain Umbelliferae the
exterior seedsaccording to Tauschare orthospermousand the central one
coelospermousand this is a character which was considered by De Candolle
to be in other species of the highest systematic importance. Professor
Braun mentions a Fumariaceous genusin which the flowers in the lower part
of the spike bear ovalribbedone-seeded nutlets; and in the upper part
of the spikelanceolatetwo-valved and two-seeded siliques. In these
several caseswith the exception of that of the well-developed rayflorets
which are of service in making the flowers conspicuous to insects
natural selection cannotas far as we can judgehave come into playor
only in a quite subordinate manner. All these modifications follow from
the relative position and inter-action of the parts; 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 conditionas are the flowers
and leaves in certain positionsall would have been modified in the same

In numerous other cases we find modifications of structurewhich are
considered by botanists to be generally of a highly important nature
affecting only some of the flowers on the same plantor occurring on
distinct plantswhich grow close together under the same conditions. As
these variations seem of no special use to the plantsthey cannot have
been influenced by natural selection. Of their cause we are quite
ignorant; we cannot even attribute themas in the last class of casesto
any proximate agencysuch as relative position. I will give only a few
instances. It is so common to observe on the same plantflowers
indifferently tetramerouspentamerousetc.that I need not give
examples; but as numerical variations are comparatively rare when the parts

are fewI may mention thataccording to De Candollethe flowers of
Papaver bracteatum offer either two sepals with four petals (which is the
common type with poppies)or three sepals with six petals. The manner in
which the petals are folded in the bud is in most groups a very constant
morphological character; but Professor Asa Gray states that with some
species of Mimulusthe aestivation is almost as frequently that of the
Rhinanthideae as of the Antirrhinideaeto which latter tribe the genus
belongs. Aug. St. Hilaire gives the following cases: the genus
Zanthoxylon belongs to a division of the Rutaceae with a single ovarybut
in some species flowers may be found on the same plantand even in the
same paniclewith either one or two ovaries. In Helianthemum the capsule
has been described as unilocular or tri-locular; and in H. mutabileUne
lame PLUS OU MOINS LARGE, s'etend entre le pericarpe et le placenta.In
the flowers of Saponaria officinalis Dr. Masters has observed instances of
both marginal and free central placentation. LastlySt. Hilaire found
towards the southern extreme of the range of Gomphia oleaeformis two forms
which he did not at first doubt were distinct speciesbut he subsequently
saw them growing on the same bush; and he then addsVoila donc dans un
meme individu des loges et un style qui se rattachent tantot a un axe
verticale et tantot a un gynobase.

We thus see that with plants many morphological changes may be attributed
to the laws of growth and the inter-action of partsindependently of
natural selection. But with respect to Nageli's doctrine of an innate
tendency towards perfection or progressive developmentcan it be said in
the case of these strongly pronounced variationsthat the plants have been
caught in the act of progressing towards a higher state of development? On
the contraryI should infer from the mere fact of the parts in question
differing or varying greatly on the same plantthat such modifications
were of extremely small importance to the plants themselvesof whatever
importance they may generally be to us for our classifications. The
acquisition of a useless part can hardly be said to raise an organism in
the natural scale; and in the case of the imperfectclosed flowersabove
describedif any new principle has to be invokedit must be one of
retrogression rather than of progression; and so it must be with many
parasitic and degraded animals. We are ignorant of the exciting cause of
the above specified modifications; but if the unknown cause were to act
almost uniformly for a length of timewe may infer that the result would
be almost uniform; and in this case all the individuals of the species
would be modified in the same manner.

>From the fact of the above characters being unimportant for the welfare of
the speciesany slight variations which occurred in them would not have
been accumulated and augmented through natural selection. A structure
which has been developed through long-continued selectionwhen it ceases
to be of service to a speciesgenerally becomes variableas we see with
rudimentary organs; for it will no longer be regulated by this same power
of selection. But whenfrom the nature of the organism and of the
conditionsmodifications have been induced which are unimportant for the
welfare of the speciesthey may beand apparently often have been
transmitted in nearly the same state to numerousotherwise modified
descendants. It cannot have been of much importance to the greater number
of mammalsbirdsor reptileswhether they were clothed with hair
feathers or scales; yet hair has been transmitted to almost all mammals
feathers to all birdsand scales to all true reptiles. A structure
whatever it may bewhich is common to many allied formsis ranked by us
as of high systematic importanceand consequently is often assumed to be
of high vital importance to the species. Thusas I am inclined to
believemorphological differenceswhich we consider as important--such as
the arrangement of the leavesthe divisions of the flower or of the
ovariumthe position of the ovulesetc.first appeared in many cases as
fluctuating variationswhich sooner or later became constant through the
nature of the organism and of the surrounding conditionsas well as
through the intercrossing of distinct individualsbut not through natural

selection; for as these morphological characters do not affect the welfare
of the speciesany slight deviations in them could not have been governed
or accumulated through this latter agency. It is a strange result which we
thus arrive atnamelythat characters of slight vital importance to the
speciesare the most important to the systematist; butas we shall
hereafter see when we treat of the genetic principle of classification
this is by no means so paradoxical as it may at first appear.

Although we have no good evidence of the existence in organic beings of an
innate tendency towards progressive developmentyet this necessarily
followsas I have attempted to show in the fourth chapterthrough the
continued action of natural selection. For the best definition which has
ever been given of a high standard of organisationis the degree to which
the parts have been specialised or differentiated; and natural selection
tends towards this endinasmuch as the parts are thus enabled to perform
their functions more efficiently.

A distinguished zoologistMr. St. George Mivarthas recently collected
all the objections which have ever been advanced by myself and others
against the theory of natural selectionas propounded by Mr. Wallace and
myselfand has illustrated them with admirable art and force. When thus
marshalledthey make a formidable array; and as it forms no part of Mr.
Mivart's plan to give the various facts and considerations opposed to his
conclusionsno slight effort of reason and memory is left to the reader
who may wish to weigh the evidence on both sides. When discussing special
casesMr. Mivart passes over the effects of the increased use and disuse
of partswhich I have always maintained to be highly importantand have
treated in my "Variation under Domestication" at greater length thanas I
believeany other writer. He likewise often assumes that I attribute
nothing to variationindependently of natural selectionwhereas in the
work just referred to I have collected a greater number of well-established
cases than can be found in any other work known to me. My judgment may not
be trustworthybut after reading with care Mr. Mivart's bookand
comparing each section with what I have said on the same headI never
before felt so strongly convinced of the general truth of the conclusions
here arrived atsubjectof coursein so intricate a subjectto much
partial error.

All Mr. Mivart's objections will beor have beenconsidered in the
present volume. The one new point which appears to have struck many
readers isThat natural selection is incompetent to account for the
incipient stages of useful structures.This subject is intimately
connected with that of the gradation of the charactersoften accompanied
by a change of functionfor instancethe conversion of a swim-bladder
into lungspoints which were discussed in the last chapter under two
headings. NeverthelessI will here consider in some detail several of the
cases advanced by Mr. Mivartselecting those which are the most
illustrativeas want of space prevents me from considering all.

The giraffeby its lofty staturemuch elongated neckfore legshead and
tonguehas its whole frame beautifully adapted for browsing on the higher
branches of trees. It can thus obtain food beyond the reach of the other
Ungulata or hoofed animals inhabiting the same country; and this must be a
great advantage to it during dearths. The Niata cattle in South America
show us how small a difference in structure may makeduring such periods
a great difference in preserving an animal's life. These cattle can browse
as well as others on grassbut from the projection of the lower jaw they
cannotduring the often recurrent droughtsbrowse on the twigs of trees which food the common cattle and horses are then driven; so
that at these times the Niatas perishif not fed by their owners. Before
coming to Mr. Mivart's objectionsit may be well to explain once again how
natural selection will act in all ordinary cases. Man has modified some of
his animalswithout necessarily having attended to special points of
structureby simply preserving and breeding from the fleetest individuals

as with the race-horse and greyhoundor as with the game-cockby breeding
from the victorious birds. So under nature with the nascent giraffethe
individuals which were the highest browsers and were able during dearths to
reach even an inch or two above the otherswill often have been preserved;
for they will have roamed over the whole country in search of food. 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 historyin
which careful measurements are given. These slight proportional
differencesdue to the laws of growth and variationare not of the
slightest use or importance to most species. But it will have been
otherwise with the nascent giraffeconsidering its probable habits of
life; for those individuals which had some one part or several parts of
their bodies rather more elongated than usualwould generally have
survived. These will have intercrossed and left offspringeither
inheriting the same bodily peculiaritiesor with a tendency to vary again
in the same manner; while the individuals less favoured in the same
respects will have been the most liable to perish.

We here see that there is no need to separate single pairsas man does
when he methodically improves a breed: natural selection will preserve and
thus separate all the superior individualsallowing them freely to
intercrossand will destroy all the inferior individuals. By this process
long-continuedwhich exactly corresponds with what I have called
unconscious selection by mancombinedno doubtin a most important
manner with the inherited effects of the increased use of partsit seems
to me almost certain that an ordinary hoofed quadruped might be converted
into a giraffe.

To this conclusion Mr. Mivart brings forward two objections. One is that
the increased size of the body would obviously require an increased supply
of foodand he considers it as "very problematical whether the
disadvantages thence arising would notin times of scarcitymore than
counterbalance the advantages." But as the giraffe does actually exist in
large numbers in Africaand as some of the largest antelopes in the world
taller than an oxabound therewhy should we doubt thatas far as size
is concernedintermediate gradations could formerly have existed there
subjected as now to severe dearths. Assuredly the being able to reachat
each stage of increased sizeto a supply of foodleft untouched by the
other hoofed quadrupeds of the countrywould have been of some advantage
to the nascent giraffe. Nor must we overlook the factthat increased bulk
would act as a protection against almost all beasts of prey excepting the
lion; and against this animalits tall neck--and the taller the better-would
as Mr. Chauncey Wright has remarkedserve as a watch-tower. It is
from this causeas Sir S. Baker remarksthat no animal is more difficult
to stalk than the giraffe. This animal also uses its long neck as a means
of offence or defenceby violently swinging its head armed with stump-like
horns. The preservation of each species can rarely be determined by any
one advantagebut by the union of allgreat and small.

Mr. Mivart then asks (and this is his second objection)if natural
selection be so potentand if high browsing be so great an advantagewhy
has not any other hoofed quadruped acquired a long neck and lofty stature
besides the giraffeandin a lesser degreethe camelguanaco and
macrauchenia? Oragainwhy has not any member of the group acquired a
long proboscis? With respect to South Africawhich was formerly inhabited
by numerous herds of the giraffethe answer is not difficultand can best
be given by an illustration. In every meadow in Englandin which trees
growwe see the lower branches trimmed or planed to an exact level by the
browsing of the horses or cattle; and what advantage would it befor
instanceto sheepif kept thereto acquire slightly longer necks? In
every district some one kind of animal will almost certainly be able to
browse higher than the others; and it is almost equally certain that this
one kind alone could have its neck elongated for this purposethrough
natural selection and the effects of increased use. In South Africa the

competition for browsing on the higher branches of the acacias and other
trees must be between giraffe and giraffeand not with the other ungulate

Whyin other quarters of the worldvarious animals belonging to this same
order have not acquired either an elongated neck or a probosciscannot 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. We are ignorant with respect
to the conditions which determine the numbers and range of each species
and we cannot even conjecture what changes of structure would be favourable
to its increase in some new country. We canhoweversee in a general
manner that various causes might have interfered with the development of a
long neck or proboscis. To reach the foliage at a considerable height
(without climbingfor which hoofed animals are singularly ill-constructed)
implies greatly increased bulk of body; and we know that some areas support
singularly few large quadrupedsfor instance South Americathough it is
so luxuriantwhile South Africa abounds with them to an unparalleled
degree. Why this should be so we do not know; nor why the later tertiary
periods should have been much more favourable for their existence than the
present time. Whatever the causes may have beenwe 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.

In order that an animal should acquire some structure specially and largely
developedit is almost indispensable that several other parts should be
modified and coadapted. Although every part of the body varies slightly
it does not follow that the necessary parts should always vary in the right
direction and to the right degree. With the different species of our
domesticated animals we know that the parts vary in a different manner and
degreeand that some species are much more variable than others. Even if
the fitting variations did ariseit 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. For instanceif the number of individuals
existing in a country is determined chiefly through destruction by beasts
of prey--by external or internal parasitesetc.--as seems often to be the
casethen natural selection will be able to do littleor will be greatly
retardedin modifying any particular structure for obtaining food.
Lastlynatural selection is a slow processand the same favourable
conditions must long endure in order that any marked effect should thus be
produced. Except by assigning such general and vague reasonswe cannot
explain whyin many quarters of the worldhoofed quadrupeds have not
acquired much elongated necks or other means for browsing on the higher
branches of trees.

Objections of the same nature as the foregoing have been advanced by many
writers. In each case various causesbesides the general ones just
indicatedhave probably interfered with the acquisition through natural
selection of structureswhich it is thought would be beneficial to certain
species. One writer askswhy 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. Oceanic islands are inhabited by bats and
sealsbut by no terrestrial mammals; yet as some of these bats are
peculiar speciesthey must have long inhabited their present homes.
Therefore Sir C. Lyell asksand assigns certain reasons in answerwhy
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 sizeand bats into
terrestrial insectivorous animals; for the former there would be no prey;
for the bats ground-insects would serve as foodbut these would already be
largely preyed on by the reptiles or birdswhich first colonise and abound
on most oceanic islands. Gradations of structurewith each stage
beneficial to a changing specieswill be favoured only under certain

peculiar conditions. A strictly terrestrial animalby occasionally
hunting for food in shallow waterthen in streams or lakesmight at last
be converted into an animal so thoroughly aquatic as to brave the open
ocean. But seals would not find on oceanic islands the conditions
favourable to their gradual reconversion into a terrestrial form. Batsas
formerly shownprobably acquired their wings by at first gliding through
the air from tree to treelike the so-called flying squirrelsfor the
sake of escaping from their enemiesor for avoiding falls; but when the
power of true flight had once been acquiredit would never be reconverted
backat least for the above purposesinto the less efficient power of
gliding through the air. Batsmightindeedlike many birdshave had
their wings greatly reduced in sizeor completely lostthrough disuse;
but in this case it would be necessary that they should first have acquired
the power of running quickly on the groundby the aid of their hind legs
aloneso as to compete with birds or other ground animals; and for such a
change a bat seems singularly ill-fitted. These conjectural remarks have
been made merely to show that a transition of structurewith each step
beneficialis a highly complex affair; and that there is nothing strange
in a transition not having occurred in any particular case.

Lastlymore than one writer has asked why have some animals had their
mental powers more highly developed than othersas such development would
be advantageous to all? Why have not apes acquired the intellectual powers
of man? Various causes could be assigned; but as they are conjecturaland
their relative probability cannot be weighedit would be useless to give
them. A definite answer to the latter question ought not to be expected
seeing that no one can solve the simpler problemwhyof two races of
savagesone has risen higher in the scale of civilisation than the other;
and this apparently implies increased brain power.

We will return to Mr. Mivart's other objections. Insects often resemble
for the sake of protection various objectssuch as green or decayed
leavesdead twigsbits of lichenflowersspinesexcrement of birds
and living insects; but to this latter point I shall hereafter recur. The
resemblance is often wonderfully closeand is not confined to colourbut
extends to formand even to the manner in which the insects hold
themselves. The caterpillars which project motionless like dead twigs from
the bushes on which they feedoffer an excellent instance of a resemblance
of this kind. The cases of the imitation of such objects as the excrement
of birdsare rare and exceptional. On this headMr. Mivart remarksAs,
according to Mr. Darwin's theory, there is a constant tendency to
indefinite variation, and as the minute incipient variations will be in ALL
DIRECTIONS, they must tend to neutralize each other, and at first to form
such unstable modifications that it is difficult, if not impossible, to see
how such indefinite oscillations of infinitesimal beginnings can ever build
up a sufficiently appreciable resemblance to a leaf, bamboo, or other
object, for natural selection to seize upon and perpetuate.

But in all the foregoing cases the insects in their original state no doubt
presented some rude and accidental resemblance to an object commonly found
in the stations frequented by them. Nor is this at all improbable
considering the almost infinite number of surrounding objects and the
diversity in form and colour of the hosts of insects which exist. As some
rude resemblance is necessary for the first startwe can understand how it
is that the larger and higher animals do not (with the exceptionas far as
I knowof one fish) resemble for the sake of protection special objects
but only the surface which commonly surrounds themand this chiefly in
colour. Assuming that an insect originally happened to resemble in some
degree a dead twig or a decayed leafand that it varied slightly in many
waysthen all the variations which rendered the insect at all more like
any such objectand thus favoured its escapewould be preservedwhile
other variations would be neglected and ultimately lost; orif they
rendered the insect at all less like the imitated objectthey would be
eliminated. There would indeed be force in Mr. Mivart's objectionif we

were to attempt to account for the above resemblancesindependently of
natural selectionthrough mere fluctuating variability; but as the case
stands there is none.

Nor can I see any force in Mr. Mivart's difficulty with respect to "the
last touches of perfection in the mimicry;" as in the case given by Mr.
Wallaceof a walking-stick insect (Ceroxylus laceratus)which resembles
a stick grown over by a creeping moss or jungermannia.So close was this
resemblancethat a native Dyak maintained that the foliaceous excrescences
were really moss. Insects are preyed on by birds and other enemies whose
sight is probably sharper than oursand every grade in resemblance which
aided an insect to escape notice or detectionwould tend towards its
preservation; and the more perfect the resemblance so much the better for
the insect. Considering the nature of the differences between the species
in the group which includes the above Ceroxylusthere is nothing
improbable in this insect having varied in the irregularities on its
surfaceand in these having become more or less green-coloured; for in
every group the characters which differ in the several species are the most
apt to varywhile the generic charactersor those common to all the
speciesare the most constant.

The Greenland whale is one of the most wonderful animals in the worldand
the baleenor whaleboneone of its greatest peculiarities. The baleen
consists of a rowon each side of the upper jawof about 300 plates or
laminaewhich stand close together transversely to the longer axis of the
mouth. Within the main row there are some subsidiary rows. The
extremities and inner margins of all the plates are frayed into stiff
bristleswhich clothe the whole gigantic palateand serve to strain or
sift the waterand thus to secure the minute prey on which these great
animals subsist. The middle and longest lamina in the Greenland whale is
tentwelveor even fifteen feet in length; but in the different species
of Cetaceans there are gradations in length; the middle lamina being in one
speciesaccording to Scoresbyfour feetin another threein another
eighteen inchesand in the Balaenoptera rostrata only about nine inches in
length. The quality of the whalebone also differs in the different

With respect to the baleenMr. Mivart remarks that if it "had once
attained such a size and development as to be at all usefulthen its
preservation and augmentation within serviceable limits would be promoted
by natural selection alone. But how to obtain the beginning of such useful
development?" In answerit may be askedwhy should not the early
progenitors of the whales with baleen have possessed a mouth constructed
something like the lamellated beak of a duck? Duckslike whalessubsist
by sifting the mud and water; and the family has sometimes been called
Criblatoresor sifters. 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. I wish only to show that this is not incredibleand
that the immense plates of baleen in the Greenland whale might have been
developed from such lamellae by finely graduated stepseach of service to
its possessor.

The beak of a shoveller-duck (Spatula clypeata) is a more beautiful and
complex structure than the mouth of a whale. The upper mandible is
furnished on each side (in the specimen examined by me) with a row or comb
formed of 188 thinelastic lamellaeobliquely bevelled so as to be
pointedand placed transversely to the longer axis of the mouth. They
arise from the palateand are attached by flexible membrane to the sides
of the mandible. Those standing towards the middle are the longestbeing
about one-third of an inch in lengthand they project fourteen onehundredths
of an inch beneath the edge. At their bases there is a short
subsidiary row of obliquely transverse lamellae. In these several respects
they resemble the plates of baleen in the mouth of a whale. But towards
the extremity of the beak they differ muchas they project inwardinstead

of straight downward. The entire head of the shovellerthough
incomparably less bulkyis about one-eighteenth of the length of the head
of a moderately large Balaenoptera rostratain which species the baleen is
only nine inches long; so that if we were to make the head of the shoveller
as long as that of the Balaenopterathe lamellae would be six inches in
lengththat istwo-thirds of the length of the baleen in this species of
whale. The lower mandible of the shoveller-duck is furnished with lamellae
of equal length with these abovebut finer; and in being thus furnished it
differs conspicuously from the lower jaw of a whalewhich is destitute of
baleen. On the other handthe extremities of these lower lamellae are
frayed into fine bristly pointsso that they thus curiously resemble the
plates of baleen. In the genus Priona member of the distinct family of
the Petrelsthe upper mandible alone is furnished with lamellaewhich are
well developed and project beneath the margin; so that the beak of this
bird resembles in this respect the mouth of a whale.

>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.
Salvin)without any great breakas far as fitness for sifting is
concernedthrough the beak of the Merganetta armataand in some respects
through that of the Aix sponsato the beak of the common duck. In this
latter species the lamellae are much coarser than in the shovellerand are
firmly attached to the sides of the mandible; they are only about fifty in
number on each sideand do not project at all beneath the margin. They
are square-toppedand are edged with translucenthardish tissueas if
for crushing food. The edges of the lower mandible are crossed by numerous
fine ridgeswhich project very little. Although the beak is thus very
inferior as a sifter to that of a shovelleryet this birdas every one
knowsconstantly uses it for this purpose. There are other speciesas I
hear from Mr. Salvinin which the lamellae are considerably less developed
than in the common duck; but I do not know whether they use their beaks for
sifting the water.

Turning to another group of the same family. In the Egyptian goose
(Chenalopex) the beak closely resembles that of the common duck; but the
lamellae are not so numerousnor so distinct from each othernor do they
project so much inward; yet this gooseas I am informed by Mr. E.
Bartlettuses its bill like a duck by throwing the water out at the
corners.Its chief foodhoweveris grasswhich it crops like the
common goose. In this latter bird the lamellae of the upper mandible are
much coarser than in the common duckalmost confluentabout twenty-seven
in number on each sideand terminating upward in teeth-like knobs. The
palate is also covered with hard rounded knobs. The edges of the lower
mandible are serrated with teeth much more prominentcoarser and sharper
than in the duck. The common goose does not sift the waterbut uses its
beak exclusively for tearing or cutting herbagefor which purpose it is so
well fitted that it can crop grass closer than almost any other animal.
There are other species of geeseas I hear from Mr. Bartlettin which the
lamellae are less developed than in the common goose.

We thus see that a member of the duck familywith a beak constructed like
that of a common goose and adapted solely for grazingor even a member
with a beak having less well-developed lamellaemight be converted by
small changes into a species like the Egyptian goose--this into one like
the common duck--andlastlyinto one like the shovellerprovided with a
beak almost exclusively adapted for sifting the water; for this bird could
hardly use any part of its beakexcept the hooked tipfor seizing or
tearing solid food. The beak of a gooseas I may addmight also be
converted by small changes into one provided with prominentrecurved
teethlike those of the Merganser (a member of the same family)serving
for the widely different purpose of securing live fish.

Returning to the whales. The Hyperoodon bidens is destitute of true teeth
in an efficient conditionbut its palate is roughenedaccording to

Lacepedewith small unequalhard points of horn. There istherefore
nothing improbable in supposing that some early Cetacean form was provided
with similar points of horn on the palatebut rather more regularly
placedand whichlike the knobs on the beak of the gooseaided it in
seizing or tearing its food. If soit 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 goosein which
case they would have been used both for seizing objects and for sifting the
water; then into lamellae like those of the domestic duck; and so onward
until they became as well constructed as those of the shovellerin which
case they would have served exclusively as a sifting apparatus. From this
stagein which the lamellae would be two-thirds of the length of the
plates of baleen in the Balaenoptera rostratagradationswhich may be
observed in still-existing Cetaceanslead us onward to the enormous plates
of baleen in the Greenland whale. Nor is there the least reason to doubt
that each step in this scale might have been as serviceable to certain
ancient Cetaceanswith the functions of the parts slowly changing during
the progress of developmentas are the gradations in the beaks of the
different existing members of the duck-family. We should bear in mind that
each species of duck is subjected to a severe struggle for existenceand
that the structure of every part of its frame must be well adapted to its
conditions of life.

The Pleuronectidaeor Flat-fishare remarkable for their asymmetrical
bodies. They rest on one side--in the greater number of species on the
leftbut in some on the right side; and occasionally reversed adult
specimens occur. The loweror resting-surfaceresembles at first sight
the ventral surface of an ordinary fish; it is of a white colourless
developed in many ways than the upper sidewith the lateral fins often of
smaller size. But the eyes offer the most remarkable peculiarity; for they
are both placed on the upper side of the head. During early youth
howeverthey stand opposite to each otherand the whole body is then
symmetricalwith both sides equally coloured. Soon the eye proper to the
lower side begins to glide slowly round the head to the upper side; but
does not pass right through the skullas was formerly thought to be the
case. It is obvious that unless the lower eye did thus travel roundit
could not be used by the fish while lying in its habitual position on one
side. The lower eye wouldalsohave 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 lifeis manifest
from several speciessuch as solesfloundersetc.being extremely
common. The chief advantages thus gained seem to be protection from their
enemiesand facility for feeding on the ground. The different members
howeverof the family presentas Schiodte remarksa long series of
forms exhibiting a gradual transition from Hippoglossus pinguis, which does
not in any considerable degree alter the shape in which it leaves the ovum,
to the soles, which are entirely thrown to one side.

Mr. Mivart has taken up this caseand remarks that a sudden spontaneous
transformation in the position of the eyes is hardly conceivablein which
I quite agree with him. He then adds: "If the transit was gradualthen
how such transit of one eye a minute fraction of the journey towards the
other side of the head could benefit the individual isindeedfar from
clear. It seemseventhat such an incipient transformation must rather
have been injurious." But he might have found an answer to this objection
in the excellent observations published in 1867 by Malm. The
Pleuronectidaewhile very young and still symmetricalwith their eyes
standing on opposite sides of the headcannot long retain a vertical
positionowing to the excessive depth of their bodiesthe small size of
their lateral finsand to their being destitute of a swim-bladder. Hence
soon growing tiredthey fall to the bottom on one side. While thus at
rest they often twistas Malm observedthe lower eye upwardto see above
them; and they do this so vigorously that the eye is pressed hard against
the upper part of the orbit. The forehead between the eyes consequently

becomesas could be plainly seentemporarily contracted in breadth. On
one occasion Malm saw a young fish raise and depress the lower eye through
an angular distance of about seventy degrees.

We should remember that the skull at this early age is cartilaginous and
flexibleso that it readily yields to muscular action. It is also known
with the higher animalseven after early youththat the skull yields and
is altered in shapeif the skin or muscles be permanently contracted
through disease or some accident. With long-eared rabbitsif one ear
flops forward and downwardits weight drags forward all the bones of the
skull on the same sideof which I have given a figure. Malm states that
the newly-hatched young of perchessalmonand several other symmetrical
fisheshave the habit of occasionally resting on one side at the bottom;
and he has observed that they often then strain their lower eyes so as to
look upward; and their skulls are thus rendered rather crooked. These
fisheshoweverare soon able to hold themselves in a vertical position
and no permanent effect is thus produced. With the Pleuronectidaeon the
other handthe older they grow the more habitually they rest on one side
owing to the increasing flatness of their bodiesand a permanent effect is
thus produced on the form of the headand on the position of the eyes.
Judging from analogythe tendency to distortion would no doubt be
increased through the principle of inheritance. Schiodte believesin
opposition to some other naturaliststhat the Pleuronectidae are not quite
symmetrical even in the embryo; and if this be sowe could understand how
it is that certain specieswhile younghabitually fall over and rest on
the left sideand other species on the right side. Malm addsin
confirmation of the above viewthat the adult Trachypterus arcticuswhich
is not a member of the Pleuronectidaerests on its left side at the
bottomand swims diagonally through the water; and in this fishthe two
sides of the head are said to be somewhat dissimilar. Our great authority
on FishesDr. Guntherconcludes his abstract of Malm's paperby
remarking that "the author gives a very simple explanation of the abnormal
condition of the Pleuronectoids."

We thus see that the first stages of the transit of the eye from one side
of the head to the otherwhich Mr. Mivart considers would be injurious
may be attributed to the habitno doubt beneficial to the individual and
to the speciesof endeavouring to look upward with both eyeswhile
resting on one side at the bottom. 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 surfacewith the jaw bones stronger and more
effective on thisthe eyeless side of the headthan on the otherfor the
sakeas Dr. Traquair supposesof feeding with ease on the ground.
Disuseon the other handwill account for the less developed condition of
the whole inferior half of the bodyincluding the lateral fins; though
Yarrel thinks that the reduced size of these fins is advantageous to the
fishas "there is so much less room for their action than with the larger
fins above." Perhaps the lesser number of teeth in the proportion of four
to seven in the upper halves of the two jaws of the plaiceto twenty-five
to thirty in the lower halvesmay likewise be accounted for by disuse.
>From the colourless state of the ventral surface of most fishes and of many
other animalswe may reasonably suppose that the absence of colour in
flat-fish on the sidewhether it be the right or leftwhich is undermost
is due to the exclusion of light. But it cannot be supposed that the
peculiar speckled appearance of the upper side of the soleso like the
sandy bed of the seaor the power in some speciesas recently shown by
Pouchetof changing their colour in accordance with the surrounding
surfaceor the presence of bony tubercles on the upper side of the turbot
are due to the action of the light. Here natural selection has probably
come into playas well as in adapting the general shape of the body of
these fishesand many other peculiaritiesto their habits of life. We
should keep in mindas I have before insistedthat the inherited effects
of the increased use of partsand perhaps of their disusewill be
strengthened by natural selection. For all spontaneous variations in the

right direction will thus be preserved; as will those individuals which
inherit in the highest degree the effects of the increased and beneficial
use of any part. How much to attribute in each particular case to the
effects of useand how much to natural selectionit seems impossible to

I may give another instance of a structure which apparently owes its origin
exclusively to use or habit. The extremity of the tail in some American
monkeys has been converted into a wonderfully perfect prehensile organand
serves as a fifth hand. A reviewerwho agrees with Mr. Mivart in every
detailremarks 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 itor favour their chance
of having and of rearing offspring." But there is no necessity for any
such belief. Habitand this almost implies that some benefit great or
small is thus derivedwould in all probability suffice for the work.
Brehm saw the young of an African monkey (Cercopithecus) clinging to the
under surface of their mother by their handsand at the same time they
hooked their little tails round that of their mother. Professor Henslow
kept in confinement some harvest mice (Mus messorius) which do not possess
a structurally prehensive tail; but he frequently observed that they curled
their tails round the branches of a bush placed in the cageand thus aided
themselves in climbing. I have received an analogous account from Dr.
Guntherwho has seen a mouse thus suspend itself. If the harvest mouse
had been more strictly arborealit would perhaps have had its tail
rendered structurally prehensileas is the case with some members of the
same order. Why Cercopithecusconsidering its habits while younghas not
become thus providedit would be difficult to say. It ishowever
possible that the long tail of this monkey may be of more service to it as
a balancing organ in making its prodigious leapsthan as a prehensile

The mammary glands are common to the whole class of mammalsand are
indispensable for their existence; they mustthereforehave been
developed at an extremely remote periodand we can know nothing positively
about their manner of development. Mr. Mivart asks: "Is it conceivable
that the young of any animal was ever saved from destruction by
accidentally sucking a drop of scarcely nutritious fluid from an
accidentally hypertrophied cutaneous gland of its mother? And even if one
was sowhat chance was there of the perpetuation of such a variation?"
But the case is not here put fairly. It is admitted by most evolutionists
that mammals are descended from a marsupial form; and if sothe mammary
glands will have been at first developed within the marsupial sack. In the
case of the fish (Hippocampus) the eggs are hatchedand the young are
reared for a timewithin a sack of this nature; and an American
naturalistMr. Lockwoodbelieves from what he has seen of the development
of the youngthat they are nourished by a secretion from the cutaneous
glands of the sack. Nowwith the early progenitors of mammalsalmost
before they deserved to be thus designatedis it not at least possible
that the young might have been similarly nourished? And in this casethe
individuals which secreted a fluidin some degree or manner the most
nutritiousso as to partake of the nature of milkwould in the long run
have reared a larger number of well-nourished offspringthan would the
individuals which secreted a poorer fluid; and thus the cutaneous glands
which are the homologues of the mammary glandswould have been improved or
rendered more effective. It accords with the widely extended principle of
specialisationthat the glands over a certain space of the sack should
have become more highly developed than the remainder; and they would then
have formed a breastbut at first without a nippleas we see in the
Ornithorhyncusat the base of the mammalian series. Through what agency
the glands over a certain space became more highly specialised than the
othersI will not pretend to decidewhether in part through compensation
of growththe effects of useor of natural selection.

The development of the mammary glands would have been of no serviceand
could not have been affected through natural selectionunless the young at
the same time were able to partake of the secretion. There is no greater
difficulty in understanding how young mammals have instinctively learned to
suck the breastthan in understanding how unhatched chickens have learned
to break the egg-shell by tapping against it with their specially adapted
beaks; or how a few hours after leaving the shell they have learned to pick
up grains of food. In such cases the most probable solution seems to be
that the habit was at first acquired by practice at a more advanced age
and afterwards transmitted to the offspring at an earlier age. But the
young kangaroo is said not to suckonly to cling to the nipple of its
motherwho has the power of injecting milk into the mouth of her helpless
half-formed offspring. On this head Mr. Mivart remarks: "Did no special
provision existthe young one must infallibly be choked by the intrusion
of the milk into the wind-pipe. But there IS a special provision. The
larynx is so elongated that it rises up into the posterior end of the nasal
passageand is thus enabled to give free entrance to the air for the
lungswhile the milk passes harmlessly on each side of this elongated
larynxand so safely attains the gullet behind it." Mr. Mivart then asks
how did natural selection remove in the adult kangaroo (and in most other
mammalson the assumption that they are descended from a marsupial form)
this at least perfectly innocent and harmless structure?It may be
suggested in answer that the voicewhich is certainly of high importance
to many animalscould hardly have been used with full force as long as the
larynx entered the nasal passage; and Professor Flower has suggested to me
that this structure would have greatly interfered with an animal swallowing
solid food.

We will now turn for a short space to the lower divisions of the animal
kingdom. The Echinodermata (star-fishessea-urchinsetc.) are furnished
with remarkable organscalled pedicellariaewhich consistwhen well
developedof a tridactyle forceps--that isof one formed of three
serrated armsneatly fitting together and placed on the summit of a
flexible stemmoved by muscles. These forceps can seize firmly hold of
any object; and Alexander Agassiz has seen an Echinus or sea-urchin rapidly
passing particles of excrement from forceps to forceps down certain lines
of its bodyin order that its shell should not be fouled. But there is no
doubt that besides removing dirt of all kindsthey subserve other
functions; and one of these apparently is defence.

With respect to these organsMr. Mivartas on so many previous occasions
asks: "What would be the utility of the FIRST RUDIMENTARY BEGINNINGS of
such structuresand how could such insipient buddings have ever preserved
the life of a single Echinus?" He addsnot 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. Mivarttridactyle forcepsesimmovably fixed at
the basebut capable of a snapping actioncertainly exist on some starfishes;
and this is intelligible if they serveat least in partas a
means of defence. Mr. Agassizto whose great kindness I am indebted for
much information on the subjectinforms me that there are other starfishes
in which one of the three arms of the forceps is reduced to a
support for the other two; and againother genera in which the third arm
is completely lost. In Echinoneusthe shell is described by M. Perrier as
bearing two kinds of pedicellariaeone resembling those of Echinusand
the other those of Spatangus; and such cases are always interesting as
affording the means of apparently sudden transitionsthrough 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. Mullerthat

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 individualas well as from a long and perfect series of
gradations in different species and generafrom simple granules to
ordinary spinesto perfect tridactyle pedicellariae. The gradation
extends even to the manner in which ordinary spines and the pedicellariae
with their supporting calcareous rodsare articulated to the shell. In
certain genera of star-fishesthe very combinations needed to show that
the pedicellariae are only modified branching spinesmay be found. Thus
we have fixed spineswith three equi-distantserratedmovable branches
articulated to near their bases; and higher upon the same spinethree
other movable branches. Now when the latter arise from the summit of a
spine they formin facta rude tridactyle pedicellariaeand 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 spineis unmistakable. It is generally admitted
that the ordinary spines serve as a protection; and if sothere 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 gradationfrom an ordinary fixed spine to
a fixed pedicellariaewould be of service.

In certain genera of star-fishes these organsinstead of being fixed or
borne on an immovable supportare placed on the summit of a flexible and
muscularthough shortstem; 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 shelland 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 gradationsas he addsmay likewise be found
between the pedicellariae of the star-fishes and the hooks of the
Ophiuriansanother group of the Echinodermata; and again between the
pedicellariae of sea-urchins and the anchors of the Holothuriaealso
belonging to the same great class.

Certain compound animalsor zoophytesas they have been termednamely
the Polyzoaare 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
miniatureseated on a neck and capable of movementas is likewise the
lower jaw or mandible. In one species observed by meall the avicularia
on the same branch often moved simultaneously backwards and forwardswith
the lower jaw widely openthrough an angle of about 90 degreesin 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 casechiefly on account of the supposed difficulty
of organsnamely the avicularia of the Polyzoa and the pedicellariae of
the Echinodermatawhich 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 polyzoaryremoving what might be noxious
to the delicate inhabitants of the cells when their tentacula are
protruded." The avicularialike the vibraculaprobably serve for
defencebut they also catch and kill small living animalswhichit is
believedare afterwards swept by the currents within reach of the
tentacula of the zooids. Some species are provided with avicularia and
vibraculasome 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 vibraculumand an avicularium like the head of a bird;
yet they are almost certainly homologous and have been developed from the
same common sourcenamely a zooid with its cell. Hencewe can understand
how it is that these organs graduate in some casesas I am informed by Mr.
Buskinto each other. Thuswith the avicularia of several species of
Lepraliathe 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 cellswithout having passed through the avicularian stage;
but it seems more probable that they have passed through this stageas
during the early stages of the transformationthe other parts of the cell
with the included zooidcould hardly have disappeared at once. In many
cases the vibracula have a grooved support at the basewhich seems to
represent the fixed beak; though this support in some species is quite
absent. This view of the development of the vibraculaif trustworthyis
interesting; for supposing that all the species provided with avicularia
had become extinctno 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 heador 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 zooidthere
is no difficulty in believing that all the gradationsby which the lip
became converted first into the lower mandible of an aviculariumand then
into an elongated bristlelikewise served as a protection in different
ways and under different circumstances.

In the vegetable kingdom Mr. Mivart only alludes to two casesnamely the
structure of the flowers of orchidsand the movements of climbing plants.
With respect to the formerhe says: "The explanation of their ORIGIN is

deemed thoroughly unsatisfactory--utterly insufficient to explain the
incipientinfinitesimal beginnings of structures which are of utility only
when they are considerably developed." As I have fully treated this
subject in another workI will here give only a few details on one alone
of the most striking peculiarities of the flowers of orchidsnamelytheir
pollinia. A polliniumwhen highly developedconsists of a mass of
pollen-grainsaffixed to an elastic foot-stalk or caudicleand 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-massesand the grains are
merely tied together by fine threads; but as these are not confined to
orchidsthey need not here be considered; yet I may mention that at the
base of the orchidaceous seriesin Cypripediumwe 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 caudicleeven when of
considerable length and highly developedwe 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 peculiaritynamelythe little mass of
viscid matter attached to the end of the caudiclea long series of
gradations can be specifiedeach of plain service to the plant. In most
flowers belonging to other orders the stigma secretes a little viscid
matter. Nowin certain orchids similar viscid matter is secretedbut in
much larger quantities by one alone of the three stigmas; and this stigma
perhaps in consequence of the copious secretionis rendered sterile. When
an insect visits a flower of this kindit rubs off some of the viscid
matterand thus at the same time drags away some of the pollen-grains.
>From this simple conditionwhich differs but little from that of a
multitude of common flowersthere are endless gradations--to species in
which the pollen-mass terminates in a very shortfree caudicle--to others
in which the caudicle becomes firmly attached to the viscid matterwith
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 threadswith the stigma differing but little from
that of the ordinary flowersto a highly complex polliniumadmirably
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 thisand in almost every other casethe enquiry may be
pushed further backwards; and it may be asked how did the stigma of an
ordinary flower become viscidbut as we do not know the full history of
any one group of beingsit is as useless to askas it is hopeless to
attempt answeringsuch questions.

We will now turn to climbing plants. These can be arranged in a long
seriesfrom those which simply twine round a supportto those which I
have called leaf-climbersand to those provided with tendrils. In these
two latter classes the stems have generallybut not alwayslost the power
of twiningthough they retain the power of revolvingwhich the tendrils
likewise possess. The gradations from leaf-climbers to tendril bearers are
wonderfully closeand certain plants may be differently placed in either
class. But in ascending the series from simple twiners to leaf-climbers
an important quality is addednamely sensitiveness to a touchby which
means the foot-stalks of the leaves or flowersor these modified and
converted into tendrilsare excited to bend round and clasp the touching
object. He who will read my memoir on these plants willI thinkadmit
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 instanceit 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-climberif the foot-stalks had possessed in any slight degree the
requisite sensitiveness to a touch.

As twining is the simplest means of ascending a supportand forms the
basis of our seriesit may naturally be asked how did plants acquire this
power in an incipient degreeafterwards to be improved and increased
through natural selection. The power of twining dependsfirstlyon the
stems while young being extremely flexible (but this is a character common
to many plants which are not climbers); andsecondlyon their continually
bending to all points of the compassone after the other in successionin
the same order. By this movement the stems are inclined to all sidesand
are made to move round and round. As soon as the lower part of a stem
strikes against any object and is stoppedthe upper part still goes on
bending and revolvingand 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 plantssingle species and
single genera possess the power of revolvingand have thus become twiners
they must have independently acquired itand cannot have inherited it from
a common progenitor. HenceI 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 predictionI
knew of only one imperfect casenamelyof the young flower-peduncles of a
Maurandia which revolved slightly and irregularlylike the stems of
twining plantsbut 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
plainlythough irregularlyand 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; anyhowthey are not
of the least use in the way of climbingwhich is the point that concerns
us. Nevertheless we can see that if the stems of these plants had been
flexibleand if under the conditions to which they are exposed it had
profited them to ascend to a heightthen the habit of slightly and
irregularly revolving might have been increased and utilised through
natural selectionuntil they had become converted into well-developed
twining species.

With respect to the sensitiveness of the foot-stalks of the leaves and
flowersand of tendrilsnearly the same remarks are applicable as in the
case of the revolving movements of twining plants. As a vast number of
speciesbelonging to widely distinct groupsare endowed with this kind of
sensitivenessit 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 movedespecially after
exposure to a hot sunwhen they were gently and repeatedly touchedor
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
distinctbut 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 Hofmeisterthe young shoots and leaves of all plants move after being
shaken; and with climbing plants it isas we knowonly during the early
stages of growth that the foot-stalks and tendrils are sensitive.

It is scarcely possible that the above slight movementsdue to a touch or
shakein the young and growing organs of plantscan be of any functional
importance to them. But plants possessin obedience to various stimuli
powers of movementwhich are of manifest importance to them; for instance
towards and more rarely from the light--in opposition toand more rarely
in the direction ofthe attraction of gravity. When the nerves and
muscles of an animal are excited by galvanism or by the absorption of

strychninethe 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 thatfrom having the power of
movement in obedience to certain stimulithey are excited in an incidental
manner by a touchor by being shaken. Hence there is no great difficulty
in admitting that in the case of leaf-climbers and tendril-bearersit is
this tendency which has been taken advantage of and increased through
natural selection. It ishoweverprobablefrom reasons which I have
assigned in my memoirthat this will have occurred only with plants which
had already acquired the power of revolvingand had thus become twiners.

I have already endeavoured to explain how plants became twinersnamelyby
the increase of a tendency to slight and irregular revolving movements
which were at first of no use to them; this movementas well as that due
to a touch or shakebeing the incidental result of the power of moving
gained for other and beneficial purposes. Whetherduring the gradual
development of climbing plantsnatural selection has been aided by the
inherited effects of useI will not pretend to decide; but we know that
certain periodical movementsfor instance the so-called sleep of plants
are governed by habit.

I have now considered enoughperhaps more than enoughof the cases
selected with care by a skilful naturalistto prove that natural selection
is incompetent to account for the incipient stages of useful structures;
and I have shownas I hopethat there is no great difficulty on this
head. A good opportunity has thus been afforded for enlarging a little on
gradations of structureoften associated with strange functions--an
important subjectwhich was not treated at sufficient length in the former
editions of this work. I will now briefly recapitulate the foregoing

With the giraffethe continued preservation of the individuals of some
extinct high-reaching ruminantwhich had the longest neckslegsetc.
and could browse a little above the average heightand the continued
destruction of those which could not browse so highwould have sufficed
for the production of this remarkable quadruped; but the prolonged use of
all the partstogether with inheritancewill have aided in an important
manner in their co-ordination. With the many insects which imitate various
objectsthere 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 selectionsince 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 variationsuntil the points were convertedfirst into
lamellated knobs or teethlike those on the beak of a goose--then into
short lamellaelike those of the domestic ducks--and then into lamellae
as perfect as those of the shoveller-duck--and finally into the gigantic
plates of baleenas in the mouth of the Greenland whale. In the family of
the ducksthe lamellae are first used as teeththen partly as teeth and
partly as a sifting apparatusand at last almost exclusively for this
latter purpose.

With such structures as the above lamellae of horn or whalebonehabit or
use can have done little or nothingas far as we can judgetowards their
development. On the other handthe transportal of the lower eye of a
flat-fish to the upper side of the headand the formation of a prehensile
tailmay be attributed almost wholly to continued usetogether with
inheritance. With respect to the mammae of the higher animalsthe 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 selectionand
concentrated into a confined areain which case they would have formed a
mamma. There is no more difficulty in understanding how the branched
spines of some ancient Echinodermwhich served as a defencebecame
developed through natural selection into tridactyle pedicellariaethan in
understanding the development of the pincers of crustaceansthrough
slightserviceable modifications in the ultimate and penultimate segments
of a limbwhich 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 orchidsthe threads which originally served to tie
together the pollen-grainscan be traced cohering into caudicles; and the
steps can likewise be followed by which viscid mattersuch as that
secreted by the stigmas of ordinary flowersand still subserving nearly
but not quite the same purposebecame attached to the free ends of the
caudicles--all these gradations being of manifest benefit to the plants in
question. With respect to climbing plantsI need not repeat what has been
so lately said.

It has often been askedif natural selection be so potentwhy has not
this or that structure been gained by certain speciesto which it would
apparently have been advantageous? But it is unreasonable to expect a
precise answer to such questionsconsidering our ignorance of the past
history of each speciesand of the conditions which at the present day
determine its numbers and range. In most cases only general reasonsbut
in some few cases special reasonscan be assigned. Thus to adapt a
species to new habits of lifemany co-ordinated modifications are almost
indispensableand 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
agencieswhich stood in no relation to certain structureswhich we
imagine would have been gained through natural selection from appearing to
us advantageous to the species. In this caseas the struggle for life did
not depend on such structuresthey could not have been acquired through
natural selection. In many cases complex and long-enduring conditions
often of a peculiar natureare necessary for the development of a
structure; and the requisite conditions may seldom have concurred. The
belief that any given structurewhich we thinkoften erroneouslywould
have been beneficial to a specieswould have been gained under all
circumstances through natural selectionis 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 consideredand the others will hereafter
be considered. They seem to me to partake little of the character of
demonstrationand to have little weight in comparison with those in favour
of the power of natural selectionaided by the other agencies often
specified. I am bound to addthat some of the facts and arguments here
used by mehave 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 instancehe 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 conclusionwhich implies great
breaks or discontinuity in the seriesappears to me improbable in the
highest degree.

Everyone who believes in slow and gradual evolutionwill of course admit
that specific changes may have been as abrupt and as great as any single
variation which we meet with under natureor even under domestication.
But as species are more variable when domesticated or cultivated than under
their natural conditionsit is not probable that such great and abrupt
variations have often occurred under natureas are known occasionally to
arise under domestication. Of these latter variations several may be
attributed to reversion; and the characters which thus reappear wereit is
probablein many cases at first gained in a gradual manner. A still
greater number must be called monstrositiessuch as six-fingered men
porcupine menAncon sheepNiata cattleetc.; and as they are widely
different in character from natural speciesthey throw very little light
on our subject. Excluding such cases of abrupt variationsthe few which
remain would at best constituteif found in a state of naturedoubtful
speciesclosely related to their parental types.

My reasons for doubting whether natural species have changed as abruptly as
have occasionally domestic racesand for entirely disbelieving that they
have changed in the wonderful manner indicated by Mr. Mivartare as
follows. According to our experienceabrupt and strongly marked
variations occur in our domesticated productionssingly and at rather long
intervals of time. If such occurred under naturethey would be liableas
formerly explainedto be lost by accidental causes of destruction and by
subsequent intercrossing; and so it is known to be under domestication
unless abrupt variations of this kind are specially preserved and separated
by the care of man. Hencein order that a new species should suddenly
appear in the manner supposed by Mr. Mivartit is almost necessary to
believein opposition to all analogythat several wonderfully changed
individuals appeared simultaneously within the same district. This
difficultyas in the case of unconscious selection by manis avoided on
the theory of gradual evolutionthrough the preservation of a large number
of individualswhich varied more or less in any favourable directionand
of the destruction of a large number which varied in an opposite manner.

That many species have been evolved in an extremely gradual mannerthere
can hardly be a doubt. 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. On every continentin proceeding from
north to southfrom lowland to uplandetc.we meet with a host of
closely related or representative species; as we likewise do on certain
distinct continentswhich we have reason to believe were formerly
connected. But in making these and the following remarksI am compelled
to allude to subjects hereafter to be discussed. Look at the many outlying
islands round a continentand see how many of their inhabitants can be
raised only to the rank of doubtful species. So it is if we look to past
timesand compare the species which have just passed away with those still
living within the same areas; or if we compare the fossil species embedded
in the sub-stages of the same geological formation. It is indeed manifest
that multitudes of species are related in the closest manner to other
species that still existor have lately existed; and it will hardly be
maintained that such species have been developed in an abrupt or sudden
manner. Nor should it be forgottenwhen we look to the special parts of
allied speciesinstead of to distinct speciesthat numerous and

wonderfully fine gradations can be tracedconnecting together widely
different structures.

Many large groups of facts are intelligible only on the principle that
species have been evolved by very small steps. For instancethe fact that
the species included in the larger genera are more closely related to each
otherand present a greater number of varieties than do the species in the
smaller genera. The former are also grouped in little clusterslike
varieties round species; and they present other analogies with varieties
as was shown in our second chapter. On this same principle we can
understand how it is that specific characters are more variable than
generic characters; and how the parts which are developed in an
extraordinary degree or manner are more variable than other parts of the
same species. Many analogous factsall pointing in the same direction
could be added.

Although very many species have almost certainly been produced by steps not
greater than those separating fine varieties; yet it may be maintained that
some have been developed in a different and abrupt manner. Such an
admissionhoweverought not to be made without strong evidence being
assigned. The vague and in some respects false analogiesas they have
been shown to be by Mr. Chauncey Wrightwhich have been advanced in favour
of this viewsuch as the sudden crystallisation of inorganic substances
or the falling of a facetted spheroid from one facet to anotherhardly
deserve consideration. One class of factshowevernamelythe sudden
appearance of new and distinct forms of life in our geological formations
supports at first sight the belief in abrupt development. But the value of
this evidence depends entirely on the perfection of the geological record
in relation to periods remote in the history of the world. If the record
is as fragmentary as many geologists strenuously assertthere is nothing
strange in new forms appearing as if suddenly developed.

Unless we admit transformations as prodigious as those advocated by Mr.
Mivartsuch as the sudden development of the wings of birds or batsor
the sudden conversion of a Hipparion into a horsehardly any light is
thrown by the belief in abrupt modifications on the deficiency of
connecting links in our geological formations. But against the belief in
such abrupt changesembryology enters a strong protest. It is notorious
that the wings of birds and batsand the legs of horses or other
quadrupedsare undistinguishable at an early embryonic periodand that
they become differentiated by insensibly fine steps. Embryological
resemblances of all kinds can be accounted foras we shall hereafter see
by the progenitors of our existing species having varied after early youth
and having transmitted their newly-acquired characters to their offspring
at a corresponding age. The embryo is thus left almost unaffectedand
serves as a record of the past condition of the species. 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. On this
view of the meaning of embryological resemblancesand indeed on any view
it is incredible that an animal should have undergone such momentous and
abrupt transformations as those above indicatedand yet should not bear
even a trace in its embryonic condition of any sudden modificationevery
detail in its structure being developed by insensibly fine steps.

He who believes that some ancient form was transformed suddenly through an
internal force or tendency intofor instanceone furnished with wings
will be almost compelled to assumein opposition to all analogythat many
individuals varied simultaneously. It cannot be denied that such abrupt
and great changes of structure are widely different from those which most
species apparently have undergone. 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 conditionshave been suddenly produced;
and of such complex and wonderful co-adaptationshe 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. To admit all this isas it seems to meto enter into the realms
of miracleand to leave those of science.



Instincts comparable with habitsbut different in their origin --
Instincts graduated -- Aphides and ants -- Instincts variable -- Domestic
instinctstheir origin -- Natural instincts of the cuckoomolothrus
ostrichand parasitic bees -- Slave-making ants -- Hive-beeits
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.

Many instincts are so wonderful that their development will probably appear
to the reader a difficulty sufficient to overthrow my whole theory. I may
here premisethat I have nothing to do with the origin of the mental
powersany more than I have with that of life itself. We are concerned
only with the diversities of instinct and of the other mental faculties in
animals of the same class.

I will not attempt any definition of instinct. It would be easy to show
that several distinct mental actions are commonly embraced by this term;
but every one understands what is meantwhen it is said that instinct
impels the cuckoo to migrate and to lay her eggs in other birds' nests. An
actionwhich we ourselves require experience to enable us to performwhen
performed by an animalmore especially by a very young onewithout
experienceand when performed by many individuals in the same waywithout
their knowing for what purpose it is performedis usually said to be
instinctive. But I could show that none of these characters are universal.
A little dose of judgment or reasonas Pierre Huber expresses itoften
comes into playeven with animals low in the scale of nature.

Frederick Cuvier and several of the older metaphysicians have compared
instinct with habit. This comparison givesI thinkan accurate notion of
the frame of mind under which an instinctive action is performedbut not
necessarily of its origin. How unconsciously many habitual actions are
performedindeed not rarely in direct opposition to our conscious will!
yet they may be modified by the will or reason. Habits easily become
associated with other habitswith certain periods of time and states of
the body. When once acquiredthey often remain constant throughout life.
Several other points of resemblance between instincts and habits could be
pointed out. As in repeating a well-known songso in instinctsone
action follows another by a sort of rhythm; if a person be interrupted in a
songor in repeating anything by rotehe is generally forced to go back
to recover the habitual train of thought: so P. Huber found it was with a
caterpillarwhich makes a very complicated hammock; for if he took a
caterpillar which had completed its hammock up tosaythe sixth stage of
constructionand put it into a hammock completed up only to the third
stagethe caterpillar simply re-performed the fourthfifthand sixth
stages of construction. Ifhowevera caterpillar were taken out of a
hammock made upfor instanceto the third stageand were put into one
finished up to the sixth stageso that much of its work was already done
for itfar from deriving any benefit from thisit was much embarrassed
andin order to complete its hammockseemed forced to start from the
third stagewhere it had left offand thus tried to complete the already
finished work.

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. If Mozartinstead of playing the pianoforte at three years
old with wonderfully little practicehad played a tune with no practice at
allbe might truly be said to have done so instinctively. But it would be
a serious error to suppose that the greater number of instincts have been
acquired by habit in one generationand then transmitted by inheritance to
succeeding generations. It can be clearly shown that the most wonderful
instincts with which we are acquaintednamelythose of the hive-bee and
of many antscould not possibly have been acquired by habit.

It will be universally admitted that instincts are as important as
corporeal structures for the welfare of each speciesunder its present
conditions of life. Under changed conditions of lifeit is at least
possible that slight modifications of instinct might be profitable to a
species; and if it can be shown that instincts do vary ever so littlethen
I can see no difficulty in natural selection preserving and continually
accumulating variations of instinct to any extent that was profitable. It
is thusas I believethat all the most complex and wonderful instincts
have originated. As modifications of corporeal structure arise fromand
are increased byuse or habitand are diminished or lost by disuseso I
do not doubt it has been with instincts. 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;--that is of variations produced by the same unknown causes which
produce slight deviations of bodily structure.

No complex instinct can possibly be produced through natural selection
except by the slow and gradual accumulation of numerousslightyet
profitablevariations. Henceas in the case of corporeal structureswe
ought to find in naturenot 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; or we ought
at least to be able to show that gradations of some kind are possible; and
this we certainly can do. I have been surprised to findmaking allowance
for the instincts of animals having been but little observedexcept in
Europe and North Americaand for no instinct being known among extinct
specieshow very generally gradationsleading to the most complex
instinctscan be discovered. Changes of instinct may sometimes be
facilitated by the same species having different instincts at different
periods of lifeor at different seasons of the yearor when placed under
different circumstancesetc.; in which case either the one or the other
instinct might be preserved by natural selection. And such instances of
diversity of instinct in the same species can be shown to occur in nature.

Againas in the case of corporeal structureand conformably to my theory
the instinct of each species is good for itselfbut has neveras far as
we can judgebeen produced for the exclusive good of others. One of the
strongest instances of an animal apparently performing an action for the
sole good of anotherwith which I am acquaintedis that of aphides
voluntarily yieldingas was first observed by Hubertheir sweet excretion
to ants: that they do so voluntarilythe following facts show. I removed
all the ants from a group of about a dozen aphides on a dock-plantand
prevented their attendance during several hours. After this intervalI
felt sure that the aphides would want to excrete. I watched them for some
time through a lensbut not one excreted; I then tickled and stroked them
with a hair in the same manneras well as I couldas the ants do with
their antennae; but not one excreted. AfterwardsI allowed an ant to
visit themand it immediately seemedby its eager way of running about to
be well aware what a rich flock it had discovered; it then began to play
with its antennae on the abdomen first of one aphis and then of another;
and eachas soon as it felt the antennaeimmediately lifted up its
abdomen and excreted a limpid drop of sweet juicewhich was eagerly
devoured by the ant. Even the quite young aphides behaved in this manner
showing that the action was instinctiveand not the result of experience.

It is certainfrom the observations of Huberthat the aphides show no
dislike to the ants: if the latter be not present they are at last
compelled to eject their excretion. But as the excretion is extremely
viscidit is no doubt a convenience to the aphides to have it removed;
therefore probably they do not excrete solely for the good of the ants.
Although there is no evidence that any animal performs an action for the
exclusive good of another speciesyet each tries to take advantage of the
instincts of othersas each takes advantage of the weaker bodily structure
of other species. So again certain instincts cannot be considered as
absolutely perfect; but as details on this and other such points are not
indispensablethey may be here passed over.

As some degree of variation in instincts under a state of natureand the
inheritance of such variationsare indispensable for the action of natural
selectionas many instances as possible ought to be given; but want of
space prevents me. I can only assert that instincts certainly do vary--for
instancethe migratory instinctboth in extent and directionand in its
total loss. So it is with the nests of birdswhich vary partly in
dependence on the situations chosenand on the nature and temperature of
the country inhabitedbut often from causes wholly unknown to us. Audubon
has given several remarkable cases of differences in the nests of the same
species in the northern and southern United States. Whyit has been
askedif instinct be variablehas 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 workas I have seenwith wax hardened
with vermilion or softened with lard. Andrew Knight observed that his
beesinstead of laboriously collecting propolisused a cement of wax and
turpentinewith which he had covered decorticated trees. It has lately
been shown that beesinstead of searching for pollenwill gladly use a
very different substancenamelyoatmeal. Fear of any particular enemy is
certainly an instinctive qualityas may be seen in nestling birdsthough
it is strengthened by experienceand by the sight of fear of the same
enemy in other animals. The fear of man is slowly acquiredas I have
elsewhere shownby the various animals which inhabit desert islands; and
we see an instance of thiseven in Englandin the greater wildness of all
our large birds in comparison with our small birds; for the large birds
have been most persecuted by man. We may safely attribute the greater
wildness of our large birds to this cause; for in uninhabited islands large
birds are not more fearful than small; and the magpieso wary in England
is tame in Norwayas is the hooded crow in Egypt.

That the mental qualities of animals of the same kindborn in a state of
naturevary muchcould be shown by many facts. Several cases could also
be adduced of occasional and strange habits in wild animalswhichif
advantageous to the speciesmight have given risethrough natural
selectionto new instincts. But I am well aware that these general
statementswithout the facts in detailcan produce but a feeble effect on
the reader's mind. I can only repeat my assurancethat I do not speak
without good evidence.


The possibilityor even probabilityof inherited variations of instinct
in a state of nature will be strengthened by briefly considering a few
cases under domestication. 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. It is notorious
how much domestic animals vary in their mental qualities. With catsfor
instanceone naturally takes to catching ratsand another miceand these
tendencies are known to be inherited. One cataccording to Mr. St. John
always brought home game birdsanother hares or rabbitsand another
hunted on marshy ground and almost nightly caught woodcocks or snipes. A
number of curious and authentic instances could be given of various shades
of disposition and tasteand likewise of the oddest tricksassociated

with certain frames of mind or periods of time. 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;
retrieving is certainly in some degree inherited by retrievers; and a
tendency to run roundinstead of ata flock of sheepby shepherd-dogs.
I cannot see that these actionsperformed without experience by the young
and in nearly the same manner by each individualperformed with eager
delight by each breedand without the end being known--for the young
pointer can no more know that he points to aid his masterthan the white
butterfly knows why she lays her eggs on the leaf of the cabbage--I cannot
see that these actions differ essentially from true instincts. If we were
to behold one kind of wolfwhen young and without any trainingas soon as
it scented its preystand motionless like a statueand then slowly crawl
forward with a peculiar gait; and another kind of wolf rushing round
instead of ata herd of deerand driving them to a distant pointwe
should assuredly call these actions instinctive. Domestic instinctsas
they may be calledare certainly far less fixed than natural instincts;
but they have been acted on by far less rigorous selectionand have been
transmitted for an incomparably shorter periodunder less fixed conditions
of life.

How strongly these domestic instinctshabitsand dispositions are
inheritedand how curiously they become mingledis well shown when
different breeds of dogs are crossed. Thus it is known that a cross with a
bull-dog has affected for many generations the courage and obstinacy of
greyhounds; and a cross with a greyhound has given to a whole family of
shepherd-dogs a tendency to hunt hares. These domestic instinctswhen
thus tested by crossingresemble natural instinctswhich in a like manner
become curiously blended togetherand for a long period exhibit traces of
the instincts of either parent: for exampleLe Roy describes a dogwhose
great-grandfather was a wolfand this dog showed a trace of its wild
parentage only in one wayby not coming in a straight line to his master
when called.

Domestic instincts are sometimes spoken of as actions which have become
inherited solely from long-continued and compulsory habitbut this is not
true. No one would ever have thought of teachingor probably could have
taughtthe tumbler-pigeon to tumble--an action whichas I have witnessed
is performed by young birdsthat have never seen a pigeon tumble. We may
believe that some one pigeon showed a slight tendency to this strange
habitand that the long-continued selection of the best individuals in
successive generations made tumblers what they now are; and near Glasgow
there are house-tumblersas I hear from Mr. Brentwhich cannot fly
eighteen inches high without going head over heels. It may be doubted
whether any one would have thought of training a dog to pointhad not some
one dog naturally shown a tendency in this line; and this is known
occasionally to happenas I once sawin a pure terrier: the act of
pointing is probablyas many have thoughtonly the exaggerated pause of
an animal preparing to spring on its prey. When the first tendency to
point was once displayedmethodical selection and the inherited effects of
compulsory training in each successive generation would soon complete the
work; and unconscious selection is still in progressas each man tries to
procurewithout intending to improve the breeddogs which stand and hunt
best. On the other handhabit alone in some cases has sufficed; hardly
any animal is more difficult to tame than the young of the wild rabbit;
scarcely any animal is tamer than the young of the tame rabbit; but I can
hardly suppose that domestic rabbits have often been selected for tameness
alone; so that we must attribute at least the greater part of the inherited
change from extreme wildness to extreme tamenessto habit and
long-continued close confinement.

Natural instincts are lost under domestication: a remarkable instance of
this is seen in those breeds of fowls which very rarely or never become

broody,that isnever wish to sit on their eggs. Familiarity alone
prevents our seeing how largely and how permanently the minds of our
domestic animals have been modified. It is scarcely possible to doubt that
the love of man has become instinctive in the dog. All wolvesfoxes
jackals and species of the cat genuswhen kept tameare most eager to
attack poultrysheep and pigs; and this tendency has been found incurable
in dogs which have been brought home as puppies from countries such as
Tierra del Fuego and Australiawhere the savages do not keep these
domestic animals. How rarelyon the other handdo our civilised dogs
even when quite youngrequire to be taught not to attack poultrysheep
and pigs! No doubt they occasionally do make an attackand are then
beaten; and if not curedthey are destroyed; so that habit and some degree
of selection have probably concurred in civilising by inheritance our dogs.
On the other handyoung chickens have lost wholly by habitthat fear of
the dog and cat which no doubt was originally instinctive in themfor I am
informed by Captain Hutton that the young chickens of the parent stockthe
Gallus bankivawhen reared in India under a henare at first excessively
wild. So it is with young pheasants reared in England under a hen. It is
not that chickens have lost all fearbut fear only of dogs and catsfor
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; and this is evidently done for the instinctive purpose of
allowingas we see in wild ground-birdstheir mother to fly away. But
this instinct retained by our chickens has become useless under
domesticationfor the mother-hen has almost lost by disuse the power of

Hencewe may conclude that under domestication instincts have been
acquired and natural instincts have been lostpartly by habit and partly
by man selecting and accumulatingduring successive generationspeculiar
mental habits and actionswhich at first appeared from what we must in our
ignorance call an accident. In some cases compulsory habit alone has
sufficed to produce inherited mental changes; in other cases compulsory
habit has done nothingand all has been the result of selectionpursued
both methodically and unconsciously; but in most cases habit and selection
have probably concurred.


We shallperhapsbest understand how instincts in a state of nature have
become modified by selection by considering a few cases. I will select
only threenamelythe instinct which leads the cuckoo to lay her eggs in
other birds' nests; the slave-making instinct of certain ants; 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.


It is supposed by some naturalists that the more immediate cause of the
instinct of the cuckoo is that she lays her eggsnot dailybut at
intervals of two or three days; so thatif she were to make her own nest
and sit on her own eggsthose 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. If this were the case the process of laying and hatching
might be inconveniently longmore especially as she migrates at a very
early period; and the first hatched young would probably have to be fed by
the male alone. But the American cuckoo is in this predicamentfor she
makes her own nest and has eggs and young successively hatchedall at the
same time. It has been both asserted and denied that the American cuckoo
occasionally lays her eggs in other birds' nests; but I have lately heard
from Dr. Merrillof Iowathat he once found in Illinois a young cuckoo
together with a young jay in the nest of a blue jay (Garrulus cristatus);
and as both were nearly full featheredthere could be no mistake in their

identification. I could also give several instances of various birds which
have been known occasionally to lay their eggs in other birds' nests. Now
let us suppose that the ancient progenitor of our European cuckoo had the
habits of the American cuckooand that she occasionally laid an egg in
another bird's nest. If the old bird profited by this occasional habit
through being enabled to emigrate earlier or through any other cause; 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
encumbered as she could hardly fail to be by having eggs and young of
different ages at the same timethen the old birds or the fostered young
would gain an advantage. 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 motherand in their turn would be apt to lay their
eggs in other birds' nestsand thus be more successful in rearing their
young. By a continued process of this natureI believe that the strange
instinct of our cuckoo has been generated. It hasalso recently been
ascertained on sufficient evidenceby Adolf Mullerthat the cuckoo
occasionally lays her eggs on the bare groundsits on them and feeds her
young. This rare event is probably a case of reversion to the long-lost
aboriginal instinct of nidification.

It has been objected that I have not noticed other related instincts and
adaptations of structure in the cuckoowhich are spoken of as necessarily
co-ordinated. But in all casesspeculation on an instinct known to us
only in a single speciesis uselessfor we have hitherto had no facts to
guide us. Until recently the instincts of the European and of the nonparasitic
American cuckoo alone were known; nowowing to Mr. Ramsay's
observationswe have learned something about three Australian species
which lay their eggs in other birds' nests. The chief points to be
referred to are three: firstthat the common cuckoowith rare
exceptionslays only one egg in a nestso that the large and voracious
young bird receives ample food. Secondlythat the eggs are remarkably
smallnot exceeding those of the skylark--a bird about one-fourth as large
as the cuckoo. 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. Thirdlythat the young cuckoosoon after birthhas the
instinctthe strength and a properly shaped back for ejecting its fosterbrothers
which then perish from cold and hunger. This has been boldly
called a beneficent arrangementin order that the young cuckoo may get
sufficient foodand 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 nestit is not rare to find two and even three eggs in
the same nest. In the bronze cuckoo the eggs vary greatly in sizefrom
eight to ten lines in length. Nowif it had been of an advantage to this
species to have laid eggs even smaller than those now laidso as to have
deceived certain foster-parentsoras is more probableto 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)then
there is no difficulty in believing that a race or species might have been
formed which would have laid smaller and smaller eggs; for these would have
been more safely hatched and reared. Mr. Ramsay remarks that two of the
Australian cuckooswhen they lay their eggs in an open nestmanifest a
decided preference for nests containing eggs similar in colour to their
own. The European species apparently manifests some tendency towards a
similar instinctbut not rarely departs from itas is shown by her laying
her dull and pale-coloured eggs in the nest of the hedge-warbler with
bright greenish-blue eggs. Had our cuckoo invariably displayed the above
instinctit would assuredly have been added to those which it is assumed
must all have been acquired together. The eggs of the Australian bronze
cuckoo varyaccording to Mr. Ramsayto an extraordinary degree in colour;
so that in this respectas well as in sizenatural selection might have
secured and fixed any advantageous variation.

In the case of the European cuckoothe offspring of the foster-parents are
commonly ejected from the nest within three days after the cuckoo is
hatched; and as the latter at this age is in a most helpless conditionMr.
Gould was formerly inclined to believe that the act of ejection was
performed by the foster-parents themselves. But he has now received a
trustworthy account of a young cuckoo which was actually seenwhile still
blind and not able even to hold up its own headin the act of ejecting its
foster-brothers. One of these was replaced in the nest by the observer
and was again thrown out. With respect to the means by which this strange
and odious instinct was acquiredif it were of great importance for the
young cuckooas is probably the caseto receive as much food as possible
soon after birthI can see no special difficulty in its having gradually
acquiredduring successive generationsthe blind desirethe strength
and structure necessary for the work of ejection; for those cuckoos which
had such habits and structure best developed would be the most securely
reared. The first step towards the acquisition of the proper instinct
might have been mere unintentional restlessness on the part of the young
birdwhen somewhat advanced in age and strength; the habit having been
afterwards improvedand transmitted to an earlier age. 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; or than in young snakes
acquiring in their upper jawsas Owen has remarkeda transitory sharp
tooth for cutting through the tough egg-shell. For if each part is liable
to individual variations at all agesand 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; and both cases must stand or fall
together with the whole theory of natural selection.

Some species of Molothrusa widely distinct genus of American birds
allied to our starlingshave parasitic habits like those of the cuckoo;
and the species present an interesting gradation in the perfection of their
instincts. The sexes of Molothrus badius are stated by an excellent
observerMr. Hudsonsometimes to live promiscuously together in flocks
and sometimes to pair. They either build a nest of their own or seize on
one belonging to some other birdoccasionally throwing out the nestlings
of the stranger. They either lay their eggs in the nest thus appropriated
or oddly enough build one for themselves on the top of it. They usually
sit on their own eggs and rear their own young; but Mr. Hudson says it is
probable that they are occasionally parasiticfor he has seen the young of
this species following old birds of a distinct kind and clamouring to be
fed by them. The parasitic habits of another species of Molothrusthe M.
bonariensisare much more highly developed than those of the lastbut are
still far from perfect. This birdas far as it is knowninvariably lays
its eggs in the nests of strangers; but it is remarkable that several
together sometimes commence to build an irregular untidy nest of their own
placed in singular ill-adapted situationsas on the leaves of a large
thistle. They neverhoweveras far as Mr. Hudson has ascertained
complete a nest for themselves. They often lay so many eggs--from fifteen
to twenty--in the same foster-nestthat few or none can possibly be
hatched. They havemoreoverthe extraordinary habit of pecking holes in
the eggswhether of their own species or of their foster parentswhich
they find in the appropriated nests. They drop also many eggs on the bare
groundwhich are thus wasted. A third speciesthe M. pecoris of North
Americahas acquired instincts as perfect as those of the cuckoofor it
never lays more than one egg in a foster-nestso that the young bird is
securely reared. Mr. Hudson is a strong disbeliever in evolutionbut he
appears to have been so much struck by the imperfect instincts of the
Molothrus bonariensis that he quotes my wordsand asksMust we consider
these habits, not as especially endowed or created instincts, but as small
consequences of one general law, namely, transition?

Various birdsas has already been remarkedoccasionally lay their eggs in

the nests of other birds. This habit is not very uncommon with the
Gallinaceaeand throws some light on the singular instinct of the ostrich.
In this family several hen birds unite and lay first a few eggs in one nest
and then in another; and these are hatched by the males. This instinct may
probably be accounted for by the fact of the hens laying a large number of
eggsbutas with the cuckooat intervals of two or three days. The
instincthoweverof the American ostrichas in the case of the Molothrus
bonariensishas not as yet been perfected; for a surprising number of eggs
lie strewed over the plainsso that in one day's hunting I picked up no
less than twenty lost and wasted eggs.

Many bees are parasiticand regularly lay their eggs in the nests of other
kinds of bees. This case is more remarkable than that of the cuckoo; for
these bees have not only had their instincts but their structure modified
in accordance with their parasitic habits; for they do not possess the
pollen-collecting apparatus which would have been indispensable if they had
stored up food for their own young. Some species of Sphegidae (wasp-like
insects) are likewise parasitic; and M. Fabre has lately shown good reason
for believing thatalthough the Tachytes nigra generally makes its own
burrow and stores it with paralysed prey for its own larvaeyet thatwhen
this insect finds a burrow already made and stored by another sphexit
takes advantage of the prizeand becomes for the occasion parasitic. In
this caseas with that of the Molothrus or cuckooI can see no difficulty
in natural selection making an occasional habit permanentif of advantage
to the speciesand if the insect whose nest and stored food are
feloniously appropriatedbe not thus exterminated.


This remarkable instinct was first discovered in the Formica (Polyerges)
rufescens by Pierre Hubera better observer even than his celebrated
father. This ant is absolutely dependent on its slaves; without their aid
the species would certainly become extinct in a single year. The males and
fertile females do no work of any kindand the workers or sterile females
though most energetic and courageous in capturing slavesdo no other work.
They are incapable of making their own nestsor of feeding their own
larvae. When the old nest is found inconvenientand they have to migrate
it is the slaves which determine the migrationand actually carry their
masters in their jaws. So utterly helpless are the mastersthat when
Huber shut up thirty of them without a slavebut with plenty of the food
which they like bestand with their larvae and pupae to stimulate them to
workthey did nothing; they could not even feed themselvesand many
perished of hunger. Huber then introduced a single slave (F. fusca)and
she instantly set to workfed and saved the survivors; made some cells and
tended the larvaeand put all to rights. What can be more extraordinary
than these well-ascertained facts? If we had not known of any other
slave-making antit would have been hopeless to speculate how so wonderful
an instinct could have been perfected.

Another speciesFormica sanguineawas likewise first discovered by P.
Huber to be a slave-making ant. This species is found in the southern
parts of Englandand its habits have been attended to by Mr. F. Smithof
the British Museumto whom I am much indebted for information on this and
other subjects. Although fully trusting to the statements of Huber and Mr.
SmithI tried to approach the subject in a sceptical frame of mindas any
one may well be excused for doubting the existence of so extraordinary an
instinct as that of making slaves. HenceI will give the observations
which I made in some little detail. I opened fourteen nests of F.
sanguineaand found a few slaves in all. Males and fertile females of the
slave-species (F. fusca) are found only in their own proper communities
and have never been observed in the nests of F. sanguinea. The slaves are
black and not above half the size of their red mastersso that the
contrast in their appearance is great. When the nest is slightly
disturbedthe slaves occasionally come outand like their masters are

much agitated and defend the nest: when the nest is much disturbedand
the larvae and pupae are exposedthe slaves work energetically together
with their masters in carrying them away to a place of safety. Henceit
is clear that the slaves feel quite at home. During the months of June and
Julyon three successive yearsI watched for many hours several nests in
Surrey and Sussexand never saw a slave either leave or enter a nest. As
during these monthsthe slaves are very few in numberI thought that they
might behave differently when more numerous; but Mr. Smith informs me that
he has watched the nests at various hours during MayJune and Augustboth
in Surrey and Hampshireand has never seen the slavesthough present in
large numbers in Augusteither leave or enter the nest. Hencehe
considers them as strictly household slaves. The masterson the other
handmay be constantly seen bringing in materials for the nestand food
of all kinds. During the year 1860howeverin the month of JulyI came
across a community with an unusually large stock of slavesand I observed
a few slaves mingled with their masters leaving the nestand marching
along the same road to a tall Scotch-fir treetwenty-five yards distant
which they ascended togetherprobably in search of aphides or cocci.
According to Huberwho had ample opportunities for observationthe slaves
in Switzerland habitually work with their masters in making the nestand
they alone open and close the doors in the morning and evening; andas
Huber expressly statestheir principal office is to search for aphides.
This difference in the usual habits of the masters and slaves in the two
countriesprobably depends merely on the slaves being captured in greater
numbers in Switzerland than in England.

One day I fortunately witnessed a migration of F. sanguinea from one nest
to anotherand it was a most interesting spectacle to behold the masters
carefully carrying their slaves in their jaws instead of being carried by
themas in the case of F. rufescens. Another day my attention was struck
by about a score of the slave-makers haunting the same spotand evidently
not in search of food; they approached and were vigorously repulsed by an
independent community of the slave species (F. fusca); sometimes as many as
three of these ants clinging to the legs of the slave-making F. sanguinea.
The latter ruthlessly killed their small opponents and carried their dead
bodies as food to their nesttwenty-nine yards distant; but they were
prevented from getting any pupae to rear as slaves. I then dug up a small
parcel of the pupae of F. fusca from another nestand put them down on a
bare spot near the place of combat; they were eagerly seized and carried
off by the tyrantswho perhaps fancied thatafter allthey had been
victorious in their late combat.

At the same time I laid on the same place a small parcel of the pupae of
another speciesF. flavawith a few of these little yellow ants still
clinging to the fragments of their nest. This species is sometimesthough
rarelymade into slavesas has been described by Mr. Smith. Although so
small a speciesit is very courageousand I have seen it ferociously
attack other ants. In one instance I found to my surprise an independent
community of F. flava under a stone beneath a nest of the slave-making F.
sanguinea; and when I had accidentally disturbed both neststhe little
ants attacked their big neighbours with surprising courage. Now I was
curious to ascertain whether F. sanguinea could distinguish the pupae of F.
fuscawhich they habitually make into slavesfrom those of the little and
furious F. flavawhich they rarely captureand it was evident that they
did at once distinguish them; for we have seen that they eagerly and
instantly seized the pupae of F. fuscawhereas they were much terrified
when they came across the pupaeor even the earth from the nestof F.
flavaand quickly ran away; but in about a quarter of an hourshortly
after all the little yellow ants had crawled awaythey took heart and
carried off the pupae.

One evening I visited another community of F. sanguineaand found a number
of these ants returning home and entering their nestscarrying the dead
bodies of F. fusca (showing that it was not a migration) and numerous

pupae. I traced a long file of ants burdened with bootyfor about forty
yards backto a very thick clump of heathwhence I saw the last
individual of F. sanguinea emergecarrying a pupa; but I was not able to
find the desolated nest in the thick heath. The nesthowevermust have
been close at handfor two or three individuals of F. fusca were rushing
about in the greatest agitationand one was perched motionless with its
own pupa in its mouth on the top of a spray of heathan image of despair
over its ravaged home.

Such are the factsthough they did not need confirmation by mein regard
to the wonderful instinct of making slaves. Let it be observed what a
contrast the instinctive habits of F. sanguinea present with those of the
continental F. rufescens. The latter does not build its own nestdoes not
determine its own migrationsdoes not collect food for itself or its
youngand cannot even feed itself: it is absolutely dependent on its
numerous slaves. Formica sanguineaon the other handpossesses much
fewer slavesand in the early part of the summer extremely few. The
masters determine when and where a new nest shall be formedand when they
migratethe masters carry the slaves. Both in Switzerland and England the
slaves seem to have the exclusive care of the larvaeand the masters alone
go on slave-making expeditions. In Switzerland the slaves and masters work
togethermaking and bringing materials for the nest: bothbut chiefly
the slavestend and milk as it may be calledtheir aphides; and thus both
collect food for the community. In England the masters alone usually leave
the nest to collect building materials and food for themselvestheir
slaves and larvae. So that the masters in this country receive much less
service from their slaves than they do in Switzerland.

By what steps the instinct of F. sanguinea originated I will not pretend to
conjecture. But as ants which are not slave-makerswillas I have seen
carry off pupae of other speciesif scattered near their nestsit is
possible that such pupae originally stored as food might become developed;
and the foreign ants thus unintentionally reared would then follow their
proper instinctsand do what work they could. If their presence proved
useful to the species which had seized them--if it were more advantageous
to this speciesto capture workers than to procreate them--the habit of
collecting pupaeoriginally for foodmight by natural selection be
strengthened and rendered permanent for the very different purpose of
raising slaves. When the instinct was once acquiredif carried out to a
much less extent even than in our British F. sanguineawhichas we have
seenis less aided by its slaves than the same species in Switzerland
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.


I will not here enter on minute details on this subjectbut 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 combso beautifully
adapted to its endwithout enthusiastic admiration. We hear from
mathematicians that bees have practically solved a recondite problemand
have made their cells of the proper shape to hold the greatest possible
amount of honeywith the least possible consumption of precious wax in
their construction. It has been remarked that a skilful workmanwith
fitting tools and measureswould find it very difficult to make cells of
wax of the true formthough this is effected by a crowd of bees working in
a dark hive. Granting whatever instincts you pleaseit seems at first
quite inconceivable how they can make all the necessary angles and planes
or even perceive when they are correctly made. But the difficulty is not
nearly so great as at first appears: all this beautiful work can be shown
I thinkto follow from a few simple instincts.

I was led to investigate this subject by Mr. Waterhousewho has shown that

the form of the cell stands in close relation to the presence of adjoining
cells; and the following view mayperhapsbe considered only as a
modification of his theory. Let us look to the great principle of
gradationand see whether Nature does not reveal to us her method of work.
At one end of a short series we have humble-beeswhich use their old
cocoons to hold honeysometimes adding to them short tubes of waxand
likewise making separate and very irregular rounded cells of wax. At the
other end of the series we have the cells of the hive-beeplaced in a
double layer: each cellas is well knownis an hexagonal prismwith the
basal edges of its six sides bevelled so as to join an inverted pyramidof
three rhombs. These rhombs have certain anglesand the three which form
the pyramidal base of a single cell on one side of the combenter into the
composition of the bases of three adjoining cells on the opposite side. In
the series between the extreme perfection of the cells of the hive-bee and
the simplicity of those of the humble-beewe have the cells of the Mexican
Melipona domesticacarefully described and figured by Pierre Huber. The
Melipona itself is intermediate in structure between the hive and humble
beebut more nearly related to the latter: it forms a nearly regular
waxen comb of cylindrical cellsin which the young are hatchedandin
additionsome large cells of wax for holding honey. These latter cells
are nearly spherical and of nearly equal sizesand are aggregated into an
irregular mass. But the important point to notice isthat 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;
but this is never permittedthe bees building perfectly flat walls of wax
between the spheres which thus tend to intersect. Henceeach cell
consists of an outer spherical portionand of twothreeor more flat
surfacesaccording as the cell adjoins twothree or more other cells.
When one cell rests on three other cellswhichfrom the spheres being
nearly of the same sizeis very frequently and necessarily the casethe
three flat surfaces are united into a pyramid; and this pyramidas Huber
has remarkedis manifestly a gross imitation of the three-sided pyramidal
base of the cell of the hive-bee. As in the cells of the hive-beeso
herethe three plane surfaces in any one cell necessarily enter into the
construction of three adjoining cells. It is obvious that the Melipona
saves waxand what is more importantlabourby this manner of building;
for the flat walls between the adjoining cells are not doublebut are of
the same thickness as the outer spherical portionsand yet each flat
portion forms a part of two cells.

Reflecting on this caseit occurred to me that if the Melipona had made
its spheres at some given distance from each otherand had made them of
equal sizes and had arranged them symmetrically in a double layerthe
resulting structure would have been as perfect as the comb of the hive-bee.
Accordingly I wrote to Professor Millerof Cambridgeand this geometer
has kindly read over the following statementdrawn up from his
informationand tells me that it is strictly correct:-

If a number of equal spheres be described with their centres placed in two
parallel layers; with the centre of each sphere at the distance of radius x
sqrt(2) or radius x 1.41421 (or at some lesser distance)from the centres
of the six surrounding spheres in the same layer; and at the same distance
from the centres of the adjoining spheres in the other and parallel layer;
thenif planes of intersection between the several spheres in both layers
be formedthere will result a double layer of hexagonal prisms united
together by pyramidal bases formed of three rhombs; 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. But I hear from Professor Wymanwho has made numerous careful
measurementsthat the accuracy of the workmanship of the bee has been
greatly exaggerated; so much sothat whatever the typical form of the cell
may beit is rarelyif everrealised.

Hence we may safely conclude thatif we could slightly modify the

instincts already possessed by the Meliponaand in themselves not very
wonderfulthis bee would make a structure as wonderfully perfect as that
of the hive-bee. We must suppose the Melipona to have the power of forming
her cells truly sphericaland of equal sizes; and this would not be very
surprisingseeing that she already does so to a certain extentand seeing
what perfectly cylindrical burrows many insects make in woodapparently by
turning round on a fixed point. We must suppose the Melipona to arrange
her cells in level layersas she already does her cylindrical cells; and
we must further supposeand this is the greatest difficultythat she can
somehow judge accurately at what distance to stand from her
fellow-labourers when several are making their spheres; but she is already
so far enabled to judge of distancethat she always describes her spheres
so as to intersect to a certain extent; and then she unites the points of
intersection by perfectly flat surfaces. By such modifications of
instincts which in themselves are not very wonderful--hardly more wonderful
than those which guide a bird to make its nest--I believe that the hive-bee
has acquiredthrough natural selectionher inimitable architectural

But this theory can be tested by experiment. Following the example of Mr.
TegetmeierI separated two combsand put between them a longthick
rectangular strip of wax: the bees instantly began to excavate minute
circular pits in it; and as they deepened these little pitsthey made them
wider and wider until they were converted into shallow basinsappearing to
the eye perfectly true or parts of a sphereand of about the diameter of a
cell. It was most interesting to observe thatwherever several bees had
begun to excavate these basins near togetherthey had begun their work at
such a distance from each other that by the time the basins had acquired
the above stated width (i.e. about the width of an ordinary cell)and were
in depth about one sixth of the diameter of the sphere of which they formed
a partthe rims of the basins intersected or broke into each other. As
soon as this occurredthe bees ceased to excavateand began to build up
flat walls of wax on the lines of intersection between the basinsso that
each hexagonal prism was built upon the scalloped edge of a smooth basin
instead of on the straight edges of a three-sided pyramid as in the case of
ordinary cells.

I then put into the hiveinstead of a thickrectangular piece of waxa
thin and narrowknife-edged ridgecoloured with vermilion. The bees
instantly began on both sides to excavate little basins near to each other
in the same way as before; but the ridge of wax was so thinthat the
bottoms of the basinsif they had been excavated to the same depth as in
the former experimentwould have broken into each other from the opposite
sides. The beeshoweverdid not suffer this to happenand they stopped
their excavations in due time; so that the basinsas soon as they had been
a little deepenedcame to have flat bases; and these flat basesformed by
thin little plates of the vermilion wax left ungnawedwere situatedas
far as the eye could judgeexactly along the planes of imaginary
intersection between the basins on the opposite side of the ridge of wax.
In some partsonly small portionsin other partslarge portions of a
rhombic plate were thus left between the opposed basinsbut the workfrom
the unnatural state of thingshad not been neatly performed. 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 waxin
order to have thus succeeded in leaving flat plates between the basinsby
stopping work at the planes of intersection.

Considering how flexible thin wax isI do not see that there is any
difficulty in the beeswhilst at work on the two sides of a strip of wax
perceiving when they have gnawed the wax away to the proper thinnessand
then stopping their work. 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; for I have noticed half-completed rhombs at the base of a
just-commenced cellwhich were slightly concave on one sidewhere I

suppose that the bees had excavated too quicklyand convex on the opposed
side where the bees had worked less quickly. In one well-marked instance
I put the comb back into the hiveand allowed the bees to go on working
for a short timeand again examined the celland I found that the rhombic
plate had been completedand had become PERFECTLY FLAT: it was absolutely
impossiblefrom the extreme thinness of the little platethat they could
have effected this by gnawing away the convex side; 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 planeand thus flatten it.

>From the experiment of the ridge of vermilion wax we can see thatif the
bees were to build for themselves a thin wall of waxthey could make their
cells of the proper shapeby standing at the proper distance from each
otherby excavating at the same rateand by endeavouring to make equal
spherical hollowsbut never allowing the spheres to break into each other.
Now beesas may be clearly seen by examining the edge of a growing comb
do make a roughcircumferential wall or rim all round the comb; and they
gnaw this away from the opposite sidesalways working circularly as they
deepen each cell. They do not make the whole three-sided pyramidal base of
any one cell at the same timebut only that one rhombic plate which stands
on the extreme growing marginor the two platesas the case may be; and
they never complete the upper edges of the rhombic platesuntil the
hexagonal walls are commenced. Some of these statements differ from those
made by the justly celebrated elder Huberbut I am convinced of their
accuracy; and if I had spaceI could show that they are conformable with
my theory.

Huber's statementthat the very first cell is excavated out of a little
parallel-sided wall of waxis notas far as I have seenstrictly
correct; the first commencement having always been a little hood of wax;
but I will not here enter on details. We see how important a part
excavation plays in the construction of the cells; 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 isalong the plane of intersection between two
adjoining spheres. I have several specimens showing clearly that they can
do this. Even in the rude circumferential rim or wall of wax round a
growing combflexures may sometimes be observedcorresponding in position
to the planes of the rhombic basal plates of future cells. But the rough
wall of wax has in every case to be finished offby being largely gnawed
away on both sides. The manner in which the bees build is curious; they
always make the first rough wall from ten to twenty times thicker than the
excessively thin finished wall of the cellwhich will ultimately be left.
We shall understand how they workby supposing masons first to pile up a
broad ridge of cementand then to begin cutting it away equally on both
sides near the groundtill a smoothvery thin wall is left in the middle;
the masons always piling up the cut-away cementand adding fresh cement on
the summit of the ridge. We shall thus have a thin wall steadily growing
upward but always crowned by a gigantic coping. From all the cellsboth
those just commenced and those completedbeing thus crowned by a strong
coping of waxthe bees can cluster and crawl over the comb without
injuring the delicate hexagonal walls. These wallsas Professor Miller
has kindly ascertained for mevary greatly in thickness; beingon an
average of twelve measurements made near the border of the comb1/352 of
an inch in thickness; whereas the basal rhomboidal plates are thicker
nearly in the proportion of three to twohaving a mean thicknessfrom
twenty-one measurementsof 1/229 of an inch. By the above singular manner
of buildingstrength is continually given to the combwith the utmost
ultimate economy of wax.

It seems at first to add to the difficulty of understanding how the cells
are madethat a multitude of bees all work together; one bee after working
a short time at one cell going to anotherso thatas Huber has stateda
score of individuals work even at the commencement of the first cell. I

was able practically to show this factby covering the edges of the
hexagonal walls of a single cellor the extreme margin of the
circumferential rim of a growing combwith an extremely thin layer of
melted vermilion wax; 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 placedand worked into the growing edges of the
cells all round. The work of construction seems to be a sort of balance
struck between many beesall instinctively standing at the same relative
distance from each otherall trying to sweep equal spheresand then
building upor leaving ungnawedthe planes of intersection between these
spheres. It was really curious to note in cases of difficultyas when two
pieces of comb met at an anglehow often the bees would pull down and
rebuild in different ways the same cellsometimes recurring to a shape
which they had at first rejected.

When bees have a place on which they can stand in their proper positions
for working--for instanceon a slip of woodplaced directly under the
middle of a comb growing downwardsso 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 hexagonin its strictly proper placeprojecting beyond the
other completed cells. 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 cellsand thenby striking imaginary spheresthey
can build up a wall intermediate between two adjoining spheres; butas far
as I have seenthey 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. This capacity in bees of laying down under certain circumstances a
rough wall in its proper place between two just-commenced cellsis
importantas it bears on a factwhich seems at first subversive of the
foregoing theory; namelythat the cells on the extreme margin of
wasp-combs are sometimes strictly hexagonal; but I have not space here to
enter on this subject. Nor does there seem to me any great difficulty in a
single insect (as in the case of a queen-wasp) making hexagonal cellsif
she were to work alternately on the inside and outside of two or three
cells commenced at the same timealways standing at the proper relative
distance from the parts of the cells just begunsweeping spheres or
cylindersand building up intermediate planes.

As natural selection acts only by the accumulation of slight modifications
of structure or instincteach profitable to the individual under its
conditions of lifeit may reasonably be askedhow a long and graduated
succession of modified architectural instinctsall tending towards the
present perfect plan of constructioncould 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 strengthand save much in labour
and spaceand in the materials of which they are constructed. With
respect to the formation of waxit is known that bees are often hard
pressed to get sufficient nectar; and I am informed by Mr. 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;
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. Moreovermany bees have to remain idle
for many days during the process of secretion. A large store of honey is
indispensable to support a large stock of bees during the winter; and the
security of the hive is known mainly to depend on a large number of bees
being supported. Hence the saving of wax by largely saving honeyand the
time consumed in collecting the honeymust be an important element of
success any family of bees. Of course the success of the species may be
dependent on the number of its enemiesor parasitesor on quite distinct
causesand so be altogether independent of the quantity of honey which the
bees can collect. But let us suppose that this latter circumstance
determinedas it probably often has determinedwhether a bee allied to

our humble-bees could exist in large numbers in any country; and let us
further suppose that the community lived through the winterand
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
togetherso as to intersect a little; for a wall in common even to two
adjoining cells would save some little labour and wax. Henceit would
continually be more and more advantageous to our humble-beesif they were
to make their cells more and more regularnearer togetherand aggregated
into a masslike the cells of the Melipona; for in this case a large part
of the bounding surface of each cell would serve to bound the adjoining
cellsand much labour and wax would be saved. Againfrom the same cause
it would be advantageous to the Meliponaif she were to make her cells
closer togetherand more regular in every way than at present; for then
as we have seenthe spherical surfaces would wholly disappear and be
replaced by plane surfaces; and the Melipona would make a comb as perfect
as that of the hive-bee. Beyond this stage of perfection in architecture
natural selection could not lead; for the comb of the hive-beeas far as
we can seeis absolutely perfect in economising labour and wax.

Thusas I believethe most wonderful of all known instinctsthat of the
hive-beecan be explained by natural selection having taken advantage of
numeroussuccessiveslight modifications of simpler instincts; natural
selection havingby slow degreesmore and more perfectly led the bees to
sweep equal spheres at a given distance from each other in a double layer
and to build up and excavate the wax along the planes of intersection. The
beesof courseno more knowing that they swept their spheres at one
particular distance from each otherthan they know what are the several
angles of the hexagonal prisms and of the basal rhombic plates; 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 larvaethis
being effected with the greatest possible economy of labour and wax; that
individual swarm which thus made the best cells with least labourand
least waste of honey in the secretion of waxhaving succeeded bestand
having transmitted their newly-acquired economical instincts to new swarms
which in their turn will have had the best chance of succeeding in the
struggle for existence.


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, as a modification in the one without
an immediate corresponding change in the other would have been fatal.The
force of this objection rests entirely on the assumption that the changes
in the instincts and structure are abrupt. To take as an illustration the
case of the larger titmouse(Parus major) alluded to in a previous
chapter; this bird often holds the seeds of the yew between its feet on a
branchand hammers with its beak till it gets at the kernel. Now what
special difficulty would there be in natural selection preserving all the
slight individual variations in the shape of the beakwhich were better
and better adapted to break open the seedsuntil a beak was formedas
well constructed for this purpose as that of the nuthatchat the same time
that habitor compulsionor spontaneous variations of tasteled the bird
to become more and more of a seed-eater? In this case the beak is supposed
to be slowly modified by natural selectionsubsequently tobut in
accordance withslowly changing habits or taste; but let the feet of the
titmouse vary and grow larger from correlation with the beakor from any
other unknown causeand 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. In this case a gradual change
of structure is supposed to lead to changed instinctive habits. 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.
Some birds build their nests of mudbelieved to be moistened with saliva;
and one of the swifts of North America makes its nest (as I have seen) of
sticks agglutinated with salivaand even with flakes of this substance.
Is it then very improbable that the natural selection of individual swifts
which secreted more and more salivashould 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. It must
howeverbe admitted that in many instances we cannot conjecture whether it
was instinct or structure which first varied.

No doubt many instincts of very difficult explanation could be opposed to
the theory of natural selection--casesin which we cannot see how an
instinct could have originated; casesin which no intermediate gradations
are known to exist; cases of instincts of such trifling importancethat
they could hardly have been acted on by natural selection; 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 progenitorand consequently must believe that they were
independently acquired through natural selection. I will not here enter on
these several casesbut will confine myself to one special difficulty
which at first appeared to me insuperableand actually fatal to the whole
theory. I allude to the neuters or sterile females in insect communities:
for these neuters often differ widely in instinct and in structure from
both the males and fertile femalesand yetfrom being sterilethey
cannot propagate their kind.

The subject well deserves to be discussed at great lengthbut I will here
take only a single casethat of working or sterile ants. How the workers
have been rendered sterile is a difficulty; but not much greater than that
of any other striking modification of structure; for it can be shown that
some insects and other articulate animals in a state of nature occasionally
become sterile; and if such insects had been socialand it had been
profitable to the community that a number should have been annually born
capable of workbut incapable of procreationI can see no especial
difficulty in this having been effected through natural selection. But I
must pass over this preliminary difficulty. The great difficulty lies in
the working ants differing widely from both the males and the fertile
females in structureas in the shape of the thoraxand in being destitute
of wings and sometimes of eyesand in instinct. As far as instinct alone
is concernedthe wonderful difference in this respect between the workers
and the perfect females would have been better exemplified by the hive-bee.
If a working ant or other neuter insect had been an ordinary animalI
should have unhesitatingly assumed that all its characters had been slowly
acquired through natural selection; namelyby individuals having been born
with slight profitable modificationswhich were inherited by the
offspringand that these again varied and again were selectedand so
onwards. But with the working ant we have an insect differing greatly from
its parentsyet absolutely sterile; so that it could never have
transmitted successively acquired modifications of structure or instinct to
its progeny. It may well be asked how it is possible to reconcile this
case with the theory of natural selection?

Firstlet it be remembered that we have innumerable instancesboth in our
domestic productions and in those in a state of natureof all sorts of
differences of inherited structure which are correlated with certain ages
and with either sex. We have differences correlated not only with one sex
but with that short period when the reproductive system is activeas in
the nuptial plumage of many birdsand in the hooked jaws of the male
salmon. We have even slight differences in the horns of different breeds
of cattle in relation to an artificially imperfect state of the male sex;
for oxen of certain breeds have longer horns than the oxen of other breeds
relatively to the length of the horns in both the bulls and cows of these
same breeds. HenceI can see no great difficulty in any character

becoming correlated with the sterile condition of certain members of insect
communities; the difficulty lies in understanding how such correlated
modifications of structure could have been slowly accumulated by natural

This difficultythough appearing insuperableis lessenedoras I
believedisappearswhen it is remembered that selection may be applied to
the familyas well as to the individualand may thus gain the desired
end. Breeders of cattle wish the flesh and fat to be well marbled
together. An animal thus characterized has been slaughteredbut the
breeder has gone with confidence to the same stock and has succeeded. Such
faith may be placed in the power of selection that a breed of cattle
always yielding oxen with extraordinarily long hornscouldit is
probablebe formed by carefully watching which individual bulls and cows
when matchedproduced oxen with the longest horns; and yet no one ox would
ever have propagated its kind. Here is a better and real illustration:
According to M. Verlotsome varieties of the double annual stockfrom
having been long and carefully selected to the right degreealways produce
a large proportion of seedlings bearing double and quite sterile flowers
but they likewise yield some single and fertile plants. These latterby
which alone the variety can be propagatedmay be compared with the fertile
male and female antsand the double sterile plants with the neuters of the
same community. As with the varieties of the stockso with social
insectsselection has been applied to the familyand not to the
individualfor the sake of gaining a serviceable end. Hencewe may
conclude that slight modifications of structure or of instinctcorrelated
with the sterile condition of certain members of the communityhave proved
advantageous; consequently the fertile males and females have flourished
and transmitted to their fertile offspring a tendency to produce sterile
members with the same modifications. This process must have been repeated
many timesuntil 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.

But we have not as yet touched on the acme of the difficulty; namelythe
fact that the neuters of several ants differnot only from the fertile
females and malesbut from each othersometimes to an almost incredible
degreeand are thus divided into two or even three castes. The castes
moreoverdo not generally graduate into each otherbut are perfectly well
defined; being as distinct from each other as are any two species of the
same genusor rather as any two genera of the same family. Thusin
Ecitonthere are working and soldier neuterswith jaws and instincts
extraordinarily different: in Cryptocerusthe workers of one caste alone
carry a wonderful sort of shield on their headsthe use of which is quite
unknown: in the Mexican Myrmecocystusthe workers of one caste never
leave the nest; they are fed by the workers of another casteand they have
an enormously developed abdomen which secretes a sort of honeysupplying
the place of that excreted by the aphidesor the domestic cattle as they
may be calledwhich our European ants guard and imprison.

It will indeed be thought that I have an overweening confidence in the
principle of natural selectionwhen I do not admit that such wonderful and
well-established facts at once annihilate the theory. In the simpler case
of neuter insects all of one castewhichas I believehave been rendered
different from the fertile males and females through natural selectionwe
may conclude from the analogy of ordinary variationsthat the successive
slightprofitable modifications did not first arise in all the neuters in
the same nestbut in some few alone; and that by the survival of the
communities with females which produced most neuters having the
advantageous modificationall the neuters ultimately came to be thus
characterized. According to this view we ought occasionally to find in the
same nest neuter-insectspresenting gradations of structure; and this we
do findeven not rarelyconsidering how few neuter-insects out of Europe
have been carefully examined. Mr. F. Smith has shown that the neuters of

several British ants differ surprisingly from each other in size and
sometimes in colour; 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. It sometimes happens that the larger or the
smaller sized workers are the most numerous; or that both large and small
are numerouswhile those of an intermediate size are scanty in numbers.
Formica flava has larger and smaller workerswith some few of intermediate
size; andin this speciesas Mr. F. Smith has observedthe larger
workers have simple eyes (ocelli)whichthough smallcan be plainly
distinguishedwhereas the smaller workers have their ocelli rudimentary.
Having carefully dissected several specimens of these workersI can affirm
that the eyes are far more rudimentary in the smaller workers than can be
accounted for merely by their proportionately lesser size; and I fully
believethough I dare not assert so positivelythat the workers of
intermediate size have their ocelli in an exactly intermediate condition.
So that here we have two bodies of sterile workers in the same nest
differing not only in sizebut in their organs of visionyet connected by
some few members in an intermediate condition. I may digress by adding
that if the smaller workers had been the most useful to the communityand
those males and females had been continually selectedwhich produced more
and more of the smaller workersuntil all the workers were in this
condition; we should then have had a species of ant with neuters in nearly
the same condition as those of Myrmica. For the workers of Myrmica have
not even rudiments of ocellithough the male and female ants of this genus
have well-developed ocelli.

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 speciesthat I gladly availed myself of Mr. F. Smith's
offer of numerous specimens from the same nest of the driver ant (Anomma)
of West Africa. The reader will perhaps best appreciate the amount of
difference in these workers by my givingnot the actual measurementsbut
a strictly accurate illustration: the difference was the same as if we
were to see a set of workmen building a houseof whom many were five feet
four inches highand many sixteen feet high; but we must in addition
suppose that the larger workmen had heads four instead of three times as
big as those of the smaller menand jaws nearly five times as big. The
jawsmoreoverof the working ants of the several sizes differed
wonderfully in shapeand in the form and number of the teeth. But the
important fact for us is thatthough the workers can be grouped into
castes of different sizesyet they graduate insensibly into each otheras
does the widely-different structure of their jaws. I speak confidently on
this latter pointas Sir J. Lubbock made drawings for mewith the camera
lucidaof the jaws which I dissected from the workers of the several
sizes. Mr. Batesin his interesting "Naturalist on the Amazons has
described analogous cases.

With these facts before me, I believe that natural selection, by acting on
the fertile ants or parents, could form a species which should regularly
produce neuters, all of large size with one form of jaw, or all of small
size with widely different jaws; or lastly, and this is the greatest
difficulty, one set of workers of one size and structure, and
simultaneously another set of workers of a different size and structure; a
graduated series having first been formed, as in the case of the driver
ant, and then the extreme forms having been produced in greater and greater
numbers, through the survival of the parents which generated them, until
none with an intermediate structure were produced.

An analogous explanation has been given by Mr. Wallace, of the equally
complex case, of certain Malayan butterflies regularly appearing under two
or even three distinct female forms; and by Fritz Muller, of certain
Brazilian crustaceans likewise appearing under two widely distinct male
forms. But this subject need not here be discussed.

I have now explained how, I believe, the wonderful fact of two distinctly
defined castes of sterile workers existing in the same nest, both widely
different from each other and from their parents, has originated. We can
see how useful their production may have been to a social community of
ants, on the same principle that the division of labour is useful to
civilised man. Ants, however, work by inherited instincts and by inherited
organs or tools, while man works by acquired knowledge and manufactured
instruments. But I must confess, that, with all my faith in natural
selection, I should never have anticipated that this principle could have
been efficient in so high a degree, had not the case of these neuter
insects led me to this conclusion. I have, therefore, discussed this case,
at some little but wholly insufficient length, in order to show the power
of natural selection, and likewise because this is by far the most serious
special difficulty which my theory has encountered. The case, also, is
very interesting, as it proves that with animals, as with plants, any
amount of modification may be effected by the accumulation of numerous,
slight, spontaneous variations, which are in any way profitable, without
exercise or habit having been brought into play. For peculiar habits,
confined to the workers of sterile females, however long they might be
followed, could not possibly affect the males and fertile females, which
alone leave descendants. I am surprised that no one has advanced this
demonstrative case of neuter insects, against the well-known doctrine of
inherited habit, as advanced by Lamarck.


I have endeavoured in this chapter briefly to show that the mental
qualities of our domestic animals vary, and that the variations are
inherited. Still more briefly I have attempted to show that instincts vary
slightly in a state of nature. No one will dispute that instincts are of
the highest importance to each animal. Therefore, there is no real
difficulty, under changing conditions of life, in natural selection
accumulating to any extent slight modifications of instinct which are in
any way useful. In many cases habit or use and disuse have probably come
into play. I do not pretend that the facts given in this chapter
strengthen in any great degree my theory; but none of the cases of
difficulty, to the best of my judgment, annihilate it. On the other hand,
the fact that instincts are not always absolutely perfect and are liable to
mistakes; that no instinct can be shown to have been produced for the good
of other animals, though animals take advantage of the instincts of others;
that the canon in natural history, of Natura non facit saltum is
applicable to instincts as well as to corporeal structure, and is plainly
explicable on the foregoing views, but is otherwise inexplicable--all tend
to corroborate the theory of natural selection.

This theory is also strengthened by some few other facts in regard to
instincts; as by that common case of closely allied, but distinct, species,
when inhabiting distant parts of the world and living under considerably
different conditions of life, yet often retaining nearly the same
instincts. For instance, we can understand, on the principle of
inheritance, how it is that the thrush of tropical South America lines its
nest with mud, in the same peculiar manner as does our British thrush; 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,
with only a small hole left in the plaster through which the males feed
them and their young when hatched; how it is that the male wrens
(Troglodytes) of North America, build cock-nests to roost in, like the
males of our Kitty-wrens,--a habit wholly unlike that of any other known
bird. Finally, it may not be a logical deduction, but to my imagination it
is far more satisfactory to look at such instincts as the young cuckoo
ejecting its foster-brothers, ants making slaves, the larvae of
ichneumonidae feeding within the live bodies of caterpillars, not as
specially endowed or created instincts, but as small consequences of one
general law leading to the advancement of all organic beings--namely,

multiply, vary, let the strongest live and the weakest die.



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

The view commonly entertained by naturalists is that species, when
intercrossed, have been specially endowed with sterility, in order to
prevent their confusion. This view certainly seems at first highly
probable, for species living together could hardly have been kept distinct
had they been capable of freely crossing. The subject is in many ways
important for us, more especially as the sterility of species when first
crossed, and that of their hybrid offspring, cannot have been acquired, as
I shall show, by the preservation of successive profitable degrees of
sterility. It is an incidental result of differences in the reproductive
systems of the parent-species.

In treating this subject, two classes of facts, to a large extent
fundamentally different, have generally been confounded; namely, the
sterility of species when first crossed, and the sterility of the hybrids
produced from them.

Pure species have of course their organs of reproduction in a perfect
condition, yet when intercrossed they produce either few or no offspring.
Hybrids, on the other hand, have their reproductive organs functionally
impotent, as may be clearly seen in the state of the male element in both
plants and animals; though the formative organs themselves are perfect in
structure, as far as the microscope reveals. In the first case the two
sexual elements which go to form the embryo are perfect; in the second case
they are either not at all developed, or are imperfectly developed. This
distinction is important, when the cause of the sterility, which is common
to the two cases, has to be considered. The distinction probably has been
slurred over, owing to the sterility in both cases being looked on as a
special endowment, beyond the province of our reasoning powers.

The fertility of varieties, that is of the forms known or believed to be
descended from common parents, when crossed, and likewise the fertility of
their mongrel offspring, is, with reference to my theory, of equal
importance with the sterility of species; for it seems to make a broad and
clear distinction between varieties and species.


First, for the sterility of species when crossed and of their hybrid
offspring. It is impossible to study the several memoirs and works of
those two conscientious and admirable observers, Kolreuter and Gartner, who
almost devoted their lives to this subject, without being deeply impressed
with the high generality of some degree of sterility. Kolreuter makes the
rule universal; but then he cuts the knot, for in ten cases in which he
found two forms, considered by most authors as distinct species, quite
fertile together, he unhesitatingly ranks them as varieties. Gartner,
also, makes the rule equally universal; and he disputes the entire

fertility of Kolreuter's ten cases. But in these and in many other cases,
Gartner is obliged carefully to count the seeds, in order to show that
there is any degree of sterility. He always compares the maximum number of
seeds produced by two species when first crossed, and the maximum produced
by their hybrid offspring, with the average number produced by both pure
parent-species in a state of nature. But causes of serious error here
intervene: a plant, to be hybridised, must be castrated, and, what is
often more important, must be secluded in order to prevent pollen being
brought to it by insects from other plants. Nearly all the plants
experimented on by Gartner were potted, and were kept in a chamber in his
house. That these processes are often injurious to the fertility of a
plant cannot be doubted; for Gartner gives in his table about a score of
cases of plants which he castrated, and artificially fertilised with their
own pollen, and (excluding all cases such as the Leguminosae, in which
there is an acknowledged difficulty in the manipulation) half of these
twenty plants had their fertility in some degree impaired. Moreover, as
Gartner repeatedly crossed some forms, such as the common red and blue
pimpernels (Anagallis arvensis and coerulea), which the best botanists rank
as varieties, and found them absolutely sterile, we may doubt whether many
species are really so sterile, when intercrossed, as he believed.

It is certain, on the one hand, that the sterility of various species when
crossed is so different in degree and graduates away so insensibly, and, on
the other hand, 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. I think no
better evidence of this can be required than that the two most experienced
observers who have ever lived, namely Kolreuter and Gartner, arrived at
diametrically opposite conclusions in regard to some of the very same
forms. 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, with the evidence from fertility adduced by different
hybridisers, or by the same observer from experiments made during different
years. It can thus be shown that neither sterility nor fertility affords
any certain distinction between species and varieties. The evidence from
this source graduates away, and is doubtful in the same degree as is the
evidence derived from other constitutional and structural differences.

In regard to the sterility of hybrids in successive generations; though
Gartner was enabled to rear some hybrids, carefully guarding them from a
cross with either pure parent, for six or seven, and in one case for ten
generations, yet he asserts positively that their fertility never
increases, but generally decreases greatly and suddenly. With respect to
this decrease, it may first be noticed that when any deviation in structure
or constitution is common to both parents, this is often transmitted in an
augmented degree to the offspring; and both sexual elements in hybrid
plants are already affected in some degree. But I believe that their
fertility has been diminished in nearly all these cases by an independent
cause, namely, by too close interbreeding. I have made so many experiments
and collected so many facts, showing on the one hand that an occasional
cross with a distinct individual or variety increases the vigour and
fertility of the offspring, and on the other hand that very close
interbreeding lessens their vigour and fertility, that I cannot doubt the
correctness of this conclusion. Hybrids are seldom raised by
experimentalists in great numbers; and as the parent-species, or other
allied hybrids, generally grow in the same garden, the visits of insects
must be carefully prevented during the flowering season: hence hybrids, if
left to themselves, will generally be fertilised during each generation by
pollen from the same flower; and this would probably be injurious to their
fertility, already lessened by their hybrid origin. I am strengthened in
this conviction by a remarkable statement repeatedly made by Gartner,
namely, that if even the less fertile hybrids be artificially fertilised
with hybrid pollen of the same kind, their fertility, notwithstanding the

frequent ill effects from manipulation, sometimes decidedly increases, and
goes on increasing. Now, in the process of artificial fertilisation,
pollen is as often taken by chance (as I know from my own experience) from
the anthers of another flower, as from the anthers of the flower itself
which is to be fertilised; so that a cross between two flowers, though
probably often on the same plant, would be thus effected. Moreover,
whenever complicated experiments are in progress, so careful an observer as
Gartner would have castrated his hybrids, and this would have insured in
each generation a cross with pollen from a distinct flower, either from the
same plant or from another plant of the same hybrid nature. And thus, the
strange fact of an increase of fertility in the successive generations of
ARTIFICIALLY FERTILISED hybrids, in contrast with those spontaneously selffertilised,
may, as I believe, be accounted for by too close interbreeding
having been avoided.

Now let us turn to the results arrived at by a third most experienced
hybridiser, namely, the Hon. and Rev. W. Herbert. 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. The difference in their
results may, I think, be in part accounted for by Herbert's great
horticultural skill, and by his having hot-houses at his command. Of his
many important statements I will here give only a single one as an example,
namely, that every ovule in a pod of Crinum capense fertilised by C.
revolutum produced a plantwhich I never saw to occur in a case of its
natural fecundation." So that here we have perfector even more than
commonly perfect fertilityin a first cross between two distinct species.

This case of the Crinum leads me to refer to a singular factnamelythat
individual plants of certain species of LobeliaVerbascum and Passiflora
can easily be fertilised by the pollen from a distinct speciesbut not by
pollen from the same plantthough this pollen can be proved to be
perfectly sound by fertilising other plants or species. In the genus
Hippeastrumin Corydalis as shown by Professor Hildebrandin various
orchids as shown by Mr. Scott and Fritz Mullerall the individuals are in
this peculiar condition. So that with some speciescertain abnormal
individualsand in other species all the individualscan actually be
hybridised much more readily than they can be fertilised by pollen from the
same individual plant! To give one instancea bulb of Hippeastrum aulicum
produced four flowers; three were fertilised by Herbert with their own
pollenand 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, and after a
few days perished entirely, whereas the pod impregnated by the pollen of
the hybrid made vigorous growth and rapid progress to maturity, and bore
good seed, which vegetated freely.Mr. Herbert tried similar experiments
during many yearsand always with the same result. These cases serve to
show on what slight and mysterious causes the lesser or greater fertility
of a species sometimes depends.

The practical experiments of horticulturiststhough not made with
scientific precisiondeserve some notice. It is notorious in how
complicated a manner the species of PelargoniumFuchsiaCalceolaria
PetuniaRhododendronetc.have been crossedyet many of these hybrids
seed freely. For instanceHerbert asserts that a hybrid from Calceolaria
integrifolia and plantagineaspecies most widely dissimilar in general
habitreproduces itself as perfectly as if it had been a natural species
from the mountains of Chile.I have taken some pains to ascertain the
degree of fertility of some of the complex crosses of Rhododendronsand I
am assured that many of them are perfectly fertile. Mr. C. Noblefor
instanceinforms me that he raises stocks for grafting from a hybrid
between Rhod. ponticum and catawbienseand that this hybrid "seeds as
freely as it is possible to imagine." Had hybridswhen fairly treated

always gone on decreasing in fertility in each successive generationas
Gartner believed to be the casethe fact would have been notorious to
nurserymen. Horticulturists raise large beds of the same hybridand such
alone are fairly treatedfor by insect agency the several individuals are
allowed to cross freely with each otherand the injurious influence of
close interbreeding is thus prevented. Any one may readily convince
himself of the efficiency of insect agency by examining the flowers of the
more sterile kinds of hybrid Rhododendronswhich produce no pollenfor he
will find on their stigmas plenty of pollen brought from other flowers.

In regard to animalsmuch fewer experiments have been carefully tried than
with plants. If our systematic arrangements can be trustedthat isif
the genera of animals are as distinct from each other as are the genera of
plantsthen we may infer that animals more widely distinct in the scale of
nature can be crossed more easily than in the case of plants; but the
hybrids themselves areI thinkmore sterile. It shouldhoweverbe
borne in mind thatowing to few animals breeding freely under confinement
few experiments have been fairly tried: for instancethe canary-bird has
been crossed with nine distinct species of finchesbutas not one of
these breeds freely in confinementwe have no right to expect that the
first crosses between them and the canaryor that their hybridsshould be
perfectly fertile. Againwith respect to the fertility in successive
generations of the more fertile hybrid animalsI hardly know of an
instance in which two families of the same hybrid have been raised at the
same time from different parentsso as to avoid the ill effects of close
interbreeding. On the contrarybrothers and sisters have usually been
crossed in each successive generationin opposition to the constantly
repeated admonition of every breeder. And in this caseit is not at all
surprising that the inherent sterility in the hybrids should have gone on

Although I know of hardly any thoroughly well-authenticated cases of
perfectly fertile hybrid animalsI have reason to believe that the hybrids
from Cervulus vaginalis and Reevesiiand from Phasianus colchicus with P.
torquatusare perfectly fertile. M. Quatrefages states that the hybrids
from two moths (Bombyx cynthia and arrindia) were proved in Paris to be
fertile inter se for eight generations. It has lately been asserted that
two such distinct species as the hare and rabbitwhen they can be got to
breed togetherproduce offspringwhich are highly fertile when crossed
with one of the parent-species. The hybrids from the common and Chinese
geese (A. cygnoides)species which are so different that they are
generally ranked in distinct generahave often bred in this country with
either pure parentand in one single instance they have bred inter se.
This was effected by Mr. Eytonwho raised two hybrids from the same
parentsbut from different hatches; and from these two birds he raised no
less than eight hybrids (grandchildren of the pure geese) from one nest.
In Indiahoweverthese cross-bred geese must be far more fertile; for I
am assured by two eminently capable judgesnamely Mr. Blyth and Captain
Huttonthat whole flocks of these crossed geese are kept in various parts
of the country; and as they are kept for profitwhere neither pure
parent-species existsthey must certainly be highly or perfectly fertile.

With our domesticated animalsthe various races when crossed together are
quite fertile; yet in many cases they are descended from two or more wild
species. From this fact we must conclude either that the aboriginal
parent-species at first produced perfectly fertile hybridsor that the
hybrids subsequently reared under domestication became quite fertile. This
latter alternativewhich was first propounded by Pallasseems by far the
most probableand canindeedhardly be doubted. It isfor instance
almost certain that our dogs are descended from several wild stocks; yet
with perhaps the exception of certain indigenous domestic dogs of South
Americaall are quite fertile together; but analogy makes me greatly
doubtwhether the several aboriginal species would at first have freely
bred together and have produced quite fertile hybrids. So again I have

lately acquired decisive evidence that the crossed offspring from the
Indian humped and common cattle are inter se perfectly fertile; and from
the observations by Rutimeyer on their important osteological differences
as well as from those by Mr. Blyth on their differences in habitsvoice
constitutionetc.these two forms must be regarded as good and distinct
species. The same remarks may be extended to the two chief races of the
pig. We mustthereforeeither give up the belief of the universal
sterility of species when crossed; or we must look at this sterility in
animalsnot as an indelible characteristicbut as one capable of being
removed by domestication.

Finallyconsidering all the ascertained facts on the intercrossing of
plants and animalsit may be concluded that some degree of sterilityboth
in first crosses and in hybridsis an extremely general result; but that
it cannotunder our present state of knowledgebe considered as
absolutely universal.


We will now consider a little more in detail the laws governing the
sterility of first crosses and of hybrids. Our chief object will be to see
whether or not these laws indicate that species have been specially endowed
with this qualityin order to prevent their crossing and blending together
in utter confusion. The following conclusions are drawn up chiefly from
Gartner's admirable work on the hybridisation of plants. I have taken much
pains to ascertain how far they apply to animalsandconsidering how
scanty our knowledge is in regard to hybrid animalsI have been surprised
to find how generally the same rules apply to both kingdoms.

It has been already remarkedthat the degree of fertilityboth of first
crosses and of hybridsgraduates from zero to perfect fertility. It is
surprising in how many curious ways this gradation can be shown; but only
the barest outline of the facts can here be given. When pollen from a
plant of one family is placed on the stigma of a plant of a distinct
familyit exerts no more influence than so much inorganic dust. From this
absolute zero of fertilitythe pollen of different species applied to the
stigma of some one species of the same genusyields a perfect gradation in
the number of seeds producedup to nearly complete or even quite complete
fertility; andas we have seenin certain abnormal caseseven to an
excess of fertilitybeyond that which the plant's own pollen produces. So
in hybrids themselvesthere are some which never have producedand
probably never would produceeven with the pollen of the pure parentsa
single fertile seed: but in some of these cases a first trace of fertility
may be detectedby the pollen of one of the pure parent-species causing
the flower of the hybrid to wither earlier than it otherwise would have
done; and the early withering of the flower is well known to be a sign of
incipient fertilisation. From this extreme degree of sterility we have
self-fertilised hybrids producing a greater and greater number of seeds up
to perfect fertility.

The hybrids raised from two species which are very difficult to crossand
which rarely produce any offspringare generally very sterile; but the
parallelism between the difficulty of making a first crossand the
sterility of the hybrids thus produced--two classes of facts which are
generally confounded together--is by no means strict. There are many
casesin which two pure speciesas in the genus Verbascumcan be united
with unusual facilityand produce numerous hybrid offspringyet these
hybrids are remarkably sterile. On the other handthere are species which
can be crossed very rarelyor with extreme difficultybut the hybrids
when at last producedare very fertile. Even within the limits of the
same genusfor instance in Dianthusthese two opposite cases occur.

The fertilityboth of first crosses and of hybridsis more easily
affected by unfavourable conditionsthan is that of pure species. But the

fertility of first crosses is likewise innately variable; for it is not
always the same in degree when the same two species are crossed under the
same circumstances; it depends in part upon the constitution of the
individuals which happen to have been chosen for the experiment. So it is
with hybridsfor their degree of fertility is often found to differ
greatly in the several individuals raised from seed out of the same capsule
and exposed to the same conditions.

By the term systematic affinity is meantthe general resemblance between
species in structure and constitution. Now the fertility of first crosses
and of the hybrids produced from themis largely governed by their
systematic affinity. This is clearly shown by hybrids never having been
raised between species ranked by systematists in distinct families; and on
the other handby very closely allied species generally uniting with
facility. But the correspondence between systematic affinity and the
facility of crossing is by no means strict. A multitude of cases could be
given of very closely allied species which will not uniteor only with
extreme difficulty; and on the other hand of very distinct species which
unite with the utmost facility. In the same family there may be a genus
as Dianthusin which very many species can most readily be crossed; and
another genusas Silenein which the most persevering efforts have failed
to produce between extremely close species a single hybrid. Even within
the limits of the same genuswe meet with this same difference; for
instancethe many species of Nicotiana have been more largely crossed than
the species of almost any other genus; but Gartner found that N. acuminata
which is not a particularly distinct speciesobstinately failed to
fertiliseor to be fertilisedby no less than eight other species of
Nicotiana. Many analogous facts could be given.

No one has been able to point out what kind or what amount of difference
in any recognisable characteris sufficient to prevent two species
crossing. It can be shown that plants most widely different in habit and
general appearanceand having strongly marked differences in every part of
the flowereven in the pollenin the fruitand in the cotyledonscan be
crossed. Annual and perennial plantsdeciduous and evergreen trees
plants inhabiting different stations and fitted for extremely different
climatescan often be crossed with ease.

By a reciprocal cross between two speciesI mean the casefor instance
of a female-ass being first crossed by a stallionand then a mare by a
male-ass: these two species may then be said to have been reciprocally
crossed. There is often the widest possible difference in the facility of
making reciprocal crosses. Such cases are highly importantfor they prove
that the capacity in any two species to cross is often completely
independent of their systematic affinitythat is of any difference in
their structure or constitutionexcepting in their reproductive systems.
The diversity of the result in reciprocal crosses between the same two
species was long ago observed by Kolreuter. To give an instance:
Mirabilis jalapa can easily be fertilised by the pollen of M. longiflora
and the hybrids thus produced are sufficiently fertile; but Kolreuter tried
more than two hundred timesduring eight following yearsto fertilise
reciprocally M. longiflora with the pollen of M. jalapaand utterly
failed. Several other equally striking cases could be given. Thuret has
observed the same fact with certain sea-weeds or Fuci. Gartnermoreover
found that this difference of facility in making reciprocal crosses is
extremely common in a lesser degree. He has observed it even between
closely related forms (as Matthiola annua and glabra) which many botanists
rank only as varieties. It is also a remarkable fact that hybrids raised
from reciprocal crossesthough of course compounded of the very same two
speciesthe one species having first been used as the father and then as
the motherthough they rarely differ in external charactersyet generally
differ in fertility in a smalland occasionally in a high degree.

Several other singular rules could be given from Gartner: for instance

some species have a remarkable power of crossing with other species; other
species of the same genus have a remarkable power of impressing their
likeness on their hybrid offspring; but these two powers do not at all
necessarily go together. There are certain hybrids whichinstead of
havingas is usualan intermediate character between their two parents
always closely resemble one of them; and such hybridsthough externally so
like one of their pure parent-speciesare with rare exceptions extremely
sterile. So again among hybrids which are usually intermediate in
structure between their parentsexceptional and abnormal individuals
sometimes are bornwhich closely resemble one of their pure parents; and
these hybrids are almost always utterly sterileeven when the other
hybrids raised from seed from the same capsule have a considerable degree
of fertility. These facts show how completely the fertility of a hybrid
may be independent of its external resemblance to either pure parent.

Considering the several rules now givenwhich govern the fertility of
first crosses and of hybridswe see that when formswhich must be
considered as good and distinct speciesare unitedtheir fertility
graduates from zero to perfect fertilityor even to fertility under
certain conditions in excess; that their fertilitybesides being eminently
susceptible to favourable and unfavourable conditionsis innately
variable; that it is by no means always the same in degree in the first
cross and in the hybrids produced from this cross; that the fertility of
hybrids is not related to the degree in which they resemble in external
appearance either parent; and lastlythat 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. This latter statement is
clearly proved by the difference in the result of reciprocal crosses
between the same two speciesforaccording as the one species or the
other is used as the father or the motherthere is generally some
differenceand occasionally the widest possible differencein the
facility of effecting an union. The hybridsmoreoverproduced from
reciprocal crosses often differ in fertility.

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. For why should the sterility be so extremely
different in degreewhen various species are crossedall of which we must
suppose it would be equally important to keep from blending together? Why
should the degree of sterility be innately variable in the individuals of
the same species? Why should some species cross with facility and yet
produce very sterile hybrids; and other species cross with extreme
difficultyand 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? Whyit may even be askedhas the production of hybrids
been permitted? To grant to species the special power of producing
hybridsand then to stop their further propagation by different degrees of
sterilitynot strictly related to the facility of the first union between
their parentsseems a strange arrangement.

The foregoing rules and factson the other handappear to me clearly to
indicate that the sterilityboth of first crosses and of hybridsis
simply incidental or dependent on unknown differences in their reproductive
systems; the differences being of so peculiar and limited a naturethat
in reciprocal crosses between the same two speciesthe male sexual element
of the one will often freely act on the female sexual element of the other
but not in a reversed direction. It will be advisable to explain a little
more fullyby an examplewhat I mean by sterility being incidental on
other differencesand 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 natureI presume that no one will suppose that this
capacity is a SPECIALLY endowed qualitybut will admit that it is
incidental on differences in the laws of growth of the two plants. We can
sometimes see the reason why one tree will not take on another from

differences in their rate of growthin the hardness of their woodin the
period of the flow or nature of their sapetc.; but in a multitude of
cases we can assign no reason whatever. Great diversity in the size of two
plantsone being woody and the other herbaceousone being evergreen and
the other deciduousand adaptation to widely different climatesdoes not
always prevent the two grafting together. As in hybridisationso with
graftingthe capacity is limited by systematic affinityfor no one has
been able to graft together trees belonging to quite distinct families;
andon the other handclosely allied species and varieties of the same
speciescan usuallybut not invariablybe grafted with ease. But this
capacityas in hybridisationis by no means absolutely governed by
systematic affinity. Although many distinct genera within the same family
have been grafted togetherin other cases species of the same genus will
not take on each other. The pear can be grafted far more readily on the
quincewhich is ranked as a distinct genusthan on the applewhich is a
member of the same genus. Even different varieties of the pear take with
different degrees of facility on the quince; so do different varieties of
the apricot and peach on certain varieties of the plum.

As Gartner found that there was sometimes an innate difference in different
INDIVIDUALS of the same two species in crossing; so Sagaret believes this
to be the case with different individuals of the same two species in being
grafted together. As in reciprocal crossesthe facility of effecting an
union is often very far from equalso it sometimes is in grafting. The
common gooseberryfor instancecannot be grafted on the currantwhereas
the currant will takethough with difficultyon the gooseberry.

We have seen that the sterility of hybrids which have their reproductive
organs in an imperfect conditionis a different case from the difficulty
of uniting two pure specieswhich have their reproductive organs perfect;
yet these two distinct classes of cases run to a large extent parallel.
Something analogous occurs in grafting; for Thouin found that three species
of Robiniawhich seeded freely on their own rootsand which could be
grafted with no great difficulty on a fourth specieswhen thus grafted
were rendered barren. On the other handcertain species of Sorbuswhen
grafted on other speciesyielded twice as much fruit as when on their own
roots. We are reminded by this latter fact of the extraordinary cases of
HippeastrumPassifloraetc.which seed much more freely when fertilised
with the pollen of a distinct species than when fertilised with pollen from
the same plant.

We thus see thatalthough 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 reproductionyet that there is a rude degree of
parallelism in the results of grafting and of crossing distinct 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 systemsso I believe that the still more
complex laws governing the facility of first crosses are incidental on
unknown differences in their reproductive systems. These differences in
both cases followto a certain extentas might have been expected
systematic affinityby which term every kind of resemblance and
dissimilarity between organic beings is attempted to be expressed. 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;
although in the case of crossingthe difficulty is as important for the
endurance and stability of specific forms as in the case of grafting it is
unimportant for their welfare.


At one time it appeared to me probableas it has to othersthat the
sterility of first crosses and of hybrids might have been slowly acquired
through the natural selection of slightly lessened degrees of fertility

whichlike any other variationspontaneously appeared in certain
individuals of one variety when crossed with those of another variety. For
it would clearly be advantageous to two varieties or incipient species if
they could be kept from blendingon the same principle thatwhen man is
selecting at the same time two varietiesit is necessary that he should
keep them separate. In the first placeit may be remarked that species
inhabiting distinct regions are often sterile when crossed; now it could
clearly have been of no advantage to such separated species to have been
rendered mutually sterileand consequently this could not have been
effected through natural selection; but it may perhaps be arguedthatif
a species was rendered sterile with some one compatriotsterility with
other species would follow as a necessary contingency. In the second
placeit is almost as much opposed to the theory of natural selection as
to that of special creationthat in reciprocal crosses the male element of
one form should have been rendered utterly impotent on a second formwhile
at the same time the male element of this second form is enabled freely to
fertilise the first form; for this peculiar state of the reproductive
system could hardly have been advantageous to either species.

In considering the probability of natural selection having come into
actionin rendering species mutually sterilethe greatest difficulty will
be found to lie in the existence of many graduated stepsfrom slightly
lessened fertility to absolute sterility. It may be admitted that it would
profit an incipient speciesif it were rendered in some slight degree
sterile when crossed with its parent form or with some other variety; for
thus fewer bastardised and deteriorated offspring would be produced to
commingle their blood with the new species in process of formation. 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 speciesand which is universal
with species which have been differentiated to a generic or family rank
will find the subject extraordinarily complex. After mature reflectionit
seems to me that this could not have been effected through natural
selection. Take the case of any two species whichwhen crossedproduced
few and sterile offspring; nowwhat is there which could favour the
survival of those individuals which happened to be endowed in a slightly
higher degree with mutual infertilityand which thus approached by one
small step towards absolute sterility? Yet an advance of this kindif the
theory of natural selection be brought to bearmust have incessantly
occurred with many speciesfor a multitude are mutually quite barren.
With sterile neuter insects we have reason to believe that modifications in
their structure and fertility have been slowly accumulated by natural
selectionfrom an advantage having been thus indirectly given to the
community to which they belonged over other communities of the same
species; but an individual animal not belonging to a social communityif
rendered slightly sterile when crossed with some other varietywould not
thus itself gain any advantage or indirectly give any advantage to the
other individuals of the same varietythus leading to their preservation.

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 principlequite independent of natural selection.
Both Gartner and Kolreuter have proved that in genera including numerous
speciesa series can be formed from species which when crossed yield fewer
and fewer seedsto species which never produce a single seedbut yet are
affected by the pollen of certain other speciesfor the germen swells. It
is here manifestly impossible to select the more sterile individualswhich
have already ceased to yield seeds; so that this acme of sterilitywhen
the germen alone is effectedcannot have been gained through selection;
and from the laws governing the various grades of sterility being so
uniform throughout the animal and vegetable kingdomswe may infer that the
causewhatever it may beis the same or nearly the same in all cases.

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. In
the case of first crossesthe greater or less difficulty in effecting a
union and in obtaining offspring apparently depends on several distinct
causes. There must sometimes be a physical impossibility in the male
element reaching the ovuleas would be the case with a plant having a
pistil too long for the pollen-tubes to reach the ovarium. It has also
been observed that when the pollen of one species is placed on the stigma
of a distantly allied speciesthough the pollen-tubes protrudethey do
not penetrate the stigmatic surface. Againthe male element may reach the
female elementbut be incapable of causing an embryo to be developedas
seems to have been the case with some of Thuret's experiments on Fuci. No
explanation can be given of these factsany more than why certain trees
cannot be grafted on others. Lastlyan embryo may be developedand then
perish at an early period. This latter alternative has not been
sufficiently attended to; but I believefrom observations communicated to
me by Mr. Hewittwho has had great experience in hybridising pheasants and
fowlsthat the early death of the embryo is a very frequent cause of
sterility in first crosses. Mr. 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; the majority of these eggs had
been fertilised; and in the majority of the fertilised eggsthe embryos
had either been partially developed and had then perishedor had become
nearly maturebut the young chickens had been unable to break through the
shell. Of the chickens which were bornmore than four-fifths died within
the first few daysor at latest weekswithout any obvious cause,
apparently from mere inability to live;so that from the 500 eggs only
twelve chickens were reared. With plantshybridized embryos probably
often perish in a like manner; at least it is known that hybrids raised
from very distinct species are sometimes weak and dwarfedand perish at an
early age; of which fact Max Wichura has recently given some striking cases
with hybrid willows. It may be here worth noticing that in some cases of
parthenogenesisthe embryos within the eggs of silk moths which had not
been fertilisedpass through their early stages of development and then
perish like the embryos produced by a cross between distinct species.
Until becoming acquainted with these factsI was unwilling to believe in
the frequent early death of hybrid embryos; for hybridswhen once born
are generally healthy and long-livedas we see in the case of the common
mule. Hybridshoweverare differently circumstanced before and after
birth: when born and living in a country where their two parents live
they are generally placed under suitable conditions of life. But a hybrid
partakes of only half of the nature and constitution of its mother; it may
thereforebefore birthas long as it is nourished within its mother's
wombor within the egg or seed produced by the motherbe exposed to
conditions in some degree unsuitableand consequently be liable to perish
at an early period; more especially as all very young beings are eminently
sensitive to injurious or unnatural conditions of life. But after allthe
cause more probably lies in some imperfection in the original act of
impregnationcausing the embryo to be imperfectly developedrather than
in the conditions to which it is subsequently exposed.

In regard to the sterility of hybridsin which the sexual elements are
imperfectly developedthe case is somewhat different. I have more than
once alluded to a large body of facts showing thatwhen animals and plants
are removed from their natural conditionsthey are extremely liable to
have their reproductive systems seriously affected. Thisin factis the
great bar to the domestication of animals. Between the sterility thus
superinduced and that of hybridsthere are many points of similarity. In
both cases the sterility is independent of general healthand is often
accompanied by excess of size or great luxuriance. In both cases the
sterility occurs in various degrees; in boththe male element is the most
liable to be affected; but sometimes the female more than the male. In
boththe tendency goes to a certain extent with systematic affinityfor
whole groups of animals and plants are rendered impotent by the same
unnatural conditions; and whole groups of species tend to produce sterile

hybrids. On the other handone species in a group will sometimes resist
great changes of conditions with unimpaired fertility; and certain species
in a group will produce unusually fertile hybrids. No one can tell till he
trieswhether any particular animal will breed under confinementor any
exotic plant seed freely under culture; nor can he tell till he tries
whether any two species of a genus will produce more or less sterile
hybrids. Lastlywhen organic beings are placed during several generations
under conditions not natural to themthey are extremely liable to vary
which seems to be partly due to their reproductive systems having been
specially affectedthough in a lesser degree than when sterility ensues.
So it is with hybridsfor their offspring in successive generations are
eminently liable to varyas every experimentalist has observed.

Thus we see that when organic beings are placed under new and unnatural
conditionsand when hybrids are produced by the unnatural crossing of two
speciesthe reproductive systemindependently of the general state of
healthis affected in a very similar manner. In the one casethe
conditions of life have been disturbedthough often in so slight a degree
as to be inappreciable by us; in the other caseor that of hybridsthe
external conditions have remained the samebut the organisation has been
disturbed by two distinct structures and constitutionsincluding of course
the reproductive systemshaving been blended into one. For it is scarcely
possible that two organisations should be compounded into onewithout some
disturbance occurring in the developmentor periodical actionor mutual
relations of the different parts and organs one to another or to the
conditions of life. When hybrids are able to breed inter sethey transmit
to their offspring from generation to generation the same compounded
organisationand hence we need not be surprised that their sterility
though in some degree variabledoes not diminish; it is even apt to
increasethis being generally the resultas before explainedof too
close interbreeding. The above view of the sterility of hybrids being
caused by two constitutions being compounded into one has been strongly
maintained by Max Wichura.

It musthoweverbe owned that we cannot understandon the above or any
other viewseveral facts with respect to the sterility of hybrids; for
instancethe unequal fertility of hybrids produced from reciprocal
crosses; or the increased sterility in those hybrids which occasionally and
exceptionally resemble closely either pure parent. Nor do I pretend that
the foregoing remarks go to the root of the matter: no explanation is
offered why an organismwhen placed under unnatural conditionsis
rendered sterile. All that I have attempted to show isthat in two cases
in some respects alliedsterility is the common result--in the one case
from the conditions of life having been disturbedin the other case from
the organisation having been disturbed by two organisations being
compounded into one.

A similar parallelism holds good with an allied yet very different class of
facts. It is an old and almost universal belieffounded on a considerable
body of evidencewhich I have elsewhere giventhat slight changes in the
conditions of life are beneficial to all living things. We see this acted
on by farmers and gardeners in their frequent exchanges of seedtubers
etc.from one soil or climate to anotherand back again. During the
convalescence of animalsgreat benefit is derived from almost any change
in their habits of life. Againboth with plants and animalsthere is the
clearest evidence that a cross between individuals of the same species
which differ to a certain extentgives vigour and fertility to the
offspring; and that close interbreeding continued during several
generations between the nearest relationsif these be kept under the same
conditions of lifealmost always leads to decreased sizeweaknessor

Hence it seems thaton the one handslight changes in the conditions of
life benefit all organic beingsand on the other handthat slight

crossesthat iscrosses between the males and females of the same
specieswhich have been subjected to slightly different conditionsor
which have slightly variedgive vigour and fertility to the offspring.
Butas we have seenorganic beings long habituated to certain uniform
conditions under a state of naturewhen subjectedas under confinement
to a considerable change in their conditionsvery frequently are rendered
more or less sterile; and we know that a cross between two forms that have
become widely or specifically differentproduce hybrids which are almost
always in some degree sterile. I am fully persuaded that this double
parallelism is by no means an accident or an illusion. He who is able to
explain why the elephantand a multitude of other animalsare incapable
of breeding when kept under only partial confinement in their native
countrywill be able to explain the primary cause of hybrids being so
generally sterile. He will at the same time be able to explain how it is
that the races of some of our domesticated animalswhich have often been
subjected to new and not uniform conditionsare quite fertile together
although they are descended from distinct specieswhich 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
which is essentially related to the principle of life; this principle
according to Mr. Herbert Spencerbeing that life depends onor consists
inthe incessant action and reaction of various forceswhichas
throughout natureare always tending towards an equilibrium; and when this
tendency is slightly disturbed by any changethe vital forces gain in


This subject may be here briefly discussedand will be found to throw some
light on hybridism. Several plants belonging to distinct orders present
two formswhich exist in about equal numbers and which differ in no
respect except in their reproductive organs; one form having a long pistil
with short stamensthe other a short pistil with long stamens; the two
having differently sized pollen-grains. With trimorphic plants there are
three forms likewise differing in the lengths of their pistils and stamens
in the size and colour of the pollen-grainsand in some other respects;
and as in each of the three forms there are two sets of stamensthe three
forms possess altogether six sets of stamens and three kinds of pistils.
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. Now I have shownand the result has been confirmed by other
observersthat in order to obtain full fertility with these plantsit is
necessary that the stigma of the one form should be fertilised by pollen
taken from the stamens of corresponding height in another form. So that
with dimorphic species two unionswhich may be called legitimateare
fully fertile; and twowhich may be called illegitimateare more or less
infertile. With trimorphic species six unions are legitimateor fully
fertileand twelve are illegitimateor more or less infertile.

The infertility which may be observed in various dimorphic and trimorphic
plantswhen they are illegitimately fertilisedthat is by pollen taken
from stamens not corresponding in height with the pistildiffers much in
degreeup to absolute and utter sterility; just in the same manner as
occurs in crossing distinct species. As the degree of sterility in the
latter case depends in an eminent degree on the conditions of life being
more or less favourableso I have found it with illegitimate unions. It
is well known that if pollen of a distinct species be placed on the stigma
of a flowerand its own pollen be afterwardseven after a considerable
interval of timeplaced on the same stigmaits action is so strongly
prepotent that it generally annihilates the effect of the foreign pollen;
so it is with the pollen of the several forms of the same speciesfor
legitimate pollen is strongly prepotent over illegitimate pollenwhen both
are placed on the same stigma. I ascertained this by fertilising several
flowersfirst illegitimatelyand twenty-four hours afterwards

legitimatelywith pollen taken from a peculiarly coloured varietyand all
the seedlings were similarly coloured; this shows that the legitimate
pollenthough applied twenty-four hours subsequentlyhad wholly destroyed
or prevented the action of the previously applied illegitimate pollen.
Againas in making reciprocal crosses between the same two speciesthere
is occasionally a great difference in the resultso the same thing occurs
with trimorphic plants; for instancethe mid-styled form of Lythrum
salicaria was illegitimately fertilised with the greatest ease by pollen
from the longer stamens of the short-styled formand yielded many seeds;
but the latter form did not yield a single seed when fertilised by the
longer stamens of the mid-styled form.

In all these respectsand in others which might be addedthe forms of the
same undoubted specieswhen illegitimately unitedbehave in exactly the
same manner as do two distinct species when crossed. This led me carefully
to observe during four years many seedlingsraised from several
illegitimate unions. The chief result is that these illegitimate plants
as they may be calledare not fully fertile. It is possible to raise from
dimorphic speciesboth long-styled and short-styled illegitimate plants
and from trimorphic plants all three illegitimate forms. These can then be
properly united in a legitimate manner. When this is donethere is no
apparent reason why they should not yield as many seeds as did their
parents when legitimately fertilised. But such is not the case. They are
all infertilein various degrees; some being so utterly and incurably
sterile that they did not yield during four seasons a single seed or even
seed-capsule. The sterility of these illegitimate plantswhen united with
each other in a legitimate mannermay be strictly compared with that of
hybrids when crossed inter se. Ifon the other handa hybrid is crossed
with either pure parent-speciesthe sterility is usually much lessened:
and so it is when an illegitimate plant is fertilised by a legitimate
plant. 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
parent-speciesso that sterility of certain illegitimate plants was
unusually greatwhile the sterility of the union from which they were
derived was by no means great. With hybrids raised from the same seedcapsule
the degree of sterility is innately variableso it is in a marked
manner with illegitimate plants. Lastlymany hybrids are profuse and
persistent flowererswhile other and more sterile hybrids produce few
flowersand are weakmiserable dwarfs; exactly similar cases occur with
the illegitimate offspring of various dimorphic and trimorphic plants.

Altogether there is the closest identity in character and behaviour between
illegitimate plants and hybrids. It is hardly an exaggeration to maintain
that illegitimate plants are hybridsproduced within the limits of the
same species by the improper union of certain formswhile ordinary hybrids
are produced from an improper union between so-called distinct species. We
have also already seen that there is the closest similarity in all respects
between first illegitimate unions and first crosses between distinct
species. This will perhaps be made more fully apparent by an illustration;
we may suppose that a botanist found two well-marked varieties (and such
occur) of the long-styled form of the trimorphic Lythrum salicariaand
that he determined to try by crossing whether they were specifically
distinct. He would find that they yielded only about one-fifth of the
proper number of seedand that they behaved in all the other above
specified respects as if they had been two distinct species. But to make
the case surehe would raise plants from his supposed hybridised seedand
he would find that the seedlings were miserably dwarfed and utterly
sterileand that they behaved in all other respects like ordinary hybrids.
He might then maintain that he had actually provedin accordance with the
common viewthat his two varieties were as good and as distinct species as
any in the world; but he would be completely mistaken.

The facts now given on dimorphic and trimorphic plants are important
because they show usfirstthat the physiological test of lessened

fertilityboth in first crosses and in hybridsis no safe criterion of
specific distinction; secondlybecause we may conclude that there is some
unknown bond which connects the infertility of illegitimate unions with
that of their illegitimate offspringand we are led to extend the same
view to first crosses and hybrids; thirdlybecause we findand this seems
to me of especial importancethat two or three forms of the same species
may exist and may differ in no respect whatevereither in structure or in
constitutionrelatively to external conditionsand yet be sterile when
united in certain ways. For we must remember that it is the union of the
sexual elements of individuals of the same formfor instanceof two longstyled
formswhich results in sterility; while it is the union of the
sexual elements proper to two distinct forms which is fertile. Hence the
case appears at first sight exactly the reverse of what occursin the
ordinary unions of the individuals of the same species and with crosses
between distinct species. It ishoweverdoubtful whether this is really
so; but I will not enlarge on this obscure subject.

We mayhoweverinfer as probable from the consideration of dimorphic and
trimorphic plantsthat the sterility of distinct species when crossed and
of their hybrid progenydepends exclusively on the nature of their sexual
elementsand not on any difference in their structure or general
constitution. We are also led to this same conclusion by considering
reciprocal crossesin which the male of one species cannot be unitedor
can be united with great difficultywith the female of a second species
while the converse cross can be effected with perfect facility. That
excellent observerGartnerlikewise concluded that species when crossed
are sterile owing to differences confined to their reproductive systems.


It may be urged as an overwhelming argument that there must be some
essential distinction between species and varieties inasmuch as the latter
however much they may differ from each other in external appearancecross
with perfect facilityand yield perfectly fertile offspring. With some
exceptionspresently to be givenI fully admit that this is the rule.
But the subject is surrounded by difficultiesforlooking to varieties
produced under natureif two forms hitherto reputed to be varieties be
found in any degree sterile togetherthey are at once ranked by most
naturalists as species. For instancethe blue and red pimpernelwhich
are considered by most botanists as varietiesare said by Gartner to be
quite sterile when crossedand he consequently ranks them as undoubted
species. If we thus argue in a circlethe fertility of all varieties
produced under nature will assuredly have to be granted.

If we turn to varietiesproducedor supposed to have been producedunder
domesticationwe are still involved in some doubt. For when it is stated
for instancethat certain South American indigenous domestic dogs do not
readily unite with European dogsthe explanation which will occur to
everyoneand probably the true oneis that they are descended from
aboriginally distinct species. Nevertheless the perfect fertility of so
many domestic racesdiffering widely from each other in appearancefor
instancethose of the pigeonor of the cabbageis a remarkable fact;
more especially when we reflect how many species there arewhichthough
resembling each other most closelyare utterly sterile when intercrossed.
Several considerationshoweverrender the fertility of domestic varieties
less remarkable. In the first placeit may be observed that the amount of
external difference between two species is no sure guide to their degree of
mutual sterilityso that similar differences in the case of varieties
would be no sure guide. It is certain that with species the cause lies
exclusively in differences in their sexual constitution. Now the varying
conditions to which domesticated animals and cultivated plants have been
subjectedhave had so little tendency towards modifying the reproductive
system in a manner leading to mutual sterilitythat we have good grounds

for admitting the directly opposite doctrine of Pallasnamelythat such
conditions generally eliminate this tendency; so that the domesticated
descendants of specieswhich in their natural state probably would have
been in some degree sterile when crossedbecome perfectly fertile
together. With plantsso far is cultivation from giving a tendency
towards sterility between distinct speciesthat in several wellauthenticated
cases already alluded tocertain plants have been affected
in an opposite mannerfor they have become self-impotentwhile still
retaining the capacity of fertilisingand being fertilised byother
species. If the Pallasian doctrine of the elimination of sterility through
long-continued domestication be admittedand it can hardly be rejectedit
becomes in the highest degree improbable that similar conditions longcontinued
should likewise induce this tendency; though in certain cases
with species having a peculiar constitutionsterility might occasionally
be thus caused. Thusas I believewe can understand whywith
domesticated animalsvarieties have not been produced which are mutually
sterile; and why with plants only a few such casesimmediately to be
givenhave been observed.

The real difficulty in our present subject is notas it appears to mewhy
domestic varieties have not become mutually infertile when crossedbut why
this has so generally occurred with natural varietiesas soon as they have
been permanently modified in a sufficient degree to take rank as species.
We are far from precisely knowing the cause; nor is this surprisingseeing
how profoundly ignorant we are in regard to the normal and abnormal action
of the reproductive system. But we can see that speciesowing to their
struggle for existence with numerous competitorswill have been exposed
during long periods of time to more uniform conditionsthan have domestic
varieties; and this may well make a wide difference in the result. For we
know how commonly wild animals and plantswhen taken from their natural
conditions and subjected to captivityare rendered sterile; 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. Domesticated productionson the
other handwhichas shown by the mere fact of their domesticationwere
not originally highly sensitive to changes in their conditions of lifeand
which can now generally resist with undiminished fertility repeated changes
of conditionsmight be expected to produce varietieswhich 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.

I have as yet spoken as if the varieties of the same species were
invariably fertile when intercrossed. But it is impossible to resist the
evidence of the existence of a certain amount of sterility in the few
following caseswhich I will briefly abstract. The evidence is at least
as good as that from which we believe in the sterility of a multitude of
species. The evidence is also derived from hostile witnesseswho in all
other cases consider fertility and sterility as safe criterions of specific
distinction. Gartner keptduring several yearsa dwarf kind of maize
with yellow seedsand a tall variety with red seeds growing near each
other in his garden; and although these plants have separated sexesthey
never naturally crossed. He then fertilised thirteen flowers of the one
kind with pollen of the other; but only a single head produced any seed
and this one head produced only five grains. Manipulation in this case
could not have been injuriousas the plants have separated sexes. No one
I believehas suspected that these varieties of maize are distinct
species; and it is important to notice that the hybrid plants thus raised
were themselves PERFECTLY fertile; so that even Gartner did not venture to
consider the two varieties as specifically distinct.

Girou de Buzareingues crossed three varieties of gourdwhich like the
maize has separated sexesand he asserts that their mutual fertilisation
is by so much the less easy as their differences are greater. How far
these experiments may be trustedI know not; but the forms experimented on

are ranked by Sagaretwho mainly founds his classification by the test of
infertilityas varietiesand Naudin has come to the same conclusion.

The following case is far more remarkableand seems at first incredible;
but it is the result of an astonishing number of experiments made during
many years on nine species of Verbascumby so good an observer and so
hostile a witness as Gartner: namelythat the yellow and white varieties
when crossed produce less seed than the similarly coloured varieties of the
same species. Moreoverhe asserts thatwhen yellow and white varieties
of one species are crossed with yellow and white varieties of a DISTINCT
speciesmore seed is produced by the crosses between the similarly
coloured flowersthan between those which are differently coloured. Mr.
Scott also has experimented on the species and varieties of Verbascum; and
although unable to confirm Gartner's results on the crossing of the
distinct specieshe finds that the dissimilarly coloured varieties of the
same species yield fewer seedsin the proportion of eighty-six to 100
than the similarly coloured varieties. Yet these varieties differ in no
respectexcept in the colour of their flowers; and one variety can
sometimes be raised from the seed of another.

Kolreuterwhose accuracy has been confirmed by every subsequent observer
has proved the remarkable fact that one particular variety of the common
tobacco was more fertile than the other varietieswhen crossed with a
widely distinct species. He experimented on five forms which are commonly
reputed to be varietiesand which he tested by the severest trialnamely
by reciprocal crossesand he found their mongrel offspring perfectly
fertile. But one of these five varietieswhen used either as the father
or motherand crossed with the Nicotiana glutinosaalways yielded hybrids
not so sterile as those which were produced from the four other varieties
when crossed with N. glutinosa. Hence the reproductive system of this one
variety must have been in some manner and in some degree modified.

>From these facts it can no longer be maintained that varieties when crossed
are invariably quite fertile. From the great difficulty of ascertaining
the infertility of varieties in a state of naturefor a supposed variety
if proved to be infertile in any degreewould almost universally be ranked
as a species; from man attending only to external characters in his
domestic varietiesand from such varieties not having been exposed for
very long periods to uniform conditions of life; from these several
considerations we may conclude that fertility does not constitute a
fundamental distinction between varieties and species when crossed. The
general sterility of crossed species may safely be looked atnot as a
special acquirement or endowmentbut as incidental on changes of an
unknown nature in their sexual elements.


Independently of the question of fertilitythe offspring of species and of
varieties when crossed may be compared in several other respects. Gartner
whose strong wish it was to draw a distinct line between species and
varietiescould find very fewandas it seems to mequite unimportant
differences between the so-called hybrid offspring of speciesand the
so-called mongrel offspring of varieties. Andon the other handthey
agree most closely in many important respects.

I shall here discuss this subject with extreme brevity. The most important
distinction isthat in the first generation mongrels are more variable
than hybrids; but Gartner admits that hybrids from species which have long
been cultivated are often variable in the first generation; and I have
myself seen striking instances of this fact. Gartner further admits that
hybrids between very closely allied species are more variable than those
from very distinct species; and this shows that the difference in the
degree of variability graduates away. When mongrels and the more fertile
hybrids are propagated for several generationsan extreme amount of

variability in the offspring in both cases is notorious; but some few
instances of both hybrids and mongrels long retaining a uniform character
could be given. The variabilityhoweverin the successive generations of
mongrels isperhapsgreater than in hybrids.

This greater variability in mongrels than in hybrids does not seem at all
surprising. For the parents of mongrels are varietiesand mostly domestic
varieties (very few experiments having been tried on natural varieties)
and this implies that there has been recent variability; which would often
continue and would augment that arising from the act of crossing. The
slight variability of hybrids in the first generationin contrast with
that in the succeeding generationsis a curious fact and deserves
attention. For it bears on the view which I have taken of one of the
causes of ordinary variability; namelythat the reproductive systemfrom
being eminently sensitive to changed conditions of lifefails under these
circumstances to perform its proper function of producing offspring closely
similar in all respects to the parent-form. Nowhybrids in the first
generation are descended from species (excluding those long cultivated)
which have not had their reproductive systems in any way affectedand they
are not variable; but hybrids themselves have their reproductive systems
seriously affectedand their descendants are highly variable.

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;
but thisif it be trueis certainly only a difference in degree.
MoreoverGartner expressly states that the hybrids from long cultivated
plants are more subject to reversion than hybrids from species in their
natural state; and this probably explains the singular difference in the
results arrived at by different observers. Thus Max Wichura doubts whether
hybrids ever revert to their parent formsand he experimented on
uncultivated species of willowswhile Naudinon the other handinsists
in the strongest terms on the almost universal tendency to reversion in
hybridsand he experimented chiefly on cultivated plants. Gartner further
states that when any two speciesalthough most closely allied to each
otherare crossed with a third speciesthe hybrids are widely different
from each other; whereas if two very distinct varieties of one species are
crossed with another speciesthe hybrids do not differ much. But this
conclusionas far as I can make outis founded on a single experiment;
and seems directly opposed to the results of several experiments made by

Such alone are the unimportant differences which Gartner is able to point
out between hybrid and mongrel plants. On the other handthe degrees and
kinds of resemblance in mongrels and in hybrids to their respective
parentsmore especially in hybrids produced from nearly related species
followaccording to Gartner the same laws. When two species are crossed
one has sometimes a prepotent power of impressing its likeness on the
hybrid. So I believe it to be with varieties of plants; and with animals
one variety certainly often has this prepotent power over another variety.
Hybrid plants produced from a reciprocal cross generally resemble each
other closelyand so it is with mongrel plants from a reciprocal cross.
Both hybrids and mongrels can be reduced to either pure parent formby
repeated crosses in successive generations with either parent.

These several remarks are apparently applicable to animals; but the subject
is here much complicatedpartly owing to the existence of secondary sexual
characters; but more especially owing to prepotency in transmitting
likeness running more strongly in one sex than in the otherboth when one
species is crossed with another and when one variety is crossed with
another variety. For instanceI think those authors are right who
maintain that the ass has a prepotent power over the horseso that both
the mule and the hinny resemble more closely the ass than the horse; but
that the prepotency runs more strongly in the male than in the female ass
so that the mulewhich is an offspring of the male ass and mareis more

like an ass than is the hinnywhich is the offspring of the female-ass and

Much stress has been laid by some authors on the supposed factthat it is
only with mongrels that the offspring are not intermediate in character
but closely resemble one of their parents; but this does sometimes occur
with hybridsyet I grant much less frequently than with mongrels. Looking
to the cases which I have collected of cross-bred animals closely
resembling one parentthe resemblances seem chiefly confined to characters
almost monstrous in their natureand which have suddenly appeared--such as
albinismmelanismdeficiency of tail or hornsor additional fingers and
toes; and do not relate to characters which have been slowly acquired
through selection. A tendency to sudden reversions to the perfect
character of either parent wouldalsobe much more likely to occur with
mongrelswhich are descended from varieties often suddenly produced and
semi-monstrous in characterthan with hybridswhich are descended from
species slowly and naturally produced. On the wholeI entirely agree with
Dr. Prosper Lucaswhoafter arranging an enormous body of facts with
respect to animalscomes to the conclusion that the laws of resemblance of
the child to its parents are the samewhether the two parents differ
little or much from each othernamelyin the union of individuals of the
same varietyor of different varietiesor of distinct species.

Independently of the question of fertility and sterilityin all other
respects there seems to be a general and close similarity in the offspring
of crossed speciesand of crossed varieties. If we look at species as
having been specially createdand at varieties as having been produced by
secondary lawsthis similarity would be an astonishing fact. But it
harmonises perfectly with the view that there is no essential distinction
between species and varieties.


First crosses between formssufficiently distinct to be ranked as species
and their hybridsare very generallybut not universallysterile. The
sterility is of all degreesand is often so slight that the most careful
experimentalists have arrived at diametrically opposite conclusions in
ranking forms by this test. The sterility is innately variable in
individuals of the same speciesand is eminently susceptible to action of
favourable and unfavourable conditions. The degree of sterility does not
strictly follow systematic affinitybut is governed by several curious and
complex laws. It is generally differentand sometimes widely different in
reciprocal crosses between the same two species. It is not always equal in
degree in a first cross and in the hybrids produced from this cross.

In the same manner as in grafting treesthe capacity in one species or
variety to take on anotheris incidental on differencesgenerally of an
unknown naturein their vegetative systemsso in crossingthe greater or
less facility of one species to unite with another is incidental on unknown
differences in their reproductive systems. 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 naturethan 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.

The sterility of first crosses and of their hybrid progeny has not been
acquired through natural selection. In the case of first crosses it seems
to depend on several circumstances; in some instances in chief part on the
early death of the embryo. In the case of hybridsit apparently depends
on their whole organisation having been disturbed by being compounded from
two distinct forms; the sterility being closely allied to that which so
frequently affects pure specieswhen exposed to new and unnatural
conditions of life. He who will explain these latter cases will be able to

explain the sterility of hybrids. This view is strongly supported by a
parallelism of another kind: namelythatfirstlyslight changes in the
conditions of life add to the vigour and fertility of all organic beings;
and secondlythat the crossing of formswhich have been exposed to
slightly different conditions of lifeor which have variedfavours the
sizevigour and fertility of their offspring. The facts given on the
sterility of the illegitimate unions of dimorphic and trimorphic plants and
of their illegitimate progenyperhaps render it probable that some unknown
bond in all cases connects the degree of fertility of first unions with
that of their offspring. The consideration of these facts on dimorphism
as well as of the results of reciprocal crossesclearly leads to the
conclusion that the primary cause of the sterility of crossed species is
confined to differences in their sexual elements. But whyin the case of
distinct speciesthe sexual elements should so generally have become more
or less modifiedleading to their mutual infertilitywe do not know; 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.

It is not surprising that the difficulty in crossing any two speciesand
the sterility of their hybrid offspringshould in most cases correspond
even if due to distinct causes: for both depend on the amount of
difference between the species which are crossed. Nor is it surprising
that the facility of effecting a first crossand the fertility of the
hybrids thus producedand the capacity of being grafted together--though
this latter capacity evidently depends on widely different
circumstances--should all runto a certain extentparallel with the
systematic affinity of the forms subjected to experiment; for systematic
affinity includes resemblances of all kinds.

First crosses between forms known to be varietiesor sufficiently alike to
be considered as varietiesand their mongrel offspringare very
generallybut notas is so often statedinvariably fertile. Nor is this
almost universal and perfect fertility surprisingwhen it is remembered
how liable we are to argue in a circle with respect to varieties in a state
of nature; and when we remember that the greater number of varieties have
been produced under domestication by the selection of mere external
differencesand that they have not been long exposed to uniform conditions
of life. It should also be especially kept in mindthat long-continued
domestication tends to eliminate sterilityand is therefore little likely
to induce this same quality. Independently of the question of fertility
in all other respects there is the closest general resemblance between
hybrids and mongrelsin their variabilityin their power of absorbing
each other by repeated crossesand in their inheritance of characters from
both parent-forms. Finallythenalthough 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
sterileyet the facts given in this chapter do not seem to me opposed to
the belief that species aboriginally existed as varieties.



On the absence of intermediate varieties at the present day -- On the
nature of extinct intermediate varieties; on their number -- On the lapse
of timeas 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.

In the sixth chapter I enumerated the chief objections which might be
justly urged against the views maintained in this volume. Most of them
have now been discussed. Onenamelythe distinctness of specific forms
and their not being blended together by innumerable transitional linksis
a very obvious difficulty. I assigned reasons why such links do not
commonly occur at the present day under the circumstances apparently most
favourable for their presencenamelyon an extensive and continuous area
with graduated physical conditions. I endeavoured to showthat the life
of each species depends in a more important manner on the presence of other
already defined organic formsthan on climateandthereforethat the
really governing conditions of life do not graduate away quite insensibly
like heat or moisture. I endeavouredalsoto show that intermediate
varietiesfrom existing in lesser numbers than the forms which they
connectwill generally be beaten out and exterminated during the course of
further modification and improvement. The main causehoweverof
innumerable intermediate links not now occurring everywhere throughout
nature dependson the very process of natural selectionthrough which new
varieties continually take the places of and supplant their parent-forms.
But just in proportion as this process of extermination has acted on an
enormous scaleso must the number of intermediate varietieswhich have
formerly existedbe truly enormous. 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; and
thisperhapsis the most obvious and serious objection which can be urged
against my theory. The explanation liesas I believein the extreme
imperfection of the geological record.

In the first placeit should always be borne in mind what sort of
intermediate forms muston the theoryhave formerly existed. I have
found it difficultwhen looking at any two speciesto avoid picturing to
myself forms DIRECTLY intermediate between them. But this is a wholly
false view; we should always look for forms intermediate between each
species and a common but unknown progenitor; and the progenitor will
generally have differed in some respects from all its modified descendants.
To give a simple illustration: the fantail and pouter pigeons are both
descended from the rock-pigeon; if we possessed all the intermediate
varieties which have ever existedwe should have an extremely close series
between both and the rock-pigeon; but we should have no varieties directly
intermediate between the fantail and pouter; nonefor instancecombining
a tail somewhat expanded with a crop somewhat enlargedthe characteristic
features of these two breeds. These two breedsmoreoverhave become so
much modifiedthatif we had no historical or indirect evidence regarding
their originit would not have been possible to have determined from a
mere comparison of their structure with that of the rock-pigeonC. livia
whether they had descended from this species or from some other allied
speciessuch as C. oenas.

So with natural speciesif we look to forms very distinctfor instance to
the horse and tapirwe have no reason to suppose that links directly
intermediate between them ever existedbut between each and an unknown
common parent. The common parent will have had in its whole organisation
much general resemblance to the tapir and to the horse; but in some points
of structure may have differed considerably from botheven perhaps more
than they differ from each other. Hencein all such caseswe should be
unable to recognise the parent-form of any two or more specieseven if we
closely compared the structure of the parent with that of its modified
descendantsunless at the same time we had a nearly perfect chain of the
intermediate links.

It is just possibleby the theorythat one of two living forms might have
descended from the other; for instancea horse from a tapir; and in this
case DIRECT intermediate links will have existed between them. But such a
case would imply that one form had remained for a very long period
unalteredwhilst its descendants had undergone a vast amount of change;

and the principle of competition between organism and organismbetween
child and parentwill render this a very rare event; for in all cases the
new and improved forms of life tend to supplant the old and unimproved

By the theory of natural selection all living species have been connected
with the parent-species of each genusby differences not greater than we
see between the natural and domestic varieties of the same species at the
present day; and these parent-speciesnow generally extincthave in their
turn been similarly connected with more ancient forms; and so on backwards
always converging to the common ancestor of each great class. So that the
number of intermediate and transitional linksbetween all living and
extinct speciesmust have been inconceivably great. But assuredlyif
this theory be truesuch have lived upon the earth.


Independently of our not finding fossil remains of such infinitely numerous
connecting linksit may be objected that time cannot have sufficed for so
great an amount of organic changeall changes having been effected slowly.
It is hardly possible for me to recall to the reader who is not a practical
geologistthe facts leading the mind feebly to comprehend the lapse of
time. He who can read Sir Charles Lyell's grand work on the Principles of
Geologywhich the future historian will recognise as having produced a
revolution in natural scienceand yet does not admit how vast have been
the past periods of timemay at once close this volume. Not that it
suffices to study the Principles of Geologyor to read special treatises
by different observers on separate formationsand to mark how each author
attempts to give an inadequate idea of the duration of each formationor
even of each stratum. We can best gain some idea of past time by knowing
the agencies at work; and learning how deeply the surface of the land has
been denudedand how much sediment has been deposited. As Lyell has well
remarkedthe extent and thickness of our sedimentary formations are the
result and the measure of the denudation which the earth's crust has
elsewhere undergone. Therefore a man should examine for himself the great
piles of superimposed strataand watch the rivulets bringing down mudand
the waves wearing away the sea-cliffsin order to comprehend something
about the duration of past timethe monuments of which we see all around

It is good to wander along the coastwhen formed of moderately hard rocks
and mark the process of degradation. The tides in most cases reach the
cliffs only for a short time twice a dayand the waves eat into them only
when they are charged with sand or pebbles; for there is good evidence that
pure water effects nothing in wearing away rock. At last the base of the
cliff is underminedhuge fragments fall downand these remaining fixed
have to be worn away atom by atomuntil after being reduced in size they
can be rolled about by the wavesand then they are more quickly ground
into pebblessandor mud. But how often do we see along the bases of
retreating cliffs rounded bouldersall thickly clothed by marine
productionsshowing how little they are abraded and how seldom they are
rolled about! Moreoverif we follow for a few miles any line of rocky
cliffwhich is undergoing degradationwe find that it is only here and
therealong a short length or round a promontorythat the cliffs are at
the present time suffering. The appearance of the surface and the
vegetation show that elsewhere years have elapsed since the waters washed
their base.

We havehoweverrecently learned from the observations of Ramsayin the
van of many excellent observers--of JukesGeikieCroll and othersthat
subaerial degradation is a much more important agency than coast-actionor
the power of the waves. The whole surface of the land is exposed to the
chemical action of the air and of the rainwaterwith its dissolved

carbonic acidand in colder countries to frost; the disintegrated matter
is carried down even gentle slopes during heavy rainand to a greater
extent than might be supposedespecially in arid districtsby the wind;
it is then transported by the streams and riverswhichwhen rapid deepen
their channelsand triturate the fragments. On a rainy dayeven in a
gently undulating countrywe see the effects of subaerial degradation in
the muddy rills which flow down every slope. Messrs. Ramsay and Whitaker
have shownand the observation is a most striking onethat the great
lines of escarpment in the Wealden district and those ranging across
Englandwhich formerly were looked at as ancient sea-coastscannot have
been thus formedfor each line is composed of one and the same formation
while our sea-cliffs are everywhere formed by the intersection of various
formations. This being the casewe are compelled to admit that the
escarpments owe their origin in chief part to the rocks of which they are
composedhaving resisted subaerial denudation better than the surrounding
surface; this surface consequently has been gradually loweredwith the
lines of harder rock left projecting. Nothing impresses the mind with the
vast duration of timeaccording to our ideas of timemore forcibly than
the conviction thus gained that subaerial agencieswhich apparently have
so little powerand which seem to work so slowlyhave produced great

When thus impressed with the slow rate at which the land is worn away
through subaerial and littoral actionit is goodin order to appreciate
the past duration of timeto consideron the one handthe masses of rock
which have been removed over many extensive areasand on the other hand
the thickness of our sedimentary formations. I remember having been much
struck when viewing volcanic islandswhich have been worn by the waves and
pared all round into perpendicular cliffs of one or two thousand feet in
height; for the gentle slope of the lava-streamsdue to their formerly
liquid stateshowed at a glance how far the hardrocky beds had once
extended into the open ocean. The same story is told still more plainly by
faults--those great cracks along which the strata have been upheaved on one
sideor thrown down on the otherto the height or depth of thousands of
feet; for since the crust crackedand it makes no great difference whether
the upheaval was suddenoras most geologists now believewas slow and
effected by many startsthe surface of the land has been so completely
planed down that no trace of these vast dislocations is externally visible.
The Craven faultfor instanceextends for upward of thirty milesand
along this line the vertical displacement of the strata varies from 600 to
3000 feet. Professor Ramsay has published an account of a downthrow in
Anglesea of 2300 feet; and he informs me that he fully believes that there
is one in Merionethshire of 12000 feet; yet in these cases there is
nothing on the surface of the land to show such prodigious movements; the
pile of rocks on either side of the crack having been smoothly swept away.

On the other handin all parts of the world the piles of sedimentary
strata are of wonderful thickness. In the CordilleraI estimated one mass
of conglomerate at ten thousand feet; and although conglomerates have
probably been accumulated at a quicker rate than finer sedimentsyet from
being formed of worn and rounded pebbleseach of which bears the stamp of
timethey are good to show how slowly the mass must have been heaped
together. Professor Ramsay has given me the maximum thicknessfrom actual
measurement in most casesof the successive formations in DIFFERENT parts
of Great Britain; and this is the result:

Palaeozoic strata (not including igneous beds)..57154
Secondary strata................................13190
Tertiary strata..................................2240

--making altogether 72584 feet; that isvery nearly thirteen and
three-quarters British miles. Some of these formationswhich are
represented in England by thin bedsare thousands of feet in thickness on
the Continent. Moreoverbetween each successive formation we havein the

opinion of most geologistsblank periods of enormous length. 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 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.

Nevertheless this impression is partly false. Mr. Crollin an interesting
paperremarks that we do not err "in forming too great a conception of the
length of geological periods but in estimating them by years. When
geologists look at large and complicated phenomena, and then at the figures
representing several million years, the two produce a totally different
effect on the mind, and the figures are at once pronounced too small. In
regard to subaerial denudation, Mr. Croll shows, by calculating the known
amount of sediment annually brought down by certain rivers, relatively to
their areas of drainage, that 1,000 feet of solid rock, as it became
gradually disintegrated, would thus be removed from the mean level of the
whole area in the course of six million years. This seems an astonishing
result, and some considerations lead to the suspicion that it may be too
large, but if halved or quartered it is still very surprising. Few of us,
however, know what a million really means: Mr. Croll gives the following
illustration: Take a narrow strip of paper, eighty-three feet four inches
in length, and stretch it along the wall of a large hall; then mark off at
one end the tenth of an inch. This tenth of an inch will represent one
hundred years, and the entire strip a million years. But let it be borne
in mind, in relation to the subject of this work, what a hundred years
implies, represented as it is by a measure utterly insignificant in a hall
of the above dimensions. Several eminent breeders, during a single
lifetime, have so largely modified some of the higher animals, which
propagate their kind much more slowly than most of the lower animals, that
they have formed what well deserves to be called a new sub-breed. Few men
have attended with due care to any one strain for more than half a century,
so that a hundred years represents the work of two breeders in succession.
It is not to be supposed that species in a state of nature ever change so
quickly as domestic animals under the guidance of methodical selection.
The comparison would be in every way fairer with the effects which follow
from unconscious selection, that is, the preservation of the most useful or
beautiful animals, with no intention of modifying the breed; but by this
process of unconscious selection, various breeds have been sensibly changed
in the course of two or three centuries.

Species, however, probably change much more slowly, and within the same
country only a few change at the same time. This slowness follows from all
the inhabitants of the same country being already so well adapted to each
other, that new places in the polity of nature do not occur until after
long intervals, due to the occurrence of physical changes of some kind, or
through the immigration of new forms. Moreover, variations or individual
differences of the right nature, by which some of the inhabitants might be
better fitted to their new places under the altered circumstance, would not
always occur at once. Unfortunately we have no means of determining,
according to the standard of years, how long a period it takes to modify a
species; but to the subject of time we must return.


Now let us turn to our richest museums, and what a paltry display we
behold! That our collections are imperfect is admitted by every one. The
remark of that admirable palaeontologist, Edward Forbes, should never be
forgotten, namely, that very many fossil species are known and named from
single and often broken specimens, or from a few specimens collected on
some one spot. Only a small portion of the surface of the earth has been
geologically explored, and no part with sufficient care, as the important
discoveries made every year in Europe prove. No organism wholly soft can
be preserved. Shells and bones decay and disappear when left on the bottom
of the sea, where sediment is not accumulating. We probably take a quite

erroneous view, when we assume that sediment is being deposited over nearly
the whole bed of the sea, at a rate sufficiently quick to embed and
preserve fossil remains. Throughout an enormously large proportion of the
ocean, the bright blue tint of the water bespeaks its purity. The many
cases on record of a formation conformably covered, after an immense
interval of time, by another and later formation, without the underlying
bed having suffered in the interval any wear and tear, seem explicable only
on the view of the bottom of the sea not rarely lying for ages in an
unaltered condition. The remains which do become embedded, if in sand or
gravel, will, when the beds are upraised, generally be dissolved by the
percolation of rain water charged with carbonic acid. Some of the many
kinds of animals which live on the beach between high and low water mark
seem to be rarely preserved. For instance, 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, with the
exception of a single Mediterranean species, which inhabits deep water and
this has been found fossil in Sicily, 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. Lastly, many great
deposits, requiring a vast length of time for their accumulation, are
entirely destitute of organic remains, without our being able to assign any
reason: one of the most striking instances is that of the Flysch
formation, which consists of shale and sandstone, several thousand,
occasionally even six thousand feet in thickness, and extending for at
least 300 miles from Vienna to Switzerland; and although this great mass
has been most carefully searched, no fossils, except a few vegetable
remains, have been found.

With respect to the terrestrial productions which lived during the
Secondary and Palaeozoic periods, it is superfluous to state that our
evidence is fragmentary in an extreme degree. For instance, until recently
not a land-shell was known belonging to either of these vast periods, with
the exception of one species discovered by Sir C. Lyell and Dr. Dawson in
the carboniferous strata of North America; but now land-shells have been
found in the lias. In regard to mammiferous remains, a glance at the
historical table published in Lyell's Manual, will bring home the truth,
how accidental and rare is their preservation, far better than pages of
detail. Nor is their rarity surprising, when we remember how large a
proportion of the bones of tertiary mammals have been discovered either in
caves or in lacustrine deposits; and that not a cave or true lacustrine bed
is known belonging to the age of our secondary or palaeozoic formations.

But the imperfection in the geological record largely results from another
and more important cause than any of the foregoing; namely, from the
several formations being separated from each other by wide intervals of
time. This doctrine has been emphatically admitted by many geologists and
palaeontologists, who, like E. Forbes, entirely disbelieve in the change of
species. When we see the formations tabulated in written works, or when we
follow them in nature, it is difficult to avoid believing that they are
closely consecutive. But we know, for instance, from Sir R. Murchison's
great work on Russia, what wide gaps there are in that country between the
superimposed formations; so it is in North America, and in many other parts
of the world. The most skilful geologist, if his attention had been
confined exclusively to these large territories, would never have suspected
that during the periods which were blank and barren in his own country,
great piles of sediment, charged with new and peculiar forms of life, had
elsewhere been accumulated. And if, in every separate territory, hardly
any idea can be formed of the length of time which has elapsed between the
consecutive formations, we may infer that this could nowhere be
ascertained. The frequent and great changes in the mineralogical
composition of consecutive formations, generally implying great changes in
the geography of the surrounding lands, whence the sediment was derived,
accord with the belief of vast intervals of time having elapsed between
each formation.

We can, I think, see why the geological formations of each region are
almost invariably intermittent; that is, have not followed each other in
close sequence. Scarcely any fact struck me more when examining many
hundred miles of the South American coasts, which have been upraised
several hundred feet within the recent period, than the absence of any
recent deposits sufficiently extensive to last for even a short geological
period. Along the whole west coast, which is inhabited by a peculiar
marine fauna, tertiary beds are so poorly developed that no record of
several successive and peculiar marine faunas will probably be preserved to
a distant age. A little reflection will explain why, along the rising
coast of the western side of South America, no extensive formations with
recent or tertiary remains can anywhere be found, though the supply of
sediment must for ages have been great, from the enormous degradation of
the coast rocks and from the muddy streams entering the sea. The
explanation, no doubt, is that the littoral and sub-littoral deposits are
continually worn away, as soon as they are brought up by the slow and
gradual rising of the land within the grinding action of the coast-waves.

We may, I think, conclude that sediment must be accumulated in extremely
thick, solid, or extensive masses, in order to withstand the incessant
action of the waves, when first upraised and during subsequent oscillations
of level, as well as the subsequent subaerial degradation. Such thick and
extensive accumulations of sediment may be formed in two ways; either in
profound depths of the sea, in which case the bottom will not be inhabited
by so many and such varied forms of life as the more shallow seas; and the
mass when upraised will give an imperfect record of the organisms which
existed in the neighbourhood during the period of its accumulation. Or
sediment may be deposited to any thickness and extent over a shallow
bottom, if it continue slowly to subside. In this latter case, as long as
the rate of subsidence and supply of sediment nearly balance each other,
the sea will remain shallow and favourable for many and varied forms, and
thus a rich fossiliferous formation, thick enough, when upraised, to resist
a large amount of denudation, may be formed.

I am convinced that nearly all our ancient formations, which are throughout
the greater part of their thickness RICH IN FOSSILS, have thus been formed
during subsidence. Since publishing my views on this subject in 1845, I
have watched the progress of geology, and have been surprised to note how
author after author, in treating of this or that great formation, has come
to the conclusion that it was accumulated during subsidence. I may add,
that the only ancient tertiary formation on the west coast of South
America, which has been bulky enough to resist such degradation as it has
as yet suffered, but which will hardly last to a distant geological age,
was deposited during a downward oscillation of level, and thus gained
considerable thickness.

All geological facts tell us plainly that each area has undergone numerous
slow oscillations of level, and apparently these oscillations have affected
wide spaces. Consequently, formations rich in fossils and sufficiently
thick and extensive to resist subsequent degradation, will have been formed
over wide spaces during periods of subsidence, 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. On the other hand, as long as
the bed of the sea remained stationary, THICK deposits cannot have been
accumulated in the shallow parts, which are the most favourable to life.
Still less can this have happened during the alternate periods of
elevation; or, to speak more accurately, the beds which were then
accumulated will generally have been destroyed by being upraised and
brought within the limits of the coast-action.

These remarks apply chiefly to littoral and sublittoral deposits. In the
case of an extensive and shallow sea, such as that within a large part of
the Malay Archipelago, where the depth varies from thirty or forty to sixty

fathoms, a widely extended formation might be formed during a period of
elevation, and yet not suffer excessively from denudation during its slow
upheaval; but the thickness of the formation could not be great, for owing
to the elevatory movement it would be less than the depth in which it was
formed; nor would the deposit be much consolidated, nor be capped by
overlying formations, 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. It has, however, been suggested by Mr. Hopkins,
that if one part of the area, after rising and before being denuded,
subsided, the deposit formed during the rising movement, though not thick,
might afterwards become protected by fresh accumulations, and thus be
preserved for a long period.

Mr. Hopkins also expresses his belief that sedimentary beds of considerable
horizontal extent have rarely been completely destroyed. But all
geologists, excepting the few who believe that our present metamorphic
schists and plutonic rocks once formed the primordial nucleus of the globe,
will admit that these latter rocks have been stripped of their covering to
an enormous extent. For it is scarcely possible that such rocks could have
been solidified and crystallised while uncovered; but if the metamorphic
action occurred at profound depths of the ocean, the former protecting
mantle of rock may not have been very thick. Admitting then that gneiss,
mica-schist, granite, diorite, etc., were once necessarily covered up, how
can we account for the naked and extensive areas of such rocks in many
parts of the world, 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. South
of the Amazon, Boue colours an area composed of rocks of this nature as
equal to that of Spain, France, Italy, part of Germany, and the British
Islands, all conjoined. This region has not been carefully explored, but
from the concurrent testimony of travellers, the granitic area is very
large: thus Von Eschwege gives a detailed section of these rocks,
stretching from Rio de Janeiro for 260 geographical miles inland in a
straight line; and I travelled for 150 miles in another direction, and saw
nothing but granitic rocks. Numerous specimens, collected along the whole
coast, from near Rio de Janeiro to the mouth of the Plata, a distance of
1,100 geographical miles, were examined by me, and they all belonged to
this class. Inland, along the whole northern bank of the Plata, I saw,
besides modern tertiary beds, only one small patch of slightly
metamorphosed rock, which alone could have formed a part of the original
capping of the granitic series. Turning to a well-known region, namely, to
the United States and Canada, as shown in Professor H.D. Rogers' beautiful
map, I have estimated the areas by cutting out and weighing the paper, and
I find that the metamorphic (excluding the semi-metamorphic") and granite
rocks exceedin the proportion of 19 to 12.5the whole of the newer
Palaeozoic formations. In many regions the metamorphic and granite rocks
would be found much more widely extended than they appear to beif all the
sedimentary beds were removed which rest unconformably on themand which
could not have formed part of the original mantle under which they were
crystallised. Henceit is probable that in some parts of the world whole
formations have been completely denudedwith not a wreck left behind.

One remark is here worth a passing notice. During periods of elevation the
area of the land and of the adjoining shoal parts of the sea will be
increased and new stations will often be formed--all circumstances
favourableas previously explainedfor the formation of new varieties and
species; but during such periods there will generally be a blank in the
geological record. On the other handduring subsidencethe inhabited
area and number of inhabitants will decrease (excepting on the shores of a
continent when first broken up into an archipelago)and consequently
during subsidencethough there will be much extinctionfew new varieties
or species will be formed; and it is during these very periods of

subsidence that the deposits which are richest in fossils have been


>From these several considerations it cannot be doubted that the geological
recordviewed as a wholeis extremely imperfect; but if we confine our
attention to any one formationit becomes much more difficult to
understand why we do not therein find closely graduated varieties between
the allied species which lived at its commencement and at its close.
Several cases are on record of the same species presenting varieties in the
upper and lower parts of the same formation. Thus Trautschold gives a
number of instances with Ammonitesand 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. Although each
formation has indisputably required a vast number of years for its
depositionseveral reasons can be given why each should not commonly
include a graduated series of links between the species which lived at its
commencement and closebut I cannot assign due proportional weight to the
following considerations.

Although each formation may mark a very long lapse of yearseach probably
is short compared with the period requisite to change one species into
another. I am aware that two palaeontologistswhose opinions are worthy
of much deferencenamely Bronn and Woodwardhave concluded that the
average duration of each formation is twice or thrice as long as the
average duration of specific forms. But insuperable difficultiesas it
seems to meprevent us from coming to any just conclusion on this head.
When we see a species first appearing in the middle of any formationit
would be rash in the extreme to infer that it had not elsewhere previously
existed. So againwhen we find a species disappearing before the last
layers have been depositedit would be equally rash to suppose that it
then became extinct. We forget how small the area of Europe is compared
with the rest of the world; nor have the several stages of the same
formation throughout Europe been correlated with perfect accuracy.

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; and when we
see a species first appearing in any formationthe probability is that it
only then first immigrated into that area. It is well knownfor instance
that several species appear somewhat earlier in the palaeozoic beds of
North America than in those of Europe; time having apparently been required
for their migration from the American to the European seas. In examining
the latest depositsin various quarters of the worldit has everywhere
been notedthat some few still existing species are common in the deposit
but have become extinct in the immediately surrounding sea; orconversely
that some are now abundant in the neighbouring seabut are rare or absent
in this particular deposit. It is an excellent lesson to reflect on the
ascertained amount of migration of the inhabitants of Europe during the
glacial epochwhich forms only a part of one whole geological period; and
likewise to reflect on the changes of levelon the extreme change of
climateand on the great lapse of timeall included within this same
glacial period. Yet it may be doubted whetherin any quarter of the
worldsedimentary depositsINCLUDING FOSSIL REMAINShave gone on
accumulating within the same area during the whole of this period. It is
notfor instanceprobable that sediment was deposited during the whole of
the glacial period near the mouth of the Mississippiwithin 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. 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
upraisedorganic remains will probably first appear and disappear at
different levelsowing to the migrations of species and to geographical
changes. And in the distant futurea geologistexamining these beds

would be tempted to conclude that the average duration of life of the
embedded fossils had been less than that of the glacial periodinstead of
having been really far greaterthat isextending from before the glacial
epoch to the present day.

In order to get a perfect gradation between two forms in the upper and
lower parts of the same formationthe deposit must have gone on
continuously accumulating during a long periodsufficient for the slow
process of modification; hencethe deposit must be a very thick one; and
the species undergoing change must have lived in the same district
throughout the whole time. But we have seen that a thick formation
fossiliferous throughout its entire thicknesscan accumulate only during a
period of subsidence; and to keep the depth approximately the samewhich
is necessary that the same marine species may live on the same spacethe
supply of sediment must nearly counterbalance the amount of subsidence.
But this same movement of subsidence will tend to submerge the area whence
the sediment is derivedand thus diminish the supplywhilst the downward
movement continues. In factthis nearly exact balancing between the
supply of sediment and the amount of subsidence is probably a rare
contingency; for it has been observed by more than one palaeontologist that
very thick deposits are usually barren of organic remainsexcept near
their upper or lower limits.

It would seem that each separate formationlike the whole pile of
formations in any countryhas generally been intermittent in its
accumulation. When we seeas is so often the casea formation composed
of beds of widely different mineralogical compositionwe may reasonably
suspect that the process of deposition has been more or less interrupted.
Nor will the closest inspection of a formation give us any idea of the
length of time which its deposition may have consumed. Many instances
could be given of bedsonly a few feet in thicknessrepresenting
formations which are elsewhere thousands of feet in thicknessand which
must have required an enormous period for their accumulation; yet no one
ignorant of this fact would have even suspected the vast lapse of time
represented by the thinner formation. Many cases could be given of the
lower beds of a formation having been upraiseddenudedsubmergedand
then re-covered by the upper beds of the same formation--factsshowing
what wideyet easily overlookedintervals have occurred in its
accumulation. In other cases we have the plainest evidence in great
fossilised treesstill standing upright as they grewof many long
intervals of time and changes of level during the process of deposition
which would not have been suspectedhad not the trees been preserved:
thus Sir C. Lyell and Dr. Dawson found carboniferous beds 1400 feet thick
in Nova Scotiawith ancient root-bearing strataone above the otherat
no less than sixty-eight different levels. Hencewhen the same species
occurs at the bottommiddleand top of a formationthe probability is
that it has not lived on the same spot during the whole period of
depositionbut has disappeared and reappearedperhaps many timesduring
the same geological period. Consequently if it were to undergo a
considerable amount of modification during the deposition of any one
geological formationa section would not include all the fine intermediate
gradations which must on our theory have existedbut abruptthough
perhaps slightchanges of form.

It is all-important to remember that naturalists have no golden rule by
which to distinguish species and varieties; they grant some little
variability to each speciesbut when they meet with a somewhat greater
amount of difference between any two formsthey rank both as species
unless they are enabled to connect them together by the closest
intermediate gradations; and thisfrom the reasons just assignedwe can
seldom hope to effect in any one geological section. Supposing B and C to
be two speciesand a thirdAto be found in an older and underlying bed;
even if A were strictly intermediate between B and Cit would simply be
ranked as a third and distinct speciesunless at the same time it could be

closely connected by intermediate varieties with either one or both forms.
Nor should it be forgottenas before explainedthat A might be the actual
progenitor of B and Cand yet would not necessarily be strictly
intermediate between them in all respects. So that we might obtain the
parent-species and its several modified descendants from the lower and
upper beds of the same formationand unless we obtained numerous
transitional gradationswe should not recognise their blood-relationship
and should consequently rank them as distinct species.

It is notorious on what excessively slight differences many
palaeontologists have founded their species; and they do this the more
readily if the specimens come from different sub-stages of the same
formation. Some experienced conchologists are now sinking many of the very
fine species of D'Orbigny and others into the rank of varieties; and on
this view we do find the kind of evidence of change which on the theory we
ought to find. Look again at the later tertiary depositswhich include
many shells believed by the majority of naturalists to be identical with
existing species; but some excellent naturalistsas Agassiz and Pictet
maintain that all these tertiary species are specifically distinctthough
the distinction is admitted to be very slight; so that hereunless we
believe that these eminent naturalists have been misled by their
imaginationsand that these late tertiary species really present no
difference whatever from their living representativesor unless we admit
in opposition to the judgment of most naturaliststhat these tertiary
species are all truly distinct from the recentwe have evidence of the
frequent occurrence of slight modifications of the kind required. If we
look to rather wider intervals of timenamelyto distinct but consecutive
stages of the same great formationwe find that the embedded fossils
though universally ranked as specifically differentyet are far more
closely related to each other than are the species found in more widely
separated formations; so that here again we have undoubted evidence of
change in the direction required by the theory; but to this latter subject
I shall return in the following chapter.

With animals and plants that propagate rapidly and do not wander much
there is reason to suspectas we have formerly seenthat their varieties
are generally at first local; 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. According to this viewthe chance
of discovering in a formation in any one country all the early stages of
transition between any two formsis smallfor the successive changes are
supposed to have been local or confined to some one spot. Most marine
animals have a wide range; and we have seen that with plants it is those
which have the widest rangethat oftenest present varietiesso thatwith
shells and other marine animalsit is probable that those which had the
widest rangefar exceeding the limits of the known geological formations
in Europehave oftenest given risefirst to local varieties and
ultimately to new species; and this again would greatly lessen the chance
of our being able to trace the stages of transition in any one geological

It is a more important considerationleading to the same resultas lately
insisted on by Dr. Falconernamelythat the period during which each
species underwent modificationthough long as measured by yearswas
probably short in comparison with that during which it remained without
undergoing any change.

It should not be forgottenthat at the present daywith perfect specimens
for examinationtwo forms can seldom be connected by intermediate
varietiesand thus proved to be the same speciesuntil many specimens are
collected from many places; and with fossil species this can rarely be
done. We shallperhapsbest perceive the improbability of our being
enabled to connect species by numerousfineintermediatefossil links
by asking ourselves whetherfor instancegeologists at some future period

will be able to prove that our different breeds of cattlesheephorses
and dogs are descended from a single stock or from several aboriginal
stocks; oragainwhether certain sea-shells inhabiting the shores of
North Americawhich are ranked by some conchologists as distinct species
from their European representativesand by other conchologists as only
varietiesare really varietiesor areas it is calledspecifically
distinct. This could be effected by the future geologist only by his
discovering in a fossil state numerous intermediate gradations; and such
success is improbable in the highest degree.

It has been asserted over and over againby writers who believe in the
immutability of speciesthat geology yields no linking forms. This
assertionas we shall see in the next chapteris certainly erroneous. As
Sir J. Lubbock has remarkedEvery species is a link between other allied
forms.If we take a genus having a score of speciesrecent and extinct
and destroy four-fifths of themno one doubts that the remainder will
stand much more distinct from each other. If the extreme forms in the
genus happen to have been thus destroyedthe genus itself will stand more
distinct from other allied genera. What geological research has not
revealedis the former existence of infinitely numerous gradationsas
fine as existing varietiesconnecting together nearly all existing and
extinct species. But this ought not to be expected; yet this has been
repeatedly advanced as a most serious objection against my views.

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. The
Malay Archipelago is about the size of Europe from the North Cape to the
Mediterraneanand from Britain to Russia; and therefore equals all the
geological formations which have been examined with any accuracyexcepting
those of the United States of America. I fully agree with Mr. Godwin-
Austenthat the present condition of the Malay Archipelagowith its
numerous large islands separated by wide and shallow seasprobably
represents the former state of Europewhile most of our formations were
accumulating. The Malay Archipelago is one of the richest regions in
organic beings; yet if all the species were to be collected which have ever
lived therehow 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. Not many of the
strictly littoral animalsor of those which lived on naked submarine
rockswould be embedded; and those embedded in gravel or sand would not
endure to a distant epoch. Wherever sediment did not accumulate on the bed
of the seaor where it did not accumulate at a sufficient rate to protect
organic bodies from decayno remains could be preserved.

Formations rich in fossils of many kindsand of thickness sufficient to
last to an age as distant in futurity as the secondary formations lie in
the pastwould generally be formed in the archipelago only during periods
of subsidence. These periods of subsidence would be separated from each
other by immense intervals of timeduring which the area would be either
stationary or rising; whilst risingthe fossiliferous formations on the
steeper shores would be destroyedalmost as soon as accumulatedby the
incessant coast-actionas we now see on the shores of South America. Even
throughout the extensive and shallow seas within the archipelago
sedimentary beds could hardly be accumulated of great thickness during the
periods of elevationor become capped and protected by subsequent
depositsso as to have a good chance of enduring to a very distant future.
During the periods of subsidencethere would probably be much extinction
of life; during the periods of elevationthere would be much variation
but the geological record would then be less perfect.

It may be doubted whether the duration of any one great period of

subsidence over the whole or part of the archipelagotogether with a
contemporaneous accumulation of sedimentwould EXCEED the average duration
of the same specific forms; and these contingencies are indispensable for
the preservation of all the transitional gradations between any two or more
species. If such gradations were not all fully preservedtransitional
varieties would merely appear as so many newthough closely allied
species. It is also probable that each great period of subsidence would be
interrupted by oscillations of leveland that slight climatical changes
would intervene during such lengthy periods; and in these cases the
inhabitants of the archipelago would migrateand no closely consecutive
record of their modifications could be preserved in any one formation.

Very many of the marine inhabitants of the archipelago now range thousands
of miles beyond its confines; and analogy plainly leads to the belief that
it would be chiefly these far-ranging speciesthough only some of them
which would oftenest produce new varieties; and the varieties would at
first be local or confined to one placebut if possessed of any decided
advantageor when further modified and improvedthey would slowly spread
and supplant their parent-forms. When such varieties returned to their
ancient homesas they would differ from their former state in a nearly
uniformthough perhaps extremely slight degreeand as they would be found
embedded in slightly different sub-stages of the same formationthey
wouldaccording to the principles followed by many palaeontologistsbe
ranked as new and distinct species.

If then there be some degree of truth in these remarkswe have no right to
expect to findin our geological formationsan infinite number of those
fine transitional formswhichon our theoryhave connected all the past
and present species of the same group into one long and branching chain of
life. We ought only to look for a few linksand such assuredly we do
find--some more distantlysome more closelyrelated to each other; and
these linkslet them be ever so closeif found in different stages of the
same formationwouldby many palaeontologistsbe ranked as distinct
species. But I do not pretend that I should ever have suspected how poor
was the record in the best preserved geological sectionshad not the
absence of innumerable transitional links between the species which lived
at the commencement and close of each formationpressed so hardly on my


The abrupt manner in which whole groups of species suddenly appear in
certain formationshas been urged by several palaeontologists--for
instanceby AgassizPictetand Sedgwickas a fatal objection to the
belief in the transmutation of species. If numerous speciesbelonging to
the same genera or familieshave really started into life at oncethe
fact would be fatal to the theory of evolution through natural selection.
For the development by this means of a group of formsall of which are
descended from some one progenitormust have been an extremely slow
process; and the progenitors must have lived long before their modified
descendants. But we continually overrate the perfection of the geological
recordand falsely inferbecause certain genera or families have not been
found beneath a certain stagethat they did not exist before that stage.
In all cases positive palaeontological evidence may be implicitly trusted;
negative evidence is worthlessas experience has so often shown. We
continually forget how large the world iscompared with the area over
which our geological formations have been carefully examined; we forget
that groups of species may elsewhere have long existedand have slowly
multipliedbefore they invaded the ancient archipelagoes of Europe and the
United States. We do not make due allowance for the enormous intervals of
time which have elapsed between our consecutive formationslonger perhaps
in many cases than the time required for the accumulation of each
formation. These intervals will have given time for the multiplication of
species from some one parent-form: and in the succeeding formationsuch

groups or species will appear as if suddenly created.

I may here recall a remark formerly madenamelythat it might require a
long succession of ages to adapt an organism to some new and peculiar line
of lifefor instanceto fly through the air; and consequently that the
transitional forms would often long remain confined to some one region; but
thatwhen this adaptation had once been effectedand a few species had
thus acquired a great advantage over other organismsa comparatively short
time would be necessary to produce many divergent formswhich would spread
rapidly and widely throughout the world. Professor Pictetin his
excellent Review of this workin commenting on early transitional forms
and taking birds as an illustrationcannot see how the successive
modifications of the anterior limbs of a supposed prototype could possibly
have been of any advantage. But look at the penguins of the Southern
Ocean; 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; for they exist in infinite
numbers and of many kinds. I do not suppose that we here see the real
transitional grades through which the wings of birds have passed; but what
special difficulty is there in believing that it might profit the modified
descendants of the penguinfirst to become enabled to flap along the
surface of the sea like the logger-headed duckand ultimately to rise from
its surface and glide through the air?

I will now give a few examples to illustrate the foregoing remarksand to
show how liable we are to error in supposing that whole groups of species
have suddenly been produced. Even in so short an interval as that between
the first and second editions of Pictet's great work on Palaeontology
published in 1844-46 and in 1853-57the conclusions on the first
appearance and disappearance of several groups of animals have been
considerably modified; and a third edition would require still further
changes. I may recall the well-known fact that in geological treatises
published not many years agomammals were always spoken of as having
abruptly come in at the commencement of the tertiary series. And now one
of the richest known accumulations of fossil mammals belongs to the middle
of the secondary series; and true mammals have been discovered in the new
red sandstone at nearly the commencement of this great series. Cuvier used
to urge that no monkey occurred in any tertiary stratum; but now extinct
species have been discovered in IndiaSouth America and in Europeas far
back as the miocene stage. Had it not been for the rare accident of the
preservation of footsteps in the new red sandstone of the United States
who would have ventured to suppose that no less than at least thirty
different bird-like animalssome of gigantic sizeexisted during that
period? Not a fragment of bone has been discovered in these beds. Not
long agopalaeontologists maintained that the whole class of birds came
suddenly into existence during the eocene period; but now we knowon the
authority of Professor Owenthat a bird certainly lived during the
deposition of the upper greensand; and still more recentlythat strange
birdthe Archeopteryxwith a long lizard-like tailbearing a pair of
feathers on each jointand with its wings furnished with two free claws
has been discovered in the oolitic slates of Solenhofen. Hardly any recent
discovery shows more forcibly than this how little we as yet know of the
former inhabitants of the world.

I may give another instancewhichfrom having passed under my own eyes
has much struck me. In a memoir on Fossil Sessile CirripedesI stated
thatfrom the large number of existing and extinct tertiary species; from
the extraordinary abundance of the individuals of many species all over the
worldfrom the Arctic regions to the equatorinhabiting various zones of
depthsfrom the upper tidal limits to fifty fathoms; from the perfect
manner in which specimens are preserved in the oldest tertiary beds; from
the ease with which even a fragment of a valve can be recognised; from all
these circumstancesI inferred thathad sessile cirripedes existed during
the secondary periodsthey would certainly have been preserved and

discovered; and as not one species had then been discovered in beds of this
ageI concluded that this great group had been suddenly developed at the
commencement of the tertiary series. This was a sore trouble to me
addingas I then thoughtone more instance of the abrupt appearance of a
great group of species. But my work had hardly been publishedwhen a
skilful palaeontologistM. Bosquetsent me a drawing of a perfect
specimen of an unmistakable sessile cirripedewhich he had himself
extracted from the chalk of Belgium. Andas if to make the case as
striking as possiblethis cirripede was a Chthamalusa very common
largeand ubiquitous genusof which not one species has as yet been found
even in any tertiary stratum. Still more recentlya Pyrgomaa member of
a distinct subfamily of sessile cirripedeshas been discovered by Mr.
Woodward in the upper chalk; so that we now have abundant evidence of the
existence of this group of animals during the secondary period.

The case most frequently insisted on by palaeontologists of the apparently
sudden appearance of a whole group of speciesis that of the teleostean
fisheslow downaccording to Agassizin the Chalk period. This group
includes the large majority of existing species. But certain Jurassic and
Triassic forms are now commonly admitted to be teleostean; and even some
palaeozoic forms have thus been classed by one high authority. If the
teleosteans had really appeared suddenly in the northern hemisphere at the
commencement of the chalk formationthe fact would have been highly
remarkable; but it would not have formed an insuperable difficultyunless
it could likewise have been shown that at the same period the species were
suddenly and simultaneously developed in other quarters of the world. It
is almost superfluous to remark that hardly any fossil-fish are known from
south of the equator; and by running through Pictet's Palaeontology it will
be seen that very few species are known from several formations in Europe.
Some few families of fish now have a confined range; the teleostean fishes
might formerly have had a similarly confined rangeand after having been
largely developed in some one seahave spread widely. 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. Even at this dayif the Malay
Archipelago were converted into landthe tropical parts of the Indian
Ocean would form a large and perfectly enclosed basinin which any great
group of marine animals might be multiplied; and here they would remain
confineduntil some of the species became adapted to a cooler climateand
were enabled to double the southern capes of Africa or Australiaand thus
reach other and distant seas.

>From these considerationsfrom our ignorance of the geology of other
countries beyond the confines of Europe and the United Statesand from the
revolution in our palaeontological knowledge effected by the discoveries of
the last dozen yearsit seems to me to be about as rash to dogmatize on
the succession of organic forms throughout the worldas it would be for a
naturalist to land for five minutes on a barren point in Australiaand
then to discuss the number and range of its productions.


There is another and allied difficultywhich is much more serious. 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. Most of the arguments which have convinced me that
all the existing species of the same group are descended from a single
progenitorapply with equal force to the earliest known species. For
instanceit cannot be doubted that all the Cambrian and Silurian
trilobites are descended from some one crustaceanwhich must have lived
long before the Cambrian ageand which probably differed greatly from any
known animal. Some of the most ancient animalsas the NautilusLingula not differ much from living species; and it cannot on our theory
be supposedthat these old species were the progenitors of all the species

belonging to the same groups which have subsequently appearedfor they are
not in any degree intermediate in character.

Consequentlyif the theory be trueit is indisputable that before the
lowest Cambrian stratum was deposited long periods elapsedas long asor
probably far longer thanthe whole interval from the Cambrian age to the
present day; and that during these vast periods the world swarmed with
living creatures. Here we encounter a formidable objection; for it seems
doubtful whether the earthin a fit state for the habitation of living
creatureshas lasted long enough. Sir W. Thompson concludes that the
consolidation of the crust can hardly have occurred less than twenty or
more than four hundred million years agobut probably not less than
ninety-eight or more than two hundred million years. These very wide
limits show how doubtful the data are; and other elements may have
hereafter to be introduced into the problem. Mr. Croll estimates that
about sixty million years have elapsed since the Cambrian periodbut this
judging from the small amount of organic change since the commencement of
the Glacial epochappears a very short time for the many and great
mutations of lifewhich have certainly occurred since the Cambrian
formation; and the previous one hundred and forty million years can hardly
be considered as sufficient for the development of the varied forms of life
which already existed during the Cambrian period. It ishowever
probableas Sir William Thompson insiststhat the world at a very early
period was subjected to more rapid and violent changes in its physical
conditions than those now occurring; and such changes would have tended to
induce changes at a corresponding rate in the organisms which then existed.

To the question why we do not find rich fossiliferous deposits belonging to
these assumed earliest periods prior to the Cambrian systemI can give no
satisfactory answer. Several eminent geologistswith Sir R. Murchison at
their headwere until recently convinced that we beheld in the organic
remains of the lowest Silurian stratum the first dawn of life. Other
highly competent judgesas Lyell and E. Forbeshave disputed this
conclusion. We should not forget that only a small portion of the world is
known with accuracy. Not very long ago M. Barrande added another and lower
stageabounding with new and peculiar speciesbeneath the then known
Silurian system; and nowstill lower down in the Lower Cambrian formation
Mr Hicks has found South Wales beds rich in trilobitesand containing
various molluscs and annelids. The presence of phosphatic nodules and
bituminous mattereven in some of the lowest azotic rocksprobably
indicates life at these periods; and the existence of the Eozoon in the
Laurentian formation of Canada is generally admitted. There are three
great series of strata beneath the Silurian system in Canadain the lowest
of which the Eozoon is found. Sir W. Logan states that their "united
thickness may possibly far surpass that of all the succeeding rocksfrom
the base of the palaeozoic series to the present time. We are thus carried
back to a period so remotethat the appearance of the so-called primordial
fauna (of Barrande) may by some be considered as a comparatively modern
event." The Eozoon belongs to the most lowly organised of all classes of
animalsbut is highly organised for its class; it existed in countless
numbersandas Dr. Dawson has remarkedcertainly preyed on other minute
organic beingswhich must have lived in great numbers. Thus the words
which I wrote in 1859about the existence of living beings long before the
Cambrian periodand which are almost the same with those since used by Sir

W. Loganhave proved true. Neverthelessthe difficulty of assigning any
good reason for the absence of vast piles of strata rich in fossils beneath
the Cambrian system is very great. It does not seem probable that the most
ancient beds have been quite worn away by denudationor that their fossils
have been wholly obliterated by metamorphic actionfor if this had been
the case we should have found only small remnants of the formations next
succeeding them in ageand these would always have existed in a partially
metamorphosed condition. But the descriptions which we possess of the
Silurian deposits over immense territories in Russia and in North America
do not support the view that the older a formation is the more invariably

it has suffered extreme denudation and metamorphism.

The case at present must remain inexplicable; and may be truly urged as a
valid argument against the views here entertained. To show that it may
hereafter receive some explanationI will give the following hypothesis.
>From the nature of the organic remains which do not appear to have
inhabited profound depthsin the several formations of Europe and of the
United States; and from the amount of sedimentmiles in thicknessof
which the formations are composedwe may infer that from first to last
large islands or tracts of landwhence the sediment was derivedoccurred
in the neighbourhood of the now existing continents of Europe and North
America. This same view has since been maintained by Agassiz and others.
But we do not know what was the state of things in the intervals between
the several successive formations; whether Europe and the United States
during these intervals existed as dry landor as a submarine surface near
landon which sediment was not depositedor as the bed of an open and
unfathomable sea.

Looking to the existing oceanswhich are thrice as extensive as the land
we see them studded with many islands; but hardly one truly oceanic island
(with the exception of New Zealandif this can be called a truly oceanic
island) is as yet known to afford even a remnant of any palaeozoic or
secondary formation. Hencewe may perhaps inferthat during the
palaeozoic and secondary periodsneither continents nor continental
islands existed where our oceans now extend; for had they existed
palaeozoic and secondary formations would in all probability have been
accumulated from sediment derived from their wear and tear; and would have
been at least partially upheaved by the oscillations of levelwhich must
have intervened during these enormously long periods. Ifthenwe may
infer anything from these factswe may infer thatwhere our oceans now
extendoceans have extended from the remotest period of which we have any
record; and on the other handthat where continents now existlarge
tracts of land have existedsubjectedno doubtto great oscillations of
levelsince the Cambrian period. The coloured map appended to my volume
on Coral Reefsled me to conclude that the great oceans are still mainly
areas of subsidencethe great archipelagoes still areas of oscillations of
leveland the continents areas of elevation. But we have no reason to
assume that things have thus remained from the beginning of the world. Our
continents seem to have been formed by a preponderanceduring many
oscillations of levelof the force of elevation. But may not the areas of
preponderant movement have changed in the lapse of ages? At a period long
antecedent to the Cambrian epochcontinents may have existed where oceans
are now spread outand clear and open oceans may have existed where our
continents now stand. Nor should we be justified in assuming that iffor
instancethe bed of the Pacific Ocean were now converted into a continent
we should there find sedimentary formationsin recognisable condition
older than the Cambrian stratasupposing such to have been formerly
deposited; for it might well happen that strata which had subsided some
miles nearer to the centre of the earthand which had been pressed on by
an enormous weight of superincumbent watermight have undergone far more
metamorphic action than strata which have always remained nearer to the
surface. The immense areas in some parts of the worldfor instance in
South Americaof naked metamorphic rockswhich must have been heated
under great pressurehave always seemed to me to require some special
explanation; 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.

The several difficulties here discussednamelythatthough we find in
our geological formations many links between the species which now exist
and which formerly existedwe do not find infinitely numerous fine
transitional forms closely joining them all together. The sudden manner in
which several groups of species first appear in our European formations
the almost entire absenceas at present knownof formations rich in

fossils beneath the Cambrian strataare all undoubtedly of the most
serious nature. We see this in the fact that the most eminent
Forbesetc.and all our greatest geologistsas LyellMurchison
Sedgwicketc.have unanimouslyoften vehementlymaintained the
immutability of species. But Sir Charles Lyell now gives the support of
his high authority to the opposite sideand most geologists and
palaeontologists are much shaken in their former belief. Those who believe
that the geological record is in any degree perfectwill undoubtedly at
once reject my theory. For my partfollowing out Lyell's metaphorI look
at the geological record as a history of the world imperfectly kept and
written in a changing dialect. Of this history we possess the last volume
alonerelating only to two or three countries. Of this volumeonly here
and there a short chapter has been preservedand of each pageonly here
and there a few lines. Each word of the slowly-changing languagemore or
less different in the successive chaptersmay represent the forms of life
which are entombed in our consecutive formationsand which falsely appear
to have been abruptly introduced. On this view the difficulties above
discussed are greatly diminished or even disappear.



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

Let us now see whether the several facts and laws relating to the
geological succession of organic beings accord best with the common view of
the immutability of speciesor with that of their slow and gradual
modificationthrough variation and natural selection.

New species have appeared very slowlyone after anotherboth on the land
and in the waters. Lyell has shown that it is hardly possible to resist
the evidence on this head in the case of the several tertiary stages; and
every year tends to fill up the blanks between the stagesand to make the
proportion between the lost and existing forms more gradual. In some of
the most recent bedsthough undoubtedly of high antiquity if measured by
yearsonly one or two species are extinctand only one or two are new
having appeared there for the first timeeither locallyoras far as we
knowon the face of the earth. The secondary formations are more broken;
butas Bronn has remarkedneither the appearance nor disappearance of the
many species embedded in each formation has been simultaneous.

Species belonging to different genera and classes have not changed at the
same rateor in the same degree. In the older tertiary beds a few living
shells may still be found in the midst of a multitude of extinct forms.
Falconer has given a striking instance of a similar factfor an existing
crocodile is associated with many lost mammals and reptiles in the
sub-Himalayan deposits. The Silurian Lingula differs but little from the
living species of this genus; whereas most of the other Silurian Molluscs
and all the Crustaceans have changed greatly. The productions of the land
seem to have changed at a quicker rate than those of the seaof which a
striking instance has been observed in Switzerland. There is some reason
to believe that organisms high in the scalechange more quickly than those
that are low: though there are exceptions to this rule. The amount of
organic changeas Pictet has remarkedis not the same in each successive

so-called formation. Yet if we compare any but the most closely related
formationsall the species will be found to have undergone some change.
When a species has once disappeared from the face of the earthwe have no
reason to believe that the same identical form ever reappears. The
strongest apparent exception to this latter rule is that of the so-called
coloniesof M. Barrandewhich intrude for a period in the midst of an
older formationand then allow the pre-existing fauna to reappear; but
Lyell's explanationnamelythat it is a case of temporary migration from
a distinct geographical provinceseems satisfactory.

These several facts accord well with our theorywhich includes no fixed
law of developmentcausing all the inhabitants of an area to change
abruptlyor simultaneouslyor to an equal degree. The process of
modification must be slowand 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
degreethus causing a greater or less amount of permanent modification
will depend on many complex contingencies--on the variations being of a
beneficial natureon the freedom of intercrossingon the slowly changing
physical conditions of the countryon 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; orif
changingshould change in a less degree. We find similar relations
between the existing inhabitants of distinct countries; for instancethe
land-shells and coleopterous insects of Madeira have come to differ
considerably from their nearest allies on the continent of Europewhereas
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 productionsby
the more complex relations of the higher beings to their organic and
inorganic conditions of lifeas explained in a former chapter. When many
of the inhabitants of any area have become modified and improvedwe can
understandon the principle of competitionand from the all-important
relations of organism to organism in the struggle for lifethat any form
which did not become in some degree modified and improvedwould be liable
to extermination. Hencewe see why all the species in the same region do
at lastif we look to long enough intervals of timebecome modified; for
otherwise they would become extinct.

In members of the same class the average amount of changeduring long and
equal periods of timemayperhapsbe nearly the same; but as the
accumulation of enduring formationsrich in fossilsdepends on great
masses of sediment being deposited on subsiding areasour 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
formationon this viewdoes not mark a new and complete act of creation
but only an occasional scenetaken almost at hazardin an ever slowly
changing drama.

We can clearly understand why a species when once lost should never
reappeareven if the very same conditions of lifeorganic 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 natureand 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 instanceit is possibleif all our fantail-pigeons were
destroyedthat 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 offspringit is
incredible that a fantailidentical with the existing breedcould be
raised from any other species of pigeonor even from any other well
established race of the domestic pigeonfor the successive variations
would almost certainly be in some degree differentand the newly-formed
variety would probably inherit from its progenitor some characteristic

Groups of speciesthat isgenera and familiesfollow the same general
rules in their appearance and disappearance as do single specieschanging
more or less quicklyand in a greater or lesser degree. A groupwhen it
has once disappearednever reappears; that isits existenceas long as
it lastsis continuous. I am aware that there are some apparent
exceptions to this rulebut the exceptions are surprisingly fewso few
that E. ForbesPictetand 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 grouphowever long it may
have lastedare the modified descendants one from the otherand all from
a common progenitor. In the genus Lingulafor instancethe species which
have successively appeared at all ages must have been connected by an
unbroken series of generationsfrom 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 factwhich if true would be fatal to my views.
But such cases are certainly exceptional; the general rule being a gradual
increase in numberuntil the group reaches its maximumand thensooner
or latera gradual decrease. If the number of the species included within
a genusor the number of the genera within a familybe represented by a
vertical line of varying thicknessascending through the successive
geological formationsin which the species are foundthe line will
sometimes falsely appear to begin at its lower endnot in a sharp point
but abruptly; it then gradually thickens upwardsoften keeping of equal
thickness for a spaceand ultimately thins out in the upper bedsmarking
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 genusand the genera of the same familycan
increase only slowly and progressively; the process of modification and the
production of a number of allied forms necessarily being a slow and gradual
processone species first giving rise to two or three varietiesthese
being slowly converted into specieswhich in their turn produce by equally
slow steps other varieties and speciesand so onlike the branching of a
great tree from a single stemtill the group becomes large.


We have as yet only spokesn incidentally of the disappearance of species
and of groups of species. On the theory of natural selectionthe
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 upeven by those geologistsas Elie de Beaumont
MurchisonBarrandeetc.whose general views would naturally lead them to
this conclusion. On the contrarywe have every reason to believefrom
the study of the tertiary formationsthat species and groups of species
gradually disappearone after anotherfirst from one spotthen from
anotherand finally from the world. In some few caseshoweveras by the
breaking of an isthmus and the consequent irruption of a multitude of new
inhabitants into an adjoining seaor 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 groupsas we
have seenhave 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 representedas
beforeby a vertical line of varying thickness the line is found to taper
more gradually at its upper endwhich marks the progress of extermination
than at its lower endwhich marks the first appearance and the early
increase in number of the species. In some caseshoweverthe
extermination of whole groupsas of ammonitestowards the close of the
secondary periodhas been wonderfully sudden.

The extinction of species has been involved in the most gratuitous mystery.
Some authors have even supposed thatas the individual has a definite
length of lifeso 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
MegatheriumToxodon and other extinct monsterswhich all co-existed with
still living shells at a very late geological periodI was filled with
astonishment; forseeing that the horsesince its introduction by the
Spaniards into South Americahas run wild over the whole country and has
increased in numbers at an unparalleled rateI 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 tooththough so like that of the existing
horsebelonged to an extinct species. Had this horse been still living
but in some degree rareno naturalist would have felt the least surprise
at its rarity; for rarity is the attribute of a vast number of species of
all classesin all countries. If we ask ourselves why this or that
species is rarewe answer that something is unfavourable in its conditions
of life; but what that something iswe can hardly ever tell. On the
supposition of the fossil horse still existing as a rare specieswe might
have felt certainfrom the analogy of all other mammalseven of the
slow-breeding elephantand from the history of the naturalisation of the
domestic horse in South Americathat 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
increasewhether some one or several contingenciesand at what period of
the horse's lifeand in what degree they severally acted. If the
conditions had gone onhowever slowlybecoming less and less favourable
we assuredly should not have perceived the factyet the fossil horse would
certainly have become rarer and rarerand 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 understoodthat 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 sizeon the contrary
would in some cases determineas has been remarked by Owenquicker
exterminationfrom the greater amount of requisite food. Before man
inhabited India or Africasome cause must have checked the continued
increase of the existing elephant. A highly capable judgeDr. Falconer
believes that it is chiefly insects whichfrom incessantly harassing and
weakening the elephant in Indiacheck 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 exterminatedeither locally or wholly
through man's agency. I may repeat what I published in 1845namelythat

to admit that species generally become rare before they become extinct--to
feel no surprise at the rarity of a speciesand yet to marvel greatly when
the species ceases to existis much the same as to admit that sickness in
the individual is the forerunner of death--to feel no surprise at sickness
butwhen the sick man diesto 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 speciesis 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 raisedit at first supplants the less improved
varieties in the same neighbourhood; when much improved it is transported
far and nearlike our short-horn cattleand takes the place of other
breeds in other countries. Thus the appearance of new forms and the
disappearance of old formsboth those naturally and artificially produced
are bound together. In flourishing groupsthe 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
increasingat least during the later geological epochsso thatlooking
to later timeswe 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 severeas formerly explained and
illustrated by examplesbetween 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 speciesthe nearest allies of
that speciesi.e. the species of the same genuswill be the most liable
to extermination. Thusas I believea number of new species descended
from one speciesthat is a new genuscomes 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 groupand thus have caused its
extermination. If many allied forms be developed from the successful
intrudermany will have to yield their places; and it will generally be
the allied formswhich will suffer from some inherited inferiority in
common. But whether it be species belonging to the same or to a distinct
classwhich have yielded their places to other modified and improved
speciesa few of the sufferers may often be preserved for a long time
from being fitted to some peculiar line of lifeor from inhabiting some
distant and isolated stationwhere they will have escaped severe
competition. For instancesome species of Trigoniaa great genus of
shells in the secondary formationssurvive in the Australian seas; and a
few members of the great and almost extinct group of Ganoid fishes still
inhabit our fresh waters. Thereforethe utter extinction of a group is
generallyas we have seena slower process than its production.

With respect to the apparently sudden extermination of whole families or
ordersas of Trilobites at the close of the palaeozoic periodand of
Ammonites at the close of the secondary periodwe 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. Moreoverwhenby sudden immigration or by unusually
rapid developmentmany species of a new group have taken possession of an
areamany of the older species will have been exterminated in a
correspondingly rapid manner; and the forms which thus yield their places
will commonly be alliedfor they will partake of the same inferiority in

Thusas it seems to methe 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 marvellet 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 actionyet seldom perceived by usthe
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;
thenand not until thenwe may justly feel surprise why we cannot account
for the extinction of any particular species or group of species.


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 climateswhere not a fragment of the mineral
chalk itself can be found; namelyin North Americain equatorial South
Americain Tierra del Fuegoat the Cape of Good Hopeand in the
peninsula of India. For at these distant pointsthe 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 samebut they belong to the same families
generaand sections of generaand sometimes are similarly characterised
in such trifling points as mere superficial sculpture. Moreoverother
formswhich are not found in the Chalk of Europebut which occur in the
formations either above or belowoccur in the same order at these distant
points of the world. In the several successive palaeozoic formations of
RussiaWestern Europe and North Americaa similar parallelism in the
forms of life has been observed by several authors; so it isaccording to
Lyellwith 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 viewthe general parallelism in the successive forms of
lifein the palaeozoic and tertiary stageswould still be manifestand
the several formations could be easily correlated.

These observationshoweverrelate 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 MegatheriumMylodon
Macraucheniaand Toxodon had been brought to Europe from La Platawithout
any information in regard to their geological positionno 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 Horseit 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 worldit must not be supposed that this
expression relates to the same yearor even to the same centuryor even
that it has a very strict geological sense; for if all the marine animals
now living in Europeand all those that lived in Europe during the
pleistocene period (a very remote period as measured by yearsincluding
the whole glacial epoch) were compared with those now existing in South
America or in Australiathe 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. Soagain
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 stagesthan to the present inhabitants
of Europe; and if this be soit 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. Neverthelesslooking to a
remotely future epochthere can be little doubt that all the more modern

MARINE formationsnamelythe upper pliocenethe pleistocene and strictly
modern beds of EuropeNorth and South Americaand Australiafrom
containing fossil remains in some degree alliedand from not including
those forms which are found only in the older underlying depositswould be
correctly ranked as simultaneous in a geological sense.

The fact of the forms of life changing simultaneously in the above large
senseat distant parts of the worldhas greatly struck those admirable
observersMM. de Verneuil and d'Archiac. After referring to the
parallelism of the palaeozoic forms of life in various parts of Europe
they addIf 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 isindeedquite futile
to look to changes of currentsclimateor other physical conditionsas
the cause of these great mutations in the forms of life throughout the
worldunder the most different climates. We mustas Barrande has
remarkedlook to some special law. We shall see this more clearly when we
treat of the present distribution of organic beingsand 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 worldis explicable on the theory of natural selection. New species
are formed by having some advantage over older forms; and the formswhich
are already dominantor have some advantage over the other forms in their
own countrygive birth to the greatest number of new varieties or
incipient species. We have distinct evidence on this headin the plants
which are dominantthat iswhich are commonest and most widely diffused
producing the greatest number of new varieties. It is also natural that
the dominantvarying and far-spreading specieswhich have already
invadedto a certain extentthe territories of other speciesshould be
those which would have the best chance of spreading still furtherand of
giving rise in new countries to other new varieties and species. The
process of diffusion would often be very slowdepending on climatal and
geographical changeson strange accidentsand on the gradual
acclimatization of new species to the various climates through which they
might have to passbut in the course of time the dominant forms would
generally succeed in spreading and would ultimately prevail. The diffusion
wouldit is probablebe slower with the terrestrial inhabitants of
distinct continents than with the marine inhabitants of the continuous sea.
We might therefore expect to findas we do finda less strict degree of
parallelism in the succession of the productions of the land than with
those of the sea.

Thusas it seems to methe parallelandtaken in a large sense
simultaneoussuccession 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 dominantowing to their having had some
advantage over their already dominant parentsas well as over other
species; and again spreadingvaryingand producing new forms. The old
forms which are beaten and which yield their places to the new and
victorious formswill generally be allied in groupsfrom inheriting some
inferiority in common; andthereforeas new and improved groups spread
throughout the worldold 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 formationsrich in

fossilswere deposited during periods of subsidence; and that blank
intervals of vast durationas far as fossils are concernedoccurred
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 extinctionand that there was much migration from other
parts of the world. As we have reason to believe that large areas are
affected by the same movementit is probable that strictly contemporaneous
formations have often been accumulated over very wide spaces in the same
quarter of the world; but we are very far from having any right to conclude
that this has invariably been the caseand that large areas have
invariably been affected by the same movements. When two formations have
been deposited in two regions during nearlybut not exactlythe same
periodwe should find in bothfrom the causes explained in the foregoing
paragraphsthe same general succession in the forms of life; but the
species would not exactly correspond; for there will have been a little
more time in the one region than in the other for modificationextinction
and immigration.

I suspect that cases of this nature occur in Europe. Mr. Prestwichin his
admirable Memoirs on the eocene deposits of England and Franceis able to
draw a close general parallelism between the successive stages in the two
countries; but when he compares certain stages in England with those in
Francealthough he finds in both a curious accordance in the numbers of
the species belonging to the same generayet the species themselves differ
in a manner very difficult to account for considering the proximity of the
two areasunlessindeedit be assumed that an isthmus separated two seas
inhabited by distinctbut contemporaneous faunas. Lyell has made similar
observations on some of the later tertiary formations. Barrandealso
shows that there is a striking general parallelism in the successive
Silurian deposits of Bohemia and Scandinavia; nevertheless he finds a
surprising amount of difference in the species. 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; in this case the several formations in the
two regions could be arranged in the same orderin accordance with the
general succession of the forms of lifeand the order would falsely appear
to be strictly parallel; nevertheless the species would not all be the same
in the apparently corresponding stages in the two regions.


Let us now look to the mutual affinities of extinct and living species.
All fall into a few grand classes; and this fact is at once explained on
the principle of descent. The more ancient any form isthe moreas a
general ruleit differs from living forms. Butas Buckland long ago
remarkedextinct species can all be classed either in still existing
groupsor between them. That the extinct forms of life help to fill up
the intervals between existing generafamiliesand ordersis certainly
true; but as this statement has often been ignored or even deniedit may
be well to make some remarks on this subjectand to give some instances.
If we confine our attention either to the living or to the extinct species
of the same classthe series is far less perfect than if we combine both
into one general system. In the writings of Professor Owen we continually
meet with the expression of generalised formsas applied to extinct
animals; and in the writings of Agassizof prophetic or synthetic types;
and these terms imply that such forms arein factintermediate or
connecting links. Another distinguished palaeontologistM. Gaudryhas
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. Cuvier ranked the Ruminants and Pachyderms as two of the

most distinct orders of mammals; but so many fossil links have been
disentombed that Owen has had to alter the whole classificationand has
placed certain Pachyderms in the same sub-order with ruminants; for
examplehe dissolves by gradations the apparently wide interval between
the pig and the camel. The Ungulata or hoofed quadrupeds are now divided
into the even-toed or odd-toed divisions; but the Macrauchenia of South
America connects to a certain extent these two grand divisions. No one
will deny that the Hipparion is intermediate between the existing horse and
certain other ungulate forms. What a wonderful connecting link in the
chain of mammals is the Typotherium from South Americaas the name given
to it by Professor Gervais expressesand which cannot be placed in any
existing order. The Sirenia form a very distinct group of the mammalsand
one of the most remarkable peculiarities in existing dugong and lamentin is
the entire absence of hind limbswithout even a rudiment being left; but
the extinct Halitherium hadaccording to Professor Floweran ossified
thigh-bone "articulated to a well-defined acetabulum in the pelvis and it
thus makes some approach to ordinary hoofed quadrupeds, to which the
Sirenia are in other respects allied. The cetaceans or whales are widely
different from all other mammals, but the tertiary Zeuglodon and Squalodon,
which have been placed by some naturalists in an order by themselves, are
considered by Professor Huxley to be undoubtedly cetaceans, and to
constitute connecting links with the aquatic carnivora."

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
manneron the one handby the ostrich and extinct Archeopteryxand on
the other hand by the Compsognathusone of the Dinosaurians--that group
which includes the most gigantic of all terrestrial reptiles. Turning to
the InvertebrataBarrande assertsa higher authority could not be named
that he is every day taught thatalthough palaeozoic animals can certainly
be classed under existing groupsyet that at this ancient period the
groups were not so distinctly separated from each other as they now are.

Some writers have objected to any extinct speciesor group of species
being considered as intermediate between any two living speciesor groups
of species. If by this term it is meant that an extinct form is directly
intermediate in all its characters between two living forms or groupsthe
objection is probably valid. But in a natural classification many fossil
species certainly stand between living speciesand some extinct genera
between living generaeven between genera belonging to distinct families.
The most common caseespecially with respect to very distinct groupssuch
as fish and reptilesseems to be thatsupposing them to be distinguished
at the present day by a score of charactersthe ancient members are
separated by a somewhat lesser number of charactersso that the two groups
formerly made a somewhat nearer approach to each other than they now do.

It is a common belief that the more ancient a form isby so much the more
it tends to connect by some of its characters groups now widely separated
from each other. This remark no doubt must be restricted to those groups
which have undergone much change in the course of geological ages; and it
would be difficult to prove the truth of the propositionfor every now and
then even a living animalas the Lepidosirenis discovered having
affinities directed towards very distinct groups. Yet if we compare the
older Reptiles and Batrachiansthe older Fishthe older Cephalopodsand
the eocene Mammalswith the recent members of the same classeswe must
admit that there is truth in the remark.

Let us see how far these several facts and inferences accord with the
theory of descent with modification. As the subject is somewhat complexI
must request the reader to turn to the diagram in the fourth chapter. We
may suppose that the numbered letters in italics represent generaand the
dotted lines diverging from them the species in each genus. The diagram is
much too simpletoo few genera and too few species being givenbut this
is unimportant for us. The horizontal lines may represent successive

geological formationsand all the forms beneath the uppermost line may be
considered as extinct. The three existing generaa14q14p14will form
a small family; b14 and f14a closely allied family or subfamily; and o14
i14m14a third family. These three familiestogether with the many
extinct genera on the several lines of descent diverging from the parent
form (A) will form an order; for all will have inherited something in
common from their ancient progenitor. On the principle of the continued
tendency to divergence of characterwhich was formerly illustrated by this
diagramthe more recent any form is the more it will generally differ from
its ancient progenitor. Hencewe can understand the rule that the most
ancient fossils differ most from existing forms. We must nothowever
assume that divergence of character is a necessary contingency; it depends
solely on the descendants from a species being thus enabled to seize on
many and different places in the economy of nature. Therefore it is quite
possibleas we have seen in the case of some Silurian formsthat a
species might go on being slightly modified in relation to its slightly
altered conditions of lifeand yet retain throughout a vast period the
same general characteristics. This is represented in the diagram by the
letter F14.

All the many formsextinct and recentdescended from (A)makeas before
remarkedone order; and this orderfrom the continued effects of
extinction and divergence of characterhas become divided into several
sub-families and familiessome of which are supposed to have perished at
different periodsand some to have endured to the present day.

By looking at the diagram we can see that if many of the extinct forms
supposed to be embedded in the successive formationswere discovered at
several points low down in the seriesthe three existing families on the
uppermost line would be rendered less distinct from each other. Iffor
instancethe genera a1a5a10f8m3m6m9were disinterredthese
three families would be so closely linked together that they probably would
have to be united into one great familyin nearly the same manner as has
occurred with ruminants and certain pachyderms. Yet he who objected to
consider as intermediate the extinct generawhich thus link together the
living genera of three familieswould be partly justifiedfor they are
intermediatenot directlybut only by a long and circuitous 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 instanceabove No. VI.--but none from beneath this line
then only two of the families (those on the left hand a14etc.and b14
etc.) would have to be united into one; and there would remain two families
which would be less distinct from each other than they were before the
discovery of the fossils. So againif the three families formed of eight
genera (a14 to m14)on the uppermost linebe supposed to differ from each
other by half-a-dozen important charactersthen the families which existed
at a period marked VI would certainly have differed from each other by a
less number of characters; for they would at this early stage of descent
have diverged in a less degree from their common progenitor. Thus it comes
that ancient and extinct genera are often in a greater or less degree
intermediate in character between their modified descendantsor between
their collateral relations.

Under nature the process will be far more complicated than is represented
in the diagram; for the groups will have been more numerous; they will have
endured for extremely unequal lengths of timeand will have been modified
in various degrees. As we possess only the last volume of the geological
recordand that in a very broken conditionwe have no right to expect
except in rare casesto fill up the wide intervals in the natural system
and thus to unite distinct families or orders. All that we have a right to
expect isthat those groups which havewithin known geological periods
undergone much modificationshould in the older formations make some
slight approach to each other; so that the older members should differ less
from each other in some of their characters than do the existing members of

the same groups; and this by the concurrent evidence of our best
palaeontologists is frequently the case.

Thuson the theory of descent with modificationthe main facts with
respect to the mutual affinities of the extinct forms of life to each other
and to living formsare explained in a satisfactory manner. And they are
wholly inexplicable on any other view.

On this same theoryit 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. 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 stageand are the
parents of those which became still more modified at the seventh stage;
hence they could hardly fail to be nearly intermediate in character between
the forms of life above and below. We musthoweverallow for the entire
extinction of some preceding formsand in any one region for the
immigration of new forms from other regionsand for a large amount of
modification during the long and blank intervals between the successive
formations. Subject to these allowancesthe fauna of each geological
period undoubtedly is intermediate in characterbetween the preceding and
succeeding faunas. I need give only one instancenamelythe manner in
which the fossils of the Devonian systemwhen this system was first
discoveredwere at once recognised by palaeontologists as intermediate in
character between those of the overlying carboniferous and underlying
Silurian systems. But each fauna is not necessarily exactly intermediate
as unequal intervals of time have elapsed between consecutive formations.

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 faunasthat certain genera offer exceptions to
the rule. For instancethe species of mastodons and elephantswhen
arranged by Dr. Falconer in two series--in the first place according to
their mutual affinitiesand in the second place according to their periods
of existence--do not accord in arrangement. The species extreme in
character are not the oldest or the most recent; nor are those which are
intermediate in characterintermediate in age. But supposing for an
instantin this and other such casesthat the record of the first
appearance and disappearance of the species was completewhich is far from
the casewe have no reason to believe that forms successively produced
necessarily endure for corresponding lengths of time. A very ancient form
may occasionally have lasted much longer than a form elsewhere subsequently
producedespecially in the case of terrestrial productions inhabiting
separated districts. To compare small things with great; if the principal
living and extinct races of the domestic pigeon were arranged in serial
affinitythis arrangement would not closely accord with the order in time
of their productionand even less with the order of their disappearance;
for the parent rock-pigeon still lives; and many varieties between the
rock-pigeon and the carrier have become extinct; and carriers which are
extreme in the important character of length of beak originated earlier
than short-beaked tumblerswhich are at the opposite end of the series in
this respect.

Closely connected with the statementthat the organic remains from an
intermediate formation are in some degree intermediate in characteris the
factinsisted on by all palaeontologiststhat fossils from two
consecutive formations are far more closely related to each otherthan are
the fossils from two remote formations. Pictet gives as a well-known
instancethe general resemblance of the organic remains from the several
stages of the Chalk formationthough the species are distinct in each
stage. This fact alonefrom its generalityseems to have shaken
Professor Pictet in his belief in the immutability of species. He who is
acquainted with the distribution of existing species over the globewill
not attempt to account for the close resemblance of distinct species in

closely consecutive formationsby the physical conditions of the ancient
areas having remained nearly the same. Let it be remembered that the forms
of lifeat least those inhabiting the seahave changed almost
simultaneously throughout the worldand therefore under the most different
climates and conditions. Consider the prodigious vicissitudes of climate
during the pleistocene periodwhich includes the whole glacial epochand
note how little the specific forms of the inhabitants of the sea have been

On the theory of descentthe full meaning of the fossil remains from
closely consecutive formationsbeing closely relatedthough ranked as
distinct speciesis obvious. As the accumulation of each formation has
often been interruptedand as long blank intervals have intervened between
successive formationswe ought not to expect to findas I attempted to
show in the last chapterin any one or in any two formationsall the
intermediate varieties between the species which appeared at the
commencement and close of these periods: but we ought to find after
intervalsvery long as measured by yearsbut only moderately long as
measured geologicallyclosely allied formsoras they have been called
by some authorsrepresentative species; and these assuredly we do find.
We findin shortsuch evidence of the slow and scarcely sensible
mutations of specific formsas we have the right to expect.


We have seen in the fourth chapter that the degree of differentiation and
specialisation of the parts in organic beingswhen arrived at maturityis
the best standardas yet suggestedof their degree of perfection or
highness. We have also seen thatas the specialisation of parts is an
advantage to each beingso natural selection will tend to render the
organisation of each being more specialised and perfectand in this sense
higher; not but that it may leave many creatures with simple and unimproved
structures fitted for simple conditions of lifeand in some cases will
even degrade or simplify the organisationyet leaving such degraded beings
better fitted for their new walks of life. In another and more general
mannernew species become superior to their predecessors; for they have to
beat in the struggle for life all the older formswith which they come
into close competition. 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 inhabitantsthe former would be beaten and
exterminated by the latteras would the secondary by the eoceneand the
palaeozoic by the secondary forms. So that by this fundamental test of
victory in the battle for lifeas well as by the standard of the
specialisation of organsmodern forms oughton the theory of natural
selectionto stand higher than ancient forms. Is this the case? A large
majority of palaeontologists would answer in the affirmative; and it seems
that this answer must be admitted as truethough difficult of proof.

It is no valid objection to this conclusionthat certain Brachiopods have
been but slightly modified from an extremely remote geological epoch; and
that certain land and fresh-water shells have remained nearly the same
from the time whenas far as is knownthey first appeared. It is not an
insuperable difficulty that Foraminifera have notas insisted on by Dr.
Carpenterprogressed in organisation since even the Laurentian epoch; for
some organisms would have to remain fitted for simple conditions of life
and what could be better fitted for this end than these lowly organised
Protozoa? Such objections as the above would be fatal to my viewif it
included advance in organisation as a necessary contingent. They would
likewise be fatalif the above Foraminiferafor instancecould be proved
to have first come into existence during the Laurentian epochor the above
Brachiopods during the Cambrian formation; for in this casethere would
not have been time sufficient for the development of these organisms up to
the standard which they had then reached. When advanced up to any given
pointthere is no necessityon the theory of natural selectionfor their

further continued process; though they willduring each successive age
have to be slightly modifiedso as to hold their places in relation to
slight changes in their conditions. The foregoing objections hinge on the
question whether we really know how old the world isand at what period
the various forms of life first appeared; and this may well be disputed.

The problem whether organisation on the whole has advanced is in many ways
excessively intricate. The geological recordat all times imperfectdoes
not extend far enough back to show with unmistakable clearness that within
the known history of the world organisation has largely advanced. Even at
the present daylooking to members of the same classnaturalists are not
unanimous which forms ought to be ranked as highest: thussome look at
the selaceans or sharksfrom their approach in some important points of
structure to reptilesas the highest fish; others look at the teleosteans
as the highest. The ganoids stand intermediate between the selaceans and
teleosteans; the latter at the present day are largely preponderant in
number; but formerly selaceans and ganoids alone existed; and in this case
according to the standard of highness chosenso will it be said that
fishes have advanced or retrograded in organisation. To attempt to compare
members of distinct types in the scale of highness seems hopeless; 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
although upon another type?" In the complex struggle for life it is quite
credible that crustaceansnot very high in their own classmight beat
cephalopodsthe highest molluscs; and such crustaceansthough not highly
developedwould stand very high in the scale of invertebrate animalsif
judged by the most decisive of all trials--the law of battle. Beside these
inherent difficulties in deciding which forms are the most advanced in
organisationwe 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
membershigh and lowat two periods. At an ancient epoch the highest and
lowest molluscoidal animalsnamelycephalopods and brachiopodsswarmed
in numbers; at the present time both groups are greatly reducedwhile
othersintermediate in organisationhave largely increased; consequently
some naturalists maintain that molluscs were formerly more highly developed
than at present; but a stronger case can be made out on the opposite side
by considering the vast reduction of brachiopodsand the fact that our
existing cephalopodsthough few in numberare more highly organised than
their ancient representatives. We ought also to compare the relative
proportional numbersat any two periodsof the high and low classes
throughout the world: iffor instanceat the present day fifty thousand
kinds of vertebrate animals existand if we knew that at some former
period only ten thousand kinds existedwe ought to look at this increase
in number in the highest classwhich implies a great displacement of lower
formsas a decided advance in the organisation of the world. We thus see
how hopelessly difficult it is to compare with perfect fairnessunder such
extremely complex relationsthe standard of organisation of the
imperfectly-known faunas of successive periods.

We shall appreciate this difficulty more clearly by looking to certain
existing faunas and floras. From the extraordinary manner in which
European productions have recently spread over New Zealandand have seized
on places which must have been previously occupied by the indigeneswe
must believethat if all the animals and plants of Great Britain were set
free in New Zealanda multitude of British forms would in the course of
time become thoroughly naturalized thereand would exterminate many of the
natives. On the other handfrom the fact that hardly a single inhabitant
of the southern hemisphere has become wild in any part of Europewe may
well doubt whetherif all the productions of New Zealand were set free in
Great Britainany considerable number would be enabled to seize on places
now occupied by our native plants and animals. Under this point of view
the productions of Great Britain stand much higher in the scale than those
of New Zealand. Yet the most skilful naturalistfrom an examination of

the species of the two countriescould not have foreseen this result.

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; and that the geological succession of
extinct forms is nearly parallel with the embryological development of
existing forms. This view accords admirably well with our theory. In a
future chapter I shall attempt to show that the adult differs from its
embryoowing to variations having supervened at a not early ageand
having been inherited at a corresponding age. This processwhilst it
leaves the embryo almost unalteredcontinually addsin the course of
successive generationsmore and more difference to the adult. Thus the
embryo comes to be left as a sort of picturepreserved by natureof the
former and less modified condition of the species. This view may be true
and yet may never be capable of proof. Seeingfor instancethat the
oldest known mammalsreptilesand fishes strictly belong to their proper
classesthough 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 dayit would be vain to look for animals having the common
embryological character of the Vertebratauntil beds rich in fossils are
discovered far beneath the lowest Cambrian strata--a discovery of which the
chance is small.


Mr. Clift many years ago showed that the fossil mammals from the Australian
caves were closely allied to the living marsupials of that continent. In
South Americaa similar relationship is manifesteven to an uneducated
eyein the gigantic pieces of armourlike those of the armadillofound
in several parts of La Plata; and Professor Owen has shown in the most
striking manner that most of the fossil mammalsburied there in such
numbersare related to South American types. This relationship is even
more clearly seen in the wonderful collection of fossil bones made by MM.
Lund and Clausen in the caves of Brazil. I was so much impressed with
these facts that I strongly insistedin 1839 and 1845on this "law of the
succession of types--on this wonderful relationship in the same
continent between the dead and the living." Professor Owen has
subsequently extended the same generalisation to the mammals of the Old
World. We see the same law in this author's restorations of the extinct
and gigantic birds of New Zealand. We see it also in the birds of the
caves of Brazil. Mr. Woodward has shown that the same law holds good with
sea-shellsbutfrom the wide distribution of most molluscsit is not
well displayed by them. Other cases could be addedas the relation
between the extinct and living land-shells of Madeira; and between the
extinct and living brackish water-shells of the Aralo-Caspian Sea.

Nowwhat does this remarkable law of the succession of the same types
within the same areas mean? He would be a bold man whoafter comparing
the present climate of Australia and of parts of South Americaunder the
same latitudewould attempt to accounton the one hand through dissimilar
physical conditionsfor the dissimilarity of the inhabitants of these two
continents; andon the other hand through similarity of conditionsfor
the uniformity of the same types in each continent during the later
tertiary periods. Nor can it be pretended that it is an immutable law that
marsupials should have been chiefly or solely produced in Australia; or
that Edentata and other American types should have been solely produced in
South America. For we know that Europe in ancient times was peopled by
numerous marsupials; and I have shown in the publications above alluded to
that in America the law of distribution of terrestrial mammals was formerly
different from what it now is. North America formerly partook strongly of
the present character of the southern half of the continent; and the
southern half was formerly more closely alliedthan it is at presentto
the northern half. In a similar manner we knowfrom Falconer and

Cautley's discoveriesthat Northern India was formerly more closely
related in its mammals to Africa than it is at the present time. Analogous
facts could be given in relation to the distribution of marine animals.

On the theory of descent with modificationthe great law of the long
enduringbut not immutablesuccession of the same types within the same
areasis at once explained; for the inhabitants of each quarter of the
world will obviously tend to leave in that quarterduring the next
succeeding period of timeclosely allied though in some degree modified
descendants. If the inhabitants of one continent formerly differed greatly
from those of another continentso will their modified descendants still
differ in nearly the same manner and degree. But after very long intervals
of timeand after great geographical changespermitting much
intermigrationthe feebler will yield to the more dominant formsand
there will be nothing immutable in the distribution of organic beings.

It may be asked in ridicule whether I suppose that the megatherium and
other allied huge monsterswhich formerly lived in South Americahave
left behind them the slotharmadilloand anteateras their degenerate
descendants. This cannot for an instant be admitted. These huge animals
have become wholly extinctand have left no progeny. 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; and
some of these fossils may have been the actual progenitors of the living
species. It must not be forgotten thaton our theoryall the species of
the same genus are the descendants of some one species; so thatif six
generaeach having eight speciesbe found in one geological formation
and in a succeeding formation there be six other allied or representative
generaeach with the same number of speciesthen we may conclude that
generally only one species of each of the older genera has left modified
descendantswhich constitute the new genera containing the several
species; the other seven species of each old genus having died out and left
no progeny. Orand this will be a far commoner casetwo 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 failing orderswith the genera and species decreasing
in numbers as is the case with the Edentata of South Americastill fewer
genera and species will leave modified blood-descendants.


I have attempted to show that the geological record is extremely imperfect;
that only a small portion of the globe has been geologically explored with
care; that only certain classes of organic beings have been largely
preserved in a fossil state; that the number both of specimens and of
speciespreserved in our museumsis absolutely as nothing compared with
the number of generations which must have passed away even during a single
formation; thatowing to subsidence being almost necessary for the
accumulation of deposits rich in fossil species of many kindsand thick
enough to outlast future degradationgreat intervals of time must have
elapsed between most of our successive formations; that there has probably
been more extinction during the periods of subsidenceand more variation
during the periods of elevationand during the latter the record will have
been least perfectly kept; that each single formation has not been
continuously deposited; that the duration of each formation is probably
short compared with the average duration of specific forms; that migration
has played an important part in the first appearance of new forms in any
one area and formation; that widely ranging species are those which have
varied most frequentlyand have oftenest given rise to new species; that
varieties have at first been local; and lastlyalthough each species must
have passed through numerous transitional stagesit is probable that the
periodsduring which each underwent modificationthough many and long as
measured by yearshave been short in comparison with the periods during
which each remained in an unchanged condition. These causestaken

conjointlywill to a large extent explain why--though we do find many
links--we do not find interminable varietiesconnecting 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
which might be foundwould be rankedunless the whole chain could be
perfectly restoredas a new and distinct species; for it is not pretended
that we have any sure criterion by which species and varieties can be

He who rejects this view of the imperfection of the geological recordwill
rightly reject the whole theory. For he may ask in vain where are the
numberless transitional links which must formerly have connected the
closely allied or representative speciesfound 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; he may
overlook how important a part migration has playedwhen the formations of
any one great regionas those of Europeare considered; he may urge the
apparentbut often falsely apparentsudden coming in of whole groups of
species. 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; but I can
answer this last question only by supposing that where our oceans now
extend they have extended for an enormous periodand where our oscillating
continents now stand they have stood since the commencement of the Cambrian
system; but thatlong before that epochthe world presented a widely
different aspect; and that the older continentsformed of formations older
than any known to usexist now only as remnants in a metamorphosed
conditionor lie still buried under the ocean.

Passing from these difficultiesthe other great leading facts in
palaeontology agree admirably with the theory of descent with modification
through variation and natural selection. We can thus understand how it is
that new species come in slowly and successively; how species of different
classes do not necessarily change togetheror at the same rateor in the
same degree; yet in the long run that all undergo modification to some
extent. The extinction of old forms is the almost inevitable consequence
of the production of new forms. We can understand whywhen a species has
once disappearedit never reappears. Groups of species increase in
numbers slowlyand endure for unequal periods of time; for the process of
modification is necessarily slowand depends on many complex
contingencies. The dominant species belonging to large and dominant groups
tend to leave many modified descendantswhich form new sub-groups and
groups. As these are formedthe species of the less vigorous groupsfrom
their inferiority inherited from a common progenitortend to become
extinct togetherand to leave no modified offspring on the face of the
earth. But the utter extinction of a whole group of species has sometimes
been a slow processfrom the survival of a few descendantslingering in
protected and isolated situations. When a group has once wholly
disappearedit does not reappear; for the link of generation has been

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
alliedbut modifieddescendants; and these will generally succeed in
displacing the groups which are their inferiors in the struggle for
existence. Henceafter long intervals of timethe productions of the
world appear to have changed simultaneously.

We can understand how it is that all the forms of lifeancient and recent
make together a few grand classes. We can understandfrom the continued
tendency to divergence of characterwhy the more ancient a form isthe
more it generally differs from those now living. Why ancient and extinct
forms often tend to fill up gaps between existing formssometimes blending
two groupspreviously classed as distinct into one; but more commonly

bringing them only a little closer together. The more ancient a form is
the more often it stands in some degree intermediate between groups now
distinct; for the more ancient a form isthe more nearly it will be
related toand consequently resemblethe common progenitor of groups
since become widely divergent. Extinct forms are seldom directly
intermediate between existing forms; but are intermediate only by a long
and circuitous course through other extinct and different forms. We can
clearly see why the organic remains of closely consecutive formations are
closely allied; for they are closely linked together by generation. We can
clearly see why the remains of an intermediate formation are intermediate
in character.

The inhabitants of the world at each successive period in its history have
beaten their predecessors in the race for lifeand arein so farhigher
in the scaleand their structure has generally become more specialised;
and this may account for the common belief held by so many
palaeontologiststhat organisation on the whole has progressed. Extinct
and ancient animals resemble to a certain extent the embryos of the more
recent animals belonging to the same classesand this wonderful fact
receives a simple explanation according to our views. The succession of
the same types of structure within the same areas during the later
geological periods ceases to be mysteriousand is intelligible on the
principle of inheritance.

Ifthenthe geological record be as imperfect as many believeand it may
at least be asserted that the record cannot be proved to be much more
perfectthe main objections to the theory of natural selection are greatly
diminished or disappear. On the other handall the chief laws of
palaeontology plainly proclaimas it seems to methat species have been
produced by ordinary generation: old forms having been supplanted by new
and improved forms of lifethe products of variation and the survival of
the fittest.



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 landand by occasional means -- Dispersal
during the Glacial period -- Alternate Glacial periods in the North and

In considering the distribution of organic beings over the face of the
globethe first great fact which strikes us isthat neither the
similarity nor the dissimilarity of the inhabitants of various regions can
be wholly accounted for by climatal and other physical conditions. Of
latealmost every author who has studied the subject has come to this
conclusion. The case of America alone would almost suffice to prove its
truth; for if we exclude the arctic and northern temperate partsall
authors agree that one of the most fundamental divisions in geographical
distribution is that between the New and Old Worlds; yet if we travel over
the vast American continentfrom the central parts of the United States to
its extreme southern pointwe meet with the most diversified conditions;
humid districtsarid desertslofty mountainsgrassy plainsforests
marsheslakes and great riversunder almost every temperature. 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. No
doubt small areas can be pointed out in the Old World hotter than any in
the New World; but these are not inhabited by a fauna different from that
of the surrounding districts; for it is rare to find a group of organisms
confined to a small areaof which the conditions are peculiar in only a

slight degree. Notwithstanding this general parallelism in the conditions
of Old and New Worldshow widely different are their living productions!

In the southern hemisphereif we compare large tracts of land in
AustraliaSouth Africaand western South Americabetween latitudes 25
and 35 degreeswe shall find parts extremely similar in all their
conditionsyet it would not be possible to point out three faunas and
floras more utterly dissimilar. Oragainwe may compare the productions
of South America south of latitude 35 degrees with those north of 25
degreeswhich consequently are separated by a space of ten degrees of
latitudeand are exposed to considerably different conditions; 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.
Analogous facts could be given with respect to the inhabitants of the sea.

A second great fact which strikes us in our general review isthat
barriers of any kindor obstacles to free migrationare related in a
close and important manner to the differences between the productions of
various regions. We see this in the great difference in nearly all the
terrestrial productions of the New and Old Worldsexcepting in the
northern partswhere the land almost joinsand whereunder a slightly
different climatethere might have been free migration for the northern
temperate formsas there now is for the strictly arctic productions. We
see the same fact in the great difference between the inhabitants of
AustraliaAfricaand South America under the same latitude; for these
countries are almost as much isolated from each other as is possible. On
each continentalsowe see the same fact; for on the opposite sides of
lofty and continuous mountain-rangesand of great deserts and even of
large riverswe find different productions; though as mountain chains
desertsetc.are not as impassableor likely to have endured so longas
the oceans separating continentsthe differences are very inferior in
degree to those characteristic of distinct continents.

Turning to the seawe find the same law. The marine inhabitants of the
eastern and western shores of South America are very distinctwith
extremely few shellscrustaceaor echinodermata in common; but Dr.
Gunther has recently shown that about thirty per cent of the fishes are the
same on the opposite sides of the isthmus of Panama; and this fact has led
naturalists to believe that the isthmus was formerly open. Westward of the
shores of Americaa wide space of open ocean extendswith not an island
as a halting-place for emigrants; here we have a barrier of another kind
and as soon as this is passed we meet in the eastern islands of the Pacific
with another and totally distinct fauna. So that three marine faunas range
northward and southward in parallel lines not far from each otherunder
corresponding climate; but from being separated from each other by
impassable barrierseither of land or open seathey are almost wholly
distinct. On the other handproceeding still further westward from the
eastern islands of the tropical parts of the Pacificwe encounter no
impassable barriersand we have innumerable islands as halting-placesor
continuous coastsuntilafter travelling over a hemispherewe come to
the shores of Africa; and over this vast space we meet with no well-defined
and distinct marine faunas. Although so few marine animals are common to
the above-named three approximate faunas of Eastern and Western America and
the eastern Pacific islandsyet many fishes range from the Pacific into
the Indian Oceanand many shells are common to the eastern islands of the
Pacific and the eastern shores of Africa on almost exactly opposite
meridians of longitude.

A third great factpartly included in the foregoing statementis the
affinity of the productions of the same continent or of the same sea
though the species themselves are distinct at different points and
stations. It is a law of the widest generalityand every continent offers
innumerable instances. Neverthelessthe naturalistin travellingfor
instancefrom north to southnever fails to be struck by the manner in

which successive groups of beingsspecifically distinctthough nearly
relatedreplace each other. He hears from closely alliedyet distinct
kinds of birdsnotes nearly similarand sees their nests similarly
constructedbut not quite alikewith eggs coloured in nearly the same
manner. The plains near the Straits of Magellan are inhabited by one
species of Rhea (American ostrich)and northward the plains of La Plata by
another species of the same genus; and not by a true ostrich or emulike
those inhabiting Africa and Australia under the same latitude. On these
same plains of La Plata we see the agouti and bizcachaanimals having
nearly the same habits as our hares and rabbitsand belonging to the same
order of Rodentsbut they plainly display an American type of structure.
We ascend the lofty peaks of the Cordilleraand we find an alpine species
of bizcacha; we look to the watersand we do not find the beaver or
muskratbut the coypu and capybararodents of the South American type.
Innumerable other instances could be given. If we look to the islands off
the American shorehowever much they may differ in geological structure
the inhabitants are essentially Americanthough they may be all peculiar
species. We may look back to past agesas shown in the last chapterand
we find American types then prevailing on the American continent and in the
American seas. We see in these facts some deep organic bondthroughout
space and timeover the same areas of land and waterindependently of
physical conditions. The naturalist must be dull who is not led to inquire
what this bond is.

The bond is simply inheritancethat cause which aloneas far as we
positively knowproduces organisms quite like each otheroras we see in
the case of varietiesnearly alike. The dissimilarity of the inhabitants
of different regions may be attributed to modification through variation
and natural selectionand probably in a subordinate degree to the definite
influence of different physical conditions. 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 preventedat
periods more or less remote--on the nature and number of the former
immigrants--and on the action of the inhabitants on each other in leading
to the preservation of different modifications; the relation of organism to
organism in the struggle for life beingas I have already often remarked
the most important of all relations. Thus the high importance of barriers
comes into play by checking migration; as does time for the slow process of
modification through natural selection. Widely-ranging speciesabounding
in individualswhich have already triumphed over many competitors in their
own widely-extended homeswill have the best chance of seizing on new
placeswhen they spread out into new countries. In their new homes they
will be exposed to new conditionsand will frequently undergo further
modification and improvement; and thus they will become still further
victoriousand will produce groups of modified descendants. On this
principle of inheritance with modification we can understand how it is that
sections of generawhole generaand even familiesare confined to the
same areasas is so commonly and notoriously the case.

There is no evidenceas was remarked in the last chapterof the existence
of any law of necessary development. As the variability of each species is
an independent propertyand will be taken advantage of by natural
selectiononly so far as it profits each individual in its complex
struggle for lifeso the amount of modification in different species will
be no uniform quantity. If a number of speciesafter having long competed
with each other in their old homewere to migrate in a body into a new and
afterwards isolated countrythey would be little liable to modification;
for neither migration nor isolation in themselves effect anything. 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. As we have seen in the last chapter that some forms have
retained nearly the same character from an enormously remote geological
periodso certain species have migrated over vast spacesand have not
become greatly or at all modified.

According to these viewsit is obvious that the several species of the
same genusthough inhabiting the most distant quarters of the worldmust
originally have proceeded from the same sourceas they are descended from
the same progenitor. In the case of those species which have undergone
during whole geological periodslittle modificationthere is not much
difficulty in believing that they have migrated from the same region; for
during the vast geographical and climatical changes which have supervened
since ancient timesalmost any amount of migration is possible. But in
many other casesin which we have reason to believe that the species of a
genus have been produced within comparatively recent timesthere is great
difficulty on this head. It is also obvious that the individuals of the
same speciesthough now inhabiting distant and isolated regionsmust have
proceeded from one spotwhere their parents were first produced: foras
has been explainedit is incredible that individuals identically the same
should have been produced from parents specifically distinct.


We are thus brought to the question which has been largely discussed by
naturalistsnamelywhether species have been created at one or more
points of the earth's surface. 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
where now found. Nevertheless the simplicity of the view that each species
was first produced within a single region captivates the mind. He who
rejects itrejects the vera causa of ordinary generation with subsequent
migrationand calls in the agency of a miracle. It is universally
admittedthat in most cases the area inhabited by a species is continuous;
and that when a plant or animal inhabits two points so distant from each
otheror with an interval of such a naturethat the space could not have
been easily passed over by migrationthe 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; andaccordinglywe find no inexplicable instances
of the same mammals inhabiting distant points of the world. No geologist
feels any difficulty in Great Britain possessing the same quadrupeds with
the rest of Europefor they were no doubt once united. But if the same
species can be produced at two separate pointswhy do we not find a single
mammal common to Europe and Australia or South America? The conditions of
life are nearly the sameso that a multitude of European animals and
plants have become naturalised in America and Australia; and some of the
aboriginal plants are identically the same at these distant points of the
northern and southern hemispheres? The answeras I believeisthat
mammals have not been able to migratewhereas some plantsfrom their
varied means of dispersalhave migrated across the wide and broken
interspaces. The great and striking influence of barriers of all kindsis
intelligible only on the view that the great majority of species have been
produced on one sideand have not been able to migrate to the opposite
side. Some few familiesmany subfamiliesvery many generaa still
greater number of sections of generaare confined to a single region; and
it has been observed by several naturalists that the most natural genera
or those genera in which the species are most closely related to each
otherare generally confined to the same countryor if they have a wide
range that their range is continuous. What a strange anomaly it would be
if a directly opposite rule were to prevail when we go down one step lower
in the seriesnamely to the individuals of the same speciesand these had
not beenat least at firstconfined to some one region!

Henceit seems to meas it has to many other naturaliststhat the view
of each species having been produced in one area aloneand having
subsequently migrated from that area as far as its powers of migration and
subsistence under past and present conditions permittedis the most
probable. Undoubtedly many cases occur in which we cannot explain how the

same species could have passed from one point to the other. But the
geographical and climatical changes which have certainly occurred within
recent geological timesmust have rendered discontinuous the formerly
continuous range of many species. So that we are reduced to consider
whether the exceptions to continuity of range are so numerousand of so
grave a naturethat we ought to give up the beliefrendered probable by
general considerationsthat each species has been produced within one
areaand has migrated thence as far as it could. It would be hopelessly
tedious to discuss all the exceptional cases of the same speciesnow
living at distant and separated points; nor do I for a moment pretend that
any explanation could be offered of many instances. Butafter some
preliminary remarksI will discuss a few of the most striking classes of
factsnamelythe existence of the same species on the summits of distant
mountain rangesand at distant points in the Arctic and Antarctic regions;
and secondly (in the following chapter)the wide distribution of fresh
water productions; and thirdlythe occurrence of the same terrestrial
species on islands and on the nearest mainlandthough separated by
hundreds of miles of open sea. 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; thenconsidering our ignorance with respect to former
climatical and geographical changesand to the various occasional means of
transportthe belief that a single birthplace is the law seems to me
incomparably the safest.

In discussing this subject we shall be enabled at the same time to consider
a point equally important for usnamelywhether the several species of a
genus which must on our theory all be descended from a common progenitor
can have migratedundergoing modification during their migration from some
one area. Ifwhen most of the species inhabiting one region are different
from those of another regionthough closely allied to themit can be
shown that migration from the one region to the other has probably occurred
at some former periodour general view will be much strengthened; for the
explanation is obvious on the principle of descent with modification. A
volcanic islandfor instanceupheaved and formed at the distance of a few
hundreds of miles from a continentwould probably receive from it in the
course of time a few colonistsand their descendantsthough modified
would still be related by inheritance to the inhabitants of that continent.
Cases of this nature are commonand areas we shall hereafter see
inexplicable on the theory of independent creation. This view of the
relation of the species of one region to those of anotherdoes not differ
much from that advanced by Mr. Wallacewho concludes that "every species
has come into existence coincident both in space and time with a
pre-existing closely allied species." And it is now well known that he
attributes this coincidence to descent with modification.

The question of single or multiple centres of creation differs from another
though allied questionnamelywhether all the individuals of the same
species are descended from a single pairor single hermaphroditeor
whetheras some authors supposefrom many individuals simultaneously
created. With organic beings which never intercrossif such existeach
speciesmust be descended from a succession of modified varietiesthat
have supplanted each otherbut have never blended with other individuals
or varieties of the same speciesso thatat each successive stage of
modificationall the individuals of the same form will be descended from a
single parent. But in the great majority of casesnamelywith all
organisms which habitually unite for each birthor which occasionally
intercrossthe individuals of the same species inhabiting the same area
will be kept nearly uniform by intercrossing; so that many individuals will
go on simultaneously changingand the whole amount of modification at each
stage will not be due to descent from a single parent. To illustrate what
I mean: our English race-horses differ from the horses of every other
breed; but they do not owe their difference and superiority to descent from
any single pairbut to continued care in the selecting and training of

many individuals during each generation.

Before discussing the three classes of factswhich I have selected as
presenting the greatest amount of difficulty on the theory of "single
centres of creation I must say a few words on the means of dispersal.


Sir C. Lyell and other authors have ably treated this subject. I can give
here only the briefest abstract of the more important facts. Change of
climate must have had a powerful influence on migration. A region now
impassable to certain organisms from the nature of its climate, might have
been a high road for migration, when the climate was different. I shall,
however, presently have to discuss this branch of the subject in some
detail. Changes of level in the land must also have been highly
influential: a narrow isthmus now separates two marine faunas; submerge
it, or let it formerly have been submerged, and the two faunas will now
blend together, or may formerly have blended. Where the sea now extends,
land may at a former period have connected islands or possibly even
continents together, and thus have allowed terrestrial productions to pass
from one to the other. No geologist disputes that great mutations of level
have occurred within the period of existing organisms. Edward Forbes
insisted that all the islands in the Atlantic must have been recently
connected with Europe or Africa, and Europe likewise with America. Other
authors have thus hypothetically bridged over every ocean, and united
almost every island with some mainland. If, indeed, the arguments used by
Forbes are to be trusted, it must be admitted that scarcely a single island
exists which has not recently been united to some continent. This view
cuts the Gordian knot of the dispersal of the same species to the most
distant points, and removes many a difficulty; but to the best of my
judgment we are not authorized in admitting such enormous geographical
changes within the period of existing species. It seems to me that we have
abundant evidence of great oscillations in the level of the land or sea;
but not of such vast changes in the position and extension of our
continents, as to have united them within the recent period to each other
and to the several intervening oceanic islands. I freely admit the former
existence of many islands, now buried beneath the sea, which may have
served as halting places for plants and for many animals during their
migration. In the coral-producing oceans such sunken islands are now
marked by rings of coral or atolls standing over them. Whenever it is
fully admitted, as it will some day be, that each species has proceeded
from a single birthplace, and when in the course of time we know something
definite about the means of distribution, we shall be enabled to speculate
with security on the former extension of the land. 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, have been continuously, or
almost continuously united with each other, and with the many existing
oceanic islands. 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, 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, as
are necessary on the view advanced by Forbes and admitted by his followers.
The nature and relative proportions of the inhabitants of oceanic islands
are likewise opposed to the belief of their former continuity of
continents. Nor does the almost universally volcanic composition of such
islands favour the admission that they are the wrecks of sunken continents;
if they had originally existed as continental mountain ranges, some at
least of the islands would have been formed, like other mountain summits,
of granite, metamorphic schists, old fossiliferous and other rocks, instead
of consisting of mere piles of volcanic matter.

I must now say a few words on what are called accidental means, but which
more properly should be called occasional means of distribution. I shall
here confine myself to plants. In botanical works, this or that plant is
often stated to be ill adapted for wide dissemination; but the greater or
less facilities for transport across the sea may be said to be almost
wholly unknown. Until I tried, with Mr. Berkeley's aid, a few experiments,
it was not even known how far seeds could resist the injurious action of
sea-water. To my surprise I found that out of eighty-seven kinds, sixtyfour
germinated after an immersion of twenty-eight days, and a few survived
an immersion of 137 days. It deserves notice that certain orders were far
more injured than others: nine Leguminosae were tried, and, with one
exception, they resisted the salt-water badly; seven species of the allied
orders, Hydrophyllaceae and Polemoniaceae, were all killed by a month's
immersion. For convenience sake I chiefly tried small seeds without the
capsules or fruit; and as all of these sank in a few days, they could not
have been floated across wide spaces of the sea, whether or not they were
injured by salt water. Afterwards I tried some larger fruits, capsules,
etc., and some of these floated for a long time. It is well known what a
difference there is in the buoyancy of green and seasoned timber; 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. Hence I was led to
dry the stems and branches of ninety-four plants with ripe fruit, and to
place them on sea-water. The majority sank quickly, but some which, whilst
green, floated for a very short time, when dried floated much longer; for
instance, ripe hazel-nuts sank immediately, but when dried they floated for
ninety days, and afterwards when planted germinated; an asparagus plant
with ripe berries floated for twenty-three days, when dried it floated for
eighty-five days, and the seeds afterwards germinated: the ripe seeds of
Helosciadium sank in two days, when dried they floated for above ninety
days, and afterwards germinated. Altogether, out of the ninety-four dried
plants, eighteen floated for above twenty-eight days; and some of the
eighteen floated for a very much longer period. So that as 64/87 kinds of
seeds germinated after an immersion of twenty-eight days; and as 18/94
distinct species with ripe fruit (but not all the same species as in the
foregoing experiment) floated, after being dried, for above twenty-eight
days, we may conclude, as far as anything can be inferred from these scanty
facts, that the seeds of 14/100 kinds of plants of any country might be
floated by sea-currents during twenty-eight days, and would retain their
power of germination. In Johnston's Physical Atlas, 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, the seeds of
14/100 plants belonging to one country might be floated across 924 miles of
sea to another country; and when stranded, if blown by an inland gale to a
favourable spot, would germinate.

Subsequently to my experiments, M. Martens tried similar ones, but in a
much better manner, for he placed the seeds in a box in the actual sea, so
that they were alternately wet and exposed to the air like really floating
plants. He tried ninety-eight seeds, mostly different from mine, but he
chose many large fruits, and likewise seeds, from plants which live near
the sea; and this would have favoured both the average length of their
flotation and their resistance to the injurious action of the salt-water.
On the other hand, he did not previously dry the plants or branches with
the fruit; and this, as we have seen, would have caused some of them to
have floated much longer. The result was that 18/98 of his seeds of
different kinds floated for forty-two days, and were then capable of
germination. 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. Therefore, it would perhaps be safer to assume that the seeds
of about 10/100 plants of a flora, after having been dried, could be
floated across a space of sea 900 miles in width, and would then germinate.
The fact of the larger fruits often floating longer than the small, is
interesting; as plants with large seeds or fruit which, as Alph. de

Candolle has shown, generally have restricted ranges, could hardly be
transported by any other means.

Seeds may be occasionally transported in another manner. Drift timber is
thrown up on most islands, even on those in the midst of the widest oceans;
and the natives of the coral islands in the Pacific procure stones for
their tools, solely from the roots of drifted trees, these stones being a
valuable royal tax. I find that when irregularly shaped stones are
embedded in the roots of trees, small parcels of earth are very frequently
enclosed in their interstices and behind 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, three dicotyledonous plants germinated: I am
certain of the accuracy of this observation. Again, I can show that the
carcasses of birds, when floating on the sea, sometimes escape being
immediately devoured; and many kinds of seeds in the crops of floating
birds long retain their vitality: peas and vetches, for instance, are
killed by even a few days' immersion in sea-water; but some taken out of
the crop of a pigeon, which had floated on artificial sea-water for thirty
days, to my surprise nearly all germinated.

Living birds can hardly fail to be highly effective agents in the
transportation of seeds. I could give many facts showing how frequently
birds of many kinds are blown by gales to vast distances across the ocean.
We may safely assume that under such circumstances their rate of flight
would often be thirty-five miles an hour; and some authors have given a far
higher estimate. I have never seen an instance of nutritious seeds passing
through the intestines of a bird; but hard seeds of fruit pass uninjured
through even the digestive organs of a turkey. In the course of two
months, I picked up in my garden twelve kinds of seeds, out of the
excrement of small birds, and these seemed perfect, and some of them, which
were tried, germinated. But the following fact is more important: the
crops of birds do not secrete gastric juice, and do not, as I know by
trial, injure in the least the germination of seeds; now, after a bird has
found and devoured a large supply of food, it is positively asserted that
all the grains do not pass into the gizzard for twelve or even eighteen
hours. A bird in this interval might easily be blown to the distance of
five hundred miles, and hawks are known to look out for tired birds, and
the contents of their torn crops might thus readily get scattered. Some
hawks and owls bolt their prey whole, and after an interval of from twelve
to twenty hours, disgorge pellets, which, as I know from experiments made
in the Zoological Gardens, include seeds capable of germination. Some
seeds of the oat, wheat, millet, canary, hemp, clover, and beet germinated
after having been from twelve to twenty-one hours in the stomachs of
different birds of prey; and two seeds of beet grew after having been thus
retained for two days and fourteen hours. Fresh-water fish, I find, eat
seeds of many land and water plants; fish are frequently devoured by birds,
and thus the seeds might be transported from place to place. I forced many
kinds of seeds into the stomachs of dead fish, and then gave their bodies
to fishing-eagles, storks, and pelicans; these birds, after an interval of
many hours, either rejected the seeds in pellets or passed them in their
excrement; and several of these seeds retained the power of germination.
Certain seeds, however, were always killed by this process.

Locusts are sometimes blown to great distances from the land. I myself
caught one 370 miles from the coast of Africa, and have heard of others
caught at greater distances. The Rev. R.T. Lowe informed Sir C. Lyell that
in November, 1844, swarms of locusts visited the island of Madeira. They
were in countless numbers, as thick as the flakes of snow in the heaviest
snowstorm, and extended upward as far as could be seen with a telescope.
During two or three days they slowly careered round and round in an immense
ellipse, at least five or six miles in diameter, and at night alighted on
the taller trees, which were completely coated with them. They then
disappeared over the sea, as suddenly as they had appeared, and have not

since visited the island. Now, in parts of Natal it is believed by some
farmers, though on insufficient evidence, that injurious seeds are
introduced into their grass-land in the dung left by the great flights of
locusts which often visit that country. In consequence of this belief Mr.
Weale sent me in a letter a small packet of the dried pellets, out of which
I extracted under the microscope several seeds, and raised from them seven
grass plants, belonging to two species, of two genera. Hence a swarm of
locusts, such as that which visited Madeira, might readily be the means of
introducing several kinds of plants into an island lying far from the

Although the beaks and feet of birds are generally clean, earth sometimes
adheres to them: in one case I removed sixty-one grains, and in another
case twenty-two grains of dry argillaceous earth from the foot of a
partridge, and in the earth there was a pebble as large as the seed of a
vetch. Here is a better case: the leg of a woodcock was sent to me by a
friend, with a little cake of dry earth attached to the shank, weighing
only nine grains; and this contained a seed of the toad-rush (Juncus
bufonius) which germinated and flowered. Mr. Swaysland, of Brighton, who
during the last forty years has paid close attention to our migratory
birds, informs me that he has often shot wagtails (Motacillae), wheatears,
and whinchats (Saxicolae), on their first arrival on our shores, before
they had alighted; and he has several times noticed little cakes of earth
attached to their feet. Many facts could be given showing how generally
soil is charged with seeds. For instance, Professor Newton sent me the leg
of a red-legged partridge (Caccabis rufa) which had been wounded and could
not fly, with a ball of hard earth adhering to it, and weighing six and a
half ounces. The earth had been kept for three years, but when broken,
watered and placed under a bell glass, no less than eighty-two plants
sprung from it: these consisted of twelve monocotyledons, including the
common oat, and at least one kind of grass, and of seventy dicotyledons,
which consisted, judging from the young leaves, of at least three distinct
species. With such facts before us, can we doubt that the many birds which
are annually blown by gales across great spaces of ocean, and which
annually migrate--for instance, the millions of quails across the
Mediterranean--must occasionally transport a few seeds embedded in dirt
adhering to their feet or beaks? But I shall have to recur to this

As icebergs are known to be sometimes loaded with earth and stones, and
have even carried brushwood, bones, and the nest of a land-bird, it can
hardly be doubted that they must occasionally, as suggested by Lyell, have
transported seeds from one part to another of the arctic and antarctic
regions; and during the Glacial period from one part of the now temperate
regions to another. In the Azores, from the large number of plants common
to Europe, in comparison with the species on the other islands of the
Atlantic, which stand nearer to the mainland, and (as remarked by Mr. H.C.
Watson) from their somewhat northern character, in comparison with the
latitude, I suspected that these islands had been partly stocked by
ice-borne seeds during the Glacial epoch. At my request Sir C. Lyell wrote
to M. Hartung to inquire whether he had observed erratic boulders on these
islands, and he answered that he had found large fragments of granite and
other rocks, which do not occur in the archipelago. Hence we may safely
infer that icebergs formerly landed their rocky burdens on the shores of
these mid-ocean islands, and it is at least possible that they may have
brought thither the seeds of northern plants.

Considering that these several means of transport, and that other means,
which without doubt remain to be discovered, have been in action year after
year for tens of thousands of years, it would, I think, be a marvellous
fact if many plants had not thus become widely transported. These means of
transport are sometimes called accidental, but this is not strictly
correct: the currents of the sea are not accidental, nor is the direction
of prevalent gales of wind. It should be observed that scarcely any means

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


The identity of many plants and animals, on mountain-summits, separated
from each other by hundreds of miles of lowlands, where Alpine species
could not possibly exist, is one of the most striking cases known of the
same species living at distant points, without the apparent possibility of
their having migrated from one point to the other. It is indeed a
remarkable fact to see so many plants of the same species living on the
snowy regions of the Alps or Pyrenees, and in the extreme northern parts of
Europe; but it is far more remarkable, that the plants on the White
Mountains, in the United States of America, are all the same with those of
Labrador, and nearly all the same, as we hear from Asa Gray, with those on
the loftiest mountains of Europe. Even as long ago as 1747, such facts led
Gmelin to conclude that the same species must have been independently
created at many distinct points; and we might have remained in this same
belief, had not Agassiz and others called vivid attention to the Glacial
period, which, as we shall immediately see, affords a simple explanation of
these facts. We have evidence of almost every conceivable kind, organic
and inorganic, that, within a very recent geological period, central Europe
and North America suffered under an Arctic climate. The ruins of a house
burnt by fire do not tell their tale more plainly than do the mountains of
Scotland and Wales, with their scored flanks, polished surfaces, and
perched boulders, of the icy streams with which their valleys were lately
filled. So greatly has the climate of Europe changed, that in Northern
Italy, gigantic moraines, left by old glaciers, are now clothed by the vine
and maize. Throughout a large part of the United States, erratic boulders
and scored rocks plainly reveal a former cold period.

The former influence of the glacial climate on the distribution of the
inhabitants of Europe, as explained by Edward Forbes, is substantially as
follows. But we shall follow the changes more readily, by supposing a new

glacial period slowly to come on, and then pass away, as formerly occurred.
As the cold came on, and as each more southern zone became fitted for the
inhabitants of the north, these would take the places of the former
inhabitants of the temperate regions. The latter, at the same time would
travel further and further southward, unless they were stopped by barriers,
in which case they would perish. The mountains would become covered with
snow and ice, and their former Alpine inhabitants would descend to the
plains. By the time that the cold had reached its maximum, we should have
an arctic fauna and flora, covering the central parts of Europe, as far
south as the Alps and Pyrenees, and even stretching into Spain. The now
temperate regions of the United States would likewise be covered by arctic
plants and animals and these would be nearly the same with those of Europe;
for the present circumpolar inhabitants, which we suppose to have
everywhere travelled southward, are remarkably uniform round the world.

As the warmth returned, the arctic forms would retreat northward, closely
followed up in their retreat by the productions of the more temperate
regions. And as the snow melted from the bases of the mountains, the
arctic forms would seize on the cleared and thawed ground, always
ascending, as the warmth increased and the snow still further disappeared,
higher and higher, whilst their brethren were pursuing their northern
journey. Hence, when the warmth had fully returned, the same species,
which had lately lived together on the European and North American
lowlands, would again be found in the arctic regions of the Old and New
Worlds, and on many isolated mountain-summits far distant from each other.

Thus we can understand the identity of many plants at points so immensely
remote as the mountains of the United States and those of Europe. We can
thus also understand the fact that the Alpine plants of each mountain-range
are more especially related to the arctic forms living due north or nearly
due north of them: for the first migration when the cold came on, and the
re-migration on the returning warmth, would generally have been due south
and north. The Alpine plants, for example, of Scotland, as remarked by Mr.

H.C. Watson, and those of the Pyrenees, as remarked by Ramond, are more
especially allied to the plants of northern Scandinavia; those of the
United States to Labrador; those of the mountains of Siberia to the arctic
regions of that country. These views, grounded as they are on the
perfectly well-ascertained occurrence of a former Glacial period, seem to
me to explain in so satisfactory a manner the present distribution of the
Alpine and Arctic productions of Europe and America, that when in other
regions we find the same species on distant mountain-summits, we may almost
conclude, without other evidence, that a colder climate formerly permitted
their migration across the intervening lowlands, now become too warm for
their existence.
As the arctic forms moved first southward and afterwards backward to the
north, in unison with the changing climate, they will not have been exposed
during their long migrations to any great diversity of temperature; and as
they all migrated in a body together, their mutual relations will not have
been much disturbed. Hence, in accordance with the principles inculcated
in this volume, these forms will not have been liable to much modification.
But with the Alpine productions, left isolated from the moment of the
returning warmth, first at the bases and ultimately on the summits of the
mountains, the case will have been somewhat different; for it is not likely
that all the same arctic species will have been left on mountain ranges far
distant from each other, and have survived there ever since; they will
also, in all probability, have become mingled with ancient Alpine species,
which must have existed on the mountains before the commencement of the
Glacial epoch, and which during the coldest period will have been
temporarily driven down to the plains; they will, also, have been
subsequently exposed to somewhat different climatical influences. Their
mutual relations will thus have been in some degree disturbed; consequently
they will have been liable to modification; and they have been modified;
for if we compare the present Alpine plants and animals of the several

great European mountain ranges, one with another, though many of the
species remain identically the same, some exist as varieties, some as
doubtful forms or sub-species and some as distinct yet closely allied
species representing each other on the several ranges.

In the foregoing illustration, I have assumed that at the commencement of
our imaginary Glacial period, the arctic productions were as uniform round
the polar regions as they are at the present day. But it is also necessary
to assume that many sub-arctic and some few temperate forms were the same
round the world, for some of the species which now exist on the lower
mountain slopes and on the plains of North America and Europe are the same;
and it may be asked how I account for this degree of uniformity of the
sub-arctic and temperate forms round the world, at the commencement of the
real Glacial period. At the present day, the sub-arctic and northern
temperate productions of the Old and New Worlds are separated from each
other by the whole Atlantic Ocean and by the northern part of the Pacific.
During the Glacial period, when the inhabitants of the Old and New Worlds
lived further southwards than they do at present, they must have been still
more completely separated from each other by wider spaces of ocean; so that
it may well be asked how the same species could then or previously have
entered the two continents. The explanation, I believe, lies in the nature
of the climate before the commencement of the Glacial period. At this, the
newer Pliocene period, the majority of the inhabitants of the world were
specifically the same as now, and we have good reason to believe that the
climate was warmer than at the present day. Hence, we may suppose that the
organisms which now live under latitude 60 degrees, lived during the
Pliocene period further north, under the Polar Circle, in latitude 66-67
degrees; and that the present arctic productions then lived on the broken
land still nearer to the pole. Now, if we look at a terrestrial globe, we
see under the Polar Circle that there is almost continuous land from
western Europe through Siberia, to eastern America. And this continuity of
the circumpolar land, with the consequent freedom under a more favourable
climate for intermigration, will account for the supposed uniformity of the
sub-arctic and temperate productions of the Old and New Worlds, at a period
anterior to the Glacial epoch.

Believing, from reasons before alluded to, that our continents have long
remained in nearly the same relative position, though subjected to great
oscillations of level, I am strongly inclined to extend the above view, and
to infer that during some earlier and still warmer period, such as the
older Pliocene period, a large number of the same plants and animals
inhabited the almost continuous circumpolar land; and that these plants and
animals, both in the Old and New Worlds, began slowly to migrate southwards
as the climate became less warm, long before the commencement of the
Glacial period. We now see, as I believe, their descendants, mostly in a
modified condition, in the central parts of Europe and the United States.
On this view we can understand the relationship with very little identity,
between the productions of North America and Europe--a relationship which
is highly remarkable, considering the distance of the two areas, and their
separation by the whole Atlantic Ocean. We can further understand the
singular fact remarked on by several observers that the productions of
Europe and America during the later tertiary stages were more closely
related to each other than they are at the present time; for during these
warmer periods the northern parts of the Old and New Worlds will have been
almost continuously united by land, serving as a bridge, since rendered
impassable by cold, for the intermigration of their inhabitants.

During the slowly decreasing warmth of the Pliocene period, as soon as the
species in common, which inhabited the New and Old Worlds, migrated south
of the Polar Circle, they will have been completely cut off from each
other. This separation, as far as the more temperate productions are
concerned, must have taken place long ages ago. As the plants and animals
migrated southward, they will have become mingled in the one great region
with the native American productions, and would have had to compete with

them; and in the other great region, with those of the Old World.
Consequently we have here everything favourable for much modification--for
far more modification than with the Alpine productions, left isolated,
within a much more recent period, on the several mountain ranges and on the
arctic lands of Europe and North America. Hence, it has come, that when we
compare the now living productions of the temperate regions of the New and
Old Worlds, we find very few identical species (though Asa Gray has lately
shown that more plants are identical than was formerly supposed), but we
find in every great class many forms, which some naturalists rank as
geographical races, and others as distinct species; and a host of closely
allied or representative forms which are ranked by all naturalists as
specifically distinct.

As on the land, so in the waters of the sea, a slow southern migration of a
marine fauna, which, during the Pliocene or even a somewhat earlier period,
was nearly uniform along the continuous shores of the Polar Circle, will
account, on the theory of modification, for many closely allied forms now
living in marine areas completely sundered. Thus, I think, we can
understand the presence of some closely allied, still existing and extinct
tertiary forms, on the eastern and western shores of temperate North
America; and the still more striking fact of many closely allied
crustaceans (as described in Dana's admirable work), some fish and other
marine animals, inhabiting the Mediterranean and the seas of Japan--these
two areas being now completely separated by the breadth of a whole
continent and by wide spaces of ocean.

These cases of close relationship in species either now or formerly
inhabiting the seas on the eastern and western shores of North America, the
Mediterranean and Japan, and the temperate lands of North America and
Europe, are inexplicable on the theory of creation. We cannot maintain
that such species have been created alike, in correspondence with the
nearly similar physical conditions of the areas; for if we compare, for
instance, certain parts of South America with parts of South Africa or
Australia, we see countries closely similar in all their physical
conditions, with their inhabitants utterly dissimilar.


But we must return to our more immediate subject. I am convinced that
Forbes's view may be largely extended. In Europe we meet with the plainest
evidence of the Glacial period, from the western shores of Britain to the
Ural range, and southward to the Pyrenees. We may infer from the frozen
mammals and nature of the mountain vegetation, that Siberia was similarly
affected. In the Lebanon, according to Dr. Hooker, perpetual snow formerly
covered the central axis, and fed glaciers which rolled 4,000 feet down the
valleys. The same observer has recently found great moraines at a low
level on the Atlas range in North Africa. Along the Himalaya, at points
900 miles apart, glaciers have left the marks of their former low descent;
and in Sikkim, Dr. Hooker saw maize growing on ancient and gigantic
moraines. Southward of the Asiatic continent, on the opposite side of the
equator, we know, from the excellent researches of Dr. J. Haast and Dr.
Hector, that in New Zealand immense glaciers formerly descended to a low
level; and the same plants, found by Dr. Hooker on widely separated
mountains in this island tell the same story of a former cold period. From
facts communicated to me by the Rev. W.B. Clarke, it appears also that
there are traces of former glacial action on the mountains of the southeastern
corner of Australia.

Looking to America: in the northern half, ice-borne fragments of rock have
been observed on the eastern side of the continent, as far south as
latitude 36 and 37 degrees, and on the shores of the Pacific, where the
climate is now so different, as far south as latitude 46 degrees. Erratic
boulders have, also, been noticed on the Rocky Mountains. In the
Cordillera of South America, nearly under the equator, glaciers once

extended far below their present level. In central Chile I examined a vast
mound of detritus with great boulders, crossing the Portillo valley, which,
there can hardly be a doubt, once formed a huge moraine; and Mr. D. Forbes
informs me that he found in various parts of the Cordillera, from latitude
13 to 30 degrees south, at about the height of 12,000 feet, deeply-furrowed
rocks, resembling those with which he was familiar in Norway, and likewise
great masses of detritus, including grooved pebbles. Along this whole
space of the Cordillera true glaciers do not now exist even at much more
considerable heights. Further south, on both sides of the continent, from
latitude 41 degrees to the southernmost extremity, we have the clearest
evidence of former glacial action, in numerous immense boulders transported
far from their parent source.

>From these several facts, namely, from the glacial action having extended
all round the northern and southern hemispheres--from the period having
been in a geological sense recent in both hemispheres--from its having
lasted in both during a great length of time, as may be inferred from the
amount of work effected--and lastly, from glaciers having recently
descended to a low level along the whole line of the Cordillera, it at one
time appeared to me that we could not avoid the conclusion that the
temperature of the whole world had been simultaneously lowered during the
Glacial period. But now, Mr. Croll, in a series of admirable memoirs, has
attempted to show that a glacial condition of climate is the result of
various physical causes, brought into operation by an increase in the
eccentricity of the earth's orbit. All these causes tend towards the same
end; but the most powerful appears to be the indirect influence of the
eccentricity of the orbit upon oceanic currents. According to Mr. Croll,
cold periods regularly recur every ten or fifteen thousand years; and these
at long intervals are extremely severe, owing to certain contingencies, of
which the most important, as Sir C. Lyell has shown, is the relative
position of the land and water. Mr. Croll believes that the last great
glacial period occurred about 240,000 years ago, and endured, with slight
alterations of climate, for about 160,000 years. With respect to more
ancient glacial periods, several geologists are convinced, from direct
evidence, that such occurred during the miocene and eocene formations, not
to mention still more ancient formations. But the most important result
for us, arrived at by Mr. Croll, is that whenever the northern hemisphere
passes through a cold period the temperature of the southern hemisphere is
actually raised, with the winters rendered much milder, chiefly through
changes in the direction of the ocean currents. So conversely it will be
with the northern hemisphere, while the southern passes through a glacial
period. This conclusion throws so much light on geographical distribution
that I am strongly inclined to trust in it; but I will first give the facts
which demand an explanation.

In South America, Dr. Hooker has shown that besides many closely allied
species, between forty and fifty of the flowering plants of Tierra del
Fuego, forming no inconsiderable part of its scanty flora, are common to
North America and Europe, enormously remote as these areas in opposite
hemispheres are from each other. On the lofty mountains of equatorial
America a host of peculiar species belonging to European genera occur. On
the Organ Mountains of Brazil some few temperate European, some Antarctic
and some Andean genera were found by Gardner which do not exist in the low
intervening hot countries. On the Silla of Caraccas the illustrious
Humboldt long ago found species belonging to genera characteristic of the

In Africa, several forms characteristic of Europe, and some few
representatives of the flora of the Cape of Good Hope, occur on the
mountains of Abyssinia. At the Cape of Good Hope a very few European
species, believed not to have been introduced by man, and on the mountains
several representative European forms are found which have not been
discovered in the intertropical parts of Africa. Dr. 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 neighbouring Cameroon Mountains, in
the Gulf of Guinea, are closely related to those on the mountains of
Abyssinia, and likewise to those of temperate Europe. It now also appears,
as I hear from Dr. Hooker, that some of these same temperate plants have
been discovered by the Rev. R.T. Lowe on the mountains of the Cape Verde
Islands. This extension of the same temperate forms, almost under the
equator, across the whole continent of Africa and to the mountains of the
Cape Verde archipelago, is one of the most astonishing facts ever recorded
in the distribution of plants.

On the Himalaya, and on the isolated mountain ranges of the peninsula of
India, on the heights of Ceylon, and on the volcanic cones of Java, many
plants occur either identically the same or representing each other, and at
the same time representing plants of Europe not found in the intervening
hot lowlands. A list of the genera of plants collected on the loftier
peaks of Java, raises a picture of a collection made on a hillock in
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. Some of these Australian forms, as I hear from Dr. Hooker, extend
along the heights of the peninsula of Malacca, and are thinly scattered on
the one hand over India, and on the other hand as far north as Japan.

On the southern mountains of Australia, Dr. F. Muller has discovered
several European species; other species, not introduced by man, occur on
the lowlands; and a long list can be given, as I am informed by Dr. Hooker,
of European genera, found in Australia, but not in the intermediate torrid
regions. In the admirable Introduction to the Flora of New Zealand by
Dr. Hooker, analogous and striking facts are given in regard to the plants
of that large island. Hence, we see that certain plants growing on the
more lofty mountains of the tropics in all parts of the world, and on the
temperate plains of the north and south, are either the same species or
varieties of the same species. It should, however, be observed that these
plants are not strictly arctic forms; for, as Mr. H.C. Watson has remarked,
in receding from polar toward equatorial latitudesthe Alpine or mountain
flora really become less and less Arctic." Besides these identical and
closely allied formsmany species inhabiting the same widely sundered
areasbelong to genera not now found in the intermediate tropical

These brief remarks apply to plants alone; but some few analogous facts
could be given in regard to terrestrial animals. In marine productions
similar cases likewise occur; as an exampleI may quote a statement by the
highest authorityProf. Danathat "it is certainly a wonderful fact that
New Zealand should have a closer resemblance in its crustacea to Great
Britainits antipodethan to any other part of the world." Sir J.
Richardsonalsospeaks of the reappearance on the shores of New Zealand
Tasmaniaetc.of northern forms of fish. Dr. Hooker informs me that
twenty-five species of Algae are common to New Zealand and to Europebut
have not been found in the intermediate tropical seas.

>From the foregoing factsnamelythe presence of temperate forms on the
highlands across the whole of equatorial Africaand along the Peninsula of
Indiato Ceylon and the Malay Archipelagoand in a less well-marked
manner across the wide expanse of tropical South Americait appears almost
certain that at some former periodno doubt during the most severe part of
a Glacial periodthe lowlands of these great continents were everywhere
tenanted under the equator by a considerable number of temperate forms. 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 latitudeor perhaps even rather cooler.
During thisthe coldest periodthe lowlands under the equator must have
been clothed with a mingled tropical and temperate vegetationlike that
described by Hooker as growing luxuriantly at the height of from four to
five thousand feet on the lower slopes of the Himalayabut with perhaps a

still greater preponderance of temperate forms. So again in the
mountainous island of Fernando Poin the Gulf of GuineaMr. Mann found
temperate European forms beginning to appear at the height of about five
thousand feet. On the mountains of Panamaat the height of only two
thousand feetDr. Seemann found the vegetation like that of Mexicowith
forms of the torrid zone harmoniously blended with those of the temperate.

Now let us see whether Mr. Croll's conclusion that when the northern
hemisphere suffered from the extreme cold of the great Glacial periodthe
southern hemisphere was actually warmerthrows any clear light on the
present apparently inexplicable distribution of various organisms in the
temperate parts of both hemispheresand on the mountains of the tropics.
The Glacial periodas measured by yearsmust have been very long; and
when we remember over what vast spaces some naturalised plants and animals
have spread within a few centuriesthis period will have been ample for
any amount of migration. As the cold became more and more intensewe know
that Arctic forms invaded the temperate regions; and from the facts just
giventhere can hardly be a doubt that some of the more vigorousdominant
and widest-spreading temperate forms invaded the equatorial lowlands. The
inhabitants of these hot lowlands would at the same time have migrated to
the tropical and subtropical regions of the southfor the southern
hemisphere was at this period warmer. On the decline of the Glacial
periodas both hemispheres gradually recovered their former temperature
the northern temperate forms living on the lowlands under the equator
would have been driven to their former homes or have been destroyedbeing
replaced by the equatorial forms returning from the south. Somehowever
of the northern temperate forms would almost certainly have ascended any
adjoining high landwhereif sufficiently loftythey would have long
survived like the Arctic forms on the mountains of Europe. They might have
survivedeven if the climate was not perfectly fitted for themfor the
change of temperature must have been very slowand plants undoubtedly
possess a certain capacity for acclimatisationas shown by their
transmitting to their offspring different constitutional powers of
resisting heat and cold.

In the regular course of events the southern hemisphere would in its turn
be subjected to a severe Glacial periodwith the northern hemisphere
rendered warmer; and then the southern temperate forms would invade the
equatorial lowlands. The northern forms which had before been left on the
mountains would now descend and mingle with the southern forms. These
latterwhen the warmth returnedwould return to their former homes
leaving some few species on the mountainsand carrying southward with them
some of the northern temperate forms which had descended from their
mountain fastnesses. Thuswe should have some few species identically the
same in the northern and southern temperate zones and on the mountains of
the intermediate tropical regions. But the species left during a long time
on these mountainsor in opposite hemisphereswould have to compete with
many new forms and would be exposed to somewhat different physical
conditions; hencethey would be eminently liable to modificationand
would generally now exist as varieties or as representative species; and
this is the case. We mustalsobear in mind the occurrence in both
hemispheres of former Glacial periods; for these will accountin
accordance with the same principlesfor the many quite distinct species
inhabiting the same widely separated areasand belonging to genera not now
found in the intermediate torrid zones.

It is a remarkable factstrongly insisted on by Hooker in regard to
Americaand by Alph. de Candolle in regard to Australiathat many more
identical or slightly modified species have migrated from the north to the
souththan in a reversed direction. We seehowevera few southern forms
on the mountains of Borneo and Abyssinia. I suspect that this preponderant
migration from the north to the south is due to the greater extent of land
in the northand to the northern forms having existed in their own homes
in greater numbersand having consequently been advanced through natural

selection and competition to a higher stage of perfectionor dominating
powerthan the southern forms. And thuswhen the two sets became
commingled in the equatorial regionsduring the alternations of the
Glacial periodsthe northern forms were the more powerful and were able to
hold their places on the mountainsand afterwards migrate southward with
the southern forms; but not so the southern in regard to the northern
forms. In the same mannerat the present daywe see that very many
European productions cover the ground in La PlataNew Zealandand to a
lesser degree in Australiaand have beaten the natives; whereas extremely
few southern forms have become naturalised in any part of the northern
hemispherethough hideswooland 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. The Neilgherrie Mountains in Indiahoweveroffer a partial
exception; for hereas I hear from Dr. HookerAustralian forms are
rapidly sowing themselves and becoming naturalised. Before the last great
Glacial periodno doubt the intertropical mountains were stocked with
endemic Alpine forms; but these have almost everywhere yielded to the more
dominant forms generated in the larger areas and more efficient workshops
of the north. In many islands the native productions are nearly equalled
or even outnumberedby those which have become naturalised; and this is
the first stage towards their extinction. Mountains are islands on the
land; and their inhabitants have yielded to those produced within the
larger areas of the northjust in the same way as the inhabitants of real
islands have everywhere yielded and are still yielding to continental forms
naturalised through man's agency.

The same principles apply to the distribution of terrestrial animals and of
marine productionsin the northern and southern temperate zonesand on
the intertropical mountains. Whenduring the height of the Glacial
periodthe ocean-currents were widely different to what they now aresome
of the inhabitants of the temperate seas might have reached the equator; of
these a few would perhaps at once be able to migrate southwardsby keeping
to the cooler currentswhile 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; in nearly the same manner as
according to Forbesisolated spaces inhabited by Arctic productions exist
to the present day in the deeper parts of the northern temperate seas.

I am far from supposing that all the difficulties in regard to the
distribution and affinities of the identical and allied specieswhich now
live so widely separated in the north and southand sometimes on the
intermediate mountain rangesare removed on the views above given. The
exact lines of migration cannot be indicated. We cannot say why certain
species and not others have migrated; why certain species have been
modified and have given rise to new formswhile others have remained
unaltered. We cannot hope to explain such factsuntil we can say why one
species and not another becomes naturalised by man's agency in a foreign
land; why one species ranges twice or thrice as farand is twice or thrice
as commonas another species within their own homes.

Various special difficulties also remain to be solved; for instancethe
occurrenceas shown by Dr. Hookerof the same plants at points so
enormously remote as Kerguelen LandNew Zealandand Fuegia; but icebergs
as suggested by Lyellmay have been concerned in their dispersal. The
existence at these and other distant points of the southern hemisphereof
specieswhichthough distinctbelong to genera exclusively confined to
the southis a more remarkable case. Some of these species are so
distinctthat we cannot suppose that there has been time since the
commencement of the last Glacial period for their migration and subsequent
modification to the necessary degree. The facts seem to indicate that
distinct species belonging to the same genera have migrated in radiating
lines from a common centre; and I am inclined to look in the southernas
in the northern hemisphereto a former and warmer periodbefore the

commencement of the last Glacial periodwhen the Antarctic landsnow
covered with icesupported a highly peculiar and isolated flora. It may
be suspected that before this flora was exterminated during the last
Glacial epocha few forms had been already widely dispersed to various
points of the southern hemisphere by occasional means of transportand by
the aidas halting-placesof now sunken islands. Thus the southern
shores of AmericaAustraliaand New Zealand may have become slightly
tinted by the same peculiar forms of life.

Sir C. Lyell in a striking passage has speculatedin language almost
identical with mineon the effects of great alternations of climate
throughout the world on geographical distribution. 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
together with the admission of the slow modification of speciesexplains a
multitude of facts in the distribution of the same and of the allied forms
of life in all parts of the globe. The living waters have flowed during
one period from the north and during another from the southand in both
cases have reached the equator; but the stream of life has flowed with
greater force from the north than in the opposite directionand has
consequently more freely inundated the south. As the tide leaves its drift
in horizontal linesrising higher on the shores where the tide rises
highestso have the living waters left their living drift on our mountain
summitsin a line gently rising from the Arctic lowlands to a great
latitude under the equator. The various beings thus left stranded may be
compared with savage races of mandriven up and surviving in the mountain
fastnesses of almost every landwhich serves as a recordfull of interest
to usof the former inhabitants of the surrounding lowlands.



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.


As lakes and river-systems are separated from each other by barriers of
landit might have been thought that fresh-water productions would not
have ranged widely within the same countryand as the sea is apparently a
still more formidable barrierthat they would never have extended to
distant countries. But the case is exactly the reverse. Not only have
many fresh-water speciesbelonging to different classesan enormous
rangebut allied species prevail in a remarkable manner throughout the
world. When first collecting in the fresh waters of BrazilI well
remember feeling much surprise at the similarity of the fresh-water
insectsshellsetc.and at the dissimilarity of the surrounding
terrestrial beingscompared with those of Britain.

But the wide ranging power of fresh-water productions canI thinkin most
cases be explained by their having become fittedin a manner highly useful
to themfor short and frequent migrations from pond to pondor from
stream to streamwithin their own countries; and liability to wide
dispersal would follow from this capacity as an almost necessary
consequence. We can here consider only a few cases; of thesesome of the
most difficult to explain are presented by fish. It was formerly believed
that the same fresh-water species never existed on two continents distant
from each other. But Dr. Gunther has lately shown that the Galaxias
attenuatus inhabits TasmaniaNew Zealandthe Falkland Islands and the

mainland of South America. This is a wonderful caseand probably
indicates dispersal from an Antarctic centre during a former warm period.
This casehoweveris 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 Islandsthough separated by a distance of
about 230 miles. On the same continent fresh-water fish often range
widelyand as if capriciously; for in two adjoining river systems some of
the species may be the same and some wholly different.

It is probable that they are occasionally transported by what may be called
accidental means. Thus fishes still alive are not very rarely dropped at
distant points by whirlwinds; and it is known that the ova retain their
vitality for a considerable time after removal from the water. Their
dispersal mayhoweverbe mainly attributed to changes in the level of the
land within the recent periodcausing rivers to flow into each other.
Instancesalsocould be given of this having occurred during floods
without any change of level. The wide differences of the fish on the
opposite sides of most mountain-rangeswhich are continuous and
consequently mustfrom an early periodhave completely prevented the
inosculation of the river systems on the two sidesleads to the same
conclusion. Some fresh-water fish belong to very ancient formsand in
such cases there will have been ample time for great geographical changes
and consequently time and means for much migration. MoreoverDr. Gunther
has recently been led by several considerations to infer that with fishes
the same forms have a long endurance. Salt-water fish can with care be
slowly accustomed to live in fresh water; andaccording to Valenciennes
there is hardly a single group of which all the members are confined to
fresh waterso that a marine species belonging to a fresh-water group
might travel far along the shores of the seaand couldit is probable
become adapted without much difficulty to the fresh waters of a distant

Some species of fresh-water shells have very wide rangesand allied
species whichon our theoryare descended from a common parentand must
have proceeded from a single sourceprevail throughout the world. Their
distribution at first perplexed me muchas their ova are not likely to be
transported by birds; and the ovaas well as the adultsare immediately
killed by sea-water. I could not even understand how some naturalised
species have spread rapidly throughout the same country. But two facts
which I have observed--and many others no doubt will be discovered--throw
some light on this subject. When ducks suddenly emerge from a pond covered
with duck-weedI have twice seen these little plants adhering to their
backs; and it has happened to mein removing a little duck-weed from one
aquarium to anotherthat I have unintentionally stocked the one with
fresh-water shells from the other. But another agency is perhaps more
effectual: I suspended the feet of a duck in an aquariumwhere many ova
of fresh-water shells were hatching; and I found that numbers of the
extremely minute and just-hatched shells crawled on the feetand clung to
them so firmly that when taken out of the water they could not be jarred
offthough at a somewhat more advanced age they would voluntarily drop
off. These just-hatched molluscsthough aquatic in their naturesurvived
on the duck's feetin damp airfrom twelve to twenty hours; and in this
length of time a duck or heron might fly at least six or seven hundred
milesand if blown across the sea to an oceanic islandor to any other
distant pointwould be sure to alight on a pool or rivulet. Sir Charles
Lyell informs me that a Dyticus has been caught with an Ancylus (a
fresh-water shell like a limpet) firmly adhering to it; and a water-beetle
of the same familya Colymbetesonce flew on board the "Beagle 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.

With respect to plants, it has long been known what enormous ranges many
fresh-water, and even marsh-species, have, both over continents and to the

most remote oceanic islands. This is strikingly illustrated, according to
Alph. de Candolle, in those large groups of terrestrial plants, which have
very few aquatic members; for the latter seem immediately to acquire, as if
in consequence, a wide range. I think favourable means of dispersal
explain this fact. I have before mentioned that earth occasionally adheres
in some quantity to the feet and beaks of birds. Wading birds, which
frequent the muddy edges of ponds, if suddenly flushed, would be the most
likely to have muddy feet. Birds of this order wander more than those of
any other; and are occasionally found on the most remote and barren islands
of the open ocean; they would not be likely to alight on the surface of the
sea, so that any dirt on their feet would not be washed off; and when
gaining the land, they would be sure to fly to their natural fresh-water
haunts. I do not believe that botanists are aware how charged the mud of
ponds is with seeds: I have tried several little experiments, but will
here give only the most striking case: I took in February three
tablespoonfuls of mud from three different points, beneath water, on the
edge of a little pond; this mud when dry weighed only 6 and 3/4 ounces; I
kept it covered up in my study for six months, pulling up and counting each
plant as it grew; the plants were of many kinds, and were altogether 537 in
number; and yet the viscid mud was all contained in a breakfast cup!
Considering these facts, 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, situated at very distant points. The same
agency may have come into play with the eggs of some of the smaller
fresh-water animals.

Other and unknown agencies probably have also played a part. I have stated
that fresh-water fish eat some kinds of seeds, though they reject many
other kinds after having swallowed them; even small fish swallow seeds of
moderate size, as of the yellow water-lily and Potamogeton. Herons and
other birds, century after century, have gone on daily devouring fish; they
then take flight and go to other waters, or are blown across the sea; and
we have seen that seeds retain their power of germination, when rejected
many hours afterwards in pellets or in the excrement. When I saw the great
size of the seeds of that fine water-lily, the Nelumbium, and remembered
Alph. de Candolle's remarks on the distribution of this plant, I thought
that the means of its dispersal must remain inexplicable; but Audubon
states that he found the seeds of the great southern water-lily (probably
according to Dr. Hooker, the Nelumbium luteum) in a heron's stomach. Now
this bird must often have flown with its stomach thus well stocked to
distant ponds, and, then getting a hearty meal of fish, analogy makes me
believe that it would have rejected the seeds in the pellet in a fit state
for germination.

In considering these several means of distribution, it should be remembered
that when a pond or stream is first formed, for instance on a rising islet,
it will be unoccupied; and a single seed or egg will have a good chance of
succeeding. Although there will always be a struggle for life between the
inhabitants of the same pond, however few in kind, 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, the competition between them will
probably be less severe than between terrestrial species; consequently an
intruder from the waters of a foreign country would have a better chance of
seizing on a new place, than in the case of terrestrial colonists. We
should also remember that many fresh-water productions are low in the scale
of nature, and we have reason to believe that such beings become modified
more slowly than the high; and this will give time for the migration of
aquatic species. We should not forget the probability of many fresh-water
forms having formerly ranged continuously over immense areas, and then
having become extinct at intermediate points. But the wide distribution of
fresh-water plants, and of the lower animals, whether retaining the same
identical form, or in some degree modified, apparently depends in main part
on the wide dispersal of their seeds and eggs by animals, more especially
by fresh-water birds, which have great powers of flight, and naturally

travel from one piece of water to another.


We now come to the last of the three classes of facts, which I have
selected as presenting the greatest amount of difficulty with respect to
distribution, on the view that not only all the individuals of the same
species have migrated from some one area, but that allied species, although
now inhabiting the most distant points, have proceeded from a single area,
the birthplace of their early progenitors. 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. This
view removes many difficulties, but it does not accord with all the facts
in regard to the productions of islands. In the following remarks I shall
not confine myself to the mere question of dispersal, but shall consider
some other cases bearing on the truth of the two theories of independent
creation and of descent with modification.

The species of all kinds which inhabit oceanic islands are few in number
compared with those on equal continental areas: Alph. de Candolle admits
this for plants, and Wollaston for insects. New Zealand, for instance,
with its lofty mountains and diversified stations, extending over 780 miles
of latitude, together with the outlying islands of Auckland, Campbell and
Chatham, 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, we must admit that
some cause, independently of different physical conditions, has given rise
to so great a difference in number. Even the uniform county of Cambridge
has 847 plants, and the little island of Anglesea 764, but a few ferns and
a few introduced plants are included in these numbers, and the comparison
in some other respects is not quite fair. We have evidence that the barren
island of Ascension aboriginally possessed less than half-a-dozen flowering
plants; yet many species have now become naturalised on it, as they have in
New Zealand and on every other oceanic island which can be named. In St.
Helena there is reason to believe that the naturalised plants and animals
have nearly or quite exterminated many native productions. He who admits
the doctrine of the creation of each separate species, will have to admit
that a sufficient number of the best adapted plants and animals were not
created for oceanic islands; for man has unintentionally stocked them far
more fully and perfectly than did nature.

Although in oceanic islands the species are few in number, the proportion
of endemic kinds (i.e. those found nowhere else in the world) is often
extremely large. If we compare, for instance, the number of endemic
land-shells in Madeira, or of endemic birds in the Galapagos Archipelago,
with the number found on any continent, and then compare the area of the
island with that of the continent, we shall see that this is true. This
fact might have been theoretically expected, for, as already explained,
species occasionally arriving, after long intervals of time in the new and
isolated district, and having to compete with new associates, would be
eminently liable to modification, and would often produce groups of
modified descendants. But it by no means follows that, because in an
island nearly all the species of one class are peculiar, those of another
class, or of another section of the same class, are peculiar; and this
difference seems to depend partly on the species which are not modified
having immigrated in a body, so that their mutual relations have not been
much disturbed; and partly on the frequent arrival of unmodified immigrants
from the mother-country, with which the insular forms have intercrossed.
It should be borne in mind that the offspring of such crosses would
certainly gain in vigour; so that even an occasional cross would produce
more effect than might have been anticipated. I will give a few
illustrations of the foregoing remarks: in the Galapagos Islands there are
twenty-six land birds; of these twenty-one (or perhaps twenty-three) are

peculiar; whereas of the eleven marine birds only two are peculiar; and it
is obvious that marine birds could arrive at these islands much more easily
and frequently than land-birds. Bermuda, on the other hand, which lies at
about the same distance from North America as the Galapagos Islands do from
South America, and which has a very peculiar soil, does not possess a
single endemic land bird; and we know from Mr. J.M. Jones's admirable
account of Bermuda, that very many North American birds occasionally or
even frequently visit this island. Almost every year, as I am informed by
Mr. E.V. Harcourt, many European and African birds are blown to Madeira;
this island is inhabited by ninety-nine kinds, of which one alone is
peculiar, though very closely related to a European form; and three or four
other species are confined to this island and to the Canaries. So that the
islands of Bermuda and Madeira have been stocked from the neighbouring
continents with birds, which for long ages have there struggled together,
and have become mutually co-adapted. Hence, when settled in their new
homes, each kind will have been kept by the others to its proper place and
habits, and will consequently have been but little liable to modification.
Any tendency to modification will also have been checked by intercrossing
with the unmodified immigrants, often arriving from the mother-country.
Madeira again is inhabited by a wonderful number of peculiar land-shells,
whereas not one species of sea-shell is peculiar to its shores: now,
though we do not know how sea-shells are dispersed, yet we can see that
their eggs or larvae, perhaps attached to seaweed or floating timber, or to
the feet of wading birds, might be transported across three or four hundred
miles of open sea far more easily than land-shells. The different orders
of insects inhabiting Madeira present nearly parallel cases.

Oceanic islands are sometimes deficient in animals of certain whole
classes, and their places are occupied by other classes; thus in the
Galapagos Islands reptiles, and in New Zealand gigantic wingless birds,
take, or recently took, the place of mammals. Although New Zealand is here
spoken of as an oceanic island, it is in some degree doubtful whether it
should be so ranked; it is of large size, and is not separated from
Australia by a profoundly deep sea; from its geological character and the
direction of its mountain ranges, the Rev. W.B. Clarke has lately
maintained that this island, as well as New Caledonia, should be considered
as appurtenances of Australia. Turning to plants, Dr. Hooker has shown
that in the Galapagos Islands the proportional numbers of the different
orders are very different from what they are elsewhere. All such
differences in number, and the absence of certain whole groups of animals
and plants, are generally accounted for by supposed differences in the
physical conditions of the islands; but this explanation is not a little
doubtful. Facility of immigration seems to have been fully as important as
the nature of the conditions.

Many remarkable little facts could be given with respect to the inhabitants
of oceanic islands. For instance, in certain islands not tenanted by a
single mammal, some of the endemic plants have beautifully hooked seeds;
yet few relations are more manifest than that hooks serve for the
transportal of seeds in the wool or fur of quadrupeds. But a hooked seed
might be carried to an island by other means; and the plant then becoming
modified would form an endemic species, still retaining its hooks, which
would form a useless appendage, like the shrivelled wings under the
soldered wing-covers of many insular beetles. Again, islands often possess
trees or bushes belonging to orders which elsewhere include only herbaceous
species; now trees, as Alph. de Candolle has shown, generally have,
whatever the cause may be, confined ranges. Hence trees would be little
likely to reach distant oceanic islands; and an herbaceous plant, which had
no chance of successfully competing with the many fully developed trees
growing on a continent, might, when established on an island, gain an
advantage over other herbaceous plants by growing taller and taller and
overtopping them. In this case, natural selection would tend to add to the
stature of the plant, to whatever order it belonged, and thus first convert
it into a bush and then into a tree.


With respect to the absence of whole orders of animals on oceanic islands,
Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts)
are never found on any of the many islands with which the great oceans are
studded. I have taken pains to verify this assertion, and have found it
true, with the exception of New Zealand, New Caledonia, the Andaman
Islands, and perhaps the Solomon Islands and the Seychelles. But I have
already remarked that it is doubtful whether New Zealand and New Caledonia
ought to be classed as oceanic islands; and this is still more doubtful
with respect to the Andaman and Solomon groups and the Seychelles. This
general absence of frogs, toads and newts on so many true oceanic islands
cannot be accounted for by their physical conditions; indeed it seems that
islands are peculiarly fitted for these animals; for frogs have been
introduced into Madeira, the Azores, and Mauritius, and have multiplied so
as to become a nuisance. But as these animals and their spawn are
immediately killed (with the exception, as far as known, of one Indian
species) by sea-water, there would be great difficulty in their transportal
across the sea, and therefore we can see why they do not exist on strictly
oceanic islands. But why, on the theory of creation, they should not have
been created there, it would be very difficult to explain.

Mammals offer another and similar case. I have carefully searched the
oldest voyages, and have not found a single instance, free from doubt, of a
terrestrial mammal (excluding domesticated animals kept by the natives)
inhabiting an island situated above 300 miles from a continent or great
continental island; and many islands situated at a much less distance are
equally barren. The Falkland Islands, which are inhabited by a wolf-like
fox, come nearest to an exception; but this group cannot be considered as
oceanic, as it lies on a bank in connection with the mainland at a distance
of about 280 miles; moreover, icebergs formerly brought boulders to its
western shores, and they may have formerly transported foxes, as now
frequently happens in the arctic regions. Yet it cannot be said that small
islands will not support at least small mammals, for they occur in many
parts of the world on very small islands, when lying close to a continent;
and hardly an island can be named on which our smaller quadrupeds have not
become naturalised and greatly multiplied. It cannot be said, on the
ordinary view of creation, that there has not been time for the creation of
mammals; many volcanic islands are sufficiently ancient, as shown by the
stupendous degradation which they have suffered, and by their tertiary
strata: there has also been time for the production of endemic species
belonging to other classes; and on continents it is known that new species
of mammals appear and disappear at a quicker rate than other and lower
animals. Although terrestrial mammals do not occur on oceanic islands,
aerial mammals do occur on almost every island. New Zealand possesses two
bats found nowhere else in the world: Norfolk Island, the Viti
Archipelago, the Bonin Islands, the Caroline and Marianne Archipelagoes,
and Mauritius, all possess their peculiar bats. Why, it may be asked, has
the supposed creative force produced bats and no other mammals on remote
islands? On my view this question can easily be answered; for no
terrestrial mammal can be transported across a wide space of sea, but bats
can fly across. Bats have been seen wandering by day far over the Atlantic
Ocean; and two North American species, either regularly or occasionally,
visit Bermuda, at the distance of 600 miles from the mainland. I hear from
Mr. Tomes, who has specially studied this family, that many species have
enormous ranges, and are found on continents and on far distant islands.
Hence, we have only to suppose that such wandering species have been
modified in their new homes in relation to their new position, and we can
understand the presence of endemic bats on oceanic islands, with the
absence of all other terrestrial mammals.

Another interesting relation exists, namely, between the depth of the sea
separating islands from each other, or from the nearest continent, and the

degree of affinity of their mammalian inhabitants. Mr. Windsor Earl has
made some striking observations on this head, since greatly extended by Mr.
Wallace's admirable researches, in regard to the great Malay Archipelago,
which is traversed near Celebes by a space of deep ocean, and this
separates two widely distinct mammalian faunas. On either side, the
islands stand on a moderately shallow submarine bank, and these islands are
inhabited by the same or by closely allied quadrupeds. I have not as yet
had time to follow up this subject in all quarters of the world; but as far
as I have gone, the relation holds good. For instance, Britain is
separated by a shallow channel from Europe, and the mammals are the same on
both sides; and so it is with all the islands near the shores of Australia.
The West Indian Islands, on the other hand, stand on a deeply submerged
bank, nearly one thousand fathoms in depth, and here we find American
forms, but the species and even the genera are quite distinct. As the
amount of modification which animals of all kinds undergo partly depends on
the lapse of time, and as the islands which are separated from each other,
or from the mainland, by shallow channels, are more likely to have been
continuously united within a recent period than the islands separated by
deeper channels, we can understand how it is that a relation exists between
the depth of the sea separating two mammalian faunas, and the degree of
their affinity, a relation which is quite inexplicable on the theory of
independent acts of creation.

The foregoing statements in regard to the inhabitants of oceanic islands,
namely, the fewness of the species, with a large proportion consisting of
endemic forms--the members of certain groups, but not those of other groups
in the same class, having been modified--the absence of certain whole
orders, as of batrachians and of terrestrial mammals, notwithstanding the
presence of aerial bats, the singular proportions of certain orders of
plants, herbaceous forms having been developed into trees, etc., seem to me
to accord better with the belief in the efficiency of occasional means of
transport, carried on during a long course of time, than with the belief in
the former connection of all oceanic islands with the nearest continent;
for on this latter view it is probable that the various classes would have
immigrated more uniformly, and from the species having entered in a body,
their mutual relations would not have been much disturbed, and
consequently, they would either have not been modified, or all the species
in a more equable manner.

I do not deny that there are many and serious difficulties in understanding
how many of the inhabitants of the more remote islands, whether still
retaining the same specific form or subsequently modified, have reached
their present homes. But the probability of other islands having once
existed as halting-places, of which not a wreck now remains, must not be
overlooked. I will specify one difficult case. Almost all oceanic
islands, even the most isolated and smallest, are inhabited by land-shells,
generally by endemic species, but sometimes by species found elsewhere
striking instances of which have been given by Dr. A.A. Gould in relation
to the Pacific. Now it is notorious that land-shells are easily killed by
sea-water; their eggs, at least such as I have tried, sink in it and are
killed. Yet there must be some unknown, but occasionally efficient means
for their transportal. Would the just-hatched young sometimes adhere to
the feet of birds roosting on the ground and thus get transported? It
occurred to me that land-shells, when hybernating and having a membranous
diaphragm over the mouth of the shell, might be floated in chinks of
drifted timber across moderately wide arms of the sea. And I find that
several species in this state withstand uninjured an immersion in sea-water
during seven days. One shell, the Helix pomatia, after having been thus
treated, and again hybernating, was put into sea-water for twenty days and
perfectly recovered. During this length of time the shell might have been
carried by a marine country of average swiftness to a distance of 660
geographical miles. As this Helix has a thick calcareous operculum I
removed it, and when it had formed a new membranous one, I again immersed
it for fourteen days in sea-water, and again it recovered and crawled away.

Baron Aucapitaine has since tried similar experiments. He placed 100 landshells,
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.


The most striking and important fact for us is the affinity of the species
which inhabit islands to those of the nearest mainland, without being
actually the same. Numerous instances could be given. The Galapagos
Archipelago, situated under the equator, lies at a distance of between 500
and 600 miles from the shores of South America. Here almost every product
of the land and of the water bears the unmistakable stamp of the American
continent. There are twenty-six land birds. Of these twenty-one, or
perhaps twenty-three, are ranked as distinct species, and would commonly be
assumed to have been here created; yet the close affinity of most of these
birds to American species is manifest in every character in their habits,
gestures, and tones of voice. So it is with the other animals, and with a
large proportion of the plants, as shown by Dr. Hooker in his admirable
Flora of this archipelago. The naturalist, looking at the inhabitants of
these volcanic islands in the Pacific, distant several hundred miles from
the continent, feels that he is standing on American land. Why should this
be so? Why should the species which are supposed to have been created in
the Galapagos Archipelago, and nowhere else, bear so plainly the stamp of
affinity to those created in America? There is nothing in the conditions
of life, in the geological nature of the islands, in their height or
climate, or in the proportions in which the several classes are associated
together, which closely resembles the conditions of the South American
coast. In fact, there is a considerable dissimilarity in all these
respects. On the other hand, there is a considerable degree of resemblance
in the volcanic nature of the soil, in the climate, height, and size of the
islands, between the Galapagos and Cape Verde Archipelagos: but what an
entire and absolute difference in their inhabitants! The inhabitants of
the Cape Verde Islands are related to those of Africa, like those of the
Galapagos to America. Facts, such as these, admit of no sort of
explanation on the ordinary view of independent creation; whereas, on the
view here maintained, it is obvious that the Galapagos Islands would be
likely to receive colonists from America, whether by occasional means of
transport or (though I do not believe in this doctrine) by formerly
continuous land, and the Cape Verde Islands from Africa; such colonists
would be liable to modification--the principle of inheritance still
betraying their original birthplace.

Many analogous facts could be given: indeed it is an almost universal rule
that the endemic productions of islands are related to those of the nearest
continent, or of the nearest large island. The exceptions are few, and
most of them can be explained. Thus, although Kerguelen Land stands nearer
to Africa than to America, the plants are related, and that very closely,
as we know from Dr. Hooker's account, to those of America: but on the view
that this island has been mainly stocked by seeds brought with earth and
stones on icebergs, drifted by the prevailing currents, this anomaly
disappears. New Zealand in its endemic plants is much more closely related
to Australia, the nearest mainland, than to any other region: and this is
what might have been expected; but it is also plainly related to South
America, which, although the next nearest continent, is so enormously
remote, that the fact becomes an anomaly. But this difficulty partially
disappears on the view that New Zealand, South America, and the other

southern lands, have been stocked in part from a nearly intermediate though
distant point, namely, from the antarctic islands, when they were clothed
with vegetation, during a warmer tertiary period, before the commencement
of the last Glacial period. The affinity, which, though feeble, I am
assured by Dr. Hooker is real, between the flora of the south-western
corner of Australia and of the Cape of Good Hope, is a far more remarkable
case; but this affinity is confined to the plants, and will, no doubt, some
day be explained.

The same law which has determined the relationship between the inhabitants
of islands and the nearest mainland, is sometimes displayed on a small
scale, but in a most interesting manner, within the limits of the same
archipelago. Thus each separate island of the Galapagos Archipelago is
tenanted, and the fact is a marvellous one, by many distinct species; but
these species are related to each other in a very much closer manner than
to the inhabitants of the American continent, or of any other quarter of
the world. This is what might have been expected, for islands situated so
near to each other would almost necessarily receive immigrants from the
same original source, and from each other. But how is it that many of the
immigrants have been differently modified, though only in a small degree,
in islands situated within sight of each other, having the same geological
nature, the same height, climate, etc? This long appeared to me a great
difficulty: but it arises in chief part from the deeply-seated error of
considering the physical conditions of a country as the most important;
whereas it cannot be disputed that the nature of the other species with
which each has to compete, is at least as important, and generally a far
more important element of success. Now if we look to the species which
inhabit the Galapagos Archipelago, and are likewise found in other parts of
the world, we find that they differ considerably in the several islands.
This difference might indeed have been expected if the islands have been
stocked by occasional means of transport--a seed, for instance, of one
plant having been brought to one island, and that of another plant to
another island, though all proceeding from the same general source. Hence,
when in former times an immigrant first settled on one of the islands, or
when it subsequently spread from one to another, it would undoubtedly be
exposed to different conditions in the different islands, for it would have
to compete with a different set of organisms; a plant, for instance, would
find the ground best-fitted for it occupied by somewhat different species
in the different islands, and would be exposed to the attacks of somewhat
different enemies. If, then, it varied, natural selection would probably
favour different varieties in the different islands. Some species,
however, might spread and yet retain the same character throughout the
group, just as we see some species spreading widely throughout a continent
and remaining the same.

The really surprising fact in this case of the Galapagos Archipelago, and
in a lesser degree in some analogous cases, is that each new species after
being formed in any one island, did not spread quickly to the other
islands. But the islands, though in sight of each other, are separated by
deep arms of the sea, in most cases wider than the British Channel, and
there is no reason to suppose that they have at any former period been
continuously united. The currents of the sea are rapid and deep between
the islands, and gales of wind are extraordinarily rare; so that the
islands are far more effectually separated from each other than they appear
on a map. Nevertheless, some of the species, both of those found in other
parts of the world and of those confined to the archipelago, are common to
the several islands; and we may infer from the present manner of
distribution that they have spread from one island to the others. But we
often take, I think, an erroneous view of the probability of closely allied
species invading each other's territory, when put into free
intercommunication. Undoubtedly, if one species has any advantage over
another, it will in a very brief time wholly or in part supplant it; but if
both are equally well fitted for their own places, both will probably hold
their separate places for almost any length of time. Being familiar with

the fact that many species, naturalised through man's agency, have spread
with astonishing rapidity over wide areas, we are apt to infer that most
species would thus spread; but we should remember that the species which
become naturalised in new countries are not generally closely allied to the
aboriginal inhabitants, but are very distinct forms, belonging in a large
proportion of cases, as shown by Alph. de Candolle, to distinct genera. In
the Galapagos Archipelago, many even of the birds, though so well adapted
for flying from island to island, differ on the different islands; thus
there are three closely allied species of mocking-thrush, each confined to
its own island. Now let us suppose the mocking-thrush of Chatham Island to
be blown to Charles Island, which has its own mocking-thrush; why should it
succeed in establishing itself there? We may safely infer that Charles
Island is well stocked with its own species, for annually more eggs are
laid and young birds hatched than can possibly be reared; and we may infer
that the mocking-thrush peculiar to Charles Island is at least as well
fitted for its home as is the species peculiar to Chatham Island. Sir C.
Lyell and Mr. Wollaston have communicated to me a remarkable fact bearing
on this subject; namely, that Madeira and the adjoining islet of Porto
Santo possess many distinct but representative species of land-shells, some
of which live in crevices of stone; and although large quantities of stone
are annually transported from Porto Santo to Madeira, yet this latter
island has not become colonised by the Porto Santo species: nevertheless,
both islands have been colonised by some European land-shells, which no
doubt had some advantage over the indigenous species. From these
considerations I think we need not greatly marvel at the endemic species
which inhabit the several islands of the Galapagos Archipelago not having
all spread from island to island. On the same continent, also,
pre-occupation has probably played an important part in checking the
commingling of the species which inhabit different districts with nearly
the same physical conditions. Thus, the south-east and south-west corners
of Australia have nearly the same physical conditions, and are united by
continuous land, yet they are inhabited by a vast number of distinct
mammals, birds, and plants; so it is, according to Mr. Bates, with the
butterflies and other animals inhabiting the great, open, and continuous
valley of the Amazons.

The same principle which governs the general character of the inhabitants
of oceanic islands, namely, the relation to the source whence colonists
could have been most easily derived, together with their subsequent
modification, is of the widest application throughout nature. We see this
on every mountain-summit, in every lake and marsh. For Alpine species,
excepting in as far as the same species have become widely spread during
the Glacial epoch, are related to those of the surrounding lowlands; thus
we have in South America, Alpine humming-birds, Alpine rodents, Alpine
plants, etc., all strictly belonging to American forms; and it is obvious
that a mountain, as it became slowly upheaved, would be colonised from the
surrounding lowlands. So it is with the inhabitants of lakes and marshes,
excepting in so far as great facility of transport has allowed the same
forms to prevail throughout large portions of the world. We see the same
principle in the character of most of the blind animals inhabiting the
caves of America and of Europe. Other analogous facts could be given. It
will, I believe, be found universally true, that wherever in two regions,
let them be ever so distant, many closely allied or representative species
occur, there will likewise be found some identical species; and wherever
many closely-allied species occur, there will be found many forms which
some naturalists rank as distinct species, and others as mere varieties;
these doubtful forms showing us the steps in the process of modification.

The relation between the power and extent of migration in certain species,
either at the present or at some former period, and the existence at remote
points of the world of closely allied species, is shown in another and more
general way. Mr. Gould remarked to me long ago, that in those genera of
birds which range over the world, many of the species have very wide
ranges. I can hardly doubt that this rule is generally true, though

difficult of proof. Among mammals, we see it strikingly displayed in Bats,
and in a lesser degree in the Felidae and Canidae. We see the same rule in
the distribution of butterflies and beetles. So it is with most of the
inhabitants of fresh water, for many of the genera in the most distinct
classes range over the world, and many of the species have enormous ranges.
It is not meant that all, but that some of the species have very wide
ranges in the genera which range very widely. Nor is it meant that the
species in such genera have, on an average, a very wide range; for this
will largely depend on how far the process of modification has gone; for
instance, two varieties of the same species inhabit America and Europe, and
thus the species has an immense range; but, if variation were to be carried
a little further, the two varieties would be ranked as distinct species,
and their range would be greatly reduced. Still less is it meant, that
species which have the capacity of crossing barriers and ranging widely, as
in the case of certain powerfully-winged birds, will necessarily range
widely; for we should never forget that to range widely implies not only
the power of crossing barriers, but the more important power of being
victorious in distant lands in the struggle for life with foreign
associates. But according to the view that all the species of a genus,
though distributed to the most remote points of the world, are descended
from a single progenitor, we ought to find, and I believe as a general rule
we do find, that some at least of the species range very widely.

We should bear in mind that many genera in all classes are of ancient
origin, and the species in this case will have had ample time for dispersal
and subsequent modification. There is also reason to believe, from
geological evidence, that within each great class the lower organisms
change at a slower rate than the higher; consequently they will have had a
better chance of ranging widely and of still retaining the same specific
character. This fact, together with that of the seeds and eggs of most
lowly organised forms being very minute and better fitted for distant
transportal, probably accounts for a law which has long been observed, and
which has lately been discussed by Alph. de Candolle in regard to plants,
namely, that the lower any group of organisms stands the more widely it

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 gen