Darwinism (Wallace)/Chapter III

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Darwinism (Wallace) by Alfred Russel Wallace
Chapter III - THE VARIABILITY OF SPECIES IN A STATE OF NATURE

CHAPTER III

THE VARIABILITY OF SPECIES IN A STATE OF NATURE

Importance of variability—Popular ideas regarding it—Variability of the lower animals—The variability of insects—Variation among lizards—Variation among birds—Diagrams of bird-variation—Number of varying individuals—Variation in the mammalia—Variation in internal organs—Variations in the skull—Variations in the habits of Animals—The Variability of plants—Species which vary little—Concluding remarks.

The foundation of the Darwinian theory is the variability of species, and it is quite useless to attempt even to understand that theory, much less to appreciate the completeness of the proof of it, unless we first obtain a clear conception of the nature and extent of this variability. The most frequent and the most misleading of the objections to the efficacy of natural selection arise from ignorance of this subject, an ignorance shared by many naturalists, for it is only since Mr. Darwin has taught us their importance that varieties have been systematically collected and recorded; and even now very few collectors or students bestow upon them the attention they deserve. By the older naturalists, indeed, varieties—especially if numerous, small, and of frequent occurrence—were looked upon as an unmitigated nuisance, because they rendered it almost impossible to give precise definitions of species, then considered the chief end of systematic natural history. Hence it was the custom to describe what was supposed to be the "typical form" of species, and most collectors were satisfied if they possessed this typical form in their cabinets. Now, however, a collection is valued in proportion as it contains illustrative specimens of all the varieties that occur in each species, and in some cases these have been carefully described, so that we possess a considerable mass of information on the subject. Utilising this information we will now endeavour to give some idea of the nature and extent of variation in the species of animals and plants.

It is very commonly objected that the widespread and constant variability which is admitted to be a characteristic of domesticated animals and cultivated plants is largely due to the unnatural conditions of their existence, and that we have no proof of any corresponding amount of variation occurring in a state of nature. Wild animals and plants, it is said, are usually stable, and when variations occur these are alleged to be small in amount and to affect superficial characters only; or if larger and more important, to occur so rarely as not to afford any aid in the supposed formation of new species.

This objection, as will be shown, is utterly unfounded; but as it is one which goes to the very root of the problem, it is necessary to enter at some length into the various proofs of variation in a state of nature. This is the more necessary because the materials collected by Mr. Darwin bearing on this question have never been published, and comparatively few of them have been cited in The Origin of Species; while a considerable body of facts has been made known since the publication of the last edition of that work.

Variability of the Lower Animals.

Among the lowest and most ancient marine organisms are the Foraminifera, little masses of living jelly, apparently structureless, but which secrete beautiful shelly coverings, often perfectly symmetrical, as varied in form as those of the mollusca and far more complicated. These have been studied with great care by many eminent naturalists, and the late Dr. W.B. Carpenter in his great work—the Introduction to the Study of the Foraminifera—thus refers to their variability: "There is not a single species of plant or animal of which the range of variation has been studied by the collocation and comparison of so large a number of specimens as have passed under the review of Messrs. Williamson, Parker, Rupert Jones, and myself in our studies of the types of this group;" and he states as the result of this extensive comparison of specimens: "The range of variation is so great among the Foraminifera as to include not merely those differential characters which have been usually accounted specific, but also those upon which the greater part of the genera, of this group have been founded, and even in some instances those of its orders."[1]

Coming now to a higher group—the Sea-Anemones—Mr. P.H. Gosse and other writers on these creatures often refer to variations in size, in the thickness and length of the tentacles, the form of the disc and of the mouth, and the character of surface of the column, while the colour varies enormously in a great number of the species. Similar variations occur in all the various groups of marine invertebrata, and in the great sub-kingdom of the mollusca they are especially numerous. Thus, Dr. S.P. Woodward states that many present a most perplexing amount of variation, resulting (as he supposes) from supply of food, variety of depth and of saltness of the water; but we know that many variations are quite independent of such causes, and we will now consider a few cases among the land-mollusca in which they have been more carefully studied.

In the small forest region of Oahu, one of the Sandwich Islands, there have been found about 175 species of land-shells represented by 700 or 800 varieties; and we are told by the Rev. J.T. Gulick, who studied them carefully, that "we frequently find a genus represented in several successive valleys by allied species, sometimes feeding on the same, sometimes on different plants. In every such case the valleys that are nearest to each other furnish the most nearly allied forms; and a full set of the varieties of each species presents a minute gradation of forms between the more divergent types found in the more widely separated localities."

In most land-shells there is a considerable amount of variation in colour, markings, size, form, and texture or striation of the surface, even in specimens collected in the same locality. Thus, a French author has enumerated no less than 198 varieties of the common wood-snail (Helix nemoralis), while of t]]he equally common garden-snail (Helix hortensis) ninety varieties have been described. Fresh-water shells are also subject to great variation, so that there is much uncertainty as to the number of species; and variations are especially frequent in the Planorbidae, which exhibit many eccentric deviations from the usual form of the species—deviations which must often affect the form of the living animal. In Mr. Ingersoll's Report on the Recent Mollusca of Colorado many of these extraordinary variations are referred to, and it is stated that a shell (Helisonia trivolvis) abundant in some small ponds and lakes, had scarcely two specimens alike, and many of them closely resembled other and altogether distinct species.[2]

The Variability of Insects.

Among Insects there is a large amount of variation, though very few entomologists devote themselves to its investigation. Our first examples will be taken from the late Mr. T. Vernon Wollaston's book, On the Variation of Species, and they must be considered as indications of very widespread though little noticed phenomena. He speaks of the curious little carabideous beetles of the genus Notiophilus as being "extremely unstable both in their sculpture and hue;" of the common Calathus mollis as having "the hind wings at one time ample, at another rudimentary, and at a third nearly obsolete;" and of the same irregularity as to the wings being characteristic of many Orthoptera and of the Homopterous Fulgoridae. Mr. Westwood in his Modern Classification of Insects states that "the species of Gerris, Hydrometra, and Velia are mostly found perfectly apterous, though occasionally with full-sized wings."

It is, however, among the Lepidoptera (butterflies and moths) that the most numerous cases of variation have been observed, and every good collection of these insects affords striking examples. I will first adduce the testimony of Mr. Bates, who speaks of the butterflies of the Amazon valley exhibiting innumerable local varieties or races, while some species showed great individual variability. Of the beautiful Mechanitis Polymnia he says, that at Ega on the Upper Amazons, "it varies not only in general colour and pattern, but also very considerably in the shape of the wings, especially in the male sex." Again, at St. Paulo, Ithomia Orolina exhibits four distinct varieties, all occurring together, and these differ not only in colour but in form, one variety being described as having the fore wings much elongated in the male, while another is much larger and has "the hind wings in the male different in shape." Of Heliconius Numata Mr. Bates says: "This species is so variable that it is difficult to find two examples exactly alike," while "it varies in structure as well as in colours. The wings are sometimes broader, sometimes narrower; and their edges are simple in some examples and festooned in others." Of another species of the same genus, H. melpomene, ten distinct varieties are described all more or less connected by intermediate forms, and four of these varieties were obtained at one locality, Serpa on the north bank of the Amazon. Ceratina Ninonia is another of these very unstable species exhibiting many local varieties which are, however, incomplete and connected by intermediate forms; while the several species of the genus Lycorea all vary to such an extent as almost to link them together, so that Mr. Bates thinks they might all fairly be considered as varieties of one species only.

Turning to the Eastern Hemisphere we have in Papilio Severus a species which exhibits a large amount of simple variation, in the presence or absence of a pale patch on the upper wings, in the brown submarginal marks on the lower wings, in the form and extent of the yellow band, and in the size of the specimens. The most extreme forms, as well as the intermediate ones, are often found in one locality and in company with each other. A small butterfly (Terias hecabe) ranges over the whole of the Indian and Malayan regions to Australia, and everywhere exhibits great variations, many of which have been described as distinct species; but a gentleman in Australia bred two of these distinct forms (T. hecabe and T. Aesiope), with several intermediates, from one batch of caterpillars found feeding together on the same plant.[3] It is therefore very probable that a considerable number of supposed distinct species are only individual varieties.

Cases of variation similar to those now adduced among butterflies might be increased indefinitely, but it is as well to note that such important characters as the neuration of the wings, on which generic and family distinctions are often established, are also subject to variation. The Rev. R.P. Murray, in 1872, laid before the Entomological Society examples of such variation in six species of butterflies, and other cases have been since described. The larvae of butterflies and moths are also very variable, and one observer recorded in the Proceedings of the Entomological Society for 1870 no less than sixteen varieties of the caterpillar of the bedstraw hawk-moth (Deilephela galii).

Variation among Lizards.

Passing on from the lower animals to the vertebrata, we find more abundant and more definite evidence as to the extent and amount of individual variation. I will first give a case among the Reptilia from some of Mr. Darwin's unpublished MSS., which have been kindly lent me by Mr. Francis Darwin.

"M. Milne Edwards (Annales des Sci. Nat., I ser., tom. xvi. p. 50) has given a curious table of measurements of fourteen specimens of Lacerta muralis; and, taking the length of the head as a standard, he finds the neck, trunk, tail, front and hind legs, colour, and femoral pores, all varying wonderfully; and so it is more or less with other species. So apparently trifling a character as the scales on the head affording almost the only constant characters."

As the table of measurements above referred to would give no clear conception of the nature and amount of the variation without a laborious study and comparison of the figures, I have endeavoured to find a method of presenting the facts to the eye, so that they may be easily grasped and appreciated. In the diagram opposite, the comparative variations of the different organs of this species are given by means of variously bent lines. The head is represented by a straight line because it presented (apparently) no variation. The body is next given, the specimens being arranged in the order of their size from No. 1, the smallest, to No. 14, the largest, the actual lengths being laid down from a base line at a suitable distance below, in this case two inches below the centre, the mean length of the body of the fourteen specimens being two inches. The respective lengths of the neck, legs, and toe of
Fig 1 - Variations of Lacerta muralis
Fig 2 - Variations of Lizards
each specimen are then laid down in the same manner at convenient distances apart for comparison; and we see that their variations bear no definite relation to those of the body, and not much to those of each other. With the exception of No. 5, in which all the parts agree in being large, there is a marked independence of each part, shown by the lines often curving in opposite directions; which proves that in those specimens one part is large while the other is small. The actual amount of the variation is very great, ranging from one-sixth of the mean length in the neck to considerably more than a fourth in the hind leg, and this among only fourteen examples which happen to be in a particular museum.

To prove that this is not an isolated case, Professor Milne Edwards also gives a table showing the amount of variation in the museum specimens of six common species of lizards, also taking the head as the standard, so that the comparative variation of each part to the head is given. In the accompanying diagram (Fig. 2) the variations are exhibited by means of lines of varying length. It will be understood that, however much the specimens varied in size, if they had kept the same proportions, the variation line would have been in every case reduced to a point, as in the neck of L. velox which exhibits no variation. The different proportions of the variation lines for each species may show a distinct mode of variation, or may be merely due to the small and differing number of specimens; for it is certain that whatever amount of variation occurs among a few specimens will be greatly increased when a much larger number of specimens are examined. That the amount of variation is large, may be seen by comparing it with the actual length of the head (given below the diagram) which was used as a standard in determining the variation, but which itself seems not to have varied.[4]

Variation among Birds.

Coming now to the class of Birds, we find much more copious evidence of variation. This is due partly to the fact that Ornithology has perhaps a larger body of devotees than any other branch of natural history (except entomology); to the moderate size of the majority of birds; and to the circumstance that the form and dimensions of the wings, tail, beak, and feet offer the best generic and specific characters and can all be easily measured and compared. The most systematic observations on the individual variation of birds have been made by Mr. J.A. Allen, in his remarkable memoir: "On the Mammals and Winter Birds of East Florida, with an examination of certain assumed specific characters in Birds, and a sketch of the Bird Faunae of Eastern North America," published in the Bulletin of the Museum of Comparative Zoology at Harvard College, Cambridge, Massachusetts, in 1871. In this work exact measurements are given of all the chief external parts of a large number of species of common American birds, from twenty to sixty or more specimens of each species being measured, so that we are able to determine with some precision the nature and extent of the variation that usually occurs. Mr. Allen says: "The facts of the case show that a variation of from 15 to 20 per cent in general size, and an equal degree of variation in the relative size of different parts, may be ordinarily expected among specimens of the same species and sex, taken at the same locality, while in some cases the variation is even greater than this." He then goes on to show that each part varies to a considerable extent independently of the other parts; so that when the size varies, the proportions of all the parts vary, often to a much greater amount. The wing and tail, for example, besides varying in length, vary in the proportionate length of each feather, and this causes their outline to vary considerably in shape. The bill also varies in length, width, depth, and curvature. The tarsus varies in length, as does each toe separately and independently; and all this not to a minute degree requiring very careful measurement to detect it at all, but to an amount easily seen without any measurement, as it averages one-sixth of the whole length and often reaches one-fourth. In twelve species of common perching birds the wing varied (in from twenty-five to thirty specimens) from 14 to 21 per cent of the mean length, and the tail from 13.8 to 23.4 per cent. The variation of the form of the wing can be very easily tested by noting which feather is longest, which next in length, and so on, the respective feathers being indicated by the numbers 1, 2, 3, etc., commencing with the outer one. As an example of the irregular variation constantly met with, the following occurred among twenty-five specimens of Dendraeca coronata. Numbers bracketed imply that the corresponding feathers were of equal length.[5]

RELATIVE LENGTHS OF PRIMARY WING FEATHERS OF

DENDRAECA CORONATA.

Longest. Second in Third in Fourth in Fifth in Sixth in
Length. Length. Length. Length. Length.
2 3 1 4 5 6
3 2 4 1 5 6
3 2 / 4 1 5 6 7
2 /3 4 1 5 6 7
2 / 1 / 3 / 4 5 6 7 8 9

Here we have five very distinct proportionate lengths of the wing feathers, any one of which is often thought sufficient to characterise a distinct species of bird; and though this is rather an extreme case, Mr. Allen assures us that "the comparison, extended in the table to only a few species, has been carried to scores of others with similar results."

Along with this variation in size and proportions there occurs a large amount of variation in colour and markings. "The difference in intensity of colour between the extremes of a series of fifty or one hundred specimens of any species, collected at a single locality, and nearly at the same season of the year, is often as great as occurs between truly distinct species." But there is also a great amount of individual variability in the markings of the same species. Birds having the plumage varied with streaks and spots differ exceedingly in different individuals of the same species in respect to the size, shape, and number of these marks, and in the general aspect of the plumage resulting from such variations. "In the common song sparrow (Melospiza melodia), the fox-coloured sparrow (Passerella iliaca), the swamp sparrow (Melospiza palustris), the black and white creeper (Mniotilta varia), the water-wagtail (Seiurus novaeboracencis), in Turdus fuscescens and its allies, the difference in the size of the streaks is often very considerable. In the song sparrow they vary to such an extent that in some cases they are reduced to narrow lines; in others so enlarged as to cover the greater part of the breast and sides of the body, sometimes uniting on the middle of the breast into a nearly continuous patch."

Mr. Allen then goes on to particularise several species in which such variations occur, giving cases in which two specimens taken at the same place on the same day exhibited the two extremes of coloration. Another set of variations is thus described: "The white markings so common on the wings and tails of birds, as the bars formed by the white tips of the greater wing-coverts, the white patch occasionally present at the base of the primary quills, or the white band crossing them, and the white patch near the end of the outer tail-feathers are also extremely liable to variation in respect to their extent and the number of feathers to which, in the same species, these markings extend." It is to be especially noted that all these varieties are distinct from those which depend on season, on age, or on sex, and that they are such as have in many other species been considered to be of specific value.

These variations of colour could not be presented to the eye without a series of carefully engraved plates, but in order to bring Mr. Allen's measurements, illustrating variations of size and proportion, more clearly before the reader, I have prepared a series of diagrams illustrating the more important facts and their bearings on the Darwinian theory.

The first of these is intended, mainly, to show the actual amount of the variation, as it gives the true length of the wing and tail in the extreme cases among thirty specimens of each of three species. The shaded portion shows the minimum length, the unshaded portion the additional length in the maximum. The point to be specially noted here is, that in each of these common species there is about the same amount of variation, and that it is so great as to be obvious at a glance.
Fig 3 - Variations of Wings and Tail.
There is here no question of "minute" or "infinitesimal" variation, which many people suppose to be the only kind of variation that exists. It cannot even be called small; yet from all the evidence we now possess it seems to be the amount which characterises most of the common species of birds.

It may be said, however, that these are the extreme variations, and only occur in one or two individuals, while the great majority exhibit little or no difference. Other diagrams will show that this is not the case; but even if it were so, it would be no objection at all, because these are the extremes among thirty specimens only. We may safely assume that these thirty specimens, taken by chance, are not, in the case of all these species, exceptional lots, and therefore we might expect at least two similarly varying specimens in each additional thirty. But the number of individuals, even in a very rare species, is probably thirty thousand or more, and in a common species thirty, or even three hundred, millions. Even one individual in each thirty, varying to the amount shown in the diagram, would give at least a million in the total population of any common bird, and among this million many would vary much more than the extreme among thirty only. We should thus have a vast body of individuals varying to a large extent in the length of the wings and tail, and offering ample material for the modification of these organs by natural selection. We will now proceed to show that other parts of the body vary, simultaneously, but independently, to an equal amount.

The first bird taken is the common Bob-o-link or Rice-bird (Dolichonyx oryzivorus), and the Diagram, Fig. 4, exhibits the variations of seven important characters in twenty male adult specimens.[6] These characters are—the lengths of the body, wing, tail, tarsus, middle toe, outer toe, and hind toe, being as many as can be conveniently exhibited in one diagram. The length of the body is not given by Mr. Allen, but as it forms a convenient standard of comparison, it has been obtained by deducting the length of the tail from the total length of the birds as given by him. The diagram has been constructed as follows:—The twenty specimens are first arranged in a series according to the body-lengths (which may be
Fig 4 - Dolichonyx oryzivorus. 20 Males.
Fig 5 - Alelæus phœniccus. 40 Males.
considered to give the size of the bird), from the shortest to the longest, and the same number of vertical lines are drawn, numbered from one to twenty. In this case (and wherever practicable) the body-length is measured from the lower line of the diagram, so that the actual length of the bird is exhibited as well as the actual variations of length. These can be well estimated by means of the horizontal line drawn at the mean between the two extremes, and it will be seen that one-fifth of the total number of specimens taken on either side exhibits a very large amount of variation, which would of course be very much greater if a hundred or more specimens were compared. The lengths of the wing, tail, and other parts are then laid down, and the diagram thus exhibits at a glance the comparative variation of these parts in every specimen as well as the actual amount of variation in the twenty specimens; and we are thus enabled to arrive at some important conclusions.

We note, first, that the variations of none of the parts follow the variations of the body, but are sometimes almost in an opposite direction. Thus the longest wing corresponds to a rather small body, the longest tail to a medium body, while the longest leg and toes belong to only a moderately large body. Again, even related parts do not constantly vary together but present many instances of independent variation, as shown by the want of parallelism in their respective variation-lines. In No. 5 (see Fig. 4) the wing is very long, the tail moderately so; while in No. 6 the wing is much shorter while the tail is considerably longer. The tarsus presents comparatively little variation; and although the three toes may be said to vary in general together, there are many divergencies; thus, in passing from No. 9 to No. 10, the outer toe becomes longer, while the hind toe becomes considerably shorter; while in Nos. 3 and 4 the middle toe varies in an opposite way to the outer and the hind toes.

In the next diagram (Fig. 5) we have the variations in forty males of the Red-winged Blackbird (Agelaeus phoeniceus), and here we see the same general features. One-fifth of the whole number of specimens offer a large amount of variation either below or above the mean; while the wings, tail, and head vary quite independently of the body. The wing and tail too,
Fig 6 - Cardinalis virginianus. 31 Males.
though showing some amount of correlated variation, yet in no less than nine cases vary in opposite directions as compared with the preceding species.

The next diagram (Fig. 6), showing the variations of thirty-one males of the Cardinal bird (Cardinalis virginianus), exhibits these features much more strongly. The amount of variation in proportion to the size of the bird is very much greater; while the variations of the wing and tail not only have no correspondence with that of the body but very little with each other. In no less than twelve or thirteen instances they vary in opposite directions, while even where they correspond in direction the amount of the variation is often very disproportionate.

As the proportions of the tarsi and toes of birds have great influence on their mode of life and habits and are often used as specific or even generic characters, I have prepared a diagram (Fig. 7) to show the variation in these parts only, among twenty specimens of each of four species of birds, four or five of the most variable alone being given. The extreme divergence of each of the lines in a vertical direction shows the actual amount of variation; and if we consider the small length of the toes of these small birds, averaging about three-quarters of an inch, we shall see that the variation is really very large; while the diverging curves and angles show that each part varies, to a great extent, independently. It is evident that if we compared some thousands of individuals instead of only twenty, we should have an amount of independent variation occurring each year which would enable almost any modification of these important organs to be rapidly effected.

In order to meet the objection that the large amount of variability here shown depends chiefly on the observations of one person and on the birds of a single country, I have examined Professor Schlegel's Catalogue of the Birds in the Leyden Museum, in which he usually gives the range of variation of the specimens in the museum (which are commonly less than a dozen and rarely over twenty) as regards some of their more important dimensions. These fully support the statement of Mr. Allen, since they show an equal amount of variability when the numbers compared are
Fig 7 - Variation of Tarsus and Toes.
Fig 8 - Birds in Leyden Museum
sufficient, which, however, is not often the case. The accompanying diagram exhibits the actual differences of size in five organs which occur in five species taken almost at random from this catalogue. Here, again, we perceive that the variation is decidedly large, even among a very small number of specimens; while the facts all show that there is no ground whatever for the common assumption that natural species consist of individuals which are nearly all alike, or that the variations which occur are "infinitesimal" or even "small."

The proportionate Number of Individuals which present a considerable amount of Variation.

The notion that variation is a comparatively exceptional phenomenon, and that in any case considerable variations occur very rarely in proportion to the number of individuals which do not vary, is so deeply rooted that it is necessary to show by every possible method of illustration how completely opposed it is to the facts of nature. I have therefore prepared some diagrams in which each of the individual birds measured is represented by a spot, placed at a proportionate distance, right and left, from the median line accordingly as it varies in excess or defect of the mean length as regards the particular part compared. As the object in this set of diagrams is to show the number of individuals which vary considerably in proportion to those which vary little or not at all, the scale has been enlarged in order to allow room for placing the spots without overlapping each other.

In the diagram opposite twenty males of Icterus Baltimore are registered, so as to exhibit to the eye the proportionate number of specimens which vary, to a greater or less amount, in the length of the tail, wing, tarsus, middle toe, hind toe, and bill. It will be noticed that there is usually no very great accumulation of dots about the median line which shows the average dimensions, but that a considerable number are spread at varying distances on each side of it.

In the next diagram (Fig. 10), showing the variation among forty males of Agelaeus phoeniceus, this approach to an equable spreading of the variations is still more apparent; while in Fig. 12, where fifty-eight specimens of Cardinalis virginianus are registered, we see a remarkable spreading out of the spots, showing in some of the characters a tendency to segregation into two or more groups of individuals, each varying considerably from the mean.

Fig 9.
In order fully to appreciate the teaching of these diagrams, we must remember, that, whatever kind and amount of variations are exhibited by the few specimens here compared, would be greatly extended and brought into symmetrical form if large numbers—thousands or millions—were subjected to the same process of measurement and registration. We know, from the general law which governs variations from a mean value, that with increasing numbers the range
Fig 10.
Fig 11.

of variation of each part would increase also, at first rather rapidly and then more slowly; while gaps and irregularities would be gradually filled up, and at length the distribution of the dots would indicate a tolerably regular curve of double curvature like those shown in Fig. 11. The great divergence of the dots, when even a few specimens are compared, shows that the curve, with high numbers, would be a flat one like the lower curve in the illustration here given. This being the case it would follow that a very large proportion of the total number of individuals constituting a species would diverge considerably from its average condition as regards each part or organ; and as we know from the previous diagrams of variation (Figs. 1 to 7) that each part varies to a considerable extent, independently, the materials constantly ready for natural selection to act upon are abundant in quantity and very varied in kind. Almost any combination of variations of distinct parts will be available, where required; and this, as we shall see further on, obviates one of the most weighty objections which have been urged against the efficiency of natural selection in producing new species, genera, and higher groups.

Fig 12.

Variation in the Mammalia.

Owing to the generally large size of this class of animals, and the comparatively small number of naturalists who study them, large series of specimens are only occasionally examined and compared, and thus the materials for determining the question of their variability in a state of nature are comparatively scanty. The fact that our domestic animals belonging to this group, especially dogs, present extreme varieties not surpassed even by pigeons and poultry among birds, renders it almost certain that an equal amount of variability exists in the wild state; and this is confirmed by the example of a species of squirrel (Sciurus carolinensis), of which sixteen specimens, all males and all taken in Florida, were measured and tabulated by Mr. Allen. The diagram here given shows, that, both the general amount of the variation and the independent variability of the several members of the body, accord completely with the variations so common in the class of birds; while their amount and their independence of each other are even greater than usual.

Variation in the Internal Organs of Animals.

In case it should be objected that the cases of variation hitherto adduced are in the external parts only, and that there is no proof that the internal organs vary in the same manner, it will be advisable to show that such varieties also occur. It is, however, impossible to adduce the same amount of evidence in this class of variation, because the great labour of dissecting large numbers of specimens of the same species is rarely undertaken, and we have to trust to the chance observations of anatomists recorded in their regular course of study.

It must, however, be noted that a very large proportion of the variations already recorded in the external parts of animals necessarily imply corresponding internal variations. When feet and legs vary in size, it is because the bones vary; when the head, body, limbs, and tail change their proportions, the bony skeleton must also change; and even when the wing or tail feathers of birds become longer or more numerous, there is sure to be a corresponding change in the bones which support and the muscles which move them. I will, however, give a few cases of variations which have been directly observed.

Mr. Frank E. Beddard has kindly communicated to me some remarkable variations he has observed in the internal
Fig 13. - Sciurus carolinensis. 32 specimens. Florida.
organs of a species of earthworm (Perionyx excavatus). The normal characters of this species are—
Setae forming a complete row round each segment.
Two pairs of spermathecae—spherical pouches without diverticulae—in segments 8 and 9.
Two pairs of testes in segments 11 and 12.
Ovaries, a single pair in segment 13.
Oviducts open by a common pore in the middle of segment 14.
Vasa deferentia open separately in segment 18, each furnished at its termination with a large prostate gland.

Between two and three hundred specimens were examined, and among them thirteen specimens exhibited the following marked variations:—

(1) The number of the spermathecae varied from two to three or four pairs, their position also varying.
(2) There were occasionally two pairs of ovaries, each with its own oviduct; the external apertures of these varied in position, being upon segments 13 and 14, 14 and 15, or 15 and 16. Occasionally when there was only the normal single oviduct pore present it varied in position, once occurring on the 10th, and once on the 11th segment.
(3) The male generative pores varied in position from segments 14 to 20. In one instance there were two pairs instead of the normal single pair, and in this case each of the four apertures had its own prostate gland.

Mr. Beddard remarks that all, or nearly all, the above variations are found normally in other genera and species.

When we consider the enormous number of earthworms and the comparatively very small number of individuals examined, we may be sure, not only that such variations as these occur with considerable frequency, but also that still more extraordinary deviations from the normal structure may often exist.

The next example is taken from Mr. Darwin's unpublished MSS. "In some species of Shrews (Sorex) and in some field-mice (Arvicola), the Rev. L. Jenyns (Ann. Nat. Hist., vol. vii. pp. 267, 272) found the proportional length of the intestinal canal to vary considerably. He found the same variability in the number of the caudal vertebrae. In three specimens of an Arvicola he found the gall-bladder having a very different degree of development, and there is reason to believe it is sometimes absent. Professor Owen has shown that this is the case with the gall-bladder of the giraffe."

Dr. Crisp (Proc. Zool. Soc., 1862, p. 137) found the gall-bladder present in some specimens of Cervus superciliaris while absent in others; and he found it to be absent in three giraffes which he dissected. A double gall-bladder was found in a sheep, and in a small mammal preserved in the Hunterian Museum there are three distinct gall-bladders.

The length of the alimentary canal varies greatly. In three adult giraffes described by Professor Owen it was from 124 to 136 feet long; one dissected in France had this canal 211 feet long; while Dr. Crisp measured one of the extraordinary length of 254 feet, and similar variations are recorded in other animals.[7]

The number of ribs varies in many animals. Mr. St. George Mivart says: "In the highest forms of the Primates, the number of true ribs is seven, but in Hylobates there are sometimes eight pairs. In Semnopithecus and Colobus there are generally seven, but sometimes eight pairs of true ribs. In the Cebidae there are generally seven or eight pairs, but in Ateles sometimes nine" (Proc. Zool. Soc., 1865, p. 568). In the same paper it is stated that the number of dorsal vertebrae in man is normally twelve, very rarely thirteen. In the Chimpanzee there are normally thirteen dorsal vertebrae, but occasionally there are fourteen or only twelve.

Variations in the Skull.

Among the nine adult male Orang-utans, collected by myself in Borneo, the skulls differed remarkably in size and proportions. The orbits varied in width and height, the cranial ridge was either single or double, either much or little developed, and the zygomatic aperture varied considerably in
Fig 14. Variation of Skull of Wolf. 10 specimens.
 size. I noted particularly that these variations bore no necessary relation to each other, so that a large temporal muscle and zygomatic aperture might exist either with a large or a small cranium; and thus was explained the curious difference between the single-crested and the double-crested skulls, which had been supposed to characterise distinct species. As an instance of the amount of variation in the skulls of fully adult male orangs, I found the width between the orbits externally to be only 4 inches in one specimen and fully 5 inches in another.

Exact measurements of large series of comparable skulls of the mammalia are not easily found, but from those available I have prepared three diagrams (Figs. 14, 15, and 16), in order to exhibit the facts of variation in this very important organ. The first shows the variation in ten specimens of the common wolf (Canis lupus) from one district in North America, and we see that it is not only large in amount, but that each part exhibits a considerable independent variability.[8]

In Diagram 15 we have the variations of eight skulls of the Indian Honey-bear (Ursus labiatus), as tabulated by the late Dr. J. E. Gray of the British Museum. For such a small number of specimens the amount of variation is very large—from one-eighth to one-fifth of the mean size,—while there are an extraordinary number of instances of independent variability. In Diagram 16 we have the length and width of twelve skulls of adult males of the Indian wild boar (Sus cristatus), also given by Dr. Gray, exhibiting in both sets of measurements a variation of more than one-sixth, combined with a very considerable amount of independent variability.[9]

The few facts now given, as to variations of the internal parts of animals, might be multiplied indefinitely by a search through the voluminous writings of comparative anatomists. But the evidence already adduced, taken in conjunction with the much fuller evidence of variation in all external organs, leads us to the conclusion that wherever variations are looked for among a considerable number of individuals of the more
Fig 15. Variation of 8 skulls (Ursus labiatus).
Fig 16.
common species they are sure to be found; that they are everywhere of considerable amount, often reaching 20 per cent of the size of the part implicated; and that they are to a great extent independent of each other, and thus afford almost any combination of variations that may be needed.

It must be particularly noticed that the whole series of variation-diagrams here given (except the three which illustrate the number of varying individuals) in every case represent the actual amount of the variation, not on any reduced or enlarged scale, but as it were life-size. Whatever number of inches or decimals of an inch the species varies in any of its parts is marked on the diagrams, so that with the help of an ordinary divided rule or a pair of compasses the variation of the different parts can be ascertained and compared just as if the specimens themselves were before the reader, but with much greater ease.

In my lectures on the Darwinian theory in America and in this country I used diagrams constructed on a different plan, equally illustrating the large amount of independent variability, but less simple and less intelligible. The present method is a modification of that used by Mr. Francis Galton in his researches on the theory of variability, the upper line (showing the variability of the body) in Diagrams 4, 5, 6, and 13, being laid down on the method he has used in his experiments with sweet-peas and in pedigree moth-breeding.[10] I believe, after much consideration, and many tedious experiments in diagram-making, that no better method can be adopted for bringing before the eye, both the amount and the peculiar features of individual variability.

Variations of the Habits of Animals.

Closely connected with those variations of internal and external structure which have been already described, are the changes of habits which often occur in certain individuals or in whole species, since these must necessarily depend upon some corresponding change in the brain or in other parts of the organism; and as these changes are of great importance in relation to the theory of instinct, a few examples of them will be now adduced. The Kea (Nestor notabilis) is a curious parrot inhabiting the mountain ranges of the Middle Island of New Zealand. It belongs to the family of Brush-tongued parrots, and naturally feeds on the honey of flowers and the insects which frequent them, together with such fruits or berries as are found in the region. Till quite recently this comprised its whole diet, but since the country it inhabits has become occupied by Europeans it has developed a taste for a carnivorous diet, with alarming results. It began by picking the sheepskins hung out to dry or the meat in process of being cured. About 1868 it was first observed to attack living sheep, which had frequently been found with raw and bleeding wounds on their backs. Since then it is stated that the bird actually burrows into the living sheep, eating its way down to the kidneys, which form its special delicacy. As a natural consequence, the bird is being destroyed as rapidly as possible, and one of the rare and curious members of the New Zealand fauna will no doubt shortly cease to exist. The case affords a remarkable instance of how the climbing feet and powerful hooked beak developed for one set of purposes can be applied to another altogether different purpose, and it also shows how little real stability there may be in what appear to us the most fixed habits of life. A somewhat similar change of diet has been recorded by the Duke of Argyll, in which a goose, reared by a golden eagle, was taught by its foster-parent to eat flesh, which it continued to do regularly and apparently with great relish.[11]

Change of habits appears to be often a result of imitation, of which Mr. Tegetmeier gives some good examples. He states that if pigeons are reared exclusively with small grain, as wheat or barley, they will starve before eating beans. But when they are thus starving, if a bean-eating pigeon is put among them, they follow its example, and thereafter adopt the habit. So fowls sometimes refuse to eat maize, but on seeing others eat it, they do the same and become excessively fond of it. Many persons have found that their yellow crocuses were eaten by sparrows, while the blue, purple, and white coloured varieties were left untouched; but Mr. Tegetmeier, who grows only these latter colours, found that after two years the sparrows began to attack them, and thereafter destroyed them quite as readily as the yellow ones; and he believes it was merely because some bolder sparrow than the rest set the example. On this subject Mr. Charles C. Abbott well remarks: "In studying the habits of our American birds—and I suppose it is true of birds everywhere—it must at all times be remembered that there is less stability in the habits of birds than is usually supposed; and no account of the habits of any one species will exactly detail the various features of its habits as they really are, in every portion of the territory it inhabits."[12]

Mr. Charles Dixon has recorded a remarkable change in the mode of nest-building of some common chaffinches which were taken to New Zealand and turned out there. He says: "The cup of the nest is small, loosely put together, apparently lined with feathers, and the walls of the structure are prolonged for about 18 inches, and hang loosely down the side of the supporting branch. The whole structure bears some resemblance to the nests of the hangnests (Icteridae), with the exception that the cavity is at the top. Clearly these New Zealand chaffinches were at a loss for a design when fabricating their nest. They had no standard to work by, no nests of their own kind to copy, no older birds to give them any instruction, and the result is the abnormal structure I have just described."[13]

These few examples are sufficient to show that both the habits and instincts of animals are subject to variation; and had we a sufficient number of detailed observations we should probably find that these variations were as numerous, as diverse in character, as large in amount, and as independent of each other as those which we have seen to characterise their bodily structure.

The Variability of Plants.

The variability of plants is notorious, being proved not only by the endless variations which occur whenever a species is largely grown by horticulturists, but also by the great difficulty that is felt by botanists in determining the limits of species in many large genera. As examples we may take the roses, the brambles, and the willows as well illustrating this fact. In Mr. Baker's Revision of the British Roses (published by the Linnean Society in 1863), he includes under the single species, Rosa canina—the common dog-rose—no less than twenty-eight named varieties distinguished by more or less constant characters and often confined to special localities, and to these are referred about seventy of the species of British and continental botanists. Of the genus Rubus or bramble, five British species are given in Bentham's Handbook of the British Flora, while in the fifth edition of Babington's Manual of British Botany, published about the same time, no less than forty-five species are described. Of willows (Salix) the same two works enumerate fifteen and thirty-one species respectively. The hawkweeds (Hieracium) are equally puzzling, for while Mr. Bentham admits only seven British species, Professor Babington describes no less than thirty-two, besides several named varieties.

A French botanist, Mons. A. Jordan, has collected numerous forms of a common little plant, the spring whitlow-grass (Draba verna); he has cultivated these for several successive years, and declares that they preserve their peculiarities unchanged; he also says that they each come true from seed, and thus possess all the characteristics of true species. He has described no less than fifty-two such species or permanent varieties, all found in the south of France; and he urges botanists to follow his example in collecting, describing, and cultivating all such varieties as may occur in their respective districts. Now, as the plant is very common almost all over Europe and ranges from North America to the Himalayas, the number of similar forms over this wide area would probably have to be reckoned by hundreds if not by thousands.

The class of facts now adduced must certainly be held to prove that in many large genera and in some single species there is a very large amount of variation, which renders it quite impossible for experts to agree upon the limits of species. We will now adduce a few striking cases of individual variation.

The distinguished botanist, Alp. de Candolle, made a special study of the oaks of the whole world, and has stated some remarkable facts as to their variability. He declares that on the same branch of oak he has noted the following variations: (1) In the length of the petiole, as one to three; (2) in the form of the leaf, being either elliptical or obovoid; (3) in the margin being entire, or notched, or even pinnatifid; (4) in the extremity being acute or blunt; (5) in the base being sharp, blunt, or cordate; (6) in the surface being pubescent or smooth; (7) the perianth varies in depth and lobing; (8) the stamens vary in number, independently; (9) the anthers are mucronate or blunt; (10) the fruit stalks vary greatly in length, often as one to three; (11) the number of fruits varies; (12) the form of the base of the cup varies; (13) the scales of the cup vary in form; (14) the proportions of the acorns vary; (15) the times of the acorns ripening and falling vary.

Besides this, many species exhibit well-marked varieties which have been described and named, and these are most numerous in the best-known species. Our British oak (Quercus robur) has twenty-eight varieties; Quercus Lusitanica has eleven; Quercus calliprinos has ten; and Quercus coccifera eight.

A most remarkable case of variation in the parts of a common flower has been given by Dr. Hermann Müller. He examined two hundred flowers of Myosurus minimus, among which he found thirty-five different proportions of the sepals, petals, and anthers, the first varying from four to seven, the second from two to five, and the third from two to ten. Five sepals occurred in one hundred and eighty-nine out of the two hundred, but of these one hundred and five had three petals, forty-six had four petals, and twenty-six had five petals; but in each of these sets the anthers varied in number from three to eight, or from two to nine. We have here an example of the same amount of "independent variability" that, as we have seen, occurs in the various dimensions of birds and mammals; and it may be taken as an illustration of the kind and degree of variability that may be expected to occur among small and little specialised flowers.[14]

In the common wind-flower (Anemone nemorosa) an almost equal amount of variation occurs; and I have myself gathered in one locality flowers varying from ⅞ inch to 1¾ inch in diameter; the bracts varying from 1½ inch to 4 inches across; and the petaloid sepals either broad or narrow, and varying in number from five to ten. Though generally pure white on their upper surface, some specimens are a full pink, while others have a decided bluish tinge.

Mr. Darwin states that he carefully examined a large number of plants of Geranium phaeum and G. pyrenaicum (not perhaps truly British but frequently found wild), which had escaped from cultivation, and had spread by seed in an open plantation; and he declares that "the seedlings varied in almost every single character, both in their flowers and foliage, to a degree which I have never seen exceeded; yet they could not have been exposed to any great change of their conditions."[15]

The following examples of variation in important parts of plants were collected by Mr. Darwin and have been copied from his unpublished MSS.:—

"De Candolle (Mem. Soc. Phys. de Genève, tom. ii. part ii. p. 217) states that Papaver bracteatum and P. orientale present indifferently two sepals and four petals, or three sepals and six petals, which is sufficiently rare with other species of the genus."

"In the Primulacae and in the great class to which this family belongs the unilocular ovarium is free, but M. Dubury (Mem. Soc. Phys. de Genève, tom. ii. p. 406) has often found individuals in Cyclamen hederaefolium, in which the base of the ovary was connected for a third part of its length with the inferior part of the calyx."

"M. Aug. St. Hilaire (Sur la Gynobase, Mem. des Mus. d'Hist. Nat., tom. x. p. 134), speaking of some bushes of the Gomphia oleaefolia, which he at first thought formed a quite distinct species, says: 'Voilà donc dans un même individu des loges et un style qui se rattachent tantôt a un axe vertical, et tantôt a un gynobase; donc celui-ci n'est qu'un axe veritable; mais cet axe est deprimé au lieu d'être vertical." He adds (p. 151), 'Does not all this indicate that nature has tried, in a manner, in the family of Rutaceae to produce from a single multilocular ovary, one-styled and symmetrical, several unilocular ovaries, each with its own style.' And he subsequently shows that, in Xanthoxylum monogynum, 'it often happens that on the same plant, on the same panicle, we find flowers with one or with two ovaries;' and that this is an important character is shown by the Rutaceae (to which Xanthoxylum belongs), being placed in a group of natural orders characterised by having a solitary ovary."

"De Candolle has divided the Cruciferae into five sub-orders in accordance with the position of the radicle and cotyledons, yet Mons. T. Gay (Ann. des Scien. Nat., ser. i. tom. vii. p. 389) found in sixteen seeds of Petrocallis Pyrenaica the form of the embryo so uncertain that he could not tell whether it ought to be placed in the sub-orders 'Pleurorhizée' or 'Notorhizée'; so again (p. 400) in Cochlearia saxatilis M. Gay examined twenty-nine embryos, and of these sixteen were vigorously 'pleurorhizées,' nine had characters intermediate between pleuro- and notor- hizées, and four were pure notorhizées."

"M. Raspail asserts (Ann. des Scien. Nat., ser. i. tom. v. p. 440) that a grass (Nostus Borbonicus) is so eminently variable in its floral organisation, that the varieties might serve to make a family with sufficiently numerous genera and tribes—a remark which shows that important organs must be here variable."

Species which vary little.

The preceding statements, as to the great amount of variation occurring in animals and plants, do not prove that all species vary to the same extent, or even vary at all, but, merely, that a considerable number of species in every class, order, and family do so vary. It will have been observed that the examples of great variability have all been taken from common species, or species which have a wide range and are abundant in individuals. Now Mr. Darwin concludes, from an elaborate examination of the floras and faunas of several distinct regions, that common, wide ranging species, as a rule, vary most, while those that are confined to special districts and are therefore comparatively limited in number of individuals vary least. By a similar comparison it is shown that species of large genera vary more than species of small genera. These facts explain, to some extent, why the opinion has been so prevalent that variation is very limited in amount and exceptional in character. For naturalists of the old school, and all mere collectors, were interested in species in proportion to their rarity, and would often have in their collections a larger number of specimens of a rare species than of a species that was very common. Now as these rare species do really vary much less than the common species, and in many cases hardly vary at all, it was very natural that a belief in the fixity of species should prevail. It is not, however, as we shall see presently, the rare, but the common and widespread species which become the parents of new forms, and thus the non-variability of any number of rare or local species offers no difficulty whatever in the way of the theory of evolution.

Concluding Remarks.

We have now shown in some detail, at the risk of being tedious, that individual variability is a general character of all common and widespread species of animals or plants; and, further, that this variability extends, so far as we know, to every part and organ, whether external or internal, as well as to every mental faculty. Yet more important is the fact that each part or organ varies to a considerable extent independently of other parts. Again, we have shown, by abundant evidence, that the variation that occurs is very large in amount—usually reaching 10 or 20, and sometimes even 25 per cent of the average size of the varying part; while not one or two only, but from 5 to 10 per cent of the specimens examined exhibit nearly as large an amount of variation. These facts have been brought clearly before the reader by means of numerous diagrams, drawn to scale and exhibiting the actual variations in inches, so that there can be no possibility of denying either their generality or their amount. The importance of this full exposition of the subject will be seen in future chapters, when we shall frequently have to refer to the facts here set forth, especially when we deal with the various theories of recent writers and the criticisms that have been made of the Darwinian theory.

A full exposition of the facts of variation among wild animals and plants is the more necessary, because comparatively few of them were published in Mr. Darwin's works, while the more important have only been made known since last edition of The Origin of Species was prepared; and it is clear that Mr. Darwin himself did not fully recognise the enormous amount of variability that actually exists. This is indicated by his frequent reference to the extreme slowness of the changes for which variation furnishes the materials, and also by his use of such expressions as the following: "A variety when once formed must again, perhaps after a long interval of time, vary or present individual differences of the same favourable nature as before" (Origin, p. 66). And again, after speaking of changed conditions "affording a better chance of the occurrence of favourable variations," he adds: "Unless such occur natural selection can do nothing" (Origin, p. 64). These expressions are hardly consistent with the fact of the constant and large amount of variation, of every part, in all directions, which evidently occurs in each generation of all the more abundant species, and which must afford an ample supply of favourable variations whenever required; and they have been seized upon and exaggerated by some writers as proofs of the extreme difficulties in the way of the theory. It is to show that such difficulties do not exist, and in the full conviction that an adequate knowledge of the facts of variation affords the only sure foundation for the Darwinian theory of the origin of species, that this chapter has been written.


  1. Foraminifera, preface, p. x.
  2. United States Geological Survey of the Territories, 1874
  3. Proceedings of the Entomological Society of London, 1875, p. vii.
  4. Ann. des Sci. Nat., tom. xvi. p. 50.
  5. See Winter Birds of Florida, p.206, Table F.
  6. See Table I, p. 211, of Allen's Winter Birds of Florida
  7. Proc. Zool. Soc., 1864, p64.
  8. J.A. Allen, on Geographical Variation among North American Mammals, Bull. U.S. Geol. and Geog. Survey, vol. ii. p. 314 (1876).
  9. Proc. Zool. Soc. Lond., 1864, p. 700, and 1868, p. 28.
  10. See Trans. Entomological Society of London,1887, p.24.
  11. Nature, vol xix. p554.
  12. Nature, vol. xvi. p. 163; and vol. xi. 227.
  13. Ibid., vol. xxxi. (1885), p.533.
  14. Nature, vol. xxvi. p. 81.
  15. Animals and Plants under Domestication, vol. ii. p. 258.