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Contributions to the Theory of Natural Selection - A Series of Essays
by Alfred Russel Wallace
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VII.

A THEORY OF BIRDS' NESTS;

SHOWING THE RELATION OF CERTAIN DIFFERENCES OF COLOUR IN FEMALE BIRDS, TO THEIR MODE OF NIDIFICATION.

The habit of forming a more or less elaborate structure for the reception of their eggs and young, must undoubtedly be looked upon as one of the most remarkable and interesting characteristics of the class of birds. In other classes of vertebrate animals, such structures are few and exceptional, and never attain to the same degree of completeness and beauty. Birds' nests have, accordingly, attracted much attention, and have furnished one of the stock arguments to prove the existence of a blind but unerring instinct in the lower animals. The very general belief that every bird is enabled to build its nest, not by the ordinary faculties of observation, memory, and imitation, but by means of some innate and mysterious impulse, has had the bad effect of withdrawing attention from the very evident relation that exists between the structure, habits, and intelligence of birds, and the kind of nests they construct.

In the preceding essay I have detailed several of these relations, and they teach us, that a consideration of the structure, the food, and other specialities of a bird's existence, will give a clue, and sometimes a very complete one, to the reason why it builds its nest of certain materials, in a definite situation, and in a more or less elaborate manner.

I now propose to consider the question from a more general point of view, and to discuss its application to some important problems in the natural history of birds.

Changed Conditions and persistent Habits as influencing Nidification.

Besides the causes above alluded to, there are two other factors whose effect in any particular case we can only vaguely guess at, but which must have had an important influence in determining the existing details of nidification. These are—changed conditions of existence, whether internal or external, and the influence of hereditary or imitative habit; the first inducing alterations in accordance with changes of organic structure, of climate, or of the surrounding fauna and flora; the other preserving the peculiarities so produced, even when changed conditions render them no longer necessary. Many facts have been already given which show that birds do adapt their nests to the situations in which they place them, and the adoption of eaves, chimneys, and boxes, by swallows, wrens, and many other birds, shows that they are always ready to take advantage of changed conditions. It is probable, therefore, that a permanent change of climate would cause many birds to modify the form or materials of their abodes, so as better to protect their young. The introduction of new enemies to eggs or young birds, might introduce many alterations tending to their better concealment. A change in the vegetation of a country, would often necessitate the use of new materials. So, also, we may be sure, that as a species slowly became modified in any external or internal characters, it would necessarily change in some degree its mode of building. This effect would be produced by modifications of the most varied nature; such as the power and rapidity of flight, which must often determine the distance to which a bird will go to obtain materials for its nest; the capacity of sustaining itself almost motionless in the air, which must sometimes determine the position in which a nest can be built; the strength and grasping power of the foot in relation to the weight of the bird, a power absolutely essential to the constructor of a delicately-woven and well-finished nest; the length and fineness of the beak, which has to be used like a needle in building the best textile nests; the length and mobility of the neck, which is needful for the same purpose; the possession of a salivary secretion like that used in the nests of many of the swifts and swallows, as well as that of the song-thrush—peculiarities of habits, which ultimately depend on structure, and which often determine the material most frequently met with or most easily to be obtained. Modifications in any of these characters would necessarily lead, either to a change in the materials of the nest, or in the mode of combining them in the finished structure, or in the form or position of that structure.

During all these changes, however, certain specialities of nest-building would continue, for a shorter or a longer time after the causes which had necessitated them had passed away. Such records of a vanished past meet us everywhere, even in man's works, notwithstanding his boasted reason. Not only are the main features of Greek architecture, mere reproductions in stone of what were originally parts of a wooden building, but our modern copyists of Gothic architecture often build solid buttresses capped with weighty pinnacles, to support a wooden roof which has no outward thrust to render them necessary; and even think they ornament their buildings by adding sham spouts of carved stone, while modern waterpipes, stuck on without any attempt at harmony, do the real duty. So, when railways superseded coaches, it was thought necessary to build the first-class carriages to imitate a number of coach-bodies joined together; and the arm-loops for each passenger to hold on by, which were useful when bad roads made every journey a succession of jolts and lurches, were continued on our smooth macadamised mail-routes, and, still more absurdly, remain to this day in our railway carriages, the relic of a kind of locomotion we can now hardly realize. Another good example is to be seen in our boots. When elastic sides came into fashion we had been so long used to fasten them with buttons or laces, that a boot without either looked bare and unfinished, and accordingly the makers often put on a row of useless buttons or imitation laces, because habit rendered the appearance of them necessary to us. It is universally admitted that the habits of children and of savages give us the best clue to the habits and mode of thought of animals; and every one must have observed how children at first imitate the actions of their elders, without any regard to the use or applicability of the particular acts. So, in savages, many customs peculiar to each tribe are handed down from father to son merely by the force of habit, and are continued long after the purpose which they originally served has ceased to exist. With these and a hundred similar facts everywhere around us, we may fairly impute much of what we cannot understand in the details of Bird-Architecture to an analogous cause. If we do not do so, we must assume, either that birds are guided in every action by pure reason to a far greater extent than men are, or that an infallible instinct leads them to the same result by a different road. The first theory has never, that I am aware of, been maintained by any author, and I have already shown that the second, although constantly assumed, has never been proved, and that a large body of facts is entirely opposed to it. One of my critics has, indeed, maintained that I admit "instinct" under the term "hereditary habit;" but the whole course of my argument shows that I do not do so. Hereditary habit is, indeed, the same as instinct when the term is applied to some simple action dependent upon a peculiarity of structure which is hereditary; as when the descendants of tumbler pigeons tumble, and the descendants of pouter pigeons pout. In the present case, however, I compare it strictly to the hereditary, or more properly, persistent or imitative, habits of savages, in building their houses as their fathers did. Imitation is a lower faculty than invention. Children and savages imitate before they originate; birds, as well as all other animals, do the same.

The preceding observations are intended to show, that the exact mode of nidification of each species of bird is probably the result of a variety of causes, which have been continually inducing changes in accordance with changed organic or physical conditions. The most important of these causes seem to be, in the first place, the structure of the species, and, in the second, its environment or conditions of existence. Now we know, that every one of the characters or conditions included under these two heads is variable. We have seen that, on the large scale, the main features of the nest built by each group of birds, bears a relation to the organic structure of that group, and we have, therefore, a right to infer, that as structure varies, the nest will vary also in some particular corresponding to the changes of structure. We have seen also, that birds change the position, the form, and the construction of their nest, whenever the available materials or the available situations, vary naturally or have been altered by man; and we have, therefore, a right to infer that similar changes have taken place, when, by a natural process, external conditions have become in any way permanently altered. We must remember, however, that all these factors are very stable during many generations, and only change at a rate commensurate with those of the great physical features of the earth as revealed to us by geology; and we may, therefore, infer that the form and construction of nests, which we have shown to be dependent on them, are equally stable. If, therefore, we find less important and more easily modified characters than these, so correlated with peculiarities of nidification as to indicate that one is probably the cause of the other, we shall be justified in concluding that these variable characters are dependent on the mode of nidification, and not that the form of the nest has been determined by these variable characters. Such a correlation I am now about to point out.

Classification of Nests.

For the purpose of this inquiry it is necessary to group nests into two great classes, without any regard to their most obvious differences or resemblances, but solely looking to the fact of whether the contents (eggs, young, or sitting bird) are hidden or exposed to view. In the first class we place all those in which the eggs and young are completely hidden, no matter whether this is effected by an elaborate covered structure, or by depositing the eggs in some hollow tree or burrow underground. In the second, we group all in which the eggs, young, and sitting bird are exposed to view, no matter whether there is the most beautifully formed nest, or none at all. Kingfishers, which build almost invariably in holes in banks; Woodpeckers and Parrots, which build in hollow trees; the Icteridae of America, which all make beautiful covered and suspended nests; and our own Wren, which builds a domed nest, are examples of the former; while our Thrushes, Warblers, and Finches, as well as the Crowshrikes, Chatterers, and Tanagers of the tropics, together with all Raptorial birds and Pigeons, and a vast number of others in every part of the world, all adopt the latter mode of building.

It will be seen that this division of birds according to their nidification, bears little relation to the character of the nest itself. It is a functional not a structural classification. The most rude and the most perfect specimens of bird-architecture are to be found in both sections. It has, however, a certain relation to natural affinities, for large groups of birds, undoubtedly allied, fall into one or the other division exclusively. The species of a genus or of a family are rarely divided between the two primary classes, although they are frequently divided between the two very distinct modes of nidification that exist in the first of them.

All the Scansorial or climbing, and most of the Fissirostral or wide-gaped birds, for example, build concealed nests; and, in the latter group, the two families which build open nests, the Swifts and the Goat-suckers, are undoubtedly very widely separated from the other families with which they are associated in our classifications. The Tits vary much in their mode of nesting, some making open nests concealed in a hole, while others build domed or even pendulous covered nests, but they all come under the same class. Starlings vary in a similar way. The talking Mynahs, like our own starlings, build in holes, the glossy starlings of the East (of the genus Calornis) form a hanging covered nest, while the genus Sturnopastor builds in a hollow tree. One of the most striking cases in which one family of birds is divided between the two classes, is that of the Finches; for while most of the European species build exposed nests, many of the Australian finches make them dome-shaped.

Sexual differences of Colour in Birds.

Turning now from the nests to the creatures who make them, let us consider birds themselves from a somewhat unusual point of view, and form them into separate groups, according as both sexes, or the males only, are adorned with conspicuous colours.

The sexual differences of colour and plumage in birds are very remarkable, and have attracted much attention; and, in the case of polygamous birds, have been well explained by Mr. Darwin's principle of sexual selection. We can, to a great extent, understand how male Pheasants and Grouse have acquired their more brilliant plumage and greater size, by the continual rivalry of the males both in strength and beauty; but this theory does not throw any light on the causes which have made the female Toucan, Bee-eater, Parroquet, Macaw and Tit, in almost every case as gay and brilliant as the male, while the gorgeous Chatterers, Manakins, Tanagers, and Birds of Paradise, as well as our own Blackbird, have mates so dull and inconspicuous that they can hardly be recognised as belonging to the same species.

The Law which connects the Colours of Female Birds with the mode of Nidification.

The above-stated anomaly can, however, now be explained by the influence of the mode of nidification, since I find that, with but very few exceptions, it is the rule—that when both sexes are of strikingly gay and conspicuous colours, the nest is of the first class, or such as to conceal the sitting bird; while, whenever there is a striking contrast of colours, the male being gay and conspicuous, the female dull and obscure, the nest is open and the sitting bird exposed to view. I will now proceed to indicate the chief facts that support this statement, and will afterwards explain the manner in which I conceive the relation has been brought about.

We will first consider those groups of birds in which the female is gaily or at least conspicuously coloured, and is in most cases exactly like the male.

1. Kingfishers (Alcedinidae). In some of the most brilliant species of this family the female exactly resembles the male; in others there is a sexual difference, but it rarely tends to make the female less conspicuous. In some, the female has a band across the breast, which is wanting in the male, as in the beautiful Halcyon diops of Ternate. In others the band is rufous in the female, as in several of the American species; while in Dacelo gaudichaudii, and others of the same genus, the tail of the female is rufous, while that of the male is blue. In most kingfishers the nest is in a deep hole in the ground; in Tanysiptera it is said to be in a hole in the nests of termites, or sometimes in crevices under overhanging rocks.

2. Motmots (Momotidae). In these showy birds the sexes are exactly alike, and the nest in a hole under ground.

3. Puff-birds (Bucconidae). These birds are often gaily coloured; some have coral-red bills; the sexes are exactly alike, and the nest is in a hole in sloping ground.

4. Trogons (Trogonidae). In these magnificent birds the females are generally less brightly coloured than the males, but are yet often gay and conspicuous. The nest is in a hole of a tree.

5. Hoopoes (Upupidae). The barred plumage and long crests of these birds render them conspicuous. The sexes are exactly alike, and the nest is in a hollow tree.

6. Hornbills (Bucerotidae). These large birds have enormous coloured bills, which are generally quite as well coloured and conspicuous in the females. Their nests are always in hollow trees, where the female is entirely concealed.

7. Barbets (Capitonidae). These birds are all very gaily-coloured, and, what is remarkable, the most brilliant patches of colour are disposed about the head and neck, and are very conspicuous. The sexes are exactly alike, and the nest is in a hole of a tree.

8. Toucans (Rhamphastidae). These fine birds are coloured in the most conspicuous parts of their body, especially on the large bill, and on the upper and lower tail coverts, which are crimson, white, or yellow. The sexes are exactly alike, and they always build in a hollow tree.

9. Plaintain-eaters (Musophagidae). Here again the head and bill are most brilliantly coloured in both sexes, and the nest is in a hole of a tree.

10. Ground cuckoos (Centropus). These birds are often of conspicuous colours, and are alike in both sexes. They build a domed nest.

11. Woodpeckers (Picidae). In this family the females often differ from the males, in having a yellow or white, instead of a crimson crest, but are almost as conspicuous. They all nest in holes in trees.

12. Parrots (Psittaci). In this great tribe, adorned with the most brilliant and varied colours, the rule is, that the sexes are precisely alike, and this is the case in the most gorgeous families, the lories, the cockatoos, and the macaws; but in some there is a sexual difference of colour to a slight extent. All build in holes, mostly in trees, but sometimes in the ground, or in white ants' nests. In the single case in which the nest is exposed, that of the Australian ground parrot, Pezoporus formosus, the bird has lost the gay colouring of its allies, and is clothed in sombre and completely protective tints of dusky green and black.

13. Gapers (Eurylaemidae). In these beautiful Eastern birds, somewhat allied to the American chatterers, the sexes are exactly alike, and are adorned with the most gay and conspicuous markings. The nest is a woven structure, covered over, and suspended from the extremities of branches over water.

14. Pardalotus (Ampelidae). In these Australian birds the females differ from the males, but are often very conspicuous, having brightly-spotted heads. Their nests are sometimes dome-shaped, sometimes in holes of trees, or in burrows in the ground.

15. Tits (Paridae). These little birds are always pretty, and many (especially among the Indian species) are very conspicuous. They always have the sexes alike, a circumstance very unusual among the smaller gaily-coloured birds of our own country. The nest is always covered over or concealed in a hole.

16. Nuthatches (Sitta). Often very pretty birds, the sexes alike, and the nest in a hole.

17.—— (Sittella). The female of these Australian nuthatches is often the most conspicuous, being white-and black-marked. The nest is, according to Gould, "completely concealed among upright twigs connected together."

18. Creepers (Climacteris). In these Australian creepers the sexes are alike, or the female most conspicuous, and the nest is in a hole of a tree.

19. Estrelda, Amadina. In these genera of Eastern and Australian finches the females, although more or less different from the males, are still very conspicuous having a red rump, or being white spotted. They differ from most others of the family in building domed nests.

20. Certhiola. In these pretty little American creepers the sexes are alike, and they build a domed nest.

21. Mynahs (Sturnidae). These showy Eastern starlings have the sexes exactly alike. They build in holes of trees.

22. Calornis (Sturnidae). These brilliant metallic starlings have no sexual differences. They build a pensile covered nest.

23. Hangnests (Icteridae). The red or yellow and black plumage of most of these birds is very conspicuous, and is exactly alike in both sexes. They are celebrated for their fine purse-shaped pensile nests.

It will be seen that this list comprehends six important families of Fissirostres, four of Scansores, the Psittaci, and several genera, with three entire families of Passeres, comprising about twelve hundred species, or about one-seventh of all known birds.

* * * * *

The cases in which, whenever the male is gaily coloured, the female is much less gay or quite inconspicuous, are exceedingly numerous, comprising, in fact, almost all the bright-coloured Passeres, except those enumerated in the preceding class. The following are the most remarkable:—

1. Chatterers (Cotingidae). These comprise some of the most gorgeous birds in the world, vivid blues, rich purples, and bright reds, being the most characteristic colours. The females are always obscurely tinted, and are often of a greenish hue, not easily visible among the foliage.

2. Manakins (Pipridae). These elegant birds, whose caps or crests are of the most brilliant colours, are usually of a sombre green in the female sex.

3. Tanagers (Tanagridae). These rival the chatterers in the brilliancy of their colours, and are even more varied. The females are generally of plain and sombre hues, and always less conspicuous than the males.

In the extensive families of the warblers (Sylviadae), thrushes (Turdidae), flycatchers (Muscicapidae), and shrikes (Laniadae), a considerable proportion of the species are beautifully marked with gay and conspicuous tints, as is also the case in the Pheasants and Grouse; but in every case the females are less gay, and are most frequently of the very plainest and least conspicuous hues. Now, throughout the whole of these families the nest is open, and I am not aware of a single instance in which any one of these birds builds a domed nest, or places it in a hole of a tree, or underground, or in any place where it is effectually concealed.

In considering the question we are now investigating, it is not necessary to take into account the larger and more powerful birds, because these seldom depend much on concealment to secure their safety. In the raptorial birds bright colours are as a rule absent; and their structure and habits are such as not to require any special protection for the female. The larger waders are sometimes very brightly coloured in both sexes; but they are probably little subject to the attacks of enemies, since the scarlet ibis, the most conspicuous of birds, exists in immense quantities in South America. In game birds and water-fowl, however, the females are often very plainly coloured, when the males are adorned with brilliant hues; and the abnormal family of the Megapodidae offers us the interesting fact of an identity in the colours of the sexes (which in Megacephalon and Talegalla are somewhat conspicuous), in conjunction with the habit of not sitting on the eggs at all.

What the Facts Teach us.

Taking the whole body of evidence here brought forward, embracing as it does almost every group of bright-coloured birds, it will, I think, be admitted that the relation between the two series of facts in the colouring and nidification of birds has been sufficiently established. There are, it is true, a few apparent and some real exceptions, which I shall consider presently; but they are too few and unimportant to weigh much against the mass of evidence on the other side, and may for the present be neglected. Let us then consider what we are to do with this unexpected set of correspondences between groups of phenomena which, at first sight, appear so disconnected. Do they fall in with any other groups of natural phenomena? Do they teach us anything of the way in which nature works, and give us any insight into the causes which have brought about the marvellous variety, and beauty, and harmony of living things? I believe we can answer these questions in the affirmative; and I may mention, as a sufficient proof that these are not isolated facts, that I was first led to see their relation to each other by the study of an analogous though distinct set of phenomena among insects, that of protective resemblance and "mimicry."

On considering this remarkable series of corresponding facts, the first thing we are taught by them seems to be, that there is no incapacity in the female sex among birds, to receive the same bright hues and strongly contrasted tints with which their partners are so often decorated, since whenever they are protected and concealed during the period of incubation they are similarly adorned. The fair inference is, that it is chiefly due to the absence of protection or concealment during this important epoch, that gay and conspicuous tints are withheld or left undeveloped. The mode in which this has been effected is very intelligible, if we admit the action of natural and sexual selection. It would appear from the numerous cases in which both sexes are adorned with equally brilliant colours (while both sexes are rarely armed with equally developed offensive and defensive weapons when not required for individual safety), that the normal action of "sexual selection" is to develop colour and beauty in both sexes, by the preservation and multiplication of all varieties of colour in either sex which are pleasing to the other. Several very close observers of the habits of animals have assured me, that male birds and quadrupeds do often take very strong likes and dislikes to individual females, and we can hardly believe that the one sex (the female) can have a general taste for colour while the other has no such taste. However this may be, the fact remains, that in a vast number of cases the female acquires as brilliant and as varied colours as the male, and therefore most probably acquires them in the same way as the male does; that is, either because the colour is useful to it, or is correlated with some useful variation, or is pleasing to the other sex. The only remaining supposition is that it is transmitted from the other sex, without being of any use. From the number of examples above adduced of bright colours in the female, this would imply that colour-characters acquired by one sex are generally (but not necessarily) transmitted to the other. If this be the case it will, I think, enable us to explain the phenomena, even if we do not admit that the male bird is ever influenced in the choice of a mate by her more gay or perfect plumage.

The female bird, while sitting on her eggs in an uncovered nest, is much exposed to the attacks of enemies, and any modification of colour which rendered her more conspicuous would often lead to her destruction and that of her offspring. All variations of colour in this direction in the female, would therefore sooner or later be eliminated, while such modifications as rendered her inconspicuous, by assimilating her to surrounding objects, as the earth or the foliage, would, on the whole, survive the longest, and thus lead to the attainment of those brown or green and inconspicuous tints, which form the colouring (of the upper surface at least), of the vast majority of female birds which sit upon open nests.

This does not imply, as some have thought, that all female birds were once as brilliant as the males. The change has been a very gradual one, generally dating from the origin of genera or of larger groups, but there can be no doubt that the remote ancestry of birds having great sexual differences of colour, were nearly or quite alike, sometimes (perhaps in most cases) more nearly resembling the female, but occasionally perhaps being nearer what the male is now. The young birds (which usually resemble the females) will probably give some idea of this ancestral type, and it is well known that the young of allied species and of different sexes are often undistinguishable.

Colour more variable than Structure or Habits, and therefore the Character which has generally been Modified.

At the commencement of this essay, I have endeavoured to prove, that the characteristic differences and the essential features of birds' nests, are dependent on the structure of the species and upon the present and past conditions of their existence. Both these factors are more important and less variable than colour; and we must therefore conclude that in most cases the mode of nidification (dependent on structure and environment) has been the cause, and not the effect, of the similarity or differences of the sexes as regards colour. When the confirmed habit of a group of birds, was to build their nests in holes of trees like the toucans, or in holes in the ground like the kingfishers, the protection the female thus obtained, during the important and dangerous time of incubation, placed the two sexes on an equality as regards exposure to attack, and allowed "sexual selection," or any other cause, to act unchecked in the development of gay colours and conspicuous markings in both sexes.

When, on the other hand (as in the Tanagers and Flycatchers), the habit of the whole group was to build open cup-shaped nests in more or less exposed situations, the production of colour and marking in the female, by whatever cause, was continually checked by its rendering her too conspicuous, while in the male it had free play, and developed in him the most gorgeous hues. This, however, was not perhaps universally the case; for where there was more than usual intelligence and capacity for change of habits, the danger the female was exposed to by a partial brightness of colour or marking might lead to the construction of a concealed or covered nest, as in the case of the Tits and Hangnests. When this occurred, a special protection to the female would be no longer necessary; so that the acquisition of colour and the modification of the nest, might in some cases act and react on each other and attain their full development together.

Exceptional Cases confirmatory of the above Explanation.

There exist a few very curious and anomalous facts in the natural history of birds, which fortunately serve as crucial tests of the truth of this mode of explaining the inequalities of sexual colouration. It has been long known, that in some species the males either assisted in, or wholly performed, the act of incubation. It has also been often noticed, that in certain birds the usual sexual differences were reversed, the male being the more plainly coloured, the female more gay and often larger. I am not, however, aware that these two anomalies had ever been supposed to stand to each other in the relation of cause and effect, till I adduced them in support of my views of the general theory of protective adaptation. Yet it is undoubtedly the fact, that in the best known cases in which the female bird is more conspicuously coloured than the male, it is either positively ascertained that the latter performs the duties of incubation, or there are good reasons for believing such to be the case. The most satisfactory example is that of the Gray Phalarope (Phalaropus fulicarius), the sexes of which are alike in winter, while in summer the female instead of the male takes on a gay and conspicuous nuptial plumage; but the male performs the duties of incubation, sitting upon the eggs, which are laid upon the bare ground.

In the Dotterell (Eudromias morinellus) the female is larger and more brightly coloured than the male; and here, also, it is almost certain that the latter sits upon the eggs. The Turnices of India also, have the female larger and often more brightly coloured; and Mr. Jerdon states, in his "Birds of India," that the natives report, that, during the breeding season, the females desert their eggs and associate in flocks, while the males are employed in hatching the eggs. In the few other cases in which the females are more brightly coloured, the habits are not accurately known. The case of the Ostriches and Emeus will occur to many as a difficulty, for here the male incubates, but is not less conspicuous than the female; but there are two reasons why the case does not apply;—the birds are too large to derive any safety from concealment, from enemies which would devour the eggs they can defend themselves by force, while to escape from their personal foes they trust to speed.

We find, therefore, that a very large mass of facts relating to the sexual colouration and the mode of nidification of birds, including some of the most extraordinary anomalies to be found in their natural history, can be shown to have an interdependent relation to each other, on the simple principle of the need of greater protection to that parent which performs the duties of incubation. Considering the very imperfect knowledge we possess of the habits of most extra-European birds, the exceptions to the prevalent rule are few, and generally occur in isolated species or in small groups; while several apparent exceptions can be shown to be really confirmations of the law.

Real or apparent Exceptions to the Law stated at page 240.

The only marked exceptions I have been able to discover are the following:—

1. King crows (Dicrourus). These birds are of a glossy black colour with long forked tails. The sexes present no difference, and they build open nests. This apparent exception may probably be accounted for by the fact that these birds do not need the protection of a less conspicuous colour. They are very pugnacious, and often attack and drive away crows, hawks, and kites; and as they are semi-gregarious in their habits, the females are not likely to be attacked while incubating.

2. Orioles (Oriolidae). The true orioles are very gay birds; the sexes are, in many Eastern species, either nearly or quite alike, and the nests are open. This is one of the most serious exceptions, but it is one that to some extent proves the rule; for in this case it has been noticed, that the parent birds display excessive care and solicitude in concealing the nest among thick foliage, and in protecting their offspring by incessant and anxious watching. This indicates that the want of protection consequent on the bright colour of the female makes itself felt, and is obviated by an increased development of the mental faculties.

3. Ground thrushes (Pittidae). These elegant and brilliantly-coloured birds are generally alike in both sexes, and build an open nest. It is curious, however, that this is only an apparent exception, for almost all the bright colours are on the under surface, the back being usually olive green or brown, and the head black, with brown or whitish stripes, all which colours would harmonize with the foliage, sticks, and roots which surround the nest, built on or near the ground, and thus serve as a protection to the female bird.

4. Grallina Australis. This Australian bird is of strongly contrasted black and white colours. The sexes are exactly alike, and it builds an open clay nest in an exposed situation on a tree. This appears to be a most striking exception, but I am by no means sure that it is so. We require to know what tree it usually builds on, the colour of the bark or of the lichens that grow upon it, the tints of the ground, or of other surrounding objects, before we can say that the bird, when sitting on its nest, is really conspicuous. It has been remarked that small patches of white and black blend at a short distance to form grey, one of the commonest tints of natural objects.

5. Sunbirds (Nectarineidae). In these beautiful little birds the males only are adorned with brilliant colours, the females being quite plain, yet they build covered nests in all the cases in which the nidification is known. This is a negative rather than a positive exception to the rule, since there may be other causes besides the need for protection, which prevent the female acquiring the gay colours of her mate, and there is one curious circumstance which tends to elucidate it. The male of Leptocoma zeylanica is said to assist in incubation. It is possible, therefore, that the group may originally have used open nests, and some change of conditions, leading the male bird to sit, may have been followed by the adoption of a domed nest. This is, however, the most serious exception I have yet found to the general rule.

6. Superb warblers (Maluridae). The males of these little birds are adorned with the most gorgeous colours, while the females are very plain, yet they make domed nests. It is to be observed, however, that the male plumage is nuptial merely, and is retained for a very short time; the rest of the year both sexes are plain alike. It is probable, therefore, that the domed nest is for the protection of these delicate little birds against the rain, and that there is some unknown cause which has led to the development of colour in the males only.

There is one other case which at first sight looks like an exception, but which is far from being one in reality, and deserves to be mentioned. In the beautiful Waxwing, (Bombycilla garrula,) the sexes are very nearly alike, and the elegant red wax tips to the wing-feathers are nearly, and sometimes quite, as conspicuous in the female as in the male. Yet it builds an open nest, and a person looking at the bird would say it ought according to my theory to cover its nest. But it is, in reality, as completely protected by its colouration as the most plainly coloured bird that flies. It breeds only in very high latitudes, and the nest, placed in fir-trees, is formed chiefly of lichens. Now the delicate gray and ashy and purplish hues of the head and back, together with the yellow of the wings and tail, are tints that exactly harmonize with the colours of various species of lichens, while the brilliant red wax tips exactly represent the crimson fructification of the common lichen, Cladonia coccifera. When sitting on its nest, therefore, the female bird will exhibit no colours that are not common to the materials of which it is constructed; and the several tints are distributed in about the same proportions as they occur in nature. At a short distance the bird would be indistinguishable from the nest it is sitting on, or from a natural clump of lichens, and will thus be completely protected.

I think I have now noticed all exceptions of any importance to the law of dependence of sexual colour on nidification. It will be seen that they are very few in number, compared with those which support the generalization; and in several cases there are circumstances in the habits or structure of the species that sufficiently explain them. It is remarkable also that I have found scarcely any positive exceptions, that is, cases of very brilliant or conspicuous female birds in which the nest was not concealed. Much less can there be shown any group of birds, in which the females are all of decidedly conspicuous colours on the upper surface, and yet sit in open nests. The many cases in which birds of dull colours in both sexes make domed or concealed nests, do not, of course, affect this theory one way or the other; since its purpose is only to account for the fact, that brilliant females of brilliant males are always found to have covered or hidden nests, while obscure females of brilliant males almost always have open and exposed nests. The fact that all classes of nests occur with dull coloured birds in both sexes merely shows, as I have strongly maintained, that in most cases the character of the nest determines the colouration of the female, and not vice versa.

If the views here advocated are correct, as to the various influences that have determined the specialities of every bird's nest, and the general colouration of female birds, with their action and reaction on each other, we can hardly expect to find evidence more complete than that here set forth. Nature is such a tangled web of complex relations, that a series of correspondences running through hundreds of species, genera, and families, in every part of the system, can hardly fail to indicate a true casual connexion; and when, of the two factors in the problem, one can be shown to be dependent on the most deeply seated and the most stable facts of structure and conditions of life, while the other is a character universally admitted to be superficial and easily modified, there can be little doubt as to which is cause and which effect.

Various modes of Protection of Animals.

But the explanation of the phenomenon here attempted does not rest alone on the facts I have been able now to adduce. In the essay on "Mimicry," it is shown how important a part the necessity for protection has played, in determining the external form and colouration, and sometimes even the internal structure of animals.

As illustrating this latter point, I may refer to the remarkable hooked, branched, or star-like spiculae in many sponges, which are believed to have the function chiefly, of rendering them unpalatable to other creatures. The Holothuridae or sea-cucumbers possess a similar protection, many of them having anchor-shaped spicules embedded in their skin, as the Synapta; while others (Cuviera squamata) are covered with a hard calcareous pavement. Many of these are of a bright red or purple colour, and are very conspicuous, while the allied Trepang, or Beche-de-mer (Holothuria edulis), which is not armed with any such defensive weapons, is of a dull sand-or mud-colour, so as hardly to be distinguished from the sea bed on which it reposes. Many of the smaller marine animals are protected by their almost invisible transparency, while those that are most brightly coloured will be often found to have a special protection, either in stinging tentacles like Physalia, or in a hard calcareous crust, as in the star fishes.

Females of some Groups require and obtain more Protection than the Males.

In the struggle for existence incessantly going on, protection or concealment is one of the most general and most effectual means of maintaining life; and it is by modifications of colour that this protection can be most readily obtained, since no other character is subject to such numerous and rapid variations. The case I have now endeavoured to illustrate is exactly analogous to what occurs among butterflies. As a general rule, the female butterfly is of dull and inconspicuous colours, even when the male is most gorgeously arrayed; but when the species is protected from attack by a disagreeable odour, as in the Heliconidae, Danaidae and Acroeidae, both sexes display the same or equally brilliant hues. Among the species which gain a protection by imitating these, the very weak and slow-flying Leptalides resemble them in both sexes, because both sexes alike require protection, while in the more active and strong-winged genera—Papilio, Pieris, and Diadema—it is generally the females only that mimic the protected groups, and in doing so often become actually more gay and more conspicuous than the males, thus reversing the usual and in fact almost universal characters of the sexes. So, in the wonderful Eastern leaf-insects of the genus Phyllium, it is the female only that so marvellously imitates a green leaf; and in all these cases the difference can be traced to the greater need of protection for the female, on whose continued existence, while depositing her eggs, the safety of the race depends. In Mammalia and in reptiles, however brilliant the colours may be, there is rarely any difference between that of the sexes, because the female is not necessarily more exposed to attack than the male. It may, I think, be looked upon as a confirmation of this view, that no single case is known either in the above-named genera—Papilio, Pieris, and Diadema—or in any other butterfly, of a male alone, mimicking one of the Danaidae or Heliconidae. Yet the necessary colour is far more abundant in the males, and variations always seem ready for any useful purpose. This seems to depend on the general law, that each species and each sex can only be modified just as far as is absolutely necessary for it to maintain itself in the struggle for existence, not a step further. A male insect by its structure and habits is less exposed to danger, and also requires less protection than the female. It cannot, therefore, alone acquire any further protection through the agency of natural selection. But the female requires some extra protection, to balance the greater danger to which she is exposed, and her greater importance to the existence of the species; and this she always acquires, in one way or another, through the action of natural selection.

In his "Origin of Species," fourth edition, p. 241, Mr. Darwin recognises the necessity for protection as sometimes being a cause of the obscure colours of female birds; but he does not seem to consider it so very important an agent in modifying colour as I am disposed to do. In the same paragraph (p. 240), he alludes to the fact of female birds and butterflies being sometimes very plain, sometimes as gay as the males; but, apparently, considers this mainly due to peculiar laws of inheritance, which sometimes continue acquired colour in the line of one sex only, sometimes in both. Without denying the action of such a law (which Mr. Darwin informs me he has facts to support), I impute the difference, in the great majority of cases, to the greater or less need of protection in the female sex in these groups of animals.

This need was seen to exist a century ago by the Hon. Daines Barrington, who, in the article already quoted (see p. 220), after alluding to the fact that singing birds are all small, and suggesting (but I think erroneously) that this may have arisen from the difficulty larger birds would have in concealing themselves if they called the attention of their enemies by loud notes, goes on thus:—"I should rather conceive it is for the same reason no hen bird sings, because this talent would be still more dangerous during incubation, which may possibly also account for the inferiority in point of plumage." This is a curious anticipation of the main idea on which this essay is founded. It has been unnoticed for near a century, and my attention was only recently called to it by Mr. Darwin himself.

Conclusion.

To some persons it will perhaps appear, that the causes to which I impute so much of the external aspect of nature are too simple, too insignificant, and too unimportant for such a mighty work. But I would ask them to consider, that the great object of all the peculiarities of animal structure is to preserve the life of the individual, and to maintain the existence of the species. Colour has hitherto been too often looked upon as something adventitious and superficial, something given to an animal not to be useful to itself, but solely to gratify man or even superior beings—to add to the beauty and ideal harmony of nature. If this were the case, then, it is evident that the colours of organised beings would be an exception to most other natural phenomena. They would not be the product of general laws, or determined by ever-changing external conditions; and we must give up all enquiry into their origin and causes, since (by the hypothesis) they are dependent on a Will whose motives must ever be unknown to us. But, strange to say, no sooner do we begin to examine and classify the colours of natural objects, than we find that they are intimately related to a variety of other phenomena, and are, like them, strictly subordinated to general laws. I have here attempted to elucidate some of these laws in the case of birds, and have shown how the mode of nidification has affected the colouring of the female sex in this group. I have before shown to how great an extent, and in how many ways, the need of protection has determined the colours of insects, and of some groups of reptiles and mammalia, and I would now call particular attention to the fact that the gay tints of flowers, so long supposed to be a convincing proof that colour has been bestowed for other purposes than the good of its possessor, have been shown by Mr. Darwin to follow the same great law of utility. Flowers do not often need protection, but very often require the aid of insects to fertilize them, and maintain their reproductive powers in the greatest vigour. Their gay colours attract insects, as do also their sweet odours and honeyed secretions; and that this is the main function of colour in flowers is shown by the striking fact, that those flowers which can be perfectly fertilized by the wind, and do not need the aid of insects, rarely or never have gaily-coloured flowers.

This wide extension of the general principle of utility to the colours of such varied groups, both in the animal and vegetable kingdoms, compels us to acknowledge that the "reign of law" has been fairly traced into this stronghold of the advocates of special creation. And to those who oppose the explanation I have given of the facts adduced in this essay, I would again respectfully urge that they must grapple with the whole of the facts, not one or two of them only. It will be admitted that, on the theory of evolution and natural selection, a wide range of facts with regard to colour in nature have been co-ordinated and explained. Until at least an equally wide range of facts can be shown to be in harmony with any other theory, we can hardly be expected to abandon that which has already done such good service, and which has led to the discovery of so many interesting and unexpected harmonies among the most common (but hitherto most neglected and least understood), of the phenomena presented by organised beings.



VIII.

CREATION BY LAW.

Among the various criticisms that have appeared on Mr. Darwin's celebrated "Origin of Species," there is, perhaps, none that will appeal to so large a number of well educated and intelligent persons, as that contained in the Duke of Argyll's "Reign of Law." The noble author represents the feelings and expresses the ideas of that large class, who take a keen interest in the progress of Science in general, and especially that of Natural History, but have never themselves studied nature in detail, or acquired that personal knowledge of the structure of closely allied forms,—the wonderful gradations from species to species and from group to group, and the infinite variety of the phenomena of "variation" in organic beings,—which are absolutely necessary for a full appreciation of the facts and reasonings contained in Mr. Darwin's great work.

Nearly half of the Duke's book is devoted to an exposition of his idea of "Creation by Law," and he expresses so clearly what are his difficulties and objections as regards the theory of "Natural Selection," that I think it advisable that they should be fairly answered, and that his own views should be shown to lead to conclusions, as hard to accept as any which he imputes to Mr. Darwin.

The point on which the Duke of Argyll lays most stress, is, that proofs of Mind everywhere meet us in Nature, and are more especially manifest wherever we find "contrivance" or "beauty." He maintains that this indicates the constant supervision and direct interference of the Creator, and cannot possibly be explained by the unassisted action of any combination of laws. Now, Mr. Darwin's work has for its main object, to show, that all the phenomena of living things,—all their wonderful organs and complicated structures, their infinite variety of form, size, and colour, their intricate and involved relations to each other,—may have been produced by the action of a few general laws of the simplest kind, laws which are in most cases mere statements of admitted facts. The chief of these laws or facts are the following:—

1. The Law of Multiplication in Geometrical Progression.—All organized beings have enormous powers of multiplication. Even man, who increases slower than all other animals, could under the most favourable circumstances double his numbers every fifteen years, or a hundred-fold in a century. Many animals and plants could increase their numbers from ten to a thousand-fold every year.

2. The Law of Limited Populations.—The number of living individuals of each species in any country, or in the whole globe, is practically stationary; whence it follows that the whole of this enormous increase must die off almost as fast as produced, except only those individuals for whom room is made by the death of parents. As a simple but striking example, take an oak forest. Every oak will drop annually thousands or millions of acorns, but till an old tree falls, not one of these millions can grow up into an oak. They must die at various stages of growth.

3. The Law of Heredity, or Likeness of Offspring to their Parents.—This is a universal, but not an absolute law. All creatures resemble their parents in a high degree, and in the majority of cases very accurately; so that even individual peculiarities, of whatever kind, in the parents, are almost always transmitted to some of the offspring.

4. The Law of Variation.—This is fully expressed by the lines:—

"No being on this earthly ball, Is like another, all in all."

Offspring resemble their parents very much, but not wholly—each being possesses its individuality. This "variation" itself varies in amount, but it is always present, not only in the whole being, but in every part of every being. Every organ, every character, every feeling is individual; that is to say, varies from the same organ, character, or feeling in every other individual.

5. The Law of unceasing Change of Physical Conditions upon the Surface of the Earth.—Geology shows us that this change has always gone on in times past, and we also know that it is now everywhere going on.

6. The Equilibrium or Harmony of Nature.—When a species is well adapted to the conditions which environ it, it flourishes; when imperfectly adapted it decays; when ill-adapted it becomes extinct. If all the conditions which determine an organism's well-being are taken into consideration, this statement can hardly be disputed.

* * * * *

This series of facts or laws, are mere statements of what is the condition of nature. They are facts or inferences which are generally known, generally admitted—but in discussing the subject of the "Origin of Species"—as generally forgotten. It is from these universally admitted facts, that the origin of all the varied forms of nature may be deduced by a logical chain of reasoning, which, however, is at every step verified and shown to be in strict accord with facts; and, at the same time, many curious phenomena which can by no other means be understood, are explained and accounted for. It is probable, that these primary facts or laws are but results of the very nature of life, and of the essential properties of organized and unorganized matter. Mr. Herbert Spencer, in his "First Principles" and his "Biology" has, I think, made us able to understand how this may be; but at present we may accept these simple laws without going further back, and the question then is—whether the variety, the harmony, the contrivance, and the beauty we perceive in organic beings, can have been produced by the action of these laws alone, or whether we are required to believe in the incessant interference and direct action of the mind and will of the Creator. It is simply a question of how the Creator has worked. The Duke (and I quote him as having well expressed the views of the more intelligent of Mr. Darwin's opponents) maintains, that He has personally applied general laws to produce effects, which those laws are not in themselves capable of producing; that the universe alone, with all its laws intact, would be a sort of chaos, without variety, without harmony, without design, without beauty; that there is not (and therefore we may presume that there could not be) any self-developing power in the universe. I believe, on the contrary, that the universe is so constituted as to be self-regulating; that as long as it contains Life, the forms under which that life is manifested have an inherent power of adjustment to each other and to surrounding nature; and that this adjustment necessarily leads to the greatest amount of variety and beauty and enjoyment, because it does depend on general laws, and not on a continual supervision and re-arrangement of details. As a matter of feeling and religion, I hold this to be a far higher conception of the Creator and of the Universe than that which may be called the "continual interference" hypothesis; but it is not a question to be decided by our feelings or convictions, it is a question of facts and of reason. Could the change, which Geology shows us has ever taken place in the forms of life, have been produced by general laws, or does it imperatively require the incessant supervision of a creative mind? This is the question for us to consider, and our opponents have the difficult task of proving a negative, if we show that there are both facts and analogies in our favour.

Mr. Darwin's Metaphors liable to Misconception.

Mr. Darwin has laid himself open to much misconception, and has given to his opponents a powerful weapon against himself, by his continual use of metaphor in describing the wonderful co-adaptations of organic beings.

"It is curious," says the Duke of Argyll, "to observe the language which this most advanced disciple of pure naturalism instinctively uses, when he has to describe the complicated structure of this curious order of plants (the Orchids). 'Caution in ascribing intentions to nature,' does not seem to occur to him as possible. Intention is the one thing which he does see, and which, when he does not see, he seeks for diligently until he finds it. He exhausts every form of words and of illustration, by which intention or mental purpose can be described. 'Contrivance'—'curious contrivance,'—'beautiful contrivance,'—these are expressions which occur over and over again. Here is one sentence describing the parts of a particular species: 'the Labellum is developed into a long nectary, in order to attract Lepidoptera, and we shall presently give reason for suspecting that the nectar is purposely so lodged, that it can be sucked only slowly in order to give time for the curious chemical quality of the viscid matter setting hard and dry.'" Many other examples of similar expressions are quoted by the Duke, who maintains that no explanation of these "contrivances" has been or can be given, except on the supposition of a personal contriver, specially arranging the details of each case, although causing them to be produced by the ordinary processes of growth and reproduction.

Now there is a difficulty in this view of the origin of the structure of Orchids which the Duke does not allude to. The majority of flowering plants are fertilized, either without the agency of insects or, when insects are required, without any very important modification of the structure of the flower. It is evident, therefore, that flowers might have been formed as varied, fantastic, and beautiful as the Orchids, and yet have been fertilized without more complexity of structure than is found in Violets, or Clover, or Primroses, or a thousand other flowers. The strange springs and traps and pitfalls found in the flowers of Orchids cannot be necessary per se, since exactly the same end is gained in ten thousand other flowers which do not possess them. Is it not then an extraordinary idea, to imagine the Creator of the Universe contriving the various complicated parts of these flowers, as a mechanic might contrive an ingenious toy or a difficult puzzle? Is it not a more worthy conception that they are some of the results of those general laws which were so co-ordinated at the first introduction of life upon the earth as to result necessarily in the utmost possible development of varied forms?

But let us take one of the simpler cases adduced and see if our general laws are unable to account for it.

A Case of Orchis-structure explained by Natural Selection.

There is a Madagascar Orchis—the Angraecum sesquipedale—with an immensely long and deep nectary. How did such an extraordinary organ come to be developed? Mr. Darwin's explanation is this. The pollen of this flower can only be removed by the base of the proboscis of some very large moths, when trying to get at the nectar at the bottom of the vessel. The moths with the longest probosces would do this most effectually; they would be rewarded for their long tongues by getting the most nectar; whilst on the other hand, the flowers with the deepest nectaries would be the best fertilized by the largest moths preferring them. Consequently, the deepest nectaried Orchids and the longest tongued moths would each confer on the other an advantage in the battle of life. This would tend to their respective perpetuation, and to the constant lengthening of nectaries and probosces. Now let it be remembered, that what we have to account for, is only the unusual length of this organ. A nectary is found in many orders of plants and is especially common in the Orchids, but in this one case only is it more than a foot long. How did this arise? We begin with the fact, proved experimentally by Mr. Darwin, that moths do visit Orchids, do thrust their spiral trunks into the nectaries, and do fertilize them by carrying the pollinia of one flower to the stigma of another. He has further explained the exact mechanism by which this is effected, and the Duke of Argyll admits the accuracy of his observations. In our British species, such as Orchis pyramidalis, it is not necessary that there should be any exact adjustment between the length of the nectary and that of the proboscis of the insect; and thus a number of insects of various sizes are found to carry away the pollinia and aid in the fertilization. In the Angraecum sesquipedale, however, it is necessary that the proboscis should be forced into a particular part of the flower, and this would only be done by a large moth burying its proboscis to the very base, and straining to drain the nectar from the bottom of the long tube, in which it occupies a depth of one or two inches only. Now let us start from the time when the nectary was only half its present length or about six inches, and was chiefly fertilized by a species of moth which appeared at the time of the plant's flowering, and whose proboscis was of the same length. Among the millions of flowers of the Angraecum produced every year, some would always be shorter than the average, some longer. The former, owing to the structure of the flower, would not get fertilized, because the moths could get all the nectar without forcing their trunks down to the very base. The latter would be well fertilized, and the longest would on the average be the best fertilized of all. By this process alone the average length of the nectary would annually increase, because, the short-nectaried flowers being sterile and the long ones having abundant offspring, exactly the same effect would be produced as if a gardener destroyed the short ones and sowed the seed of the long ones only; and this we know by experience would produce a regular increase of length, since it is this very process which has increased the size and changed the form of our cultivated fruits and flowers.

But this would lead in time to such an increased length of the nectary that many of the moths could only just reach the surface of the nectar, and only the few with exceptionally long trunks be able to suck up a considerable portion.

This would cause many moths to neglect these flowers because they could not get a satisfying supply of nectar, and if these were the only moths in the country the flowers would undoubtedly suffer, and the further growth of the nectary be checked by exactly the same process which had led to its increase. But there are an immense variety of moths, of various lengths of proboscis, and as the nectary became longer, other and larger species would become the fertilizers, and would carry on the process till the largest moths became the sole agents. Now, if not before, the moth would also be affected, for those with the longest probosces would get most food, would be the strongest and most vigorous, would visit and fertilize the greatest number of flowers, and would leave the largest number of descendants. The flowers most completely fertilized by these moths being those which had the longest nectaries, there would in each generation be on the average an increase in the length of the nectaries, and also an average increase in the length of the probosces of the moths; and this would be a necessary result from the fact that nature ever fluctuates about a mean, or that in every generation there would be flowers with longer and shorter nectaries, and moths with longer and shorter probosces than the average. No doubt there are a hundred causes that might have checked this process before it had reached the point of development at which we find it. If, for instance, the variation in the quantity of nectar had been at any stage greater than the variation in the length of the nectary, then smaller moths could have reached it and have effected the fertilization. Or if the growth of the probosces of the moths had from other causes increased quicker than that of the nectary, or if the increased length of proboscis had been injurious to them in any way, or if the species of moth with the longest proboscis had become much diminished by some enemy or other unfavourable conditions, then, in any of these cases, the shorter nectaried flowers, which would have attracted and could have been fertilized by the smaller kinds of moths, would have had the advantage. And checks of a similar nature to these no doubt have acted in other parts of the world, and have prevented such an extraordinary development of nectary as has been produced by favourable conditions in Madagascar only, and in one single species of Orchid. I may here mention that some of the large Sphinx moths of the tropics have probosces nearly as long as the nectary of Angraecum sesquipedale. I have carefully measured the proboscis of a specimen of Macrosila cluentius from South America, in the collection of the British Museum, and find it to be nine inches and a quarter long! One from tropical Africa (Macrosila morganii) is seven inches and a half. A species having a proboscis two or three inches longer could reach the nectar in the largest flowers of Angraecum sesquipedale, whose nectaries vary in length from ten to fourteen inches. That such a moth exists in Madagascar may be safely predicted; and naturalists who visit that island should search for it with as much confidence as Astronomers searched for the planet Neptune,—and I venture to predict they will be equally successful!

Now, instead of this beautiful self-acting adjustment, the opposing theory is, that the Creator of the Universe, by a direct act of his Will, so disposed the natural forces influencing the growth of this one species of plant as to cause its nectary to increase to this enormous length; and at the same time, by an equally special act, determined the flow of nourishment in the organization of the moth, so as to cause its proboscis to increase in exactly the same proportion, having previously so constructed the Angraecum that it could only be maintained in existence by the agency of this moth. But what proof is given or suggested that this was the mode by which the adjustment took place? None whatever, except a feeling that there is an adjustment of a delicate kind, and an inability to see how known causes could have produced such an adjustment. I believe I have shown, however, that such an adjustment is not only possible but inevitable, unless at some point or other we deny the action of those simple laws which we have already admitted to be but the expressions of existing facts.

Adaptation brought about by General Laws.

It is difficult to find anything like parallel cases in inorganic nature, but that of a river may perhaps illustrate the subject in some degree. Let us suppose a person totally ignorant of Modern Geology to study carefully a great River System. He finds in its lower part, a deep broad channel filled to the brim, flowing slowly through a flat country and carrying out to the sea a quantity of fine sediment. Higher up it branches into a number of smaller channels, flowing alternately through flat valleys and between high banks; sometimes he finds a deep rocky bed with perpendicular walls, carrying the water through a chain of hills; where the stream is narrow he finds it deep, where wide shallow. Further up still, he comes to a mountainous region, with hundreds of streams and rivulets, each with its tributary rills and gullies, collecting the water from every square mile of surface, and every channel adapted to the water that it has to carry. He finds that the bed of every branch, and stream, and rivulet, has a steeper and steeper slope as it approaches its sources, and is thus enabled to carry off the water from heavy rains, and to bear away the stones and pebbles and gravel, that would otherwise block up its course. In every part of this system he would see exact adaptation of means to an end. He would say, that this system of channels must have been designed, it answers its purpose so effectually. Nothing but a mind could have so exactly adapted the slopes of the channels, their capacity, and frequency, to the nature of the soil and the quantity of the rainfall. Again, he would see special adaptation to the wants of man, in broad quiet navigable rivers flowing through fertile plains that support a large population, while the rocky streams and mountain torrents, were confined to those sterile regions suitable only for a small population of shepherds and herdsmen. He would listen with incredulity to the Geologist, who assured him, that the adaptation and adjustment he so admired was an inevitable result of the action of general laws. That the rains and rivers, aided by subterranean forces, had modelled the country, had formed the hills and valleys, had scooped out the river beds, and levelled the plains;—and it would only be after much patient observation and study, after having watched the minute changes produced year by year, and multiplying them by thousands and ten thousands, after visiting the various regions of the earth and seeing the changes everywhere going on, and the unmistakable signs of greater changes in past times,—that he could be made to understand that the surface of the earth, however beautiful and harmonious it may appear, is strictly due in every detail to the action of forces which are demonstrably self-adjusting.

Moreover, when he had sufficiently extended his inquiries, he would find, that every evil effect which he would imagine must be the result of non-adjustment does somewhere or other occur, only it is not always evil. Looking on a fertile valley, he would perhaps say—"If the channel of this river were not well adjusted, if for a few miles it sloped the wrong way, the water could not escape, and all this luxuriant valley, full of human beings, would become a waste of waters." Well, there are hundreds of such cases. Every lake is a valley "wasted by water," and in some cases (as the Dead Sea) it is a positive evil, a blot upon the harmony and adaptation of the surface of the earth. Again, he might say—"If rain did not fall here, but the clouds passed over us to some other regions, this verdant and highly cultivated plain would become a desert." And there are such deserts over a large part of the earth, which abundant rains would convert into pleasant dwelling-places for man. Or he might observe some great navigable river, and reflect how easily rocks, or a steeper channel in places, might render it useless to man;—and a little inquiry would show him hundreds of rivers in every part of the world, which are thus rendered useless for navigation.

Exactly the same thing occurs in organic nature. We see some one wonderful case of adjustment, some unusual development of an organ, but we pass over the hundreds of cases in which that adjustment and development do not occur. No doubt when one adjustment is absent another takes its place, because no organism can continue to exist that is not adjusted to its environment; and unceasing variation with unlimited powers of multiplication, in most cases, furnish the means of self-adjustment. The world is so constituted, that by the action of general laws there is produced the greatest possible variety of surface and of climate; and by the action of laws equally general, the greatest possible variety of organisms have been produced, adapted to the varied conditions of every part of the earth. Tho objector would probably himself admit, that the varied surface of the earth—the plains and valleys, the hills and mountains, the deserts and volcanoes, the winds and currents, the seas and lakes and rivers, and the various climates of the earth—are all the results of general laws acting and re-acting during countless ages; and that the Creator does not appear to guide and control the action of these laws—here determining the height of a mountain, there altering the channel of a river—here making the rains more abundant, there changing the direction of a current. He would probably admit that the forces of inorganic nature are self-adjusting, and that the result necessarily fluctuates about a given mean condition (which is itself slowly changing), while within certain limits the greatest possible amount of variety is produced. If then a "contriving mind" is not necessary at every step of the process of change eternally going on in the inorganic world, why are we required to believe in the continual action of such a mind in the region of organic nature? True, the laws at work are more complex, the adjustments more delicate, the appearance of special adaptation more remarkable; but why should we measure the creative mind by our own? Why should we suppose the machine too complicated, to have been designed by the Creator so complete that it would necessarily work out harmonious results? The theory of "continual interference" is a limitation of the Creator's power. It assumes that he could not work by pure law in the organic, as he has done in the inorganic world; it assumes that he could not foresee the consequences of the laws of matter and mind combined—that results would continually arise which are contrary to what is best, and that he has to change what would otherwise be the course of nature, in order to produce that beauty, and variety, and harmony, which even we, with our limited intellects, can conceive to be the result of self-adjustment in a universe governed by unvarying law. If we could not conceive the world of nature to be self-adjusting and capable of endless development, it would even then be an unworthy idea of a Creator, to impute the incapacity of our minds to him; but when many human minds can conceive, and can even trace out in detail some of the adaptations in nature as the necessary results of unvarying law, it seems strange that, in the interests of religion, any one should seek to prove that the System of Nature, instead of being above, is far below our highest conceptions of it. I, for one, cannot believe that the world would come to chaos if left to Law alone. I cannot believe that there is in it no inherent power of developing beauty or variety, and that the direct action of the Deity is required to produce each spot or streak on every insect, each detail of structure in every one of the millions of organisms that live or have lived upon the earth. For it is impossible to draw a line. If any modifications of structure could be the result of law, why not all? If some self-adaptations could arise, why not others? If any varieties of colour, why not all the varieties we see? No attempt is made to explain this, except by reference to the fact that "purpose" and "contrivance" are everywhere visible, and by the illogical deduction that they could only have arisen from the direct action of some mind, because the direct action of our minds produces similar "contrivances"; but it is forgotten that adaptation, however produced, must have the appearance of design. The channel of a river looks as if made for the river, although it is made by it; the fine layers and beds in a deposit of sand, often look as if they had been sorted, and sifted, and levelled, designedly; the sides and angles of a crystal exactly resemble similar forms designed by man; but we do not therefore conclude that these effects have, in each individual case, required the directing action of a creative mind, or see any difficulty in their being produced by natural Law.

Beauty in Nature.

Let us, however, leave this general argument for a while, and turn to another special case, which has been appealed to as conclusive against Mr. Darwin's views. "Beauty" is, to some persons, as great a stumbling-block as "contrivance." They cannot conceive a system of the Universe, so perfect, as necessarily to develop every form of Beauty, but suppose that when anything specially beautiful occurs, it is a step beyond what that system could have produced, something which the Creator has added for his own delectation.

Speaking of the Humming Birds, the Duke of Argyll says: "In the first place, it is to be observed of the whole group, that there is no connection which can be traced or conceived, between the splendour of the humming birds and any function essential to their life. If there were any such connection, that splendour could not be confined, as it almost exclusively is, to only one sex. The female birds are, of course, not placed at any disadvantage in the struggle for existence by their more sombre colouring." And after describing the various ornaments of these birds, he says: "Mere ornament and variety of form, and these for their own sake, is the only principle or rule with reference to which Creative Power seems to have worked in these wonderful and beautiful birds.... A crest of topaz is no better in the struggle for existence than a crest of sapphire. A frill ending in spangles of the emerald is no better in the battle of life than a frill ending in spangles of the ruby. A tail is not affected for the purposes of flight, whether its marginal or its central feathers are decorated with white.... Mere beauty and mere variety, for their own sake, are objects which we ourselves seek when we can make the Forces of Nature subordinate to the attainment of them. There seems to be no conceivable reason why we should doubt or question, that these are ends and aims also in the forms given to living organisms" ("Reign of Law," p. 248).

Here the statement that "no connection can be conceived between the splendour of the humming birds and any function essential to their life," is met by the fact, that Mr. Darwin has not only conceived but has shown, both by observation and reasoning, how beauty of colour and form may have a direct influence on the most important of all the functions of life, that of reproduction. In the variations to which birds are subject, any more brilliant colour than usual would be attractive to the females, and would lead to the individuals so adorned leaving more than the average number of offspring. Experiment and observation have shown, that this kind of sexual selection does actually take place; and the laws of inheritance would necessarily lead to the further development of any individual peculiarity that was attractive, and thus the splendour of the humming birds is directly connected with their very existence. It is true that "a crest of topaz may be no better than a crest of sapphire," but either of these may be much better than no crest at all; and the different conditions under which the parent form must have existed in different parts of its range, will have determined different variations of tint, either of which were advantageous. The reason why female birds are not adorned with equally brilliant plumes is sufficiently clear; they would be injurious, by rendering their possessors too conspicuous during incubation. Survival of the fittest, has therefore favoured the development of those dark green tints on the upper surface of so many female humming birds, which are most conducive to their protection while the important functions of hatching and rearing the young are being carried on. Keeping in mind the laws of multiplication, variation, and survival of the fittest, which are for ever in action, these varied developments of beauty and harmonious adjustments to conditions, are not only conceivable but demonstrable results.

The objection I am now combating is solely founded on the supposed analogy of the Creator's mind to ours, as regards the love of Beauty for its own sake; but if this analogy is to be trusted, then there ought to be no natural objects which are disagreeable or ungraceful in our eyes. And yet it is undoubtedly the fact that there are many such. Just as surely as the Horse and Deer are beautiful and graceful, the Elephant, Rhinoceros, Hippopotamus, and Camel are the reverse. The majority of Monkeys and Apes are not beautiful; the majority of Birds have no beauty of colour; a vast number of Insects and Reptiles are positively ugly. Now, if the Creator's mind is like ours, whence this ugliness? It is useless to say "that is a mystery we cannot explain," because we have attempted to explain one-half of creation by a method that will not apply to the other half. We know that a man with the highest taste and with unlimited wealth, practically does abolish all ungraceful and disagreeable forms and colours from his own domains. If the beauty of creation is to be explained by the Creator's love of beauty, we are bound to ask why he has not banished deformity from the earth, as the wealthy and enlightened man does from his estate and from his dwelling; and if we can get no satisfactory answer, we shall do well to reject the explanation offered. Again, in the case of flowers, which are always especially referred to, as the surest evidence of beauty being an end of itself in creation, the whole of the facts are never fairly met. At least half the plants in the world have not bright-coloured or beautiful flowers; and Mr. Darwin has lately arrived at the wonderful generalization, that flowers have become beautiful solely to attract insects to assist in their fertilization. He adds, "I have come to this conclusion from finding it an invariable rule, that when a flower is fertilized by the wind it never has a gaily-coloured corolla." Here is a most wonderful case of beauty being useful, when it might be least expected. But much more is proved; for when beauty is of no use to the plant it is not given. It cannot be imagined to do any harm. It is simply not necessary, and is therefore withheld! We ought surely to have been told how this fact is consistent with beauty being "an end in itself," and with the statement of its being given to natural objects "for its own sake."

How new Forms are produced by Variation and Selection.

Let us now consider another of the popular objections which the Duke of Argyll thus sets forth:—

"Mr. Darwin does not pretend to have discovered any law or rule, according to which new Forms have been born from old Forms. He does not hold that outward conditions, however changed, are sufficient to account for them.... His theory seems to be far better than a mere theory—to be an established scientific truth—in so far as it accounts, in part at least, for the success and establishment and spread of new Forms when they have arisen. But it does not even suggest the law under which, or by or according to which, such new Forms are introduced. Natural Selection can do nothing, except with the materials presented to its hands. It cannot select except among the things open to selection.... Strictly speaking, therefore, Mr. Darwin's theory is not a theory on the Origin of Species at all, but only a theory on the causes which lead to the relative success or failure of such new forms as may be born into the world." ("Reign of Law," p. 230.)

In this, and many other passages in his work, the Duke of Argyll sets forth his idea of Creation as a "Creation by birth," but maintains that each birth of a new form from parents differing from itself, has been produced by a special interference of the Creator, in order to direct the process of development into certain channels; that each new species is in fact a "special creation," although brought into existence through the ordinary laws of reproduction. He maintains therefore, that the laws of multiplication and variation cannot furnish the right kinds of materials at the right times for natural selection to work on. I believe, on the contrary, that it can be logically proved from the six axiomatic laws before laid down, that such materials would be furnished; but I prefer to show there are abundance of facts which demonstrate that they are furnished.

The experience of all cultivators of plants and breeders of animals shows, that when a sufficient number of individuals are examined, variations of any required kind can always be met with. On this depends the possibility of obtaining breeds, races, and fixed varieties of animals and plants; and it is found, that any one form of variation may be accumulated by selection, without materially affecting the other characters of the species; each seems to vary in the one required direction only. For example, in turnips, radishes, potatoes, and carrots, the root or tuber varies in size, colour, form, and flavour, while the foliage and flowers seem to remain almost stationary; in the cabbage and lettuce, on the contrary, the foliage can be modified into various forms and modes of growth, the root, flower, and fruit remaining little altered; in the cauliflower and brocoli the flower heads vary; in the garden pea the pod only changes. We get innumerable forms of fruit in the apple and pear, while the leaves and flowers remain undistinguishable; the same occurs in the gooseberry and garden currant. Directly however, (in the very same genus) we want the flower to vary in the Ribes sanguineum, it does so, although mere cultivation for hundreds of years has not produced marked differences in the flowers of Ribes grossularia. When fashion demands any particular change in the form or size, or colour of a flower, sufficient variation always occurs in the right direction, as is shown by our roses, auriculas, and geraniums; when, as recently, ornamental leaves come into fashion sufficient variation is found to meet the demand, and we have zoned pelargoniums, and variegated ivy, and it is discovered that a host of our commonest shrubs and herbaceous plants have taken to vary in this direction just when we want them to do so! This rapid variation is not confined to old and well-known plants subjected for a long series of generations to cultivation, but the Sikim Rhododendrons, the Fuchsias, and Calceolarias from the Andes, and the Pelargoniums from the Cape are equally accommodating, and vary just when and where and how we require them.

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