Chapter XI: The Special Colours of Plants: Their Origin and Purpose (2)

The rapidity and comparative certainty with which such changes as are here supposed do really take place, are well shown by the great differences in floral structure, as regards the mode of fertilisation, in allied genera and species, and even in some cases in varieties of the same species. Thus in the Ranunculaceae we find the conspicuous part of the flower to be the petals in Ranunculus, the sepals in Helleborus, Anemone, etc., and the stamens in most species of Thalictrum. In all these we have a simple regular flower, but in Aquilegia it is made complex by the spurred petals, and in Delphinium and Aconitum it becomes quite irregular. In the more simple class self-fertilisation occurs freely, but it is prevented in the more complex flowers by the stamens maturing before the pistil. In the Caprifoliaceae we have small and regular greenish flowers, as in the moschatel (Adoxa); more conspicuous regular open flowers without honey, as in the elder (Sambucus); and tubular flowers increasing in length and irregularity, till in some, like our common honeysuckle, they are adapted for fertilisation by moths only, with abundant honey and delicious perfume to attract them. In the Scrophulariaceae we find open, almost regular flowers, as Veronica and Verbascum, fertilised by flies and bees, but also self-fertilised; Scrophularia adapted in form and colour to be fertilised by wasps; and the more complex and irregular flowers of Linaria, Rhinanthus, Melampyrum, Pedicularis, etc., mostly adapted to be fertilised by bees.

In the genera Geranium, Polygonum, Veronica, and several others there is a gradation of forms from large and bright to small and obscure coloured flowers, and in every case the former are adapted for insect fertilisation, often exclusively, while in the latter self-fertilisation constantly occurs. In the yellow rattle (Rhinanthus Crista-galli) there are two forms (which have been named _major_ and _minor_), the larger and more conspicuous adapted to insect fertilisation only, the smaller capable of self-fertilisation; and two similar forms exist in the eyebright (Euphrasia officinalis). In both these cases there are special modifications in the length and curvature of the style as well as in the size and shape of the corolla; and the two forms are evidently becoming each adapted to special conditions, since in some districts the one, in other districts the other is most abundant.[159]

These examples show us that the kind of change suggested above is actually going on, and has presumably always been going on in nature throughout the long geological epochs during which the development of flowers has been progressing. The two great modes of gaining increased vigour and fertility--intercrossing and dispersal over wider areas--have been resorted to again and again, under the pressure of a constant struggle for existence and the need for adaptation to ever-changing conditions. During all the modifications that ensued, useless parts were reduced or suppressed, owing to the absence of selection and the principle of economy of growth; and thus at each fresh adaptation some rudiments of old structures were re-developed, but not unfrequently in a different form and for a distinct purpose.

The chief types of flowering plants have existed during the millions of ages of the whole tertiary period, and during this enormous lapse of time many of them may have been modified in the direction of insect fertilisation, and again into that of self-fertilisation, not once or twice only, but perhaps scores or even hundreds of times; and at each such modification a difference in the environment may have led to a distinct line of development. At one epoch the highest specialisation of structure in adaptation to a single species or group of insects may have saved a plant from extinction; while, at other times, the simplest mode of self-fertilisation, combined with greater powers of dispersal and a constitution capable of supporting diverse physical conditions, may have led to a similar result. With some groups the tendency seems to have been almost continuously to greater and greater specialisation, while with others a tendency to simplification and degradation has resulted in such plants as the grasses and sedges.

We are now enabled dimly to perceive how the curious anomaly of very simple and very complex methods of securing cross-fertilisation--both equally effective--may have been brought about. The simple modes may be the result of a comparatively direct modification from the more primitive types of flowers, which were occasionally, and, as it were, accidentally visited and fertilised by insects; while the more complex modes, existing for the most part in the highly irregular flowers, may result from those cases in which adaptation to insect-fertilisation, and partial or complete degradation to self-fertilisation or to wind-fertilisation, have again and again recurred, each time producing some additional complexity, arising from the working up of old rudiments for new purposes, till there have been reached the marvellous flower structures of the papilionaceous tribes, of the asclepiads, or of the orchids.

We thus see that the existing diversity of colour and of structure in flowers is probably the ultimate result of the ever-recurring struggle for existence, combined with the ever-changing relations between the vegetable and animal kingdoms during countless ages. The constant variability of every part and organ, with the enormous powers of increase possessed by plants, have enabled them to become again and again readjusted to each change of condition as it occurred, resulting in that endless variety, that marvellous complexity, and that exquisite colouring which excite our admiration in the realm of flowers, and constitute them the perennial charm and crowning glory of nature.

_Flowers the Product of Insect Agency._

In his _Origin of Species_, Mr. Darwin first stated that flowers had been rendered conspicuous and beautiful in order to attract insects, adding: "Hence we may conclude that, if insects had not been developed on the earth, our plants would not have been decked with beautiful flowers, but would have produced only such poor flowers as we see on our fir, oak, nut, and ash trees, on grasses, docks, and nettles, which are all fertilised through the agency of the wind." The argument in favour of this view is now much stronger than when he wrote; for not only have we reason to believe that most of these wind-fertilised flowers are degraded forms of flowers which have once been insect fertilised, but we have abundant evidence that whenever insect agency becomes comparatively ineffective, the colours of the flowers become less bright, their size and beauty diminish, till they are reduced to such small, greenish, inconspicuous flowers as those of the rupture-wort (Herniaria glabra), the knotgrass (Polygonum aviculare), or the cleistogamic flowers of the violet. There is good reason to believe, therefore, not only that flowers have been developed in order to attract insects to aid in their fertilisation, but that, having been once produced, in however great profusion, if the insect races were all to become extinct, flowers (in the temperate zones at all events) would soon dwindle away, and that ultimately all floral beauty would vanish from the earth.

We cannot, therefore, deny the vast change which insects have produced upon the earth's surface, and which has been thus forcibly and beautifully delineated by Mr. Grant Allen: "While man has only tilled a few level plains, a few great river valleys, a few peninsular mountain slopes, leaving the vast mass of earth untouched by his hand, the insect has spread himself over every land in a thousand shapes, and has made the whole flowering creation subservient to his daily wants. His buttercup, his dandelion, and his meadow-sweet grow thick in every English field. His thyme clothes the hillside; his heather purples the bleak gray moorland. High up among the alpine heights his gentian spreads its lakes of blue; amid the snows of the Himalayas his rhododendrons gleam with crimson light. Even the wayside pond yields him the white crowfoot and the arrowhead, while the broad expanses of Brazilian streams are beautified by his gorgeous water-lilies. The insect has thus turned the whole surface of the earth into a boundless flower-garden, which supplies him from year to year with pollen or honey, and itself in turn gains perpetuation by the baits that it offers for his allurement."[160]

_Concluding Remarks on Colour in Nature._

In the last four chapters I have endeavoured to give a general and systematic, though necessarily condensed view of the part which is played by colour in the organic world. We have seen in what infinitely varied ways the need of concealment has led to the modification of animal colours, whether among polar snows or sandy deserts, in tropical forests or in the abysses of the ocean. We next find these general adaptations giving way to more specialised types of coloration, by which each species has become more and more harmonised with its immediate surroundings, till we reach the most curiously minute resemblances to natural objects in the leaf and stick insects, and those which are so like flowers or moss or birds' droppings that they deceive the acutest eye. We have learnt, further, that these varied forms of protective colouring are far more numerous than has been usually suspected, because, what appear to be very conspicuous colours or markings when the species is observed in a museum or in a menagerie, are often highly protective when the creature is seen under the natural conditions of its existence. From these varied classes of facts it seems not improbable that fully one-half of the species in the animal kingdom possess colours which have been more or less adapted to secure for them concealment or protection.

Passing onward we find the explanation of a distinct type of colour or marking, often superimposed upon protective tints, in the importance of easy recognition by many animals of their fellows, their parents, or their mates. By this need we have been able to account for markings that seem calculated to make the animal conspicuous, when the general tints and well-known habits of the whole group demonstrate the need of concealment. Thus also we are able to explain the constant symmetry in the markings of wild animals, as well as the numerous cases in which the conspicuous colours are concealed when at rest and only become visible during rapid motion. In striking contrast to ordinary protective coloration we have "warning colours," usually very conspicuous and often brilliant or gaudy, which serve to indicate that their possessors are either dangerous or uneatable to the usual enemies of their tribe. This kind of coloration is probably more prevalent than has been hitherto supposed, because in the case of many tropical animals we are quite unacquainted with their special and most dangerous enemies, and are also unable to determine whether they are or are not distasteful to those enemies. As a kind of corollary to the "warning colours," we find the extraordinary phenomena of "mimicry," in which defenceless species obtain protection by being mistaken for those which, from any cause, possess immunity from attack. Although a large number of instances of warning colour and of mimicry are now recorded, it is probably still an almost unworked field of research, more especially in tropical regions and among the inhabitants of the ocean.

The phenomena of sexual diversities of coloration next engaged our attention, and the reasons why Mr. Darwin's theory of "sexual selection," as regards colour and ornament, could not be accepted were stated at some length, together with the theory of animal coloration and ornament we propose to substitute for it. This theory is held to be in harmony with the general facts of animal coloration, while it entirely dispenses with the very hypothetical and inadequate agency of female choice in producing the detailed colours, patterns, and ornaments, which in so many cases distinguish the male sex.

If my arguments on this point are sound, they will dispose also of Mr. Grant Allen's view of the direct action of the colour sense on the animal integuments.[161] He argues that the colours of insects and birds reproduce generally the colours of the flowers they frequent or the fruits they eat, and he adduces numerous cases in which flower-haunting insects and fruit-eating birds are gaily coloured. This he supposes to be due to the colour-taste, developed by the constant presence of bright flowers and fruits, being applied to the selection of each variation towards brilliancy in their mates; thus in time producing the gorgeous and varied hues they now possess. Mr. Allen maintains that "insects are bright where bright flowers exist in numbers, and dull where flowers are rare or inconspicuous;" and he urges that "we can hardly explain this wide coincidence otherwise than by supposing that a taste for colour is produced through the constant search for food among entomophilous blossoms, and that this taste has reacted upon its possessors through the action of unconscious sexual selection."

The examples Mr. Allen quotes of bright insects being associated with bright flowers seem very forcible, but are really deceptive or erroneous; and quite as many cases could be quoted which prove the very opposite. For example, in the dense equatorial forests flowers are exceedingly scarce, and there is no comparison with the amount of floral colour to be met with in our temperate meadows, woods, and hillsides. The forests about Para in the lower Amazon are typical in this respect, yet they abound with the most gorgeously coloured butterflies, almost all of which frequent the forest depths, keeping near the ground, where there is the greatest deficiency of brilliant flowers. In contrast with this let us take the Cape of Good Hope--the most flowery region probably that exists upon the globe,--where the country is a complete flower-garden of heaths, pelargoniums, mesembryanthemus, exquisite iridaceous and other bulbs, and numerous flowering shrubs and trees; yet the Cape butterflies are hardly equal, either in number or variety, to those of any country in South Europe, and are utterly insignificant when compared with those of the comparatively flowerless forest-depths of the Amazon or of New Guinea. Neither is there any relation between the colours of other insects and their haunts. Few are more gorgeous than some of the tiger-beetles and the carabi, yet these are all carnivorous; while many of the most brilliant metallic buprestidae and longicorns are always found on the bark of fallen trees. So with the humming-birds; their brilliant metallic tints can only be compared with metals or gems, and are totally unlike the delicate pinks and purples, yellows and reds of the majority of flowers. Again, the Australian honey-suckers (Meliphagidae) are genuine flower-haunters, and the Australian flora is more brilliant in colour display than that of most tropical regions, yet these birds are, as a rule, of dull colours, not superior on the average to our grain-eating finches. Then, again, we have the grand pheasant family, including the gold and the silver pheasants, the gorgeous fire-backed and ocellated pheasants, and the resplendent peacock, all feeding on the ground on grain or seeds or insects, yet adorned with the most gorgeous colours.

There is, therefore, no adequate basis of facts for this theory to rest upon, even if there were the slightest reason to believe that not only birds, but butterflies and beetles, take any delight in colour for its own sake, apart from the food-supply of which it indicates the presence. All that has been proved or that appears to be probable is, that they are able to perceive differences of colour, and to associate each colour with the particular flowers or fruits which best satisfy their wants. Colour being in its nature diverse, it has been beneficial for them to be able to distinguish all its chief varieties, as manifested more particularly in the vegetable kingdom, and among the different species of their own group; and the fact that certain species of insects show some preference for a particular colour may be explained by their having found flowers of that colour to yield them a more abundant supply of nectar or of pollen. In those cases in which butterflies frequent flowers of their own colour, the habit may well have been acquired from the protection it affords them.

It appears to me that, in imputing to insects and birds the same love of colour for its own sake and the same aesthetic tastes as we ourselves possess, we may be as far from the truth as were those writers who held that the bee was a good mathematician, and that the honeycomb was constructed throughout to satisfy its refined mathematical instincts; whereas it is now generally admitted to be the result of the simple principle of economy of material applied to a primitive cylindrical cell.[162]

In studying the phenomena of colour in the organic world we have been led to realise the wonderful complexity of the adaptations which bring each species into harmonious relation with all those which surround it, and which thus link together the whole of nature in a network of relations of marvellous intricacy. Yet all this is but, as it were, the outward show and garment of nature, behind which lies the inner structure--the framework, the vessels, the cells, the circulating fluids, and the digestive and reproductive processes,--and behind these again those mysterious chemical, electrical, and vital forces which constitute what we term Life. These forces appear to be fundamentally the same for all organisms, as is the material of which all are constructed; and we thus find behind the outer diversities an inner relationship which binds together the myriad forms of life.

Each species of animal or plant thus forms part of one harmonious whole, carrying in all the details of its complex structure the record of the long story of organic development; and it was with a truly inspired insight that our great philosophical poet apostrophised the humble weed--

Flower in the crannied wall,
I pluck you out of the crannies,
I hold you here, root and all, in my hand,
Little flower--but _if_ I could understand
What you are, root and all, and all in all,
I should know what God and man is.

FOOTNOTES:

[Footnote 136: Burchell's _Travels_, vol. i. p. 10.]

[Footnote 137: _Nature_, vol. iii. p. 507.]

[Footnote 138: _Flowers, Fruits, and Leaves_, p. 128 (Fig. 79).]

[Footnote 139: For a popular sketch of these, see Sir J. Lubbock's _Flowers, Fruits, and Leaves_, or any general botanical work.]

[Footnote 140: _Nature_, vol. xv. p, 117.]

[Footnote 141: Grant Allen's _Colour Sense_, p. 113.]

[Footnote 142: Canon Tristram's _Natural History of the Bible_, pp. 483, 484.]

[Footnote 143: For a complete historical account of this subject with full references to all the works upon it, see the Introduction to Hermann Müller's _Fertilisation of Flowers_, translated by D'Arcy W. Thompson.]

[Footnote 144: For the full detail of his experiments, see _Cross-and Self-Fertilisation of Plants_, 1876.]

[Footnote 145: See Darwin's _Fertilisation of Orchids_ for the many extraordinary and complex arrangements in these plants.]

[Footnote 146: The English reader may consult Sir John Lubbock's _British Wild Flowers in Relation to Insects_, and H. Müller's great and original work, _The Fertilisation of Flowers_.]

[Footnote 147: Müller's _Fertilisation of Flowers_, p. 248.]

[Footnote 148: "Alpenblumen," by D.H. Müller. See _Nature_, vol. xxiii. p. 333.]

[Footnote 149: This peculiarity of local distribution of colour in flowers may be compared, as regards its purpose, with the recognition colours of animals. Just as these latter colours enable the sexes to recognise each other, and thus avoid sterile unions of distinct species, so the distinctive form and colour of each species of flower, as compared with those that usually grow around it, enables the fertilising insects to avoid carrying the pollen of one flower to the stigma of a distinct species.]

[Footnote 150: See H. Müller's _Fertilisation of Flowers_, p. 18.]

[Footnote 151: The above examples are taken from Rev. G. Henslow's paper on "Self-Fertilisation of Plants," in _Trans. Linn. Soc._ Second series, _Botany_, vol. i. pp. 317-398, with plate. Mr. H.O. Forbes has shown that the same thing occurs among tropical orchids, in his paper "On the Contrivances for insuring Self-Fertilisation in some Tropical Orchids," _Journ. Linn. Soc._, xxi. p. 538.]

[Footnote 152: These are the numbers given by Darwin, but I am informed by Mr. Hemsley that many additions have been since made to the list, and that cleistogamic flowers probably occur in nearly all the natural orders.]

[Footnote 153: For a full account of cleistogamic flowers, see Darwin's _Forms of Flowers_, chap. viii.]

[Footnote 154: Henslow's "Self-Fertilisation," _Trans. Linn. Soc._ Second series, _Botany_, vol. i. p. 391.]

[Footnote 155: The Rev. George Henslow, in his _Origin of Floral Structures_, says: "There is little doubt but that all wind-fertilised angiosperms are degradations from insect-fertilised flowers.... _Poterium sanguisorba_ is anemophilous; and _Sanguisorba officinalis_ presumably was so formerly, but has reacquired an entomophilous habit; the whole tribe Poterieae being, in fact, a degraded group which has descended from Potentilleae. Plantains retain their corolla but in a degraded form. Junceae are degraded Lilies; while Cyperaceae and Gramineae among monocotyledons may be ranked with Amentiferae among dicotyledons, as representing orders which have retrograded very far from the entomophilous forms from which they were possibly and probably descended" (p. 266).

"The genus Plantago, like _Thalictrum minus_, Poterium, and others, well illustrate the change from an entomophilous to the anemophilous state. _P. lanceolata_ has polymorphic flowers, and is visited by pollen-seeking insects, so that it can be fertilised either by insects or the wind. _P. media_ illustrates transitions in point of structure, as the filaments are pink, the anthers motionless, and the pollen grains aggregated, and it is regularly visited by _Bombus terrestris_. On the other hand, the slender filaments, versatile anthers, powdery pollen, and elongated protogynous style are features of other species indicating anemophily; while the presence of a degraded corolla shows its ancestors to have been entomophilous. _P. media_, therefore, illustrates, not a primitive entomophilous condition, but a return to it; just as is the case with _Sanguisorba officinalis_ and _Salix Caprea_; but these show no capacity of restoring the corolla, the attractive features having to be borne by the calyx, which is purplish in Sanguisorba, by the pink filaments of Plantago, and by the yellow anthers in the Sallow willow" (p. 271).

"The interpretation, then, I would offer of inconspicuousness and all kinds of degradations is the exact opposite to that of conspicuousness and great differentiations; namely, that species with minute flowers, rarely or never visited by insects, and habitually self-fertilised, have primarily arisen through the neglect of insects, and have in consequence assumed their present floral structures" (p. 282).

In a letter just received from Mr. Henslow, he gives a few additional illustrations of his views, of which the following are the most important: "Passing to Incompletae, the orders known collectively as 'Cyclospermeae' are related to Caryophylleae; and to my mind are degradations from it, of which Orache is anemophilous. Cupuliferae have an inferior ovary and rudimentary calyx-limb on the top. These, as far as I know, cannot be interpreted except as degradations. The whole of Monocotyledons appear to me (from anatomical reasons especially) to be degradations from Dicotyledons, and primarily through the agency of growth in water. Many subsequently became terrestrial, but retained the effects of their primitive habitat through heredity. The 3-merous [sic] perianth of grasses, the parts of the flower being in whorls, point to a degradation from a sub-liliaceous condition."

Mr. Henslow informs me that he has long held these views, but, as far as he knows, alone. Mr. Grant Allen, however, set forth a similar theory in his _Vignettes from Nature_ (p. 15) and more fully in _The Colours of Flowers_ (chap. v.), where he develops it fully and uses similar arguments to those of Mr. Henslow.]

[Footnote 156: H. Müller gives ample proof of this in his _Fertilisation of Flowers_.]

[Footnote 157: _Cross-and Self-Fertilisation_, p. 27.]

[Footnote 158: _Animals and Plants_, vol. ii. p. 145.]

[Footnote 159: Müller's _Fertilisation of Flowers_, pp. 448, 455. Other cases of recent degradation and readaptation to insect-fertilisation are given by Professor Henslow (see footnote, p. 324).]

[Footnote 160: _The Colour Sense_, by Grant Allen, p. 95.]

[Footnote 161: _The Colour Sense_, chap. ix.]

[Footnote 162: See _Origin of Species_, sixth edition, p. 220.]