Of all the external characters of animals, the most beautiful, the most varied, and the most generally attractive are the brilliant colours and strange yet often elegant markings with which so many of them are adorned. Yet of all characters this is the most difficult to bring under the laws of utility or of physical connection. Mr. Darwin—as you are well aware—has shown how wide is the influence of sex on the intensity of coloration; and he has been led to the conclusion that active or voluntary sexual selection is one of the chief causes, if not the chief cause, of all the variety and beauty of colour we see among the higher animals. This is one of the points on which there is much divergence of opinion even among the supporters of Mr. Darwin, and one as to which I myself differ from him. I have argued, and still believe, that the need of protection is a far more efficient cause of variation of colour than is generally suspected; but there are evidently other causes at work, and one of these seems to be an influence depending strictly on locality, whose nature we cannot yet understand, but whose effects are everywhere to be seen when carefully searched for.
Although the careful experiments of Sir John Lubbock have shown that insects can distinguish colours—as might have been inferred from the brilliant colours of the flowers which are such an attraction to them—yet we can hardly believe that their appreciation and love of distinctive colours is so refined as to guide and regulate their most powerful instinct—that of reproduction. We are therefore led to seek some other cause for the varied colours that prevail among insects; and as this variety is most conspicuous among butterflies—a group perhaps better known than any other—it offers the best means of studying the subject. The variety of colour and marking among these insects is something marvellous. There are probably about ten thousand different kinds of butterflies now known, and about half of these are so distinct in colour and marking that they can be readily distinguished by this means alone. Almost every conceivable tint and pattern is represented, and the hues are often of such intense brilliance and purity as can be equalled by neither birds nor flowers.
Any help to a comprehension of the causes which may have concurred in bringing about so much diversity and beauty must be of value; and this is my excuse for laying before you the more important cases I have met with of a connection between colour and locality.
The influence of Locality on Colour in Butterflies and Birds.—Our first example is from tropical Africa, where we find two unrelated groups of butterflies belonging to two very distinct families (Nymphalidæ and Papilionidæ) characterized by a prevailing blue-green colour not found in any other continent.[29] Again, we have a group of African Pieridæ which are white or pale yellow with a marginal row of bead-like black spots; and in the same country one of the Lycænidæ (Leptena erastus) is coloured so exactly like these that it was at first described as a species of Pieris. None of these four groups are known to be in any way specially protected, so that the resemblance cannot be due to protective mimicry.
[29] Romaleosoma and Euryphene (Nymphalidæ), Papilio zalmoxis and several species of the Nireus-group (Papilionidæ).
In South America we have far more striking cases; for in the three subfamilies Danainæ, Acræinæ, and Heliconiinæ, all of which are specially protected, we find identical tints and patterns reproduced, often in the greatest detail, each peculiar type of coloration being characteristic of separate geographical subdivisions of the continent. Nine very distinct genera are implicated in these parallel changes—Lycorea, Ceratinia, Mechanitis, Ithomia, Melinæa, Tithorea, Acræa, Heliconius, and Eueides, groups of three or four (or even five) of them appearing together in the same livery in one district, while in an adjoining district most or all of them undergo a simultaneous change of coloration or of marking. Thus in the genera Ithomia, Mechanitis, and Heliconius we have species with yellow apical spots in Guiana, all represented by allied species with white apical spots in South Brazil. In Mechanitis, Melinæa, and Heliconius, and sometimes in Tithorea, the species of the Southern Andes (Bolivia and Peru) are characterized by an orange and black livery, while those of the Northern Andes (New Granada) are almost always orange-yellow and black. Other changes of a like nature, which it would be tedious to enumerate but which are very striking when specimens are examined, occur in species of the same groups inhabiting these same localities, as well as Central America and the Antilles. The resemblance thus produced between widely different insects is sometimes general, but often so close and minute that only a critical examination of structure can detect the difference between them. Yet this can hardly be true mimicry, because all are alike protected by the nauseous secretion which renders them unpalatable to birds.
In another series of genera (Catagramma, Callithea, and Agrias) all belonging to the Nymphalidæ, we have the most vivid blue ground, with broad bands of orange, crimson or a different tint of blue or purple, exactly reproduced in corresponding, yet unrelated species, occurring in the same locality; yet, as none of these groups are known to be specially protected, this can hardly be true mimicry. A few species of two other genera in the same country (Eunica and Siderone) also reproduce the same colours, but with only a general resemblance in the markings. Yet again, in tropical America we have species of Apatura which, sometimes in both sexes, sometimes in the female only, exactly imitate the peculiar markings of another genus (Heterochroa) confined to America: here, again, neither genus is protected, and the similarity must be due to unknown local causes.
But it is among islands that we find some of the most striking examples of the influence of locality on colour, generally in the direction of paler, but sometimes of darker and more brilliant hues, and often accompanied by an unusual increase of size. Thus in the Moluccas and New Guinea we have several Papilios (P. euchenor, P. ormenus, and P. tydeus) distinguished from their allies by a much paler colour, especially in the females which are almost white. Many species of Danais (forming the subgenus Ideopsis) are also very pale. But the most curious are the Euplœas, which in the larger islands are usually of rich dark colours, while in the small islands of Banda, Ké, and Matabello at least three species not nearly related to each other (E. hoppferi, E. euripon, and E. assimilata) are all broadly banded or suffused with white, their allies in the larger islands being all very much darker. Again, in the genus Diadema, belonging to a distinct family, three species from the small Aru and Ké islands (D. deois, D. hewitsonii, and D. polymena) are all more conspicuously white-marked than their representatives in the larger islands. In the beautiful genus Cethosia, a species from the small island of Waigiou (C. cyrene) is the whitest of the genus. Prothoë is represented by a blue species in the continental island of Java, while those inhabiting the ancient insular groups of the Moluccas and New Guinea are all pale yellow or white. The genus Drusilla, almost confined to these islands, comprises many species which are all very pale; while in the small island of Waigiou is found a very distinct genus, Hyantis, which, though differing completely in the neuration of the wings, has exactly the same pale colours and large ocellated spots as Drusilla.
Equally remarkable is the increase of size in some islands. The small island of Amboina produces larger butterflies than any of the much larger islands which surround it. This is the case with at least a dozen butterflies belonging to many distinct genera,[30] so that it is impossible to attribute the fact to other than some local influence. In Celebes, as I have elsewhere pointed out,[31] we have a peculiar form of wing and much larger size running through a whole series of distinct butterflies; and this seems to take the place of any speciality in colour.
[30] Ornithoptera priamus, O. helena, Papilio deiphobus, P. ulysses, P. gambrisius, P. codrus, Iphias leucippe, Euplœa prothoë, Hestia idea, Athyma jocaste, Diadema pandarus, Nymphalis pyrrhus, N. euryalus, Drusilla jairus.
[31] “Contributions to the Theory of Natural Selection,” pp. 168–173.
In a very small collection of insects recently brought from Duke-of-York Island (situated between New Britain and New Ireland) are several of remarkably white or pale coloration. A species of Euplœa is the whitest of all known species of that extensive genus; while a beautiful diurnal moth is much whiter than its ally in the larger island of New Guinea. There is also a magnificent longicorn beetle almost entirely of an ashy white colour.[32]
[32] These insects are described and figured in the “Proceedings of the Zoological Society,” for 1877, p. 139. Their names are Euplœa browni, Alcides aurora, and Batocera browni.
From the Fiji Islands we have comparatively few butterflies; but there are several species of Diadema of unusually pale colours, some almost white.
The Philippine Islands seem to have the peculiarity of developing metallic colours. We find there at least three species of Euplœa[33] not closely related, and all of more intense metallic lustre than their allies in other islands. Here also we have one of the large yellow Ornithopteræ (O. magellanus), whose hind wings glow with an intense opaline lustre not found in any other species of the entire group; and an Adolias[34] is larger and of more brilliant metallic colouring than any other species in the archipelago. In these islands also we find the extensive and wonderful genus of weevils (Pachyrhynchus), which in their brilliant metallic colouring surpass anything found in the whole eastern hemisphere, if not in the whole world.
[33] Euplœa hewitsonii, E. diocletiana, E. lætifica.
[34] Adolias calliphorus.
In the Andaman Islands in the Bay of Bengal there are a considerable number of peculiar species of butterflies differing slightly from those on the continent, and generally in the direction of paler or more conspicuous colouring. Thus two species of Papilio which on the continent have the tails black, in their Andaman representatives have them either red or white-tipped.[35] Another species[36] is richly blue-banded where its allies are black; while three species of distinct genera of Nymphalidæ[37] all differ from their allies on the continent in being of excessively pale colours as well as of somewhat larger size.
[35] Papilio rhodifer (near P. doubledayi), and Papilio charicles (near P. memnon).
[36] Papilio mayo.
[37] Euplœa andamanensis, Cethosia biblis, Cyrestis cocles.
In Madagascar we have the very large and singularly white-spotted Papilio antenor; while species of three other genera[38] are very white or conspicuous as compared with their continental allies.
[38] Danais nossima, Melanitis massoura, Diadema dexithea.
Passing to the West-Indian Islands and Central America (which latter country has formed a group of islands in very recent times) we have similar indications. One of the largest of the Papilios inhabits Jamaica,[39] while another, the largest of its group, is found in Mexico.[40] Cuba has two of the same genus whose colours are of surpassing brilliancy;[41] while the fine genus Clothilda—confined to the Antilles and Central America—is remarkable for its rich and showy colouring.
[39] Papilio homerus.
[40] P. daunus.
[41] P. gundlachianus, P. villiersi.
Persons who are not acquainted with the important structural differences that distinguish these various genera of butterflies can hardly realize the importance and the significance of such facts as I have now detailed. It may be well, therefore, to illustrate them by supposing parallel cases to occur among the Mammalia. We might have, for example, in Africa, the gnus, the elands, and the buffaloes, all coloured and marked like zebras, stripe for stripe over the whole body exactly corresponding. So the hares, marmots, and squirrels of Europe might be all red with black feet, while the corresponding species of Central Asia were all yellow with black heads. In North America we might have raccoons, squirrels, and opossums, in particoloured livery of white and black, so as exactly to resemble the skunk of the same country; while in South America they might be black with a yellow throat-patch, so as to resemble with equal closeness the tayra of the Brazilian forests. Were such resemblances to occur in anything like the number and with the wonderful accuracy of imitation met with among the Lepidoptera, they would certainly attract universal attention among naturalists, and would lead to the exhaustive study of the influence of local causes in producing such startling results.
One somewhat similar case does indeed occur among the Mammalia, two singular African animals, the Aard-wolf (Proteles) and the hyæna-dog (Lycaon), both strikingly resembling hyænas in their general form as well as in their spotted markings. Belonging as they all do to the Carnivora, though to three distinct families, it seems quite an analogous case to those we have imagined; but as the Aard-wolf and the hyæna-dog are both weak animals compared with the hyæna, the resemblance may be useful, and in that case would come under the head of mimicry. This seems the more probable because, as a rule, the colours of the Mammalia are protective, and are too little varied to allow of the influence of local causes producing any well-marked effects.
When we come to birds, however, the case is different; for although they do not exhibit such distinct marks of the influence of locality as do butterflies—probably because the causes which determine colour are in their case more complex—yet there are distinct indications of some effect of the kind, and we must devote some little time to their consideration.
One of the most curious cases is that of the parrots of the West-Indian Islands and Central America, several of which have white heads or foreheads, occurring in two distinct genera,[42] while none of the more numerous parrots of South America are so coloured. In the small island of Dominica we have a very large and richly-coloured parrot (Chrysotis augusta) corresponding to the large and richly-coloured butterfly (Papilio homerus) of Jamaica.
[42] Pionus albifrons and Chrysotis senilis (C. America), Chrysotis sallæi (Hayti).
The Andaman Islands are equally remarkable, at least six of the peculiar birds differing from their continental allies in being much lighter, and sometimes with a large quantity of pure white in the plumage,[43] exactly corresponding to what occurs among the butterflies.
[43] Kittacincla albiventris, Geocichla albigularis, Sturnia andamanensis, Hyloterpe grisola var., Ianthœnas palumboides, Osmotreron chloroptera.
In the Philippines this is not so marked a feature; yet we have here the only known white-breasted king-crow (Dicrurus mirabilis); the newly discovered Eurylæmus steerii, wholly white beneath; three species of Diceum, all white beneath; several species of Parus, largely white-spotted; while many of the pigeons have light ashy tints. The birds generally, however, have rich dark colours, similar to those which prevail among the butterflies.
In Celebes we have a swallow-shrike and a peculiar small crow allied to the jackdaw,[44] whiter than any of their allies in the surrounding islands; but otherwise the colours of the birds call for no special remark.
[44] Artamus monachus, Corvus advena.
In Timor and Flores we have white-headed pigeons,[45] and a long-tailed flycatcher almost entirely white.[46]
[45] Ptilopus cinctus, P. albocinctus.
[46] Tchitrea affinis, var.
In Duke-of-York Island east of New Guinea we find that the four new species figured in the “Proceedings of the Zoological Society,” for 1877, are all remarkable for the unusual quantity of white in their plumage. They consist of a flycatcher, a diceum, a wood-swallow, and a ground pigeon;[47] all equalling if not surpassing their nearest allies in whiteness, although some of these, from the Philippines, Moluccas and Celebes, are sufficiently remarkable in this respect.
[47] Monarcha verticalis, Diceum eximium, Artamus insignis, Phlogœnas johannæ.
In the small Lord Howe’s Island we have the recently extinct white rail (Notornis alba), remarkably contrasting with its allies in the larger islands of New Zealand.
We cannot, however, lay any stress on isolated examples of white colour, since these occur in most of the great continents; but where we find a series of species of distinct genera all differing from their continental allies in a whiter coloration, as in the Andaman Islands, Duke-of-York Island, and the West Indies, and, among butterflies, in the smaller Moluccas, the Andamans, and Madagascar, we cannot avoid the conclusion that in these insular localities some general cause is at work.
There are other cases, however, in which local influences seem to favour the production or preservation of intense crimson or a very dark coloration. Thus in the Moluccas and New Guinea alone we have bright red parrots belonging to two distinct families,[48] and which therefore most probably have been independently produced or preserved by some common cause. Here, too, and in Australia we have black parrots and pigeons;[49] and it is a most curious and suggestive fact that in another insular subregion—that of Madagascar and the Mascarene Islands—these same colours reappear in the same two groups.[50]
[48] Lorius, Eos (Trichoglossidæ), Eclectus (Palæornithidæ).
[49] Microglossus, Calyptorhynchus, Turacœna.
[50] Coracopsis, Alectrœnas.
Sense-perception influenced by Colour of the Integuments.—Some very curious physiological facts bearing upon the presence or absence of white colours in the higher animals have lately been adduced by Dr. Ogle.[51] It has been found that a coloured or dark pigment in the olfactory region of the nostrils is essential to perfect smell, and this pigment is rarely deficient except when the whole animal is pure white. In these cases the creature is almost without smell or taste. This, Dr. Ogle believes, explains the curious case of the pigs in Virginia adduced by Mr. Darwin, white pigs being killed by a poisonous root which does not affect black pigs. Mr. Darwin imputed this to a constitutional difference accompanying the dark colour, which rendered what was poisonous to the white-coloured animals quite innocuous to the black. Dr. Ogle, however, observes that there is no proof that the black pigs eat the root, and he believes the more probable explanation to be that it is distasteful to them; while the white pigs, being deficient in smell and taste, eat it and are killed. Analogous facts occur in several distinct families. White sheep are killed in the Tarentino by eating Hypericum crispum, while black sheep escape; white rhinoceroses are said to perish from eating Euphorbia candelabrum; and white horses are said to suffer from poisonous food where coloured ones escape. Now it is very improbable that a constitutional immunity from poisoning by so many distinct plants should, in the case of such widely different animals, be always correlated with the same difference of colour; but the facts are readily understood if the senses of smell and taste are dependent on the presence of a pigment which is deficient in wholly white animals. The explanation has, however, been carried a step further, by experiments showing that the absorption of odours by dead matter, such as clothing, is greatly affected by colour; black being the most powerful absorbent; then blue, red, yellow, and lastly white. We have here a physical cause for the sense-inferiority of totally white animals which may account for their rarity in nature; for few, if any, wild animals are wholly white. The head, the face, or at least the muzzle or the nose, are generally black; the ears and eyes are also often black; and there is reason to believe that dark pigment is essential to good hearing, as it certainly is to perfect vision. We can therefore understand why white cats with blue eyes are so often deaf, a peculiarity we notice more readily than their deficiency of smell or taste.
[51] “Medico-Chirurgical Transactions,” vol. liii. (1870).
If, then, the prevalence of white coloration is generally associated with some deficiency in the acuteness of the most important senses, this colour becomes doubly dangerous; for it not only renders its possessor more conspicuous to its enemies, but at the same time makes it less ready in detecting the presence of danger. Hence, perhaps, the reason why white appears more frequently in islands, where competition is less severe and enemies less numerous and varied. Hence, also, a reason why albinoism, although freely occurring in captivity, never maintains itself in a wild state, while melanism does. The peculiarity of some islands in having all their inhabitants of dusky colours (as the Galapagos) may also perhaps be explained on the same principles; for poisonous fruits may there abound which weed out all white- or light-coloured varieties, owing to their deficiency of smell and taste. We can hardly believe, however, that this would apply to white-coloured butterflies; and this may be a reason why the effect of an insular habitat is more marked in these insects than in birds or mammals.
It is even possible that this relation of sense-acuteness with colour may have had some influence on the development of the higher human races. If light tints of the skin were generally accompanied by some deficiency in the senses of smell, hearing, and vision, the white could never compete with the darker races so long as man was in a very low or savage condition, and wholly dependent for existence on the acuteness of his senses. But as the mental faculties became more fully developed and more important to his welfare than mere sense-acuteness, the lighter tints of skin and hair and eyes would cease to be disadvantageous whenever they were accompanied by superior brain-power. Such variations would then be preserved; and thus may have arisen the Xanthochroic race of mankind, in which we find a high development of intellect accompanied by a slight deficiency in the acuteness of the senses as compared with the darker forms.
Relations of Insular Plants and Insects.—I have now to ask your attention to a few remarks on the peculiar relations of plants and insects as exhibited in islands.
Ever since Mr. Darwin showed the immense importance of insects in the fertilization of flowers, great attention has been paid to the subject, and the relation of these two very different classes of natural objects has been found to be more universal and more complex than could have been anticipated. Whole genera and families of plants have been so modified as, first to attract and then to be fertilized by, certain groups of insects; and this special adaptation seems in many cases to have determined the more or less wide range of the plants in question. It is also known that some species of plants can be fertilized only by particular species of insects; and the absence of these from any locality would necessarily prevent the continued existence of the plant in that area.
In this direction, I believe, will be found the clue to much of the peculiarity of the floras of oceanic islands; since the methods by which these have been stocked with plants and with insects will be often quite different. Many seeds are, no doubt, carried by oceanic currents, others probably by aquatic birds. Mr. H. N. Moseley informs me that the albatrosses, gulls, puffins, tropic birds and many others, nest inland, often amidst dense vegetation; and he believes they often carry seeds, attached to their feathers, from island to island for great distances. In the tropics they often nest on the mountains far inland, and may thus aid in the distribution even of mountain-plants. Insects, on the other hand, are mostly conveyed by aerial currents, especially by violent gales; and it may thus often happen that totally unrelated plants and insects may be brought together, in which case the former must often perish for want of suitable insects to fertilize them. This will, I think, account for the strangely fragmentary nature of these insular floras, and the great differences that often exist between those which are situated in the same ocean; as well as for the preponderance of certain orders and genera.
In Mr. Pickering’s valuable work on the “Geographical Distribution of Animals and Plants” (founded on his researches during the United States exploring expedition), he gives a list of no less than sixty-six natural orders of plants unexpectedly absent from Tahiti, or which occur in many of the surrounding lands; some being abundant in other islands—as the Labiatæ at the Sandwich Islands. In these latter islands the flora is much richer, yet a large number of families which abound in other parts of Polynesia are totally wanting. Now much of the poverty and exceptional distribution of the plants of these islands is probably due to the great scarcity of flower-frequenting insects. Lepidoptera and Hymenoptera are exceedingly scarce in the eastern islands of the Pacific, and it is almost certain that many plants which require these insects for their fertilization have been thereby prevented from establishing themselves. In the western islands, such as the Fijis, several species of butterflies occur in tolerable abundance, and no doubt some flower-haunting Hymenoptera accompany them; and in these islands the flora appears to be much more varied, and especially to be characterized by a much greater variety of showy flowers, as may be seen by examining the plates of Dr. Seeman’s “Flora Vitiensis.”
Darwin and Pickering both speak of the great preponderance of ferns at Tahiti; and Mr. Moseley, who spent several days in the interior of the island, informs me that “at an elevation of from 2,000 to 3,000 feet the dense vegetation is composed almost entirely of ferns. A tree fern (Alsophila tahitensis) forms a sort of forest to the exclusion of almost every other tree, and, with huge plants of two other ferns (Angiopteris evecta and Aspelenium nidus), forms the main mass of the vegetation.” And he adds, “I have nowhere seen ferns in so great proportionate abundance.” This unusual proportion of ferns is a general feature of insular as compared with continental floras; but it has, I believe, been generally attributed to favourable conditions, especially to equable climate and perennial moisture. In this respect, however, Tahiti can hardly differ greatly from many other islands, which yet have no such vast preponderance of ferns. This is a question that cannot be decided by mere lists of species, since it is probable that in Tahiti they are less numerous than in some other islands where they form a far less conspicuous feature in the vegetation. The island most comparable with Tahiti in this respect is Juan Fernandez. Mr. Moseley writes to me:—“In a general view of any wide stretch of the densely clothed mountainous surface of the island, the ferns, both tree ferns and the unstemmed forms, are seen at once to compose a very large proportion of the mass of foliage.” As to the insects of Juan Fernandez, Mr. Edwyn C. Reed, who made two visits and spent several weeks there, has kindly furnished me with some exact information. Of butterflies there is only one (Pyrameis carie), and that rare—a Chilian species and probably an accidental straggler. Four species of moths of moderate size were observed (all Chilian), and a few larvæ and pupæ. Of bees there were none, except one very minute species (allied to Chilicola), and of other Hymenoptera a single specimen of Ophion luteus a cosmopolitan ichneumon. About twenty species of flies were observed, and these formed the most prominent feature of the entomology of the island.
Now, as far as we know, this extreme entomological poverty agrees closely with that of Tahiti; and there are probably no other portions of the globe equally favoured in soil and climate, and with an equally luxuriant vegetation, where insect-life is so scantily developed. It is curious, therefore, to find that these two islands also agree in the wonderful predominance of ferns over the flowering plants—in individuals even more than in species; and there is no difficulty in connecting the two facts. The excessive minuteness and great abundance of fern-spores causes them to be far more easily distributed by winds than the seeds of flowering plants; and they are thus always ready to occupy any vacant places in suitable localities, and to compete with the less vigorous flowering plants. But where insects are so scarce, all plants which require insect-fertilization, whether constantly to enable them to produce seed at all or occasionally to keep up their constitutional vigour by crossing, must be at a great disadvantage; and thus the scanty flora which oceanic islands must always possess, peopled as they usually are by waifs and strays from other lands, is rendered still more scanty by the weeding out of all such as depend largely on insect-fertilization for their full development. It seems probable, therefore, that the preponderance of ferns in islands (considered in mass of individuals rather than in number of species) is largely due to the absence of competing phænogamous plants, and that this is in great part due to the scarcity of insects. In other oceanic islands, such as New Zealand and the Galapagos, where ferns, although tolerably abundant, form no such predominant feature in the vegetation, but where the scarcity of flower-haunting insects is almost equally marked, we find a great preponderance of small, green, or otherwise inconspicuous flowers, indicating that only such plants have been enabled to flourish there as are independent of insect-fertilization. In the Galapagos (which are perhaps even more deficient in flying insects than Juan Fernandez) this is so striking a feature that Mr. Darwin speaks of the vegetation as consisting in great part of “wretched-looking weeds,” and states that “it was some time before he discovered that almost every plant was in flower at the time of his visit.” He also says that he “did not see one beautiful flower” in the islands. It appears, however, that Compositæ, Leguminosæ, Rubiaceæ, and Solanaceæ form a large proportion of the flowering plants; and as these are orders which usually require insect-fertilization, we must suppose, either that they have become modified so as to be self-fertilized, or that they are fertilized by the visits of the minute Diptera and Hymenoptera which are the only insects recorded from these islands.
In Juan Fernandez, on the other hand, there is no such total deficiency of showy flowers. I am informed by Mr. Moseley that a variety of the Magnoliaceous winter-bark abounds and has showy white flowers, and that a Bignoniaceous shrub with abundance of dark blue flowers was also plentiful; while a white-flowered Liliaceous plant formed large patches on the hill-sides. Besides these, there were two species of woody Compositæ with conspicuous heads of yellow blossoms, and a species of white-flowered myrtle also abundant; so that, on the whole, flowers formed a rather conspicuous feature in the aspect of the vegetation of Juan Fernandez.
But this fact—which at first sight seems entirely at variance with the view we are upholding of the important relation between the distribution of insects and plants—is well explained by the existence of two species of humming-birds in Juan Fernandez, which, in their visits to these large and showy flowers, fertilize them as effectually as bees, moths, or butterflies. Mr. Moseley informs me that “these humming-birds are extraordinarily abundant, every tree or bush having one or two darting about it.” He also observed that “nearly all the specimens killed had the feathers round the base of the bill and front of the head clogged and coloured yellow with pollen.” Here, then, we have the clue to the perpetuation of large and showy flowers in Juan Fernandez; while the total absence of humming-birds in the Galapagos may explain why no such large-flowered plants have been able to establish themselves in those equatorial islands.
This leads to the observation that many other groups of birds also, no doubt, aid in the fertilization of flowers. I have often observed the beaks and faces of the brush-tongued lories of the Moluccas covered with pollen; and Mr. Moseley noted the same fact in a species of Artamus, or swallow-shrike, shot at Cape York, showing that this genus also frequents flowers and aids in their fertilization. In the Australian region we have the immense group of the Meliphagidæ, which all frequent flowers; and as these range over all the islands of the Pacific, their presence will account for a certain proportion of showy flowers being found there, such as the scarlet Metrosideros, one of the few conspicuous flowers in Tahiti. In the Sandwich Islands, too, there are forests of Metrosideros; and Mr. Charles Pickering writes me, that they are visited by honey-sucking birds, one of which is captured by sweetened bird-lime, against which it thrusts its extensile tongue. I am also informed that a considerable number of flowers are occasionally fertilized by humming-birds in North America; so that there can, I think, be little doubt that birds play a much more important part in this respect than has hitherto been imagined. It is not improbable that in Tropical America, where the humming-bird family is so enormously developed, many flowers will be found to be expressly adapted to fertilization by them, just as so many in our own country are specially adapted to the visits of certain families or genera of insects.[52]
[52] The probable influence of fertilization by birds on the flowers of the Auckland Isles has been referred to at p. 238. Mr. Darwin, in his book on Cross and Self-Fertilisation of Plants (p. 371), gives in a note numerous cases in which birds are known to fertilise flowers, the most important being that of several species of Abutilon in South Brazil, which, according to Fritz Müller, are sterile unless fertilised by humming-birds. This proves, not only that birds fertilise flowers in the same manner as insects, but that the two classes of organisms have become so correlated as to be mutually necessary to each other; and it completely justifies us in imputing the fertilization of flowers to flower-frequenting birds wherever these are present and suitable insects are notoriously scarce, as is the case in so many of the islands here referred to.
It must also be remembered, as Mr. Moseley has suggested to me, that a flower which has acquired a brilliant colour to attract insects might, on transference to another country and becoming so modified as to be capable of self-fertilization, retain the coloured petals for an indefinite period. Such is probably the explanation of the Pelargonium of Tristan d’Acunha, which forms masses of bright colour near the shore during the flowering season; while most of the other plants of the island have colourless flowers in accordance with the almost total absence of winged insects. The presence of many large and showy flowers among the indigenous flora of St. Helena must be an example of a similar persistence. Mr. Melliss indeed states it to be “a remarkable peculiarity that the indigenous flowers are, with very slight exceptions, all perfectly colourless;”[53] but although this may apply to the general aspect of the remains of the indigenous flora, it is evidently not the case as regards the species, since the interesting plates of Mr. Melliss’s volume show that about one third of the indigenous flowering plants have more or less coloured or conspicuous flowers, while several of them are exceedingly showy and beautiful. Among these are a Lobelia, three Wahlenbergias, several Compositæ, and especially the handsome red flowers of the now almost extinct forest-trees, the ebony and redwood (species of Melhania, Byttneriaceæ). We have every reason to believe, however, that when St. Helena was covered with luxuriant forests, and especially at that remote period when it was much more extensive than it is now, it must have supported a certain number of indigenous birds and insects, which would have aided in the fertilization of these gaily-coloured flowers. The researches of Dr. Hermann Müller have shown us by what minute modifications of structure or of function, many flowers are adapted for partial insect and self-fertilization in various degrees; so that we have no difficulty in understanding how, as the insects diminished and finally disappeared, self-fertilization may have become the rule, while the large and showy corollas remain to tell us plainly of a once different state of things.