The Polynesian species are for the most part restricted to the Pacific islands, but there are only two species that range over the South Pacific as far east as Tahiti, namely, Ficus prolixa, the Tahitian banyan, and F. tinctoria. Some species are confined to Western Polynesia, such as F. obliqua, the Fijian banyan, F. scabra, and F. aspera, the last occurring in East Australia. Among the individual groups Fiji possesses probably fourteen or fifteen species, of which, perhaps, a third would be peculiar. According to Dr. Warburg, as cited in Dr. Reinecke’s paper, Samoa owns eight species, of which six may be endemic. In Rarotonga and Tahiti we find only F. prolixa and F. tinctoria. The species in the groups where they are best represented belong to three or four sections of the genus.
The banyans of the South Pacific are represented by three or four species, namely, Ficus prolixa, the Tahitian banyan, found all over the tropical groups of the South Pacific from the New Hebrides and New Caledonia to Tahiti, the Marquesas and Pitcairn Island (Maiden); F. obliqua, the Fijian banyan, confined to the islands of the Western Pacific from the New Hebrides to Tonga; and two new banyans in Samoa, as described by Dr. Warburg in Dr. Reinecke’s paper. In my paper on Polynesian plant-names it is shown that the banyans possess two names in the Pacific, one being “aoa,” the Polynesian name, found in all the groups from Samoa eastward, and connected linguistically with the Malayan and Malagasy banyan-words; the other, the Melanesian name typified in the Fijian “mbaka,” and represented in a variety of forms in the New Hebrides and neighbouring groups.
It is probable that the Pacific islanders have assisted in the dispersal of one or two of the species of Ficus, such as F. tinctoria, which they employ for different purposes, but, generally speaking, birds are active agents in distributing the genus. I need scarcelysay that the agency of the currents is quite insufficient to explain the distribution of Ficus. When in Fiji I experimented on three or four different species of Ficus belonging to the sections of the genus there represented. The fruits may float at first, but within a week or ten days they break down, and the seeds escape and sink. Beneath a tree of F. scabra growing on the banks of the Wai Tonga in Viti Levu, I noticed a number of its fruits floating in a sodden condition among the reeds at the river-side.
It is with the banyans that the dispersal of the seeds by frugivorous birds becomes most evident. This is at once indicated by the frequent occurrence of these trees in the interior of coral islets in the Western Pacific, as in Fiji and in the Solomon Islands. Fruit-pigeons roost in their branches, and birds shot on these islets often contain the fruits in their crops (Bot. Chall. Exped., iv, 310). The process may also be seen in operation in Krakatoa. Professor Penzig found in 1897 that three species of Ficus had established themselves there since the eruption of 1883 through the agency of frugivorous birds. Besides pigeons, we find that parrots, hornbills, honey-eaters, &c., feed on these fruits, and I possess a large number of references to this subject. The Messrs. Layard in New Caledonia, Dr. Meyer in Celebes, Mr. Everett in Borneo, Dr. Forbes in Sumatra, and several other contributors toIbismight be here mentioned. Dr. Beccari, in hisWanderings in the Great Forests of Borneo, speaks of “the facile dissemination of the various species of Ficus through the agency of birds,” and he arrives at certain important conclusions which are discussed inChapter XXXIII.
I have before alluded to the absence of Ficus from Hawaii. This group possesses the Honey-Eaters (Meliphagidæ), birds well suited for dispersing species of Ficus over Polynesia; but this family of birds is only represented by peculiar genera in Hawaii, and therein lies the explanation. At the time when the Honey-Eaters roamed over Polynesia, the genus Ficus had not arrived from Malaya. The connection between the bird and the plant is well shown on Fernando Noronha, which possesses a peculiar species of Ficus and a peculiar species of dove, the only fruit-eating bird in the island (Ridley).
Generally speaking, all the “difficult” genera which puzzle the student of plant-dispersal in Fiji and Hawaii are absent from the Tahitian region. Those with stone-fruits and with large seeds,where the stone or seed is an inch in size and over, are absent from Tahiti. Thus the genera Canarium, Dracontomelon, Myristica, Sterculia, and others, of which the three first-named are known to be dispersed by fruit-pigeons, have not advanced into the Pacific eastward of the Fijian region. We miss in the Tahitian islands the large-fruited palms of Fiji, such as the Veitchias with fruits two to two and a half inches (5 to 6 cm.) long, and we find in their place a Ptychosperma, evidently very rare, and the widely spread Pritchardia pacifica, that may have been introduced by man, both with drupes not far exceeding half an inch (1·2 cm.) in size. The islands of the Tahitian region also lack the Coniferæ; and genera like Dammara, Dacrydium, and Podocarpus that give such a character to the Fijian forests are not to be found. In this region we do not find many of the large-seeded Leguminous genera, such as Cynometra, Storckiella, and Afzelia, that occur in Fiji, the only large-seeded genera that it possesses being such as are brought by the currents, namely, Mucuna, Strongylodon, Cæsalpinia. The difficulties presented by the occurrence of the inland species of Canavalia and Mezoneuron in Hawaii do not offer themselves in Tahiti (seeChapter XV). Tahiti also lacks, as often before observed, the mangroves and most of the plants of the mangrove-formation.
As above remarked, the Fijian trees with large “stones” and heavy seeds an inch in size are not to be reckoned amongst the indigenous Tahitian plants, “size” being an important determining factor in the exclusion. The occurrence of Elæocarpus in Rarotonga presents no real difficulty, as I have explained inChapter XXVI.An apparent exception is presented by the existence in Tahiti of Calophyllum spectabile, where the stones are about an inch across; but since its fruits can float in sea-water for nearly a month, and on account of the value placed on its timber by the Polynesians, we cannot altogether exclude the agencies of man and the currents. One seeming exception is also offered by the presence of Serianthes myriadenia, a tree which in Fiji grows both in the forests and on the banks of the tidal estuaries. Its seeds, which are six to seven-tenths of an inch (15 to 18 mm.) in length, have no buoyancy, and the pods float only two or three weeks. The case of Lepinia tahitensis is alluded to elsewhere, but it may be added that these and other difficulties await further investigation.
A great many Fijian plants are not found in the Tahitian region, such as Micromelum, those of the order Meliaceæ, the Melastomaceous genus Medinilla, Myrmecodia, Ophiorrhiza, &c.,which are often quite as well fitted for over-sea transport as are several of the plants already established there. But it should be remembered that crowding out would often come into play in such a contracted region. The area, however, has been very generously dealt with as regards plant genera. Though the total land-surface cannot be more than one-fourth or one-third that of Fiji or Hawaii, it possesses more than half the number of genera found in Fiji, and four-fifths of the number found in Hawaii.
We have already in some degree dealt with Fiji in so far as the partial dispersal of genera over the Pacific islands is concerned. We have seen that it possesses very few genera (not a score in all) in common with Hawaii that are not found in the Tahitian region, and it is assumed that in most cases such genera reached Hawaii independently and not through the South Pacific. On the other hand, excluding the grasses, sedges, and vascular cryptogams, Fiji owns in common with Tahiti between sixty and seventy genera that do not occur in Hawaii. This shows unmistakably the trend of plant migration in the Pacific islands. Several interesting features in plant-distribution have been already brought out, and notably the fact that Indo-Malayan genera with large seeds or “stones” an inch in size have been arrested in the Fijian region in their passage into the South Pacific. Thus Canarium, Dracontomelon, Myristica, and Sterculia have not extended eastward of the Fijian area.
Yet a very large proportion of the Fijian genera, quite half of the total number, are not represented either in the Tahitian or in the Hawaiian region; and of many of them it is obvious that they are as well fitted to be carried over the Pacific as are those that have actually reached Tahiti and Hawaii. Take, for instance, Begonia, which has not extended east of Fiji, though Hillebrandia, a genus of the order, is peculiar to Hawaii. Nor can we explain why with three genera like Geissois, Dolicholobium, and Alstonia, possessing seeds dispersed by the winds, only the last-named has passed beyond Fiji. However, as before remarked, it is probable that lack of opportunity rather than capacity for dispersal has determined the matter, and we must, therefore, assume that many of the genera havehalted in the Fijian region because they entered the Pacific after the age of active general dispersal over that ocean.
Occasionally we notice in this region that which we have observed in the case of Cyrtandra in different Pacific groups, namely, a sudden development of what Hillebrand terms “formative energy” in a genus, such as we find in the case of Elatostema in Samoa, and in that of Psychotria in Fiji and Samoa. The principle of polymorphism in the development of species is also illustrated by Micromelum and by Limnanthemum. In the last case we possess a typical polymorphous species in Limnanthemum indicum that has played in this respect therôleof Naias marina in the warm waters of the globe.
With several genera that like Gnetum, Myristica, and Sterculia occur both in the Old and the New World, it is evident that in explaining their distribution we are dealing with something more than questions of means of dispersal. With these genera, and with others like Lindenia, it seems almost futile to talk of means of dispersal, when to all appearance their existing distribution is but the remnant of an age of general dispersion over the greater part of the warm regions of the world. These genera, with others, might be cited in favour of the continental hypothesis relating to the islands of the Western Pacific. Trees with stone-fruits, such as Canarium, Couthovia, Dracontomelon, and Veitchia, where the stones are an inch and more in length, might be also adduced by some in evidence of this theory. But in these cases the lesson of Elæocarpus (Chapter XXVI) should always be remembered, since the “stones” of drupes may vary greatly in size amongst the different species of a genus, and species seemingly “impossible” from the standpoint of dispersal in one group may be represented in other groups by species where the size of the “stone” presents no difficulty in attributing the dispersal of the genus to frugivorous birds.
The problem connected with the presence of this genus in Fiji is but a part of the still more difficult problem connected with the dispersal of the genus over the tropics. The riddle presented by the Fijian species seems, indeed, difficult enough; but it merely presents in miniature the great mystery surrounding the whole genus. According to theIndex Kewensisno other species have been found in oceanic islands except those occurring in the Western Pacific, as in Fiji, the New Hebrides, and New Caledonia, and most of these seem to be confined to those islands. We havehere a genus that repeats the Dammara difficulty of the Western Pacific.
The trees are common in places in the Vanua Levu forests, where the large, woody, open follicles may be seen lying in numbers on the ground, empty and in all stages of decay. The seeds of one species, near Sterculia vitiensis, were nearly an inch long and sank like stones. The unopened follicles will float for weeks; but it is evident that Nature does not disperse the genus in this fashion, since the fruits before dehiscence remain on the tree. It is also noteworthy that Gaudichaud, when describing the floating drift of the Molucca seas, refers to the open follicles of two or three species of Sterculia (Bot. Chall. Exped., iii, 279). The fruits never came under my notice in the drift of Fiji. The seeds of a Fijian species examined by me were four-fifths of an inch (2 cm.) long. They had a thin, brittle, outer skin and crustaceous inner test, and, being edible, might attract birds; but such birds would be ground feeders, like the Megapod, and the Goura pigeon of New Guinea, and the Nicobar pigeon, birds of this habit being rare in Fiji. I should doubt whether the seeds are sufficiently protected to be preserved from injury in a bird’s stomach during a long sea-passage; and they may thus be placed in the same category with the seeds of Myristica, a genus that has also failed to reach Tahiti and Hawaii.
But the distribution of Sterculia raises other more important questions than that connected with its occurrence in Fiji, which involves an over-sea passage of only 500 or 600 miles. As in Podocarpus amongst the Coniferæ, which has a similar distribution in the Western Pacific, we have to explain the existence of the genus in the three great continental masses of Africa, Asia, and America, now separated by oceans several thousands of miles across. Here also we must look far back into the ages for a common centre of diffusion in the extreme north, such as is in a sense suggested by the occurrence of the order in the Eocene beds of Europe.
As showing unmistakably that Fiji received its species from the Old World, it may be observed that one of its trees, Sterculia vitiensis, is very closely allied to S. fœtida, widely spread in tropical Asia, in Malaya, and Australia, as well as in Africa.
There are only two species of this tree recorded in theIndex Kewensis, one in Java, and one in Fiji as well as in Samoa. Thefruit is a capsule with small, flat seeds, margined by long hairs, that might possibly attach themselves to a bird’s feathers.
This small genus of tropical Asia, Malaya, tropical Australia and the islands of the Western Pacific, has one species, Micromelum pubescens, possessing the range of the genus with other species that are restricted to different localities. We thus have apparently another illustration of the part played by a wide-ranging polymorphous plant in providing new species. The red berries would easily attract frugivorous birds; but the seed-tests seem too delicate to allow the seeds to remain more than a few hours in a bird’s stomach without injury.
This tree, which is cultivated in many places in tropical Asia and Malaya, but is certainly indigenous, according to the authors of theFlora Indica, in Ava and Tenasserim, has apparently extended into the Pacific by cultivation. But though much valued by the natives on account of its fragrant flowers, and in consequence often planted by them near their villages, it grows in some localities in Fiji and Samoa as an indigenous plant. The berries are especially suited for dispersal by frugivorous birds, their flat seeds, 8 mm. in length, possessing hard crustaceous tests that would enable them to pass unharmed in a bird’s droppings. According to Reinecke the fruits are sought after by pigeons, and particularly by Didunculus strigirostris, the Samoan Tooth-Billed Pigeon. The tree has not travelled eastward of Tonga and Samoa, with the exception of its occurrence in Rarotonga; and according to Mr. Cheeseman the Rarotongans received it from Samoa several years ago.
This genus of seven or eight known species is found in Australia, New Caledonia, the New Hebrides, and Fiji. Since New Caledonia possesses four species, it may be considered the home of the genus. To the Fijian endemic species, G. ternata, I paid special attention. The capsules dehisce on the tree and allow the small seeds to escape. These seeds, which are very light, 150 to 200 going to a grain, are 3 to 4 mm. long and are winged at one end. They could no doubt be carried some distance by strongwinds; but they possess no buoyancy. Large bats probably aid in their dispersal. The Fijians assert that these animals are in the habit of visiting the trees for the sake of the honey furnished by the conspicuous red flowers. When they see a bat flying towards these trees, they are wont to remark that it is going to drink the “se ni vota,” that is, to suck the flowers of the Vota tree. It is very likely that seeds would sometimes be carried in their fur for considerable distances.
Before the discovery of Hillebrandia, a new genus of the Begoniaceæ, in Hawaii, the order was not known from Polynesia. However, in 1878 Mr. Horne collected a species of Begonia in Fiji, and it was probably this species that frequently came under my notice in the rain-forests of the Vanua Levu mountains. In 1883 I collected a Begonia in the Solomon Islands, which I gave to Baron F. von Mueller, who informed me that it was the first record of the genus east of New Guinea, the description of Mr. Horne’s Fijian plant apparently not having been published (see Guppy’sSolomon Islands, p. 288). It is not easy to explain why a genus with such minute seeds, which are apparently as well fitted for dispersal as those of the orchids, should have such a limited distribution in the Pacific.
In theIndex Kewensisthis genus, containing five species, is restricted to Fiji. It must, however, be more generally distributed in the Western Pacific, since the genus was identified at Kew among my Solomon Island collections, and it is recorded in the list given in my book on that group (pages 283, 288, 297).
The showy, large, white, fragrant flowers of these small trees recall those of Lindenia, with which Dolicholobium is often associated in Fiji by the sides of streams and rivers. As Horne observes, the Fijian Dolicholobiums range from the sea-shores and the heads of the estuaries to the tops of the highest mountains. As noticed by me in the Solomon Islands they affected the same station, being especially common on the banks of streams. The genus has a long, narrow capsule six inches or more in length. The linear seeds, though very light, are an inch or more long, the coats being drawn out into a long tail at either end, and thus differing greatly from those of Lindenia, the other Rubiaceous genus, with which these plants are so frequently associated at theriver-side. I can only suppose that the seeds are transported by the winds. The history of the genus is suggested in my remarks on Lindenia.
Respecting its distribution in the Pacific, this genus of showy river-side shrubs takes the same place amongst the plants that Galaxias takes among the fishes. It is full of mystery. Of the four species known, two grow on the river-banks of Central America and two in similar stations in the islands of the Western Pacific. Of the last-named both occur in New Caledonia, one of them being endemic, whilst the other, Lindenia vitiensis, is found also in Fiji and Samoa. Reinecke seemingly records no Samoan species, but in the list of additions at the end of hisFlora Vitiensis, Seemann refers to the Fijian species as having been found in Samoa by Dr. Graeffe.
Lindenia vitiensis, as Horne aptly remarks, adorns the rocky banks of many Fijian streams with its cream-coloured flowers, which impregnate the air with their sweet odour. I found it in Vanua Levu, both at the heads of the estuaries and beside the stream and the torrent in the heart of the mountains. It was often associated with a species of Dolicholobium, which it resembled strangely in its large, showy, scented flowers and in the form of the leaf. Seemann says it is also accompanied at the river-side in Viti Levu by Ficus bambusæfolia and Acalypha rivularis. It is noteworthy that all the four plants here mentioned as being associated river-side plants in Fiji possess the long, narrow leaves of the willow type, a subject that is discussed innote 79.
The capsules of Lindenia vitiensis contain numbers of small, angular seeds about 1·5 mm. across, some 400 of them when well dried going to a grain. The seeds float buoyantly by reason of their outer covering of crisp, air-bearing, cellular tissue. When this outer covering is stripped off, the minute nucleus, or seed proper, which is barely a millimetre across and is but slightly protected, sinks at once. As the seeds float on the surface of a stream they might readily get on the plumage of an aquatic bird; but they have no special means of attachment; though, if they dried on the feathers they might adhere to some extent. That they could be carried in mud adhering to a bird across an ocean’s breadth I think most unlikely; and it should be remembered in this connection that only the dead or sickly seeds would be found at the bottom of a stream.
The most reasonable explanation of the extraordinary distribution of Lindenia is that it was in a past age found over the tropical regions of both America and the Old World, and that it has died out over the greater part of its original area. To study the means of dispersal of plants with such a distribution seems almost futile. I am inclined to think that the limited range of Dolicholobium, so frequently its station-companion in Fiji, may be similarly explained.
This interesting genus of aquatic plants is dispersed over the tropical and temperate regions of the globe, but with the exception of Fiji and the New Hebrides it is not found in oceanic groups, though it occurs in large continental islands like New Caledonia and Cuba. About twenty species are enumerated in theIndex Kewensis, but it is stated in theGenera Plantarumthat they can probably be reduced to ten, the reduction being chiefly applicable to the tropical species, nearly all of which are reducible to varieties of L. indicum, the temperate species being often very distinct. It would thus appear that although dispersal is still active in the tropics, it is in part suspended in the temperate zone, and we seem to possess in L. indicum a typical polymorphous species that has played therôleof Naias marina in the warm, fresh waters of the globe (see page368).
Although some of the temperate species, like Limnanthemum nymphæoides in Europe and Northern Asia, have a wide range, it is probable that this is connected not so much with means of dispersal, as with its relation to present and past drainage-areas. Rivers in the lapse of ages change their courses and carry their aquatic floras with them, leaving, however, a few of their plants around the springs and in the lakes which serve still as centres of dispersal. Rivers may even exchange their plants in flood-time in extensive level districts. Nor is the occurrence of the genus in the Old and New Worlds in the northern hemisphere to be connected with questions of dispersal across an ocean. Except in the case of small-seeded plants, like Nasturtium and Lythrum, where the dispersal could be carried on by water-fowl, the plant-species being often identical on both sides of the Atlantic, it is probable that most of the large-seeded river-side genera common to Europe and North America, such as Iris and Acorus, had in past ages their home in the extreme north, whence the plants spread as from afocus into the continents of America and Eurasia. It is also to be doubted whether even in the tropics there has been much over-sea dispersal of Limnanthemum without the aid of man, and reasons will be given for the belief that probably in Fiji, in the New Hebrides, and in New Caledonia the seeds of the first plants were unintentionally introduced by the aborigines.
Following Bentham we may regard the species of the Western Pacific Islands as a form of the wide-ranging Limnanthemum indicum. These plants in Fiji do not play the part in river-vegetation that they do in the temperate regions, as for instance in the Upper Thames. They are not common except in places, and seem to be chiefly confined to Viti Levu, particularly to ponds in the Rewa delta, where theirrôleis that of an Indian tank plant. In the Rewa delta they may be sometimes seen thriving in brackish water having a density of 1·005.
Looking at the mode of dispersal to which the Limnanthemums owe their existence in the Western Pacific, we cannot disregard, especially in Fiji, the possibility of the seeds having been unintentionally transported by the natives when they carried in their migrations their edible tubers, such as Colocasia antiquorum, Alocasia indica, and Cyrtosperma edulis, that are cultivated in wet places. It is in the ponds around which these plants grow that the Limnanthemums thrive. The Chinese, with their peculiar methods of cultivation, are now carrying with them strange water-plants over the warmer regions of the globe; and it would be surprising if the Pacific islanders in their migrations did not do the same. If such an introduction, however, took place, it must have happened before the time of Captain Cook, when the plant was found in New Caledonia. (It may be remarked in this connection that the seeds of the genus will germinate after being kept dry for years. Seeds of the British species which I had kept dry for two and a half years germinated healthily when placed in water.)
Some years ago I ascertained that the seeds of the British plants were enabled, by means of their fringe of hairs, to attach themselves firmly to the downy plumage of a bird’s breast. This could not happen with the Fijian plant as the seeds are naked, and the same may be said of some species described by Gray and Chapman as widely spread over the United States. The seeds of the genus appear quite unsuited for safe transport inside the body of a bird. The Fijians give the plants a variety of names, nearly all of which are associated with the word for a duck, and none of them bear an ancient impress. Thus we find such names as“Ndambe-ndambe-ni-nga” and “Vothe-vothe-ni-nga,” meaning respectively “the duck’s seat” and “the duck’s paddle.”
This wonderful aquatic has been dispersed over most of the globe; but I will only mention its occurrence in oceanic islands, such as Fiji, Samoa, the Bermudas, and the Azores, to indicate the necessity of attributing its distribution in islands to birds. Several years ago I made a careful study in England of the habits and mode of germination of this plant, the results of which are given inScience Gossipfor November, 1894; but reference can only be made here to such points as bear on the occurrence of the plant in the Pacific islands.
It is well known that in our English ponds and rivers the plant propagates itself, as a rule, by budding; and that it is only in unusually hot and dry summers, such as that of 1893, when many ponds became very low and were excessively heated, that the fruits mature in any quantity. My observations clearly showed that a higher temperature is required for the completion of maturation than for the early stage of the fruiting process and for the flowering. After a comparison of my river and pond temperatures, I formed the conclusion that whilst in water 12 to 18 inches deep this plant requires for a week or more an average daily maximum water temperature of 70° F. to produce its flowers, a warmth of 80° and over is necessary to mature its fruit, a condition to be found in England only in shallow ponds, where the plants may fruit abundantly, but not in rivers, where they flower and rarely mature the fruit (see also for the thermometric conditions my paper inProc. Roy. Phys. Soc. Edin., xii, 296). Since a yet lower temperature (an average maximum water temperature of 66° for a week or more) is sufficient for germination, it follows that the thermal conditions of our English climate will allow Ceratophyllum to germinate and to flower, though but rarely to mature the fruit.
Even in Fiji we can notice the distinction between the cooler river and the superheated ponds and swamps of the Rewa delta as regards the maturation of the fruit. In 1897 I found Ceratophyllum thriving in the main channel of the Lower Rewa where the water was quite fresh; whilst lower down where the water was often brackish its place was taken by Ruppia maritima. In the main river, where the water unmixed with sea-water rarely acquiresa temperature of 80° F., the reading being usually 78° to 79°, I never found the plants in fruit, and it is only in the superheated shallow waters of the swamps and back-waters that they mature their fruits.
Since Ceratophyllum even in tropical climates would probably only mature its fruits in the superheated waters of shallow ponds, tanks, and ditches, it follows that its dispersal by birds is confined to warm regions. In the cold waters of the Siberian lakes and rivers it would never mature its seeds, and could only be propagated by budding. If it existed in the head-springs of the sources of a river in these latitudes, it would be distributed by means of its floating shoots and fragments along the length of the river basin, and in the times of flood it might pass in the lower plains from one river system to another. When rivers changed their courses it would be left behind in the lakes and ponds and springs, and would also be carried away to the new region. In this manner it would in the course of ages be distributed over a continent without the aid of seed, propagating itself in a vegetative fashion.
In the case of oceanic islands, however, we have to appeal to the seed. Since the fruits sink in sea-water even after prolonged drying, and since a few days’ immersion in sea-water, as I found, kills the floating plant, we are driven to the agency of birds. The fruits, which without appendages are a quarter of an inch (6 mm.) in length, are too large and heavy to be carried in dry mud adhering to birds. The chances of their becoming entangled in a bird’s feathers by means of their basal spines and terminal style seem small, since they would be lying usually on the mud under the water. They are quite fitted for safe transport in the stomach and intestines of birds, such as is established inChapter XXXIIIfor Potamogeton and Sparganium in the case of ducks. As my experiments show, drying for a period of three months does not injure the germinating capacity of the seeds.
This is a genus accredited in theIndex Kewensiswith eight species, of which three belong to Borneo, one to Sumatra, one to Java, one to the Philippines, and two to Fiji, all the species being restricted in their range. My observations were confined to D. vitiense, Engler (D. sylvestre in Seemann’s work), the Tarawau of the Fijians, who regard it as a tree that is planted by the dead in Naithombothombo, the place of departed spirits, according to thelegend given by Hazlewood in his Fijian Dictionary. Its method of dissemination in the Fijian forests is, however, far more prosaic. Pigs and fruit-pigeons assist in the dispersal of the seeds in these islands. Pigs are often found in the vicinity of a Tarawau tree; and evidently they much appreciate the fallen fleshy fruits, which are about11⁄3inch (3·3 cm.) across and inclose a large stone7⁄8inch (2·2 cm.) in diameter. The entire fruit and the detached stone sink in sea-water, the last floating only a few hours, even after drying for four years. Mr. Hemsley regards the genus as probably dispersed by the currents, since a stone was found amongst the floating drift collected by theChallengerExpedition off the coast of New Guinea. The stone, however, is described as seedless, which may explain its buoyancy. It is, however, to the fruit-pigeon that we must look for the dispersal of this genus. In the crop of one of these birds shot in Fiji I found the entire fruit of a Tarawau tree.
This genus of trees, to which nearly a hundred species are referred in theIndex Kewensis, belongs mainly to tropical Asia and Malaya, a few species occurring in tropical Africa, Madagascar, the Mascarene Islands, and Polynesia. Its great home is in Malaya, to which two-thirds of the species are confined; but its distribution in the oceanic islands of the Indian and Pacific Oceans is especially interesting, Mauritius, Bourbon, Fiji, Tonga, and Samoa (Horne) each possessing a species.
The large drupes of the genus, as I found in Fiji, have no capacity for dispersal by currents; and we are, therefore, compelled to appeal to the agency of the frugivorous bird. Yet to a person unaccustomed to the ways of fruit-pigeons the transportation across a broad tract of ocean of large heavy “stones,” an inch and more in size, would seem impossible; and even to a student of dispersal improbable. Unless, however, we prefer to accept the Lemurian theory for the Indian Ocean and the theory of a Melanesian continent for the Pacific we are compelled to appeal to these birds; and it can scarcely be said that our appeal is without some justification. Both in the Solomon Islands and in the Fijis I was familiar with the dispersal of the stones of these trees by fruit-pigeons; and Wallace, amongst other writers, observed the same long ago in the Malayan Islands (Malay Archipelago). Stones obtained from the crops of Fijian pigeons measured12⁄10× 1 inch (3 × 2·5 cm.). In the Solomon Islands these birds stock theinterior of the coral islets with trees of the genus, and the ground below the trees is often strewn with the disgorged stones (Bot. Chall. Exped., iv, 310; Guppy’sSolomon Islands, p. 85).
Although the difficulty concerned with the transport of the seeds across a broad tract of ocean seems very great, it is quite possible that further investigation will enable us to overcome this objection, just as we have done inChapter XXVIwhen explaining how the genus Elæocarpus may have reached Hawaii. It is, indeed, not unlikely that, as with Elæocarpus, the stones of the drupes may in some species be much smaller and far more fitted for being carried in a bird’s body over several hundred miles of ocean.
Reference is here made to this genus because its mode of dispersal is known, and because I was familiar with it in Fiji. Seemann gives two species for Fiji, C. corynocarpa and C. seemanni, and the few other species known seem to be confined to the Western Pacific. Solereder gives a third species, C. densiflora, for Kaiser-Wilhelmsland in New Guinea (Engler’sPflanz. Fam.teil 4, abth. 2); and a Solomon Island species, nearly allied to, if not a variety of, the Fijian species, C. seemanni, is referred to in the list of plants from that group given in my book on those islands. I found C. corynocarpa not infrequently growing on the banks of small rivers in the heart of Vanua Levu. Its drupes, which float for a few days in sea-water, are, according to Seemann, eaten by fruit-pigeons. The “stone” varies from 2 to 4 centimetres(3⁄4-11⁄2inch) in length; and from the standpoint of dispersal the genus ranks with Canarium and Dracontomelon. Seemann describes and figures this species, which was constituted by Gray, in hisFlora Vitiensis; but, apparently through an error, it is in theIndex Kewensisaccredited to Hawaii. Hillebrand makes no reference to the genus in his book on the Hawaiian flora.
This genus of palms is closely allied to Ptychosperma, a Malayan genus also represented in Fiji. TheIndex Kewensisnames four species, one New Hebridean, and three Fijian. The fruits of two of the last-named species tested by me had no floating power. The seed is about an inch long, and the genus would be likely to be spread by fruit-pigeons. From the standpoint of dispersal the genus would be placed with Canarium and Couthovia; but possiblyits presence in the Pacific may be indicative of an ancient Western Pacific continent.
This genus of some eighty known species is almost entirely Australian, with the exception of a few species found in New Caledonia, Tasmania, and apparently also in the Mascarene Islands. Horne was the first to record a species from Fiji, where it grows commonly in the “talasinga” plains on the lee sides of the islands, and also on the scantily vegetated mountain summits. In Vanua Levu I often found these plants growing on the rocky peaks of the highest mountains of the island, as on Mbatini, 3,500 feet, and on Mariko, 2,900 feet. Their presence on these isolated peaks can only be attributed to birds. The carpels contain one or two seeds, which have a membranous aril; but in the plains the seeds are usually destroyed by grubs.
These two genera of epiphytes, distributed over Malaya and extending to the islands of the Western Pacific, possess tuber-like stems, which are extensively chambered by ants that find a home in the interior. They were familiar to me in the Solomon Islands, where they frequently grow on the mangroves and on other littoral trees. They do not form such a feature in the shore vegetation of Fiji, and judging from the observations of Dr. Seemann and myself they occur most often on the wooded mountain-peaks. The berries of these plants would attract frugivorous birds; and their pyrenes, which in a Fijian Myrmecodia I found to be 4 millimetres long, appear quite suitable for dispersal through this agency. It would seem that germination may occur in the berry on the plant. A specimen of Myrmecodia in fruit, that had been lying overlooked for a fortnight between newspapers during one of my mountain journeys, displayed on examination the pyrenes in a germinating condition, the process being subsequently completed. The reader will find these interesting plants described and illustrated in the English edition of Schimper’s work onPlant-Geography, pp. 149, 150.
The Nutmeg trees, though principally at home in Indo-Malaya, are found also in the warm regions of Africa and America, as wellas in the islands of the Western Pacific from the Solomon group eastward to Fiji, Tonga, and Samoa. The Tongan and Samoan groups possess two species in common, whilst Fiji seems to possess its own species, four or five in number.
The seeds of this genus have long been known to be dispersed by fruit-pigeons. Mr. Moseley, in hisNotes of a Naturalist, and in theJournal of the Linnean Society(vol. xv), tells us how at one time these birds in their dissemination of the seeds in the Banda Islands were active opponents of the policy of the Dutch Government in preserving their monopoly of the cultivation of the nutmeg of commerce. He found numbers of wild nutmegs in the crops of these birds in the Admiralty Islands, some of which were partially digested and others seemingly sound; and Mr. Hemsley includes the genus as amongst those dispersed in the Western Pacific by birds (Bot. Chall. Exped., Introd. 46; iv, 229, 308). In my book on the Solomon Islands I refer to the occurrence of these seeds in the crops of fruit-pigeons; and I found that the seeds were similarly dispersed by these birds in Fiji. It is likely that the absence of the genus from Eastern Polynesia is to be partially connected with the insufficient protection of the seeds against injury during such a long ocean passage in a bird’s body.
Gaudichaud, as quoted by Hemsley, refers to the occurrence of the fruits of three or four species of Myristica in the drift floating in the Molucca Sea. When in the Solomon Islands I noticed that the unopened fruits of a species floated in sea-water. In later years in Fiji I tested this point, and found that whilst the fruits just before dehiscing will float between three and seven days in sea-water, the seeds sink. As I have pointed out in the chapter on Drift, rivers carry down to the sea an abundance of seeds and fruits that can float a few days but do not imply dispersal by currents.
Although, as I have above remarked, the localised range of the genus in Polynesia may be in part connected with the insufficient protection of the seed, it is apparent that in the case of a genus found in Asia, Africa, and America we are brought into contact with questions other than those of means of dispersal. No one would pretend that Myristica seeds could be carried by birds uninjured across the Pacific Ocean; and to explain the present distribution of the genus we must recall cases of a similar kind, such as Podocarpus, where the genus in past ages had a home in the north, from which, as from a focus of dispersion, it extended into the continents of the Old and the New World (see p.302).
Rhaphidophora(Araceæ)
This genus of climbing aroids, which gives a character to the forests of Indo-Malaya as well as to those of the Western Pacific, is represented in the New Hebrides, Fiji, Tonga, and Rarotonga by a variety of the widely spread R. pertusa that ranges over Indo-Malaya and Eastern Australia. The ripe berries would readily attract birds; and the seeds, 4·5 millimetres long in the case of a Fijian plant, appear hard enough to pass unharmed through a bird’s digestive canal. We seem here to have evidence of a somewhat recent connection between Indo-Malaya and Polynesia through the agency of frugivorous birds. That the genus has been long established in Polynesia is, however, indicated by the occurrence there of a species seemingly peculiar to Fiji. We are disappointed that in Engler’s recent contribution to thePflanzenreich(in his volume on the Araceæ-Pothoideæ) he has not been able to include this genus in the field of his studies.
This Gymnospermous genus, which is found in the warm regions both of the Old and the New World, is represented in Fiji by a Malayan species, Gnetum gnemon, which exists also in the Solomon group with other species of the genus (Guppy’sSolomon Islands, pp. 288, 301). I was familiar with this species in both Fiji and the Solomon group; but in the first-named locality it is seemingly restricted to the borders of Wainunu Bay on the south side of Vanua Levu, where Dr. Harvey first found it. It grows there abundantly in young wood.
It seems almost idle to discuss the mode of dispersal of a genus that is placed in a class apart with the African Welwitschia and the European Ephedra, possessing with them a history of which we know nothing. Yet it is ranked by Mr. Hemsley amongst those genera that are dispersed in Polynesia by birds, and he produces better evidence in support of this view than we possess for many other plants. Thus a fruit of a species of Gnetum, perhaps G. gnemon, has been found in a New Guinea fruit-pigeon; and the fruits of two species of the genus were found in the crops of fruit-pigeons shot by Mr. Moseley in the Admiralty Islands (Bot. Chall. Exped., Introd. 46; iv, 308). The red drupes of Gnetum gnemon of Fiji would readily attract birds, and their nut-like stones, about 8 millimetres long, are well suited for thismode of dispersal. My experiments in Fiji show that neither the drupe nor the stone of this species floats in sea-water; and it is probable that the fruits of this genus referred to by Mr. Hemsley as having been picked up on the beach in the Aru Islands possessed only a temporary buoyancy.
This genus presents us with the same puzzling question put to us by several Fijian genera, such as Myristica and Podocarpus, that occur in both Asia and America; and until we answer that query it seems almost futile to study modes of dispersal.
This genus of annual and perennial herbs belongs to the tropical regions of the Old World. It is represented in Samoa by fifteen known species and by at least four or five in Fiji, whilst with the exception of a solitary Tahitian species it is not recorded from East Polynesia. Reference is here made to it particularly on account of its great development in Samoa. We have here a genus that, like Psychotria in Fiji and Cyrtandra in Fiji, Samoa, and Hawaii, runs riot in respect to the production of species (see p.317). Dr. Reinecke describes fifteen Samoan species, of which, with the exception of two found in Malaya, all seem to be described for the first time. So sensitive, he remarks, is the genus to external conditions that station-forms abound; and he points out that if we were to follow the dividing lines usually recognised between species, we should account every station-form a new species. It is, of course, obvious that the polymorphism of the Samoan Elatostemas depends primarily not on the varying influence of station but on their sensitiveness to external conditions. One might put the question to the Samoan Elatostemas that Hillebrand put to the Hawaiian Cyrtandras, and ask why nature in this particular genus in this particular locality thus luxuriates in formative energy. Almost every Pacific group in respect of some of its plants presents the problem so well stated by Dr. Reinecke for this genus in Samoa. It is noteworthy that Schimper, in his work onPlant-Geography(English edition, pp. 291, 297, 299), especially singles out Elatostema and Cyrtandra as growing socially in the tropical rain-forests of Java and of the Asiatic mainland.
As far as I can gather, this giant-sedge has not been previously recorded from Fiji; but it is included in the Samoan flora, and hasalso been found at Penang and Singapore, as well as in Borneo, Java, and Queensland. In Samoa, as we learn from Reinecke, it grows both in the coast swamps and on dry ground. In Fiji it is very common in the mangrove-swamps at the mouths of rivers, especially in the Lower Rewa; but in Vanua Levu it is also frequent in the marshy localities of inland plateaux, 700 to 800 feet above the sea, as well as by the side of streams in swampy districts on the lower hill slopes. This double station in the salt-water swamp of the coast and in the fresh-water marsh of the interior seems to be repeated in Java, where the plant was first discovered by Zippelius on the banks of torrents in mountainous regions and in swampy places.
The genus comprises, according to theIndex Kewensis, only this species, though variations are to be observed in plants from different localities. The species was described by Kurz in theJournal of the Asiatic Society of Bengal(vol. 38, 1869) and by Bentham in hisFlora Australiensis; and an illustration is given by Miquel in hisIllustrations de la Flore de l’Archipel Indien(1871). The plant is so common in Fiji that one can only suppose that its resemblance to a stemless Pandanus, from which, as Kurz observes, it is with difficulty distinguished except when in flower or fruit, led to its being overlooked by both Seemann and Horne. Its leaves, from 9 to 12 feet in length, are commonly used for making mats and for thatching, both in Fiji and Samoa. The plant usually attains a height of 3 to 5 feet.
The fruits occur abundantly in the floating and stranded river and sea drift in Fiji, a circumstance that led to my discovery of the parent plant in the swamps. The fruit, which is about half an inch (12 mm.) long, consists of a hard, stony nut invested by a thick ribbed cork-like covering, to which it owes its buoyancy, since the nut sinks. The detached fruit is perforated at the base through both coverings, and only a little soft tissue closes the aperture in the inner shell, the protection against the entry of sea-water in the case of floating fruits being quite inadequate. This explains also why the stranded fruits were so frequently found by me germinating on the beach, where, as my observations showed, they never established the plant. This early germination would prove to be an advantage in the case of fruits stranded in a suitable locality.
But though the perforation in the fruit favours its early germination, it lessens its ability to withstand a long sea-passage without injury to the embryo. I found in different experiments onfruits of plants growing in the mangrove swamps, that when placed in sea-water 40 per cent. sank during the first fortnight, whilst 15 per cent. floated after five or six weeks, but all were at the bottom in two months. On the other hand, fruits from plants of the swamps of the inland plateaux displayed much feebler floating power, in some cases sinking at once, in others floating for a few days, and in others again floating for a week or two. In this case the outer cork-like covering proved to have lost most of its floating power.
From the number of empty seed-vessels found, both in the floating and stranded drift, it appeared evident that the seed had often rotted away during the flotation. It is apparent from these observations and experiments that Scirpodendron costatum is not suited for dispersal by currents over wide tracts of ocean. The fruits might be able to float unharmed for a few weeks, but they would be unable to accomplish much more than the 500 or 600 miles intervening between Fiji and the nearest groups to the west.
This order, judging from the writings of Hegelmaier, Schenck, and Hemsley, is represented by one or other of the common species, Lemna minor, L. gibba, L. polyrrhiza, in various Atlantic islands, as in the Bermudas, the Azores, Madeira, the Canary Islands, and St. Helena; but doubts frequently arise as to their being truly indigenous. Lemna trisulca is regarded by Hemsley as indigenous in the Bermudas. Lemna minor has been introduced in recent years into Hawaii, where I observed it flowering and sometimes fruiting abundantly in the heated waters of the ponds. Two species found in other regions were recorded by Seemann from Fiji, and I have come upon few other records of the occurrence of the order in the tropical islands of the open Pacific. I am inclined to the opinion, based not only on the facts of distribution, but also on the results of numerous experiments on the means of dispersal, that this order has in most cases reached oceanic islands with man’s assistance.
Some years ago I made a systematic study of the habits of the British Lemnæ, most of the results being published in theLinnean Society’s Journal(vols. xxix and xxx), as far as concerned Lemna minor, L. gibba, and L. polyrrhiza. During this inquiry I ascertained that with these species, as well as with L. trisulca, the chances of a bird’s carrying their fronds uninjured in its plumageover a wide extent of ocean were small. None of them survived twenty-four hours’ drying in fine weather, whether in the sun or in the shade; but in rainy weather they withstand an exposure of one or two days. It is, therefore, unlikely, even if the fronds were entangled by their rootlets in a bird’s feathers, that they would be able under ordinary conditions to reproduce the plants after a day’s flight of some five hundred miles across the sea. It must also be remembered that the drying capacity of the air when a bird is in full flight in ordinary weather would be that displayed during a gale of wind with a velocity of at least thirty to forty miles an hour. For this reason I do not think with Kerner that under usual conditions drops of water would be a factor of importance in causing the adherence of minute seeds of any kind to birds’ plumage. Where the seeds are not available, it is most probable that birds disperse the duckweeds by their fronds over short distances, but not across broad seas. This would certainly apply to temperate latitudes, where these plants rarely seed. Thus with Lemna, as with Ceratophyllum, it would seem that the dispersal of the seeds by birds takes place normally only in warm latitudes. Those of the duckweeds could be transported in adherent mud over land-areas.
According to Hegelmaier, the two species of Lemna found in Fiji are L. paucicostata, an Asiatic species, and a variety of an Australian species, L. oligorrhiza, possessing dark root-sheaths. These plants mostly came under my notice in the Rewa delta. They were rarely seen in Vanua Levu, where in one locality I found the typical Lemna minor. The first species is also Samoan.
In 1897 and in 1899, in a pool near Notho in the Rewa delta, in Viti Levu, Fiji, I found a great abundance of a species of Wolffia, specimens of which were sent to Prof. Schimper with my mangrove collections, but his death intervened, and I have not been able to follow up the matter. On comparing the specimens with Hegelmaier’s descriptions and plates, it would seem that the species is near W. arrhiza and W. brasiliensis, but differs from both in the greater length of the fronds. As concerning the means of dispersal of the genus, I may add that the fronds were killed after being allowed to dry for eighteen hours.
A species of this genus, apparently near Marsilea villosa, was common in the ditches and ponds around Notho, in the Rewadelta, Fiji, in 1897-99. The genus is included by Horne in his list of Fijian plants; but is not given by Seemann. The villous sporocarps, when dry, are very light and readily catch in cloth and in feathers. Hillebrand includes in the Hawaiian flora M. villosa and M. crenulata. The first-named, which was collected by Chamisso and Gaudichaud, finds (he says on the authority of Braun) its nearest relative in a species from Oregon and California. The other has been collected in the Liukiu Islands, the Philippines, Mauritius, and Bourbon. It is very probable that the occurrence of the genus in oceanic islands is due to the agency of birds.
(1) We are here concerned with the more restricted distribution of non-endemic tropical genera over the Pacific. The general trend eastward of these genera is well brought out in the fact that whilst Fiji possesses some sixty or seventy genera in common with Tahiti to the exclusion of Hawaii, it does not possess a score in common with Hawaii to the exclusion of Tahiti. The grasses and sedges and the mountain genera are not here included; and we are comparing the flora of the Hawaiian lowlands below 4,000 feet with the floras in mass of Fiji and Tahiti.
(2) Hawaii possesses very few genera (less than thirty) that are not found either in Fiji or in Tahiti, or in both; and of these quite a third are to be traced to America.
(3) From two of these genera, Embelia, a land genus, and Naias, an aquatic genus, we obtain two important indications, namely, that specific differentiation has taken place to much the same extent in a water plant as in a land plant, whether in a continent or in an island. In other words, new species have been developed or are developing independently of the immediate environment and of isolating influences.
(4) The interchange of plants between the regions of Hawaii and Tahiti to the exclusion of Fiji has been very slight. Probably not half a dozen genera belong to this category.
(5) Excluding plants brought by man and by the currents, Tahiti possesses very few that present any difficulty from the standpoint of dispersal, plants with seeds or “stones” an inch in size being, as a rule, absent.
(6) With the genera (60-70) common to Fiji and Tahiti, and distributed, therefore, over the South Pacific, the wide-ranginghighly variable plant is an important factor in the development of peculiar species in the different groups, just as it has been shown to be in the previous chapter in the case of genera dispersed over the whole Pacific. Therôleof the polymorphous species has always been an important one in this region.
(7) In the case of several Fijian genera it seems almost futile to talk of existing means of dispersal, since the present distribution of genera like Sterculia and Gnetum, that occur on both sides of the Pacific, in America and in Asia, is not to be thus explained.
(8) On account of the large size of their seeds and “stones” it might be argued that certain of the Fijian plants afford evidence of a previous continental condition of the islands of the Western Pacific, since it is not easy to understand how such large seeds and “stones” could have been transported over broad seas by birds. It is, however, pointed out that in these respects the species of a genus may vary greatly, and that the seeds and stones may be large in some species and small in others.
(9) The greater number of the genera that have entered the Pacific from the Old World have not advanced eastward of the Fijian region, half of the Fijian genera not occurring in the Hawaiian and Tahitian regions; and the explanation of this is to be found not in any lack of capacities for dispersal, but in a want of opportunities. The story of plant-distribution in the Pacific is bound up with the successive stages of decreasing activity in the dispersing agencies. The area of active dispersion that at first comprised the whole of the tropical Pacific was afterwards restricted to the South Pacific, and finally to the Western Pacific only. The birds that in an early age carried seeds all over this ocean became more and more restricted in their ranges, probably on account of increasing diversity of climatic conditions. The plants of necessity responded to the ever narrowing conditions of bird-life in this ocean, and the differentiation of the plant and of the bird have taken place together.