The Megapodidæ of the Western Pacific are a family of birds that suggest themselves in this connection. Their distribution corresponds with that of Pandanus in the Western Pacific, excepting the littoral species; and like Pandanus the Megapodes have “differentiated” in every group. The limited powers of flight possessed by existing species would unfit them for crossing wide tracts of sea; but the parent form or forms of all these species must have been able to traverse broad tracts of ocean. These birds subsist on fallen fruits, seeds, &c.; but I have no data relating to them as seed-dispersers.

It is evident from the endemic character of most of the species of Pandanus in oceanic islands that, except with a few widely-spread littoral species, the dispersal of the genus has been for ages suspended. Whether the explanation is to be found in the isolation and differentiation of the extinct Columbæ of the Mascarene Islands, where the endemic species of Pandanus are most numerous, has yet to be established. It seems to offer the only way out of the difficulty, unless we accept the old view concerned with the continent of Lemuria.

Barringtonia.

There are two littoral species of this genus in the Pacific, B. speciosa and B. racemosa, both widely spread over the Old World, but only the first is generally distributed over the Polynesian region reaching east to Ducie Island, whilst the second does not extend east of Fiji and Samoa. With the exception of one or two inland species in Fiji and Samoa no inland species have been recorded from the groups of the open Pacific, and the genus is not represented at all in Hawaii. If it were not for a suspicion that the aborigines may have aided in the distribution of the inland species, the advocate of the previous continental connections of the islands of the Western Pacific would receive from their occurrence in these islands considerable support for his views. The fruits of the inland Fijian species are large, the smallest being three inches in length; and the agency of birds seems to be out of the question.

The fruits of the littoral species possess dry buoyant husks that enable them to be carried by the currents over wide tracts of ocean. Those of the Fijian inland species display only a trace of these buoyant coverings and the floating power is much diminished or absent altogether. These inland species are two or three in number. One of them, described as a new species by Seemann under the name of B. edulis, has edible kernels and is sometimes cultivated. A species that I found growing in the plantations of the Solomon Islanders in Bougainville Straits may be near the Fijian tree just named (Solomon Islands, pp. 85, 297). Its kernels are edible; and I may add that the Solomon Islanders cultivate other species with edible fruits. We cannot, therefore, exclude the agency of the aborigines in the distribution of the inland species of this genus. Horne found an undescribed species in Fiji, which may be that which I found on the slopes of Mount Seatura in Vanua Levu, as described in [Note 50]; and it is quite possible that it was originally a cultivated tree, though not necessarily within the memory of the later generations of the aborigines.

This retrocession to the wild state of cultivated plants and the resulting production of apparently new species is a point on which Dr. Beccari lays considerable stress in the English edition of his book on the Great Forests of Borneo. He takes the case of Nephelium and other fruit-trees and shows how in old clearings, long since abandoned, they have undergone singular alteration in characters. For these reasons, therefore, Barringtonia can scarcely be regarded as offering in its inland species unequivocal evidence of a previous continental condition of the islands of the Western Pacific. Nor, as shown in [Note 50], should we be justified in establishing a genetic connection between the inland and coast species; but a great deal of research is needed before we can handle the numerous interesting problems connected with the genus; and indeed it cannot be said that the specific limits of the inland Polynesian trees have been definitely determined, or the species themselves diagnosed.

Section V.

In this section are included those genera where within the same genus some inland species have been derived from the coast species whilst others have been originally brought by birds. Guettarda alone belongs here. In this genus we find, as is so frequently the case, a littoral tree (G. speciosa) widely spread in the Old World and ranging over the whole tropical Pacific as far east as Pitcairn and Elizabeth islands, but absent from Hawaii. Here also as with Pandanus it is only in the Western Pacific that we find inland endemic species so distinct in character from the littoral tree that they may be regarded as of independent origin.

Since, however, there is an inland form of the coast species in Tahiti (Guettarda speciosa, var. tahitensis) which, according to Drake del Castillo, is distinguished only by its more rounded leaves and by the more marked pubescence of the under leaf-surfaces, we evidently have there an inland species in process of development from the littoral species. This inland tree is found at elevations as great as 600 metres or almost 2,000 feet above the sea; and indeed if we follow Nadeaud the specific differentiation is complete. However, there is no doubt raised as to its close affinity to the beach tree; and we are almost compelled for another reason to regard it as a derivative of the shore species, because, as pointed out in [Chapter XXVII.], there are very few inland plants in the Tahitian flora possessing fruits as large as those of Guettarda that owe their presence in those islands to frugivorous birds.

Of the two inland species of the genus found in Fiji, G. inconspicua and G. vitiensis, it may at once be said that, as indicated in Dr. Seemann’s work, their characters are far from suggesting any connection in origin with G. speciosa, the shore-species, the inland and littoral plants belonging to different sections of the genus. In their case we can only look to the frugivorous bird for the explanation of their existence in the group. The fruits would be probably small; and in this connection it is to be noted that Mr. H. N. Ridley in his paper on the flora of Fernando Noronha evidently looks to birds to account for the presence of a species of Guettarda on the island, a species not found elsewhere.