My further remarks on these Tahitian genera found in Fiji but not in Hawaii will be limited to some general observations from the standpoint of dispersal. I will first discuss some of those genera that possess only peculiar species. They belong to an era of dispersal that, as far as Tahiti is concerned, is passing or has passed away. Here we have the species of each genus more or less localised in the various South Pacific archipelagoes; but, as with Meryta, Alstonia, and Loranthus, it is often apparent that, although the Tahitian region is mainly outside the zone of present dispersal, the different groups of the Western Pacific are kept in touch by the possession of species in common. This testifies to the activity of dispersal in that region after it had become suspended in Eastern Polynesia. The connection between the isolated endemic species of Eastern Polynesia and a species ranging over the Western Pacific can sometimes be shown, as in the case of Loranthus, where a species confined to the Society Islands and to the Marquesas is very closely related to L. insularum, a widely-ranging West Polynesian species that reaches eastward as far as Rarotonga.
We next have those genera like Grewia, Nelitris, Melastoma, Randia, Geniostoma, Tabernæmontana, Fagræa, Bischoffia, Macaranga, and Ficus, that possess in Polynesia one or more widely-ranging species. The agency of the polymorphous species, which I have described in the preceding chapter in connection with the general dispersal of Malayan plants over the whole of Polynesia, is evidently also active when the work of dispersal is restricted to the South Pacific. Its operation is to be distinctly traced in all the genera above named except in Fagræa and Ficus. Thus, in the genera Grewia, Melastoma, Randia, Geniostoma, and Macaranga we find a single variable species ranging over the South Pacific from Fiji to Tahiti, keeping all the groups in touch, but associated in each, as a rule, with one or more peculiar species. A yet earlier stage in the process is to be seen in those genera which, like Nelitris, Tabernæmontana, and Bischoffia, possess only a solitary species ranging over the South Pacific, varying in each group, but not usually associated with endemic species. As with Melastoma, Macaranga, and others, we can often trace the widely-ranging species of Polynesia back to its home in Malaya, and with these and other genera the connection between a species confined to a group and a variable species ranging through all the archipelagoes of the South Pacific can sometimes be detected in the affinity of their characters.
It is thus seen that one of the principal determining causes of the differentiation of species in Polynesia lies in the failure of the dispersing agencies, a widely-ranging species becoming in consequence gradually isolated in the various groups. With some genera, as with Ophiorrhiza, it is possible to show that the resulting endemic species pass into each other by intermediate forms.
My further remarks on the Tahitian genera occurring in Fiji but not in Hawaii will be devoted mainly to those with which I was most familiar from the standpoint of dispersal.
The Tiliaceous genus Grewia offers a good example of those Polynesian genera which possess in the South Pacific a single widely-ranging species associated often with endemic species in the individual groups. It is likely that a polymorphous form, including most of the Polynesian species, could be here constituted. The fruits are dryish drupes, becoming black and moist when over-ripe, and containing three or four pyrenes suitable for distribution by birds and five or six millimetres in size.
The berries of Nelitris, a genus of the Myrtaceæ, contain a few hard seeds that are well fitted for dispersal by frugivorous birds. I am inclined to follow Drake del Castillo, who considers that there is only one varying species, N. vitiensis (Gray), which is distributed over the whole of the South Pacific from the Solomon Islands to Tahiti. The tendency of this widely-ranging species to vary in different groups is indicated in the fact that some botanists have distinguished other species within these limits. It is noteworthy that N. paniculata in Indo-Malaya and N. vitiensis in the Pacific cover the whole range of the genus. It would be interesting to establish a connection between them.
Melastoma, an Old World genus of forty and more species, has one very variable species, M. denticulatum, which, as defined by Bentham, has the range of the genus from tropical Asia across the Pacific to Tahiti. This plant is associated in some groups, as in Fiji, Tonga, and Samoa, with other more or less localised species, and it affords a good example of the principle of polymorphism in species-making. The berry-like fruits contain an abundance of minute seeds, half a millimetre in size, which, when rendered adhesive by adherent pulp, might readily stick to feathers, or they might pass unharmed through the alimentary canal of a bird. It is noteworthy that amongst the plants regarded by Prof. Penzig as introduced by frugivorous birds into Krakatoa since the eruption is a species of Melastoma.
Few genera in these islands would better repay a careful study of their species with regard both to the influence of station on specific characters and to the question of “mutations” than Ophiorrhiza. I found the three Fijian species of this Rubiaceous genus so often in close association, that I cannot doubt there is some connection between them. Seemann and Gray, indeed, characterise two of them as confluent species. The island of Tahiti alone possesses five peculiar species, and it is evident that this island has been a centre of development for species of Ophiorrhiza, just as Samoa has become the birthplace of many species of the Urticaceous genus Elatostema. The minute angular seeds of Ophiorrhiza would probably be transported in a bird’s feathers or in adherent soil. As my experiments showed, they do not become adhesive when wet.
The genus Loranthus as distributed in the South Pacific has already been briefly noticed. There is a species confined to the Tahitian region, and there is another peculiar to Samoa, whilst one widely-ranging species, L. insularum, that connects these regions together, reaching east to Rarotonga, is closely related with the Tahitian species. There was no doubt originally a single polymorphous plant that ranged over the tropical South Pacific. With regard to the mode of dispersal of the seeds of this genus of parasites, I should at once refer to the systematic and careful observations made by Mr. F. W. Keeble in Ceylon (Trans. Linn. Soc., v. 1895-1901). He formed the opinion that the seeds of Loranthus usually reach their host without passing through the alimentary canal of a bird, being merely wiped off its bill. This method would never carry the seeds to Tahiti or even to Fiji; and since this observer remarks that, although most of the seeds in the droppings were completely rotten, some of them “possibly pass through the gut uninjured,” we may accept this possibility as sufficient for the purpose of dispersal in the Pacific Ocean. Mr. Keeble notes the observation in Teil 3 of Engler’s Die Natürlichen Pflanzenfamilien that the seeds may germinate after passing through a bird’s intestine; and we may therefore infer that whilst the method he describes is typical of local dispersal, the other method is required in the instance of oceanic dispersal.
Alstonia, an Apocynaceous genus of tropical Asia and Australia, yields the caoutchouc of Fiji. Besides possessing in Fiji and Samoa peculiar species, the islands of Western Polynesia have in A. plumosa a species common to Fiji, Samoa, and New Caledonia. Another species, A. costata, is restricted to Eastern Polynesia, occurring in the different islands of the Tahitian Group as well as in Rarotonga. It is possible that the Pacific species may be connected with A. scholaris, a species possessing the range of the genus with the exception of Polynesia. The long ciliated or hairy seeds, six to nine millimetres in length, are fitted for transport by the winds and in birds’ plumage. The follicles dehisce on the tree, and, according to Horne, the light seeds are distributed locally by the wind. It is probable that the thick white juice oozing from a broken branch would at times aid the adhesion of the seeds to a bird’s feathers.