In this chapter we have a study of Leguminous strand plants that are of great interest. It can be safely said that the student of plant-dispersal in the Pacific will be brought into contact with the problems here involved wherever he goes.
Afzelia bijuga (Gray).
This Old World tree, which belongs to the sub-family Cæsalpiniæ, is of great interest to the student of plant-dispersal. It is one of that large group of Indo-Malayan plants that extend into the Western Pacific, and give the prevailing character to the floras of such archipelagoes as that of Fiji. It is a large tree yielding a valuable timber used by the Fijians and Samoans for many purposes, such as for canoes, house-posts, clubs, kava bowls, &c., but it has not been recorded from the Tahitian region, and is unknown from Hawaii. In the fact of its being a littoral as well as an inland tree, it possesses a peculiar interest from the standpoint of plant-dispersal, and especially since this difference in station is associated with a difference in buoyancy, the seeds of the inland trees usually sinking, whilst those of the coast trees usually float, and often for a period of months.
A glance at the distribution of the genus will enable us to appreciate some of the points that will be touched upon in the following discussion; and it may be here remarked that the explanation of the distribution of these Leguminous trees will go far to make clear some of the most difficult points in plant-geography. Of the eleven species enumerated in the Index Kewensis, five belong to tropical Africa, occurring on both the east and west coasts as well as in the interior, three are confined to the mainland of tropical Asia, and two are peculiar to Malaya. In the last place we have the wide-ranging Afzelia bijuga, which, if it does not actually occur on the east coast of Africa, is found at all events in Madagascar and in the Seychelles, and is to be followed by the way of the Chagos Archipelago to the Malayan Islands and Queensland, and eastward to Fiji and Samoa.
The most suggestive feature in the distribution of the genus is to be seen in the frequent station of the species by rivers. We learn from Oliver’s Flora of Tropical Africa that these trees find a home along river-courses on both sides of the continent, as on the banks of the Congo, the Niger, the rivers of Senegambia, and the Zambesi, the Zambesi species being found also on the shores of Lake Nyassa. Since tropical Africa possesses about half of the species, it would seem highly probable that it is the home of the genus, and that from the rain-forests in the heart of the continent rivers flowing east and west have borne the buoyant seeds of the wandering species to the coasts of the Atlantic and Pacific Oceans. The operation that I witnessed on a miniature scale in the case of a species of Entada (E. scandens) in the Isthmus of Panama, as described in a later page of this chapter, has been in progress through the ages with the genus of Afzelia in the breadth of the African continent. According to the principle illustrated by Afzelia bijuga in the forests of Fiji, the seeds of the African forest-trees would, as a rule, possess no floating power; but now and then in the lapse of long periods of time buoyancy in some species would be developed, and such species would ultimately, through their buoyant seeds, find their station along the lower courses of the rivers.
To sustain this view it is not necessary that continuous rain-forests should now clothe the elevated regions in the interior of tropical Africa; but it is requisite that there should be sometimes a generic similarity between the plants of the East African and West African rain-forests; and it is evident that this is the case. Pechuel-Lösche, as quoted by Schimper (Plant-Geography, p. 299), describes the rain-forest on the Loango coast as covering the mountain ranges and as extending to the river-plains. In such a locality the operation would be rapid. In advancing this hypothesis I am referring to the possibility, however, of such an operation having effected the distribution of Afzelia in tropical Africa in the past rather than in the present. I would suggest that botanists in other habitats of the genus, as for instance in Queensland, might put it to the test of observation and experiment.
The interest that attaches itself to the story of the genus in its African home may be extended to the species that forms its outpost in the Pacific, and we shall see there a littoral species that doubtless had its home in the interior of a continent endeavouring, with a considerable measure of success, to become again an inland plant. Horne (p. 112), who was familiar with Afzelia bijuga at the two extremes of its range, namely, in the Mascarene Islands and in Fiji, speaks of it as characteristic of the shores of tropical regions; and Schimper, who includes it in the Indo-Malayan strand-flora, implies that it is more or less exclusively confined to the coast and its immediate vicinity (pages 121, 191-2). In the Seychelles, according to Mr. Button, this tree attains gigantic dimensions on the sandy flats. Still larger trees occur in the coral islands of the Chagos Archipelago; but in the atoll of Diego Garcia, as we learn from Mr. Bourne, it is almost extinct only some four or five trees existing there about twenty years ago, the increase of the tree being prevented through the destruction of the fallen seeds by the rats (Journ. Linn. Soc. Bot., vol. 22, 1887).
Afzelia bijuga may, therefore, be safely regarded as a littoral tree. We shall now see the importance of this conclusion when we come to consider its station in the Pacific islands, where it grows both inland and at the coast, and we have to decide to which station we must assign the priority. Speaking of its occurrence in Fiji, Dr. Seemann says it is “common in the forests all over Viti,” but makes no allusion to it as a littoral tree either in Fiji or elsewhere. On the other hand, Mr. Horne (p. 112) describes it as “generally growing on the shore or sandy beaches, and in rocky clefts, and by the sides of streams in the interior of Viti Levu and Vanua Levu.” It was on or near the coast in Fiji that the present writer was most familiar with this tree, sometimes bordering the sandy beach, at other times growing behind the mangrove-belt, or again thriving in the half sandy and half swampy soil of some low islet off the mouth of the Rewa. Especially is it to be found on those parts of the coast where the hill-slopes descend rapidly to the beach, or where some lofty spur from the mountains of the interior reaches the shore. It is also not uncommon on the banks of rivers both in their lower and upper courses. But it is as a forest-tree of the interior that it is most valued by both the white men and the natives on account of the superior quality of its timber in that station. There, far removed from stream or river, the Vesi, as the Fijians name Afzelia bijuga, takes its place amongst the lofty forest-trees, such as the Ndamanu (Calophyllum), the Ndakua (Dammara), and the Wathi-wathi (Sterculia). It is not often that one finds a tree in these islands that, like the Vesi, is able to make its home in almost any station, excepting, however, the “talasinga” or “sun-burnt” regions of the plains. Wherever tall trees grow gregariously in Vanua Levu, one will probably find Afzelia bijuga, whether beside a sandy beach, or bordering a swamp, or on a river’s bank, or on some rocky declivity, or on the great forest-clad mountain-slopes and plateaux of the interior. No doubt the same diversity of station is displayed in Samoa, where, according to Dr. Reinecke, the tree is most frequent in the “coast-bush.”
From the variety in station it might be expected that corresponding variations in character would be found. There are differences, such as in the quality of the timber and in the size of the seeds between coast and inland trees; but the most important distinction in connection with the study of the dispersal of the species is to be found in the circumstance that whilst the seeds of the coast trees are, as a rule, buoyant, and often float for months, those of the inland trees usually sink, even after being kept for three or four years. I made a considerable number of experiments on the buoyancy of the seeds of this tree in Vanua Levu, and found that with the coast trees, as a rule, either all the seeds or the majority of them floated in sea-water, whilst with the inland trees either all of them or the majority of them sank. The buoyant seeds are able in most cases to float for a long time. Thus, in one experiment half were afloat after two months, and in another half were afloat after five months. It is probable that several of the exceptions, where inland seeds float, will prove to be connected with an inland station by a river. (I experimented on eight sets of seeds of coast trees from eight different localities, and found 70 to be the mean percentage of buoyant seeds. In the same way, four sets of seeds from four different inland localities gave 13 as the mean percentage of buoyant seeds.)
As in the case of Entada scandens, there is a rather fine adjustment between the mean specific weight of seeds and the density of water. If we place a number of the buoyant seeds in sea-water and begin to lower the density, some of the seeds will at once commence to float heavily and afterwards sink; and when the density has been lowered to approximately that of fresh water, usually about a third will be found at the bottom of the vessel. Out of 100 coast seeds, 70 will, as a rule, float in the sea and about 47 in the river; whilst of the same number of inland seeds, 13 on the average will float in sea-water and 8 or 9 in fresh water. The bearing of facts of this kind is especially discussed in [Chapter X.]