(3) The first group is best represented by the Convolvulaceous and the Leguminous types. In the former, which is well illustrated by Ipomœa pes capræ, the seed-cavity is imperfectly filled by the crumpled embryo, the result of the shrinking process during the final setting of the seed. In the latter, which is exemplified by Entada scandens and Cæsalpinia bonducella, the seed displays a large central cavity produced by the arching outward of the cotyledons during the shrinking process accompanying the last stage of the maturation of the seed. As an instance of fruits belonging to the group, those of Heritiera littoralis may be cited. An uncommon type is presented in the “stones” of the drupes of Premna taitensis, and in the pyrenes of Morinda citrifolia, where the buoyancy arises from empty seed-cavities resulting from the failure of some of the seeds.

(4) The second group with buoyant kernels includes mostly widespread Leguminous species, such as Canavalia obtusifolia and Sophora tomentosa.

(5) The third or “adaptive” group comprises many of the characteristic littoral trees and shrubs of the Pacific islands, such as Barringtonia speciosa, Guettarda speciosa, Terminalia katappa, Tournefortia argentea, &c., that contain in their fruit-coverings a buoyant cork-like material often bound together by fibres, but which proves on examination to resemble cork only in appearance. In another type, illustrated by the fruits of Cycas circinalis and the seeds of Anona paludosa, the buoyant tissue forms a layer inside the shell of the seed or “stone.”

(6) Some fruits like those of Ximenia americana and Calophyllum inophyllum illustrate both the so-called mechanical and adaptive principles in their structure; whilst with the first-named species they are as well adapted for dispersal by frugivorous birds and are known to be a favourite food of fruit-pigeons. The same difficulty arises with the fruits of some other characteristic littoral plants, as with Scævola Kœnigii, the drupes of which are equally well fitted for dispersal by birds and currents.

(7) The same general principles have been at work in determining the structures concerned with the buoyancy of the fruits and seeds of British littoral plants. Although the species are few in number they exhibit in this respect great variety, eight species illustrating six or seven types of buoyant structure.

CHAPTER XIII
ADAPTATION AND SEED-BUOYANCY

The question of the operation of Natural Selection.—Are there two principles at work?—The presence of buoyant tissue in the seed-tests and fruit-coats of inland plants, both wild and cultivated.—Useless buoyancy.—The buoyancy of seeds and fruits is not concerned with adaptation.—Summary.

When we speak of a certain structure as an adaptation to dispersal by currents through the agency of Natural Selection, it is necessary at the outset to be quite clear as to what is implied. Professor Schimper, who brought his great and varied knowledge of many other phases of plant-life to bear on this subject, is careful to clear the ground of preliminary erroneous conceptions in such a perspicuous and impartial manner that we cannot do better than follow his guidance. There are, he observes (p. 178), many mechanisms or contrivances in plants, which, though they seem to have arisen with a fixed purpose, can in no wise be regarded as having been developed for that end, since they were produced in quite a different connection and have merely acquired a new or supplementary function, of which they are the cause and not the effect.

This is very much the position that I have taken up for the whole subject of the relation between plants and their dispersing agencies, and it will be found discussed in [Chapter XI.] It involves, as I venture to think, a dominant principle in the organic world, which it is one of the objects of this work to emphasise, namely, that Nature in dispersing plants habitually makes use of structures and capacities that were originally developed in quite another connection. Behind this change of function, this new purpose, lies the secret of the organic world. There is for me no more pregnant fact in plant-life than the thistle-seed blown before the wind, or the seed of our sea-convolvulus floating in the sea. It proves to my mind that the evolutionary power in nature is checked and hampered by conditions not of its own creation, and that two opposing forces are ever at work, the one creating and the other limiting the creative power, the actual mode of dispersal being but a blind and accidental result of the struggle.

The question of the operation of Natural Selection is not raised, as Professor Schimper indicates, until we consider whether the new function has had any bettering influence on the structure or mechanism with which it has come to be concerned. If such a modification is thus brought about it might be legitimately claimed as a result of this agency, and the term “adaptation” could be used. But if there is no evident change produced, we should be compelled to assign very subordinate limits to the capacity of Natural Selection; and in the instance of buoyant fruits and seeds it would be restricted to determining a plant’s station by the water-side and in increasing its area. It is only in the first case that we could speak of them as adaptations in the meaning attached to this term in the language of the Selection Theory. It would at first sight seem easy to ascertain whether the characters of fruits and seeds, to which the buoyancy is due, are adaptations in this sense of the word; but in reality it is far from being so. We can, however, proceed with unanimity up to a certain stage in the argument; but there agreement ends.