Soc." 1907, and especially E.A.N. Arber and J. Parkin, "On the Origin of Angiosperms", "Journal Linn. Soc." (Bot.) Vol. XXXVIII. page 29, 1907.)The great complexity of the Bennettitean flower, the earliest known fructification to which the word "flower" can be applied without forcing the sense, renders it probable, as Wieland and others have pointed out, that the evolution of the flower in Angiosperms has consisted essentially in a process of reduction, and that the simplest forms of flower are not to be regarded as the most primitive. The older morphologists generally took the view that such simple flowers were to be explained as reductions from a more perfect type, and this opinion, though abandoned by many later writers, appears likely to be true when we consider the elaboration of floral structure attained among the Mesozoic Cycadophyta, which preceded the Angiosperms in evolution.
If, as now seems probable, the Angiosperms were derived from ancestors allied to the Cycads, it would naturally follow that the Dicotyledons were first evolved, for their structure has most in common with that of the Cycadophyta. We should then have to regard the Monocotyledons as a side-line, diverging probably at a very early stage from the main dicotyledonous stock, a view which many botanists have maintained, of late, on other grounds. (See especially Ethel Sargant, "The Reconstruction of a Race of Primitive Angiosperms", "Annals of Botany", Vol. XXII. page 121, 1908.) So far, however, as the palaeontological record shows, the Monocotyledons were little if at all later in their appearance than the Dicotyledons, though always subordinate in numbers. The typical and beautifully preserved Palm-wood from Cretaceous rocks is striking evidence of the early evolution of a characteristic monocotyledonous family. It must be admitted that the whole question of the evolution of Monocotyledons remains to be solved.
Accepting, provisionally, the theory of the cycadophytic origin of Angiosperms, it is interesting to see to what further conclusions we are led. The Bennettiteae, at any rate, were still at the gymnospermous level as regards their pollination, for the exposed micropyles of the ovules were in a position to receive the pollen directly, without the intervention of a stigma. It is thus indicated that the Angiosperms sprang from a gymnospermous source, and that the two great phyla of Seed-plants have not been distinct from the first, though no doubt the great majority of known Gymnosperms, especially the Coniferae, represent branch-lines of their own.
The stamens of the Bennettiteae are arranged precisely as in an angiospermous flower, but in form and structure they are like the fertile fronds of a Fern, in fact the compound pollen-sacs, or synangia as they are technically called, almost exactly agree with the spore-sacs of a particular family of Ferns--the Marattiaceae, a limited group, now mainly tropical, which was probably more prominent in the later Palaeozoic times than at present. The scaly hairs, or ramenta, which clothe every part of the plant, are also like those of Ferns.
It is not likely that the characters in which the Bennettiteae resemble the Ferns came to them directly from ancestors belonging to that class; an extensive group of Seed-plants, the Pteridospermeae, existed in Palaeozoic times and bear evident marks of affinity with the Fern phylum. The fern-like characters so remarkably persistent in the highly organised Cycadophyta of the Mesozoic were in all likelihood derived through the Pteridosperms, plants which show an unmistakable approach to the cycadophytic type.
The family Bennettiteae thus presents an extraordinary association of characters, exhibiting, side by side, features which belong to the Angiosperms, the Gymnosperms and the Ferns. ii. ORIGIN OF SEED-PLANTS.
The general relation of the gymnospermous Seed-plants to the Higher Cryptogamia was cleared up, independently of fossil evidence, by the brilliant researches of Hofmeister, dating from the middle of the past century. (W. Hofmeister, "On the Germination, Development and Fructification of the Higher Cryptogamia", Ray Society, London, 1862. The original German treatise appeared in 1851.) He showed that "the embryo-sac of the Coniferae may be looked upon as a spore remaining enclosed in its sporangium; the prothallium which it forms does not come to the light."(Ibid. page 438.) He thus determined the homologies on the female side.
Recognising, as some previous observers had already done, that the microspores of those Cryptogams in which two kinds of spore are developed, are equivalent to the pollen-grains of the higher plants, he further pointed out that fertilisation "in the Rhizocarpeae and Selaginellae takes place by free spermatozoa, and in the Coniferae by a pollen-tube, in the interior of which spermatozoa are probably formed"--a remarkable instance of prescience, for though spermatozoids have not been found in the Conifers proper, they were demonstrated in the allied groups Cycadaceae and Ginkgo, in 1896, by the Japanese botanists Ikeno and Hirase. A new link was thus established between the Gymnosperms and the Cryptogams.
It remained uncertain, however, from which line of Cryptogams the gymnospermous Seed-plants had sprung. The great point of morphological comparison was the presence of two kinds of spore, and this was known to occur in the recent Lycopods and Water-ferns (Rhizocarpeae) and was also found in fossil representatives of the third phylum, that of the Horsetails. As a matter of fact all the three great Cryptogamic classes have found champions to maintain their claim to the ancestry of the Seed-plants, and in every case fossil evidence was called in. For a long time the Lycopods were the favourites, while the Ferns found the least support.
The writer remembers, however, in the year 1881, hearing the late Prof.
