Ancient Plants – Past Histories of Plant Families:The Cycads

CHAPTER XI
PAST HISTORIES OF PLANT FAMILIES IV. The Cycads

The group of the Cycadales, which has a systematic value equivalent to the Ginkgoales, contains a much larger variety of genera and species than does the latter. There are still living nine genera, with more than a hundred and fifty species, which form (though a small one compared with most of the prime groups) a well-defined family. They are the most primitive Gymnosperms, the most primitive seed-bearing plants now living, and in their appearance and characters are very different from any other modern type. Their external resemblance to the group of the Bennettitales, however, is very striking, and indeed, without the fructifications it would be impossible to distinguish them.

The best known of the genera is that of Cycas, of which an illustration is given in fig. 74. The thick, stumpy stem and crown of “palm”-like leaves give it a very different appearance from any other Gymnosperm. Commonly the plants reach only a few feet in height, but very old specimens may grow to the height of 30 ft. or more. The other genera are smaller, and some have short stems and a very fern-like appearance, as, for example, the genus Stangeria, which was supposed to be a fern when it was first discovered and before fruiting specimens had been seen.

The large compound leaves are all borne directly on the main stem, generally in a single rosette at its apex, and as they die off they leave their fleshy leaf bases, which cover the stem and remain for an almost indefinite number of years.

The wood of the main trunks differs from that of the other Gymnosperms in being very loosely built, with a large pith and much soft tissue between the radiating bands of wood. There is a cambium which adds zones of secondary tissue, but it does not do its work regularly, and the cross section of an old Cycad stem shows disconnected rings of wood, accompanied by much soft tissue. The cells of the wood have bordered pits on their walls, and in the main axis the wood is usually all developed in a centrifugal direction, but in the axis of the cones some centripetal wood is found (refer to c, fig. 65.

fig74

Fig. 74.:Plant of Cycas, showing the main stem with the crown of leaves and the irregular branches which come on an old plant

In their fructifications the Cycads stand even further apart from the rest of the Gymnosperms. One striking point is the enormous size of their male cones. The male cones consist of a stout axis, round which are spiral series of closely packed simple scales covered with pollen-bearing sacs (which bear no inconsiderable likeness to fern sporangia), the whole cone reaching 1½ ft. in length in some genera, and weighing several pounds. All the other Gymnosperms, except the Araucareæ, where they are an inch or two long, have male cones but a fraction of an inch in length.

fig75

Fig. 75.:Seed-bearing Scale of Cycas, showing its lobed and leaflike character

s, Seeds attached on either side below the divisions of the sporophyll.

In all the members of the family, excepting Cycas itself, the female fructifications also consist of similarly organized cones bearing a couple of seeds on each scale instead of the numerous pollen sacs. In Cycas the male cones are like those of the other genera, and reach an enormous size; but there are no female cones, for the seeds are borne on special leaflike scales. These are illustrated in fig. 75, which shows also that there are not two seeds (as in the other genera with cones) to each scale, but an indefinite number.

The leafy nature of the seed-bearing scale is an important and interesting feature. Although theoretically botanists are accustomed to accept the view that seeds are always borne on specially modified leaves (so that to a botanist even the “shell” of a pea-pod and the box of a poppy capsule are leaves), yet in Cycas alone among living plants are seeds really found growing on a large structure which has the appearance of a leaf. Hence, from this point of view, however, for a caution against concluding that the whole plant is similarly lowly organized), Cycas is the most primitive of all the living plants that bear seeds, and hence presumably the likest to the fossil ancestors of the seed-bearing types. In this character it is more primitive than the fossil group of the Cordaiteæ, and comes very close to an intermediate group of fossils to be considered in the next chapter.

fig76

Fig. 76.:Seed of Cycas cut open

n, The nucellus, fused at the level l to the coat c; sc, stony layer of coat; p.c, pollen chamber in apex of the nucellus; S, “spore”, filled with endosperm, in which lies the embryo e.

To enter into the detailed anatomy of the seeds would lead us too far into the realms of the specialist, but we must notice one or two points about them. Firstly, their very large size, for ripe seeds of Cycas are as large as peaches (and peaches, it is to be noted, are fruits, not seeds), and particularly the large size they attain before they are fertilized and have an embryo. Among the higher plants the young seeds remain very minute until an embryo is secured by the act of fertilization, but in the Cycads the seeds enlarge and lay in a big store of starch in the endosperm before the embryo appears, so that in the cases in which fertilization is prevented large, sterile “seeds” are nevertheless produced. This must be looked on as a want of precision in the mechanism, and as a wasteful arrangement which is undeniably primitive. An even more wasteful arrangement appears to have been common to the “seeds” of the Palæozoic period, for, though many fossil “seeds” are known in detail from the old rocks, not one is known to have any trace of an embryo. A general plan of the Cycas seed is shown in fig. 76, which should be compared with that of Ginkgo (fig. 68). The large size of the endosperm and the thick and complex seed-coats are characteristic features of both these structures. Another point that makes the Cycad seeds of special interest is the fact that the male cells (as in Ginkgo) are developed as active, free-swimming sperms, which swim towards the female cell in the space provided for them in the seed.

The characters of the Cycads as they are now living prove them to be an extremely primitive group, and therefore presumably well represented among the fossils; and indeed among the Mesozoic rocks there is no lack of impressions which have been described as the leaves of Cycads. There is, however, very little reliable material, and practically none which shows good microscopic structure. Leaf impressions alone are most unsafe:more unsafe in this group, perhaps, than in any other:for reasons that will be apparent later on, and the conclusions that used to be drawn about the vast number of Cycads which inhabited the globe in the early Mesozoic must be looked on with caution, resulting from the experience of recent discoveries proving many of these leaves to belong to a different family.

There remain, however, many authentic specimens which show that Cycas certainly goes back very far in history, and specimens of this genus are known from the older Mesozoic rocks. We cannot say, however, as securely as used to be said, that the Mesozoic was the “Age of Cycads”, although it was doubtless the age of plants which had much of the external appearance of Cycads.

From the Palæozoic we have no reliable evidence of the existence of Cycads, though the plants of that time included a group which has an undoubted connection with them.

Indeed, so far as fossil evidence goes, we must suppose that the Cycads, since their appearance, possibly at the close of the Palæozoic, have never been a dominant or very extensive family, though they grew in the past all over the world, and in Europe seem to have remained till the middle of the Tertiary epoch.