During recent years a number of fossil (Carboniferous and Permian) plants have been very thoroughly investigated in the light of modern anatomical knowledge, and as a result it has become st i s clear that in those times a large series of plants etisted ear ys intermediate in structure between the modern ferns tern of Cycaand the modern Gymnosperms (especially Cycads), dofiices.
Among the Cycadofilices a series of stages is found leading from the primitive fern-protostele to the type of siphonostele characteristic of the Cycads which agrees in essentials in all the Spermophytes.
Among the existing Cycads, though the type of vascular system conforms on the whole with that of the other existing seed-plants, peculiar structures are often found (e.g.
In Monoblepliaris, one of the lower Fungi, in some Algae, in the Vascular Cryptograms, in Cycads (Zamia and Cycas), and in Ginkgo, an isolated genus of Gymnosperms, the male cell is a motile spermatozoid with two or more cilia.
This body has been called a blepharoplast, and in the Pteridophytes, Cycads and Ginkgo it gives rise to the spiral band on which the cilia are formed.
Cycads are represented by Cycas itself, which in several species ranges from southern India to Polynesia.
Amongst Cycads, Zamia is confined to the New World, and amongst Conifers, Araucaria, limited to the southern hemisphere, has scarcely less antiquity; Pinus reaches as far south as Cuba and Nicaragua.
Conifers are scantily represented by Callitris and Podocarpus, which is common to all three sub-regions; and Cycads by the endemic Encephalartos and Stangeria.
Of Cycads, Australia and New Caledonia have Cycas, and the former the endemic Macrozamia and Bowenia.
A distinct connexion between the flora of the peninsula and Ceylon and that of eastern tropical Africa is observable not only in the great similarity of many of the more truly tropical forms, and the identity of families and genera found in both regions, but in a more remarkable manner in the likeness of the mountain flora of this part of Africa to that of the peninsula, in which several species occur believed to be identical with Abyssinian forms. This connexion is further established by the absence from both areas of oaks, conifers and cycads, which, as regards the first two families, is a remarkable feature of the flora of the peninsula and Ceylon, as the mountains rise to elevations in which both of them are abundant to the north and east.
They are characterized by fine cycads (Zamites arcticus and Glossozamites Hoheneggeri), which also occur in the Urgonian strata of Wernsdorff.
Cordaites, a tall plant (20-30 ft.) with yucca-like leaves, was related to the cycads and conifers; the catkin-like inflorescence, which bore yew-like berries, is called Cardiocarpus.
Among other trees and shrubs may be mentioned the sumach, the date-palm, the plantain, various bamboos, cycads and the dwarf-palm, the last of which grows in some parts of Sicily more profusely than anywhere else, and in the desolate region in the south-west yields almost the only vegetable product of importance.
The plants of the Lower Gondwanas consist chiefly of acrogens(Equisetaceae and ferns)and gymnogens (cycads and conifers), the former being the more abundant.
The same classes of plants occur in the Upper Gondwanas; but there the proportions are reversed, the conifers, and still more the cycads, being more numerous than the ferns, whilst the Equisetaceae are but sparingly found.
It consists partly of sandstones with impressions of plants (cycads, ferns, &c.), and partly of clay-beds with coal.
It was the English botanist Robert Brown who first recognized this important distinguishing feature in conifers and cycads in 1825; he established the gymnospermy of these seed-bearing classes as distinct from the angiospermy of the monocotyledons and dicotyledons.
It is needless to discuss at length the origin of the Gymnosperms. The two views which find most favour in regard to the Coniferales and Cycadophyta are: (I) that both have been derived from remote filicinean ancestors; (2) that the cycads are the descendants of a fern-like stock, while conifers have been evolved from lycopodiaceous ancestors.
The line of descent of recent cycads is comparatively clear in so far as they have undoubted affinity with Palaeozoic plants which combined cycadean and filicinean features; but opinion is much more divided as to the nature of the phylum from which the conifers are derived.
The cycads constitute a homogeneous group of a few living members confined to tropical and sub-tropical regions.
In the genus Cycas the female flower is peculiar among cycads in consisting of a terminal crown of separate leaf-like carpels several inches in length; the apical portion of each carpellary leaf may be broadly triangular in form, and deeply dissected on the margins into narrow woolly appendages like rudimentary pinnae.
The male flower of Cycas conforms to the type of structure characteristic of the cycads, and consists of a long cone of numerous sporophylls bearing many oval pollen-sacs on their lower faces.
The stems of cycads are often described as unbranched; it is true that in comparison with conifers, in which the numerous branches, Stem springing from the main stem, give a characteristic form to the tree, the tuberous or columnar stem of the Cyca daceae constitutes a striking distinguishing feature.
Probably the oldest example of this genus in cultivation is in the Botanic Garden of Amsterdam, its age is considered by Professor de Vries to be about two thousand years: although an accurate determination of age is impossible, there is no doubt that many cycads grow very slowly and are remarkable for longevity.
The thick armour of petiole-bases enveloping the stem is a characteristic C y cad c an feature; in Cycas the alternation of scale-leaves and fronds is more clearly shown than in other cycads; in Encephalartos, Dioon, &e., From a photograph of a plant in the Peradeniya the persistent scale - leaf Gardens, Ceylon, by Professor R.
The Cyas type of frond, except as regards the presence of a midrib in each pinna, characterizes the cycads generally, except Bowenia and Stangeria.
In rare cases the pinnae of cycads are lobed or branched: in Dioon spinulosum (Central America) the margin of the segments bears numerous spinous processes; in some species of Encephalartos, e.g.
A feature of interest in connexion with the phylogeny of cycads is the presence of long hairs clothing the scale-leaves, and forming a cap on the summit of the stem-apex or attached to the bases of petioles; on some fossil cycadean plants these outgrowths have the form of scales, and are identical in structure with the ramenta (paleae) of the majority of ferns.
The male flowers of cycads are constructed on a uniform plan, and in all cases consist of an axis bearing crowded, spirally disposed sporophylls.
The cones of cycads attain in some cases (e.g.
Webber, and more recent work enables us to assume that all cycads produce ciliated male gametes.
Pollination in cycads has always been described as anemophilous, but according to recent observations by Pearson on South African species it seems probable that, at least in some cases, the pollen is conveyed to the ovules by animal agency.
The anatomical structure of the vegetative organs of recent cycads is of special interest as affording important evidence of relationship with extinct types, and with other groups of recent plants.
The leaftraces of cycads are remarkable both on account of their course and their anatomy.
The complicated girdle-like course is characteristic of the leaf-traces of most recent cycads, but in some cases, e.g.
In Zamia floridana, the traces are described by Wieland in his recent monograph on American fossil cycads (Carnegie Institution Publications, 1906) as possessing a more direct course similar to that in Mesozoic genera.
Similarly in the sporophylls of some cycads the bundles are endarch near the base and mesarch near the distal end of the stamen or carpel.
This is of interest from the point of view of the comparison of recent cycads with extinct species (Bennettites), in which the leaf-traces follow a much more direct course than in modern cycads.
Palaeozoic genera, has not entirely disappeared from the stems of modern cycads; but the mesarch bundle is now confined to the leaves and peduncles.
The roots of some cycads resemble the stems in producing several cambiumrings; they possess 2 to 8 protoxylem-groups, and are characterized by a broad pericyclic zone.
A common phenomenon in cycads is the production of roots which grow upwards (apogeotropic), and appear as coralline branched structures above the level of the ground; some of the cortical cells of these roots are hypertrophied, and contain numerous filaments of blue-green Algae (Nostocaceae), which live as endoparasites in the cell-cavities.
Surrounding the pitted wall of the ovum there is a definite layer of large cells, no doubt representing a tapetum, which, as in cycads and conifers, plays an important part in nourishing the growing egg-cell.
The spermatozoids constitute the most striking link with both cycads and ferns.
Cordaites is an extinct type which in certain respects resembles Ginkgo, cycads and the Araucarieae, but its agreement with true conifers is probably too remote to justify our attri buting much weight to the bearing of the morphology of its female flowers on the interpretation of that of the Coniferae.
(1898); Wieland, " American Fossil Cycads," Carnegie Institution Publication (1906); Stopes, " Beitrage zur Kenntnis der Fortpflanzungsorgane der Cycadeen," Flora (1904); Caldwell, " Microcycas Calocoma," Bot.
Xliv., 1907 (also papers on this and other Cycads in the Bot.