lived as hermits in forests, the Greeks also called them Hylobioi (cf. the Vāna-prasthās in Sanskrit writings). Diogenes Laërtius (ix. 61 and 63) refers to them, and asserts that Pyrrho of Elis, the founder of pure scepticism, came under their influence, and on his return to Elis imitated their habits of life, to what extent does not appear. Strabo (xv. 711, 714) divides them into Brahmans and Sarmans (or Shamans). See Jains.
GYMNOSPERMS, in Botany. The Gymnosperms, with the Angiosperms, constitute the existing groups of seed-bearing plants or Phanerogams: the importance of the seed as a distinguishing feature in the plant kingdom may be emphasized by the use of the designation Spermophyta for these two groups, in contrast to the Pteridophyta and Bryophyta in which true seeds are unknown. Recent discoveries have, however, established the fact that there existed in the Palaeozoic era fern-like plants which produced true seeds of a highly specialized type; this group, for which Oliver and Scott proposed the term Pteridospermae in 1904, must also be included in the Spermophyta. Another instance of the production of seeds in an extinct plant which further reduces the importance of this character as a distinguishing feature is afforded by the Palaeozoic genus Lepidocarpon described by Scott in 1901; this lycopodiaceous type possessed an integumented megaspore, to which the designation seed may be legitimately applied (see Palaeobotany: Palaeozoic).
As the name Gymnosperm (Gr. γυμνός, naked, σπέρμα, seed) implies, one characteristic of this group is the absence of an ovary or closed chamber containing the ovules. 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. As Sachs says in his history of botany, “no more important discovery was ever made in the domain of comparative morphology and systematic botany.” As Coulter and Chamberlain express it, “the habitats of the Gymnosperms to-day indicate that they either are not at home in the more genial conditions affected by Angiosperms, or have not been able to maintain themselves in competition with this group of plants.”
These naked-seeded plants are of special interest on account of their great antiquity, which far exceeds that of the Angiosperms, and as comprising different types which carry us back to the Palaeozoic era and to the forests of the coal period. The best known and by far the largest division of the Gymnosperms is that of the cone-bearing trees (pines, firs, cedars, larches, &c.), which play a prominent part in the vegetation of the present day, especially in the higher latitudes of the northern hemisphere; certain members of this class are of considerable antiquity, but the conifers as a whole are still vigorous and show but little sign of decadence. The division known as the Cycadophyta is represented by a few living genera of limited geographical range and by a large number of extinct types which in the Mesozoic era (see Palaeobotany: Mesozoic) played a conspicuous part in the vegetation of the world. Among existing Cycadophyta we find surviving types which, in their present isolation, their close resemblance to fossil forms, and in certain morphological features, constitute links with the past that not only connect the present with former periods in the earth’s history, but serve as sign-posts pointing the way back along one of the many lines which evolution has followed.
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: (1) 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 Cordaitales (see Palaeobotany: Palaeozoic) are represented by extinct forms only, which occupied a prominent position in the Palaeozoic period; these plants exhibit certain features in common with the living Araucarias, and others which invite a comparison with the maidenhair tree (Ginkgo biloba), the solitary survivor of another class of Gymnosperms, the Ginkgoales (see Palaeobotany: Mesozoic). The Gnetales are a class apart, including three living genera, of which we know next to nothing as regards their past history or line of descent. Although there are several morphological features in the three genera of Gnetales which might seem to bring them into line with the Angiosperms, it is usual to regard these resemblances as parallel developments along distinct lines rather than to interpret them as evidence of direct relationship.
Gymnospermae.—Trees or shrubs; leaves vary considerably in size and form. Flowers unisexual, except in a few cases (Gnetales) without a perianth. Monoecious or dioecious. Ovules naked, rarely without carpellary leaves, usually borne on carpophylls, which assume various forms. The single megaspore enclosed in the nucellus is filled with tissue (prothallus) before fertilization, and contains two or more archegonia, consisting usually of a large egg-cell and a small neck, rarely of an egg-cell only and no neck (Gnetum and Welwitschia). Microspore spherical or oval, with or without a bladder-like extension of the exine, containing a prothallus of two or more cells, one of which produces two non-motile or motile male cells. Cotyledons two or several. Secondary xylem and phloem produced by a single cambium, or by successive cambial zones; no true vessels (except in the Gnetales) in the wood, and no companion-cells in the phloem.
| I. | Pteridospermae (see Palaeobotany, Palaeozoic). | |
| II. | Cycadophyta. | |
| A. | Cycadales (recent and extinct). | |
| B. | Bennettitales (see Palaeobotany: Mesozoic). | |
| III. | Cordaitales (see Palaeobotany: Palaeozoic). | |
| IV. | Ginkgoales (recent and extinct). | |
| V. | Coniferales. | |
| A. | Taxaceae. | |
| B. | Pinaceae. | |
There is no doubt that the result of recent research and of work now in progress will be to modify considerably the grouping of the conifers. The family Araucarieae, represented by Araucaria and Agathis, should perhaps be separated as a special class and a rearrangement of other genera more in accord with a natural system of classification will soon be possible; but for the present its twofold subdivision may be retained.
| VI. | Gnetales. | |
| A. | Ephedroideae. | |
| B. | Gnetoideae. | |
| C. | Welwitschioideae (Tumboideae). | |
Cycadophyta.—A. Cycadales.—Stems tuberous or columnar, not infrequently branched, rarely epiphytic (Peruvian species of Zamia); fronds pinnate, bi-pinnate in the Australian genus Bowenia. Dioecious; flowers in the form of cones, except the female flowers of Cycas, which consist of a rosette of leaf-like carpels at the apex of the stem. Seeds albuminous, with one integument; the single embryo, usually bearing two partially fused cotyledons, is attached to a long tangled suspensor. Stems and roots increase in diameter by secondary thickening, the secondary wood being produced by one cambium or developed from successive cambium-rings.
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Fig. 1.—Stem of Cycas. F, foliage-leaf bases; S, scale-leaf bases. |
The cycads constitute a homogeneous group of a few living members confined to tropical and sub-tropical regions. As a fairly typical and well-known example of the Cycadaceae, a species of the genus Cycas (e.g. C. circinalis, C. revoluta, &c.) is briefly described. The stout columnar stem may reach a height of 20 metres, and a diameter of half a metre; it remains either unbranched or divides near the summit into several short and thick branches, each branch terminating in a crown of long pinnate leaves. The surface of the stem is covered with rhomboidal areas, which represent the persistent bases of foliage- and scale-leaves. In some species of Cycas there is a well-defined alternation of transverse zones on the stem, consisting of larger areas representing foliage-leaf bases, and similar but smaller areas formed by the bases of scale-leaves (F and S, fig. 1). The scale-leaves clothing the terminal bud are linear-lanceolate in form, and of a brown or yellow colour; they are pushed aside as the stem-axis elongates and becomes shrivelled, finally falling off, leaving projecting bases which are eventually cut off at a still lower level. Similarly, the dead fronds fall off, leaving a ragged petiole, which is afterwards separated from the stem by an absciss-layer a short distance above the base. In some species of Cycas the leaf-bases do not persist as a permanent covering to the stem, but the surface
