Part 38

3. The sporangia are situated in continuous lines just inside the margin of the leaf.--Pteris: the sporangia form a continuous line along the entire margin of the leaf (Fig. 211 C), which bends over and covers the sporangia, forming a “false-indusium.” Pteridium has linear sori situated on a marginal vascular bundle, covered by two linear basal indusia, of which the outer is bent over like the edge of a leaf.--P. aquilinum (Bracken) has a wide-spreading rhizome with large alternate leaves, placed on opposite sides, at some distance apart. Only one leaf is developed from each branch every year.

Adiantum (Maiden-hair): sori on the underside of small portions of the edge of the leaf, which are bent over (false indusium). Cryptogramme (Allosorus), Cheilanthes.

4. The sori are oval or linear, situated on one side of the vascular bundle.--Asplenium (Fig. 212 A): sori linear; indusium with one of its edges attached at the external side. A. ruta muraria (Wall-Rue); A. septentrionale; A. trichomanes.--Athyrium: sori linear or curved; A. filix-fœmina (Lady-Fern).--Scolopendrium (Fig. 212 B): sori as in Asplenium, but situated in pairs across the lanceolate, entire leaves. Each sorus is covered on the external side by an indusium, whose free edges are parallel and approach each other. S. vulgare (Hart’s-tongue).--~Blechnum (B. spicant, Hard Fern; the fertile leaves differ from the barren, the pinnæ being narrower, while the underside is almost entirely covered with sori, and hence they are of a much darker brownish hue than the barren ones).--Ceterach: indusium rudimentary or absent.~

5. Sori circular and covered by a shield-like, or reniform indusium.--Aspidium (Fig. 211 B); the leaves wither away and leave no scar upon the root-stock. A. filix-mas (Male-Fern); A. spinulosum.--Phegopteris has no indusium, the withered bases of the leaf-stalks are persistent; P. dryopteris and P. polypodioides.

6. The indusium is situated below the sori, and has the shape of a one-sided scale (Cystopteris, Struthiopteris), or of a cup or cupule, which in Woodsia is sometimes fimbriate (Fig. 212 C, D).

7. The sori are situated on the margin of the leaf, and at the end of a vascular bundle. Indusium, semi-cupular. Davallia. Principally tropical species. 1 in S. Europe.

This order is the greatest, comprising about 2,800 species, the majority being perennial plants. A few are large, and known as Tree-Ferns.

As plants in conservatories and rooms the following are cultivated: species of Gymnogramme (tropical America), Lomaria, Nephrolepis, Pteris (P. serrulata, cretica).

Officinal. Aspidium filix-mas, rhizome and the withered petioles.--Species of Alsophila and Cibotium give Penghawar Djambi. The rhizome of Pteridium aquilinum, var. esculentum, contains so much starch that it is used as food.

The other orders of true Ferns deviate from the Polypodiaceæ, especially in the formation of the annulus, the bursting of the sporangium and its mode of attachment and development, and in the differences in the formation of the prothallium, etc. The principal are:--

Order 2. =Hymenophyllaceæ.= To this order belong the lowest and most Moss-like Ferns; the leaves, with the exception of the veins, are most frequently formed of only one layer of cells, and consequently stomata are wanting; the formation of the prothallium also somewhat resembles the Mosses. Sori marginal, on the extremities of the vascular bundles, and surrounded by a cupular indusium. The sporangia are sessile, with equatorial annulus. Hymenophyllum (H. tunbridgense, European). Trichomanes (T. speciosum, European). Species about 200, which live especially on rocks and trees in damp and shady tropical forests. Some have no roots.

Order 3. =Cyatheaceæ.= Annulus complete and oblique. To this order belong, principally, the tree-like Ferns with palm-like habit. The number of species is about 200, they are all tropical and form forests in some regions of Australia. Cibotium and Dicksonia have marginal sori, with cupular, basal indusium. (The stem of D. antarctica is covered with aerial roots.) Alsophila (without indusium); Cyathea with cupular, inferior indusium (Fig. 212 E).

Order 4. =Gleicheniaceæ.= Sporangia with equatorial annulus, and longitudinal dehiscence, most frequently groups of 3–4 in sori without indusium (Fig. 213). Gleichenia: the apical growth of the leaves continues for a long time.

Order 5. =Schizæaceæ.= Annulus apical. To this order belongs Aneimia, which is so commonly cultivated in conservatories. The two lowest pinnæ are metamorphosed, having no leaf parenchyma and being covered with sporangia. Schizæa. Mohria. Lygodium, a climber, whose leaves have unlimited growth and attain a length of several metres. About 70 species. Tropical.

Order 6. =Osmundaceæ.= The sporangia have at the apex a lateral group of strongly thickened cells, which gradually pass over into the ordinary cells. The sporangia open by a longitudinal cleft. Indusium wanting. Osmunda bears the sporangia upon peculiar, branched pinnæ, without parenchyma (the uppermost in the leaf). O. regalis (Royal-Fern): European.

Sub-Class 2. =Hydropterideæ= (formerly Rhizocarpeæ), =Water Ferns=.

The following further characteristics must be added to those given on page 205:--

=Sexual generation.= The MICROSPORES produce an extremely rudimentary prothallium, formed of only a single cell, and having also a very much reduced bicellular antheridium with a small number of spermatozoid mother-cells in each cell (in Salvinia 4, in Marsilia and Pilularia 16). In Salvinia the microspores remain embedded in a hard mucilaginous mass (at first frothy) which fills up the cavity of the sporangium. The prothallium must therefore grow out through this slime and also through the wall of the sporangium (Fig. 214), and it thus terminates in a relatively long cell.

In Marsilia the microspores are set free from the microsporangium, and the prothallia, with the antheridia, remain in them until the spermatozoids are liberated. The latter are spirally-twisted threads.

The MACROSPORES, on germination, give rise to a very reduced prothallium, which in Salvinia bears 3 archegonia; but, if these are not fertilised, the prothallium may continue to grow and become a fairly large, green body with several archegonia (Fig. 215 A, B). In Marsilia the prothallium is still more reduced, it is enclosed in the macrospore, and only bears one archegonium. The archegonia are similar in structure to those of the Ferns, but are smaller, and sunk more deeply in the tissue of the prothallium.

=The asexual generation= is developed from the fertilised egg-cell. It is a dorsiventral, horizontal shoot. In Salvinia it bears at first a shield-like leaf, the scutiform leaf (Fig. 215 C, a), which is succeeded by the ordinary foliage-leaves. The young plants of Marsilia, likewise, have less perfect leaves in the very early stage.

The formation of the sporangium is the same as in the Leptosporangiate Ferns. (The 16 spore-mother-cells originate from one central, tetrahedric archesporium.)

The Hydropterideæ are divided into 2 orders, the chief differences between them being found in the asexual generation.

Order 1. =Salviniaceæ.= This order more nearly approaches the true Ferns, especially so on account of the form of the indusium. Only one species is found in Europe, Salvinia natans (Fig. 217). This is a small, floating, annual, aquatic plant, entirely destitute of roots. The dorsiventral, horizontal stem bears two kinds of leaves, which are arranged in whorls of three. Two of these which turn upwards are oval, entire, “aerial foliage-leaves” (Fig. 217 B, b^2-b^3); the third, the “water-leaf” (b^1) is submerged and divided into a number of hair-like segments, similar to the submerged leaves in many aquatic plants, for instance, Water-buttercup (see also Fig. 215 C). The whorls of leaves alternate with each other; there are thus 4 rows of dorsally-placed aerial leaves, and two rows of ventrally-placed submerged leaves. The sporangia are situated in sori, each sorus being borne on a small column (receptacle or placenta) and enveloped by a cupular, but entirely closed indusium (Fig. 218). The sori are situated on the submerged leaves (Fig. 217 B, s-s) and are unisexual, i.e. each sorus contains microsporangia only, or macrosporangia.

Azolla belongs to this order. It is a very small, floating, tropical water-plant (America and East India), with horizontal, root-bearing stem. The stem branches profusely by lateral buds, and bears the two rows of leaves on its dorsal side, the roots on the ventral side. Each leaf is bifid, and divided into an upper dorsal, and a lower ventral portion. The upper segments float on the surface of the water and are arranged like tiles on a roof, each one overlapping its neighbour. In each floating segment a large cavity is found, in which Anabæna is always present. The lower segments are submerged.

Order 2. =Marsiliaceæ.= The characteristic feature of this order, and one not possessed by other Fern-like plants, is that the sori (2–many) are enveloped in leaf-segments which close round them and form a “sporocarp,” just in the same manner as the carpels, in the Angiospermous Flowering-plants, close round the ovules and form ovaries. The sori contain both micro-and macrosporangia. When the spores are ripe, the sporocarp opens in order to disperse the spores (Fig. 220).

The two genera (with 57 species, Temperate, Tropics) are land-and marsh-plants, whose dorsiventral, creeping stem bears roots on the under surface, and the leaves in two rows on the upper side (Figs. 219, 221). The leaves of Marsilia are compound, and divided into four small leaflets springing from the apex of the petiole (Fig. 219), and resemble the leaves of Oxalis. In the bud the leaves are circinate (Fig. 219 b), and at night they exhibit the well-known sleep-movements. The sporocarps are borne on the petioles of the fertile leaves, near their bases (Fig. 219 f); they are oblong and resemble small beans, the outer cells being hard and sclerenchymatous, while the inner ones are divided into a number of loculi arranged in two rows. On germination, water is absorbed, the two sides separate slightly, as valves (Fig. 220 A), and a long vermiform mass of gelatinous, parenchymatous cells (Fig. 220), swollen by the water, emerges, bearing a large number of sori arranged pinnately. Each sorus (sr) is covered by a thin indusium. (The thin covering may be considered an indusium physiologically, though not morphologically).

Marsilia quadrifolia, in Europe. Many species are found in Australia. The nutritious sporocarps of M. salvatrix were the means of saving the Burke expedition in the interior of Australia, and hence this species has earned its specific name.