HYDROPTERIDEAE. Two very distinct orders of heterosporous Filicales, the Salviniaceae and the Marsiliaceae, are included in this group. The difficulty of determining their exact relationship to the other orders of Ferns is increased by the more or less completely aquatic habit of the plants and the modifications and reductions in structure associated with this. The absence of an annulus from their indehiscent sporangia makes it impossible to compare them with the other Ferns in respect of this important character. It has been suggested with considerable probability that the Marsiliaceae are allied to the Schizaeaceae, while the Salviniaceae may possibly be related to the Hymenophyllaceae or to some other family of the Gradatae. Space will only permit of a brief general account of the more obvious features of the several genera, the structure and lifehistory of which are known in great detail. Unlike as they are in many respects, the two orders agree in being heterosporous. The microspores on germination produce a small, greatly reduced male prothallus bearing one or two antheridia which give rise to a number of spirally coiled, multiciliate spermatozoids. The single large megaspore contained in each megasporangium produces a small prothallus, which bears one or a few archegonia; these are exposed on the surface of the prothallus at the summit of the germinated megaspore (fig. i, i).
i. The Salviniaceae include the two genera Salvinia (fig. 10) and Azolla. The small dorsiventral plants are in both cases floating aquatics. Azolla has roots depending from the lower surface of the stem into the water, while these organs are completely wanting in Salvinia, their place being taken functionally by highly divided leaves borne on the ventral surface of the stem. Nostoc colonies are constantly present in a special cavity of the dorsal lobe of the leaf in Azolla. The sporangia in both genera are associated in sori enclosed by indusia springing from the base of the receptacle. In Salvinia (fig. 2, h) the sori are borne towards the base of the submerged leaves, in Azolla on the reduced ventral lobe of the leaf. They consist either of microspprangia or megasporangia, which are arranged in basipetal succession on the receptacle. In the megasorus of Azolla there is only the one terminal, functional sporangium. The microspores are united by means of hardened protoplasm into one or more manses, while the solitary megasporcs have a more or less complicated episporium.
(Reduced. After BLschoff from Strasburger's Lehrbuch der Botatuk.)
FIG. 10. Salvinia natans. A, From the side. B, From above.
2. The Marsiliaceaealso include two genera, Marsilia and Pilularia, the latter of which is found in Britain. The plants grow as a rule in marshy places, though some species of Marsilia are xerophytic. The creeping stem produces roots from the ventral surface and leaves from the dorsal surface; the leaves when young are circinately coiled. The leaves are simple and linear in Pilularia, but in Marsilia bear a pinnate four-lobed lamina. The highly specialized sporocarps are borne on the basal portions of the leaves, as a rule singly, but in some species of Marsilia in numbers. The development of the sporocarp snows that it corresponds to a pinna, although when mature it may appear to occupy a -ventral position in relation to the vegetative portion of the leaf. It has a complicated structure in both genera; in Pilularia its shape is nearly spherical, while in Marsilia it is elongated and bean-shaped. The sori are developed in depressions and are thus protected within the resistent outer wall of the sporocarp. There are usually four sori in Pilularia, while in Marsilia they form two longitudinal rows. Each sorus includes both microsporangia, with numerous spores, and megasporangia, each of which contains a single megaspore with a complicated wall. Enclosed within the sporocarp they can endure a period of drought, but on the return of moist conditions are extruded from the sporocarp by the swelling of a special mucilaginous tissue and the spores become free. The development of the prothalli is in general similar to that of the S.ilviniureae, though the resemblance may behomoplastic. The stem in the less reduced forms is solenostelic with sclerenchymatous ground tissue occupying the centre of the stele.
In the absence of direct evidence from Palaeobotany, and bearing in mind the modifications associated with adaptation to an aquallw life in other plants, the recognition of any more definite affinity for these heterosporous ferns than that indicated above appears to be inadvisable. Further evidence is necessary before they can be removed from such a position of convenience as is assigned to them here and placed in proper relation to the series of the Filicaceae.
The several phyla of Pteridophyta having now been briefly described, their relationship to one another remains for conPhyhgeay sideration. The available evidence does not suffice to solve this question, although certain indications exist. In the earliest land vegetations of which we have any sufficient record specialized forms of Equisetales, Lycopodiales, Sphenophyllales and Filicales existed, so that we are reduced to hypotheses founded on the careful comparison of the recent and extinct members of these groups. In this connexion it may be pointed out that the fuller study of the extinct forms has as yet been of most use in emphasizing the difficulty of the questions at issue. It has thus led to a condition of uncertainty as regards the relationship of the great groups of Vascular Cryptogams, in which, however, lies the hope of an ultimate approach to a satisfactory solution. The study of the Sphenophyllales, however, as has been pointed out above, appears to indicate that the Equisetales and Lycopodiales may be traced back to a common ancestry. As to the relationship of the Filicales to the other phyla, evidence from extinct plants appears to be wanting.
If, as has been suggested by Bower, the strobiloid types are relatively primitive, the large-leaved Pteridophyta must be supposed to have arisen early from such forms. The question cannot be discussed fully here, but enough has been said above to show that in the light of our present knowledge the main phyla of the Vascular Cryptogams cannot be placed in any serial relationship to one another.
It may even be regarded as an open question whether some of them may not have arisen independently and represent parallel lines of evolution from Bryophytic or Algal forms. This leads us to consider the question whether any indications exist as to the manner in which the Pteridophyta arose. It will be evident that no direct record of this evolution can be expected, and recourse must be had to hypotheses founded on the indirect evidence available. There appears to be no reason to doubt that the sexual generation is homologous with the thallus of a Liverwort, or of such an Alga as Coleochaete. It is with regard to the origin of the spore-bearing generation of the Pteridophyta that differences of opinion exist. This, though at first dependent on the prothallus, soon becomes independent. It may be regarded as derived from a wholly dependent sporogonium not unlike that of some of the simpler Bryophyta; the latter are assumed to have arisen from primitive Algal forms, in which, as the first step in the interpolation of the second generation in the life cycle, the fertilized ovum gave rise to a group of swarm spores, each of which developed into a new sexual plant. On this view the origin of the sporophyte is looked for in the gradual development of sterile tissue in the generation arising from the fertilized ovum, and a consequent postponement of spore- formation. Certain green Algae (e.g. Oedogonium, Coleochaete), the Bryophyta, and the simpler Pteridophyta, such as Phylloglossum, have been regarded as illustrating the method of progression, though there is no reason to regard the existing forms as constituting an actual series. For a discussion of this view, which regards the alternation of generations in Pteridophytes as antithetic and the. two generations as not homologous with one another, reference may be made to the works of Celakovsky and Bower. Although the antithetic theory is supported by many facts regarding the lifehistory and structure of the group of plants under consideration, it is quite possible that a stage in which the sporophyte was wholly dependent on the gametophyle may never have been passed through in their evolution. The spore-bearing generation may throughout its phylogenetic history have been independent at one part of its life, and have been derived by modification of individuals homologous with those of the sexual generation, and not by the progressive sterilization of a structure the whole of which was originally devoted to asexual reproduction. A number of facts regarding the Algae, and also those relating to such deviations from the normal life cycle as apogamy or apospory, may be regarded as lending support to this view, which, in contrast to the theory of antithetic alternation, has been called that of homologous alternation. Without entering further into the discussion of these alternative theories, for which the literature of the subject must be consulted, it may be pointed out that on the latter view the strobiloM forms of Pteridophyta would not necessarily be regarded as primitive relatively to the large-leaved forms, and also that the early stages of the origin of the sporophyte in the two cases may have proceeded on different lines.
Another question of great interest, which can only be touched upon here and may fitly close the consideration of this division of the Vegetable Kingdom, concerns the evidence as to the derivation of higher groups from the Pteridophyta. The most important positive evidence on this point indicates that the most ancient Gymnosperms were derived from the Filicales rather than from any other phylum of the Vascular Cryptogams. Extinct forms are known intermediate between the Ferns and the Cycads, and a number of these have been shown to bear seeds and must be classed as Pteridospermae. These forms will, however, be found discussed in the articles treating of extinct plants and the Gymnosperms, but their recognition will serve to emphasize, in conclusion, the important position the Pteridophyta hold with regard to the existing flora.
Cultivation. Numerous species of ferns, both temperate and tropical, are cultivated as valued ornamental plants. Species of the other groups are occasionally grown for scientific purposes in the larger botanic gardens, but their cultivation, which often presents special difficulties, need not be referred to here. While a number of ferns can be multiplied vegetatively, by buds formed on the leaves and in other ways, the regular mode of propagation is by sowing the spores shed from the ripe sporangia. The spores should be thinly sprinkled on the surface of the soil in well-drained pots, which should stand in saucers filled with water and be covered with glass plates. After the prothalli have attained some size and bear sexual organs the pots should be occasionally sunk in water so as to flood the prothalli for a few minutes and facilitate fertilization. The young plants developed on the prothalli should be carefully pricked out into other pans and later transferred to 3-in. pots. When the pots are fairly filled with roots the plants may be shifted into larger ones.
The best time for a general repotting of ferns is in spring, just before growth commences. Those with creeping rhizomes can be propagated by dividing these into well-rooted portions, and, if a number of crowns is formed, they can be divided at that season. In most cases this can be performed with little risk, but the Gleichenias, for example, must only be cut into large portions, as small divisions of the rhizomes are almost certain to die; in such cases, however, the points of the rhizomes can be led over and layered into small pots, several in succession, and allowed to remain unsevered from the parent plant until they become well rooted. In potting the well-established plants, and all those of considerable size, the soil should be used in a rough turfy state, not sifted but broken, and one-sixth of broken crocks or charcoal and as much sand as will insure free percolation should be mixed with it.
The stove ferns require a day temperature of 65 to 75, but do not thrive in an excessively high or close dry atmosphere. They require only such shade as will shut out the direot rays of the Sun, and, though abundant moisture must be supplied, the atmosphere should not be loaded with it. The water used should always be at or near the temperature of the house in which the plants are growing. Some ferns, as the different kinds of Gymnogrammae and Ctieilanthes, prefer a drier atmosphere than others, and the former do not well bear a lower winter temperature than about 60 by night. Most other stove ferns, if dormant, will bear a temperature as low as 55 by night and 60 by day from November to February. About the end of the latter month the whole collection should be turned out of the pots and redrained or repotted into larger pots as required. This should take place before growth has commenced. Towards the end of March the night temperature may be raised to 60, and the day temperature to 70 or 75, the plants being shaded in bright weather. Such ferns as Gymnogrammas, which have their surface covered with golden or silver powder, and certain species of scalysurfaced Cheilanlhes and Nothochlaena, as they cannot bear to have their fronds wetted, should never be syringed ; but most other ferns may have a moderate sprinkling occasionally (not necessarily daily) and as the season advances sufficient air and light must be admitted.
AUTHORITIES. Scott, Structural Botany: Flowerless Plants (London, 1896), Studies in Fossil Botany (Edinburgh, 1900) ;* Campbell, Mosses and Ferns (London, 1895);* Engler and Prantl, Die naturlichen Pflanzenfamilien (Theil i. Abth. 4.; Leipzig, 1898-1902); Bower, The Origin of a Land Flora (London, 1908); Goebel, Organography of Plants (Oxford, 1905) ; Hooker and Baker, Synopsis Filicum (London, 1874); Baker, Fern Allies (London, 1887); Christ, Die F,ankrduter der Erde (Jena, 1897); Seward, Fossil Botany, vol. i. (Cambridge, 1898). In those works marked with an asterisk copious references to the recent literature of the subject will be found.
(W. H. L.)
Note - this article incorporates content from Encyclopaedia Britannica, Eleventh Edition, (1910-1911)