Fig. 18.—Melampsora salicina.
Cæomacei.—Of far greater interest are the Coniomycetous parasites on living plants. The present order includes those in which the spore[] is reduced to a single cell; and here we may observe that, although many of them are now proved to be imperfect in themselves, and only forms or conditions of other fungals, we shall write of them here without regard to their duality. These originate, for the most part, within the tissues of living plants, and are developed outwards in pustules, which burst through the cuticle. The mycelium penetrates the intercellular passages, and may sometimes be found in parts of the plants where the fungus does not develop itself. There is no proper excipulum or peridium, and the spores spring direct from a more compacted portion of the mycelium, or from a cushion-like stroma of small cells. In Lecythea, the sub-globose spores are at first generated at the tips of short pedicels, from which they are ultimately separated; surrounding these spores arise a series of barren cells, or cysts, which are considerably larger the true spores, and colourless, while the spores are of some shade of yellow or orange.[c] In Trichobasis, the spores are of a similar character, sub-globose, and at first pedicellate; but there are no surrounding cysts, and the colour is more usually brown, although sometimes yellow. In Uredo, the spores are at first generated singly, within a mother cell; they are globose, and either yellow or brown, without any pedicel. In Coleosporium, there are two kinds of spores, those of a pulverulent nature, globose, which are sometimes produced alone at the commencement of the season, and others which originate as an elongated cell; this becomes septate, and ultimately separates at the joints. During the greater part of the year, both kinds of spores are to be found in the same pustule. In Melampsora, the winter spores are elongated and wedge-shaped, compacted together closely, and are only matured during winter on dead leaves; the summer spores are pulverulent and globose, being, in fact, what were until recently regarded as species of Lecythea. In Cystopus, the spores are sub-globose, or somewhat angular, generated in a moniliform manner, and afterwards separating at the joints. The upper spore is always the oldest, continuous production of spores going on for some time at the base of the chain. Under favourable conditions of moisture, each of these spores, or conidia, as De Bary terms them, is capable of producing within itself a number of zoospores;[d] these ultimately burst the vesicle, move about by the aid of vibratile cilia, and at last settle down to germinate. Besides these, other reproductive bodies are generated upon the mycelium, within the tissues of the plant, in the form of globose oogonia, or resting spores, which, when mature, also enclose great numbers of zoospores. Similar oogonia are produced amongst the Mucedines in the genus Peronospora, to which De Bary considers Cystopus to be closely allied. At all events, this is a peculiarity of structure and development not as yet met with in any other of the Cæomacei. In Uromyces is the nearest approach to the Pucciniæi; in fact, it is Puccinia reduced to a single cell. The form of spore is usually more angular and irregular than in Trichobasis, and the pedicel is permanent. It may be remarked here, that of the foregoing genera, many of the species are not autonomous that have hitherto been included amongst them. This is especially true of Lecythea, Trichobasis, and, as it now appears, of Uromyces.[e]
Fig. 19.—Cystopus candidus.
Fig. 20.—Xenodochus carbonarius.
Pucciniæi.—This group differs from the foregoing chiefly in having septate spores. The pustules, or sori, break through the cuticle in a similar manner, and here also no true peridium is present. In Xenodochus, the highest development of joints is reached, each spore being composed of an indefinite number, from ten to twenty cells. With it is associated an unicellular yellow Uredine, of which it is a condition. Probably, in every species of the Pucciniæi, it may hereafter be proved, as it is now suspected, that an unicellular Uredine precedes or is associated with it, forming a condition, or secondary form of fruit of that species. Many instances of that kind have already been traced by De Bary,[f] Tulasne, and others, and some have been a little too rashly surmised by their followers. In Phragmidium, the pedicel is much more elongated than in Xenodochus, and the spore is shorter, with fewer and a more definite number of cells for each species; Mr. Currey is of opinion that each cell of the spore in Phragmidium has an inner globose cell, which he caused to escape by rupture of the outer cell wall as a sphæroid nucleus,[g] leading to the inference that each cell has its own individual power of germination and reproduction. In Triphragmium, there are three cells for each spore, two being placed side by side, and one superimposed. In one species, however, Triphragmium deglubens (North American), the cells are arranged as in Phragmidium, so that this represents really a tricellular Phragmidium, linking the present with the latter genus. In Puccinia the number of species is by far the most numerous; in this genus the spores are uniseptate, and, as in all the Pucciniæi, the peduncles are permanent. There is great variability in the compactness of the spores in the sori, or pulvinules. In some species, the sori are so pulverulent that the spores are as readily dispersed as in the Uredines, in others they are so compact as to be separated from each other with great difficulty. As might be anticipated, this has considerable effect on the contour of the spores, which in pulverulent species are shorter, broader, and more ovate than in the compact species. If a section of one of the more compact sori be made, it will be seen that the majority of the spores are side by side, nearly at the same level, their apices forming the external surface of the sori, but it will not be unusual to observe smaller and younger spores pushing up from the hymenial cells, between the peduncles of the elder spores, leading to the inference that there is a succession of spores produced in the same pulvinule. In Podisoma, a rather anomalous genus, the septate spores are immersed in a gelatinous stratum, and some authors have imagined that they have an affinity with the Tremellini, but this affinity is more apparent than real. The phenomena of germination, and their relations to Rœstelia, if substantiated, establish their claim to a position amongst the Pucciniæi.[h] It seems to us that Gymnosporangium does not differ generically from Podisoma. In a recently-characterized species, Podisoma Ellisii, the spores are bi-triseptate. This is, moreover, peculiar from the great deficiency in the gelatinous element. In another North American species, called Gymnosporangium biseptatum, Ellis, which is distinctly gelatinous, there are similar biseptate spores, but they are considerably broader and more obtuse. In other described species they are uniseptate.
Fig. 21.—Phragmidium bulbosum.