In the last family, the Ascomycetes, we shall meet with a very great variety of forms, all agreeing in producing sporidia contained in certain cells called asci, which are produced from the hymenium. In some of these, the asci are evanescent, but in the greater number are permanent. In Onygenei, the receptacle is either club-shaped or somewhat globose, and the peridium is filled with branched threads, which produce asci of a very evanescent character, leaving the pulverulent sporidia to fill the central cavity. The species are all small, and singular for their habit of affecting animal substances, otherwise they are of little importance. The Perisporiacei, on the other hand, are very destructive of vegetation, being produced, in the majority of cases, on the green parts of growing plants. To this order the hop mildew, rose mildew, and pea mildew belong. The mycelium is often very much developed, and in the case of the maple, pea, hop, and some others, it covers the parts attacked with a thick white coating, so that from a distance the leaves appear to have been whitewashed. Seated on the mycelium, at the first as little orange points, are the perithecia, which enlarge and become nearly black. In some species, very elegant whitish appendages radiate from the sides of the perithecia, the variations in which aid in the discrimination of species. The perithecia contain pear-shaped asci, which spring from the base and enclose a definite number of sporidia.[R] The asci themselves are soon dissolved. Simultaneously with the development of sporidia, other reproductive bodies are produced direct from the mycelium, and in some species as many as five different kinds of reproductive bodies have been traced. The features to be remembered in Perisporiacei, as forming the basis of their classification, are, that the asci are saccate, springing from the base of the perithecia, and are soon absorbed. Also that the perithecia themselves are not perforated at the apex.
The four remaining orders, though large, can be easily characterized. In Tuberacei, all the species are subterranean, and the hymenium is mostly sinuated. In Elvellacei, the substance is more or less fleshy, and the hymenium is exposed. In Phacidiacei, the substance is hard or leathery, and the hymenium is soon exposed. And in Sphæriacei, although the substance is variable, the hymenium is never exposed, being enclosed in perithecia with a distinct opening at the apex, through which the mature spores escape. Each of these four orders must be examined more in detail. The Tuberacei, or subterranean Ascomycetes, are analogous to the Hypogæi of the Gasteromycetes. The truffle is a familiar and highly prized example. There is a kind of outer peridium, and the interior consists of a fleshy hymenium, more or less convoluted, sometimes sinuous and confluent, so as to leave only minute elongated and irregular cavities, and sometimes none at all, the two opposing faces of the hymenium meeting and coalescing.[S] Certain privileged cells of the hymenium swell, and ultimately become asci, enclosing a definite number of sporidia. The sporidia in many cases are large, reticulated, echinulate or verrucose, and mostly somewhat globose. In the genus Elaphomyces, the asci are more than commonly diffluent.
The Elvellacei are fleshy in substance, or somewhat waxy, sometimes tremelloid. There is no peridium, but the hymenium is always exposed. There is a great variety of forms, some being pileate, and others cup-shaped, as there is also a great variation in size, from the minute Peziza, small as a grain of sand, to the large Helvella gigas, which equals in dimensions the head of a child. In the pileate forms, the stroma is fleshy and highly developed; in the cup-shaped, it is reduced to the external cells of the cup which enclose the hymenium. The hymenium itself consists of elongated fertile cells, or asci, mixed with linear thread-like barren cells, called paraphyses, which are regarded by some authors as barren asci. These are placed side by side in juxtaposition with the apex outwards. Each ascus contains a definite number of sporidia, which are sometimes coloured. When mature, the asci explode above, and the sporidia may be seen escaping like a miniature cloud of smoke in the light of the mid-day sun. The disc or surface of the hymenium is often brightly coloured in the genus Peziza; tints of orange, red, and brown having the predominance.
In Phacidiacei, the substance is hard and leathery, intermediate between the fleshy Elvellacei and the more horny of the Sphæriacei. The perithecia are either orbicular or elongated, and the hymenium soon becomes exposed. In some instances, there is a close affinity with the Elvellacei, the exposed hymenium being similar in structure, but in all the disc is at first closed. In orbicular forms, the fissure takes place in a stellate manner from the centre, and the teeth are reflexed. In the Hysteriacei, where the perithecia are elongated, the fissure takes place throughout their length. As a rule, the sporidia are more elongated, more commonly septate, and more usually coloured, than in Elvellacei. Only a few solitary instances occur of individual species that are parasitic on living plants.
Fig. 42.—Sphæria aquila.
In the Sphæriacei, the substance of the stroma (when present) and of the perithecia is variable, being between fleshy and waxy in Nectriei, and tough, horny, sometimes brittle, in Hypoxylon. A perithecium, or cell excavated in the stroma which fulfils the functions of a perithecium, is always present. The hymenium lines the inner walls of the perithecium, and forms a gelatinous nucleus, consisting of asci and paraphyses. When fully mature, the asci are ruptured and the sporidia escape by a pore which occupies the apex of the perithecium. Sometimes the perithecia are solitary or scattered, and sometimes gregarious, whilst in other instances they are closely aggregated and immersed in a stroma of variable size and form. Conidia, spermatia, pycnidia, &c., have been traced to and associated with some species, but the history of others is still obscure. Many of the coniomycetous forms grouped under the Sphæronemei are probably conditions of the Sphæriacei, as are also the Melanconiei, and some of the Hyphomycetes. A very common fungus, for instance, which is abundant on sticks and twigs, forming rosy or reddish pustules the size of a millet seed, formerly named Tubercularia vulgaris, is known to be the conidia-bearing stroma of the sphæriaceous fungus, Nectria cinnabarina;[T] and so with many others. The following are the technical characters of the family:—
Fruit consisting of sporidia, mostly definite, contained in asci, springing from a naked or enclosed stratum of fructifying cells and forming a hymenium or nucleus = Ascomycetes.
If the characters of the different families are borne in mind, there will be but little difficulty in assigning any fungus to the order to which it belongs by means of the foregoing remarks. For more minute information, and for analytical tables of the families, orders, and genera, we must refer the student to some special systematic work, which will present fewer difficulties, if he keeps in mind the distinctive features of the families.[U]
To assist in this we have given on the following page an analytical arrangement of the families and orders, according to the system recognized and adopted in the present volume. It is, in all essential particulars, the method adopted in our “Handbook,” based on that of Berkeley’s “Introduction” and “Outlines.”