Lichens

by Annie Lorrain Smith

The conception of Graphidineae as a whole is puzzling, but one or other characteristic has brought the various members within the series. It is in this respect an epitome of the lichen class of which the different groups, with all their various origins and affinities, yet form a distinct and well-defined section of the vegetable kingdom.

d. Cyclocarpineae. This is by far the largest series of lichens. The genera are associated with algae belonging both to the Myxophyceae and the Chlorophyceae, and from the many different combinations are produced great variations in the form of the vegetative body. The fruit is an emergent, round or roundish disc or open apothecium in all the members of the series except Pertusariaceae, where it is partially immersed in thalline “warts.” In its most primitive form, described as “biatorine” or “lecideine,” it may be soft and waxy (Biatora) or hard and carbonaceous (Lecidea), in the latter the paraphyses being mostly coloured at the tips; these are either simple or but sparingly branched, so that the epithecium is a comparatively slight structure. The outer sterile tissue forms a protective wall or “proper margin” which may be entirely pushed aside, but generally persists as a distinct rim round the disc.

A great advance within the series arose when the gonidial elements of the thallus took part in fruit-formation. In that case not only is the hymenium generally subtended by a layer of algae, but thalline tissue containing algae grows up around the fruit, and forms a second wall or thalline margin. This type of apothecium, termed “lecanorine,” is thus intimately associated with the assimilating tissue and food supply, and it gains in capacity of ascus renewal and of long duration. This development from non-marginate to marginate ascomata is necessarily an accompaniment of symbiosis.

There is no doubt that the Cyclocarpineae derive from some simple form or forms of Discomycete in the Patellariaceae. The relationship between that family and the lower Lecideae is very close. Rehm[993] finds the direct ancestors of Lecidea itself in the fungus genus, Patinella, in which the apothecia are truly lecideine in character—open, flat and slightly margined, the hypothecium nearly always dark-coloured and the paraphyses branched, septate, clavate and coloured at the tips, forming a dark epithecium. More definitely still he describes Patinella atroviridis, a new species he discovered, as in all respects a Lecidea, but without gonidia.

In the crustaceous Lecideaceae, a number of genera have been delimited on spore characters—colourless or brown, and simple or variously septate. In Patellariaceae as described by Rehm are included a number of fungus genera which correspond to these lichen genera. Only two of them—Patinella and Patellaria—are saprophytic; in all the other genera of the family, the species with very few exceptions are parasitic on lichens: they are parasymbionts sharing the algal food supply; in any case, they thrive on a symbiotic thallus.

Rehm unhesitatingly derives the corresponding lichen genera from these fungi. He takes no account of the difficulty that if these parasitic (or saprophytic) fungi are primitive, they have yet appeared either later in time than the lichens on which they exist, or else in the course of ages they have entirely changed their substratum.

He has traced, for instance, the lichen, Buellia, to a saprophytic fungus species, Karschia lignyota, to a genus therefore in which most of the species are parasitic on lichens and have generally been classified as parasitic lichens. There is no advance in apothecial characters from the fungus, Karschia, to Buellia, merely the change to symbiosis. It therefore seems more in accordance with facts to regard Buellia as a genus evolved within the lichen series from Patinella through Lecidea, and to accept these species of Karschia on the border line as parasitic, or even as saprophytic, reversions from the lichen status. We may add that while these brown-spored lichens are fairly abundant, the corresponding athalline or fungus forms are comparatively few in number, which is exactly what might be expected from plants with a reversionary history.

Occasionally in biatorine or lecideine species with a slight thalline development all traces of the thallus disappear after the fructification has reached maturity. The apothecia, if on wood or humus, appear to be saprophytic and would at first sight be classified as fungi. They have undoubtedly retained the capacity to live at certain stages, or in certain conditions, as saprophytes.

The thallus disappears also in some species of the crustaceous genera that possess apothecia with a thalline margin, and the fruits may be left stranded and solitary on the normal substratum, or on some neighbouring lichen thallus where they are more or less parasitic; but as the thalline margin persists, there has been no question as to their nature and affinity.

Rehm suggests that many species now included among lichens may be ultimately proved to be fungi; but it is equally possible that the reverse may be the case, as for instance Bacidia flavovirescens, held by Rehm and others to be a parasitic fungus species, but since proved by Tobler[994] to be a true lichen.