I. GENERAL STATEMENT

A. Origin of Lichens

Though lichens are very old members of the vegetable kingdom, as symbiotic plants they yet date necessarily from a time subsequent to the evolution of their component symbionts. Phylogeny of lichens begins with symbiosis.

The algae, which belong to those families of Chlorophyceae and Myxophyceae that live on dry land, had become aerial before their association with fungi to form lichens. They must have been as fully developed then as now, since it is possible to refer them to the genus or sometimes even to the species of free-living forms. The fungus hyphae have combined with a considerable number of different algae, so that, even as regards the algal symbiont, lichens are truly polyphyletic in origin.

The fungus is, however, the dominant partner, and the principal line of development must be traced through it, as it provides the reproductive organs of the plant. Representatives of two great groups of fungi are associated with lichens: Basidiomycetes, found in only a few genera, and Ascomycetes which form with the various algae the great bulk of lichen families. In respect of their fungal constituents lichens are also polyphyletic, and more especially in the Ascolichens which can be traced back to several starting points. But though lichens have no common origin, the manner of life is common to them all and has influenced them all in certain directions: they are fitted for a much longer existence than that of the fungi from which they started; and both the thallus and the fruiting bodies—at least in the sub-class Ascolichens—can persist through great climatic changes, and can pass unharmed through prolonged periods of latent or suspended vitality.

Another striking note of similarity that runs through the members of this sub-class, with perhaps the exception of the gelatinous lichens, is the formation of lichen-acids which are excreted by the fungus. These substances are peculiar to lichens and go far to mark their autonomy. The production of the acids and the many changes evolved in the vegetative thallus suggest the great antiquity of lichens.

B. Algal Ancestors

It is unnecessary to look far for the algae as they have persisted through the ages in the same form both without and within the lichen thallus. By many early lichenologists the free-living algae, similar in type to lichen algae, were even supposed to be lichen gonidia in a depauperate condition and were, for that reason, termed by Wallroth “unfortunate brood-cells.” In the condition of symbiosis they may be considerably modified, but they revert to their normal form, and resume their normal life-history of spore production, etc., under suitable and free culture. The different algae taking part in lichen-formation have been treated in an earlier chapter[980].

C. Fungal Ancestors

a. Hymenolichens. The problem of the fungal origin in this sub-class is comparatively simple. It contains but three genera of tropical lichens which are all associated with Myxophyceae, and the fungus in them, to judge from the form and habit of the plants, is a member of the Thelephoraceae. It may be that Hymenolichens are of comparatively recent origin and that the fungi belonging to the Basidiomycetes had, in the course of time, become less labile and less capable of originating a new method of existence. Whatever the reason, they lag immeasurably behind Ascomycetes in the formation of lichens.

b. Ascolichens. Lichens are again polyphyletic within this sub-class. The main groups from which they are derived are evident. Whether there has been a series of origins within the different groups or a development from one starting point in each it would be difficult to determine. In any case great changes have taken place after symbiosis became established.

The main divisions within the Ascolichens are related to fungi thus:

Series1.Pyrenocarpineae}to Pyrenomycetes.
2.Coniocarpineae}
3.Graphidineaeto Hysteriaceae.
4.Cyclocarpineaeto Discomycetes.