One lichen only, Strigula complanata, a tropical species, has been proved to be truly and constantly parasitic. It grows on the surface of thick leathery leaves such as those of Camellia[967], etc. and the alga and fungus both penetrate the epidermis and burrow beneath the cuticle and outer cells, causing them to become brown. It undoubtedly injures the leaves.
Friedrich[968] has given an isolated instance of the hold-fast hyphae of Usnea piercing through the cortex to the living tissue of the host, and not only destroying the middle lamella by absorption, but entering the cells. The Usnea plant was characterized by exceptionally vigorous growth. Practically all corticolous lichens are epiphytic and the injury they cause is of an accidental nature. Crustaceous species on the outer bark occupy the dead cortical layers and seem to be entirely harmless[969]. The larger foliose and fruticose forms are not so innocuous: by their abundant enveloping growth they hinder the entrance of air and moisture, and thus impede the life of the higher plant. Gleditsch[970], one of the earliest writers on Forestry, first indicated the possibly harmful effect of lichens especially on young trees and “in addition,” he says, “they serve as cover for large numbers of small insects which are hurtful in many ways to the trees.” Lindau[971] pointed out the damage done to pine-needles by Xanthoria parietina which grew round them like a cuff and probably choked the stomata, the leaves so clothed being mostly withered. Dufrenoy[972] states that he found the hyphae of a Parmelia entering a pine-needle by the stomata, and that the starch disappeared from the neighbouring parenchyma the cells of which tended to disintegrate.
It is no uncommon sight to see neglected fruit trees with their branches crowded with various lichens, Evernia prunastri, Ramalina farinacea, etc. Such lichens often find the lenticels a convenient opening for their hold-fasts and exercise a smothering effect on the trees. Lilian Porter[973] distinctly states that Ramalinae by their penetrating bases damage the tissues of the trees. The presence of lichens is however generally due to unhealthy conditions already at work. Friedrich[974] reported of a forest which he examined, in which the atmospheric moisture was very high, with the soil water scarce, that those trees that were best supplied with soil water were free from lichens, while those with little water at the base bore dead branches which gave foothold to a rich growth of the epiphytes.
Experiments to free fruit trees from their coating of lichens were made by Waite[975]. With a whitewash brush he painted over the infested branches with solutions of Bordeaux mixture of varying strength, and found that this solution, commonly in use as a fungicide, was entirely successful. The trees were washed down about the middle of March, and some three weeks later the lichens were all dead, the fruticose and foliose forms had changed in colour to a yellowish or brownish tint and were drooping and shrivelled.
Waite was of opinion that the lichens did considerable damage to the trees, but it has been held by others that in very cold climates they may provide protection against severe frost. Instances of damage are however asserted by Bouly de Lesdain[976]. The bark of willows he found was a favourite habitat of numerous lichens: certain species, such as Xanthoria parietina, completely surrounded the branches, closing the stomata; others, such as Physcia ascendens, by the mechanical strain of the rhizoids, first wet and then dry, gradually loosened the outer bark and gave entry to fungi which completed the work of destruction.
H. Gall-Formation
Several instances of gall-formation to a limited extent have been already noted as caused by parasitic fungi or lichens. Greater abnormality of development is induced in a few species by the presence of minute animals, mites, wood-lice, etc. Zopf[977] noted these deformations of the thallus in specimens of Ramalina Kullensis collected on the coasts of Sweden. The fronds were frequently swollen in a sausage-like manner, and branching was hindered or altogether prevented; apothecia were rarely formed, though pycnidia were abundant. Here and there, on the swollen portions of the thallus, small holes could be detected and other larger openings of elliptical outline, about 1-1-1/2 mm. in diameter, the margins of which had a nibbled appearance. Three types of small articulated animals were found within the openings: species of mites, spiders and wood-lice. Mites were the most constant and were more or less abundant in all the deformations; frequently a minute Diplopodon belonging to the genus Polyxenus was also met with.
Zopf came to the conclusion that the gall-formation was mainly due to the mites: they eat out the medulla and possibly through some chemical irritation excite the algal zone and cortex to more active growth, so that an extensive tangential development takes place. The small spiders may exercise the same power; evidently the larger holes were formed by them.
Later Zopf added to gall-deformed plants Ramalina scopulorum var. incrassata and R. cuspidata var. crassa. He found in the hollow swollen fronds abundant evidence of mites, but whether identical with those that attacked R. Kullensis could not be determined. These two Ramalinae are maritime species; they are morphologically identical, as are also the deformed varieties, and the presence of mites, excreta, etc., are plainly visible in our British specimens.
Bouly de Lesdain[978] found evidence of mite action in Ramalina farinacea collected from Pinus sylvestris on the dunes near Dunkirk. The cortex had been eaten off either by mites or by a small mollusc (Pupa muscorum) and the fronds had collapsed to a more or less convex compact mass. Somewhat similar deformations, though less pronounced, were observed in other Ramalinae.