The natural increase of lichen plants may primarily be sought for in the dispersal of the spores produced in the fruiting-bodies. These are ejected, as in fungi, by the pressure of the paraphyses on the mature ascus. The spores are then carried away by wind, water, insects, etc. In a few lichens gonidia are enclosed in the hymenium and are ejected along with the spores, but, in most, the necessary encounter with the alga is as fortuitous, and generally as certain, as the pollination of anemophilous flowers. A case of dispersal in Sagedia microspora has been described by Miyoshi[915] in which entire fruits, small round perithecia, were dislodged and carried away by the wind. The addition of water caused them to swell enormously and brought about the ejection of the spores. Areas covered by the thallus are also being continually enlarged by the spreading growth of the hypothallus.
a. Dispersal of Crustaceous Lichens. These lichens are distributed fairly equally on trees or wood (corticolous) and on rocks (saxicolous). Some species inhabit both substrata. As regards corticolous lichens that live on smooth bark such as hazel or mountain-ash, the vegetative body or thallus is generally embedded beneath the epidermis of the host. Soredia are absent and the thallus is protected from dispersal. In these lichens there is rather an abundant and constant formation of apothecia or perithecia.
Other species that affect rugged bark and are more superficial are less dependent on spore production. The thallus is either loosely granular, or is broken up into areolae. The areolae are each a centre of growth, and with an accession of moisture they swell up and exert pressure on each other. Parts of the thallus thus become loosened and are dislodged and carried away. If anchored on a suitable substratum they grow again to a complete lichen plant. Sorediate lichens are dependent almost wholly on these bud-like portions for increase in number; soredia are easily separated from the parent plant, and easily scattered. Darbishire[916] noted frequently that small Poduridae in moving over the surface of Pertusaria amara became powdered with soredia and very evidently took a considerable part in the dissemination of the species.
Crustaceous rock lichens are rarely sorediate, but they secure vegetative propagation[917] by the dispersal of small portions of the thallus. The thalli most securely attached are cracked into small areolae which, by unequal growth, become very soon lop-sided, or, by intercalary increase, form little warts and excrescences on their surface. These irregularities of development give rise to more or less tension which induces a loosening of the thallus from the substratum. Weather changes act similarly and gradually the areolae are broken off. Loosening influence is also exercised by the developing fruits, the expanding growth of which pushes aside the neighbouring tissues. Wind or water then carries away the thalline particles which become new centres of growth if a suitable substratum is reached.
b. Dispersal of Foliose Lichens. It is a matter of common observation that, in foliose lichens where fruits are abundant, there are few or no soredia and vice versa. In either case propagation is ensured. In addition to these obvious methods of increase many lichens form isidia, outgrowths from the thallus which are easily detached. Bitter[918] considers for instance that the coralloid branchlets, which occur in compact tufts on the thallus of Umbilicaria pustulata, are of immense service as organs of propagation. Apothecia and pycnidia are rarely present in that species, and the plant thus falls back on vegetative production. Slender crisp thalline outgrowths, easily separable, occur also on the edges of lobes, as in species of Peltigera, Platysma, etc.
Owing to the gelatinous character of lichen hyphae, the thallus quickly becomes soft with moisture and is then easily torn and distributed by wind, animals, etc. The action of lichens on rocks has been shown to be of a constantly disintegrating character, and the destruction of the supporting rock finally entails the scattering of the plant. This cause of dispersal is common to both crustaceous and foliose species. The older central parts of a lichen may thus have disappeared while the areolae on lobes of the circumference are still intact and in full vigour.
As in crustaceous lichens the increase in the area of growth may take place by means of the lichen mycelium which, originating from the rhizinae in contact with the substratum, spreads as a hypothallus under the shelter of the lobes and far beyond them. When algae are encountered a new lobe begins to form. The process can be seen perhaps most favourably in lichens on decaying wood which harbours moisture and thus enables the wandering hyphae to retain life.
c. Dispersal of Fruticose Lichens. Many of these lichens are abundantly fruited; in others soralia are as constantly developed. Species of Usnea, Alectoria, Ramalina and many Cladoniae are mainly propagated by soredia. They are all peculiarly liable to be broken and portions of the thallus scattered by the combined action of wind and rain.
Peirce[919] found that Ramalina reticulata ([Fig. 65]), of which the fronds are an open network, was mainly distributed by the tearing of the lichen in high wind. This takes place during the winter rains, when not only the lichen is wet and soft in texture, but when the deciduous trees are bare of leaves, at a season, therefore, when the drifting thalline scraps can again catch on to branch or stem. A series of observations on the dispersal of forms of long pendulous Usneas was made by Schrenk[920]. In the Middle and North Atlantic States of America these filamentous species rarely bear apothecia. The high winds break and disperse them when they are in a wet condition. They generally grow on Spruces and Firs, because the drifting filaments are more easily caught and entangled on short needles. The successive wetting and drying causes them to coil and uncoil, resulting in a tangle impossible to unravel, which holds them securely anchored to the support.