Bible Encyclopedias
Lichens

Conidia

Contrary to the behaviour of the non-lichen forming Ascomycetes the lichen-fungi show very few cases of ordinary conidial formation. Bornet describes free conidia in Arnoldia minitula, and Placodium decipiens and Conidia-formation has been described by Neubner in the Caliciae.

Spermatia

In the majority of genera of lichens small flask-shaped structures are found embedded in the thallus (fig. 13). These were investigated by Tulasne in 1853, who gave them the name spermogonia The lower, ventral portion of the sperm09' gonium is lined by delicate hyphae, the sterigmata, which give origin to minute colourless cells, the spermatia. The sterigmata are either simple (fig. 13, C) or septate - the socalled arthrosterigmata (fig. 13, B). The spermogonia open by a small pore at the apex, to wards which the sterig- ca mata converge and through which the sper matia escape (fig. 13). There are two views as to the nature of the spermatia. In one view they are mere asexual conidia, and the term pycnoconidia is accordingly applied since they are borne in structures like the non-sexual pycnidia of other fungi. In the other view the spermatia are the male sexual cells and thus A, Optical longitudinal section of the ex are rightly named; it tremity of a thin branch of the thallus should, however, be which has become transparent in pointed out that this solution of potash.

was not the view of B, Transverse section through a stronger Tulasne, though we owe branch with the point of origin of an to him the designation adventitious branch (sa). which carries with it r, Cortical layer.

the sexual significance. m, Medullary layer.

The question is one x, Stout axile strand.

very difficult to settle g, The algal zone (Cystococcus). owing to the fact that s, Apex of the branch.

the majority of sper matia appear to be functionless. In favour of the conidial view is the fact that in the case of Collema and a few other forms the spermatia have been made to germinate in artificial cultures, and in the case of Calicium parietinum Moller succeeded in producing a spermogonia bearing thallus from a spermatium. For the germination of the spermatia in nature there is only the observation of Hedlund, that in Catillaria denigrata and C. prasena a thallus may be derived from the spermatia under natural conditions. In relation to the view that the spermatia are sexual cells, or at least were primitively so, it must be pointed out that although the actual fusion of the spermatial nucleus with a female nucleus has not been observed, yet in a few cases the spermatia have been seen to fuse with a projecting portion (trichogyne) of the ascogonium, as in Collema and Physcia, and there is very strong circumstantial evidence that fertilization takes place (see later in section on development of ascocarp). The resemblance of the spermatia and spermogonia to those of Uredineae should be pointed out, where also there is considerable evidence for their original sexual nature, though they appear in that group to be functionless in all cases. The observations of Moller, &c., on the germination cannot be assumed to negative the sexual hypothesis for the sexual cells of Ulothrix and Ectocarpus, for example From Strasburger's Lehrbuch der Botanik, by permission of Gustav Fischer.

FIG. 6. - Cladonia rangiferina. (Nat. size.) A, Sterile.

B, With ascus-fruit at the ends of the branches.

From Strasburger's Lehrbuch der Botanik, by permission of Gustav Fischer.

FIG. 5. - Usnea barbata. (Nat. size.) ap, Apothecium.

After Sachs, from De Bary's Vergleichende Morphologie and Biologie der Pilze, Mycetozoen and Bacterien, by permission of Wilhelm Engelmann.

FIG. 8. - Usnea barbata. (Mag. nearly 100 times.) are able to develop with or without fusion. The most satisfactory view in the present state of our knowledge seems to be that the sper matia are male cells which, while retaining their fertiliz ing action in a few cases are now mainly functionless. The female sexual organs, the ?, ascogonia, would thus in the majority of cases develop by the aid of some reduced sexual process or the ascocarps be developed without relation to sexual organs. A further argument in support of this view is that it is in complete agreement with what we know of the sexuality of the ordinary, free-living ascomycetes, where we find both normal and reduced forms (see Fungi).

Fruit Bodies

We find two chief types of fruit bodies in the lichens, the perithecium and apothecium; the first when the fungal element is a member of the Pyrenomycetes division of the Ascomycetes, the second when the fungus belongs to the Discomycetes division. In the two genera of lichens - the Basidiolichens - i n which the fungus is a member of the Basidiomycetes, we have the fructification characteristic of that class of fungi: these are dealt with separately. The perithecium is very constant in form and since the gonidia take no part variations are of value in classification some more details may be added.

They present various shapes, of which the following are the principal: (a) peltate, which are large, rounded, without any distinct thalline margin 1 (e.g. Usnea, Peltigera); (b) lecanorine, or scutelliform, which are orbicular and surrounded by a distinct, more or less prominent thalline margin (e.g. Parmelia, Lecanora), having sometimes also in addition a proper one 1 (e.g. Thelotrema, Urceolaria); (c) lecideine, or patelliform, which are typically orbicular, with only a proper margin (e.g. Lecidea), sometimes obsolete, and which are occa sionally irregular in shape, angular or flexuose (e.g. Lecidea jurana, L. myrmecina), or complicated and gyrose (e.g. Gyro- phora), and even After Schwendener, from De Bary's Vergleichende Mor s t i p i t a t e (e.g. phologie permiss on und Biologic Wilhelm deer Pilz Engelmann. Mycetozoen and Bacterien, by Baeomyces); (d) lirelliform, which FIG. 12. - Usnea barbata. (Mag. more than are of very, irregu500 times.) lar figure, elonc, An isolated mature soredium, with an algal gated, branched or cell (Pleurococcus) in the envelope or hyphae. gate flexuose, with only d, Another with several algal cells in optical a proper margin longitudinal section.

(e. g. Xylographa, e, f, Two soredia in the act of germinating; the Graphis, &c.) or hyphal envelope has grown out below into G none (e.g. some rhizoid branches, and above shows already rthoniae), and the structure of the apex of the thallus (see A often very variable fig. 9).

even in the same species. In colour the apothecia are extremely variable, and it is but rarely that they are the same colour as the thallus (e.g. Usnea, Ramalina). Usually they are of a different colour, and may be black, brown, yellowish, or also less frequently rose-coloured, rusty-red, orange-reddish, saffron, or of various intermediate shades. Occasionally in the same species their colour is very variable (e.g. Lecanora metaboloides, Lecidea decolorans), while sometimes they are white or glaucous, rarely greenish, pruinose. Lecideine apothecia, which are not black, but otherwise variously coloured, are termed biatorine. The two principal parts of which an apothecium consists are the hypothecium and the hymenium, or thecium. The hypothecium is the basal part of the apothecium on which the hymenium is borne; the latter consists of asci (thecae) with ascospores, and paraphyses. The paraphyses; (which may be absent entirely in the Pyrenolichens) are erect, colourless filaments which are After Tulasne, from De Bary's Vergleichende Morphologie and Biologie der Pilze, Mycetozoen and Bacterien, by permission of Wilhelm Engelmann.

FIG. 13. - A, B, Gyrophora cylindrica. (A mag. 90, B 390 times, C highly magnified.) A, A vertical median sec- w, Its wall from which tion through a sperproceed sterigmata mogonium imbedded with rod-like sper in the thallus. matia (s).

o, Upper rind. m, Medullary hyphae of u, Under rind. [thallus. the thallus.

m, Medullary layer of the C, Cladonia novae Angliae, B, Portion of a very thin Delise; sterigmata section from the base with spermatia from of the spermogonium. the spermogonium. usually dilated and coloured at the apex; the apices are usually cemented together into a definite layer, the epithecium (fig. 14). The spores themselves may be unicellular without a septum or multicellular with one or more septa. Sometimes the two cavities are restricted to the two ends of the spore, the polaribilocular type and the two loculi may be united From Beitrcge zur Wissenschaftlichen Botanik. FIG. 9. - Section of Heteromerous Lichen Thallus.

a, Upper cortical layer. d, Lower cortical layer.

c, Medullary layer.

b, Gonidial layer.

After Bornet, from De Bary's Vergleichende Bacterien, by permission of Wilhelm Engelmann.

FIG. i i. - Lichen-forming Algae. (A, C, D, E mag. 950, B 650 times.) The alg"; s in all cases indicated by the letter g, the assailing hyphae by h. A, Pleurococcus, Ag. (Cystococcus, Nag.) C, Nostoc from the thallus of Physma attacked by the germ-tube from a chalazanum. spore of Physica parietina. D, Gloeocapsa from the thallus of B, Scytonema from the thallus of Synalissa Symphorea. Stereocaulon ramulosum. E, Pleurococcus Sp. (Cystococcus) from the thallus of Cladonia furcata. in the formation of this organ or that of the apothecium it has the general structure characteristic of that division of fungi. The apothecia, though of the normal fungal type and usually disk-shaped, are somewhat more variable, and since the Morphologie and Biologie der Pilze, Mycetozoen und 1 The thalline margin (margo thallinus) is the projecting edge of a special layer of thallus, the amphithecium, round the actual apothecium; the proper margin (margo proprius) is the projecting edge of the apothecium itself.

by a narrow channel (fig. 15). At other times the spores are divided by both transverse and longitudinal septa producing the muriform (murali-divided) spore so called from the resemblance of the individual chambers to the stones in a wall. The very large single spores of Pertusaria have been shown to contain numerous nuclei and when they germinate develop a large number of germ tubes.

Development of the Ascocarps

As the remarks on the nature of the spermatia show, the question of the sexuality of the lichens has been hotly disputed in common with that of the rest of the Ascomycetes. As indicated above, the weight of evidence seems to favour what has been put forward in the case of the non-lichenforming fungi (see Fungi), that in some cases the ascogonia develop as a result of a previous fertilization by spermatia, in other cases the ascogonia develop without such a union, while in still other Epithecium Asci  :Thecium (Hymenium) Paraphyses Parathecium, - Cortex gn,go mphitheciam - __ __MeddllzC-- _Hypoitienum==?=_` After Darbishire, from Berichte der deutschen botanischen Gesellschaft, by permission of Borntraeger & Co.

FIG. 14. - Diagram showing Apothecium in Section and surrounding Portion of Thallus, and special terms used to designate these parts.

cases the reduction goes still farther and the ascogenous hyphae instead of developing from the ascogonia are derived directly from the vegetative hyphae.

The first exact knowledge as to the origin of the ascocarp was the work of Stahl on Collema in 1877. He showed that the archicarp consisted of two parts, a lower coiled portion, the ascogonium, and an upper portion, the trichogyne, which projected from the thallus. Only when a spermatium was found attached to the trichogyne did the further development of the ascogonium take place. From these observations he drew the natural conclusion that the spermatium was a male, sexual cell. This view was hotly contested by many workers and it was sought to explain the trichogyne - without much success - as a respiratory organ, or as a boring organ which made a way for the developing apothecium. It was not till 1898, however, that Stahl's work received confirmatio and addition at the hands }$?_ of Baur (fig. 16). The latter :F' s showed that in Collema crispum there are two kinds <71f? T L of thalli, one with numerous apothecia, the other quite sterile or bearing only a few. The sterile thalli possessed no spermogonia, but were found to show sometimes as many as 1000 archicarps with trichogynes; yet none or very few came to maturity to bear bear ieitherllspermogonia or to be in immediate con nexion with spermogonia bearing thalli. Furthermore Baur showed that after the fusion of the spermatium with the trichogyne the transverse walls of that organ became perforated.

There was thus very strong circumstantial evidence in favour of fertilization, although the male nucleus was not traced. The further work of Baur, and that of Darbishire, Funfstuck and Lindau, have shown that in a number of other cases trichogynes are present. Thus ascogonia with trichogynes have been observed in Endocarpon, Collema, Pertusaria, Lecanora, Gyrophora, Parmelia, Ramalina, Physcia, Anaptychia and Cladonia. In Nephroma, Peltigera, Peltidea and Solorana a cogonia without trichogynes have been observed. In Collema and a form like Xanthoria parietina it is probable that actual fertilization takes place, and possibly also in some of the other forms. It is probable, however, that in the majority of cases the ascogonia develop without normal fertilization, as is necessarily the case where the ascogonia have no trichogynes or the spermatia are absent. In these cases we should expect to find some reduced process of fertilization similar to that of Humaria granulata among the ordinary Ascomycetes, where in the absence of the antheridia the female nuclei fuse in pairs. In other lichens we should expect to find the ascogenous hyphae arising directly from the vegetative hyphae as in Humaria rutilans among the ordinary fungi, where the process is associated with the fusion of vegetative nuclei. It is possible that Solorina saccata belongs to this class. Cytological details of nuclear behaviour among the lichens are, however, difficult to obtain owing 'to the slow growth of these forms and the often refractory t' nature of the material in the matter of preparation for microscopical ex amination.

Ejection of Spores

The spores are ejected from the apothecia and peri thecia as in the fungi by forcible ejacu lation from the asci. In the majority of forms it is clear that the soredia rather than the ascospore must play.

the more important part in lichen distribution as the development of the r1012444 ordinary spores is dependent on their ?,O.° finding the proper alga on the sub- I t.? stratum on which they happen to fall. - In a number of forms (Endocarpon pusillum, Stigmaatonima cataleptum, various species of Staurothele), however, there is a special arrangement by which the spores are, on ejection, associated with gonidia. In these forms gonidia are found in connexion with the young fruit; such algal cells undergo numerous divisions becoming very small in size and penetrating into the hymenium among the asci and paraphyses. When the spores are thrown out some of these hymenial gonidia, as they are called, are carried with them. When the spores germinate the germ-tubes surround the algal cells, which now increase in size and become the normal gonidia of the thallus.

Basidiolichens. As is clear from the above, nearly all the lichens are produced by the association of an ascomycetous fungus with algae. For some obscure reason the Basidiomycetes do not readily form lichens, so that only a few forms are known in which the fungal element is a member of this family. The two bestknown genera are Cora and Dictyonema; Corella, whose hymenium is unknown, is also placed here by Wainio. The so-called Gasterolichens, Trichocoma and Emeri- cella, have been shown to be merely ascomycetous fungi. Clavaria mucida, however, h a s apparently some claims to be considered as a Basidiolichen, since the base of the fruit body and the thallus from which it arises, according to Coker, always shows a mixture of hyphae and algae.

The best-known species is Cora pavonia, which is found in tropical regions growing on the bare earth and on trees; the gonidia belong to the genus Chroococcus while the fungus belongs, apparently, to the Thelephoreae (see Fungi). This lichen seems unique in the fact that the fungal element is also found growing and fruiting entirely devoid of algae, while in the TRaiiine margin  ? ??.: ??0411 - ?

=G illIi ' 'Proper margin ~?.

FIG. 15. - Vertical Section of Apothecium of Xanthoria parietina. a, Paraphyses.

b, Asci (thecae) with bilocular spores.

c, Hypothecium.

A After E..Baur, from Strasburger's Lehrbuch der Botanik, by permission of Gustav Fischer.

FIG. Collema crispum. A, Carpogonium, c, with its trichogyne t (X405).

B, Apex of the trichogyne with the spermatium, s, (X405) attached (>0125).

A From Strasburger's Lehrbuch permission of Gustav Fischer.

FIG. 17. - Cora pavonia. A, Viewed from above; B, From below; hym, hymenium. (Nat. size.) der Botanik, by ascolichens the fungus portion seems to have become so specialized to its symbiotic mode of life that it is never found growing independently.

The genus Dictyonema has gonidia belonging to the bluegreen alga, Scytonema. When the fungus predominates in the thallus it has a bracket-like mode of growth and is found projecting from the branches of trees with the hymenium on the under side. When the alga is predominant it forms felted patches on the bark of trees, the Laudatea form. It is said that the fungus of Cora pavonia and of Dictyonema is identical, the difference being in the nature of the alga.

Mode of Life. Lichens are found growing in various situations such as bare earth, the bark of trees, dead wood, the surface of stones and rocks, where they have little competition to fear from ordinary plants. As is well known, the lichens are often found in the most exposed and arid situations; in the extreme polar regions these plants are practically the only vegetable forms of life. They owe their capacity to live under the most inhospitable conditions to the dual nature of the organism, and to their capacity to withstand extremes of heat, cold and drought without destruction. On a bare rocky surface a fungus would die from want of organic substance and an alga from drought and want of mineral substances. The lichen, however, is able to grow as the alga supplies organic food material and the fungus has developed a battery of acids (see below) which enable it actually to dissolve the most resistant rocks. It is owing to the power of disintegrating by both mechanical and chemical means the rocks on which they are growing that lichens play such an important part in soil-production. The resistance of lichens is extraordinary; they may be cooled to very low temperatures and heated to high temperatures without being killed. They may be dried so thoroughly that they can easily be reduced to powder yet their vitality is not destroyed but only suspended; on being supplied with water they absorb it rapidly by their general surface and renew their activity. The life of many lichens thus consists of alternating periods of activity when moisture is plentiful, and completely suspended animation under conditions of dryness. Though so little sensitive to drought and extremes of temperature lichens appear to be very easily affected by the presence in the air of noxious substances such as are found in large cities or manufacturing towns. In such districts lichen vegetation is entirely or almost entirely absent. The growth of lichens is extremely slow and many of them take years before they arrive at a spore-bearing stage. Xanthoria parietina has been known to grow for fortyfive years before bearing apothecia. This slowness of growth is associated with great length of life and it is probable that individuals found growing on hard mountain rocks or on the trunks of aged trees are many hundreds of years old. It is possible that specimens of such long-lived species as Lecidea geographica actually outrival in longevity the oldest trees.

Relation of Fungus and Alga. The relation of the two constituents of the lichen have been briefly stated in the beginning of this article. The relation of the fungus to the alga, though it may be described in general terms as one of symbiosis, partakes also somewhat of the nature of parasitism. The algal cells are usually controlled in their growth by the hyphae and are prevented from forming zoospores, and in some cases, as already described, the algal cells are killed sooner or later by the fungus. The fungus seems, on the other hand, to stimulate the algal cells to special development, for those in the lichen are larger than those in the free state, but this is not necessarily adverse to the idea of parasitism, for it is well known that an increase in the size of the cells of the host is often the result of the attacks of parasitic fungi. It must be borne in mind that the exact nutritive relations of the two constituents of the lichen have not been completely elucidated, and that it is very difficult to draw the line between symbiosis and parasitism. The lichen algae are not alone in their specializa tion to the symbiotic (or parasitic) mode of life, for, as stated earlier, the fungus appear in the majority of cases to have completely lost the power of independent development since with very rare exceptions they are not found alone. They also differ very markedly from free living fungi in their chemical reactions.

Chemistry of Lichens. The chemistry of lichens is very complex, not yet fully investigated and can only be very briefly dealt with here. The wall of the hyphae of the fungus give in the young state the ordinary reactions of cellulose but older material shows somewhat different reactions, similar to those of the so-called fungus-cellulose. In many lichenfungi the wall shows various chemical modifications. In numerous lichens, e.g. Cetraria islandica, the wall contains Lichenin (C6H1005), a gummy substance which swells in cold water and dissolves in hot. Besides this substance, a very similar one, Isolichenin, is also found which is distinguished from lichenin by the fact that it dissolves in cold water and turns blue under the reaction of Iodine. Calcium oxalate is a very common substance, especially in crustaceous lichens; fatty oil in the form of drops or as an infiltration in the membrane is also common; it sometimes occurs in special cells and in extreme cases may represent 90% of the dry substance as in Verrucaria calciseda, Biatora immersa. Colouring Matters. - Many lichens, as is well known, exhibit a vivid colouring which is usually due to the incrustation of the hyphae with crystalline excretory products. These excretory products have usually an acid nature and hence are generally known as lichen-acids. A large number of these acids, which are mostly benzene derivatives, have been isolated and more or less closely investigated. They are characterized by their insolubility or very slight solubility in water; as examples may be mentioned erythrinic acid in Roccella and Lecanora; evernic acid in species of Evernia, Ramalina and Cladonia; lecanoric acid in Lecanora, Gyrophora. The so-called chrysophanic acid found in Xanthoria (Physcia) parietina is not an acid but a quinone and is better termed physcion.

Colour Reactions of Lichens

The classification of lichens is unique in the fact that chemical colour reactions are used by many lichenologists in the discrimination of species, and these reactions are included in the specific diagnoses. The substances used as tests in these reactions are caustic potash and calcium hypochlorite; the former being the substance dissolved in an equal weight of water and the latter a saturated extract of bleaching powder in water. These substances are represented by lichenologists by the signs K and CaC1 respectively, and the presence or absence of the colour reactions are represented thus, K+, CaCI+, or K -, CaC1 -. If the cortical layer should exhibit positive reaction and the medulla of the same species a negative reaction with both reagents, the result is represented thus, K CaCI i. If a reaction is only produced after the consecutive addition of the two reagents, this is symbolized by K(CaC1) +. A solution of iodine is also used as a test owing to the blue or wine-red colour which the thallus, hymenium or spores may give with this reagent. The objection to the case of these colour reactions is due to the indefinite nature of the reaction and the doubt as to the constant presence of a definite chemical compound in a given species. A yellow colour with caustic potash solution is produced not only by atranoric acid but also by evernic acid, thamnolic acid, &c. Again in the case of Xanthoria parietina vulpinic acid is only to be found in young thalli growing on sandstone; in older forms or in those growing on another substratum it is not to be detected. A similar relation between oil formation and the nature of the substratum has been observed in many lichens. Considerations such as these should make one very wary in placing reliance on these colour reactions for the purposes of classification.

Economic Uses of Lichens. In the arts, as food and as medicine, many lichens have been highly esteemed, though others are not now employed for the same purposes as formerly.

i. Lichens Used in the Arts. - Of these the most important are such as yield, by maceration in ammonia, the dyes known in commerce as archil, cudbear and litmus. These, however, may with propriety be regarded as but different names for the same pigmentary substance, the variations in the character of which are attributable to the different modes in which the pigments are manufactured. Archil proper is derived from several species of Roccella (e.g. R. Montaguei, R. tinctoria), which yield a rich purple dye; it once fetched a high price in the market. Of considerable value is the " perelle " prepared from Lecanora parella, and used in the preparation of a red or crimson dye. Inferior to this is " cudbear," derived from Lecanora tartarea, which was formerly very extensively employed by the peasantry of north Europe for giving a scarlet or purple colour to woollen cloths. By adding certain alkalies to the other ingredients used in the preparation of these pigments, the colour becomes indigoblue, in which case it is the litmus of the Dutch manufacturers. Amongst other lichens affording red, purple or brown dyes may be mentioned Ramalina scopulorum, Parmelia, saxatilis and P. omphalodes, Umbilicaria pustulate and several species of Gyrophora, Urceolaria scruposa, all of which are more or less employed as domestic dyes. Yellow dyes, again, are derived from Chlorea vulpina, Platysma juniperinum, Parmelia caperata and P. conspersa, Physcia jlavicans, Ph. parietina and Ph. lychnea, though like the preceding they do not form articles of commerce, being merely used locally by the natives of the regions in which they occur most plentifully. In addition to these, many exotic lichens, belonging especially to Parmelia and Sticta (e.g. Parmelia tinctorum, Sticta argyracea), are rich in colouring matter, and, if obtained in sufficient quantity, would yield a dye in every way equal to archil. These pigments primarily depend upon special acids contained in the thalli of lichens, and their presence may readily be detected by means of the reagents already noticed. In the process of manufacture, however, they undergo various changes, of which the chemistry is still but little understood. At one time also some species were used in the arts for supplying a gum as a substitute for gum-arabic. These were chiefly Ramalina fraxinea, Evernia prunastri and Parmelia physodes, all of which contain a considerable proportion of gummy matter (of a much inferior quality, however, to gum-arabic), and were employed in the process of calico-printing and in the making of parchment and cardboard. In the i 7th century some filamentose and fruticulose lichensviz. species of Usnea and Ramalina, also Evernia furfuracea and Cladonia rangiferina, were used in the art of perfumery. From their supposed aptitude to imbibe and retain odours, their powder was the basis of various perfumes, such as the celebrated " Poudre de Cypre " of the hairdressers, but their employment in this respect has long since been abandoned.

2. Nutritive Lichens

Of still greater importance is the capacity of many species for supplying food for man and beast. This results from their containing starchy substances, and in some cases a small quantity of saccharine matter of the nature of mannite. One of the most useful nutritious species is Cetraria islandica, " Iceland moss," which, after being deprived of its bitterness by boiling in water, is reduced to a powder and made into cakes, or is boiled and eaten with milk by the poor Icelander, whose sole food it often constitutes. Similarly Cladonia rangiferina and Cl. sylvatica, the familiar " reindeer moss," are frequently eaten by man in times of scarcity, after being powdered and mixed with flour. Their chief importance, however, is that in Lapland and other northern countries they supply the winter food of the reindeer and other animals, who scrape away the snow and eagerly feed upon them. Another nutritious lichen is the " Tripe de Roche " of the arctic regions, consisting of several species of the Gyrophorei, which when boiled is often eaten by the Canadian hunters and Red Indians when pressed by hunger. But the most singular esculent lichen of all is the " manna lichen," which in times of drought and famine has served as food for large numbers of men and cattle in the arid steppes of various countries stretching from Algiers to Tartary. This is derived chiefly from Lecanora esculents, which grows unattached on the ground in layers from 3 to 6 in. thick over large tracts of country in the form of small irregular lumps of a greyish or white colour. In connexion with their use as food we may observe that of recent years in Scandinavia and Russia an alcoholic spirit has been distilled from Cladonia rangiferina and extensively consumed, especially in seasons when potatoes were scarce and dear. Formerly also Sticta pulmonaria was much employed in brewing instead of hops, and it is said that a Siberian monastery was much celebrated for its beer which was flavoured with the bitter principle of this species.

3. Medicinal Lichens

During the middle ages, and even in some quarters to a much later period, lichens were extensively used in medicine in various European countries. Many species had a great repute as demulcents, febrifuges, astringents, tonics, purgatives and anthelmintics. The chief of those employed for one or other, and in some cases for several, of these purposes were Cladonia pyxidata, Usnea barbata, Ramalina farinacea, Evernia prunastri, Cetraria islandica, Sticta pulmonaria, Parmelia saxatilis, Xanthoria parietina and Pertusaria amara. Others again were believed to be endowed with specific virtues, e.g. Peltigera canin g , which formed the basis of the celebrated " pulvis antilyssus " of Dr Mead, long regarded as a sovereign cure for hydrophobia; Platysma juniperinum, lauded as a specific in jaundice, no doubt on the similia similibus principle from a resemblance between its yellow colour and that of the jaundiced skin; Peltidea aphthosa, which on the same principle was regarded by the Swedes, when boiled in milk, as an effectual remedy for the aphthae or rash on their children. Almost all of these virtues, general or specific, were imaginary; and at the present day, except perhaps in some remoter districts of northern Europe, only one of them is employed as a remedial agent. This is the " Iceland moss " of the druggists' shops, which is undoubtedly an excellent demulcent in various dyspeptic and chest complaints. No lichen is known to be possessed of any poisonous properties to man, although Chlorea vulpina is believed by the Swedes to be so. Zukal has considered that the lichen acids protect the lichen from the attacks of animals; the experiments of Zopf, however, have cast doubt on this; certainly lichens containing very bitter acids are eaten by mites though some of the acids appear to be poisonous to frogs.

Classification. The dual nature of the lichen thallus introduces at the outset a classificatory difficulty. Theoretically the lichens may be classified on the basis of their algal constituent, on the basis of their fungal constituent, or they may be classified as if they were homogeneous organisms. The first of these systems is impracticable owing to the absence of algal reproductive organs and the similarity of the algal cells (gonidia) in a large number of different forms. The second system is the most obvious one, since the fungus is the dominant partner and produces reproductive organs. The third system was that of Nylander and his followers, who did not accept the Schwenderian doctrine of duality. In actual practice the difference between the second and third methods is not very great since the fungus is the producer of the reproductive organs and generally the main constituent. Most systems agree in deriving the major divisions from the characters of the reproductive organs (perithecia, apothecia, or basidiospore bearing fructification), while the characters of the algal cells and those of the thallus generally are used for the minor divisions. The difference between the various systems lies in the relative importance given to the reproductive characters on the one hand and the vegetative characters on the other. In the system (1854-1855) of Nylander the greater weight is given to the latter, while in more modern systems the former characters receive the more attention.

A brief outline of a system of classification, mainly that of Zahlbruckner as given in Engler and Prantl's Pjlanzenfamilien, is outlined below.

There are two main divisions of lichens, Ascolichenes and Basidiolichenes, according to the nature of the fungal element, whether an ascomycete or basidiomycete. The Ascolichenes are again divided into Pyrenocarpeae or Pyrenolichenes and Gymnocarpeae or Discolichenes; the first having an ascocarp of the nature of a perithecium, the second bearing their ascospores in an open apothecium.

Pyrenolichenes Series I. Perithecium simple not divided.

a. With Pleurococcus or Palmella gonidia.

Moriolaceae, Verrucariaceae, Pyrenothamnaceae.

b. With Chroolepus gonidia.

Pyrenulaceae, Paratheliaceae.

c. With Phyllactidium or Cephaleurus gonidia. Strigulaceae.

d. With Nostoc or Scytonema gonidia.

Pyrenidiaceae.

Series II. Perithecia divided or imperfectly divided by cross-walls. Mycoporaceae with Palmella_or Chroolepus gonidia.

Series I. Coniocarpineae. The paraphyses branch and form a network (capillitium) over the asci, the capillitium and ejected spores forming a long persistent powdery mass (mazaedium).

Caliciaceae, Cypheliaceae, Sphaerophoraceae.

Series II. Graphidineae. Apothecia seldom round, usually elongatedellipsoidal, no capillitium.

Arthoniaceae, Graphidiaceae, Roccellaceae.

Series III. Cyclocarpineae, Apothecium usually circular, no capillitium.

A. Spores usually two-celled, either with a strongly thickened cross-wall often perforated by a narrow canal or with crosswall only slightly thickened. In the first case the spores are usually colourless, the second case always brown.

Buelliaceae, Physciaceae.

B. Spores unicellular, parallel-multicellular or muriform, usually colourless, cross-walls usually thin.

a Thallus in moist state more or less gelatinous.

Gonidia always belonging to the Cyanophyceae, Lichinaceae, Ephebaceae, Collemaceae, Pyrenopsidaceae.

(3 Thallus not gelatinous.

Coenogoniaceae, Lecideaceae, Cladoniaceae, Lecanoraceae, Pertusariaceae, Peltigeraceae, Stictaceae, Pannariaceae, Gyrophoraceae, Parmeliaceae, Cladoniaceae, Usneaceae.

Basidiolichenes (Hymenolichenes) Cora, Dictyonema (incl. Laudatea), Corella (doubtfully placed here as the hymenium is unknown).

Habitats and Distribution of Lichens. 1. Habitats. - These are extremely varied, and comprise a great number of very different substrata. Chiefly, however, they are the bark of trees, rocks, the ground, mosses and, rarely, perennial leaves. (a) With respect to corticolous lichens, some prefer the rugged bark of old trees (e.g. Ramalina, Parmelia, Stictei) and others the smooth bark of young trees and shrubs (e.g. Graphidei and some Lecideae). Many are found principally in large forests (e.g. Usnea, Alectoria jubata); while a few occur more especially on trees by roadsides (e.g. Physcia parietina and Ph. pulverulenta). In connexion with corticolous lichens may be mentioned those lignicole species which grow on decayed, or decaying wood of trees and on old pales (e.g. Caliciei, various Lecideae, Xylographa), (b) As to saxicolous lichens, which occur on rocks and stones, they may be divided into two sections, viz. calcicolous and calcifugous. To the former belong such as are found on calcareous and cretaceous rocks, and the mortar of walls (e.g. Lecanora calcarea, Lecidea calcivora and several Verrucariae), while all other saxicolous lichens may be regarded as belonging to the latter, whatever may be the mineralogical character of the substratum. It is here worthy of notice that the apothecia of several calcicolous lichens (e.g. Lecanora Prevostii, Lecidea calcivora) have the power of forming minute cavities in the rock, in which they are partially buried. (c) With respect to terrestrial species, some prefer peaty soil (e.g. Cladonia, Lecidea decolorans), others calcareous soil (e.g. Lecanora crassa, Lecidea decipiens), others sandy soil or hardened mud (e.g. Collema limosum, Peltidea venosa); while many may be found growing on all kinds of soil, from the sands of the sea-shore to the granitic detritus of lofty mountains, with the exception of course of cultivated ground, there being no agrarian lichens. (d) Muscicolous lichens again are such as are most frequently met with on decayed mosses and Jungermannia, whether on the ground, trees or rocks (e.g. Leptogium muscicola, Gomphillus calicioides). (e) The epiphyllous species are very peculiar as occurring upon perennial leaves of certain trees and shrubs, whose vitality is not at all affected by their presence as it is by that of fungi. In so far, however, as is known, they are very limited in number (e.g. Lecidea, Bouteillei, Strigula). Sometimes various lichens occur abnormally in such unexpected habitats as dried dung of sheep, bleached bones of reindeer and whales, old leather, iron and glass, in districts where the species are abundant. It is apparent that in many cases lichens are quite indifferent to the substrata on which they occur, whence we infer that the preference of several for certain substrata depends upon the temperature of the locality or that of the special habitat. Thus in the case of saxicolous lichens the mineralogical character of the rock has of itself little or no influence upon lichen growth, which is influenced more especially and directly by their physical properties, such as their capacity for retaining heat and moisture. As a rule lichens grow commonly in open exposed habitats, though some are found only or chiefly in shady situations; while, as already observed, scarcely any occur where the atmosphere is impregnated with smoke. Many species also prefer growing in moist places by streams, lakes and the sea, though very few are normally and probably none entirely, aquatic, being always at certain seasons exposed for a longer or shorter period to the atmosphere (e.g. Lichina, Leptogium rivulare, Endocarpon fiuviatile, Verrucaria maura). Some species are entirely parasitical on other lichens (e.g. various Lecideae and Pyrenocarpei), and may be peculiar to one (e.g. Lecidea vitellinaria) or common to several species (e.g. Habrothallus parmeliarum). A few, generally known as erratic species, have been met with growing unattached to any substratum (e.g. Parmelia revoluta, var. concentrica, Lecanora esculenta); but it can hardly be that these are really free ab initio (vide Crombie in Journ. Bot., 1872, p. 306). It is to the different characters of the stations they occupy with respect to exposure, moisture, &c., that the variability observed in many types of lichens is to be attributed. 2. Distribution. - From what has now been said it will readily be inferred that the distribution of lichens over the surface of the globe is regulated, not only by the presence of suitable substrata, but more especially by climatic conditions. At the same time it may safely be affirmed that their geographical range is more extended than that of any other class of plants, occurring as they do in the coldest and warmest regions - on the dreary shores of arctic and antarctic seas and in the torrid valleys of tropical climes, as well as on the greatest mountain elevations yet attained by man, on projecting rocks even far above the snow line (e.g. Lecidea geographica). In arctic regions lichens form by far the largest portion of the vegetation, occurring everywhere on the ground and on rocks, and fruiting freely; while terrestrial species of Cladonia and Stereocaulon are seen in the greatest luxuriance and abundance spreading over extensive tracts almost to the entire exclusion of other vegetation. The lichen flora of temperate regions again is essentially distinguished from the preceding by the frequency of corticolous species belonging to Lecanora, Lecidea and Graphidei. In intertropical regions lichens attain their maximum development (and beauty) in the foliaceous Stictei and Parmeliei, while they are especially characterized by epiphyllous species, as Strigula, and by many peculiar corticole Thelotremei, Graphidei and Pyrenocarpei. Some lichens, especially saxicolous ones, seem to be cosmopolitan (e.g. Lecanora subfusca, Cladonia pyxidata); and others, not strictly cosmopolitan, have been observed in regions widely apart. A considerable number of species, European and exotic, seem to be endemic, but further research will no doubt show that most of them occur in other climatic regions similar to those in which they have hitherto alone been detected. To give any detailed account, however, of the distribution of the different genera (not to speak of that of individual species) of lichens would necessarily far exceed available limits.

Bibliography

General: Engler and Prantl, Die natiirlichen Pflanzenfamilien, Teil I, Abt. 1 * where full literature will be found up to 1898. M. Funfstuck, " Der gegenwartige Stand der Flechtenkunde," Refer. Generalvers. d. deut. bot. Ges. (1902). Dual Nature: J. Baranetzky, " Beitrge zur Kenntnis des selbststandigen Lebens der Flechtengonidien," Prings. Jahrb. f. wiss. Bot. vii. (1869); E. Bornet, " Recherches sur les gonidies des lichens," Ann. de sci. nat. bot., 5 ser. n. 17 (1873); G. Bonnier, " Recherches sur la synthbse des lichens," Ann. de sci. nat. bot., 7 ser. n. 9 (1889); A. Famintzin and J. Baranetzky, "Zur Entwickelungsgeschichte der Gonidien u. Zoosporenbildung der Lichenen," Bot. Zeit. (1867, p. 189, 1868, p. 169); S. Schwendener, Die Algentypen der Flechtengonidien (Basel, 1869); A. Moller, Ober die Kultur flechtenbildender Ascomyceten ohne Algen. (Munster, 1887). Sexuality: E. Stahl,. Beitrage zur Entwickelungsgeschichte der Flechten (Leipzig, 1877); G. Lindau, Ober Anlage and Entwickelung einiger Flechtenapothecien (Flora, 1888); E. Baur, " Zur Frage nach der Sexualitat der Collemaceae," Ber. d. deut. bot. Ges. (1898); " Ober Anlage und.

xvi. 19 a Entwickelung einiger Flechtenapothecien " (Flora, Bd. 88, 1901); " Untersuchungen fiber die Entwickelungsgeschichte der Flechtenapothecien," Bot. Zeit. (1904); 0. V. Darbishire," Ober die Apothecium-entwickelung der Flechte, Physcia pulverulenta," Nyl. Prings. Jahrb. (Bd.034, 1900). Chemistry. - W. Zopf, " Vergleichende Produkte," Beitr. z. bot. Centralbl. (Bd. 14, 1903); Die Flechtenstoffe (Jena, 1907). (J. M. C.; V. H. B.)