  Since the time when lichens were first recognized as a separate class—as members of the genus Lichen by Tournefort a. Dillenius and Linnaeus. The first systematists were chiefly impressed by their likeness to mosses, hepatics or algae. Dillenius b. Acharius. Early in the nineteenth century Acharius, the Swedish Lichenologist, worked a revolution in the classification of lichens. He gave first place to the form of the thallus, but he also noted the fundamental differences in fruit-formation: his new system appeared in the Methodus Lichenum Diagnoses of twenty-three genera are given with their included species. The work was further extended and emended in Lichenographia Universalis
The orders are thus based on the form of the fruit; the genera in the Synopsis number 41. Large genera such as Lecanora with 132 species are divided into sections, many of which have in turn been established as genera, by S. F. Gray in 1821, and later by other systematists. The Synopsis was the text-book adopted by succeeding botanists for some 40 years with slight alterations in the arrangement of classes, genera, etc. Wallroth c. Schaerer. This valiant lichenologist worked continuously during the first half of the nineteenth century, but with very partial use of the microscope. His last publication in 1850, an Enumeration of Swiss Lichens, was the final declaration of the older school that relied on field characters. His classification is as follows:
An “Appendix” contains descriptions of Crustacei and Fruticulosi, all sterile forms, except Coniocarpon and Arthonia, which seem out of place, and finally a “Corollarium” of gelatinous lichens all classified under one genus Collema. d. Massalongo and Koerber. As a result of their microscopic studies, these two workers proposed many changes based on fruit and spore characters, and Koerber in the Systema Lichenum Germaniae (1855) gave expression to these views in his classification. He also made use of Wallroth’s distinctions of “homoiomerous” and “heteromerous,” thus dividing lichens at the outset into those mostly with blue-green and those with bright-green gonidia. The following is the main outline of Koerber’s classification:
With the exception of Order V all are subdivided into two sections, “gymnocarpi” with open fruits and “angiocarpi” with closed fruits, a distinction that had long been recognized both in lichens and in fungi. e. Nylander. The above writers had been concerned with the interrelationships of lichens; Nylander, who was now coming forward as a lichenologist of note, gave a new turn to the study by dwelling on their relation to other classes of plants. Without for a moment conceding that they were either algal or fungal, he yet insisted on their remarkable affinity to algae on the one hand, and to fungi on the other, and he sought to make evident this double connection by his very ingenious scheme of classification In his scheme, which is the one followed by Leighton and Crombie, the “family” represents the highest division; series, tribe, genus and species come next in order. We have thus:
This last family, which includes the great bulk of lichens, is divided into the following series: I. Epiconiodei; II. Cladoniodei; III. Ramalodei; IV. Phyllodei; V. Placodei; VI. Pyrenodei. It is an ascending series up to the Phyllodei, or foliaceous lichens, which he considers higher in development than the fruticose or filamentous Ramalodei. The Placodei include four tribes on a descending scale, the Lecanorei, Lecidinei, Xylographidei and Graphidei. The classification is almost wholly based on thalline form, except for the Pyrenodei in which are represented genera with closed fruits, there being one tribe only, the Pyrenocarpei. Nylander claims however to have had regard equally to the reproductive system and was the first to give importance to the spermogonia. The classification is coherent and easy to follow, though, like all classifications f. MÜller(-Argau). In preparing his lists of Genevan lichens (1862), MÜller realized that Nylander’s series was unnatural, and he found as he studied more deeply that lichens must be ranged in parallel or convergent but detached groups. He recognized three main groups:
He suggested that, in relation to other plants, Eulichens approach Pezizae, Hysteriaceae and Sphaeriaceae; Epiconiaceae have affinity with Lycoperdaceae, while Collemaceae are allied to the algal family Nostocaceae. These three groups of Eulichens, he held, advanced on somewhat parallel lines, but reached a very varied development, the Discocarpeae attaining the highest stage of thalline form. MÜller accepted as characters of generic importance the form and structure of the fruiting body, the presence or absence of paraphyses, and the septation, colour, etc. of the spores. A few years later (1867) the composite nature of the lichen thallus was announced by Schwendener, and, after some time, was acknowledged by most botanists to be in accordance with the facts of nature. Any system of classification, therefore, that claims to be a natural one, must, while following as far as possible the line of plant development, take into account the double origin of lichens both from algae and fungi, the essential unity and coherence of the class being however proved by the recurring similarity between the thalline types of the different phyla. As MÜller had surmised: “they are a series of parallel detached though convergent groups.” g. Reinke. The arrangement of Ascolichens on these lines was first seriously studied by Reinke The Coniocarpi are a group apart, but as their fruit is at first entirely closed—at least in some of the genera—the more natural position for them is between Discocarpi and Pyrenocarpi. It is in the arrangement of the Discocarpi that variation occurs. Reinke’s arrangement of orders and families in that sub-class is as follows: Subclass 2. Discocarpi.
The orders represent generally the principal phyla or groups, the families subordinate parallel phyla within the orders. The first three orders are stages of advance as regards fruit development; the Cyanophili are a group apart. Wainio h. Zahlbruckner. The systematist of greatest weight in recent times is A. Zahlbruckner, who is responsible for the systematic account of lichens in Engler and Prantl’s NatÜrlichen Pflanzenfamilien. It is difficult to express the very great service he has rendered to Lichenology, in that and other world-wide studies of lichens. The sketch of lichen phylogeny as given in the present volume owes a great deal to the sound and clear guidance of his work, though his conclusions may not always have been accepted. The classification in the Pflanzenfamilien is the one now generally followed. The class Lichenes is divided by Zahlbruckner
These are again broken up into families, and in the arrangement and sequence of the families Zahlbruckner indicates his view of development and relationship. They occur in the following order: Series 1. PYRENOCARPEAE
Series 2. GYMNOCARPEAE
Subseries 1. CONIOCARPINEAEThis is a well-defined group, peculiar in the disappearance of the asci at an early stage so that the spores lie like a powder in the globose partly closed fruits. Algal cells, bright-green; Protococcaceae. There are only three families:
Subseries 2. GRAPHIDINEAEThis subseries comes next in the form of fruit development; generally the apothecia are elongate, with a narrow slit-like opening, so that a transverse section shows almost a perithecial outline. Algal cells are mostly Trentepohlia.
Subseries 3. CYCLOCARPINEAEA large and very varied group! In most of the families the algal cells are bright-green (Chlorophyceae), in some they are blue-green (Cyanophyceae), these latter corresponding to Reinke’s order Cyanophili. The apothecia, as the name implies, are round and open; the “Cyanophili” have been placed by Zahlbruckner after those families in which the The first family of the Cyclocarpineae, the Lecanactidaceae, is often placed under Graphidineae; in any case it forms a link between the two subseries. 1. Lecideine group (apothecia without a thalline margin).
2. Cyanophili group. In this group the classification depends almost entirely on the nature of the algal constituents. The apothecia are in most genera provided with a thalline margin. a. More or less gelatinous when moist.
b. Not gelatinous when moist.
3. Lecanorine group (apothecia with a thalline margin). The remaining families have all bright-green gonidia and nearly always apothecia with a thalline margin. The group includes several distinct phyla:
Subclass 2. Hymenolichens.There are only three closely related genera of Hymenolichens, Cora, Corella and Dictyonema with Chroococcus or Scytonema algae. There is reason to dissent from the arrangement in one or two instances which will be pointed out in the following examination of families and genera. B. Families and Genera of AscolichensThe necessity for a well-reasoned and well-arranged system of classification is self-evident: without a working knowledge of the plants that are the subject of study no progress can be made. The recognition of plants as isolated individuals is not sufficient, it must be possible to place them in relation to others; hence the importance of a natural system. In identifying species artificial aids, such as habitat and substratum, are also often of great value, and a good working system should take account of all characteristics. Lichen development is the result of two organisms mutually affecting each other, but as the fungus provides the reproductive system, it is the dominant partner: the main lines of classification are necessarily determined Zahlbruckner’s system gives due weight to the inheritance from both fungus and alga with, however, the fungus as the chief factor in development, and as his work is certain to be generally followed by modern lichenologists, it is the one of most immediate interest. His scheme has been accepted in the following more detailed account of families and genera, and for the benefit of home workers those that have not so far been recorded from the British Isles have been marked with an asterisk. It cannot be affirmed that nomenclature is as yet firmly established in lichenology. Both on historical grounds and on those of convenience, the subject is one of extreme importance, and interest in it is one of the main avenues by which we secure continuity with the past, and by which we are able to realize not only the difficulty, but the romance of pioneer work. Besides, there can be no exchange of opinion between students nor assured knowledge of plants, until the names given to them are beyond dispute. According to the ruling of the Brussels Botanical Congress in 1910, Linnaeus’s In the following necessarily brief account of families and genera, the “spermogonia” or “pycnidia” have in most cases been left out of account, as in many instances they vary within the family and occasionally even within the genus. Their taxonomic value is not without importance, but, in the general systematic arrangement, they are only subsidiary characters. An account of them has already been given, and for more detailed statements the student is referred to purely systematic works. There are two main types of spore production in the “pycnidia” which have been shortly described by Steiner Subclass I. ASCOLICHENS Series I. PYRENOCARPINEAEIt is on mycological grounds that Pyrenocarpineae are placed at the base of lichen classification. There is no evidence that the series was first in time. I. MoriolaceaeThis family was described by Norman The thallus is crustaceous. The algal cells, Protococcaceae, occur either in groups (sometimes stalked) surrounded by a plectenchymatous wall and called by Norman “goniocysts,” or they form nests in the thallus termed “nuclei” which are surrounded by a double wall of plectenchyma, colourless in the interior and brown outside. Norman invented the term “Allelositismus,” which may be rendered “mutualism,” to indicate this peculiar form of thallus. The species of Spheconisca are fairly numerous on poplars, willows and conifers:
II. EpigloeaceaeThe family consists of but one genus and one species, Epigloea bactrospora, and, according to Zahlbruckner, further examination is necessary to make certain as to the lichenoid nature of the plant. Zukal The small globose pycnidia contain simple sporophores and acrogenous straight or slightly bent rod-like spores.
III. VerrucariaceaeIn all the genera of this family the thallus is crustaceous, and, with very few exceptions, the species are saxicolous or terricolous. The thallus is variable within the crustaceous limits, and may be superficial and very conspicuous, almost imperceptible, or wholly immersed in the substratum. The algal cells are Protococcaceae, and in two of the genera the green cells penetrate the hymenium and grow in rows alongside of the asci. The perithecia are small roundish structures scattered over the thallus, the base immersed, but the upper portion generally projecting. An outer dark-coloured wall surrounds the whole perithecium (entire) or only the upper exposed portion (dimidiate); it opens above by a pore or ostiole more or less prominent. In some of the genera the paraphyses become dissolved at an early stage, and somewhat similar filaments near the ostiole, termed periphyses, aid in the expulsion of the spores. The spores vary in septation, colour and size, and these variations have served to delimit the genera which have been formed from the original very large genus Verrucaria. The ascus may be 1-, 2-, 4- or 8-spored. In only one genus is it many-spored (Trimmatothele). The genera are as follows:
IV. DermatocarpaceaeIn this family there is a much more advanced thalline development—generally squamulose or with some degree of foliose structure, though in the genus Endocarpon, some of the species are little more than crustaceous. The least developed in structure is Normandina: the thallus of the single species consists of delicate shell-like squamules which are non-corticate above and below. In the other genera there is a cortex of plectenchyma. The perithecia are almost wholly immersed, and open above by a straight ostiole. The fructification of Dacampia is considered by some lichenologists to be only a parasite on the white thickish squamulose thallus with which it is associated.
V. PyrenothamniaceaeThallus more or less fruticose and corticate on both surfaces. Algal cells Protococcaceae. Only two genera are included in this family: Nylanderiella with one species from New Zealand, with a small laciniate thallus up to 15 mm. in height, partly upright, partly decumbent, and attached to the substratum by basal rhizinae; the other small genus, Pyrenothamnia, belongs to N. America; the thallus has a short rounded stalk which expands above to an irregular frond. The perithecia are immersed in the fronds.
VI. MastoideaceaeA family containing one genus and one species, with a wide distribution, having been found in Siberia, on the Antarctic continent (Graham’s Land), as also in Tierra del Fuego, South Georgia, South Shetland Islands and Kerguelen. The thallus is foliose, of small thin lobes, and without rhizinae. Algal cells Prasiola
VII. PyrenulaceaeThis family of crustaceous lichens differs from Verrucariaceae chiefly in the gonidium which is a species of Trentepohlia. Genera and species are largely corticolous and the thallus is inconspicuous, often developing within the substratum. The perithecia, like those of Verrucariae, are immersed or partly emergent and have an entire or dimidiate outer wall. They are scattered over the thallus except in Anthracothecium where they are often coalescent. This genus is tropical or subtropical except for one species which inhabits S.W. Ireland. Paraphyses are variable, and in some species tend to disappear, but do not dissolve in mucilage. The spores are generally colourless, only in one monotypic genus, Coccotrema, are they simple. The cells into which the spore is divided differ in form according to the genus.
VIII. ParatheliaceaeThis family is peculiar in that the perithecia open by a somewhat elongate ostiole that slants at an oblique angle. The algal cells are Trentepohlia. Genera and species are endemic in tropical or subtropical regions of the Western hemisphere, though a species of Pleurotrema has been found in subantarctic America. They are corticolous and the thallus is either
IX. TrypetheliaceaeThis and the following two families are distinguished by the pseudostroma or compound fruit, a character rare among lichens, though the true stroma is frequent in Pyrenomycetes in such genera as Dothidea, Valsa, etc. The genera are crustaceous and corticolous and occur with few exceptions in tropical or subtropical regions, mostly in the Western Hemisphere. Several grow on officinal bark (Cinchona, etc.). Algal cells are Trentepohlia. As in many tropical lichens, the spores are large. The genera are based chiefly on spore characters, on septation, and on the form of the spore cells:
X. AstrotheliaceaeThe perithecia are either upright or inclined, and occur usually in radiate groups. They are free or united in a stroma, and the elongate ostioles open separately or coalesce in a common canal. The genera are all crustaceous, with Trentepohlia gonidia. They are tropical or subtropical, mostly in the Western Hemisphere; but species of Parmentaria and Astrothelium have been recorded also from Australia. The spores are all many-celled and the form of their cells is a generic character:
XI. MycoporaceaeA small family with only two genera which are found in both Hemispheres; species of both occur in Great Britain. They are all corticolous. The perithecia are united into a partially chambered fruiting body surrounded by a common wall, but opening by separate ostioles. The thallus is thinly crustaceous, with Palmella gonidia in Mycoporum, and Trentepohlia in Mycoporellum. The spores are colourless or brown in both genera:
XII. PhyllopyreniaceaeThallus foliose with both surfaces corticate and attached by rhizinae. Algal cells Trentepohlia. There is but one genus, Lepolichen, which has a laciniate somewhat upward growing thallus. Two species, both from South America, have been described, L. granulatus MÜll.-Arg. and L. coccophora Hue. The latter has been recently examined by Hue The perithecia are immersed in thalline tubercles:
XIII. StrigulaceaeA family of epiphyllous lichens inhabiting and disfiguring coriaceous evergreen leaves, or occasionally fern leaves in tropical or subtropical regions. The algae associated are Mycoidea and Phycopeltis (Phyllactidium). The only truly parasitic lichen, Strigula, belongs to this family: the alga precedes the lichen on the leaves and is gradually invaded by the hyphae of the lichen and altered in character. The small black perithecia are scattered over the surface. In Strigula the lichen retains the spreading rounded form of the alga. The other genera are more irregular.
XIV. PyrenidiaceaeThe only family of Pyrenocarpineae associated with blue-green algae. The genera of Pyrenidiaceae are all monotypic, only one is common and of wide distribution, Coriscium (Normandina Nyl.). Pyrenidium is the only member that has a fruticose thallus, and that is of minute dimensions. Eolichen Heppii, found and described by Zukal, is a doubtful lichen. “Lophothelium” Stirton is a case of parasitism of a fungus, Ticothecium, on the squamules of Stereocaulon condensatum.
Series II. GYMNOCARPEAESubseries 1. Coniocarpineae This small subseries is marked by the peculiar “mazaedium” type of fruit with its disappearing asci. It forms a connecting link between the families with perithecia and those with apothecia. The thallus is crustaceous or fruticose, often poorly developed and sometimes absent. The algal cells are Protococcaceae or rarely Trentepohlia. XV. CaliceaceaeThe thallus is thinly crustaceous, sometimes brightly coloured, sometimes absent, taking no part in the formation of the fruits; these have upright stalks with a small capitulum, and often look like minute nails. One genus, Sphinctrina, is parasitic on the thallus of other lichens, mostly Pertusariae.
XVI. CypheliaceaeThallus crustaceous. Algal cells Protococcaceae or Trentepohlia. Apothecia sessile, more widely open than in the previous family; in some genera the thallus forms an outer apothecial margin. The genera Farriola from Norway and Tylophorella from New Granada are monotypic. The British genus Cyphelium has been known as Trachylia.
XVII. SphaerophoraceaeThe most highly evolved family of the subseries, as regards the thallus. Algal cells Protococcaceae. In Tholurna, a small lichen endemic in Scandinavia, there is a double thallus: one of horizontal much-divided squamules, the other swollen, upright, terminating in the capitulum. The fruit is lateral in Calycidium, a squamulose form from New Zealand, and in Pleurocybe from Madagascar, with stiff strap-shaped fronds. All the genera are monotypic except Sphaerophorus, of which genus ten species are recorded, some of them with a world-wide distribution. The spores are brown and simple or 1-septate.
Subseries 2. Graphidineae In this subseries are included five families that differ rather widely from each other both in thallus and apothecia; the latter are more or less carbonaceous and mostly with a proper margin only. Families and genera are widely distributed, though most abundant in warm regions. Algal cells mostly Trentepohlia. A comprehensive study of the apothecia of this series by Bioret XVIII. ArthoniaceaeThe thallus of Arthoniaceae is corticolous with few exceptions and is very inconspicuous, being largely embedded in the substratum. The apothecia (ardellae) are round, irregular or stellate, without any margin, the hymenium being protected by the dense branching of the paraphyses at the tips. Arthonia is abundant everywhere. The species of the other genera belong mostly to tropical or subtropical countries. Arthoniopsis is similar to Arthonia in the character of the fruits, but the gonidium is a Phycopeltis, and it is only found on leaves. Synarthonia with peculiar stromatoid fructification is monotypic; it occurs in Costa Rica.
XIX. GraphidaceaeThallus crustaceous, inconspicuous, partly immersed, mainly growing on bark but occasionally on dead wood or stone. Algal cells chiefly Trentepohlia, very rarely Palmella or Phycopeltis (epiphyllous). Apothecia (lirellae) carbonaceous more or less linear, opening by a narrow slit with a well-developed proper margin except in Gymnographa, a monotypic Australian genus. In two genera, the fruit is of a compound nature, several parallel discs occurring in one lirella: these are Ptychographa (on bark in Scotland) and Diplogramma (Australia), both are monotypic. They must not be confused with Graphis elegans and allied species in which the sterile carbonaceous margin is furrowed. Two tropical genera associated with Phycopeltis are epiphyllous. Graphidaceae are among the oldest recorded lichens, attention having been drawn to them since early times by the resemblance of the lirellae on the bark of trees to hieroglyphic writing.
XX. ChiodectonaceaeSpecially distinguished in this subseries by the grouping of the somewhat rudimentary apothecia in pseudostromata in which they are almost wholly immersed. In form they are roundish or linear; the spores are septate or muriform. The thallus is thinly crustaceous and continuous: in Glyphis, Sarcographa and Sarcographina there is an amorphous upper cortex, the other genera are non-corticate. Algal cells are Trentepohlia with the exception of two epiphyllous genera associated with Phycopeltis. Genera and species are mostly tropical. Sclerophyton with five species is represented in Europe by a single British specimen, S. circumscriptum. The form of the paraphyses is a distinguishing character of the genera.
XXI. DirinaceaeA small family, which is associated with and often included under Graphidaceae. The thallus is crustaceous and corticate on the upper surface, the cortex being formed of palisade hyphae. Algal cells Trentepohlia. Apothecia are rounded or with a tendency to elongation, and, in addition to a thin proper margin, possess a stout thalline margin; the hypothecium is thick and carbonaceous. There are two genera: Dirina with twelve species has a wide distribution; Dirinastrum is monotypic and occurs on maritime rocks in Australia. In both the spores are elongate-septate, differing only in colour:
XXII. RoccellaceaeThe Roccellaceae differ from the preceding Dirinaceae chiefly in the fruticose thallus which is more or less characteristic of all the genera, though in Roccellographa it expands into foliose dimensions and in Roccellina is reduced to short podetia-like processes from a crustose base. The fronds—mostly long and strap-shaped—are protected in most of the genera by a cortex of compact palisade hyphae; in a few the outer hyphae are parallel with the long axis. The medulla is of parallel hyphae, either loose or compact. The algal cells are Trentepohlia. The apothecia are lateral except in Roccellina where they occur at the tips of the short upright fronds, and only in Roccellaria is there no thalline margin. They are superficial in all of the genera except Roccellographa, in which they are immersed and almost closed, recalling the perithecia-like The affinity of Dirinaceae and Roccellaceae with Graphidaceae was first indicated by Reinke In most of the genera only one species is recorded. Roccella, represented by twelve species, is well known for its dyeing properties, and has a wide distribution. Like other Graphidineae they are mainly plants of warm regions, many of them exclusively maritime rock-dwellers. The following synopsis of the genera is the one given by Darbishire in his monograph.
Subseries 3. Cyclocarpineae This last subseries includes the remaining twenty-nine families of Ascolichens. They are very varied both in the fungal and the algal symbionts. The fruit is more or less a discoid open apothecium. The gonidia belong to different genera of Myxophyceae and Chlorophyceae, but the most frequent are Protococcaceae. Families are based largely on thalline structure. XXIII. LecanactidaceaeBy many systematists this family is included under Graphidineae on account of the fruit structure which in some of the forms is carbonaceous and almost lirellate, and also because the algal symbiont is Trentepohlia. The thallus is primitive, being thinly crustaceous and non-corticate; the apothecium has a black carbonaceous hypothecium in two of the genera, Lecanactis and Schismatomma (Platygrapha); in the third genus, Melampydium, it is colourless. The latter is monotypic, and the spores become muriform. In the other genera they are elongate and multi-septate.
XXIV. PilocarpaceaeA small family with but one genus, Pilocarpon. It is distinguished as one of the few epiphyllous genera of lichens associated with Protococcaceous gonidia and with a distribution extending far beyond the tropics. The best known species, P. leucoblepharum, encircles the base of pine-needles with a white felted crust, or inhabits coriaceous evergreen leaves. Another species lives on fern leaves. The fruit is a discoid apothecium with a dark carbonaceous hypothecium and proper margin, and with a second thalline margin. The paraphyses are branched and interwoven above.
XXV. ChrysotrichaceaeThis family now, according to Hue
XXVI. ThelotremaceaeA tropical or subtropical family of which the leading characteristic is the deeply sunk disc of the apothecium: it has a proper hyphal margin, and, round that, an overarching thalline margin. The apothecia occur singly, or they are united in a kind of pseudostroma: in Tremotylium several grow together, while in Polystroma each new apothecium develops as an outgrowth from the thalline margin of the one already formed, so that an upright, branching succession of fruits is built up. It is a very unusual type of lichen fructification, with one species, P. Ferdinandezii, found in Spain and in Guiana. The thallus in all the genera is crustaceous with an amorphous (decomposed) cortex; or it is non-corticate. The algal cells are Trentepohlia except in Phyllophthalmaria, an epiphyllous genus associated with the alga Phycopeltis. In Polystroma the alga is unknown. Only one genus is represented in the British Isles.
XXVII. DiploschistaceaeScarcely differing from the preceding family except in the gonidia which are Protococcaceous algae. The thallus is crustaceous and non-corticate. The apothecia have a double margin but the outer thalline margin is less overarching than in Thelotremaceae. The spores in the two genera are somewhat peculiar: in Conotrema they are exceedingly long and divided by parallel septa into thirty to forty small cells; in Diploschistes (Urceolaria) they are large, muriform and brown. Conotrema contains two corticolous species; Diploschistes about thirty species mostly saxicolous. Both genera are represented in the British Isles.
XXVIII. EctolechiaceaeA family of tropical epiphyllous lichens that are associated with Protococcaceous gonidia. The thallus is primitive in character, mostly a weft of hyphae with intermingled algal cells, described as homoiomerous. The apothecia are without a thalline margin, and with a scarcely developed proper margin: their affinity is with the Lecideaceae, though in two genera, Lecaniella and Arthotheliopsis, there are gonidia below the hypothecium, a character of Lecanoraceae. The genera are nearly all monotypic; in Sporopodium has been included Lecidea phyllocharis Wainio (Sect. Gonothecium), which is distinguished by hymenial gonidia.
XXIX. GyalectaceaeThe algal cells in this family are filamentous; either Myxophyceae (Scytonema) or Chlorophyceae (Trentepohlia or Phyllactidium). The thallus is crustaceous, and in some cases homoiomerous, as in Petractis, where the alga, Scytonema, penetrates the substratum as deeply as the hyphae. Monophiale, a tropical genus, possesses two kinds of gonidia: the species that grow on bark or mosses are associated with Trentepohlia; others that have invaded the surface of leathery evergreen leaves resemble most epiphyllous lichens in being associated with the leaf alga Phyllactidium (Phycopeltis). Some species of Trentepohlia exhale when moist an odour of violets. This scent is retained in at least one genus, Jonaspis. The apothecia are superficial, and are soft, waxy and bright-coloured, with prominent margins which are however entirely hyphal: the affinity is therefore with Lecideaceae. In one genus, Sagiolechia, the fruit is carbonaceous and dark coloured. The spores of all the genera are colourless.
XXX. CoenogoniaceaeThere are only two genera in this small family, Coenogonium with Trentepohlia gonidia, and Racodium with Cladophora. Both genera follow the algal form and are filamentous. In Coenogonium the filaments are sometimes matted into a loose felted expansion. The genus is mainly tropical or subtropical and mostly rather light-coloured. There is only one British species, C. ebeneum Racodium includes 2 (?) species. One of these, R. rupestre, is sterile and resembles C. ebeneum in form and colour. The apothecia of Coenogonium are waxy and light-coloured; they are borne laterally on the filaments; the spores are simple or 1-septate.
XXXI. LecideaceaeOne of the largest lichen families as regards both genera and species, and of world-wide distribution. The algal cells are Protococcaceae. The thallus is mostly crustaceous but it becomes squamulose in Psora, a section of Lecidea; and in Sphaerophoropsis, a Brazilian genus, there are small upright fronds or stalks with lateral apothecia. The prevailing colour of the thallus is some shade of grey, but it ranges from white or yellow to dark-brown or almost black. Cephalodia appear in some of the species. The apothecia have a proper margin only, no gonidia taking part in the fruit-formation. They may be soft and waxy (biatorine) or hard and carbonaceous (lecideine). The genera are mainly based on spore characters which are very varied. The arrangement of genera given below follows that of Zahlbruckner; in several instances, both as to the limitations of genera and to the nomenclature, it differs from that of British text-books, though the general principle of classification is the same.
XXXII. PhyllopsoraceaeA small family of exotic lichens with a somewhat more developed thallus than that of the Lecideaceae, being in both of the genera squamulose or almost foliose. The apothecia are without a thalline margin; they are biatorine or lecideine; the hypothecium is formed of plectenchyma and is purple-red in one species, Phyllopsora furfuracca. The two genera differ only in spore characters. There are fifteen species, mostly corticolous, belonging to Phyllopsora; only one, from New Zealand, is recorded for Psorella.
XXXIII. CladoniaceaeAssociated with Lecideaceae in the type of apothecium, but differing widely in thallus formation. The latter is of a twofold type: the primary thallus is crustaceous, squamulose, or very rarely foliose; the secondary thallus or podetium, upright, simple or branched, is terminated by the apothecia, or broadens upwards to cup-like scyphi. Algal cells, Protococcaceae, according to Chodat, Cystococcus. Much attention has been given to the origin and development of the podetia in this family. They are superficial on granule or squamule except in the monotypic Himalayan genus Gymnoderma where they are marginal on the large leaf-like lobes. Though in origin the podetia are doubtless fruit stalks, they have become in most cases vegetative in function. The fruits are coloured yellowish, brown or red (or dark and carbonaceous in Pilophorus), and are borne on the tips of the branches or on the margins of the scyphi. In Glossodium and Thysanothecium—the former from New Granada, the latter from Australia—the apothecia occupy one side of the widened surface at the tips. Cephalodia are developed on the primary thallus of Pilophorus, and on the podetia of Stereocaulon and Argopsis.
XXXIV. GyrophoraceaeA small family of foliose lichens allied to Lecideaceae by the character of the fruit—a superficial apothecium in the formation of which the gonidia take no share. There are only three genera, distinguished by differences in spore and other characters. Dermatiscum has light-coloured thallus and fruits; of the two species, one occurs in Central Europe, the other in North America. Umbilicaria and Gyrophora are British; they are dark-coloured rock-lichens and are extremely abundant in Northern regions where they are known as “tripe de roche.” Algal cells Protococcaceae. Umbilicaria, Dermatiscum, and some species of Gyrophora are attached to the substratum by a central point. Other species of Gyrophora are rhizinose. In all there is a cortex of plectenchyma above and below. In Gyrophora the thallus may be monophyllous as in Umbilicaria, or polyphyllous and with or without rhizinae. New lobes frequently arise from protuberances or warts on the older parts of the thallus. At the periphery, in most species, growth is equal along the margins, in G. erosa There is abundant detritus formation in this family; the outer layers of the cortex are continually being sloughed, the dead tissues lying on the upper surface as a dark gelatinous layer, continuous or in small patches. On the under surface the cast-off cortex gathers into a loose confused mass of dead tissues.
XXXV. AcarosporaceaeThallus foliose, squamulose or crustaceous, sometimes scarcely developed. Algal cells Protococcaceae. Into this family Zahlbruckner has gathered the genera in which the asci are many-spored, as he considers that a character of great importance in determining relationship, but he has in doing so overlooked other very great differences. The fruit-bodies are round and completely enclosed in a thalline wall in Thelocarpon, which has however no perithecial wall. They have a proper margin only (lecideine) in Biatorella, and a thalline margin (lecanorine) in the remaining genera. In Acarospora the apothecia are sunk in the thallus. Stirton’s genus Cryptothecia
XXXVI. EphebaceaeA family of very simple structure either filamentous, foliose or crustaceous. The algal cells which give a dark colour to the thallus are Stigonema or Scytonema, members of the blue-green Myxophyceae, and consist of minute simple or branched filaments—single cell-rows in Scytonema, compound in Stigonema. In some of the genera the lichen hyphae travel within the gelatinous sheath of the filaments, both algae and hyphae increasing by apical growth so that filaments many times the length of the alga are formed as in Ephebe. In others the filaments scarcely increase beyond the normal size of the alga as in Thermutis (Gonionema); or the gelatinous algal cells may be distributed in a stratum of hyphae. The apothecia are minute and almost closed; they may be embedded in swellings of the thallus, or are more or less superficial. The spores are rather small, colourless and simple or 1-septate. The lichens of this family are rock-dwellers and are mostly to be found in hilly or Alpine regions. A tropical species, Leptogidium dendriscum, occurs in sterile condition in south-west Ireland. There are few species in any of the genera.
XXXVII. PyrenopsidaceaeIn this family are included gelatinous lichens of which the gonidium is a blue-green alga with a thick gelatinous coat, either Gloeocapsa (including Xanthocapsa) or Chroococcus. In Gloeocapsa and Chroococcus the gelatinous envelope is often red, in Xanthocapsa it is yellow, and these colours persist more or less in the lichens, especially in the outer layers. The thallus is in many cases a formless gelatinous crust of hyphal filaments mingling with colonies of algal cells as in Pyrenopsis; but small fruticose tufts are characteristic of Synalissa, and larger foliose and fruticose thalli appear in some exotic genera. A plectenchymatous cortex is formed on the thallus of Forssellia, a crustaceous genus from Central Europe, with two species only; the whole thallus is built up of a kind of plectenchyma in some others, but in most of the genera there is no tissue formed. The apothecia, as in Ephebaceae, are generally half-closed.
XXXVIII. LichinaceaeThe only family of lichens associated with Rivularia gonidia, the trichomes of which retain their filamentous form to some extent in the more highly developed genera; they lie parallel to the long axis of the squamule or of the frond except in Lichinella in which genus they are vertical to the surface. The thallus may be crustaceous, or minutely foliose, or fruticose; in all cases it is dark-brown in colour, and the gelatinous character is evident in the moist condition. The best known British genus is Lichina which grows on rocks by the sea. The apothecia are more or less immersed in the tissue; in Pterygium and Steinera they are open and superficial (the latter monotypic genus confined to Kerguelen). They are also open in Lichinella and Homopsella, both very rare genera. The spores are colourless and simple except in Pterygium and Steinera where they are elongate, and 1-3-septate.
XXXIX. CollemaceaeThe most important family of the gelatinous lichens and the most numerous. Collema is historically interesting as having first suggested the composite thallus. Algal cells, Nostoc, which retain the chain-like form except in Leprocollema, a doubtful member of the family. The thallus varies from indeterminate crusts to lobes of considerable size; occasionally the lobes are narrow and erect, forming minute fruticose structures. In the more primitive genera the thallus is non-corticate, but in the more evolved, the apical cells of the hyphae coalesce to form a continuous cellular cortex, one or more cells thick, well marked in some species, in others rudimentary; the formation of plectenchyma also occurs occasionally in the apothecial tissues of some non-corticate species. The apothecia are superficial except in Pyrenocollema, a monotypic genus of unknown locality. They are generally lecanorine, with gonidia entering into the formation of the apothecium: in some genera they are lecideine or biatorine, being formed of hyphae alone. The spores are colourless and vary in form, size and septation.
XL. HeppiaceaeA family belonging to the “blue-green” series as it is associated with a gelatinous alga, Scytonema, but is of almost entirely cellular structure and is non-gelatinous. The thallus is squamulose or minutely foliose, or is formed of narrow almost fruticose lobes; the apothecia are semi-immersed; the asci are 4-many-spored. Heppia is a wide-spread genus both in northern and tropical regions with about forty species that live on soil or rock. So far, no representative has been recorded in our Islands.
XLI. PannariaceaeThe members of this family are also non-gelatinous, though for the most part associated with blue-green gelatinous algae, Nostoc or Scytonema. The gonidia are bright-green in the genera Psoroma and Psoromaria, the former often included under Lecanora, but too closely resembling Pannaria to be dissociated from that genus. The thallus varies from being crustaceous to squamulose or foliose, and has a cortex of plectenchyma on the upper and sometimes also on the lower surface. The apothecia are superficial or lateral and with or without a thalline margin (lecanorine or biatorine), the spores are colourless. Zahlbruckner has included Hydrothyria in this family. It is a monotypic aquatic genus found in North America and very closely allied to Peltigera. The British species of the genus, familiarly known as Coccocarpia, have been placed under Parmeliella, the former name being restricted to the tropical or subtropical species first assigned to Coccocarpia and distinguished by the cortex, the hyphae forming it lying parallel with the surface though forming a regular plectenchyma. An Antarctic lichen Thelidea corrugata with Palmella gonidia is doubtfully included: the thallus is foliose, the apothecia biatorine with colourless 1-septate spores.
XLII. StictaceaeThallus foliose, mostly horizontal, with a plectenchymatous cortex on both surfaces, a tomentum of hair-like hyphae taking the place of rhizinae on the lower surface. Algal cells Protococcaceae or Nostoc. Cephalodia and cyphellae or pseudocyphellae often present. Apothecia superficial or lateral; spores colourless or brown, variously septate. The highly organized cortex and the presence of aeration organs—cyphellae or pseudocyphellae—which are almost solely confined to the genus Sticta give this family a high position as regards vegetative development. The two genera are of wide distribution, but Sticta is more abundant in the Southern Hemisphere. Lobaria pulmonaria is one of our largest lichens.
XLIII. PeltigeraceaeA family of heteromerous foliose lichens containing in some instances blue-green (Nostoc), in others bright-green (Protococcaceae) gonidia, and thus representing a transition between these two series. They have large or small lobes and grow on the ground or on trees. Cephalodia, either ectotrophic (Peltidea) or endotrophic (Solorina), occur in the family and further exemplify the capacity of the fungus hyphae to combine with different types of algae. The upper surface is a wide cortex of plectenchyma, which in some forms (Nephromium) is continued below. In the non-corticate under surface of Peltigera, the lower hyphae grow out in hairs or rhizinae, very frequently brown in colour. Intercalary growth of the upper tissues stretches the thallus and tears apart the lower under surface so that the hair-bearing areas become a network of veins, with the white exposed medulla between. In Peltigera canina there is further growth and branching of the hyphae in the veins, adding to the bulk of the interlacing ridges. From all other foliose lichens Peltigeraceae are distinguished by the flat wholly appressed or peltate apothecia without a thalline margin which arise mostly on the upper surface, but in Nephromium on the extreme Bitter
XLIV. PertusariaceaeThallus crustaceous, often rather thick and with an amorphous cortex on the upper surface. Algal cells Protococcaceae. Apothecia solitary or several immersed in thalline warts, generally with a narrow opening which barely exposes the disc, and which in one genus, Perforaria, is so small as almost to constitute a perithecium; spores are often very large and with thick walls; some if not all are multinucleate and germinate at many points. In the form of the fruit, this family stands between Pyrenocarpeae and Gymnocarpeae, though more akin to the latter. Perforaria, with two species, belongs to New Zealand and Japan. Pertusaria has a world-wide distribution, and Varicellaria, a monotypic genus, with a very large two-celled spore, is an Alpine plant, recorded from Europe and from Antarctic America.
XLV. LecanoraceaeThallus mostly crustaceous, occasionally squamulose or very rarely minutely fruticulose. The squamulose thallus is corticate above, the under surface appressed and attached to the substratum by penetrating hyphae, often effigurate at the circumference. Algal cells Protococcaceae. Apothecia well distinguished by the thalline margin; spores colourless, simple or variously septate or muriform. Lecanora, Ochrolechia, Lecania, Haematomma and Phlyctis are cosmopolitan genera, some of them with a very large number of species; the other genera are more restricted in distribution and generally with few species. The genus Candelariella is of uncertain position; the spores are 8 or many in the ascus and are simple or 1-septate, and not unfrequently become polarilocular as in Caloplacaceae, but there is no parietin present.
XLVI. ParmeliaceaeA very familiar family of foliose lichens. Genera and species are dorsiventral and stratose in structure, though some Cetrariae are fruticose in habit. Algal cells are Protococcaceae; in Physcidia they are Palmellae. In every case the upper surface of the thallus is corticate and generally of plectenchyma, the lower being somewhat similar, but in Heterodea and Physcidia, monotypic Australasian genera, the upper cortex is of branching hyphae parallel with the surface, the lower surface being non-corticate. The Parmeliae are mostly provided with abundant rhizinae; in Cetrariae and Nephromopsis these are very sparingly present, while in Anzia (including Pannoparmelia) the medulla passes into a wide net-like structure of anastomosing hyphae. In Heterodea, cyphellae occur on the under surface as in Stictaceae; and in Cetraria islandica bare patches have been described as pseudocyphellae. The latter lichen is one of the few that are of value as human food. Special aeration structures are present on the upper cortex of Parmelia aspidota.
XLVII. UsneaceaeThis also is a familiar family of lichens, Usnea barbata the “bearded moss” being one of the first lichens noted and chronicled. Algal cells Protococcaceae. Structure radiate, the upright or pendulous habit characteristic of the family securing all-round illumination. Special adaptations of the cortex or of the internal tissues have been evolved to strengthen the thallus against the strains incidental to their habit of growth as they are attached in nearly all cases by one point only, by a special sheath, or by penetrating hold-fasts. Apothecia are superficial or marginal and sometimes shortly stalked; spores are simple or variously septate. Ramalina and Usnea, the most numerous, are cosmopolitan genera; Alectoria inhabits northern or hilly regions. The genus Evernia, also cosmopolitan, represents a transition between foliose and fruticose types; the fronds of the two species, though strap-shaped and generally upright, are dorsiventral and stratose, the gonidia for the most part lying beneath one surface; the other (lower) surface is either white or very dark-coloured. Everniopsis, formed of thin branching strap-shaped fronds, is also dorsiventral. A number of genera, Thamnolia, Siphula, etc. are of podetia-like structure, generally growing in swards. Several of them have been classified with Cladoniae, but they lack the double thallus. One of these, Endocena, a sterile monotypic Patagonian lichen, with stiff hollow coralloid fronds, was classified by Hue
XLVIII. CaloplacaceaeIn this family Zahlbruckner has included the squamulose or crustaceous lichens with colourless polarilocular spores, relegating those with more highly developed thallus or with brown spores to other families. He has also substituted the name Caloplaca for the older Placodium, the latter being, as he considers, less well defined. Algal cells are Protococcaceae. The thallus is mostly light-coloured, generally some shade of yellow, and, with few exceptions, contains parietin, which gives a purple colour on the application of potash. The squamulose forms are closely appressed to the substratum, and have often a definite rounded outline (effigurate). The spores have a thick median septum with a loculus at each end and a connecting canal In Blastenia the outer thalline margin is obscure or absent—though gonidia are frequently present below the hymenium. Caloplacaceae occur all over the globe; they are among the most brilliantly coloured of all lichens. Polycauliona Hue
XLIX. TeloschistaceaePolarilocular colourless spores are the distinguishing feature of this family as of the Caloplacaceae. Algal cells Protococcaceae. The thallus of Teloschistaceae is more highly developed, being either foliose or fruticose, though never attaining to very large dimensions. The cortex of Xanthoria (foliose) is plectenchymatous, that of Teloschistes (fruticose) is fibrous. The species of both genera are yellow or greenish-yellow due to the presence of the lichen-acid parietin. Both genera have a wide distribution over the globe, more especially in maritime regions.
L. BuelliaceaeA family of crustaceous lichens distinguished by the brown two-celled spores. Algal cells Protococcaceae. Zahlbruckner has included here Buellia and Rinodina; the former with a distinctly lecideine fruit and with thinly septate spores; the latter lecanorine and with spores of the polarilocular type, with a very wide central septum pierced in most of the species by a canal which may or may not traverse the middle lamella of the wall. Rinodina is closely allied to Physciaceae, while Buellia has more affinity with Lecideaceae and is near to Rhizocarpon. Both genera are of world-wide distribution.
LI. PhysciaceaeThallus foliose or partly fruticose, and generally attached by rhizinae. Algal cells Protococcaceae. The spores resemble those of Rinodina, dark-coloured with a thick septum and reduced cell-lumina. As in that species there may be a second septum in each cell, giving a 3-septate spore; but that is rare. Pyxine, a tropical or subtropical genus, is lecanorine only in the very early stages; it soon loses the thalline margin. Anaptychia is differentiated from Physcia by the subfruticose habit, though the species are nearly all dorsiventral in structure, only a few of them being truly radiate and corticate on both surfaces. The upper cortex of Anaptychia is fibrous, but that character appears also in most species of Physcia either on the upper or the lower side. Physcia and Anaptychia are widely distributed.
C. *HymenolichensFungus a Basidiomycete, akin to Thelephora. Algal cells Scytonema or Chroococcus. Thallus crustaceous, squamulose or foliose. Spores colourless, produced on basidia, on the under surface of the free thallus. The Hymenolichens
II. NUMBER AND DISTRIBUTION OF LICHENS1. ESTIMATES OF NUMBERCalculations have been made and published, once and again, as to the number of lichen species occurring over the globe or in definite areas. In 1898 FÜnfstÜck stated that about 20,000 different species had been described, but as many of them had been proved to be synonyms, and since many must rank as forms or varieties, the number of well-authenticated species did not then, according to his estimate, exceed 4000. Many additional genera and species have, however, been discovered since then. In Engler and Prantl’s Pflanzenfamilien, over 50 families and nearly 300 genera find a place, but even in these larger groupings opinions differ as to the limits both of genera and families, and lichenologists would not all accept the arrangement given in that volume. FÜnfstÜck has reckoned that of his estimated 4000, about 1500 are European and of these at least 1200 occur in Germany. Probably this is too low an estimate for that large country. Leighton in 1879 listed, in his British Lichen Flora, 1710 in all, and, as the compilation includes varieties, it cannot be considered as very far astray. On comparing it with Olivier’s A general idea of the comparative numbers of the different types of lichens may be gathered from Hue’s compilation of exotic lichens 2. GEOGRAPHICAL DISTRIBUTIONA. General SurveyThe larger foliose and fruticose lichens are now fairly well known and described for Europe, and the knowledge of lichens in other continents is gradually increasing. It is the smaller crustaceous forms that baffle the investigator. The distribution of all lichens over the surface of the earth is controlled by two principal factors, climate and substratum; for although lichens as a rule require only support, they are most of them restricted to one or another particular substratum, either organic or inorganic. As organisms which develop slowly, they require an unchanging substratum, and as sun-plants they avoid deeply shaded woodlands: their occurrence thus depends to a large extent on the configuration and general vegetation of the country. Though so numerous and so widely distributed, lichens have not evolved that great variety of families and genera characteristic of the allied fungi and algae. They conform to a few leading types of structure, and thus the Orders and Families are comparatively few, and more or less universal. They are most of them undoubtedly very old plants and were probably wide-spread before continents and climates had attained their present stability. Arnold During the Ice age of the Quaternary period, not many new species can have arisen, and such forms as were not killed off must have been driven towards the south. As the ice retreated the valleys were again stocked with southern forms, and northern species were left behind on mountain tops all over the globe. In examining therefore the distribution of lichens, it will be found that the distinction between different countries is relative, certain families being more or less abundant in some regions than others, but, in general, nearly all being represented. Certain species are universal, where similar conditions prevail. This is especially true of those species adapted to extreme cold, as that condition, normal in polar regions, recurs even on the equator if the mountains reach the limit of perpetual snow; the vertical distribution thus follows on the lines of the horizontal. In all the temperate countries we find practically the same families, with some few exceptions; there is naturally more diversity of genera and species. Genera that are limited in locality consist, as a rule, of one or few species. In this category, however, are not included the tropical families or genera which may be very rich in species: these are adapted to extreme conditions of heat and often of moisture, and cannot exist outside tropical or subtropical regions, extreme heat being more restricted as to geographical position than extreme cold. In the study of distribution the question which arises as to the place of origin of such widely distributed plants is one that is difficult to solve. Wainio The distribution of Phanerogams is connected, as Wainio points out, with causes anterior to the present geological era, but this cannot be the case in a genus so labile and probably so recent as Cladonia, though some of the species have existed long enough to spread and establish themselves from pole to pole. Endemic species, or those that are confined to a comparatively limited area, are easily traced to their place of origin, that being generally the locality where they are found in most abundance, and as a general rule in the centre of that area, though there may be exceptions: a plant for instance that originated on a mountain would migrate only in one direction—towards the regions of greater cold. The difficulty of determining the primitive stations of cosmopolitan, or of widely spread, species is much greater, but generally they also may be referred to their area of greatest abundance. Thus a species may occur frequently in one continent and but rarely in another, even where the conditions of climate, etc., are largely comparable. It may therefore be inferred that the plant has not yet reached the full extent of possible distribution in the less frequented area. As examples of this, Wainio cites, among other instances, Cladonia papillaria, which has a very wide distribution in Europe, A detailed account of species distribution according to these indications is given by Wainio and is full of interest. No such attempt has been made to deal with any other group, and the distribution of genera and species can only be suggested. An exhaustive comparison of the lichens of different regions is beyond the purpose of our study and is indeed impossible as, except in some limited areas, or for certain species, the occurrence and distribution are not fully known. It is in any case only tentatively that genera or species can be described as local or rare, until diligent search has been made for them over a wider field. The study of lichens from a floristic point of view lags behind that of most other groups of plants. The larger lichen forms have received more attention, as they are more evident and more easily collected; but the more minute species are not easily detected, and, as they are largely inseparable from their substratum of rocks, or trees, etc., on which they grow, they are often difficult to collect. They are also in many instances so indefinite, or so alike in outward form, that they are liable to be overlooked, only a microscopic examination revealing the differences in fruit and vegetative structure. Though much remains to be done, still enough is known to make the geographical distribution of lichens a subject of extreme interest. It will be found most instructive to follow the usual lines of treatment, which give the three great divisions: the Polar, the Temperate and the Tropical regions of the globe. B. Lichens of Polar RegionsStrictly speaking, this section should include only lichens growing within the Polar Circles; but in practice the lichens of the whole of Greenland and those of Iceland are included in the Arctic series, as are those of Alaska: the latitudinal line of demarcation is not closely adhered to. With the northern lichens may also be considered those of the Antarctic continent, as well as those of the islands just outside the Antarctic Circle, the South Cold winds are a great feature of both poles, and the lichens that by structure or habit can withstand these are the most numerous; those that have a stout cortical layer are able to resist the low temperatures, or those that grow in tufts and thus secure mutual protection. In Arctic and Subarctic regions, 495 lichens have been recorded, most of them crustaceous. Among the larger forms the most frequently met are certain species of Peltigera, Parmelia, Gyrophora, Cetraria, Cladonia, Stereocaulon and Alectoria. Among smaller species Lecanora tartarea spreads everywhere, especially over other vegetation, Lecanora varia reaches the farthest limits to which wood, on which it grows, has drifted, and several species of Placodium occur constantly, though not in such great abundance. Over the rocks spread also many crustaceous Lecideaceae too numerous to mention, one of the most striking being the cosmopolitan Rhizocarpon geographicum. Wainio Wainio determined a large number of lichens with many new species, but the region is colder than that of Lappland, and trees with tree-lichens are absent, with the exception of those given above. In Arctic Siberia, Elenkin The Antarctic Zone and the neighbouring lands are less hospitable to plant life than the northern regions, and there is practically no accumulation of detritus. Collections have been made by explorers, and several lists have been published which include a marvellous number of species common to both Poles, if the subantarctic lands are included in the survey. An analytic study of the various lists has been published by Darbishire A somewhat different view of the Antarctic lichen flora is indicated by Hue Hue determined 126 species, but as 15 of these came from the Magellan regions only 111 were truly Antarctic. Of these 90 are new species, 29 of them belonging to the genus Buellia. Hue considers, therefore, that in Antarctica there is a flora that, with the exception of cosmopolitan species, C. Lichens of the Temperate ZonesRegions outside the Polar Circles which enjoy, on the whole, cool moist climates, are specially favourable to lichen growth, and the recorded numbers are very large. The European countries are naturally those in which the lichen flora is best known. Whereas polar and high Alpine species are stunted in growth and often sterile, those in milder localities grow and fruit well, and the more highly developed species are more frequent. Parmeliae, Nephromae, Usneae and Ramalinae become prominent, especially in the more northern districts. Many Arctic plants are represented on the higher altitudes. A comparison has been made between the lichens of Greenland and those of Germany: of 286 species recorded for the former country, 213 have been found in Germany, the largest number of species common to both countries being crustaceous. Lindsay There is an astonishing similarity of lichens in the Temperate Zone all round the world. Commenting on a list of Chicago lichens by Calkins If we contrast American lichens with these collected in South Siberia near Lake Baikal A marked difference as regards species is noted between the Flora of Minnesota and that of California. Herre With the Japanese Flora may be compared a list It is unnecessary to describe in detail the British lichens. Some districts have been thoroughly worked, others have barely been touched. The flora as a whole is of a western European type showing the influence of the Gulf Stream, though there is also a representative boreal growth on the moorlands and higher hills, especially in Scotland. Such species as Parmelia pubescens, P. stygia and P. alpicola recall the Arctic Circle while Alectoriae, Cetrariae and Gyrophorae represent affinity with the colder temperate zone. In the southern counties such species as Sticta aurata, S. damaecornis, Phaeographis Lyellii and Lecanora (Lecania) holophaea belong to the flora of the Atlantic seaboard, while in S.W. Ireland the tropical genera Leptogidium and Anthracothecium are each represented by a single species. The tropical or subtropical genus Coenogonium occurs in Great Britain and in Germany, with one sterile species, C. ebeneum. Enterographa crassa is another of our common western lichens which however has travelled eastwards as far as Wiesbaden. Roccella is essentially a maritime genus of warm climates: two species, R. fuciformis and R. fucoides, grow on our south and west coasts. The famous R. tinctoria is a Mediterranean plant, though it is recorded also from a number of localities outside that region and has been collected in Australia. In the temperate zones of the southern hemisphere are situated the great narrowing projections of South Africa and South America with Australia and New Zealand. As we have seen, the Antarctic flora prevails more or less in the extreme southern part of America, and the similarity between the lichens of that country and those of New Zealand is very striking, especially in the fruticulose forms. There is a very abundant flora in the New Zealand islands with their cool moist climate and high mountains. Churchill Babington A warm current from the tropical Pacific Ocean passes southwards along the East Coast of Australia, and Wilson The continent of Africa on the north and east is in so close connection with Europe and Asia that little peculiarity in the flora could be expected. In comparing small representative collections of lichens, 37 species from Egypt and 20 from Palestine, MÜller The vast African continent—more especially the central region—has been but little explored in a lichenological sense; but in 1895 Stizenberger Wainio’s D. Lichens of Tropical RegionsIn the tropics lichens come under the influence of many climates: on the high mountains there is a region of perpetual snow, lower down a gradual change to temperate and finally to tropical conditions of extreme heat, and, in some instances, extreme moisture. There is thus a bewildering variety of forms. By “tropical” however the warmer climate is always implied. Several families and genera seem to flourish best in these warm moist conditions and our familiar species grow there to a large size. Among crustaceous families Thelotremaceae and Graphidaceae are especially abundant, and probably originated there. In the old comprehensive genus Graphis, 300 species were recorded from the tropics. It should be borne in mind that Trentepohlia, the alga that forms the gonidia of these lichens, is very abundant in the tropics. Coenogonium, a genus containing about twelve species and also associated with Trentepohlia, is scarcely found in Europe, except one sterile species, C. ebeneum. Other species of the genus have been recorded as far north as Algeria in the Eastern Hemisphere and Louisiana in the Western, while one species, C. implexum, occurs in the southern temperate zone in Australia and New Zealand. Of exclusively tropical lichens, the Hymenolichens are the most noteworthy. They include three genera, Cora, Corella and Dictyonema, the few species of which grow on trees or on the ground both in eastern and western tropical countries. Other tropical or subtropical forms are Oropogon loxensis, similar to Alectoria in form and habit, but with one brown muriform spore in the ascus; it is only found in tropical or subtropical lands. Physcidia Wrightii (Parmeliaceae) is exclusively a Cuban lichen. Several small genera of Pyrenopsidaceae such as Jenmania (British Guiana), Paulia (Polynesia) and Phloeopeccania (South Arabia) seem to be confined to very hot localities. On the other hand Collemaceae are rare: Wainio records from Brazil only four species of Collema, with nine of Leptogium. Among Pyrenolichens, Paratheliaceae, Mycoporaceae and Astrotheliaceae are almost exclusively of tropical distribution, and finally the leaf lichens with very few exceptions. These follow the leaf algae, Mycoidea, Phycopeltis, etc., which are so abundant on the coriaceous long-lived green leaves of a number of tropical Phanerogams. All the Strigulaceae are epiphytic lichens. Phyllophthalmaria (Thelotremaceae) is also a leaf genus; one of the species, Ph. coccinea, has beautiful carmine-red apothecia. The genera of the tropical family Ectolechiaceae also inhabit leaves, but they are associated with Protococcaceae; one of the genera Sporopodium Pilocarpon, an epiphytic genus, is associated with Protococcaceae; one of the species, P. leucoblepharum, spreads from the bark to the leaves of pine-trees; it is widely distributed and has also been reported in the Caucasus A series of lichens from the warm temperate region of Transcaucasia investigated by Steiner Much of Asia lies within tropical or subtropical influences. Several regions have received some amount of attention from collectors. From Persia there has been published a list of 59 species determined by MÜller A small collection of 53 species from India, near to Calcutta, published by Nylander Leighton On the whole it is safe to conclude from the above and other publications that the exceptional conditions of the tropics have produced many distinctive lichens, but that a greater abundance both of species and individuals is now to be found in temperate and cold climates. III. FOSSIL LICHENSIn pronouncing on the great antiquity of lichens, proof has been adduced from physiological rather than from phytogeological evidence. It would have been of surpassing interest to trace back these plants through the ages, The reason for this is fairly obvious: not only are the primitive thalline forms too indistinct for recognizable preservation, but all lichens are characterized by the gelatinous nature of the hyphal or of the algal membranes which readily imbibe water. They thus become soft and flaccid and unfit to leave any impress on sedimentary rocks. It has also been pointed out by Schimper The few records that have been made are here tabulated in chronological order: 1. Palaeozoic. Schimper decides that there are no records of lichens in the earlier epochs. Any allusions 2. Mesozoic. Braun 3. Cainozoic. In the brown-coal formations of Saxony Engelhardt Other records on brown coal or lignite are: Verrucarites geanthricis Schimper has questioned, as he well might, Ludwig’s More authentic also are the lichens from the amber beds of KÖnigsberg and elsewhere collected by Goeppert and others. These deposits are Cainozoic and have been described by Goeppert and Menge In a later work Goeppert and Menge Another series of lichens found in recent deposits in North Europe has been described by Sernander Of especial lichenological interest in these northern regions was the Calcareous Tufa or Calc-sinter in which Sernander also found subfossil lichens—distinct impressions of Peltigera spp. and the foveolae of endolithic calcicolous species. In another category he has placed Ramalina fraxinea, Graphis sp. and Opegrapha sp., traces of which were embedded with drift in the Tufa. In the two Graphideae the walls of apothecia and pycnidia were preserved. Sernander considers their presence of interest as testifying to warmer conditions than now prevail in these latitudes. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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