Fig. 107.Erysiphe cichoracearum: a mycelium-threads; b sterile hypha (“pollinodium”); c fertile hypha (ascogone or archicarp); d and e young ascocarps.

A characteristic feature of the Mildew-Fungi is the thin, pseudo-parenchymatous covering of the ascocarp, enclosing one (Podosphæra and Sphærotheca; compare Thelebolus among the Hemiasci) or a few asci (Fig. 108 c), which do not form any hymenium, but are irregularly placed. The cells of the ascocarp-envelope are often prolonged into hair-like appendages. The ascocarps are developed from the mycelium at places where two hyphæ cross each other (Fig. 107). At these places two short and erect hyphæ are produced side by side. The one from the lower hypha (Fig. 107 c) assumes an ellipsoidal shape, and is known as the archicarp or ascogone, while the other (“pollinodium”) arches over the ascogone. From the latter one ascus may be at once developed (Sphærotheca, etc.), or after its division several asci may be produced, each developed from one division. The sterile hypha (termed “pollinodium,” since it was formerly, but erroneously, supposed to fertilise the ascogone) produces a number of branches, and forms the pseudo-parenchymatous envelope of one cell in thickness, enclosing the asci.

Fig. 108.Erysiphe communis. A small portion of a leaf with this Fungus growing upon it (considerably magnified). The hyphæ b and d do not belong to this Fungus, but are reproductive organs of a pyrenomycetous Fungus parasitic upon it (Cicinnobolus).

Many plants, both cultivated and wild, are attacked by various species of Mildew. A common means of prevention against their attacks is to dust the diseased parts with sulphur.

Fig. 109.Eurotium glaucum: α portion of mycelium lying horizontally; β vertically-placed conidiophore; the mycelium gives rise to another branch near α; the conidia are abstricted from short flask-shaped cells; b a ripe conidium; c, d germinating conidia; e spirally-twisted hypha, commencement of an ascocarp; f a stage later; g still later, the hypha at the base of the coil has given off branches which are applied to it; h, i sections of young ascocarps.

Sphærotheca pannosa occurs on the leaves of Roses, and on the fruit of Peaches and Apricots. S. castagnei on Humulus, Cucumis, etc.—Erysiphe tuckeri grows on the leaves and fruit of the Vine; it spins its hyphæ over the bunches of grapes, curtails their growth, and causes them to burst, and to become decayed and rotten (Grape-disease). The Fungus was first noticed in England in 1845, and later was found in all countries where grapes are grown. It is only known in the conidial form (“Oidium tuckeri”). Many other species of Erysiphe are found on herbaceous plants.—Microsphæra has appendages which are repeatedly forked at their extremities. M. grossulariæ on Ribes grossularia.—Uncinula has appendages with spirally-coiled extremities; on Salix and Acer.—Phyllactinia has a circle of bristle-like appendages with dilated bases. P. guttata on Corylus, Fraxinus, Fagus, etc.

Order 2. Perisporiaceæ, Moulds and Mildews. A group of Fungi widely distributed and found in all situations. Usually they have a well-developed surface mycelium, and small, round, seldom conspicuous ascocarps, containing ovoid, pulley-like spores. They are partly saprophytic, partly parasitic, in the latter condition having a brown mycelium.

Fig. 110.Eurotium glaucum: a longitudinal section of a half-ripe ascocarp, bounded externally by a well-defined layer of cells, enclosing asci in various stages of development; b a semi-ripe, c an almost ripe ascus; d and e spores seen from the edge and side; f germinating spore twenty-two hours after been sown in plum juice.

Eurotium glaucum (= E. herbariorum, Figs. 109, 110) and E. repens live on dead organic matter, preserved fruits, etc. The conidial forms of both species are known as “Moulds” (Fig. 109), and formerly were described under the name “Aspergillus glaucus.” The conidia for some time remain attached to each other in chains (Fig. 109 a); they are abstricted from sterigmata arranged radially on the spherical, swollen end of the conidiophore. The small yellow or brownish ascocarps are frequently found in herbaria, especially when the specimens have been insufficiently dried. Aspergillus fumigatus and others are pathogenic, causing mycosis in warm-blooded animals.

Fig. 111.Penicillium crustaceum: a conidia (× 300); b germination of conidia; c small portion of mycelium, produced from a conidium at *, with five conidiophores; d young conidiophore (× 630), a flask-shaped cell is abstricting a conidium; e the same conidiophore after 9–10 hours.

Fig. 112.Penicillium crustaceum: a two spirally-coiled hyphæ arise from the mycelium, from one of which (archicarp) the asci are produced; b a further step in the development of the ascocarp; the branching archicarp is surrounded by sterile hyphæ; c section of young ascocarp; the larger hyphæ in the centre are the ascogenous hyphæ; these are enclosed by a pseudo-parenchyma of sterile hyphæ (× 300); d series of ripe asci with spores; e four ascopores seen laterally; f germinating ascospores (× 800).

Penicillium crustaceum (P. glaucum, Figs. 111, 112) is an exceedingly common “Mould.” Its mycelium appears very frequently on any organic matter which is permitted to remain untouched, and soon covers it with a dense mass of blue-green conidiophores. These branch at their summits and bear flask-shaped cells from which the conidia are abstricted. The ascocarps which, both in size and colour, resemble grains of sand, have only been obtained in luxuriant cultivation with a limited supply of oxygen.

Capnodium salicinum (Fumago salicina, Cladosporium fumago), a common Mildew, forms dark overgrowths on the leaves and branches of various shrubs (Poplars, Elms, Willows) and on Hops. The conidia appear in various forms, as on conidiophores, in conidiocarps with large multicellular conidia, and in conidiocarps with small unicellular conidia; in nutritive solutions yeast-like conidia are also developed.—Apiosporium pinophilum produces mildew on the leaves of Abies alba and Picea excelsa. (The conidial-forms were formerly described as “Antennaria pinophila”).

Order 3. Tuberaceæ, Truffles. The Fungi belonging to this order are entirely subterranean. The mycelium is filamentous, and partly parasitic upon the roots of plants, especially trees, in its neighbourhood; it is then known as Mycorhiza. The fruit-body is relatively large, in some cases about the size of a hen’s egg. Internally it is traversed by a number of winding passages (Fig. 113 a), the walls of which are coated with the asci. The asci (b) contain only a small number of spores, and these are set free by the putrefaction of the fruit-body. Conidia are unknown.

Fig. 113.Tuber melanosporum: a fruit-body (nat. size), a portion having been removed to show the internal structure; b an ascus with ascospores.

Tuber melanosporum, T. brumale, T. æstivum, and other species are edible. Terfezia leonis and Choiromyces mæandriformis are also edible. The Truffles are always found in woods and under trees, and disappear when these are destroyed. France and Italy produce the best and the largest number of Truffles, which are hunted by specially trained dogs and pigs.

In Elaphomyces (Stag-Truffle) the fruit-body has a corky external layer, and is inedible. Some of the species are found in this country. E. granulatus is parasitic on the roots of the Fir.

Family 3. Pyrenomycetes.

In this family the hymenium is enclosed in small fruit-bodies, perithecia (Fig. 120 b), which appear to the naked eye as small dots. In shape they resemble a globe or a flask with a narrow mouth, through which the spores are ejected (peronocarpic ascocarps). Different kinds of reproduction—conidia, pycnidia (chiefly with microconidia), chlamydospores, and perithecia—are found in the same species. The various stages in the life-history of these Fungi are so dissimilar, that formally they were considered to be different genera. Ergot furnishes a very good example.

Fig. 114.—A small portion of an ovary attacked with Claviceps purpurea (Sphacelia).

Fig. 115.—An ovary with the conidial stage of Claviceps purpurea (Sphacelia).

This family may be subdivided into 3 sub-families.

Sub-Family 1. Hypocreales.

The perithecia are pale, fleshy, brightly coloured, and generally aggregated on a stroma. Conidia and chlamydospores occur very frequently. Only one order.

Order. Hypocreaceæ. In this order the majority are parasites upon Flowering-plants (Nectria, Polystigma, Epichloë, Claviceps); but some are parasites upon Fungi (Hypomyces, Melanospora), or upon insects (Cordyceps).

Fig. 116.Claviceps purpurea. A Sclerotium with stromata (cl) (× by 2). B Stroma divided longitudinally to show the perithecia (cp). C A perithecium with the surrounding hyphæ (hy). D An ascus ruptured, with the eight filamentous ascospores emerging.

The most important member of this order is the Ergot (Claviceps purpurea, Figs. 114, 115, 116). This Fungus is found in the flowers of many species of Grasses, especially the Rye, attacking and destroying the ovaries. In the FIRST or CONIDIAL STAGE of the attack, the ovaries are found covered with a white, irregularly folded mycelium (Fig. 114 m, Fig. 115), formed of numerous hyphæ woven together and penetrating the wall of the ovary. From these a number of hyphæ (Fig. 114 a) project into the air and abstrict from their apices the conidia (b) which serve as reproductive organs. The mycelium also secretes a sticky, stinking fluid (honey-dew) in which the conidia are embedded in great numbers. The honey-dew exudes from the bases of the glumes, and is greedily sought by flies, which thus carry the conidia to other ovaries. In this manner fresh ears are infected, which might escape were the conidia only distributed by the wind. This stage formerly was regarded as an independent Fungus, known as Sphacelia segetum (Fig. 115). On germination, the conidia produce either a new mycelium (Fig. 114 d, c), or new conidia. The SECOND or SCLEROTIUM STAGE is the one in which the Fungus passes the winter. The mycelium penetrates deeper and deeper into the attacked ovaries, their tissues are destroyed and replaced by the hyphæ, which gradually become more and more felted together. A firm, pseudo-parenchymatous mass of hyphæ is thus formed at the base of the loosely-woven Sphacelia, which is in part transformed into the hard sclerotium, and the remainder thrown off. A dark, hard, poisonous body, longer than the natural grain, is thus formed; these bodies are known as Ergots, and were formerly considered to be a distinct species,—Sclerotium clavus (“Secale cornutum,” Ergot, Fig. 116 A, c). The THIRD STAGE, described as Claviceps purpurea, is developed in the following spring from the germinating sclerotium, which produces dark-red stromata with short stalks. In the stroma numerous perithecia with asci and ascospores are produced. The latter may infect young flowers of the cereals, in which the disease is then developed as before.

Fig. 117.Nectria cinnabarina: a branch of Acer pseudoplatanus, with conidial-layers and perithecia (nat. size); b a conidial-layer (Tuberculoria vulgaris); c, a mass of perithecia. (b and c × 8.)

Several species of the genus Nectria, with blood-red perithecia, are found as dangerous parasites, especially N. ditissima, which causes “Canker” in the Beech, Ash, and Apple, etc.; N. cucurbitula, which appears on Pine-trees, and N. cinnabarina (Fig. 117), whose conidial form was formerly named Tubercularia vulgaris.—Polystigma rubrum forms shining red spots on the green leaves of Prunus-species.—Epichloë typhina is parasitic on the sheaths of Grasses, on which it first forms a white conidial-layer, later on a yellow layer of perithecia.—Cordyceps (Chrysalis Fungus, Figs. 118, 119) lives in and destroys insects, and after compassing their death produces the club-formed, generally yellow, stromata, one part of which bears conidia (Isaria) and another perithecia. C. militaris (Fig. 118) on the chrysalides and caterpillars of moths, is the most common.

The so-called Botrytis bassiana, which produces the disease known by the name of “Muscardine,” in silkworms, is probably a conidial form belonging to Cordyceps.

Fig. 118.Cordyceps militaris. I Stromata with conidiophores (Isaria farinosa). II A larva, with stromata, bearing perithecia. III A spore.

Fig. 119.Cordyceps robertii on the larva of Hepialus virescens: a stalk of stroma; b perithecia.

Sub-Family 2. Sphæriales.

To this sub-family belong the majority of the Pyrenomycetes. The perithecia are of a firm consistence (tough, leathery, woody or carbonaceous), and of a dark colour. Their covering is quite distinct from the stroma when this structure is present. The stromata are sometimes very large, and may be either cushion-like, crustaceous, upright and club-like, or branched bodies. In general, small, inconspicuous Fungi, living on dead vegetable matter, sometimes parasites. Free conidiophores and conidiocarps are known in many species, and in several, chlamydospore-like forms of reproduction. Orders 3–18 constitute the Sphæriaceæ of older systematists.

Fig. 120.Strickeria obducens: a a portion of an Ash-branch with the bark partly thrown off; on the wood are numerous black perithecia (× 20); b longitudinal section through a perithecium; c a spore; d longitudinal section through a pycnidium whose ascospores are being ejected; e portion of the same, with hyphæ and spores.

Order 1. Sordariaceæ.—Fungi living on dung with fragile perithecia, either aerial or buried in the substratum. The dark brown or black spores have either a mucilaginous envelope (Sordaria, etc.) or mucilaginous appendages (Podospora), by means of which their expulsion and distribution are promoted.

Order 2. Chætomiaceæ. Perithecia fragile, free, bearing on the summit a tuft of hairs. Chætomium, on decaying vegetable matter.

Orders 3–7. Perithecia scattered or aggregated, situated from the commencement on the surface of the substratum. Stroma wanting.

Order 3. Trichosphæriaceæ. Trichosphæria parasitica (Fig. 121), on Abies alba; Herpotrichia nigra on Picea excelsa and Pinus montana.

Fig. 121.Trichosphæria parasitica: a a twig of Abies alba, with epiphytic mycelium; b a leaf with mycelium and sporangia (magnified); c a sporangium (× 60); d an ascus with spores (× 550).

Order 4. Melanommaceæ. Rosellinia quercina lives in the roots of 1–3-year-old Oaks, and destroys the plants.

Order 5. Ceratostomaceæ.

Order 6. Amphisphæriaceæ. Strickeria obducens (Fig. 120) has brick-like spores, and lives aggregated on the hard branches of Fraxinus.

Order 7. Lophiostomaceæ.

Order 8. Cucurbitariaceæ. Perithecia tufted, at first embedded, then breaking through, often situated upon an indistinct stroma.

Orders 9–13. The perithecia remain embedded, and are only liberated by the casting off of the covering layers of the substratum. Stroma wanting.

Order 9. Sphærellaceæ. The species of Sphærella have colourless, bicellular spores. They live upon the leaves of many plants, and develope spherical perithecia upon the fallen leaves.

Order 10. Pleosporaceæ. The conidial-forms of Pleospora herbarum and P. vulgaris form a black covering on various plants, known as “smuts.”—Venturia ditricha occurs on the underside of dry Birch leaves, and perhaps to this belongs the conidial-form, Fusicladium pirinum, which causes the “Rust spots” on Apples and Pears.

Order 11. Massariaceæ.

Order 12. Clypeosphæriaceæ.

Order 13. Gnomoniaceæ. Perithecia, with peak-like aperture. Gnomonia erythrostoma in the leaves of Prunus avium, which turn brown and do not fall in autumn.

Orders 14–18. Stroma generally well developed. The perithecia are embedded in the stroma, but when this is rudimentary, in the substratum.

Order 14. Valsaceæ. Valsa.

Order 15. Diatrypaceæ. Diatrype.

Order 16. Melanconidaceæ.

Order 17. Melogrammataceæ.

Order 18. Xylariaceæ. This order is the most highly developed of the Sphæriales. The stroma arises on the surface of the substratum, which is generally dead or decorticated wood; it is well-developed, crustaceous, hemispherical or upright. In the younger conditions it is covered with a layer of conidia, and later on it bears the perithecia, arranged in a layer immediately beneath its surface. The ascospores are of a dark colour. Often also there are free conidiophores.

Fig. 122.Xylaria hypoxylon (nat. size) on a tree stump: a younger, b an older stroma, both of which, with the exception of the black lower portion, are covered with white conidia; n, spot where the perithecia are developed; c an old stroma with upper part fallen off; d, e large branched stromata; k conidia.

Hypoxylon and Ustulina have a cushion-like or crustaceous stroma.—Xylaria has a club-shaped or branched stroma, often several centimetres high. X. hypoxylon (Fig. 122) and X. polymorpha occur on old tree stumps.—Poronia grows on old horse dung, and has a conical stroma.

Sub-Family 3. Dothideales.

The perithecia are always embedded in a black stroma, and are not distinctly separated from it. The accessory forms of reproduction are: conidiophores, conidiocarps, and yeast-like conidia. The majority are parasites. One order.

Order Dothideaceæ. Phyllachora graminis produces scab-like patches on the leaves of the Grasses.—Scirrhia rimosa grows on the leaf-sheathes of Phragmites.—Rhopographus pteridis on Pteridium aquilinum.

Family 4. Hysteriales.

This family, like the following, has hemiangiocarpic ascocarps (apothecia). These are closed in the early stages, but when ripe open in a valvular manner by a longitudinal fissure; they are black, oblong, and often twisted. Some species are parasites, especially upon the Coniferæ.

Fig. 123.Lophodermium (Hypoderma) nervisequium: a two leaves of Abies alba seen from above with pycnidia; b a leaf seen from the underside with apothecia; c an ascus with ascospores. (× 500.)

Fig. 124.—Three leaves of the Red-pine with Lophodermium macrosporum: a under side of the leaves with apothecia; b a leaf from upper side with pycnidia. (× about 2.)

Fig. 125.Lophodermium pinastri: a leaves of Pinus sylvestris with apothecia (nat. size); b two paraphyses and an ascus with filamentous spores.

Order 1. Hysteriaceæ. Hysterium pulicare upon the ruptured bark of many trees.

Order 2. Hypodermaceæ. The species of Lophodermium live upon the leaves of Conifers, and are the cause of their falling off (blight). L. pinastri (Fig. 125), on the leaves of Pinus and Picea; the leaves become red-brown and fall off; at first conidiocarps are formed, and later apothecia; L. nervisequium (Fig. 123), on Abies alba; L. macrosporum (Fig. 124), on Picea excelsa; L. brachysporum, on Pinus strobus.

Order 3. Dichænaceæ.

Order 4. Acrospermaceæ.

Family 5. Discomycetes.

The ascocarps (apothecia) are at first closed, and only open at the time of their ripening, not valvularly, but more or less like a saucer or cup, so that the hymenium lies exposed on their upper surface. In the first three sub-families, and generally also in the fourth, the apothecia are formed inside the substratum. The apothecia are, in contrast to the Pyrenomycetes, light and brightly coloured, and their size varies very much, and may be several centimetres in diameter. Paraphyses are often present between the asci; they often contain colouring matter, and give to the disc its characteristic colour. The tissue on which the asci are borne is known as the hypothecium. The shape and colour of the spores is not so varied as in the Pyrenomycetes. The accessory forms of reproduction are conidia (sometimes of two forms), chlamydospores, and oidia. The family is divided into 5 sub-families.

Sub-Family 1. Phacidiales.

The apothecia are developed in the interior of the substratum, which they break through, and in general dehisce apically. The envelope is tough and black. Hypothecium inconspicuous; hymenium flat.

Order 1. Euphacidiaceæ. Phacidium abietinum, on the leaves of Abies alba.—Rhytisma; the pycnidia are found in the summer on the green leaves, while the apothecia are developed on the fallen leaves and dehisce in the following spring. R. acerinum causes black spots on the leaves of the Sycamore, and R. salicinum on Willows.

Order 2. Pseudophacidiaceæ.

Sub-Family 2. Stictidales.

The apothecia when ripe break through the substratum which forms a border round them. Hymenium generally saucer-shaped.

Order 1. Stictidaceæ. Stictis.

Order 2. Ostropaceæ. Ostropa.

Sub-Family 3. Tryblidiales.

The apothecia are embedded in the substratum in the early stages, and then are raised high above it. Hypothecium thick. Hymenium cup-shaped.

Order 1. Tryblidiaceæ. Tryblidium.

Order 2. Heterosphæriaceæ. Heterosphæria patella on the dead stalks of Umbellifers.

Sub-Family 4. Dermateales.

The apothecia in the early stages are embedded in the substratum and then break through it, or are from the first situated on the surface of the substratum. Hypothecium thick.

Order 1. Cenangiaceæ. Cenangium.

Order 2. Dermateaceæ. Dermatea.

Order 3. Patellariaceæ. Patellea, Biatorella, Patellaria.

Order 4. Caliciaceæ. Calicium, Coniocybe, etc., on the bark of trees.

Order 5. Arthoniaceæ. Arthonia on the bark of several trees. Celidium stictarum on the apothecia of Sticta pulmonaria.

Order 6. Bulgariaceæ. Apothecia gelatinous under moist conditions, and horny when dried.—Calloria fusarioides; the red apothecia break out in the spring on the dried stalks of Urtica dioica; a gelatinous reproductive form of the Fungus is found before the apothecia, which consists of oidia (formerly described as “Dacryomyces urticæ”).—Bulgaria inquinans on the living or fallen trucks of Oaks and Beeches.

Fig. 126.Botrytis cinerea: a slightly magnified; b more highly magnified; c germinating conidium.

Fig. 127.Sclerotinia fuckeliania: a sclerotium with conidiophores; b with apothecia; c section through sclerotium and apothecium; d ascus with eight ascospores. (× 390.)

Sub-Family 5. Pezizales.

The apothecia are developed on the surface of the substratum and are waxy or fleshy; at the commencement closed, and covered with a saucer- or cup-shaped, seldom flat, hymenium. The hypothecium is generally well developed. This sub-family is the richest in species of the Discomycetes and contains forms of very different habit. They grow upon dead wood, upon the ground, and upon dung. A few are parasites.

Order 1. Helotiaceæ. Apothecia with waxy envelope of colourless, or yellowish prosenchymatous cells.—Chlorosplenium æruginosum is found on decaying wood (particularly Oak and Birch), to which it gives a green colour. Sclerotinia has sclerotia which are developed upon the host-plant and from which, after a period of rest, the long, brown-stalked apothecia arise. S. ciborioides (S. trifoliorum, Fig. 128) is parasitic on Clover; S. sclerotiorum, on Daucus-roots, Phaseolus, etc.; S. baccarum, on the berries of Vaccinium myrtillus; “Botrytis cinerea” is a common parasite and is probably the conidial form of S. fuckeliania (Fig. 127).—Helotium herbarum lives on dry plant stems.—Dasyscypha willkommii (Fig. 129) produces Larch-canker on the bark of the Larch.

Fig. 128.Sclerotinia ciborioides: a sclerotium with three apothecia slightly magnified; b ascus with eight ascospores; c germinating ascospore.

Fig. 129.Dasyscypha willkommii: a portion of bark of Larix decidua with sessile, cup-shaped apothecia (nat. size); b two paraphyses on either side of an ascus with eight ascospores.

Order 2. Mollisiaceæ. Mollisia cinerea, principally on decaying wood.

Order 3. Pezizaceæ. This order contains the largest and morphologically the highest forms of the Discomycetes. Apothecia fleshy, and in the later conditions generally saucer-shaped.

Peziza, with sessile apothecia, growing on the ground; P. cochleata is brown, and coiled like a snail-shell; P. coccinea is scarlet; P. aurantia occurs as an orange-coloured expansion on the ground.

Order 4. Ascobolaceæ. Apothecia fleshy; in the later stages flat or convex. The asci are, comparatively speaking, large, and often contain a great number of spores which escape by the casting off of a lid on the summit of the ascus. Generally living on dung.—Ascobolus furfuraceus, etc.

Family 6. Helvellales.

These Fungi have the appearance of clubs, bells, or mushrooms, consisting of an upright stalk bearing a large and fleshy head, on the exterior surface of which the hymenium is spread. The ascocarps are probably gymnocarpic from the beginning, and on this account these plants are placed in a separate family. The development of the ascocarps is unknown. The Morchella (Morell) grows on the ground; some species are edible. 1 order.

Order. Helvellaceæ. Spathulea is yellow and club-shaped, and forms “fairy rings” in woods.—Geoglossum (Earth-tongue) projects above the ground as a black tongue, or as a club-shaped body. Several species are found in meadows and on heaths.—Helvella has a stalk, bearing an irregularly folded head, on the external surface of which is the hypothecium.—Morchella (Morell, Fig. 130), the stalk bears on its summit the conical or spherical head, the external surface of which is reticulate and bears the asci.—Mitrula. Verpa.

Fig. 130.Morchella esculenta: a an entire specimen, about one half natural size; b longitudinal section through the head.

Appendix to the Ascomycetes:

Family 7. Ascolichenes (Lichen-forming Ascomycetes).

The Lichens were formerly classed among the Thallophyta as a group quite distinct from the Algæ and Fungi. Investigations during the last twenty-five years, however, have conclusively proved that the Lichens are Fungi which reproduce in the same manner as the Ascomycetes, or, more rarely, the Basidiomycetes, and have entered into a peculiar symbiotic relation with Algæ, especially the Cyanophyceæ and Protococcoideæ, with which they associate, and without which they would be unable to exist. The Fungus forms the largest portion of the Lichen, enclosing the Alga with which it may be said to be commensal. The Fungus especially produces reproductive bodies and absorbs the inorganic nourishment through the rhizoids, whilst the Alga supplies it with the organic materials. In consequence of this the Lichens, in contradistinction to other Fungi, need light for the development of their nutritive organs, and are therefore, in any case internally, of a more or less greenish colour. The form and condition of the thallus is unusual among the Fungi, and they can grow upon rocks and in other places where no dead organic matter, such as would be required by other Fungi, is obtainable.

Fig. 131.—Transverse section through the thallus of Sticta fuliginosa (× 500): r-r rhizoid-strands, which arise from the under side; g-g gonidial layer; m medullary layer; o upper, u lower cortex.

Two cellular forms are therefore to be found in each Lichen:

1. The cells which belong to the Fungus. These are generally septate, branched hyphæ without any trace of chlorophyll. In the thallus of the majority of Lichens there may be found a medullary layer (Fig. 131 m) of loosely-woven hyphæ, between which there are large air chambers; and an external layer (cortex) (Fig. 131 o, u) formed of closely-woven hyphæ without any intercellular spaces. In some Lichens (Collemaceæ) the hyphæ wind about in the thallus, being equally distributed throughout, without forming any decided strata. These Lichens moreover become gelatinous when exposed to moisture (Fig. 132), on account of the swelling of the walls of the Algæ. The hyphæ contain protoplasm with drops of oil, but never starch; their walls easily swell when exposed to damp after having been dried, and in some (e.g. Cetraria islandica) they become gelatinous when cooked. Certain strata of hyphæ become blue on treatment with iodine alone, from which it is inferred that the wall is allied, in its chemical nature, to starch.

2. The enclosed Algæ, termed “gonidia.” Some belonging to the Cyanophyceæ, Protococcoideæ, (especially Pleurococcus) and Chroococcaceæ, are spherical and are found isolated, or in irregular groups of cells (Fig. 131 g); some belonging to Nostoc (Fig. 132 g), Lyngbyaceæ, etc., are placed in cell-rows. Each Lichen, as a rule, has only one definite Algal-form for its gonidium.

The gonidia either lie together in a certain stratum between the cortex and the medullary layer (Fig. 131 g), or are scattered irregularly throughout the entire thallus (Fig. 132). The thallus is in the first instance termed “heteromerous,” in the second instance, “homoiomerous.” The Fungal-hyphæ embrace the gonidia and apply themselves closely to, or even penetrate them, and hence it has been difficult to decide whether the one cellular form does or does not develop from the other (Figs. 134, 135).

Fig. 132.Collema microphyllum. Transverse section through the thallus; g Nostoc-chains; h hyphæ.

This theory regarding the symbiosis of Fungi and Algæ to form a Lichen is termed the Schwendenerian theory, after the first scientist who advanced it with any weight. It had been already indicated by De Bary, and further arguments in its support have at a later time been adduced by Bornet, Stahl, Treub, Frank, Bonnier, Alfr. Möller and others.

Fig. 133.Ephebe pubescens. The apex of a branch of the thallus with two lateral branches (s): h its hyphæ; g the apical gonidium of the main branch.

Fig. 134.Nostoc lichenoides, which is attached by a germinating thread (h) of Collema glaucescens.

Fig. 135.A Germinating spore of Physcia parietina with Protococcus viridis. B Synalissa symphorea with Glæocapsa. C Cladonia furcata with Protococcus.

The thallus of the Lichen appears mainly under three forms:—

1. The Crustaceous, which adheres firmly to the substratum (bark, stone) throughout its entire surface, without being raised into any free patches or lobes. It has, in many instances, no definite outline, and hyphal-branches from it often penetrate deeply into the substratum. It grows at the circumference and sometimes dies away in the centre (Figs. 138, 139, 140).

2. The Foliaceous. This also lies flat upon the substratum, but is not firmly attached to and has a definite outline. It grows at the margin, and raises itself a little by free outgrowths and lobes (Fig. 141). The rhizoid-strands spring out from its whitish under surface (Fig. 131, r).

Fig. 136.—Portion of a hymenium: d a thin stratum on which the asci (s) are situated.

Fig. 137.—Spores of, a Cladonia, Lecanora and Pertusaria; b Bæomyces; c Sphinctrina; d, e, f various species of Parmelia; g, h Verrucaria in its younger and older condition; i, k species of Leptogium.

3. The Fruticose, which is attached to its substratum at a small point from which it projects freely, either erect or pendulous. It is more or less tufted, in the form of a bush (Figs. 142, 143). These three thallus-forms gradually pass over by many intermediate forms into one another.