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(J press.) XV. FUNGI; tHeir Nature, INFLUENCE, AND Uses. By M. C. Cooxr, M.A., LL.D. Edited by Rev. M. J. BerKELey, M. A., F.L.S. With og Illustrations. (Jz press.) XVI. OPTICS. By Proressor LommeEL, University of Erlangen. (/# press.) THE INTERNATIONAL SCIENTIFIC SERIES. Ho NGL: THEIR RATUBREH AND Vers. BY M. C. COOKE, M. A., LL.D. EDITED BY Tar Ruv. M. J. BERKELEY, M.A., F.L.S. 2A NEW YORK: D. APPLETON AND COMPANY, 549 AND 551 BROADWAY. 1875. PREFACE BY THE EDITOR. As my name appears on the title-page of this volume, it is necessary that I should exactly state what part I had in its preparation, I had no doubt originally engaged to undertake the work myself; but finding, from multiplicity of engagements and my uncertain health, that I could not accomplish it satis- factorily, I thought the best course I could take was to recom- mend Mr. Cooke to the publishers; a gentleman well known, not only in this country, but in the United States. The whole of the work has therefore been prepared by himself, the manu- script and proof sheets being submitted to me from time to time, in which I merely suggested such additions as seemed needful, subjoining occasionally a few notes. As the work is intended for students, the author has had no hesitation in vi PREFACE. repeating what has been stated in former chapters where it has been thought to prove useful. I have no doubt that the same high character will justly apply to this as to Mr. Cooke’s former publications, and especially to his ‘“ Handbook of British Fungi.” M. J. BERKELEY. SipbeRrort, November 23, 1874. CONTENTS. camer ecie I, PAGE Nature or Fonst a ; é ‘ . A . e e Il. ScrvucrurE . fe $ . : . ° e e ° » IF HI. CLASSIFICATION « ‘ . . . ‘ * . . > » 64 e IV. Uses . ° ° . - . . ° . . e « 82 Vv. Notasie PHENOMENA . . . 7 ; . ‘ . . - 105 VI. 7 Tas Spore anp 1Ts DIssEMINATION : 2 . r 5 . 119 vili GERMINATION AND GROWTH . Sexual REPRODUCTION . a PoLy MORPHISM * . . InFLUENCES AND ErFects . Hasitats . . . . CULTIVATION » . . GxrocRAPHICAL DisTRIBUTION . CoLLECTION AND PRESERVATION CONTENTS. VIII. . 163 . 209 LIST OF ILLUSTRATIONS. ae ae FIG 1. Agaric in process of growth . . A . . . 2. Scetion of common mushroom . . . . 176 101. Tilletia caries with conjugating cells . : . 2 ‘ - 178 102. Aspergillus glaucus and Eurotium 2 . . . 3 - 189 108. Erysiphe cichoraccarum, receptacle and mycelium . 7 . . 191 104. Twig with Tubcrcularia and Nectria . 7 . . . . . 193 105. Section of Tubercularia with conidia . 7 a s . . 194 106. Nectria with Tubercularia, ascus and paraphyses . 5 a » 195 107. Cells and pseudospores of Mcidium Berberidis ‘ é . - 201 108. Cells and pseudospores of Acidium graveolens . . . + 201 109. Torrubia militaris on pupa of amoth , . . . i » 243 FUNGI THEIR NATURE, USES, INFLUENCES, ETC. —1— I, NATURE OF FUNGI. 2 Tue most casual observer of Nature recognizes in almost every instance that comes under his notice in every-day life, without the aid of logical definition, the broad distinctions between an animal, a plant, and a stone. To him, the old definition that an animal is possessed of life and locomotion, a plant of life with- out locomotion, and a mineral deficient in both, seems to be sufficient, until some day he travels beyond the circuit of diurnal routine, and encounters a sponge or a zoophyte, which possesses only one of his supposed attributes of animal life, but which he is assured is nevertheless a member of the animal kingdom. Such an encounter usually perplexes the neophyte at first, but rather than confess his generalizations to have been too gross, he will tenaciously contend that the sponge must be a plant, until the evidence produced is so strong that he is compelled to desert his position, and seek refuge in the declaration that one kingdom runs into the other so imper- ceptibly that no line of demarcation can be drawn between them. Between these two extremes of broad distinction, and no distinction, lies the ground occupied by the scientific student, who, whilst admitting that logical definition fails in assigning briefly and tersely the bounds of the three kingdoms, contends 4 2 FUNGI. that such limits exist so positively, that the universal scientific mind accepts the recognized limit without controversy or con- tradiction. In like manner, if one kingdom be made the subject of in- quiry, the ‘same difficulties will arise. A flowering plant, as represented by a rose or a lily, will be recognized as distinct from a fern, a seaweed, or a fungus. Yet there are some flower- ing plants which, at first sight, and without examination, simu- late cryptogams, as, for example, many Balanophore, which the unscientific would at once class with fungi. It is never- theless true that even the incipient botanist will accurately separate the phanerogams from the cryptogams, and by means of alittle more, but still elementary knowledge, distribute the ‘latter amongst ferns, mosses, fungi, lichens, and alge, with comparatively few exceptions. It is true that bétween fungi and lichens there exists so close an affinity that difficulties arise, and doubts, and disputations, regarding certain small groups or a few species; but these are the exception, and not the rule. Botanists generally are agreed in recognizing the five principal groups of Cryptogamia, as natural and distinct. In proportion as we advance from comparison of members of the three king- doms, through that of the primary groups in one kingdom, to a comparison of tribes, alliances, and orders, we shall require closer observation, and more and more education of the eye to see, and the mind to appreciate, relationships and distinctions. We have already assumed that fungi are duly and universally admitted, as plants, into the vegetable kingdom. But of this fact some have even ventured to doubt. This doubt, however, has been confined to one order of fungi, except, perhaps, amongst the most illiterate, although now the animal nature of the Myxogastres has scarcely a serious advocate left. In this order the early condition of the plant is pulpy and gelatinous, and consists of a substance more allied to sarcode than cellulose. De Bary insinuated affinities with Ameta,* whilst Tulasne * De Bary, ‘‘Des Myxomycétes,” in ‘ Ann. des Sci. Nat.” ‘4 sér, xi. p. 153; ‘*Bot. Zeit.” xvi. p. 357. De Bary’s views are controverted by M. Wigand.in ‘Ann, des Sci. Nat.” 4 sér. (Bot.) xvi. p, 255, &c. NATURE OF FUNGI. 5 affirmed that the outer coat in some of these productions con- tained so much carbonate of lime that strong effervescence took place on the application of sulphuric acid. Dr. Henry Carter is ‘well known as an old and experienced worker amongst amoeboid forms of animal life, and, when in Bombay, he devoted himself to the examination of the Myzxogastres in their early stage, and the result of his examinations has been a firm conviction that there is no relationship whatever between the Myxogastres and the lower forms of animal life. De Bary has himself very much modified, if not wholly abandoned, the views once propounded by him on this subject. When mature, and the dusty spores, mixed with threads, sometimes spiral, are produced, the Myw«ogastres are so evidently close allies of the Lycoperdons, or Puffballs, as to leave no doubt of their affinities, It is scarcely necessary to remark that the presence of zoospores is no proof of animal nature, for not only do they occur in the white rust (Cystopus), and in such moulds as Peronospora,* but are common in alge, the vegetable nature of which has never been disputed. There is another equally important, but more complicated subject to which we must allude in this connection. This is the probability of minute fungi being developed without the ‘intervention of germs, from certain solutions. The observations of M. Trécul, in a paper laid before the French Academy, have thus been summarized:—1l. Yeast cells may be formed in the must of beer without spores being previously sown. 2. Cells of the same form as those of yeast, but with different contents, arise spontaneously in simple solution of sugar, or to which a little tartrate of ammonia has been added, and these cells are capable of producing fermentation in certain liquids under favourable conditions. 38. The cells thus formed produce Peni- cillium like the cells of yeast. 4; On the other hand, the spores of Penicillium are capable of being transformed into yeast. + The interpretation of this is, that the mould Penicillium may be "* De Bary, ‘Recherches sur le Developpement de quelques Champiguons Parasites,” in ‘‘ Ann. des Sci. Nat.” 4 sér. (Bot.) xx. p. 5, + ‘Popular Science Review,” vol. viii. p. 96. 4 FUNGI. - produced from a sugar solution by “spontaneous generation,” and without spore or germ of any kind. The theory is, that a molecular mass which is developed in certain solutions or infu- sions, may, under the influence of different circumstances, pro- duce either animalcules or fungi. “In all these cases, no kind of animalcule or fungus is ever seen to originate from pre- existing cells or larger bodies, but always from molecules.” * The molecules are said to form small masses, which soon melt together to constitute a globular body, from which a process juts out on one side. These are the so-called Torwle,t which give off buds which are soon transformed into jointed tubes of various diameters, terminating in rows of sporules, Penicil- lium, or capsules containing numerous globular seeds, Aspergil- lus (sic). This is but another mode of stating the same thing as above referred to by M. Trécul, that certain cells, resembling yeast cells (Zorula), are developed spontaneously, and that these ultimately pass through the form of mould called Penicillium to the more complex Mucor (which the writer evidently has confounded with Aspergillus, unless he alludes to the ascigerous form of Asper- gillus, long known as Eurotium). From what is now known of the polymorphism of fungi, there would be litfle difficulty in believing that cells resembling yeast cells would develop: into Penicillium, as they do in fact in what is called the “ vine- gar plant,” and that the capsuliferous, or higher condition of this mould may be a Mucor, in which the sporules are produced in capsules. The difficulty arises earlier, in the supposed spon- taneous origination of yeast cells from molecules, which result from the peculiar conditions of light, temperature, c&c., in which certain solutions are placed. It would be impossible to review all the arguments, or tabulate all the experiments, which have been employed for and against this theory. It could not be passed over in silence, since it has been one of the stirring ques- tions of the day. The great problem how to exclude all germs * Dr. J. H. Bennett ‘‘ On the Molecular Origin of Infusoria,” p. 56. + They have, however, no close relation with real Torule, such as T. mont- lioides, &c,—Cooxe’s Handbook, p: 477. NATURE OF FUNGI. 5 from the solutions experimented upon, and to keep them ex- cluded, lies at the foundation of the theory. It must ever, as we think, be matter of doubt that all germs were not excluded or destroyed, rather than one of belief that forms known to be developed day by day from germs should under other conditions originate spontaneously. Fungi are veritably and unmistakably plants, of a low or- ganization, it is true, but still plants, developed from germs, - somewhat analogous, but not wholly homologous, to the seeds of higher orders. The process of fertilization is still obscure, but facts are slowly and gradually accumulating, so that we may hope at some not very distant period to comprehend what as yet are little removed from hypotheses. Admitting that fungi are independent plants, much more complex in their relations and development than was formerly ‘supposed, it will be ex- pected that certain forms should be comparatively permanent, that is, that they should constitute good species. Here, also, efforts have been made to develop a theory that there are no legitimate species amongst fungi, accepting the terms as hither- to applied to flowering plants. In this, as in allied instances, too hasty generalizations have been based on a few isolated facts, without due comprehension of the true interpretation of such facts and phenomena. Polymorphism will hereafter receive special illustration, but meantime it may be well to state that, be- cause some forms of fungi which have been described, and which have borne distinct names as autonomous species, are now proved to be only stages or conditions of other species, there is no reason for concluding that no forms are autonomous, or that fungi which appear and are developed in successive stages are not, in their entirety, good species. Instead, therefore, of insinuating that there are no good species, modern investigation tends rather to the establishment of good species, and the elimination of those that are spurious. It is chiefly amongst the microscopic species that polymorphism has been determined. In the larger and fleshy fungi nothing has-been discovered which can shake our faith in the species described half a century, or more, ago. In the Agarics, for instance, the forms seem to be as permanent and 6 FUNCI. as distinct as in the flowering plants. In fact, there is still no reason to dissent, except to a very limited extent, from what was written before polymorphism was accredited, that, “with a few exceptions only, it may without doubt be asserted that more certain species do not exist in any part of the organized world than amongst fungi. The same species constantly recur in the same places, and if kinds not ‘hitherto detected present them- selves, they are either such as are well known in other districts, or species which have been overlooked, and which are found on better experience to be widely diffused. There is nothing like chance about their characters or growth.” * The parasitism of numerous minute species on living and growing plants has its parallel even amongst phanerogams in the mistletoe and broom-rape and similar species. Amongst fungi a large number are thus parasitic, distorting, and in many. cases ultimately destroying, their host, burrowing within the tissues, and causing rust and smut in corn and grasses, or even more destructive and injurious in such moulds as those of the potato disease and its allies. A still larger number of fungi are developed from decayed or decaying vegetable matter. These are found in winter on dead leaves, twigs, branches, rotten wood, the remains of herbaceous plants, and soil largely charged with disintegrated vegetables. As soon as a plant begins to decay it becomes the source of a new vegetation, which hastens its destruction, and a new cycle of life com- mences. In these instances, whether parasitic on living plants or developed on dead ones, the source is still vegetable. But this is not always the case, so that it cannot be predicated that fungi are wholly epiphytal. Some species are always found on animal matter, leather, horn, bone, &c., and some affect such unpromising substances as minerals, from which it would be supposed that no nourishment could be obtained, not only hard gravel stones, fragments of rock, but also metals, such as iron and lead, of which more may be said when we come to treat of the habitats of fungi. Although in general terms fungi may be deséribed as “hysterophytal or epiphytal mycetals deriving * Berkelcy’s ‘ Outlines of British Fungology,” p. 24. NATURE OF FUNGI. 7 nourishment by means of a mycelium from the matrix,’ * there are exceptions to this rule with which the majority accord. Of the fungi found on animal substances, none are more extraordinary than those species which attack insects. The white mould which in autumn proves so destructive to the common house-fly may for the present be omitted, as it is probably a condition of one of the Saprolegnici, which some authors include with fungi, and others with alge. Wasps, spiders, moths, and butterflies become enveloped in a kind of mould named Isaria, which constitutes the conidia of Torrubia, a genus of club-shaped Spherie afterwards developed. Some species of Isaria and Torrubia also affect the larvse and pup» of moths and butterflies, converting the whole interior into a mass of mycelium, and fructifying ina clavate head. It has been subject for discussion whether in such instances the fungus commenced its development during the life of the in- sect, and thus hastened its death, or whether it resulted after death, and was subsequent to the commencement of decay.t The position in which certain large moths are found standing on leaves when infested with Jsaria resembles so closely that of the house-fly when succumbing to Sporendonema Musce, would lead to the conclusion that certainly in some cases the insect was attacked by the fungus whilst still living; whilst in the case of buried caterpillars, such as the New Zealand or British Hepialus, it is difficult to decide. Whether in life or death in these instances, it is clear that the silk-worm diseasc Muscardine attacks the living insect, and causes death. In the case of the Guépes vé;étantes, the wasp is said to fly about with the fungus partially developed. In all fungi we may recognize a vegetative and a reproductive system : sometimes the first only becomes developed, and then the fungus is imperfect, and sometimes the latter is far more prominent than the former. There is usually an agglomeration of delicate threads, either jointed or not, which are somewhat analogous to the roots of higher plants. These delicate threads * Berkeley's ‘(Introduction to Cryptogamic Botany,” p. 235. + Gray, ‘‘ Notices of Insects which form the Basis of Fungoid Parasites.” 8 FUNGI. permeate the tissues of plants attacked by parasitic fungi, or they run over dead leaves forming whitened patches, formerly bearing the name of Himantia, but really the mycelium of some species of Marasmius. If checked or disturbed, the process stops here, and only a mycelium of interwoven threads is produced. In this condition the mycelium of one species so much resembles that of another, that no accurate determination can be made. If the process goes on, this mycelium gives rise to the stem and cap of an agaricoid fungus, completing the vegetative system. This in turn gives origin to a spore-bearing surface, and ultimately the fruit is formed, and then the fungus is complete ; no fungus can be regarded as perfect or complete without its reproductive system being developed. In some this is very simple, in others it is as complex. In many of the moulds we have miniature representatives of higher plants in the mycelium or roots, stem, branches, and at length capsules bearing sporidia, which correspond to seeds. It is true that leaves are absent, but these are sometimes compensated by lateral processes or abortive branchlets. A tuft of mould is in miniature a forest of. trees. Although such a definition may be deemed more poetic than accurate, more figurative than literal, yet few could believe in the marvellous beauty of a tuft of mould if they never saw it as exhibited under the microscope. In sucha condition no doubt could be entertained of its vegetable character. But there is a lower phase in which these plants are sometimes encountered ; they may consist only of single cells, or strings of cells, or threads of simple structure floating in fluids. In such conditions only the vegetative system is probably developed, and that imperfectly, yet some have ventured to give names to isolated cells, or strings of cells, or threads of mycelium, which really in them- selves possess none of the elements of correct classification—the vegetative system, even, being imperfect, and consequently the reproductive is absent. As already observed, no fungus is per- fect without fruit of some kind, and the peculiarities of structure and development of fruit form one of the most important elements in classification. To attempt, therefore, to give names to such imperfect fragments of undeveloped plants is almost as absurd * NATURE OF FUNGI. 9 as to name a flowering plant froma stray fragment of a root- fibril accidentally cast out of the ground—nay, even worse, for identification would probably be easier. It is well to protest at all times against attempts to push science to the verge of absurdity; and such must be the verdict upon endeavours to determine positively such incomplete organisms as floating cells, or hyaline threads which may belong to any one of fifty species of moulds, or after all to an alga. This leads us to remark, in passing, that there are forms and conditions under which fungi may be found when, fructification being absent—that is, the vegetative system alone developed—they approximate so closely to algie that it is almost impossible to aay to which group the organisms belong. Finally, it is a great characteristic of fungi in general that they are very rapid in growth, and rapid in decay. Ina night a puffball will grow prodigiously, and in the same short period a mass of paste may be covered with mould. Ina few hours a gelatinous mass of Reticularia will pass into a bladder of dust, or a Coprinus will be dripping into decay. Remembering this, mycophagists will take note that a fleshy fungus which may be good eating at noon may undergo such changes in a few hours as to be anything but good eating at night. Many instances have been recorded of the rapidity of growth in fungi; it may also be accepted as an axiom that they are, in many instances, equally as rapid in decay. The affinity between lichens and fungi has long been re- cognized to its full and legitimate extent by lichenologists and mycologists.* In the “ Introduction to Cryptogamic Botany,” it * On the relation’ or connection between fungi and lichens, H. C. Sorby has some pertinent remarks in his communication to the Royal Society on ‘Comparative Vegetable Chromatology” (Proceedings Royal Society, vol. xxi. 1873, p. 479), as one result of his spectroscopic examinations. He says, **Such being the relations between the organs of reproduction and the foliage, it is to some extent possible to understand the connection between parasitic plants like fungi, which do not derive their support from the constructive energy of their fronds, and those which are self-supporting and possess true fronds. In the highest classes of plants the flowers are connected with the leaves,’ more especially by means of xanthophyll and yellow xanthophyll, 10 FUNGI. was proposed to unite them in one alliance, under the name of Mycetales, in the same manner as the late Dr. Lindley had united allied orders under alliances in his “ Vegetable Kingdom ;”’ but, beyond this, there was no predisposition towards the theory since propounded, and which, like all new théories, has collected a small but zealous circle of adherents. It will be necessary briefly to summarize this theory and the arguments by which it is supported and opposed, inasmuch as it is intimately connected with our subject. As recently as 1868, Professor Schwendener first propounded bis views,* and then briefly and vaguely, that all and every individual lichen was but an algal, which had collected about it a parasitic fungal growth, and that those peculiar bodies which, under the name of gonidia, were considered as special organs of lichens, were only imprisoned algw. In language which the Rev. J. M. Crombie + describes as “ pictorial,” this author gave the general conclusion at which he had arrived, as follows :— “As the result of my researches, all these growths are not simple plants, not individuals in the usual sense of the term; they are rather colonies, which consist of hundreds and thousands of individuals, of which, however, only one acts as master, while the others, in perpetual captivity, provide nourishment for them- selves and their master. This master is a fungus of the order Ascomycetes, a parasite which is accustomed to live upon the work of others; its slaves are green algw, which it has sought out, or indeed caught hold of, and forced into its service. It surrounds whereas in the case of lichens the apothecia contain very little, if any, of those substances, but a large amount of the lichenoxanthines so characteristic of the class. Looking upon fungi from this chromatological point of view, they bear something like the same relation to lichens that the petals of a leafless parasitic plant would bear to the foliage of one of normal'character—that is to say, they are, as it were, the coloured organs of reproduction of parasitic plants of a type closely approaching that of lichens, which, of course, is in very close, if not in absolute agreement with the conclusions drawn by botanists from entirely different data.” : * Schwendener, ‘‘ Untersuchungen wither den Flechtenthallus.”’ + Crombie (J. M.) ‘On the Lichen-Gonidia Question,” in ‘‘ Popular Science Review” for July, 1874. NATURE Ol’ FUNGI. 11 them, as a spider does its prey, with a fibrous net of narrow meshes, which is gradually converted into an impenetrable covering. While, however, the spider sucks its prey and leaves it lying dead, the fungus incites the alge taken in its net to more rapid activity; nay, to more vigorous increase.” This hypothesis, ushered upon the world with all the prestige of the Professor’s name, was not long in meeting with adherents, and the cardinal. points insisted upon were—Ist. That the generic relationship of the coloured “ gonidia”’ to the colourless fila- ments which compose the lichen thallus, had only been assumed, and not proved; 2nd. That the membrane of the gonidia was chemically different from the membrane of the other tissues, inasmuch as the first had a reaction corresponding to that. ot alge, whilst the second had that of fungi; 3rd. That the different forms and varieties of gonidia corresponded with parallel types of alge; 4th. That as the germination of the spore had not been followed further than the development of a hypothallus, it might be accounted for by the absence of the essential algal on which the new organism should become para- sitic; 5th. That there is a striking correspondence between the development of the fruit in lichens and in some of the sporidii- ferous fungi (Pyrenomycetes). These five points have been combated incessantly by lichen- ologists, who would really be supposed by ordinary minds to be the most practically acquainted with the structure and develop- ment of these plants, in opposition to the theorists. It is a fact which should have some weight, that no lichenologist of repute has as yet accepted the theory. In 1873 Dr. E. Bornet* came to the aid of Schwendener, and almost exhausted the subject, but failed to convince either the practised lichenologist or mycologist. The two great points sought to be established are these, that what we call lichens are compound organisms, not simple, independent vegetable entities ; and that this compound organism consists of unicellular alge, with a fungus parasitic upon them. ‘he coloured gonidia which are found in the * Bornet, (E.), ‘‘ Recherches sur les Gonidies des Lichens,” in ‘‘ Ann. des Sci. Nat.” 1873, 5 sér. vol. xvii. 12 FUNGI. substance, or thallus of lichens, are the supposed alge ; and the cellular structure which surrounds, encloses, and imprisons the gonidia is the parasitic fungus, which is parasitic on something infinitely smaller than itself, and which it entirely and absolutely isolates from all external influences. Dr. Bornet believed himself to have established that every gonidium of a lichen may be referred to a species of alge, and that the connection between the hypha and gonidia is of such a. nature as to exclude all possibility of the one organ being pro- duced by the other. This he thinks is the only way in which it can be accounted for that the gonidia of diverse lichens should be almost identical. ; Dr. Nylander, in referring to this hypothesis of an imprisoned algal,* writes: “The absurdity of such an hypothesis is evident from the very consideration that it cannot be the case that an organ (gonidia) should at the same time be a parasite on the body of which it exercises vital functions; for with equal propriety it might be contended that the liver or the spleen constitutes parasites of the mammifere. Parasite existence is autonomous, living upon a foreign body, of which nature prohibits it from being at the same time an organ. This is an elementary axiom of general physiology. But observation directly made teaches that the green matter originally arises within the primary chlorophyll- or phycochrom-bearing cellule, and consequently is not intruded from any external quarter, nor arises in any way from any parasitism of anykind. The cellule at first is observed to be empty, and then, by the aid of secretion, green matter is gradually produced in the cavity and assumes a definite form. It can, therefore, be very easily and evidently demonstrated that the origin of green matter in lichens is en- tirely the same as in other plants.” On another occasion, and in another place, the same eminent lichenologist remarks,t as to the supposed algoid nature of gonidia—‘ that such an unnatural existence as they would thus pass, enclosed in a prison and * Nylander, ‘*On the Algo-Lichen Hypothesis,” &., in ‘‘Grevillea,” vol. ii. (1874), No. 22, p. 146. + In Regensburg ‘‘ Flora,” 1870, p. 92. NATURE OF FUNGI. 138 deprived of all autonomous liberty, is not at all consonant with the manner of existence of the other algw, and that it has no parallel in nature, for nothing physiologically analogous occurs anywhere else. Krempelhuber has argued that there are no conclusive reasons against the assumption that the lichen-gonidia may be self-developed organs of the lichen proper rather than alge, and that these gonidia can continue to vegetate separately, and so be mistaken for unicellular alge.” In this Th, Fries seems substantially to concur. But there is one strong argu- ment, or rather a repetition of an argument already cited, placed in a much stronger light, which is employed by Nylander in the following words :—“ So far are what are called alge, according to the turbid hypothesis of Schwendener, from constituting true alge, that on the contrary it may be affirmed that they have a lichenose nature, whence it follows that these pseudo-algw are in a systematic arrangement to be referred rather to the lichens, and that the class of algze hitherto so vaguely limited should be circumscribed by new and truer limits. As to another phase in this question, there are, as Krempel- huber remarks, species of lichens which in many countries do not fructify, and whose propagation can only be carried on by means of the soredia, and the hyphee of such could in themselves alone no more serve for propagation than the hyphw from the pileus or stalk of an Agaric, while it is highly improbable that they could acquire this faculty by interposition of a foreign algal. On the other hand he argues: “It is much more con- formable to nature that the gonidia, as self-developed organs of the lichens, should, like the spores, enable the hyphe proceeding from them to propagate the individual.* A case in point has been adducedt in which gonidia were produced by the hypha, and the genus Hmericella,t which is allied to Husseia in the Trichogastres, shows a structure in tho stem exactly resembling Palmella botryoides of Greville, and to what occurs in Synalyssa. Emericella, with one or two other * Rev. J. M. Crombie, in ‘‘ Popular Science Review,” July, 1874. + Berkeley’s “ Introduction to Cryptogamic Botany,” p. 373, fig. 78a. + Berkeley’s “Introduction,” p. 341, fig. 76. 2 14 FUNGI. genera, must, however, be considered as connecting Trichogastres with lichens, and the question cannot be considered as satis- factorily decided till a series of experiments has been made on the germination of lichen spores and their relation to free alg considered identical with gonidia. Mr. Thwaites was the first to point out* the relation of the gonidia in the different sections of lichens to different types of supposed alge. The question cannot be settled by mere @ priori notions. It is, perhaps, worthy of remark that in Chionyphe Cartert the threads grow over the cysts exactly as the hypha of lichens is represented as growing over the gonidia. Recently, Dr. Thwaites has communicated his views on one phase of this controversy,f which will serve to illustrate the question as seen from the mycological side. As is well known, this writer has had considerable experience in the study of the anatomy and physiology of all the lower cryptogamia, and any suggestion of his on such a subject will at least commend itself to a patient consideration. “ According to our experience,” he writes, “I think parasitic fungi invariably produce a sad effect upon the tissues they fix themselves upon or in. These tissues become pale in colour, and in every respect sickly in appearance. But who has ever seen the gonidia of lichens the worse for having the ‘hypha’ growing amongst them? These gonidia are always in the plumpest state, and with the freshest, healthiest colour possible. Cannot it enter into the heads of these most patient and ex- cellent observers, that a cryptogamic plant may have two kinds of tissue growing side by side, without the necessity of one being parasitic upon the other, just as one of the higher plants may have half a dozen kinds of tissue making up its organiza- tion? The beautifully symmetrical growth of the same lichens has scemed to me a sufficient argument against one portion being parasitic upon another, but when we see all harmony and robust health, the idea that one portion is subsisting parasitically upon another appears to me to be a perfect absurdity.” * “Annals and Magazine of Natural History,” April, 1849. + In ‘Gardener's Chronicle” for 1873, p. 1341. NATURE OF FUNGI. 15 It appears to us that a great deal of confusion and a large number of errors which creep into our modern generalizations and hypotheses, may be traced to the acceptance of analogies for identities. How many cases of mistaken identity has the improvement of microscopes revealed during the past quarter of acentury. This should at least serve asa caution for the future. Apart, however, from the “ gonidia,” whatever they may be, is the remainder of the lichen a genuine fungus? Nylander writes, “‘The anatomical filamentose elements of lichens are distinguished by various characters from the hyphe of fungi. They are firmer, elastic, and at once present themselves in the texture of lichens. On the other hand, the hyphe of fungi are very soft, they possess a thin wall, and are not at all gelatinous, while they are immediately dissolved by the application of hrdrate of potash, &c.* Our own experience is soniewhat to the effect, that there are some few lichens which are doubtful as to whether they are fangi or lichens, but, in by far the majority of cases, there is not the slightest difficulty in determining, from the peculiar firmness and elasticity of the tissues, minute peculiarities which the practised hand can detect rather than describe, and even the general character of the fruit that they differ materially from, though closely allied to fungi. We have only experience to guide us in these matters, but that is something, and we have no experience in fungi of anything like a Cladonia, however much it may resemble a Zorrubia or Clavaria. We have Pezize with asubiculum in the section Zapesia, but the veriest tyro would not confound them with species of Parmelia. It is true that a great number of lichens, at first sight, and casually, resemble species of the Hysteriacei, but it is no less strange than true, that lichenologists and mycologists know their own sufficiently not to commit depredations on each other. Contributions are daily being made to this controversy, and already the principal arguments on both sides have appeared in * © Grevillea,” vol. ii. p. 147, in note. 16 FUNGI. an English dress,* hence it will be unnecessary to repeat those which are modifications only of the views already stated, our own conclusions being capable of a very brief summary: that lichens and fungi are closely related the one to the other, but that they are not identical; that the ‘“ gonidia” of lichens are part of the lichen-organization, and consequently are not alge, or any introduced bodies; that there is no parasitism ; aud that the lichen thallus, exclusive of gonidia, is wholly unknown amongst fungi. The Rev. J. M. Crombie has therefore our synipathies in the remark with which his summary of the gonidia controversy closes, in which he characterizes it as a “‘ sensational romance of lichenology,”’ of the ‘ unnatural union between a captive algal damsel and a tyrant fungal master.” * W. Archer, in “Quart. Journ. Micr. Sci.” vol. xiii. p. 217; vol. xiv. p. 115. Translation of Schwendener's ‘‘ Nature of the Gonidia of Lichens,” in same journal, vol. xiii. p. 235, I. STRUCTURE. Wirsour some knowledge of the structure of fungi, it is scarcely possible to comprehend the principles of classification, or to appreciate the curious phenomena of polymorphism. Yet there is so great a variety in the structure of the different groups, that this subject cannot be compressed within a few paragraphs, neither do we think that this would be desired if practicable, seeing that the anatomy and physiology of plants is, in itself, sufficiently important and interesting to warrant a rather ex- tended and explicit survey. In order to impart as much prac- tical utility as possible to this chapter, it seems advisable to treat some of the most important and typical orders and sub- orders separately, giving prominence to the features which are chiefly: characteristic of those sections, following the order of systematists as much as possible, whilst endeavouring to render each section independent to a considerable extent, and complete in itself. Some groups naturally present more noteworthy features than others, and will consequently seem to receive more than their proportional share of attention, but this seem- ing inequality could scarcely have been avoided, inasmuch as hitherto some groups have been more closcly investigated than others, are more intimately associated with other questions, or are more readily and satisfactorily examined under different aspects of their life-history. AGaricin1.—For the structure that prevails in the order to which the mushroom belongs, an examination of that species will be almost sufficient. Here we shall at once recognize 18 FUNGI. : three distinct parts requiring elucidation, viz. the , rooting slender fibres that traverse the soil, and termed the mycelium, or spawn, the stem and cap or pileus, which together con- stitute what is called the hymenophore, and the plates or gill on the under surface of the cap, which bear the hymenium. The earliest condition in which the mushroom can be recognized as a vegetable entity is in that of the “spawn” or mycelium, which is essentially an agglomeration of vegetating spores. Its normal form is that of branched, slender, entangled, anasto- mosing, hyaline threads. At certain privileged points of the my- celium, the threads seem to be aggregated, and become centres of vertical extension. At first only a small nearly globose bad- Fic. 1.—Agaric in Process of Growth. ding, like a grain of mustard seed, is visible, but this after- wards increases rapidly, and other similar buddings or swellings appear at the base.* These are the young hymenophore. As * A curious case occurred some years since at Bury St. Edmunds, which may be mentioned here in connection with the development of these nodules. Two children had died under suspicious circumstances, and an examination of the body of the latter after exhumation was made, a report having arisen that the child died after eating mushrooms. As certain white nodules appeared on the inner surface of the intestines, it was at once hastily concluded that the spores of the mush- room had germinated, and that the nodules were infant mushrooms. This appeared to one of us so strange, that application was made for specimens which were kindly forwarded, and a cursory glance was enough to convince iia that they were not fungoid. An examination under the microscope further con- firmed the diagnosis, and the application of nitric acid showed that the nodules were merely due to chalk mixture, which had been given to the child for the diarrhetic symptoms under which he succumbed, STRUCTURE. 19 it pushes through the soil, it gradually loses its globose form, becomes more or less elongated, and in this condition a longitu- dinal section shows the position of the future gills in a pair of opposite crescent-shaped darker-coloured spots near the apex. The dermal membrane, or outer skin, seems to be continuous over the stem and the globose head. At present, there is no external evidence of an expanded pileus and gills; a longitu- dinal section at this stage shows that the gills are being deve- loped, that the pileus is assuming its cap-like form, that the membrane stretching from the stem to the edge of the young pileus is separating from the edge of the gills, and forming a veil, which, in course of time, will separate below and leave the gills exposed. When, therefore, the mushroom has arrived almost at maturity, the pileus §=——~~—_——~ expands, and in this act the veil is torn away from the margin of the cap, and re- mains for a time like a collar around the stem. Fragments of the veil often remain at- tached to the margin of the pileus, and the collar adhe- rent to the stem falls back, and thenceforth is known as the annulus or ring. We have in this stage the fully- developed hymenophore, — the stem with its ring, sup- porting an expanded cap or pileus, with gills on the under surface. bearing the hyme- Fic, 2.—Section of Common Mushroom. nium.* A longitudinal section cut through the pileus and down * Ebrenberg compared the whole structure of an Agaric with that of a mould, ‘the mycelium’ corresponding with the hyphasma, the stem and pileus with the flocci, and the hymenium with the fructifying branchlets. The comparison is no less ingenious than true, and gives a lively idea of the connection of the more noble with the more humble fungi.—Zhrb. de Mycetogenesi. 20 TUNGI. the stem, gives the -best notion of the arrangement of the parts, and their relation to the whole. By this means it will be seen that the pileus is continuous with the stem, that the sub- stance of the pileus descends into the gills, and that relatively the substance of the stem is more fibrous than that of the pileus. In the common mushroom the ring is very distinct surrounding the stem, a little above the middle, like a collar. In some Agarics the ring is very fugacious, or absent altogether. The form of the gills, their mode of attachment to the stem, their colour, and more especially the colour of the spores, are all very important features to be attended to in the discrimination of species, since they vary in different species. The whole substance of the Agaric is cellular. A longitudinal slice from the stem will exhibit under the microscope delicate tubular cells, the general direction of which is lengthwise, with lateral branches, the whole interlacing so intimately that it is diffi- cult to trace any individual thread very far in its course. It will be evident that the structure is less compact as it approaches the centre of the stem, which in many species is hollow. The hymenium is the spore-bearing surface, which is exposed or naked, and spread over the gills. These plates are covered on all sides with a delicate membrane, upon which the reproductive organs are developed. If it were possible to remove this membrane in one entire piece and spread it out flat, it would cover an immense surface, as compared with the size of the pileus, for it is plaited or folded like a lady’s fan over the whole of the gill- plates, or lamella, of the fungus.* If the stem of a mushroom be cut off close to the gills, and the cap laid upon a sheet of paper, with the gills downwards, and left there for a few hours, when removed a number of dark radiating lines.will be deposited upon the paper, each line corresponding with the interstices between one pair of gills. These lines are made up of spores which have fallen from the hymenium, and, if placed under the microscope, their character will at once’ be made evident. If a fragment of the hymenium be also submitted to a similar examination, it will be found that the whole surface is studded * In Pawillus involutus the hymenium may be readily torn off and unfolded. STRUCTURE. 91 with spores. The first peculiarity which will be observed is, that these spores are almost uniformly in groups of four together. The next feature to be observed is, that each spore is borne upon a slender stalk or sterigma, and that four of these sterigmata proceed from the apex of a thicker projection, from the hymenium, called a basidiwm, each basidium being the sup- porter of four sterigmata, and each sterigma of a spore.* A closer examination of the hymenium will reveal the fact that the basidia are accompained by other bodies, often larger, but without sterigmata or spores; these have been termed cystidia, and their structure and functions have been the subject of much controversy. Both kinds of bodies are produced on the hymenium of most, if not all, the Agaricini. The basidia are usually expanded upwards, so as to have more or less of a clavate form, surmounted by four slender points, or tubular processes, each supporting a spore ; the contents of these cells are granular, mixed apparently with oleaginous particles, which communicate through the slender tubes of the spicules with the interior of the spores. Corda ar ek a ae states that, although only one spore is sidia.” c. Cystidium. From Gon produced at a time on each sporo-” ES ans: : phore, when this falls away others are produced in succession for a limited period. As the spores approach maturity, the con- nection between their contents and the contents of the basidia diminishes and ultimately ceases. When the basidium which bears mature spores is still well: charged with granular matter, it may be presumed that the production of a second or third * This was well delineated in ‘‘ Flora Danica,” plate 834. as observed in Coprinus comatus as long ago as 1780. + A. de Bary, ‘‘ Morphologie und Physiologie der Pilze,” in ‘‘Hofmeister’s Hand- buch,” vol. ii. cap. 5, 1866, translated in ‘‘ Grevillea,” vol. i. p. 181. 22 FUNGI. series of spores is quite possible. Basidia exhausted entirely of their contents, and which have become quite hyaline, may often be observed. The cystidia are usually larger than the basidia, varying in size and form in different species. They present the appearance of large sterile cells, attenuated upwards, sometimes into a slender neck. Corda was of opinion that these were male organs, and gave them the name of pollinaires. Hoffmann has also described* both these organs under the names of pollinaria and spermatia, but does not appear to recognize in them the sexual elements which those names would indicate ; whilst de Seynes suggests that the cystidia are only organs returned to vegetative functions by a sort of hypertrophy of the basidia.t This view seems to be supported by the fact that, in the section Pluteus and some others, the cystidia are surmounted by short horns resembling sterigmata. Hoffmann has also indicated t . the passage of cystidia into basidia. The evidence s¢ems to be in favour of regarding the cystidia as barren conditions of basidia. There are to be found upon the hymenium of Agarics a third kind of elongated cells, called by Corda§ basilary cells, and by Hoffmann “ sterile cells,’ which are either equal in size or smaller than the basidia, with which also their structure agrees, except- ing in the development of spicules. These are the “ proper cclls of the hymenium” of Léveillé, and are simply. the terminal cells of the gill structure—cells which, under vigorous conditions, might be developed into basidia, but which are commonly arrested in their development. As suggested by de Seynes, the hymenium seems to be reduced to great simplicity, “ one sole and self-same organ is the basis of it; according as it experiences an arrest of development, as it grows and fructifies, or as it- becomes hypertrophied, it gives us a paraphyse, a basidium, or a cystidium—ain other terms, atrophied basidium, normal basi- * “Die Pollinarien und Spermatien von Agaricus,” in ‘‘ Botanische Zcitung,”” Feb. 29 and March 7, 1856. + ‘Essai d'une Flore mycologique de la Région de Montpellier.” Paris, 1863. t Hoffmann, ‘‘ Botanische Zeitung,” 1856, p. 139. ; § Corda, ‘‘ Icones Fungorum hucusque cognitorum,” iii. p. 41, Prague, 1839, STRUCTURE. 23 dium, and hypertrophied basidium ; these are the three elements which form the hymenium.’’* The only reproductive organs hitherto demonstrated in Agarics are the spores, or, as sometimes called, from their method of production, basidiospores.+| These are at first colourless, but afterwards acquire the colour peculiar to the species. In size ‘and form they are, within certain limits, exceedingly variable, although form and size are tolerably constant in the same species. Atfirst allare globose; as they mature, the majority are ovoid or elliptic ; some are fusiform, with regularly attenuated extremities. In Hygrophorus they are rather irregular, reniform, or compressed in the middle. Sometimes the external surface is rough with more or less projecting warts. Some mycologists are of opinion that the covering of the spore is double, consist- ing of an ewospore and an endospore, the latter being very fine and delicate. In other orders the double coating of the spore has been demonstrated. When the spore is coloured, the exter- nal membrane alone appears to pos- sess colour, the endospore being con- stantly hyaline. It may be added here, that in this order the spore is simple and unicellular. In Lactarius and Russula the trama, or inner substance, isvesicular. True latex vessels occur occasionally in Agaricus, though not filled with milk as in Lactarius. Potyrorer.—In this order the gill plates are replaced by tubes or pores, the interior of which is lined by the 5, i errs hymenium ; indications of this struc- duced). ture having already been exhibited in some of the lower i i i hy a My A ll ‘< * Cooke, M. C., ‘‘ Anatomy of a Mushroom,” in ‘‘ Popular Science Review,” vol. viii. p. 380. ¢ An attempt was made to show that, in Agaricus melleus, distinct asci were found, in a certain stage, on the gills or lamella. We have in vain examined the gills in various conditions, and could never detect anything of the kind. It is probable that the asci belonged to some species of Mypomyces, a genus of para- sitic Spheeriaceous fungi. . : 24 ‘FUNGI. Agaricini. In many cases the stem is suppressed. The sub- stance is fleshy in Boletus, but in Polyporus the greater number of species are leathery or corky, and more persistent. The basidia, spicules, and quaternate spores agree with those of Agaricini.* In fact there are no features of importance which “relate to the hymenium in any order of Hymenomycetes (the Tremellini excepted) differing from the same organ in Agaricim, unless it be the absence of eystidia. Hypnet.—Instead of pores, ~~ in this order the hymenium Wy. : oa “Ti a