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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 thet the same high character will justly apply to this ag to Mr. Cooke's former publications, and especially to his “Handbook of British Fungi.” M. J. BERKELEY. SIBDERTOFT, Movember 23, 1874. C O N T E N T S. —º-º- I. PAGE NATURE OF FUNGI * gº { } ſº g tº © * © o II. STRUCTUR: tº Q g © º © º º Q Q . 17 III, CLASSIFICATION . © g Jº © sº & * tº * ... 64 IV. USES & © © tº tº & º o tº sº © © * , S2 W *. NoTABLE PHENOMENA . © ë e ſº º gº º Q. ... 105 WI. The Spore. AND ITS DISSEMINATION . • • tº . . . 119 viii CONTENTS, GERMINATION * GROWTH . SExUAL REPRODUCTION . PolyMoRPHISM * > Q tº INFLUENCES AND EFFECTs tº HABITATS • º o dº CULTIVATION . . . . GEOGRAPHICAL DISTRIBUTION . CoLLECTION AND PRESERVATION WII. VIII. IX. X. XI. XII. XIII. XIV. PAGE . 137 . 163 • 209 • 233 , 253 , 266 . 287 LIST OF II, LUSTRATIONS. —£4– FIG PAGE 1. Agaric in process of growth .. º * º • . C . 18 2. S.ction of common mushroom . © © º g e & • 19 3. Sterile cells, basidia, and cystidia, from Gomphidius . tº º . 21 4. Polyporus giganteus (reduced). * * & e * . 23 5. Hydnwm, repandwm (section) . o tº tº • . e. ſº . . 24 6. Calocera viscosa . tº tº tº tº & e te & . 25 7. Tremella mesenterica. tº * e e tº © ſº g . 25 8. Basidia and spores of Phallus . o º tº tº tº tº . 28 9. Basidia and spores of Lycoperdon . . . . . . . a 30 10. Threads of Trichia . . . . . . . . . . 32 II. Arcyria incarnata, with threads and spore o g o tº . 33 12. Diachaea elegans . tº tº º tº g º tº e . 34 13. Cyathus vernicosus . . . . . . . . . . . 84 14. Cyathus, sporangia and spores . * e º © & g . 35 15. Asterosporium Hoffmann? º tº ge º e • wº . 36 16. Barren cysts and pseudospores of Lecythea * * tº © º . 37 17. Coleosporium Tussilaginis . tº te gº º º * . 37 18. Melampsora salicina, pseudospores of e tº tº © e . 37 19. Cystopus candidus, conidia of . © © º wº g e . 38 20. Xenodochus carbonarius, pseudospore © º ſº e . 39 21. Phragmidium bulbosum, pseudospores . . e e º . 39 22. Pseudospores of Puccinia * © tº e sº g {} . 40 X. * LIST OF ILLUSTRATIONS. FIGH, 23. Thecaphora hyalina, pseudospores . tº 24. Alcidium Berberidis, peridia of 25. Helminthosporium molle, threads and spores 26. Acrothecium simplea, . gº * > ſº 27. Peronospora Arenariæ . ( ) tºo 28. Polyactis cinerea . g & 29. Peziza Fuckeliana, with ascus and sporidia. 30. Penicillium chartarwm . & g º 31. Mucor mucedo, with sporangia & tº 32. Small portion of Botrytis Jonesi: . e 33. Section of cup of .48cobolus . & & 34. Asci, sporidia, and paraphyses of Ascobolus 35. Perithecium of Sphaeria. 36. Uncinula adunca, conceptacle with appendages 37. Agaricus nudus . wº e e 38. Scleroderma vulgare . . ſº e 39. Ceuthospora phacidioides . e 40. Rhopalomyces candidus. gº 41. Mucor caninus . tº & 42. Sphaeria aquila, cluster of perithecia 43. Morchella gigaspora, from Kashmir 44, Cyttaria Gunnii . g g © 45. Spores of Agarics . tº " ſº * - e. 46. Spores of Lactarius . e dº e 46*. Spores of Gomphidius . º ge 47. Spores of Polyporus, Boletus, and IIydnum 48. Diachaea elegans, capillitium of - 49. Spore of Hendersonia polycystis . & 50. Spores of Dilophospora gramin's . 51. Spores of Discosia {e * e e 52. Spore of Prosthemium betw!inum . © PAGE 41 41 43 44 44 45 48 50 5] 53 57 59 61 62 66 69 70 74 78 99 101 121 121 122 122 123 53. Spore of Stegonosporium cellulosum © 124 124 124 124 125 IIST OF ILLUSTRATIONS. FIG. 54. Stylospores of Coryneum disciforme . . g 55. Spores of Asterosporium Hoffmanni tº 56. Spores of Pestalozzia. . 83. . Bispora monilioides, concatenate spores . 58. Pseudospores of Thecaphora hyalina. º 59. Pseudospores of Puccinia . ſº * o 60. Pseudospores of Triphragmium . e tº 61. Pseudospores of Phragmidium bulbosum. © 62. Winter spores of Melampsora salicina, 63. Spores of Helicocoryne . tº sº º 64. Sporidium of Genea, verrucosa * o 65. Alveolate sporidium of Tuber . * & 66. Asci, sporidia, and paraphyses of Ascobolus 67. Sporidium of ostreichnion Americanum . tº 68. Ascus and sporidia of Hypocrea. . o 69. Sporidium of Sphaeria ulnaspora . Cº. * 70. Sporidia of Walsa profusa . e º 71. Sporidia of Massaria fabdans . . © tº 72. Sporidium of Melanconis bicornis . e & 73. Caudate sporidia of Sphaeria ſimiseda . º 74. Sporidia of Walsa thelebola. . . te º 75. Sporidia of Walsa taleola . o e º 76. Sporidia of Sporormia intermedia . - 78. Sporidium of Sphaeria putaminwm, . . Dasidia and spores of Exidia spiculosa . Germinating spore and corpuscles of Dacrymyces . Germination of Æcidium Euphorbide . Germinating pseudospores of Coleosporium Sonchi Germinating pseudospores of Melannpsora betwlina 77. Asci and sporidia of Sphaeria (Pleospora) kerbacum. 84. Germinating pseudospore of Uromyces appendiculatus 85. Germinating pseudospore of Puccinia Moliniae e PAGE 125 25 126 126 127 127 127 127 127 129 130 130 131 132 xii LIST OF ILLUSTRATIONS. FIG. * PAGF) 86. Germinating pseudospore of Thriphragmium Ulmariat . g . 146 87. Germinating pseudospore of Phragmidium bulbosum. . © . 147 88. Germinating pseudospores of Podisoma Juniper; . º ſº . 148 89. Germinating pseudospore of Tilletia caries . † & tº . 150 b0. Pseudospore of Ustilago receptaculorum in germination, and secondary spores in conjugation . ge ſº º * {º º . 151 91. Conidia and zoospores of Cystopus candidus . g e e . 151 . 92. Resting spore of Cystopus candidus with zoospores . . . Ç . 152 93. Zygospores of Mucor phycomyces . . cº wº ſº g . 158 94. Sporidium of Ascobolus germinating . tº tº tº tº . 161 95. Zygospore of Mucor . te o tº ſº { } gº e , 165 96. Zygospore of Rhizopus in different stages gº § * e . 167 97. Conjugation in Achlya racemosa . g ę ſº g e . . 169 98. Conjugation in Peronospora . g & º & Q e . 171 99. Antheridia and oogonium in Peronospora . tº G is . 172 100. Conjugation in Peziza omphalodes. o © tº º & . 175 100*. Formation of conceptacle in Erysiphe . * º dº © . 176 101. Tilletia caries with conjugating cells . © wº de gº . 178 102. Aspergillus glaucus and Eurotium ſº © e & . . . 189 103. Erysiphe cichoraccarum, receptacle and mycelium . & e • 191 104. Twig with Tubercularia and Nectria . º g g g . 193 . 105. Section of Tubercularia with conidia . • • * s . 194 106. Wectria with Tubercularia, ascus and paraphyses g sº e . 195 107. Cells and pseudospores of Æcidium Berberidis g © to . 201 108. Cells and pseudospores of Æcidium graveolens © º tº . 201 109. Torrubia militaris on pupa of a moth . . . . . . 243 F U N G I THEIR NATURE, USES, INFLUENCES, ETC. —4–4–s I. NATURE OF FUNGI. THE most casual observer of Nature regognizes 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 2 EUNGI. 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 Balanophora, 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 a little more, but still elementary knowledge, distribute thé latter amongst ferns, mosses, fungi, lichens, and algae, with comparatively few exceptions. It is true that between 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 Myanogastres 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 Amoeta,” whilst Tulasne * De Bary, “Des Myxomycetes,” 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. {} 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 Myxogastres in their early stage, and the result of his examinations has been a firm conviction that there is no relationship whatever between the Myaogastres 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 Aſya.cgastres 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 algae, 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:–1. 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. 3. 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.f The interpretation of this is, that the mould Penicillium may be * De Bary, “Recherches sur le Developpement de quelques Champignons 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 Torulae, f 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 (Torula), 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 little 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., 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 Torulae, such as T. moni- lioides, &c.—Cooke's Handbook, p. 477. NATURF: 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 FUNGI, 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 described as “hysterophytal or epiphytal mycetals deriving *Berkeley's “Outlines of British Fungology,” p. 24. NATURE OF FINGI. 7 nourishment by means of a mycelium from the matrix,” thero 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 Saprolegniei, which some authors include with fungi, and others with algae. 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 Sphaeria afterwards developed. Some species of Isaria and Torrubia also affect the larvae and pupae of moths and butterflies, converting the whole interior into a mass of mycelium, and fructifying in a 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 Isaria resembles so closely that of the house-fly when succumbing to Sporendonema Musca, 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 disease Muscardine attacks the living insect, and causes death. In the case of the Guépes végétantes, 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 such a 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 from a 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 algae that it is almost impossible to say 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. In a night a puffball will grow prodigiously, and in the same short period a mass of paste may be covered with mould. In a 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 theories, 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 his 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 algae. In language which the Rev. J. M. Crombie # describes as “pictorial,” this author gave the gene:al 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 algae, 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 über den Flechtenthallus.” + Crombie (J. M.) “On the Lichen-Gonidia Question,” in “Popular Science Review” for July, 1874. NATURE OF 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 algae 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—1st. 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 of algae, whilst the second had that of fungi; 3rd. That the different forms and varieties of gonidia corresponded with parallel types of algae; 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 algae, with a fungus parasitic upon them. The 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 algae; 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 algae, 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 mammiferae. 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 any kind. 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,f 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,” &c., in “Grevillea,” vol. ii. (1874), No. 22, p. 146. t In Regensburg “Flora,” 1870, p. 92. NATURE OF FUNGI. 13 deprived of all autonomous liberty, is not at all consonant with the manner of existence of the other algae, 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 algae, and that these gonidia can continue to vegetate separately, and so be mistaken for unicellular algae.” 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 algae, according to the turbid hypothesis of Schwendener, from constituting true algae, that on the contrary it may be affirmed that they have a lichenose nature, whence it follows that these pseudo-algae are in a systematic arrangement to be referred rather to the lichens, and that the class of algae 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 hyphae of such could in themselves alone no more serve for propagation than the hyphae 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 hyphae 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 Emericella, f which is allied to Husseia in the Trichogastres, shows a structure in the stem exactly resembling Palmella botryoides of Greville, and to what occurs in Synalyssa. I'mericella, 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 algae 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 algae. The question cannot be settled by mere à priori notions. It is, perhaps, worthy of remark that in Chion/phe Carteri 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,t 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 P 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 as seemed 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 a century. This should at least serve as a caution for the future. Apart, however, from the “gonidia,” whatever they may be, is the remainder of the lichen a genuine fungus P. Nylander writes, “The anatomical filamentose elements of lichens are distinguished by various characters from the hyphae of fungi. They are firmer, elastic, and at once present themselves in the texture of lichens. On the other hand, the hyphae 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 hydrate of potash, &c.” - Our own experience is somewhat to the effect, that there are some few lichens which are doubtful as to whether they are fungi 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 Torrubia or Clavaria. We have Peziza with a subiculum in the section Tapesia, 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 rusal. 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 algae, or any introduced bodies; that there is no parasitism; and that the lichen thallus, exclusive of gonidia, is wholly “unknown amongst fungi. The Rev. J. M. Crombie has therefore our sympathies 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. II. STRUCTURE. WITHOUT 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 closely investigated than others, are more intimately associated with other questions, or are more readily and satisfactorily examined under different aspects of their life-history. AGARICINI-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 gills 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 bud- FIG. 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 us 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. I 9 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- FIG. 2.-Section of Common Mushroom. nium.* A longitudinal section cut through the pileus and down * Ehrenberg 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. —Ehrb. de Mycetogenesi. 20 FUNGI, 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 lamellae, 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 Pazillus involutus the hymenium may be readily torn off and unfolded. STRUCTURE. 21 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 basidium, 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.t 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 FIG. 3.-a. Sterile cells. b. Ba- states that, although only one spore is sidia, c. Cystidium. From Gom- º phidius (de Seynes). produced at a time on each sporo- 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.f 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 ; the passage of cystidia into basidia. The evidence seems 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 Cordaš 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 cells 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—in other terms, atrophied basidium, normal basi- * “Die Pollinarien und Spermatien von Agaricus,” in “Botanische Zeitung,” Feb. 29 and March 7, 1856. + “Essai d'une Flore mycologique de la Région de Montpellier.” Paris, 1863. it 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. i. 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. At first all are 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 eacospore 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- sº I'll), stantly hyaline. It may be added here, tº "\, (". e - º e j | ſº !!!, \l that in this order the spore is simple N sº !"/. and unicellular. In Lactarius and §§§ º [. | '. ſt ºš % /. 3. S Nº. Russula the trama, or inner substance, is vesicular. True latex vessels occur occasionally in Affaricus, though not filled with milk as in Lactarius. PolypoREI.-In this order the gill $$. :, plates are replaced by tubes or pores, Q C the interior of which is lined by the rº, 4.—Polyporus gigantents 're- hymenium ; indications of this struc- duced). ture having already been exhibited in some of the lower º - &º • *º t . . .3 % £º § % w º /. º z .** Pº Ǻ *-** * Cooke, M. C., “Anatomy of a Mushroom,” in “Popular Science Review,” vol. viii. p. 380. . + Am attempt was made to show that, in Agaricus melleus, distinct asci were found, in a certain stage, on the gills or lamellae. 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 Hypomyces, a genus of para- sitic Sphoeriaceous fungi. 24 - FUNGI. Agartevni. 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 Agaricini, unless it be the absence of cystidia. HYDNEI.—Instead of pores, ~. $ in this order the hymenium "ºl ~sº is spread over the surface of |N| % ) ºš spines, prickles, or warts.} % is sº sº * | ſh N AURICULARINI.-The hyme nium is more or less even, Š=== % and in— OO CLAVARIEI the whole fungus C) } is club-shaped, or more or less intricately branched, with FIG. 5.-Hydrºwn repandwin. the hy menium covering the outer surface. TREMELLINI.-In this order we have a great departure from the character of the substance, external appearance, and internal structure of the other orders in this family. Here we have a gelatinous substance, and the form is lobed, folded, convolute, often resembling the brain of some animal. The internal struc- * It is not intended that the spores are always quaternate in Agaricini, though that number is constant in the more typical species. They sometimes exceed four, and are sometimes reduced to one. + The species long known as Hydnºm gelatinosum was examined by Mr. F. Currey in 1860 (Journ. Linn. Soc.), and he came to the conclusion that it was not a good Hydmum. Since then it has been made the type of a new genus (ffydnoglaca B. and Br. or, as called by Fries, in the new edition of “Epicrisis,” Tremellodon, Pers. Myc. Eur.), and transferred to the Tremellini. Currey says, upon examining the fructification, he was surprised to find that, although in its external characters it was a perfect Hydmum, it bore the fruit of a Tremella. If one of the teeth be examined with the microscope, it will be seen to consist of threads bearing four-lobed sporophores, and spores exactly similar to Tremella. It will thus be seen, he adds, that the plant is exactly intermediate between IIydnei and Tremellini, forming, as it were, a stepping-stone from one to the other. STRUCTURE. 25 ture has been specially illustrated by M. Tulasne,” through the common species, Tremella mesenterica. This latter is of a fine golden yellow colour, and rather large size. It is uniformly composed throughout of a colourless mucilage, with no appreciable texture, in which are distributed very fine, diversely branched and anastomosing filaments. Towards the surface, the ultimate branches of this filamentous network give birth, both at their summits and laterally, to globular cells, which ac- - quire a comparatively large size. FIG. 6. —Calocera viscosa. These cells are filled with a protoplasm, to which the plant owes its orange colour. When they have attained their normal dimensions, they elongate at the summit into two, three, or four distinct, thick, obtuse tubes, into which the protoplasm gradually passes. The development º • * > \;\ § of these tubes is unequal and In Ot § $º simultaneous, so that one will often tº ºù attain its full dimensions, equal, per- haps, to three or four times the dia- meter of the generative cell, whilst the others are only just appearing. By degrees, as each tube attains its full size, it is attenuated into a fine point, the extremity of which swells into a spheroidal cell, which ulti- mately becomes a spore. Sometimes these tubes, or spicules, send out one or two lateral branches, each terminated by a spore. These spores (about 006 to ‘008 mm. diameter) are smooth, and deposit themselves, like a fine white dust, on the surface of the Tremella and on its matrix. M. Léveillé+ was of opinion that FIG. 7.—Tremella mesentericat. * Tulasne, L. R. and C., “Observations on the Organization of the Tremellini,” in “Ann, des Sci. Nat.” 3" sér. xix. (1853), pp. 193, &c. + M. Léveillé, in “Ann. des Sci. Nat.” 2ne sér. viii. p. 328; 3me sér. ix. p. 127; also Bonorden, “‘Handbuch der Mycologie,” p. 151. 26 FUNGI. the basidia of the Tremellini were monosporous, whilst M. Tulasne has demonstrated that they are habitually tetrasporous, as in other of the Hymenomycetes. Although agreeing in this, they differ in other features, especially in the globose form of the basidia, mode of production of the spicules, and, finally, the division of the basidia into two, three, or four cells by septa which cut each other in their axis. This division precedes the growth of the spicules. It is not rare to see these cells, formed at the expense of an unilocular basidium, become partly isolated from each other ; in certain cases they seem to have separated very early, they then become larger than usual, and are grouped on the same filament so as to represent a kind of buds. This phenomenon usually takes place below the level of the fertile cells, at a certain depth in the mucous tissue of the Tremella. Besides the reproductive system here described, Tulasne also made known the existence of a series of filaments which produce spermatia. These filaments are often scattered and confused with those which produce the basidia, and not distinguishable from them in size or any other apparent characteristic, except the manner in which their extremities are branched in order to produce the spermatia. At other times the spermatia-bearing surface covers exclusively certain portions of the fungus, espe- cially the inferior lobes, imparting thereto a very bright orange colour, which is communicated by the layer of spermatia, unmixed with spores. These spots retain their bright colour, while the remainder of the plant becomes pale, or covered with a white dust. The spermatia are very small, spherical, and smooth, scarcely equalling ‘002 mm. They are sessile, some- times solitary, sometimes three or four together, on the slightly swollen extremities of certain filaments of the weſt of the fungus.* Tulasne found it impossible to make these cor- puscles germinate, and in all essential particulars they agreed with the spermatia found in ascomycetous fungi. In the genus Dacrymyces, the same observer found the structure * Tulasne, in “Ann. des Sci. Nat.” (loc. cit.) xix. pl. x, fig. 29. Tulasne, “INew Notes upon Tremellinous Iungi,” in “Journ. Linn. Soc.” vol. xiii. (1871), p. 31. STRUCTURE. 27 to have great affinity with that of Tremella. The spores in the species examined were of a different form, being oblong, very obtuse, slightly curved ('013–1019 × 1004–006 mm.), at first unilocular, but afterwards triseptate. The basidia are cylin- drical or clavate, filled with coloured granular matter; each of these bifurcates at the summit, and gradually elongates into two very open branches, which are attenuated above, and ultimately each is crowned by a spore. There are to be found also in the species of this genus globose bodies, designated “sporidioles'' by M. Léveillé, which Tulasne took considerable care to trace to their source. He thus accounts for them :—Each of the cells of the spore emits exteriorly one or several of these corpuscles, supported on very short and very slender pedicels, which remain after the corpuscles are detached from them, new corpuscles succeeding the first as long as there remains any plastic matter within the spore. The pedicels are not all on the same plane; they are often implanted all on the same, and oftenest on the convex side of the reproductive body. These corpuscles, though placed under the most favourable conditions, never gave the least sign of vegetation, and Tulasne concludes that they are spermatia, analogous to those produced in Tremella. The sporos which produce spermatia are not at all apt to germinate, whilst those which did not produce spermatia germinated freely. Hence it would appear that, although all spores seem to be perfectly iden- tical, they have not all the same function. The same observer detected also amongst specimens of the Dacrymyces some of a darker and reddish tint, always bare of spores or spermatia on the surface, and these presented a somewhat different structure. Where the tissue had turned red it was sterile, the constituent filaments, ordinarily colourless, and almost empty of solid matter, were filled with a highly-coloured protoplasm; they were of less tenuity, more irregularly thick, and instead of only rarely pre- senting partitions, and remaining continuous, as in other parts of the plant, were parcelled out into an infinity of straight or curved pieces, angular and of irregular form, especially towards the surface of the fungus, where they compose a sort of pulp, varying in cohesion according to the dry or moist condition of 28 FUNGI. the atmosphere. All parts of these reddish individuals seemed more or less infected with this disintegration, the basidia divided by transverse diaphragms into several cylindrical or oblong pieces, which finally become free. Transitional conditions were also observed in mixed individuals. This sterile condition is called by Tulasne “gemmiparous,” and he believes that it has ere now given origin to one or more spurious species, and misled mycologists as to the real structure of perfect and fruitful Dacrymyces. PHALLOIDEI.—In this order the hymenium is at first enclosed within a sort of peridium or universal volva, maintaining a somewhat globose or egg-shape. This envelope consists of an outer and inner coat of somewhat similar texture, and an inter- mediate gelatinous layer, often of considerable thickness. When a section is made of the fungus, whilst still enclosed in the Volva, the hymenium is found to present numerous cavities, in which basidia are developed, each surmounted by spicules (four to six) bearing oval or oblong spores.* It is very difficult to observe the structure of the hy- menium in this order, on account of its deliques- cent nature. As the hymenium approaches ma- turity, the volva is ruptured, and the plant rapidly enlarges. In Phallus, a long erect cellular stem bears the cap, over which the hymenium is a.º.º spread, and this expands enormously after escap- lw8. ing the restraint of the volva. Soon after expo- sure, the hymenium deliquesces into a dark mucilage, coloured by the minute spores, which drips from the pileus, often diffus- ing a most loathsome odour for a considerable distance. In Clathrus, the receptacle forms a kind of network. In Aserüe, the pileus is beautifully stellate. In many the attractive forms would be considered objects of beauty, were it not for their deliquescence, and often foetid odour.t * Berkeley, M. J., “On the Fructification of Lycoperdon, Phallus, &c.,” in “Ann. Nat. Hist.” 1840, vol. iv. p. 158, pl. 5. Berkeley, M. J., “Introduc- tion Crypt. Bot.” p. 346. ‘i Tulasne, L. R. and C., “Fungi Hypogoei.” Paris. Berkeley and Broome, STRUCTURE. 29 PoDAxINEI.-This is a small but very curious group of fungi, in which the peridium resembles a volva, which is more or less confluent with the surface of the pileus. They assume hymeno- mycetal forms, some of them looking like Agarics, Boleti, or species of Hydnum, with deformed gills, pores, or spines; in Montagmites, in fact, the gill structure is very distinct. The spores are borne in definite clusters on short pedicels in such of the genera as have been examined.* HYPOGOEI.—These are subterranean puff-balls, in which some- times a distinct peridium is present; but in most cases it consists entirely of an external series of cells, continuous with the in- ternal structure, and cannot be correctly estimated as a peridium. The hymenium is sinuous and convolute, bearing basidia with sterigmata and spores in the cavities. Sometimes the cavities are traversed by threads, as in the Myanogastres. The spores are in many instances beautifully echinulate, sometimes globose, at others elongated, and produced in such numbers as to lead to the belief that their development is successive on the spicules. When fully matured, the peridia are filled with a dusty mass of spores, so that it is scarcely possible in this condition to gain any notion of the structure. This is, indeed, the case with nearly all Gasteromycetes. The hypogoeous fungi are curiously connected with Phalloidei by the genus Hysterangium. TRICHOGASTRES..+-In their early stages the species contained in this group are not gelatinous, as in the Myaogastres, but are rather fleshy and firm. Very little has been added to our knowledge of structure in this group since 1839 and 1842, when one of us wrote to the following effect:—If a young plant of Lycoperdon caelatum or L. gemmatum be cut through and examined with a common pocket lens, it will be found to consist of a fleshy mass, “British Hypogoeous Fungi,” in “Ann. Nat. Hist.” 1846, xviii. p. 74. Corda, “Icones Fungorum,” vol. vi. pl. vii. viii. * Tulasne, “Sur le Genre Secotium,” in “Ann. des Sci. Nat.” (1845), 3” sér. vol. iv. p. 169, plate 9. # Tulasne, L. R. and C., “De la Fructification des Scleroderma comparée a celle des Lycoperdon et des Borista,” in “Ann. des Sci. Nat.” 1842, xvii. p. 5. Tulasne, L. R. and C., “Sur les Genres Polysaccum et Geaster,” in ‘‘Ann. des Sci. Nat.” 1842, xviii. p. 129, pl. 5 and 6. 30 FUNGI. perforated in every direction with minute elongated, reticulated, anastomosing, labyrinthiform cavities. The resemblance of these to the tubes of Boleti in an early stage of growth, first led me to suspect that there must be some very close connection between them. If a very thin slice now be taken, while the mass is yet firm, and before there is the slightest indication of a change of colour, the outer stratum of the walls of these cavities is found to consist of pellucid obtuse cells, placed parallel to each other like the pile of velvet, exactly as in the young hymenium of an Agaric or Boletus. Occasionally one or two filaments cross from one wall to another, and once I have seen these anastomose. At a more advanced stage of growth, four little spicules aro developed at the tips of the sporo- phores, all of which, as far as I have been able to observe, are fertile and of equal height, and on each of these 'spicules a globose spore is seated. It is clear that we have here a structure identical with that of the true Hy- menomycetes, a circumstance which accords well with the fleshy habit and mode of growth. There is some diffi- culty in ascertaining the exact struc- Fig. 9–Basidia and spores ture of the species just noticed, as of Lycoperdon. the fruit-bearing cells, or sporophores, are very small, and when the spicules are developed the substance becomes so flaccid that it is difficult to cut a proper slice, even with the sharpest lancet. I have, however, satisfied myself as to the true structure by repeated observations. But should any difficulty arise in verifying it in the species in question, there will be none in doing so in Lycoperdon giganteum. In this species the fructifying mass consists of the same sinuous cavities, which are, however, smaller, so that the substance is more com- pact, and I have not seen them traversed by any filaments. In an early stage of growth, the surface of the hymenium, that is of the walls of the cavities, consists of short threads composed of two or three articulations, which are slightly constricted at the STRUCTUTE. 31 joints, from which, especially from the last, spring short branch- lets, often consisting of a single cell. Sometimes two or more branchlets spring from the same point. Occasionally the threads are constricted without any dissepiments, the terminal articula- tions are obtuse, and soon swell very much, so as greatly to exceed in diameter those on which they are seated. When arrived at their full growth, they are somewhat obovate, and produce four spicules, which at length are surmounted each with a glo- bose spore. When the spores are fully developed, the sporophores wither, and if a solution of iodine be applied, which changes the spores to a rich brown, they will be seen still adhering by their spicules to the faded sporophores. The spores soon become free, but the spicule often still adheres to them ; but they are not attached to the intermingled filaments. In Bovista plumbea, the spores have very long peduncles.” As in the Hymenomycetes, the prevailing type of reproductive organs consisted of quaternary spores borne on spicules; so in Gastero- mycetes, the prevailing type, in so far as it is yet known, is very similar, in some cases nearly identical, consisting of a definite number of minute spores borne on spicules seated on basidia. In a very large number of genera, the minute structure and development of the fructification (beyond the mature spores) is almost unknown, but from analogy it may be concluded that a method prevails in a large group like the Myaogastres which does not differ in essential particulars from that which is known to exist in other groups. The difficulties in the way of studying the development of the spores in this are far greater than in the previous order. MyxoGASTRES.–At one time that celebrated mycologist, Pro- fessor De Bary, seemed disposed to exclude this group from the vegetable kingdom altogether, and relegate them to a companion- ship with amoeboid forms. But in more recent works he seems to have reconsidered, and almost, if not entirely, abandoned, that disposition. These fungi, mostly minute, are characterized in their early stages by their gelatinous nature. The substance * Berkeley, “On the Fructification of Lycoperdon, &c.,” in “Annals of Natural History” (1840), iv. p. 155. 32 FUNGI. of which they are then composed bears considerable resemblance to sarcode, and, did they never change from this, there might be some excuse for doubting as to their vegetable nature; but as the species proceed towards maturity they lose their mucilaginous texture, and become a mass of spores, intermixed with threads, surrounded by a cellular peridium. Take, for instance, the genus Trichia, and we have in the matured specimens a somewhat globose peridium, not larger than a mustard seed, and some- times nearly of the same colour; this ultimately ruptures and exposes a mass of minute yellow spherical spores, intermixed with threads of the same colour.” These threads, when highly magnified, exhibit in themselves a spiral arrangement, which has been the basis of some controversy, and in some species these threads are externally spinulose. The chief controversy Q. FIG. 10.-a. Threads of Trichia. b. Portion further magnified, with spores. c. Por- tion of spinulose thread. on these threads has been whether the spiral markings are external or internal, whether caused by twisting of the thread or by the presence of an external or internal fibre. The spiral appearance has never been called in question, only the structure from whence it arises, and this, like the striae of diatoms, is very much an open question. Mr. Currey held that the spiral * Wigand, “Morphologie des Genres Trichia et Arcyria,” in “Ann. des Sci. Nat.” 4me sér. xvi. p. 223. STRUCTURE. 33 appearance may be accounted for by supposing the existence of an accurate elevation in the wall of the cell, following a spiral direction from one end of the thread to the other. This supposition would, he thinks, accord well with the optical appearances, and it would account exactly for the undulations of outline to which he alludes. He states that he had in his possession a thread of Trichia chrysosperma, in which the spiral appearance was so manifestly caused by an elevation of this nature, in which it is so clear that no internal spiral fibre exists, that he did not think there could be a doubt in the mind of any person carefully examining it with a power of 500 diameters that the cause of the spiral appearance was not a ... spiral fibre. In Areyria, threads of a different kind are present; #º : º § º ; hº Miſſillºw/ sInWº. FIG. 11.-Arcyria incarnata, with portion of threads and spore, magnified. they mostly branch and anastomose, and are externally furnished with prominent warts or spines, which Mr. Currey + holds are also arranged in a spiral manner around the threads. In other Myxogastres, threads are also present without any appreciable spiral markings or spines. In the mature condition of these fungi, they so clearly resemble, and have such close affinities with, the Trichogastres that one is led almost to doubt whether it was not on hasty grounds, without due examination or consideration, that proposals were made to remove them from the society of their kindred. Very little is known of the development of the spores in this group; in the early stages the whole substance is so pulpy, and in the latter so dusty, whilst the transition from one to * Currey, “On Spiral Threads of Trichia,” in “Quart. Journ. Micr Science " (1855), iii. p. 17. 34 FUNGI. the other is so rapid, that the relation between the spores and threads, and their mode of attachment, has never been definitely made out. It has been supposed that the spinulose projections Ǻyº Ö º § ; \ §§ § X25 X ;j ? () 2 & §3 § à º K $3 Q C ºº i i & ſº § () C É. º : CU d |º § ; ; ; § C C O NC º § |ë i ºſ º º § º | K. ſº ; ſ § É Q ; Q § U º : i.| § § K. º : C º § § : | | º from the capillitium in some species are the remains of pedicels from which the spores have fallen, but there is no evidence beyond this supposition in its favour, whilst on the other hand, in Stemonitis, for instance, there is a profuse interlacing capillitium, and no spines have been detected. In order to strengthen the supposition, spines should be more commonly present. The threads, or capillitium, form a beautiful reticulated network in Stemonitis, Cribra- ria, Diachaea, Dictydium, &c. In Spumaria, Reticu- laria, Lycogala, &c., they are almost obsolete.” In no group is the examination of the development of structure more difficult, for the reasons already alleged, than in the Myxogastres. NIDULARIACEI.—This small group departs in Some Fig. 12 pia- important particulars from the general type of struc- ** ture present in the rest of the Gasteromycetes.t The plants here included may be described under three parts, the mycelium, the peridium, and the sporangia. The mycelium FIG. 13.−Cyathws vermicosus. is often plentiful, stout, rigid, interlacing, and coloured, running over the surface of the soil, or amongst the vegetable débris on which the fungi establish themselves. The peridia are seated upon this mycelium, and in most instances are at length open above, taking the form of cups, or beakers. These organs consist of three strata of tissue vary- ing in structure, the external being fibrous, and sometimes hairy, the interior cellular and delicate, the inter- mediate thick and at length tough, coriaceous, and resistant. * In some of the genera, as, for instance, in Badhamia, Enerthenema, and Reticularia, the spores are produced within delicate cells or cysts, which are afterwards absorbed. t Tulasne, “Essai d'une Monographie des Nidulariées,” in “Ann. des Sci. Nat.” (1844), i. 41 and 64. STRUCTURE. 35 When first formed, the peridia are spherical, they then elongate and expand, the mouth being for some time closed by a veil, or diaphragm, which ultimately disappears. Within the cups lentil-shaped bodies are attached to the base and sides by elastic cords. These are the sporangia. Each of these has a com- plicated structure; externally there is a filamentous tunic, composed of interlaced fibres, sometimes called the peridiole; beneath this is the cortex, of compact homogenous structure, then follows a cellular thicker stratum, bearing, towards the centre of the sporangia, delicate branched threads, or sporo- phores, on which, at their extremities, the ovate spores are generated, some- times in pairs, but normally, it would seem that they are quaternary on spicules, the threads being true basidia. The whole structure is exceedingly interesting and peculiar, and may be studied in detail in Tulasne's memoir on this group. SPHERONEMEI.-In this very large and, within certain limits, variable order, there is but little of interest as regards struc- ture, which is not better illustrated else- where; as, for instance, some sort of peri- ºtº lºgº º sº thecium is always present, but this can phore, d. Spores. be better studied in the Sphaeriacei. The spores are mostly very minute, borne on delicate sporophores, which originate from the inner surface of the perithecia, but the majority of so-called species are undoubtedly conditions of sphaeriaceous fungi, either spermatogonia or pycnidia, and are of much more interest when studied in connection with the higher forms to which they belong.” Probably the number of complete. and autonomous species are very few. MELANCONIEI.—Here, again, are associated together a great number of what formerly were considered good species of fungi, but which are now known to be but conditions of other forms. * Berkeley, M. J., “Introduction, Crypt. Bot.” p. 330. 36 FUNGI. One great point of distinction between these and the preceding is the absence of any true perithecium, the spores being pro- duced in a kind of spurious receptacle, or from a sort of stroma. The spores are, as a rule, larger and much more attractive than in Sphaeronemei, and, in some instanceſ, arc cithel very ſine, or very curious. Under this head we may mention the multi- septate spores of Coryneum; the tri-radiate spores of Astero- * sporium ; the curious crested spores of Pestalozzia; the doubly crested spores of Dilophospora; and the scarcely less sin- gular gelatinous coated spores of Cheiro- spora. In all cases the fructification is abundant, and the spores frequently ooze Out in tendrils, or form a black mass above the spurious receptacle from which they issue.* Fia, is ºrium had TortAce1–In this order there seems at first to be a considerable resemblance to the Dematiei, except that the threads are almost obsolete, and the plant is reduced to chains of spores, without trace of perithe- cium, investing cuticle, or definite stroma. Sometimes the spores are simple, in other cases septate, and in Sporochisma are at first produced in an investing cell. In most cases simple threads at length become septate, and are ultimately differentiated into spores, which separate t the joints when fully mature. CAEOMACEI.-Of far greater interest are the Coniomycetous parasites on living plants. The present order includes those in which the spore f is reduced to a single cell; and here we may observe that, although many of them are now proved to be imperfect in themselves, and only forms or conditions of other fungals, we shall write of them here without regard to their duality. These originate, for the most part, within the tissues of living plants, and are developed outwards in pustules, which burst through the cuticle. The mycelium penetrates the inter- * Berkeley, M. J., “Introduction, Crypt. Bot.” p. 329. # In the Cabomace; and Puccinited the term “pseudospore ' would be much more accurate. STRUCTURE. 37 cellular passages, and may sometimes be found in parts of the plants where the fungus does not develop itself. There is no proper excipulum or peridium, and the spores spring direct from a more compacted portion of the mycelium, or from a cushion-like stroma of small cells. In Lecythea, the sub-globose spores are at first generated at the tips of short pedicels, from which they are ulti- mately separated; surrounding these spores arise a series of barren cells, 77/. or cysts, which are considerably larger Fig. 16–Barren cysts and pseudo- than the true spores, and colourless, Spores of Lecytheat. while the spores are of some shade of yellow or orange.* In Trichobasis, the spores are of a similar character, sub-globose, and at first pedicellate; but there are no surrounding cysts, and the colour is more usually brown, al- though sometimes yellow. In Uredo, the spores are at first generated singly, within a mother cell; they are globose, and either yellow or brown, without any pedicel. In Coleosporium, there are two kinds of spores, those of a pulverulent nature, globose, which are sometimes produced alone at the com- mencement of the season, and others Fig. 17.-Coleosporium Tussilagini, which originate as an elongated cell; Lev. this becomes septate, and ultimately separates at the joints. During the greater part of the year, both kinds of spores are to be found in the same pustule. In Melampsora, the winter spores are elongated and wedge-shaped, com- pacted together closely, and are only matured during winter on dead leaves; the summer spores are pulverulent and globose, being, in fact, what were until recently regarded FIG. 18.-Melampsora Salicina. * Léveillé, “Sur la Disposition Méthodique des Urédinées,” in “Ann. des Sci. Nat.” (1847), vol. viii. p. 369. - e 38 FUNGI. as species of Lecythea. In Cystopus, the spores are sub-globose, or somewhat angular, generated in a moniliform manner, and afterwards separating at the joints. The upper spore is always the oldest, continuous production of spores going on for some timue at the base uſ the claim. Under ſavourable conditions of moisture, each of these spores, or conidia, as De Bary terms them, is capable of producing within itself a number of zoospores; * these ultimately burst the vesicle, move about by the aid of vibratile cilia, and at last settle down to germinate. Besides these, other reproductive bodies are generated upon the mycelium, within the tissues of the plant, in the form of globose oogonia, or resting spores, which, when mature, also enclose great numbers of zoospores. Similar Oogonia are produced amongst the Mucedines in the genus Peronospora, to which De Bary con- siders Cystopus to be closely allied. At all events, Fig. 19.-Cystopus this is a peculiarity of structure and development * not as yet met with in any other of the Caomacei. In Uromyces is the nearest approach to the Pucciniai ; in fact, it is Puccinia reduced to a single cell. The form of spore is usually more angular and irregular than in Trichobasis, and the pedicel is permanent. It may be remarked here, that of the foregoing genera, many of the species are not autonomous that have hitherto been included amongst them. This is especially true of Lecythea, Trichobasis, and, as it now appears, of Uromyces.t - PUCCINIAEI.—This group differs from the foregoing chiefly in having septate spores. The pustules, or sori, break through the cuticle in a similar manner, and here also no true peridium is present. In A enodochus, the highest development of joints is reached, each spore being composed of an indefinite number, from ten to twenty cells. With it is associated an unicellular * De Bary, “Champignons Parasites,” in “Ann. des Sci. Nat.” 4me sér. vol. xx. + Tulasne, “Mémoire sur les Urédinées, &c.,” in “Ann. des Sci. Nat.” (1854), vol. ii. p. 78. STRUCTURE. 39 yellow Uredine, of which it is a condition. Probably, in every species of the Pucciniaei, it may hereafter be proved, as it is now suspected, that an unicellular Uredine precedes or is associated with it, forming a condition, or secondary form of fruit of that species. Many instances of that kind have already been traced by De Bary,” Tulasne, and others, and some have been a little too rashly surmised by their followers. In Phragmidium, the pedicel is much more elongated than in Xenodochus, and the spore is shorter, with fewer and a more definite number of cells for each species; Mr. Currey is of opinion that each cell of the spore in Phragmidium has an inner globose cell, Fig. 20-xenodochus car which he caused to escape by rupture of the bonariws. outer cell wall as a sphaeroid nucleus,f leading to the inference that each cell has its own individual power of germination and reproduction. In Triphragmium, there are three cells for each spore, two being placed side by side, and one superimposed. In one species, however, Triphragmium deglubens (North American), the cells are arranged as in Phragmidium, so that this represents really a tricellular Phragmidium, linking the pre- sent with the latter genus. In Puccinia the number of species is by far the most numerous; in this genus the spores are uni- septate, and, as in all the Pucciniaei, the peduncles are permanent. There is great variability in the compactness of the spores in the Sori, or pulvinules. In some species, the sori are so pulverulent that the spores Flo ºn.” are as readily dispersed as in the Uredines, in others they are so compact as to be separated from each * De Bary, “Ueber die Brandpilze,” Berlin, 1853. • + Currey, in “Quart. Journ. Micr. Sci.” (1857), vol. v. p. 119, p. 8, fig 13. 40 FUNGI. other with great difficulty. As might be anticipated, this has considerable effect on the contour of the spores, which in pul- verulent species are shorter, broader, and more ovate than in the compact species. If a section of one of the more compact sori be made, it will be seen that the majority of the spores are side by side, nearly at the same level, their apices forming the external surface of the sori, but it will not be unusual to observe smaller and younger spores pushing up from the Fig. 22–Pseudospores hymenial cells, between the peduncles of of Paccinia. the elder spores, leading to the inference that there is a succession of spores produced in the same pulvi- nule. In Podisoma, a rather anomalous genus, the septate spores are immersed in a gelatinous stratum, and some authors have imagined that they have an affinity with the Tremellini, but this affinity is more apparent than real. The phenomena of germination, and their relations to Raestelia, if substantiated, establish their claim to a position amongst the Pucciniai.” It seems to us that Gymnosporangium does not differ generically from Podisoma. In a recently-characterized species, Podisoma I'llisii, the spores are bi-triseptate. This is, moreover, peculiar from the great deficiency in the gelatinous element. In another North American species, called Gymnosporangium biseptatum, Ellis, which is distinctly gelatinous, there are similar biseptate spores, but they are considerably broader and more obtuse. In other described species they are uniseptate. USTILAGINEI.—These fungi are now usually treated as distinct from the Caomacei, to which they are closely related. They are also parasitic on growing plants, but the spores are usually black or sooty, and never yellow or orange; on an average much smaller than in the Caeomacei. In Tilletia, the spores are spherical and reticulated, mixed with delicate threads, from * Cooke, “On Podisoma,” in “Journal of Quekett Microscopical Club,” vol. ii. p. 255. ºf Tulasne, “Mémoire sur les Ustilaginées,” in “Ann. des Sci. Nat.” (1847), vii. pp. 12 and 73. STRUCTURE. 41 whence they spring. In the best known species, Tilletia caries, they constitute the “bunt" of wheat. The peculiarities of germination will be alluded to hereafter. In Ustilago, the minute sooty spores are developed either on delicate threads or in compacted cells, arising first from a sort of semi-gelati- nous, grumous stroma. It is very difficult to detect any threads associated with the spores. The species attack the flowers and anthers of composite and polygonaceous plants, the leaves, culms, and germen of grasses, &c., and are popularly known as “smuts.” In Urocystis and Thecaphora, the spores are united together into sub-globose bodies, form- ing a kind of compound spore. In some species of Urocystis, the union which subsists between them is com- paratively slight. In Thecaphora, on the contrary, the complex spore, or agglomeration of spores, is compact, being at first apparently enclosed in a delicate cyst. In Tubur- cinia, the minute cells are compacted into a hollow sphere, having lacunae communicating with the interior, and often exhi- biting the remains of a pedicel. AECIDIACEI.-This group differs from the foregoing three groups prominently in the presence of a cellular peridium, which encloses the spores; hence some mycologists have not hesitated to propose their association with the Gasteromycetes, although every other feature in their structure seems to indicate a close affinity with the Caeomacei. The pretty cups in the genus AEcidium are sometimes scat- tered and sometimes collected in clus- ters, either with spermogonia in the centre or on the opposite surface. The cups are usually white, composed of regularly arranged bordered cells at length bursting at the apex, with the margins turned back and split into radiating teeth. The spores are commonly of a bright orange or golden yellow, sometimes white or brownish, and are produced in chains, or moniliform FIG. 23. – Thecaphora hyalina. FIG. 24.—AEcidium Berberidis. 42 FUNGI. strings, slightly attached to each other,” and breaking off at thc summit at the same time that they continue to be produced at the base, so that for some time there is a successive production of spores. The spermogonia are not always readily detected, as they are much smaller than the peridia, and sometimes precede them. The spermatia are expelled from the lacerated and fringed apices, and are very minute and colourless. In Raestelia the peridia are large, growing in company, and splitting longi- tudinally in many cases, or by a lacerated mouth. In most in- stances, the spores are brownish, but in a splendid species from North America (Roºstelia aurantiaca, Peck), recently charac- terized, they are of a bright orange. If GErsted is correct in his observations, which await confirmation, these species are all related to species of Podisoma as a secondary form of fruit.f In the Roºstelia of the pear-tree, as well as in that of the moun- tain ash, the spermogonia will be found either in separate tufts on discoloured spots, or associated with the Raesfelia. In Peri- dermium there is very little structural difference from Raesfelia, and the species are all found on coniferous trees. In Fndo- phyllum, the peridia are immersed in the succulent substance of the matrix; whilst in Graphiola, there is a tougher and withal double peridium, the inner of which form a tuft of erect threads resembling a small brush.; IIYPHOMYCETES.–The predominant feature in the structure of this order has already been intimated to consist in the develop- ment of the vegetative system under the form of simple or branched threads, on which the fruit is generated. The common name of mould is applied to them perhaps more generally than to other groups, although the term is too vague, and has been too vaguely applied to be of much service in giving an idea of the characteristics of this order. Leaving the smaller groups, and confining ourselves to the Dematiei and the Mucedines, we * Corda, “Icones Fungorum,” vol. iii. fig. 45. + Cooke, “On Podisoma,” in “Quekett Journal,” vol. ii. p. 255. # It may be a question whether Graphiola is not more nearly allied to Trichocoma (Jungh Fl. Crypt. Javae, p. 10, f. 7) than to the genera with which it is usually associated —M. J. B. STRUCTURE. 43 shall obtain some notion of the prevalent structure. In the former the threads are more or less carbonized, in the latter nearly colourless. One of the largest genera in Dematiei is Helminthosporium. It appears on decaying herbaceous plants, and on old wood, forming effused black velvety patches. The mycelium, of coloured jointed threads, overlays and penetrates the matrix ; from this arise erect, rigid, and usually jointed threads, of a dark brown, nearly black colour - at the base, but paler towards the apex. In most cases these threads have an externally cortical layer, which imparts rigidity; usually from the apex, but sometimes laterally, the spores are produced. Although some- times colourless, these are most com- monly of some shade of brown, more or less elongated, and divided trans- versely by few or many septa. In Helminthosporium Smithii, the spores much exceed the dimensions of the threads; * in other species they are smaller. In Dendryphium, the threads and spores are very similar, except that the threads are branched at their apex, and the spores are often pro- duced one at the end of another in a short chain.t In Septosporium again, the threads and spores are similar, but the spores are pedicellate, and at- tached at or near the base; whilst in Acrothecium, with similar threads and spores, the latter are clustered together at the apex of the threads. In Triposporium, the threads are similar, but the spores are tri-radiate ; and in Helicoma, the spores are twisted spirally. Thus, we might pass FIG. 25.---Helminthosporium molle. * Cooke, “On Microscopic Moulds,” in “Quekett Journal,” vol. ii. plate 7. f See “Dendryphium Fumosum,” in “Quekett Journal,” vol. ii. plate 8; or, “Corda Prachtflora,” plate 22. 44 FUNGT. through all the genera to illustrate this chief feature of coloured, septate, rather rigid, and mostly erect threads, bearing at some point spores, which in most in- stances are elongated, coloured, and septate. MUCEDINEs.—Here, on the other hand, the threads, if coloured at all, are still delicate, more flexuous, with much thinner walls, and never invested with an external cortical layer. One of the most important and highly developed genera is Peronospora, the members of which are parasitic upon and destructive of living vegetables. It is to this genus that the mould of the too famous potato disease belongs. Professor De Bary has done more than any other mycologist in the investigation and eluci- Q9 % FIG. 27.-Peronospora Arenariae. Bary terms them—conidia. dation of this genus; and his mono- graph is a masterpiece in its way.* He was, however, preceded by Mr. Berkeley, and more especially by Dr. Montagne, by many years in eluci- dation of the structure of the flocci and conidia in a number of species # In this genus, there is a delicate mycelium, which penetrates the in- tercellular passages of living plants, giving rise to erect branched threads, which bear at the tips of their ultimate ramuli, sub-globose, ovate, or elliptic spores, or, as De Deeply seated on the mycelium, within the substance of the foster plant, other reproductive bodies, called oogonia, originate. These are spherical, more or * e º * e $ * De Bary, “Champignons Parasites,” in “Ann. des Sci. Nat.” 4me sér. vol. xx. + Berkeley, “On the Potato Murrain,’ in “Journ. of IIort. Soc. of London,” vol. i. (1846), p. 9. STRUCTURE. " 45 less warted and brownish, the contents of which become dif. ferentiated into vivacious zoospores, capable, when expelled, of moving in water by the aid of vibratile cilia. A similar struc- ture has already been indicated in Cystopus, otherwise it is rare in fungi, if the Saprolegniei be excluded. In Botrytis and in Polyactis, the flocci and spores are similar, but the branches of the threads are shorter and more compact, and the septa are more common and numerous; the oogonia also are absent. De Bary has selected Polyactis cinerea, as it occurs on dead vine leaves, to illustrate his views of the dual- ism which he believes himself to have discovered in this species. “It spreads its mycelium in the tissue which is becom- ing brown,” he writes, “and this shows at first essentially the same construc- tion and growth as that of the mycelium filaments of Aspergillus.” On the my- Celium soon appear, besides those which are spread over the tissue of the leaves, strong, thick, mostly fasciculate branches, which stand close to one another, break- ing forth from the leaf and rising up per- pendicularly, the conidia-bearers. They grow about 1 mm. long, divide them- selves, by successively rising partitions, into some prominent cylindrical linked cells, and then their growth is ended, / and the upper cell produces near its a' point three to six branches almost stand- . Fºrſ: ºre. ing rectangularly. Of these the under ones are the longest, and they again shoot forth from under their ends one or more still shorter little branches. The nearer they are to the top, the shorter are the branches, and less divided ; the upper ones are quite branchless, and their length scarcely exceeds the breadth of the principal stem. Thus a system of branches appears, upon which, on a small scale, a bunch of grapes is represented. All the twigs soon end their 46 FUNGI. growth ; they all separate their inner space from the principal stem, by means of a cross partition placed close to it. All the ends, and also that of the principal stem, swell about the same time something like a bladder, and on the upper free half of each swelling appear again, simultaneously, several fine pro- tuberances, close together, which quickly grow to little oval bladders filled with protoplasm, and resting on their bearers with a sub-sessile, pedicellate, narrow basis, and which at length separate themselves through a partition as in Aspergillus. The detached cells are the conidia of our fungus; only one is formed on each stalk. When the formation is completed in the whole of the panicle, the little branches which compose it are de- prived of their protoplasm in favour of the conidia; it is the same with the under end of the principal stem, the limits of which are marked by a cross partition. The delicate wall of these parts shrinks up until it is unrecognizable ; all the conidia of the panicle approach one another to form an irregular grape- like bunch, which rests loosely on the bearer, and from which it easily falls away as dust. If they be brought into water they fall off immediately; only the empty, shrivelled, delicate skins are to be found on the branch which bore them, and the places on which they are fixed to the principal stem clearly appear as round circumscribed hilums, generally rather arched towards the exterior. The development of the main stem is not ended here. It remains solid and filled with protoplasm as far as the portion which forms the end through its conidia. Its end, which is to be found among these pieces, becomes pointed after the ripening of the first panicle, pushes the end of the shrivelled member on one side, and grows to the same length as the height of one or two panicles, and then remains still, to form a second panicle similar to the first. This is later equally per- foliated as the first, then a third follows, and thus a large number of panicles are produced after and over one another on the same stem. In perfect specimens, every perfoliated panicle hangs loosely to its original place on the surface of the stem, until by shaking or the access of water to it, it falls immediately into the single conidia, or the remains of branches, and the STRUCTURE. 47 already-mentioned oval hilums are left behind. Naturally, the stem becomes longer by every perfoliation; in luxuriant speci- mens the length can reach that of some lines. Its partition is already, by the ripening of the first panicle from the beginning of its foundation, strong and brown; it is only colourless at the end which is extending, and in all new formations. During all these changes the filament remains either unbranched, except as regards the transient panicles, or it sends out here and there, at the perfoliated spots, especially from the lower ones, one or two strong branches, standing opposite one another and resembling the principal stem. The mycelium, which grows so exuberantly in the leaf, often brings forth many other productions, which are called sclerotia, and are, according to their nature, a thick bulbous tissue of mycelium filaments. Their formation begins with the profuse ramification of the mycelium threads in some place or other; generally, but not always, in the veins of the leaf; the inter- twining twigs form an uninterrupted cavity, in which is often enclosed the shrivelling tissue of the leaf. The whole body swells to a greater thickness than that of the leaf, and protrudes on the surface like a thickened spot. Its form varies from circular to fusiform ; its size is also very unequal, ranging between a few lines and about half a millimetre in its largest diameter. At first it is colourless, but afterwards its outer layers of cells become round, of a brown or black colour, and it is surrounded by a black rind, consisting of round cells, which separate it from the neighbouring tissue. The tissue within the rind remains colourless; it is an entangled uninterrupted tissue of fungus filaments, which gradually obtain very solid, hard, cartilaginous coats. . The sclerotium, which ripens as the rind becomes black, loosens itself easily from the place of its forma- tion, and remains preserved after the latter is decayed. The sclerotia are, here as in many other fungi, biennial organs, designed to begin a new vegetation after a state of apparent quietude, and to send forth special fruit-bearers. They may in this respect be compared to the bulbs and perennial roots of under shrubs. The usual time for the development of 48 FUNGT. the sclerotia is late in the autumn, after the fall of the vine leaves. As long as the frost does not set in, new ones continu- ally spring up, and each one attains to ripeness in a few days. If frost appears, it can lie dry a whole year, without losing its power of development. This latter commences when the sclerotium is brought into contact with damp ground during the usual temperature of our warmer seasons. If this occur soon, at the latest some weeks after it is ripe, new vegetation grows very quickly, generally after a few days; in several parts the colourless filaments of the inner tissue begin to send out clusters of strong branches, which, breaking through the black rind, stretch themselves up perpendicularly towards the surface, separate from one another, and then take all the characteristics of the conidia-bearers. Many such clusters can be produced on one Sclerotium, so that soon the greater part of the surface is covered by filamentous conidia- bearers with their panicles. The colourless tissue of the sclerotium disappears in the same degree as the conidia-bearers grow, and at last the black rind remains behind empty and shrivelled. If we bring, after many months, for the first time, the ripe sclerotium, in damp ground, in summer or autumn, after it has ripened, the further *ś. development takes place In OPé slowly, and .*.*.eu... in an essentially different form. It is true ridia. that from the inner tissue numerous fila- mentous branches shoot forth at the cost of this growing fascicle, and break through the black rind, but its filaments remain strongly bound, in an almost parallel situation, to a cylindrical cord, which for a time lengthens itself and spreads out its free end to a flat plate-like disc. This is always formed of strongly united threads, ramifications of the cylindrical cord. On the free upper surface of the disc, the filaments shoot forth innumerable branches, which, growing to the same height, thick and parallel with one another, cover the before-named disc. STRUCTURE. 49 Some remain narrow and cylindrical, are very numerous, and produce fine hairs (paraphyses); others, also very numerous, take the form of club-like ampulla cells, and each one forms in its interior eight free swimming oval sporos. Those ampulla cells are sporidiiferous asci. After the spores have become ripe, the free point of the utricle bursts, and the spores are scattered to a great distance by a mechanism which we will not here further describe. New ampullas push themselves between those which are ripening and withering; a disc can, under favourable circum- stances, always form new asci for weeks at a time. The num- ber of the already described utricle-bearers is different, accord- ing to the size of the sclerotium ; smaller specimens usually produce only one, larger two to four. The size is regulated by that of the sclerotia, and ranges, in full-grown specimens, between one and more millimetres for the length of the stalk, and a half to three (seldom more) millimetres for the breadth of the disc.* For some time the conidia form, belonging to the Mucedines, has been known as Botrytis cinerea (or Polyactis cinerea). The compact mycelium, or sclerotium, as an im- perfect fungus, bore the name of Sclerotium echinatum, whilst to the perfect and cup-like form has been given the name of Peziza Puckeliana. We have reproduced De Bary's life-history of this mould here, as an illustration of structure in the Mucedines, but hereafter we shall have to write of similar transformations when treating of polymorphism. - The form of the threads, and the form and disposition of the spores, vary according to the genera of which this order is com- posed. In Oidium the mostly simple threads break up into joints. Many of the former species are now recognized as con- ditions of Erysiphe. In Aspergillus, the threads are simple and erect, with a globose head, around which are clustered chains of simple spores. In Penicillium, the lower portion of the threads is simple, but they are shortly branched at the apex, the branches being terminated by necklaces of minute spores. In Dactylium, * De Bary, “On Mildew and Fermentation,” p. 25, reprinted from “German Quarterly Magazine,” 1872; De Bary, “Morphologie und Physiologie der Pilze,” (1866), 201 50 FUNGI. the threads are branched, but the spores are collected in clusters usually, and are moreover septate. In other genera similar distinctions prevail. These two groups of black moulds and white moulds are the noblest, and contain the largest muuibel of genera aud speuies amongst the Hyphomycetes. There is, how- ever, the small group of Tsariacei, in which the threads are compacted, and a semblance . of such hymenomycetal forms as Clavaria and Pterula is the result, but it is doubtful if this group contains many autonomous species. In another small group, the Stil- bacei, there is a composite character in the head, or receptacle,” and in the stem when the latter is present. Many of these, again, as Tubercularia, Volutella, Fusarium, &c., - contain doubtful species. In Sepedoniei and Fig. 30, Peniçilliºn char. Trichodermacei, the threads are reduced to a tarwmv, Cooke. tº ſº minimum, and the spores are such a dis- tinctive element that through these groups the Hyphomycetes are linked with the Coniomycetes. These groups, however, are not of sufficient size or importance to demand from us, in a work of this character, anything more than the passing allusion which we have given to them. We come now to consider the structure in the Sporidiifera, in which the fructifying corpuscles or germs, whether called spores or sporidia, are generated within certain privileged cysts, usually in definite numbers. In systematic works, these are included under two orders, the Physomycetes and the Ascomycetes. The former of these consists of cyst-bearing moulds, and from their nearest affinity to the foregoing will occupy the first place. PHYSOMYCETEs include, especially amongst the Mucorini, many most interesting and instructive species for study, whigh even very lately have occupied the attention of continental mycolo- gists. Most of these phenomena are associated more or less with reproduction, and as such will have to be adverted to again, * Cooke, ‘‘Handbook of British Fungi,” vol. ii. p. 552. STRUCTURE. 51 but there are points in the structure which can best be alluded to here. Again taking Professor de Bary's researches as our guide,” we will illustrate this by the common Mucor mucedo: If we bring quite fresh horse-dung into a damp confined atmosphere, for example, under a bell-glass, there appears on its surface, after a few days, an immense white mildew. Upright strong filaments of the breadth of a hair raise themselves over the surface, each of them soon shows at its point a round little head, which gradually becomes black, and a closer examination shows us that in all principal points it perfectly agrees with the sporangia of other species. Each of these white filaments is a sporangia-bearer. They spring from a mycelium which is spread in the dung, and appear singly upon it. Cer- tain peculiarities in the form of the sporangium, and the little long cylindrical spores, which, when examined sepa- rately, are quite flat and co- lourless, are characteristic of - the species. If the latter be º S sown in a suitable medium, for example, in a solution of % ple, FIG. 31.-Mucor ºwcedo, with three sporangia. Sugar, they swell, and shoot a. Portion of frill with sporangiola. forth germinating utricles, which quickly grow to mycelia, which bear sporangia. This is easily produced on the most various organic bodies, and Mucor mucedo is therefore found sponta- neously on every substratum which is capable of nourishing mildew, but on the above-named the most perfect and exuberant specimens are generally to be found. The sporangia-bearers are at first always branchless and without partitions. After the sporangium is ripe, cross partitions in irregular order and number often appear in the inner space, and on the upper * De Bary, “On Mildew and Fermentation,” in “Quarterly German Magazine,” or 1872. 52 FUNGI. surface branches of different number and size, each of which forms a sporangium at its point. The sporangia which are formed later are often very similar, but sometimes very different, to those which first appeared, because their partition is very thick and does not fall to pieces when it is ripe, but irregu- larly breaks off, or remains entire, enclosing the spores, and at last falls to the ground, when the fungus withers. The cross partition which separates the sporangia from its bearers is in those which are first formed (which are always relatively thicker sporangia) very strongly convex, while those which follow later are often smaller, and in little weak specimens much less arched, and sometimes quite straight. After a few days, similar filaments generally show themselves on the dung between the sporangia- bearers, which appear to the naked eye to be provided with deli- cate white frills. Where such an one is to be found, two to four rectangular expanding little branches spring up to the same height round the filament. Each of these, after a short and simple process, branch out into a furcated form; the furcations being made in such a manner that the ends of the branch at last so stand together that their surface forms a ball. Finally, each of the ends of a branch swells to a little round sporangium, which is limited by a partition (called sporangiolum, to distin- guish it from the larger ones), in which some, generally four, spores are formed in the manner already known. When the sporangiola are alone, they have such a peculiar appearance, with their richly-branched bearers, that they can be taken for some- thing quite different to the organs of the Mucor mucedo, and were formerly not considered to belong to it. That they really belong to the Mucor is shown by the principal filament which it bears, not always, but very often, ending with a large sporan- gium, which is characteristic of the Mucor mucedo; it is still more evident if we sow the spores of the sporangiolum, for, as it germinates, a mycelium is developed, which, near a simple bearer, can form large sporangia, and those form sporangiola, the first always considerably preponderating in number, and very often exclusively. If we examine a large number of speci- mens, we find every possible middle form between the simple STRUCTURE. 53 or less branched sporangia-bearers and he typical sporangiola frills; and we arrive at last at the conclusion simply to place the latter among the varieties of form which the sporangia-bearer of the Mucor mucedo shows, like every other typical organic form within certain limits. On the other hand, propagation organs, differing from those of the sporangia and their products, belong to Mucor mucedo, which may be termed conidia. On the dung (they are rare on any other substance) these appear at the same time, or generally somewhat later, than the sporangia- bearers, and are not unlike those to the naked eye. In a more accurate examination, they appear different; a thicker, partition- less filament rises up and divides itself, generally three-forked, at the length of one millimetre, into several series of branchlets. The forked branches of the last series bear under their points, which are mostly capillary, short erect little ramuli, and these, with which the ends of the principal branches ar- ticulate on their somewhat broad tops, several spores and conidia, near one another; about fifteen to twenty are ------> formed at the end of each little ra- - -— mulus. The peculiarities and varia- ~ tions which so often appear in the Fig. 32.--Small portion of Botrytis g tº e Jonesii. ramification need not be discussed here. After the articulation of the conidia, their bearers sink together by degrees, and are quite destroyed. The ripe conidia are round like a ball, their surface is scarcely coloured, and almost wholly smooth. These conidioid forms were at first described as a separate species under the name of Botrytis Jonesii. How, then, do they belong to the Mucor # * That they appear grega- riously is as little proof of an original relation to one another, here as elsewhere. Attempts to prove that the conidia and spo- rangia-bearers originate on one and the same mycelium filament may possibly hereafter succeed. Till now this has not been the case, * We are quite aware that Von Tieghem and Le Monnier, in “Ann, des Sci. Nat.” 1873, p. 335, dispute that this belongs to Mucor mucedo, and assert that Chaetocladium Jonesii is itself a true Mucor, with monosporous Sporangla. 54 - FUNGI. and he who has ever tried to disentangle the mass of filaments which exuberantly covers the substratum of a Mucor vegetation, which has reached so far as to form conidia, will not be surprised that all attempts have hitherto proved abortive. The suspicion of the connection founded on the gregariously springing up, and external resemblance, is fully justified, if we sow the conidia in a suitable medium, for example, in a solution of sugar. They here germinate and produce a mycelium which exactly re- sembles that of the Mucor mucedo, and, above all, they pro- duce in profusion the typical sporangia of the same on its bearers. The latter are till now alone reproductions of conidia- bearers, and have never been observed on mycelia which have grown out of conidia. These phenomena of development appear in the Mucor when it dwells on a damp substance, which must naturally contain the necessary nourishment for it, and is exposed to the atmo- spheric air. Its mycelium represents at first strong branched utricles without partitions; the branches are of the higher order, mostly divided into rich and very fine-pointed ramuli. In old mycelium, and also in the sporangia-bearers, the contents of which are mostly used for the formation of spores, and the substratum of which is exhausted for our fungus, short stationary pieces, filled with protoplasm, are very often formed into cells through partitions in order to produce spores, that is, grow to a new fruitful mycelium. These cells are called gemmules, brooding cells, and resemble such vegetable buds and sprouts of foliaceous plants which remain capable of develop- ment after the organs of vegetation are dead, in order to grow, under suitable circumstances, to new vegetating plants, as, for example, the bulbs of onions, &c. If we bring a vegetating mycelium of Mucor mucedo into a medium which contains the necessary nourishment for it, but excluded from the free air, the formation of sporangia takes place very sparingly or not at all, but that of gemmules is very abun- dant. Single interstitial pieces of the ramuli, or even whole systems of branches, are quite filled with a rich greasy proto- plasm; the short pieces and ends are bound by partitions which STRUCTURE, -. 55 form particular, often fun-like or globular cells; the longer ones are changed, through the formation of cross partitions, into chains of similar cells; the latter often attain by degrees strong, thick walls, and their greasy contents often pass into innumerable drops of a very regular globular form and of equal size. Similar appearances show themselves after the sowing of spores, which are capable of germinating in the medium already described, from which the air is excluded. Either short germinating utricles shoot forth, which soon form themselves into rows of gemmules, or the spores swell to large round bladders filled with protoplasm, and shoot forth on various parts of their surface innumerable protuberances, which, fixing themselves with a narrow basis, soon become round vesiculate cells, and on which the same sprouts which caused their production are re- peated, formations which remind us of the fungus of fermenta- tion called globular yeast. Among all the known forms of gemmules we find a variety which are intermediate, all of which show, when brought into a normal condition of development, the same proportion, and the same germination, as those we first described. - - We have detailed rather at length the structure and develop- ment of one of the most common of the Mucors, which will serve as an illustration of the order. Other distinctions there may be which are of more interest as defining the limits of genera, except such as may be noticed when we come to writo more specially of reproduction. ASCOMYCETES.—Passing now to the Ascomycetes, which are especially rich in genera and species, we must first, and but super- ficially, allude to Tuberacei, an order of sporidiiferous fungi of subterranean habit, and rather peculiar structure.” In this order an external stratum of cells forms a kind of perithecium, which is more or less developed in different genera. This encloses the hymenium, which is sinuous, contorted, and twisted, often forming lacunae. The hymenium in some genera consists of elongated, nearly cylindrical asci, enclosing a definite number of sporidia; in the true truffles and their immediate allies, the asci are broad * Wittadini, “Monographia Tuberacearum,” 1831. 56 FUNGI, sacs, containing very large and beautiful, often coloured, sporidia. These latter have either a smooth, warted, spinulose, or lacunose epispore, and, as will be seen from the figures in Tulasne's Monograph, * or those in the last volume of Corda’s great work,+ are attractive microscopical objects. In some cases, it is not difficult to detect paraphyses, but in others they would seem to be entirely absent. A comparatively large number have been discovered and recorded in Great Britain,; but of these none are more suitable for study of general structure than the ordi- mary truffle of the markets. The structure of the remaining Ascomycetes can be studied under two groups, i.e., the fleshy Ascomycetes, or, as they have been termed, the Discomycetes, and the hard,or carbonaceous Asco- mycetes, sometimes called the Pyrenomycetes. Neither of these names gives an accurate idea of the distinctions between the two groups, in the former of which the discoid form is not universal, and the latter contains somewhat fleshy forms. But in the Dis- comycetes the hymenium soon becomes more or less exposed, and in the latter it is enclosed in a perithecium. The Discomy- cetes are of two kinds, the pileate and the cup-shaped. Of the pileate such a genus as Gyromitra or Helvella is, in a certain sense, analogous to the Agarics amongst Hymenomycetes, with a superior instead of an inferior hymenium, and enclosed, not naked, spores. Again, Geoglossum is somewhat analogous to Clavaria. Amongst the cup-shaped, Peziza is an Ascomycetous Cyphella. But these are perhaps more fanciful than real analogies. Recently Boudier has examined one group of the cup-shaped Discomycetes, the Ascobolei, and, by making a somewhat free use of his Memoir, $ we may arrive at a general idea of the struc- ture in the cupulate Discomycetes. They present themselves at * Tulasne, “Fungi Hypogaei,” 1851. + Corda, “Icones Fungorum,” vol. vi. † Berkeley and Broome, in “Ann. of Nat. Hist.” 1st ser, vol. xviii. (1846), p. 73; Cooke, in “Seem. Journ. Bot.” § Boudier (E.), “Mémoire sur les Ascobolés,” in “Ann, des Sci. Nat.” 5me sér. vol. x. (1869). STRUCTURE. 57 first under the form of a small rounded globule, and almost entirely cellular. This small globule, the commencement of the receptacle, is not long in increasing, preserving its rounded form up to the development of the asci. At this period, under the influence of the rapid growth of these organs, it soon produces at its summit a fissure of the external mem- brane, which becomes a more marked depression in the mar- ginate species. The receptacle thus formed increases rapidly, becomes plane, more convex, or more or less undulated at the margin, if at all of large size. Fixed to the place where it is generated by some more or less abundant mycelioid filaments, the receptacle becomes somewhat cup-shaped and either stipitate or sessile, composed of the receptacle proper and the hymenium. 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