fo) > 5 VERS 8 1761 3 Digitized by the Internet Archive in 2007 with funding from | Microsoft Corporation a as ' Al) A ee ae __ http:/www.archive.org/details/diseasesofplantsO0tubeuoft , Ar Rig * DISEASES OF PLANTS INDUCED BY CRYPTOGAMIC PARASITES i +. DISEASES OF PLANTS Induced by Cryptogamic Parasites Introduction to the Study of Pathogenic Fungi, Sliime-Fungt, Bacteria, & Algae BY Dr. KARL FREIHERR von TUBEUF PRIVATDOZENT IN THE UNIVERSITY OF MUNICH English Edition bp WILLIAM G. SMITH, B.Sc., PH.D. LECTURER ON PLANT PHYSIOLOGY UNIVERSITY OF EDINBURGH THREE HUNDRED AND THIRTY ILLUSTRATIONS LONGMANS, GREEN ©& CO. LONDON, NEW YORK, AND BOMBAY 1897 All rights reserved 2c * , & em ® Pr ‘ oF * ‘ are . GUASGOW Oe gic h Sy ee” whey Map? ta ie Printed at the Anibersity Press by eee ROBERT MACLEHOSE AND CO. ; . %. AUTHOR’S PREFACE, In my research work, and in connection with my lectures at the University and Technical School of Munich, I have for some time felt convinced that there existed a very evident gap in the literature relating to the diseases of plants. There was need of a newer and more complete work on cryptogamic para- sites and the diseases induced by them on higher plants, a work furnished with many accurate illustrations, with a survey of the newer literature, and with a general part wherein parasitism and the relations between parasite and host are discussed from a botanical standpoint. Therefore, I have undertaken to write a book intended to supply in’ some degree this pressing want. Here the attempt has been made for the first time to review in a general and comparative manner the biological, physiological, and anatomical relationships accompanying the phenomena of parasitism. Already De Bary has considered the varying degrees of parasitism and the phenomena of symbiosis in his celebrated Morphology and Biology of the Fungi; while Wakker has laid the foundations of our knowledge of the alterations in the anatomy of plants diseased by the agency of fungi, more especially, however, those alterations accompanying ‘hypertrophy.’ I venture to continue this difficult and comprehensive chapter of plant physiology, because for ten years I have devoted my time to the study of plant pathology. The book may be all the more acceptable since I have confirmed a large number of the observations and added the results of my own investigations, many of them now published for the first time. The present time is favourable to my work. The great Sylloge Fungorum of Saccardo (with its appendices in Vols. IX. and x.) has been recently completed; the classic investigations vi AUTHOR’S PREFACE. of Brefeld in the domain of mycology, and containing his classification of the fungi, are now well advanced; the Krypto- gamen-Flora of Rabenhorst is nearly completed; and the newer literature and observations are now periodically reviewed in the Zeitschrift fiir Pflanzenkrankheiten, and other magazines. The recent publication of several investigations on the influence of parasites on the anatomy of their host-plants greatly facilitated the compilation of the general part of the work. [ have here attempted to summarize in a systematic manner the preventive and combative agencies available against the more important diseases of economic plants. In many cases these are supported by facts given in the chapters on the natural and artificial infection of host-plants, and their disposition towards diseases produced by lower organisms. As already indicated in the title-page, the book deals only with those diseases of plants produced by the cryptogams and other lower organisms of the vegetable kingdom. The large number of parasites which attack such lower plants as algae and lichens, although not altogether neglected, have as a rule been omitted, otherwise the book could not have been brought within the limits of a single volume. In the second or systematic part of the book, the pathological phenomena are considered along with the description of the organism producing them. Where the diseases are of economic importance, measures for pre- vention and extermination are also suggested. Notices of greater length are given to such parasites and diseases as have formed the subject of special investigations. We could only aim at a complete list for Germany and the neighbouring countries, yet we have included many species of interest occurring in other parts of the world, notably in America. Though it will be possible to identify most of the more important parasites by the aid of this book, we do not intend it to replace the systematic works; we purpose rather to add to the descriptions given in Rabenhorst, Saccardo, and similar works. This book is intended above all to be, in the terms of its title-page, “an Introduction”; hence it seeks to orient in a general way, to give a summary of our knowledge, and to indicate the way to more detailed records. On this account great care has been taken in the citation of home and foreign literature, not only up to the time of finishing the manuscript AUTHOR’S PREFACE. vil (Easter, 1894), but also during the time of proof-reading up till the following Christmas. Reference to the book will be rendered easier by the numerous illustrations, which are almost exclusively the work of the author, and reproduced either from drawings or from photographs of the living objects, in many cases taken in situ. I consider it more essential to illustrate the habitus of pathological objects rather than to give drawings of microscopic subjects; those one may find in other works. Some of the illustrations are copied from the excellent plates of Tulasne, Woronin, De Bary, Klebs, Reess, Cohn, and Robert Hartig; while a number of woodcuts have been borrowed from the well-known Lehrbuch der Bawmkrank- heiten of the last named author. The grouping of the ‘Fungi imperfecti, which have not yet been worked up for the German flora, is based on Saccardo’s Sylloge; hence the arrangement into Hyalosporae, ete., which is intended for the benefit of those having access to Saccardo. Particular attention has been paid in the two Indices to the scientific names of both parasites and hosts, to popular names, and to technical expressions. In my labours I received great assistance from the following sources: From the collection of pathological material begun by Professor Robert Hartig, and now carried on with my help in the Botanical Institute of the Royal Bavarian School of Forestry in Munich; from the facilities for research and photography afforded by the laboratories of the same institution; from the Royal Library of Munich, the Library of the University, and the private pathological library of Professor Hartig. Living material for investigation has been kindly sent to me from many sources, particularly from the following gentlemen: Herr Lehrer Schnabl of Munich, Geh. Oberregierungsrath Prof. Kiihn of Halle, Hofgiirtner Kaiser of Munich, Prof. Dr. Fries of Upsala, Forstrath von Plonnies and Oberforster Losch at Amorbach. Preserved material came from Herr Hauptlehrer Allescher of Munich, Director Dr. Goethe and Dr. Wortmann in Geisenheim, Prof. Dr. Stahl of Jena, Prof. Dr. Magnus of Berlin, Prof. Dr. Grasmann and Prof. Dr. Loew of Tokio, Dr. Bruns of Erlangen kindly photographed some specimens in the botanical museuin there. Numerous botanists have greatly assisted me by sending papers, especially Dr. Dietel of Leipzig; I have also to Vili AUTHOR’S PREFACE. thank him for valuable aid with the Uredineae. To Prof. Dr, Soxhlet I am indebted for literature and the opportunity given to establish a museum of pathological material in connection with the agricultural division of the Munich Technical School. Dr. Solla of Trieste, while working in our laboratory here, very kindly translated the earlier fascicles of the ‘Funghi parasitici’ of Briosi and Cavara as far as they were then published. Prof. Dr. Wollny allowed me to carry out some researches on his experimental fields. Very opportune were the investigations of my pupils, Dr. Woernle and Dr. W. G. Smith, on the anatomical changes in plants attacked by Gymnosporangeae and Exoasceae respectively. To all these gentlemen, and to many more who sent me material, but whom it is impossible to name individually in this place, I here express my warmest thanks. The reproduction of my drawings and photographs has been most carefully carried out by Herr O. Consée of Munich. I am also deeply indebted to the publisher, Herr Springer, for the excellent manner in which he has done his work; this will no doubt be also appreciated by the reader. v. TUBEUF. Monicu, December, 1894. NOTE TO THE ENGLISH EDITION. SINCE the publication of this work, I have received a large contribution of original papers. Though there was no time to embody all these in the English edition, yet many of them have been used for its correction and amplification. Some were of such a kind as to necessitate the re-writing of whole sections, notably those on the genera EHxoaseus and Gymnosporangium. The remainder will be thoroughly revised if a second German edition be called for. I again take the opportunity of thanking all those who have sent me literature, and I shall be grateful if they will continue to do so in the future. v. TUBEUF. Monicu, December, 1895. PREFACE TO THE ENGLISH EDITION. My justification for placing another translation in our libraries is that no such book as this exists in the English language, and that I could not, for some considerable time, see my way to collect so many observations on the cryptogamic parasites of higher plants, or to find so many suitable subjects for the pictorial illustration of their habits and structure, as Dr. von Tubeuf has given us. The work was undertaken all the more willingly, because, while working under the guidance of the author, I had seen the book take shape in his hands, and even added some items to its pages. The aims of the book are sufficiently set forth in the author’s preface, and in the preparation of an English edition these have been kept in view. The first or general part and the more important descriptions in the second part are practically translations, but a certain amount of modification was found necessary in adapting the work to the requirements of English readers. With this object many additions were made both by the author and myself. Those which I have inserted are in most cases indicated by the use of (Edit.); this has, how- ever, been entirely omitted in the group ‘Fungi imperfecti, and nearly so in the Uredineae, on account of the number of changes found necessary. I also thought it advisable to indicate whether the different species of fungi had been recorded for Britain and North America; this has been done generally by the use of brackets,—(Britain and U.S. America.) The records for Britain are taken from the works of Plowright, Massee, and others; those of three groups,—the Uredineae, Basidiomycetes, and ‘Fungi imperfecti’ were, however, revised by Professor J. W. H. Trail of Aberdeen, a well-known authority. For America x PREFACE TO THE ENGLISH EDITION, the records of economic interest are selected chiefly from Farlow and Seymour’s Host-Jndex, which contains the complete list. I here take the opportunity of expressing my thanks to Professor I. Bayley Balfour for valuable aid and advice; to Professor J. W. H. Trail for kindly revising important parts of the proofs; to my brother, Robert Smith, for assistance in proof- reading, and to other friends who have aided me. The difficulties of translation are well known; in the present case they have been increased by the technical nature of the subject, and by the modification which the original has under- gone. Faults there must be; for those I ask the indulgence of the reader. W. G. SMITH. Royat Boranic GARDEN, Epriypureu, October, 1896. BIBLIOGRAPHY. Tue following are some of the more important general works and text books. Books and papers on special subjects are given throughout the text as foot-notes :-— GENERAL WorkKS ON FUNGI. TuLasneE. Selecta fungorum carpologia. 1861-1865. De Bary. Comparative Morphology and Biology of the Fungi, Mycetozoa, and Bacteria. English edition. 1887. Zorr. ‘Die Pilze.” Schenk’s Handbuch der Botanik. 1890. Lupwie. Lehrbuch der niederer Kryptogamen. 1892. BreFELD. Untersuchungen aus dem Gesammtgebiete der Mycologie. 1872-1891. Tavet. Vergleichende Morphologie der Pilze. 1892. Saccarpo. Sylloge fungorum. 1882-1893. Winter, Fiscuer, and Ream. “ Die Pilze.” Rabenhorst’s Kryptogamen- flora. ScuroeTer. “ Die Pilze.” Cohn’s Kryptogamenflora von Schlesien. 1885- 1894. (Incomplete.) Scuroeter. “ Die Schleimpilze und die Pilze.” Engler-Prantl natiirlichen Pflanzenfamilien. 1892-1894. (Not yet complete.) Cooke. Handbook of British Fungi. 1871. Piowricut. British Ustilagineae and Uredineae. 1889. Fartow and Seymour. Host-Index of the Fungi of the United States. 1888-1891. Massez. British Fungus-Flora. 1892-1895. Works ON DISEASES OF PLANTS. Uncer. Die Exantheme der Pflanzen und einige mit diesen verwandte Krankheiten der Gewiichse. 1833. Wiremann. Die Krankheiten und krankhaften Misbildungen der Gewiichse. 1839. Meyen. Pflanzenpathologie. 1841. xil BIBLIOGRAPHY. De Bary. Untersuchungen iiber die Brandpilze und die durch sie verur- sachten Krankheiten der Pflanzen. 1853. Ktuy. Die Krankheiten der Kulturgewichse und ihre Verhiitung. 1858. Hauer. Phytopathologie ; die Krankheiten der Kulturgewichse. 1868. Hartia, R. Wichtige Krankheiten der Waldbiume. 1874. Harrie, R. Die Zersetzungserscheinungen des Holzes. 1878. Frank. Die Krankheiten der Pflanzen. I. Aufl. 1880, II. Aufi. 1894-1896. Harrie, R. Lehrbuch der Baumkrankheiten. I. Aufl. 1882, II. Aufl. 1889. (Editions in English, French, and Russian.) Smirn, Worrn. G. Diseases of Field and Garden Crops. 1884. Soraver. Handbuch der Pflanzenkrankheiten. II. Aufl. 1886. Wotr and Zorr. Krankheiten der landwirthschaftlichen Nutzpflanzen durch Schmarotzerpflanzen. 1887. Soraver. Die Schiiden der einheimischen Kulturpflanzen durch thierische und pflanzliche Schmarotzer. 1888. MarsHatt Warp. ‘Timber and some of its Diseases. 1889. Kircuner. Die Krankheiten und Beschidigungen unserer landwirthschaft- lichen Kulturpflanzen. 1890. Frank and Soraver. Pflanzenschutz. 1892. Prituieux. Maladies des plantes agricoles et des arbres fruitiers et forestiers causées par des parasites vegetaux. 1895. Zeitschrift fiir Pflanzenkrankheiten (since 1891). The Publications of the Division of Vegetable Pathology of the Department of Agriculture, U.S. America, issued from Washington. The Bulletins of the Agricultural Experimental Stations, issued by many of the States and Universities of the United States. Exsiccata of Parasitic Fungi, by (2) Briosi and Cavara, (b) Eriksson. “Economic Fungi” of U.S. America, by Seymour and Earle; Exsiccata begun 1890, (still being issued). Etc., ete. wp OP a ws CT GL GL Ca DH oO > Do =e CONTENTS. PART FIRST. CHAPTER I. THE PARASITIC FUNGI. PAGE DEFINITION OF THE PaRASITISM OF FUNGI, - - - CLASSIFICATION OF PARASITES AND SAPROPHYTES, - . Mopkr or Lire or THE Parasitic Funal, - - - CHAPTER ILI. REACTION OF HOST TO PARASITIC ATTACK. Errect oF Parasitic FunGI on THEIR Host, - - - Errect oF Parasitic FunGI on THE Form oF THE Host-PLANt, Errect oF Parasitic Fune1 oN CELL-ConTEN Ts, - . Errect oF Parasitic Funel on THE CEeLL-WALL, - Errecr or Parasitic FuNGI ON THE TISSUES OF THE’ CHAPTER III. Errecrt OF THE SUBSTRATUM ON THE PARASITE, - - - XIV CONTENTS. CHAPTER IV. $10. NATURAL AND ARTIFICIAL INFECTION, - - CHAPTER V. $11. DISPOSITION OF PLANTS TO DISEASE, - - CHAPTER VI. PREVENTIVE AND COMBATIVE MEASURES, - - wee _ bo I. EXTERMINATION AND REMOVAL OF THE PARASITIC FUNGI ALONE, - - - = 2 II. RemovaL AND DeEstRUCTION OF DISEASED PLANTS, - III. AvorpANCE OR REMOVAL OF CONDITIONS WHICH FAVOUR INFECTION, - - - . - . IV. Sevection or Harpy VARIETIES, - - - CHAPTER. VIL § 13. ECONOMIC IMPORTANCE OF DISEASES OF PLANTS, CHAPTER VIII. SYMBIOSIS: MUTUALISM, . - - : : ‘ CHAPTER IX. NUTRICISM, - - - : 4 ‘ Hic Mycoruiza, - 2 : : ~ R 1c Mycoruiza, - - - - - 40DOM \ OF THE ALDER, ETC., - - - - ¥CODOMA, OF THE LEGUMINOSAE, - - - : PAGE 58 83 86 92 93 97 99 101 CONTENTS. XV PART SECOND. SYSTEMATIC ARRANGEMENT OF THE CRYPTOGAMIC PARASITES. PAGE I. THE PATHOGENIC FUNGI OF PLANTS, - - - 104 A. Lower Funei (PHycomycereEs), - - - - 106 (1) CuyTrIDIACEAE, - - - - - 106 (2) ZyGoMyceTEs, - - - - - - 114 (3) OomyceTEs : PERONOSPOREAE, - - - - 115 B. HicHer Funai (Mycomyceress), - - - - 135 ASCOMYCETES, - - - - - - 136 A, GYMNOASCI, - - - - - 137 ‘THe Parasitic EXoAscEak, - - - 144 B. CARPoasct, - - - - - - 168 PERISPORIACEAE, - - - - 170: PYRENOMYCETES, : : 2 - 183 HYSTERIACEAE, - - - - - 232 DiIscoMYCETES, - - - - - 240; UsTILAGINEAE, - - : - - - 275 UREDINEAE, - - - - - - - 328 BASIDIOMYCETES, - - - - - - 421 Funer Iwrerrecti—I. SPHAEROPSIDEAE, - - - 463 II. MELANCONIEAE, - : - 482 III. Hypuomyceres,-— - - - 496 Il. THE PATHOGENIC SLIME-FUNGI, - - - 522 Ill. THE PATHOGENIC BACTERIA, - < ; 3 530 IV. THE PATHOGENIC ALGAE, - - - - 539 INDEX OF PARASITES, - - - - = . 556 GENERAL INDEX, - . - - - - - 580: ERRATA. 9, Fig. 1, for “‘ Erysipheae ” vead ‘‘ Erysiphe.” line 11 from foot, for ‘‘ tyrosin” read ‘‘ trypsin.” 35, 181, 185, 195, 256, 305, 312, 337, 403, 404, 420, 29 9) 24, for *‘ quercinium ” read ‘‘ quercinum.”’ 6, for ‘* Nectrina”’ read ‘‘ Nectria.” 3, for ‘‘setuloso”’ read ‘‘ setulosa.” 6, for ‘‘ Belionella”’ vead ‘* Beloniella.”’ 11 from foot, for Tolysporium ” read ‘‘ Tolyposporium.” 16, for ‘heloscladii” and ‘‘* Heloscladium” respectively, read *‘ helosciadii ” and ‘* Helosciadium.” 10, for ‘‘ Onybrychis ” read ‘* Onobrychis.” 25, for “* Cichoria” read ‘‘ Cichorium.” 9, for ‘‘ Cypressus”’ read ‘* Cupressus.” 5, for *‘ Escheveria” read ‘‘ Echeveria.”’ 3, for *‘ Thecospora ” read ‘‘ Thecopsora.” PART FIRST. CHAPTER I. THE PARASITIC FUNGI. THE true Fungi, together with the Myxomycetes or Slime-fungi, and the Schizomycetes or Bacteria, constitute a group of the Cryptogams characterized by lack of chlorophyll. In consequence, the members of the group are unable to utilize light as a source of energy, and must obtain their food as organized material, complex in comparison with the simple substances required by green plants. These fungi, in short, are, in common with animals, ultimately dependent for the greater portion of their support on living or dead chlorophyllous plants. According as they obtain nutriment from dead organic remains or from living plants or animals, we distinguish them as Saprophytes and Parasites respectively. The same mode of nutrition is found in the case of most non-chlorophyllous Phanerogams, and also in a few chlorophyllous plants, both Cryptogams and Phanero- gams. When parasitic Fungi, Bacteria, and other lower organisms attack higher plants, they, as a rule, endeavour to penetrate the living organs of their host. It is only when this penetration has taken place to some extent, and the parasite has thereby come into more or less close contact with the tissues of its host, that conditions suitable to a parasitic mode of nutrition are established. To deal with the lower forms of vegetable parasites, with their relations to their respective hosts, and with the structural altera- tions which they bring into existence in the latter, is our object in the present book. 2 THE PARASITIC FUNGI. § 1. DEFINITION OF THE PARASITISM OF FUNGI. Parasitic Fungi are those which, stimulated by the cell- contents of another living plant, penetrate wholly or partially into its tissues, and draw their nutriment from that source. Saprophytic Fungi are those which make no attempt to penetrate the tissues of living plants, but derive their nutriment from a dead substratum. Intermediate between these two extremes come those fungi which, in consequence of some stimulus, attempt to effect an entrance into the tissues of living plants by the secretion of some fluid or ferment, but only attain their object after first killing the part they attack (e.g. Sclerotinia sclerotiorum). A special position’ must also be ascribed to certain forms which inhabit the wood of trees, but have not the power to penetrate through the outer tissues; they depend on first gaining entrance through wounds into dead parts of the bark or wood, and, after living there for a time as saprophytes, extend into the living elements and cause their death. Many parasites may be artificially cultivated so as to pass some part of their life-history on dead pabulum, and even in natural conditions many of them regularly live for a season in a saprophytic manner. On this account it appears to me more correct, in distinguishing between parasites and saprophytes, to lay less weight on the adaptation to nutrition and more on their response to the stimuli exerted by living plant-cells. The nature of this stimulus which affects parasitic hyphae has not as yet been investigated. It appears probable, however, especially from the investigations of Pfeffer and Miyoshi, that the influence is primarily a chemical one, and that the nutritive value of the stimulating substance is not a measure of the ensuing effect. Biisgen states that the formation of adhesive-dises by germinating spores is induced by a stimulus due to contact, whereas the production and penetration of the first haustorium is independent of contact, and is probably due to some chemical stimulus (see: p. 9). Miyoshi’s investigations have also proved that saprophytic fungi are capable of penetrating into living plant-organs, even 1 Miyoshi. ‘‘ Ueber Chemotropismus d. Pilze.” Botan. Zeitung, 1894; also ‘‘ Die Durchbohrung von Membranen durch Pilzfaden.” Pringsheim’s Jahrbuch, 1895 Senta 166.1} ) . Z = fd : J se Pfeffer. Ueber Election organischer Nahrstoffe.” Pringsheim’s Jahrbuch, 895. ; DEFINITION OF THE PARASITISM OF FUNGI. 3 of boring through cell-walls, if the part be impregnated with a stimulating solution. They behave here completely as parasites. For example, hyphae of Penicillium glaucum penetrate into living cells of a leaf injected with a two per cent. solution of cane sugar, while without previous injection of the leaf they have never been observed to do so. Penicillium is also known, in certain circumstances, to become parasitic. Many species of fungi are capable of passing the whole or a part of their life as parasites on living plants. Conspicuous in this respect are the Uredineae and Ustilagineae, many Ascomy- cetes, including all Exoasceae and Erysipheae; and amongst the lower fungi, most of the Chytridiaceae and all the Peronosporeae. Nor does this exhaust the list, for amongst the remaining fungi we may find isolated families, genera, and even species occurring as parasites, while forms closely related to them are saprophytic. To classify the parasites, saprophytes, and intermediate forms, we shall adopt that arrangement proposed by Van Tieghenr and De Bary. § 2. CLASSIFICATION OF PARASITES AND SAPROPHYTES. 1. True saprophytes are such as regularly pass through their whole life-history in a saprophytic manner. They may derive their nourishment from different kinds of pabulum, or be limited to some definite substratum. The true saprophytes do not come within the scope of this book.! 2. Hemi-saprophytes (the ‘facultative parasites’ of De Bary) are wont to pass through their whole development as saprophytes, but on occasion are capable of existing wholly or partially as parasites. Amongst them are included particularly such species as may be designated “occasional parasites,’ which commonly occur as saprophytes, and only under certain conditions become parasitic. 3. True parasites (the ‘obligate parasites’ of De Bary). These undergo no part of their development as saprophytes, but live in every stage of existence as parasites. 4. Hemi-parasites (the ‘facultative saprophytes’ of De Bary) are capable, if need be, of becoming saprophytes for a season 'Johow proposes the term Holo-saprophytes for those non-chlorophyllous Phanerogams which live exclusively saprophytic on organic debris, in contrast to those possessing chlorophyll, which he names Hemi-saprophytes. 4 THE PARASITIC FUNGI. but as a rule they live throughout their whole development as parasites. Within each of these four divisions one may introduce a number of subdivisions. Hemi-saprophytes. The majority of saprophytes are never parasitic, yet there are a number which become so occasionally. Thus some species of Mucor and Penicillium can penetrate into thin-skinned fruits, and this they do the more easily, the further the fruits are from the condition of full vital energy, to use De Bary’s expression. Related to these are other fungi which, although incapable of effecting entrance into plants in active life, may yet do so as the plant, though still living, begins to wither. In such cases the parasitism is somewhat difficult to prove. In particular, the so-called ‘Fungi imperfecti’ contain forms of this kind. Amongst the hemi-saprophytes we may include the species of Botrytis, which are able to penetrate into unfolding parts of plants, but not into the older parts. We may specially mention Botrytis Douglasii as a form more generally known as a sapro- phyte, but which becomes parasitic on immature organs, and which penetrates young needles of various conifers to kill them, whereas it 1s unable to attack older needles. In this case the thickness of the membranes would seem to act as a protection, | just as the vital energy of the plant does in the preceding cases. In Sclerotinia sclerotiorum, Sel. ciborioides, and Sel. Fuckeliana, a saprophytic existence must, as in the example just mentioned, precede the parasitic condition; in fact De Bary holds that these forms can only become parasites after their mycelium has been saprophytically strengthened; the parasitic condition is not necessary to them, for they can go through their whole develop- ment on a dead substratum. Pythium De Baryanum is also to be regarded as a hemi-saprophyte which attacks and _ kills seedlings of many plants as a parasite, but otherwise vegetates on dead plant remains. Cladosporium herbarum, one of the commonest of saprophytes, behaves similarly, but it is of less frequent occurrence than Pythiwm, and in fact its parasitism has only been suspected quite recently. ‘This has been confirmed by Davaine (Compt. rend. Lx111., 1866, pp. 277 and 344) and Brefeld (Sitzungsber. d. naturforsch. Fr. zu Berlin, 1875). CLASSIFICATION OF PARASITES AND SAPROPHYTES. 5) As further examples of fungi, capable, as parasites, of killing living cells, but which pass through more or less of their life as saprophytes, may be taken species whose mycelium inhabits the wood of trees and shrubs. Amongst these are numerous’ Polyporeae, which find admission only by wounds in the wood. At first these destroy and derive nourishment from the substance of dead parts of the wood, but later they begin to attack the parenchyma of the living wood, and extending outwards kill, as they go, cambium, bast, and rind, till they reach the exterior, and there develop sporophores. As examples we may take those species investigated by R. Hartig of Munich, eg. Poly- porus fomentarius, P. igniarius, P. Hartigii, P. sulphureus, Stereum hirsutum, Trametes pini.’ The heart-wood is a part of the tree generally avoided by insects, which would in very short time destroy the sap-wood with its rich starch-content, eg. Annobiae in oak. Again, the heart-wood resists the influence of certain saprophytic fungi much longer than the sap-wood, hence it is preferred as the timber used for railway sleepers. Although in these cases we might describe the heart-wood as possessing antiseptic properties, yet this would scarcely be accurate, since it is just this very heart-wood which is always first attacked by the wound-parasites of trees, and gives them a hold on the tree as parasites. See also Chap. V. Since these dangerous tree-fungi can live wholly as sapro- phytes in the heart-wood, and in the sap-wood partly as such, partly as parasites, they are also able to vegetate further, and to reproduce themselves on felled stems, especially when the ‘necessary moisture is provided. Thus, for example, Agaricus adiposus, 2 wound-parasite of the silver fir, produces its yellow sporophores on felled stems and split wood during the whole summer in moist parts of the forest, while in a cellar or other moist chamber the development of sporophores may continue over a year. In fact, I have found that a billet of beech-wood, after being placed under a glass and allowed to lie completely dry, on again being soaked from time to time, continued to produce a crop of toadstools annually for five years. Some wound-parasites occur occasionally as typical sapro- phytes on dead wood. Thus Polyporus annosus, perhaps better 1R. Hartig, Zersetzunyserscheinungen des Holzes, 1878, and other works. 6 THE PARASITIC FUNGI. known as Z'rametes radiciperda, is an undoubted parasite of pines, spruces, and other trees, yet on timber in mines! it grows luxuriantly, and reproduces abundantly from sporophores, which, however, differ somewhat from the typical form. Again, the rhizomorph-strands of Agaricus melleus grow under dead bark, in the earth, in mines, and in wooden water-pipes, while other forms of its mycelium are completely parasitic; thus the apices of the rhizomorphs penetrate the bark of young conifers, and, in the form of a mycelium, live parasitic on rind, bast, and cambium. Polyporus vaporarius, a true parasite on living Scots pine, is also an enemy of timber in newly-built structures, or in subterranean spaces and cellars, so long as it can obtain the necessary moisture. Polyporus sulphureus produces sporophores on the bark of living trees, as well as on the dead stools of felled trees. Many other related forms would probably be able to live on dead timber if they were not dependent on a certain degree of moisture, and could submit to drying-up as easily as, for example, Polyporus abietinus, a true saprophyte, and one of the most common enemies of old wooden bridges. Fungi from other groups are also known to effect an entrance into the wood of trees through wounds only, yet when once in, they spread rapidly, and at leneth bring about the death of their host. The spores of Cucurbitaria laburni were demonstrated by me to germinate on the laburnum, on wounds produced by hail and otherwise, and to send into the wood so exposed a mycelium, which spread through the vessels and into the rind, killing all the tissues on its way. Similarly Weetria cinnabarina, after it has killed its host, lives thereon as a saprophyte, and develops patches of conidia and perithecia on the dead _ bark. Pexiza Willkommii, although really a strict parasite on the living rind, yet continues to grow and to reproduce itself on the dead branches. Hemi-parasites. If the examples already given, 7c. Mucor, Penicillium, Lotrytis, Pythium, are typical of hemi-saprophytes, then there may arise a doubt whether the remainder, the wood-destroying Polyporeae, Nectria, Cucurbitaria, and Agaricus melleus, should not be regarded ‘Harz, Botan. Centralblatt, 1888, Vol. xxxv1.; Magnus, Botan. Verein d. Prov. Brandenburg, 1SS8. CLASSIFICATION OF PARASITES AND SAPROPHYTES. 7 as hemi-parasites. They must, however, be included amongst the hemi-saprophytes, because doubtless they are capable of going through their whole development as saprophytes. The hemi-parasites include, amongst others, the Ustilagineae, all of which live for a time as parasites, and cannot, even by artificial cultivation, be made to complete their life-history as saprophytes. While, however, many of the Ustilagineae are adapted to a com- pletely parasitic life, others can, in the form of sprouting conidia, live and multiply saprophytically. The conidia of Hvobasidiwm and Hzvoascus continue to bud off conidia for a considerable time in nutritive solutions, yet in nature, the spores probably produce infecting hyphae at once, and the fungus is but little suited to sustain a saprophytic mode of life. Phytophthora infestans is more easily reared as a saprophyte, and occurs in nature as such, hence it approaches somewhat towards the hemi-saprophytes. True Parasites. The Uredineae may be taken as the most typical of the true parasites; they constantly pass through their whole life-history on living plants, and cannot be cultivated on a dead substratum. So also the Erysipheae, although frequently their spores only reach maturity on a dead substratum, as do also those of Rhytisma and Polystigma. Ergot of grain and the Sclerotinia inhabiting berries, are also truly parasitic, even though their apothecia or perithecia are produced from hibernating sclerotia, and though their conidia can be saprophytically cultivated on dead pabulum. The Peronosporeae and Protomyces are also true parasites. In many other forms the development of germ-tubes, or the sprouting of conidia, may be obtained in artificial nutritive solutions by exclusion of rival fungi and bacteria, yet it is doubtful whether this takes place in nature. § 3. MODE OF LIFE OF THE PARASITIC FUNGI. The parasitic fungi may be divided according to the place of their occurrence and their mode of attack on the host, into two categories, which may be designated epiphytic and endophytic 8 THE PARASITIC FUNGI. parasites. The former have their vegetative mycelium spread over the surface of the host-plant, the latter penetrate into the plant and there develop their mycelium. Both receive nourish- ment from the cells of the host-plants, generally by means of special absorptive organs inserted into the cells of the host, the so-called haustoria. We may distinguish the following groups of parasites accord- ing to the degree of their penetration into the organs of the host-plant they attack: 1. Epiphytes: (~) with haustoria which only sink into the outer membranes of the host; (b) with haustoria penetrating into the cavity of the host- cells. 2. Endophytes : (~) with a mycelium which grows in the walls of the host-cell, and is generally nourished without the aid of haustoria ; (b) with a mycelium which grows in the intercellular spaces only, and is nourished with or without haustoria ; (c) with a mycelium which penetrates into the host-cells and becomes an intracellular mycelium ; (7) lower fungi which live completely in a host-cell. 1. Acquisition of nutriment by the epiphytic parasitic fungi. The simplest mode of acquiring nutriment is found in yeasts (Saccharomyces apiculatus, etc.) which frequent the outside of living fruits, and live on the drops of sugary solution which diffuse therefrom.’ 1 Epiphytic parasites always produce their reproductive organs outside their host-plant. In the case of endophytic parasites, the reproductive organs of some are produced inside the host-tissue, e.g. the zygospores and oospores of Chytridiaceae and Peronosporeae, the chlamydospores of the Usti/agineae; others form their sporocarps wholly or partially embedded, the spores and conidia only being discharged externally ; while a large number form sporocarps on the surface after the epidermis has been torn. Conidia are generally abjointed from the free surface of the host-plant. The terms epiphytic and endophytic parasites have been chosen with regard to the development of the parasitic food-absorbing mycelium. Some authors regard epiphytism somewhat differently, and include amongst endophytes those forms which live on the surface of the host and penetrate only by haustoria. If this be accepted, epiphytism is very exceptional amongst parasites of the higher plants. Zopt (‘* Die Pilze”) gives as examples of this condition only the following: the Laboulbeniaceae inhabiting the chitinous skeleton of certain insects, and J/elano- spora parasitica on filaments of species of Zsaria ; these have no communication between the mycelium and their host. Species of Chaetocladium parasitic on fungi and absorbing the cell-wall of the host at the point of contact, could. strictly speaking, no longer be classed as epiphytes. *Biisgen. ‘‘ Ueber einige Eigenschaften d. Keimlinge parasitischer Pilze.” Botan. Zeitung, 1893. MODE OF LIFE OF THE PARASITIC FUNGI. 9 I can however hardly regard as parasites, fungi like these which live on an accidental outflow from plants or plant-cells, even though they regularly frequent places where an outflow is to be expected. They exert no influence on the host-plant, and they are nourished by substances which can no longer be regarded as belonging to the host. I would rather include them amongst non-parasitic epiphytes which, without specially adapting themselves, settle on any part of a living plant where sugary solutions suitable for their nutriment may occur. » One might imagine however such epiphytes inducing a diffusion of nutritive substance from the cells of the host-epidermis to the closely adherent fungal hyphae; then we should have the simplest mode of parasitic acquisition of nutriment on the part of epiphytes. They would take up food-material from the epi- dermal cells in much the same manner as many intercellular hyphae do from the adjoining walls of the host-cell.+ Epiphytic parasites frequenting the surface of plant-organs generally endeavour to increase their supply of nutriment from the host-cells by formation of haustoria, which pierce the cuticle or the whole cell-wall. Biisgen has shown experimentally that the adhesive discs, often formed on the germination of a spore, owe their origin to a contact-stimulus; the formation and direction of the infecting hyphae, on the other hand, though depending on this, are much more determined by a stimulus originating from the host-cell itself. In this we have a confirma- tion of the accuracy of our definition of parasite and saprophyte. The appressoria, adhesion-organs or adhesive discs just mentioned, are char- acteristic of many parasites. They are formed chiefly on epiphytic mycelia, but also accompany the earlier life of other fungi. In the case of epiphytes, pores are formed on definite places of a ere at such an adhesive-disc, and from these wntetliverarwm germinating on the i" 3 epidermis of a host-plant; an ad- haustoria are developed, or a hypha is _ hesion-discandhaustorium have been : formed. (After De Bary.) given off and enters the host-plant to : form a mycelium. The appressoria of the Erysipheae are very characteristic; in many they are broad lobed dises (Fig. 1); in t— ? Compare those cases of parasites on insects and fungi already given, p. 8 (note). 10 THE PARASITIC FUNGI. others, like Podosphaera castagnei, they take the form of broadened closely-clinging hyphae with haustoria. Frank describes a swell- ing of the germ-tube of Fusicladium tremulae just before the infecting hypha pierces the cell-walls of its host. A similar phenomenon can be observed in Polystigma rubrum, in Gnomonia erythrostoma, and in the germinating aecidiospores of MJelampsora Goeppertiana. Some other examples will be mentioned in our next section. Haustoria of the epiphytic Parasites. The most inconspicuous haustoria are those of Herpotrichia nigra and Trichosphaeria parasitica, described by Rk. Hartig. lA i} LL Vy en ee 8 ee Pe Fic. 2.—Haustoria of Trichosphaeria parasitica. (Details on Fig. 88.) (After R. Hartig.) They are tiny hyphal processes resting on the host-epidermis, and sunk into the outer walls of the epidermal cells, so as to pierce the cuticle but not the whole wall (Fig. 2, d, e; also Fig. 90). The Erysipheae are typical epiphytes, which weave a mycelium over the surface of plants they attack; the mycelium retains its hold by adhesion-discs or appressoria, and from certain parts of these a fine thread-like process is given off, which, after piercing the epidermal wall of the host, swells inside to a simple or branched sac, the haustorium. The ‘Lehrbuch d. Baumkrankheiten, II, Aufl. English translation by Professor Somerville. Macmillan & Co., 1894. MODE OF LIFE OF THE PARASITIC FUNGI. ll haustoria of Podosphavra castagnei (Fig. 71) are bladder-like, those of Oidium Tuckeri are lobed. The simpleste formation of haustoria consists in an outgrowth of the mycelium which depresses the cell-wall of the host without piercing it (e.g. Peronospora densa). In other cases the cell-wall, at first only depressed, becomes ultimately broken through. Certain lower fungi live parasitic on other fungi and adhere to their hyphae by means of well-developed adhesion-dises from which haustorial structures are formed inside the hyphae of the host. Thus Piptocephalis fresenia is parasitic on hyphae of some species of Mucor, and produces from a swollen bulb-like appres- sorium a tuft of very fine haustoria inside the Mucor-hypha. Syncephalis proceeds even further, for the haustorial process grows and branches inside the host, becoming, in fact, an endophytic mycelium. A further advance towards endophytic parasitism is presented by the Chytridiaceae, low forms of fungi living on algae or fungi; some send haustorial structures into their host, others develop a mycelium whose attack however is directed against only one host-cell. Fischer, in his “ Phycomy- cetes,” thus describes the latter forms: “The vegetative body, a resting swarmspore, consists of a spherical or ellipsoidal part which becomes a sporangium, and of a filamentous vegetative portion which spreads through the host-cell as a haustorium or mycelium and dies away after the formation of the sporangium. This primitive mycelium is uniceliniar, and may be unbranched or very finely branched.” 2. Acquisition of nutriment by the endophytic parasitic fungi. The simplest case of the endophytic mode of life is presented by those fungi which vegetate in the epidermal membranes of their hosts, aud derive their nutriment osmotically through the inner cell-walls. They live covered by the cuticle, which must have been penetrated by an infecting hypha at the time of first attack. This mode of life is exhibited by many fungi, particularly by the Zroasceae; the mycelium of these vegetates under the cuticle of the host plant, and ruptures it at the time of ascus-formation. In spite of their limited distri- bution the species of this group so influence the development of their hosts as to induce pustule-like outgrowths, crumpling and distortion of leaves, and even “witches’ brooms.” In some 12 THE PARASITIC FUNGI. of the Exoasceae the bases of the asci penetrate deeply between the walls of the epidermal cells, so forming an intermediate stage leading to other Hvoasceae and endophytic fungi, with a mycelium growing between, or in the cells of tissues which le deeper than the epidermis. The mycelium of Cycloconium oleaginum grows in the epi- dermal cell-membranes, branching dichotomously under the cuticle and sending through it erect hyphal branches for pro- duction of conidia The germinating conidia of Sphaceloma ampelinum are said by De Bary to penetrate the cuticle, and to produce a mycelium which spreads thereunder and breaks out just before formation of conidia. Jycoidea parasitica, an alga, lives under the cuticle of leaves of Thea and Camellia. We have next to consider fungi with a mycelium which lives and multiples in the intercellular spaces of living plants. Like the Hvoasceae just mentioned, they push their way between neighbouring cells and spread through the already existing intercellular spaces. Numerous Uredineae behave in this way, and towards the period of reproduction the mycelium is capable of increasing so much that the cells of the host-tissues become isolated and even displaced. The various species of Hysteriwm have an intercellular mycelium, which kills those cells with which it comes in contact. Certain forms, eg. Caeoma pinitor- quum and Peridermium pint (Fig. 247) possess a mycelium which, while still intercellular, sends off here and there little lateral branches into the host-cells. It is an easy step from forms like these to forms whose mycelium is no longer strictly intercellular, but derives nutriment by means of specialised haustoria. Haustoria of the endophytic Parasites. A large number of endophytic parasites frequenting hosts which do not immediately succumb to their attack, possess “haustoria” or special organs for the acquisition of nutriment from the cells of the host. The haustoria are lateral outgrowths of the mycelium with a limited period of growth and a more or less constant form. They are more varied in form, but otherwise quite comparable with haustoria of the epiphytes, especially with those of the Erysipheae. One of the simplest forms of ‘Figures in Funghi Parasitti, Cavara and Briosi. MODE OF LIFE OF THE PARASITIC FUNGI. 13 haustorium on an endophytic mycelium is that exhibited by the parasite Cystopus; the hyphae send off very fine filaments which penetrate the walls of a host-cell and swell up to little button-like sacs. Many Peronosporeae (P. pygmaea, P. nivea, P. viticola and Phytophthora omnivora) have haustoria of the form just described, whereas others have them thread-like and branched (P. calotheca of the woodruff), or crenately lobed (P. parasitica). Amongst the species of Uredineae and Ustilagineae, haustoria are not uncommon and present many varied forms. They are, however, few in number, or confined to certain parts of the mycelium, so that they may be easily overlooked. Haustoria in the form of long sacs of various lengths are produced by Melampsora Goeppertiana in the tissues of both cowberry and fir-needle. Gymnosporangium in juniper has occasionally very delicate button-like haustoria. Lndophyllum sempervivt in the house-leek has haustorial branches which, according to Zopf, are coiled together and anastomose frequently with each other. TZuburcinia amongst the Ustilagineae possesses short branched haustoria resembling one-sided clusters, and Melanotaenium endogenum has similar haustorial-tufts even more branched! Uvrocystis pompholygodes in Hepatica triloba has spirally coiled haustorial hyphae, while TZi/letia endophylla, Sorosporiwm saponariae,? and many species of Ustilago, have haustoria with the form of knotted hyphae. Amongst the Hymenomycetes, Evobasidium vaceinii forms a mycelium which permeates the host-tissues with numerous hyphae, but the only haustoria are hyphae which here and there penetrate into a cell. No haustoria have as _ yet been found amongst the Basidiomycetes,? Pyrenomycetes, or Discomycetes. The two groups last-mentioned have an inter- cellular or intracellular mycelium, which as a rule quickly kills all cells with which it comes in contact. ' Senckenbergische naturforsch. Ges, Abhandl. 1880. Plates I. and IV, 2 Pringsheim's Jahrbuch, 1869. Plates VII., VIII. *Sarauw has figured haustoria in mycorhiza of beech, without however determining exactly whether they belonged to a Hymenomycete. Reess also figures similar organs on mycorhiza produced by one of the Tuberaceae. CHAPLER GIL REACTION OF HOST TO PARASITIC ATTACK. THE reaction of the host to the attacks of parasitic fungi is fairly constant for the same host and fungus. The various fungi, however, exert on the same host-plant each an influence of its own, while different host-plants behave very differently under attacks of the same fungus. § 4, EFFECT OF PARASITIC FUNGI ON THEIR HOST." A. Kuivuine or Host-CE.ts.? 1. Absorption of living cell-content by parasitic fungi. The lower fungi give us examples of the simplest mode in which fungus-parasites draw nutriment from their host-cells ; particularly those forms parasitic on algae or other fungi. The most primitive of all are numerous species which, applying themselves to a host-cell, bore through its walls and enter the cavity. There they derive nutriment at the cost of the living cell-content,—the plasma, cell-sap, chloroplasts, starch grains, etc.—and finally kill the cell. The host-cell does not survive the later development and reproduction of the parasite. The effect of the fungus is however limited to the ' Billroth (‘‘iiber die Einwirkungen lebender Pflanzen und Thierzellen aufeinan- der,” Sammlung Medic. Schriften. Wiener klin. Wochenblatt, 1890), compares in a masterly way the effects of micro-organisms and of injuries on animal and vege- table tissues. He employs Virchow’s terms ‘‘ formative stimulus ” and ‘‘ formative irritability ” ; the former to denote the capacity of micro-organisms in producing outgrowths of definite form or the formation of new tissues; the latter, the capacity of the tissues to react to such stimuli, and to produce outgrowths, etc. A comparison of the external phenomena of fungoid diseases in the case of animals and plants recently formed the subject of a short paper by Lewin. * Perniciasmus. EFFECT OF PARASITIC FUNGI ON THEIR HOST. 15 cell attacked which is at once killed before it can enlarge or, otherwise react to the influence of the intruder.* Good examples of such parasites are presented by some of the Chytridiaceae —the Archimycetes of Fischer—which, as a rule, inhabit only isolated cells of their respective host-plants. This mode of nutrition is equivalent to that of the Myxomycetes and Mycetozoa, which absorb the cell-contents after completely enveloping the living cell, or after slipping inside or sending a haustorial process into it. A second series of parasites consists of those which live on the contents of the host-cell, and give it time to react to the stimulus exerted by the intruder. The reaction generally results in a cell-enlargement or fungus-gall, which in the simpler cases includes one cell only. The gall harbours one or more parasites, which gradually use up the cell-contents. As examples we have Olpidium tumaefaciens and O. uredinis,) Pseudolpidium saprolegniae, Olpidiopsis saprolegniae, Rhizomyxa hypogaea, ete. A specially striking case is that of Pleotrachelus fulgens, which causes the rudiment of the sporangiophore of Pilobolus Kleinii to become hypertrophied and gall-like.* We have as a third series those parasites which penetrate into living cells and absorb their contents, at the same time stimulating the host-cell to abnormal and increased growth, as well as some surrounding cells not directly in contact with the fungus. In this case the parasite exerts a far-reaching effect, and produces a gall composed of more than one cell. Species of Synchytrium are examples. The fungus itself penetrates into one cell only, which enlarges; but simultaneously the surrounding cells grow and multiply to form a wall or rampart enclosing the cell originally attacked. Other parasites do not absorb the’ host-contents as a whole, but only withdraw osmotic substances by means of delicate processes of the fungus-hyphae. These haustoria penetrate the wall of the host-cell, but the fungal protoplasm inside them remains separated from the host-proto- plasm by a delicate membrane. In the case of the vine-mildew and some other Erysipheae, the cells thus preyed on turn brown and die. With other related forms (e.g. Sphaerotheca castagnet), ‘See Fischer's Phycomycetes. * This causes a slight swelling of the root-hairs of various plants and absorbs their content. *Zopf, Beitrdge zur Physiol. u. Morphol. nied. Organismen, 11. 1892, 16 REACTION OF HOST TO PARASITIC ATTACK. absorption by haustoria results in a deformation and distortion of attacked organs, which embraces even cells far distant from the point of attack, yet without death following directly to any cell. 2. Absorption of cells or tissues by parasitic fungi. The total absorption of cells or tissues by parasitic fungi constitutes a special form of cell-destruction. Cases of this kind occur particularly amongst the Ustilagineae. Thus Urocystis violae so stimulates the cells of Viola that they divide and produce a delicate tissue, rich in protoplasm; this nutritive tissue is used up when spores are formed, but without any great detriment to the host-plant. At the time of spore-formation of other Ustila- gineae a great destruction of the host-tissues may, however, take place; this is especially marked in attacks of Ustilago maydis, U. avenae, Tilletia tritici, on the ovaries of their respective hosts, as well as in other cases to be considered later. 3. Killing of host-cells and tissues by fungi which excrete ferments. The simplest case under this heading is presented by species of Sclerotinia studied by De Bary, e.g. Sel. selerotiorwm. The mycelium of these, while still lying on the outer surface of the host-plant, excretes a ferment which sinks through the mem- branes into the cell-cavities, causing death to the protoplasm and even destruction of whole tissues. A similar process may be assumed in the case of numerous fungi with a mycelium which grows only in the intercellular spaces, yet causes immediate death to any cell it may touch. This is the case with many leaf-spot diseases, like those due to Cercospora, Hysterium, etc. So also do the apices of rhizomorph- strands kill portions of the bast of living Conifers with which they may come in contact. The rapid death of tissue following the attack of such deadly fungi as Phytophthora is probably due not altogether to the deprivation of nutriment, but also to the effects of a poisonous excretion. This, however, has not as yet been satisfactorily ascertained. B. KiLuine or Organs oR WHOLE PLANTS. A large number of fungi have a mycelium which never ex- tends beyond a very short distance round the point of first infection, and causes only local disease, frequently with no EFFECT OF PARASITIC FUNGI ON THEIR HOST. 17 perceptible disturbing effect on the host. Such is the case - particularly with leaf-spot diseases ; the tissues of isolated spots are killed and fall out, the leaf appearing as if perforated by shot, but otherwise exhibiting no discoloration or other symptom of disease. In contrast to these there are fungi which, directly or indirectly, bring about death of their host or some part of it. The simplest example of parasitic fungi killing their host directly is presented by one-celled or few-celled plants, which soon succumb to attack even on a single cell. Where, however, the host is a highly organized plant, its organs will resist the attack of the parasite for some time. Thus with Phytophthora fagi, the mycelium spreads rapidly through the tissues of a seedling, so that death ensues in a few days. Similarly species of Peronospora rapidly kill leaves, branches, and fruits; likewise Cladosporium, Septoria parasitica, and others. Somewhat different in their action are those fungi which kill some tender part of a plant directly, and thereby in- directly further the death of other parts dependent thereon., As examples, take Pestalozzia Hartigii (Fig. 301) and Phoma abietina (Fig. 293), which kill only some small portion of a young plant or branch, but thereby cause drying-up of higher or distal parts. Gibbera vaceinii on stems of cowberry (Fig. 95) is another example. Similarly cankers arising from Nectria ditissima (Fig. 80), or Peziza Willkommi. Again, Agaricus melleus and Trametes radiciperda kill roots or lower portions of the stem, and bring about the death of trees of all ages. The case varies somewhat with certain wound-parasites like Nectria cinnabarina and Cucurbitaria laburni. There the my- celium extends so vigorously in the water-conducting organs, as to kill them and fill up the vessels, causing thereby so serious a disturbance in conduction, that branches or whole plants wither away in summer. The wood-destroying Polyporeac and Agaricini act similarly, although more slowly; they attack large branches and stems, destroying all parts of the wood, duramen as well as sap-wood, and finally the bark. There are also cases where organs of the attacked host remain alive, but suffer on account of the hypertrophy of other parts. In this way portions of a plant may be killed although not directly the seat of the parasite. This is particularly the case where hypertrophied organs undergo increased growth and B 18 REACTION OF HOST TO PARASITIC ATTACK. utilize the water which would otherwise have ascended to higher parts of the branch-system (Fig. 3). It must indeed - be assumed that the latter are preyed on by the hypertrophied parts and give up plastic material, which they would otherwise have utilized themselves or stored up as reserve material. On branches attacked by mistletoe and other phanerogamous parasities, it can easily be observed, particularly on broad- leaved trees, that a supporting branch grows vigorously in the parts under the influence of the root-system of the parasite, whereas the distal parts of the same branch-system remain stunted and finally die. The mistletoe-bush thus comes to form the termination of the supporting branch. If, in consequence of this, the branch ceases to produce the leaves necessary in preparing food for it, then like every other leafless branch it dies. Such branches carrying leaves of the mistletoe alone may frequently be found on firs, pines, and _ broad-leaved trees ; even whole tree-summits have been seen on the silver fir with every branch terminated by a mistletoe-tuft, not unlike some huge candelabrum. In a similar manner a witches’ broom, developed from a lateral bud, exhibits throughout an increased growth, while the branch supporting it remains thin and dies from the insertion of the broom outwards. So also in attacks of Gymnosporangium on juniper it may be observed that the parts attacked have their growth much accelerated and many of their dormant buds developed, while the distal parts of the same branch die off. In all such cases it is quite probable that, as the distal parts die back, any food material which they may contain finds its way into the hypertrophied region. C. SHORTENING OF LIFE. Many fungi inhabit a plant without disturbing the develop- ment of any part or causing immediate death, yet with such effect that the vegetative period of the organ in question terminates earlier than normally. A very striking example of this is presented by the needles of silver fir on the witches’ brooms caused by aoe or £8,896,364 was depreciated by rust. Oats reached ' The California Vine-Disease. U.S. Dept. of Agriculture, Bull. 2,, 1892, p. 15. * Doppelcentner = 100 kilogramme. ECONOMIC IMPORTANCE OF DISEASES OF PLANTS. 85 32,165,473 dc. at 16 marks, of which 10,325,124 de. or £8,158,023 falls to be deducted on account of rust. Thus on the crops wheat, rye, and oats, the loss reached the sum of £20,628,147 sterling, or almost a third of the total value of the crops. The year 1891 was a very unfavourable one, but even taking the estimate at the half of the above sum we have a yearly loss by rust amounting to £10,000,000 sterling. In Australia the loss in the wheat harvest of 1890-91, on account of rust, has been estimated at £2,500,000 sterling. Consideration of the loss of sums of money like these, which might be considerably reduced if energetic and universal measures were employed against fungoid plant-diseases, will serve to em- phasize the importance of remedial measures. It must also he borne in mind that the use of diseased fodder, especially hay, grass or grain, infested by rust or smut-fungi, is productive of serious results to the various animals of the farm; while the use of meal or flour contaminated with smuts, stinking-smuts, or ergot is dangerous for mankind. 'Frohner, Lehrbuch d. Toxikologie f. Thiertirzte, 1890. CHAPTER VIIL SYMBIOSIS. § 14. MUTUALISM. Mutualism, or Symbiosis in the stricter sense,' has been distinguished as a special case of parasitism. This condition occurs when a _ parasite and its host mutually work for the benefit of one another, each contributing to the other’s nourishment. The lichens furnish the most conspicuous example. Here funeus-hyphae unite with algal cells, the algae furnishing the fungi with assimilated organic nutriment, the fungi pro- viding water and dissolved salts for the algae. While it is by no means uncommon to find two organisms taking a mutual advantage of each other, yet mutualism in its strictest sense is a rare phenomenon. For it generally happens, and is indeed to be expected, that one or both symbiotic organisms modify in some degree their mode of life to suit the altered conditions necessary for their mutual support. Thus amongst the lichens, as a result of the union of fungus and alga, a living organism originates, which in form, necessities, and mode of life is quite new, and differs completely from either of its components. In the lichen-community, the fungus alone reproduces itself; yet the alga occurs as a free organism in nature, while the fungus can only be reared in artificial culture. This combination might perhaps be compared with that of oxygen and hydrogen to form water, also to a certain extent with the union of the sexual cells to produce a new ‘The term Symbiosis was applied by De Bary, (who introduced it), by Frank and others, to denote those cases where a cohabitation or partnership was: observed to take place between two different organisms. (Frank, Lehrbuch d. Sotanik, 1892). ‘* Mutualism” was first used by Van Beneden. MUTUALISM. 87 individual. These, and other examples, will serve to illustrate how we have in the lichen an organism with peculiarities of structure and of life, widely differing from those of either an alga or a fungus. This unification of two living beings into an individual whole, I have designated “ Zndividuation.” } In the case of the lichen-symbiosis, the chlorophyllous part consists of minute algal cells, completely enclosed in a_ tissue of fungus-hyphae, and the lichen lives as a perfectly isolated and independent plant. The case is, however, different where the fungus enters into parasitic relationship with the green cells of a large plant. Union may then take place, so that the fungus lives on, or inside its host, and removed from contact with any other substratum. The fungus is, however, not in a position to convey any nourishment to its host, and in fact is absolutely dependent on it for the organic substance and water necessary for growth. Where, however, the relationship is such that the parasitic fungus is still in contact with some other substratum, then it may be assumed that, in spite of its parasitism, it takes up nutriment from this source, and shares it with its host. This, as has already been pointed out, is the state of things in the lichens, where the fungus completely envelopes the small isolated algae, and must, as a condition of the growth of the lichen, remain in direct contact with the substratum: the fungus is believed to take from the substratum water and inorganic food-material with which it supplies the algae, while it receives in return plastic organic substance to be used in its own growth. Of course cases do occur amongst the lichens, where, in moist places, the alga is not dependent on the fungus, or, on the other hand, where the fungus can itself take up organic substance from its substratum. Another example of the case is the union of fungi with non-chlorophyllous plants which inhabit humus (e.g. Monotropa). Here the fungus takes up organic nourishment from the substratum and supplies it to the higher plant, which, in consequence of its lack of chlorophyll, is directly dependent on the plastic organized substance from the soil, supplied through the agency of the fungus. The latter, however, receives nothing in return; it requires nothing, since its substratum offers it the most favourable conditions for nutrition. This form of ' Individualismus. re) SYMBIOSIS. io 2) symbiosis, in which the fungus becomes the nurse or feeder, I distinguish as Nutricism. Between the case just cited and that in which the fungus is a pronounced root-parasite on green plants, there exists every possible intermediate stage. Before nutricism is considered in detail it would be well to exemplify brietly from the ranks of plant-parasites, that pheno- menon of individuation so sharply defined in the lichens. A large number of parasitic fungi cause local cell-enlargement and cell-increase, with the frequent result that an attacked plant- organ becomes very much enlarged and its form much changed. Qne speaks in such cases of hypertrophy and _ hypertrophied organs. It is quite evident that in cases of hypertrophy the attacked part must be better nourished, otherwise it could never sustain the great Increase in number and size of its cells. The hypertrophied organ is, in fact, imdebted to the surrounding healthy parts for its additional nourishment; in other words, the place of demand draws to itself the materials it requires. This is all the more necessary when the region of increased erowth is deficient in, or altogether devoid of, chlorophyll, and thus quite dependent on the assimilating green parts. This is frequently the case, as in the scales of alder catkins attacked by Lxoascus alni incanae, in the needles of silver fir deformed by Aecidium elatinwm, or in the yellow needles on spruce resulting from t My Fic. 17.—Spruce seedling in third year, grown in clay-loam. Typical coral-like mycorhiza are absent. The strong root to the right shows, on its newer parts and on all lateral roots, only root-hairs and no fungus. The remaining roots are not modified in any way—some are covered with loose fungal caps, others have both fungal caps and root-hairs, while others are quite free from fungi. (v. Tubeuf phot,) 96 SYMBIOSIS. in Monotropa. The root-system of a tree has not only to secure nourishment, but also the rigidity and stability of the tree. This latter can only be attained by a wide distribution of roots in the firm subsoil free from humus, where normal roots with root-hairs will be formed. The nursing function of the mycorhiza seems thus to be less important than in the case of Monotropa. Fic. 18.—Mycorhiza of Pinus Cembra. A, Typical mycorhiza. B, Root showing clusters of mycorhiza as well as portions clad with fine root-hairs. @, Rootlet exhibiting button-like thickenings externally devoid of a fungoid mantle, but internally completely destroyed by mycelium. D, Section through a thickened branch of a mycorhiza-cluster: a, fungoid mantle; /, fungoid tissue between the cells of the root, rendering them unrecognizable except by their large nuclei ; the inner parts contain no fungi. (v. Tubeuf del.) My newest investigations on this subject? show that, amongst the gymnospermous forest-trees, the Abietineae alone have roots externally clothed with a fungus; the remaining groups have all endophytic mycorhiza. The Abietineae have frequently only a fine mautle of fungus on their rootlets, and do not produce the tufts of short, branched roots so characteristic of mycorhiza in general. Frank does not seem to be altogether correct in his view that the Abietineae are almost or quite incapable of multi- plication by slips, because they would then require to exist for a time without mycorhiza. Probably there is some other reason for this, because the Salicaceae (e.g. Poplars), which have typical coral-branched mycorhiza, are almost exclusively multiplied by slips. 1 Hoveler, (‘‘iib. die Verwerthung d. Humus bei d. Ernahrung d. chlorophyll- fiihrenden Pflanzen.’ Inaug. Diss., Berlin, 1892}, states that roots are able to utilize the soil-constituents without aid of fungi. 2Tubeuf, Forst.-naturwiss. Zeitschrift, 1896. ECTOTROPHIC MYCORHIZA. 97 After the mycorhiza have functioned as such for some time, the fungoid sheath, as well as the hyphae contained in the cortex of the root outside the endodermis, are thrown off by internal cork-formation. This is, however, not always the case, for the fungus may penetrate further and develop injurious para- sitie characteristics; this is so with Polysaccum' and Elaphomyces.? Endotrophic Mycorhiza. (1) On non-chlorophyllous plants living in humus. Certain Orchideae—WNeottia Nidus avis, Epipogon Gmelini, Goodyera repens, ete., as well as some Gentianeae,* possess roots developed as endotrophic mycorhiza. In Coralliorhiza the fungus frequents the short coral-like rhizomes. The fungus in these cases penetrates into the cells of the root-cortex, and there forms a ball or coil of hyphae; it neither covers the roots externally nor inhabits the epidermal cells, so that the production of root-hairs goes on quite normally. From the circumstance that the hyphal coils become emptied and only the remains of walls are left in the still living root-cells, Frank concludes that the fungus after being nourished for a time by the root-cells is ultimately deprived of its contents by them. On this account he calls these roots “ fungus-traps,’ and the plants possessing them “ fungus-digesting plants.” It must be remarked, however, that the fungus grows onwards from older parts of the roots to younger, so that here, as in many other cases, the contents of the hyphae may pass from the older into the younger hyphae. Frank himself suggests* the possibility that the roots take up nutriment without aid from the enclosed fungus, and also that the latter receives its food parasitically from the former. What advantage the roots may receive from reabsorption of food, which they have previously supplied to the fungus, has not been closely investigated, nor has the question whether the roots are in a position to nourish the plants equally well without fungi. The root-fungi of Orchideae have long been known, and Pfetter® ' Bruns, ‘‘ Beitrag z. Kenntniss d. Gattung Polysaccum,’’ Flora, 1894. * Reess, ‘‘ Untersuch. tiber d. Hirschtriiffel,” Bib/iog. Botan. 1887. * Pringsheim’s Jahrbuch, xvi. and Xx. 4 Frank, Lehrbuch d. Botanik p. 267. ® Landwirth. Jahrbuch, 1877. G 98 SYMBIOSIS. suggested that the reduced formation of hairs on their roots was due to the fungus-hyphae behaving physiologically as root-hairs, Johow, in opposition to Frank, states that the non-chloro- phyllous Wullschlaegelia, a relative of Neottia, shows no trace of Fia, 19.—-Coralliorhiza innata Br. (v. Tubeuf phot.) Fic. 20.—Neottia Nidus avis Rich. (v. Tubeuf phot.) fungal hyphae about its roots, and yet derives nourishment direct from humus, The same author found among roots of the Bur- manniaceac, some free from fungi, and some with the rind, and even the epidermis full of mycelium. (2) On chlorophyllous plants living amongst humus. Accordmeg to the investigations of Frank, all our Ericaceae, Epacrideae, and Empetraceae, living in the humus of moor, heath, ENDOTROPHIC MYCORHIZA. 99 or wood, possess endotrophic mycorhiza. These appear as fine, elongated rootlets whose epidermal cells never develop as root- hairs, but become filled with coils of fungoid hyphae. Schlicht mentions a large number of herbaceous plants out of the most widely separated genera of Angiosperms, the finer roots of which he found regularly developed as mycorhiza. These, how- ever, possess in addition normal root-hairs, which without doubt function as such. The endotrophic coils of fungi are situated in the inner cells of the cortex surrounding the conductive tissues, and Schlicht regards them as important in transmitting to the conducting tissues substances taken up by the root-hairs. Since, however, the fungus inhabits living cortical cells, it is quite possible that these transmit the food-materials direct without the aid of the fungus. Schlicht found endotrophic mycorhiza on Leguminosae, while Frank found them on the alder, both being distinct from the well-known tubercles of these plants. Kiihn! and Goebel? found endophytic root-fungi on Marat- tiaceae, Ophioglosseae, and Lycopodium; Kiihn also found spores which resembled those of Schinzia. Endotrophic mycorhiza are also present in saprophytic green orchids, as well as in hemi-saprophytic orchids without green colour; and Meineke* found hyphae passing through the little cells of the endodermis of the aerial roots of orchids into the mucilage-masses of the rind-parenchyma. Schimper found fungi present on the adherent side of the roots of epiphytic orchids. Mycodomatia of Myricaceae, Elaeagnaceae, and the Alder.' The above-mentioned plants possess a well-developed and normal root-system, and also characteristic lateral outgrowths, ' Kiihn, ‘‘ Untersuchungen iib. d. Anat. d. Marattiaceen,” Flora, 1889. * Goebel, Botan. Zeitung, 1887. ® Meineke, ‘‘ Beitr. z. Anat. d. Luftwurzeln d. Orchideen,” Flora, 1894. * Bibliography—Woronin, Mém. de Vacad. des sci. de St. Pétersburg, 1866. Brunchorst, Ber. d. deutsch. botan. Ges., 1885. Brunchorst, Unters. aus d. botan. Inst. Tiihingen, 1886. Moller, Ber. d. deutsch. botan, Ges., 1885 and 1890, Frank, Ber. d. deutsch. botan. Ges., 1887 and 1889. Atkinson, ‘‘ The genus Frankia in U.S. America,” Torrey Club Bulletin, 1892, p. 171, with plate. 100 SYMBIOSIS. which may increase to very large tubers, with surfaces resemb- ling a bunch of grapes (Fig. 21). In the large cells of the middle layers of the primary root-cortex of these growths, coils of very fine fungus-threads are sheltered; these extend year after year into the younger parts of the enlarging tuber- cles, and gradually disappear in the older parts. What may be the significance of these structures for plants possessing 7 Mi ea N iii Fig. 21.—Frankia alni. Root-tubercles on the Alder. (v. Tubeuf phot.) chlorophyll and furnished with normal roots is as yet unknown. Plants which have grown well for years in water-cultures do not show them. On account of the cork-covering with which these tubercles are furnished, it would seem that they are not adapted for taking nourishment out of the soil. Woronin described them first on the alder, Warming on Klaeagnaceae, while Moller proved their fungal origin. The species of fungi which produce these tubercles have been MYCODOMATIA OF THE ALDER, ETC. 101 provisionally distinguished as Frankia alni (Wor.) on alder, and Frankia Brunchorstii (Moll.) on Myrica Gale. _ Hiltner, after a series of experiments, states that first-year alders without tubercles do not thrive in soil free from nitrogen, nor do they take up nitrogen from the atmosphere; when, however, provided with root-tubercles they assimilate nitrogen. The tubercles also functionate in water, and soil rich in nitrogen has the affect of slightly increasing the assimilation of that element. The tubercle-fungus is at first parasitic on the alder, and is only of use to the plant after the tubercles have fully developed. Mycodomatia of the Leguminosae. All Leguminosae growing in their native soils exhibit the so-called tubercles. These are accessory formations of the primary root-rind and are furnished with vascular bundles connected with the root- bundles ; they consist of a cortex of normal cells surrounding an inner large-celled parenchyma with turbid cell-contents con- sisting of numbers of bacteria, (Bacterium radicola, Beyerink, or Rhizobium leguininos- arum, Frank.) * Frank describes minutely the formation of these tubercles.* The short rod-shaped microbe forces its way into a_ root-hair or epidermal cell, multiplies there, and is conducted to the inner cortical cells by plasma-threads continuous through — the cell-walls. A rapid division of the inner cortical cells is set up, till a tubercle is Fic. 22.—Rhizobium leguin- formed, which may still further increase jjgsarum. Root-tubercles on by continued cell-division from a meristem he slices apiariten at its apex. The bacteria multiply simul- taneously, and are transferred into the new cells where a great change comes over most of them; they enlarge very ' Hiltner, ‘Ueber d. Bedeutung d. Wurzelknéllchen v. Alnus glutinosa,” Nobbe’s Landwirtschaft. Versuchs-stationen, 1895. *Woronin, Mém. de lacad. des sci, de St. Pétershurg, 1886. 4 Lehrbuch d. Botanik, p. 271. 102 SYMBIOSIS. much and become club-shaped or dichotomously branched bodies without power of division, which may be designated “bacteroids.”! Brunchorst found the contents of the bacteroids Fic. 23.—Rhizobium leguninosarum. Root-tubercles on Robinia Pseudacacia (v. Tubeuf phot.) to disappear at the time of the fruit-formation of the host- plant. A small number of microbe-bodies still remain, according to Frank’s observations, capable of division, and these, after 1 According to Méller, they undergo fatty degeneration. MYCODOMATIA OF THE LEGUMINOSAE, 103 decay and break-up of the tubercles, reach the soil ready to bring about new infections. The great importance of the tubercles: of Leguminosae is that the plants bearing them are capable of taking up free nitrogen from the atmosphere and utilizing it, while without the tubercles they could not do so.) If Leguminosae be grown in soil rich in nitrogenous food-substances, the tubercles are not so well developed. According to Schneider,? the host-plant under the influence of the Rhizobium produces cellulose tubes, which become filled with the fungus. According to Beyerink,® these tubes consist of bacterial slime secreted by the Rhizobium. The epidermal tissue of the tubercles consists of a loose layer of cork with many intercellular spaces; this arrangement is stated by Frank* to facilitate the usual transpiration.® 'Hellriegel u. Wilfarth, Berichte d. deutsch. botan. Ges., 1889; also Zeitschrift f. d. Riibenzucker-Industrie, 1888. * Ber. d. deutsch. botan. Ges., 1894, p. 11. *Centralbl. f. Bacterologie u. Parasitenkunde, 1894. 4 Ber. d. deutsch. botan. Ges., 1892. * Schneider (Bulletin of the Torrey Club, 1892), gives a short account of American Rhizobia, and refers to the chief works on this subject. (Edit.) PART SECOND. SYSTEMATIC ARRANGEMENT OF THE CRYPTOGAMIC PARASITES. I. THE PATHOGENIC FUNGI OF PLANTS. THE vegetative body of the Fungi is a thalloid structure known as a mycelium, and composed of one or more hyphae. The hyphae are cells included in a firm wall of fungus-cellulose of varying composition; they grow apically, and hence are always filamentous in shape. In the simpler cases, the mycelium is a non-septate tube unbranched or branched; in the more complex forms, it consists of a system of hyphae divided into cells by cross-septa. By the union and anastomosing of numerous hyphae, a tissue may be formed not unhke the parenchyma of higher plants, hence receiving the name _ pseudo-parenchyma. From this tissue may arise distinct structures of many kinds, such as the sporophores of the Polyporeae, or strands of tissue like the well-known rhizomorphs of Agaricus melleus, or masses of resting-mycelium lke the sclerotia of Claviceps. It is also not uncommon to find a differentiation in the structure of the vegetative mycelium in the form of lateral outgrowths of the hyphae, developed as organs for the collection of nutri- ment—the haustoria,—or as organs of attachment—appressoria. Reproduction may take place sexually by the union of two cells or nuclei, the product of which is a spore or zygote capable of germination; or asexually by means of endogenous spores or swarm-spores, or by the abjunction of conidia of different kinds. Sexual reproduction is common amongst the lower fungi, but in the higher forms, if existent at all, it is very THE PATHOGENIC FUNGI OF PLANTS. 105 obscure and is replaced by numerous and complex modes of asexual multiplication. The lower forms of fungi, in the structure of the thallus, mode of reproduction, and adaptation to an aquatic life, exhibit distinct relationship with the Algae, particularly with the Siphoneae. Since the fungi do not possess chlorophyll, their nutrition is carried out by the absorption of organized material in a saprophytic or parasitic manner. Parasitic fungi are the cause of numerous and dangerous diseases of plants, whereas they only rarely bring about a diseased condition of the animal body. Bacteria on the other hand, which cause so many animal diseases, seldom affect plants injuriously. While many parasites are strictly limited to a parasitic mode of life, a large number naturally spend a part of their lives as saprophytes, and others may be made to do so artificially on nutritive substrata under suitable conditions. The latter method forms in fact a valuable aid for completing our imperfect knowledge of the life-histories of parasitic forms. In addition to the well-marked parasitic fungi, there are many saprophytic forms which become parasitic for a relatively short time or under special conditions of environment. ; The Fungi are divisible into two large groups, the lower fungi (Phycomycetes) and higher fungi (Mycomycetes). The systems instituted by various investigators differ not a little from each other. Three of the principal are: Der Bary. Zorr. BREFELD. (1) Phycomycetes (1) Phycomycetes (1) Phycomycetes (2) Ustilagineae (2) Mycomycetes (2) Higher Fungi’* (3) Ascomycetes (a) Basidiomycetes (a) Mesomycetes (4) Uredineae (>) Uredineae Hemiasci— Hemibasiii (5) Basidiomycetes (c) Ustilagineae (b) Mycomycetes (7) Ascomycetes Ascomycetes—Basidio- mycetes We shall in the present work consider the Fungi in the following order : Lower Fungi or Phycomycetes : Chytridiaceae, Zygomycetes, Oomycetes. Higher Fungi or Mycomycetes : Ascomycetes. Ustilagineae, Uredineae, Basidiomycetes. 106 PHYCOMYCETES. A. Lower Funer (PHycomMyceETEs).! The lower fungi possess, at least in their earlier stages, single-celled mycelia, which may in the higher families become branched. They reproduce sexually by oospores or zygospores, asexually by conidia. The Phycomycetes are divided into: Chytridiaceae, Zygomycetes, and Oonvycetes. (1) CHYTRIDIACEAE. The fungi of this family are chiefly parasites on aquatic plants, or on land-plants inhabiting moist places. The my- celium is one-celled, very rudimentary, or altogether absent. Asexual reproduction takes place by the formation of zoo- sporangia which usually produce uniciliate swarm-spores. Sexual reproduction is rare, and is effected by fructification of one cell by a fertilization-tube from another; the resulting bodies are zoosporangia which on germination set free swarm-spores. Hibernation is effected by resting-spores produced from sporangia in which the formation of swarm-spores is suppressed, and which become clothed in a thick membrane. Some of the species cause interesting deformations on the organs of plants. The Chytridiaceae include the families of Olpidiaceae, Synchy- triaceae, Cladochytriaceac, Rhizidiaceae, Hypochytriaceac, and Oochy- triaceae. Of these, only the first three contain species parasitic on higher plants. They occur epidemic only in moist situations, and rarely cause great damage to cultivated plants. OLPIDIACEAE. The whole vegetative body becomes a single zoosporangium or a resting-spore. Sexual reproduction is very rare. Olpidium. The vegetative body consists of a naked mass of protoplasm, the product of a single spore. This becomes later enveloped in a thin wall of cellulose, and forms a zoosporangium with a long neck through which the cell-contents are ejected as uniciliate swarm-spores. The cellulose membrane may become thicker and a resting-spore (sporangium) result, which in course of time germinates and gives off swarm-spores. ! Bibliography—A. Fischer in Rabenhorst’s Kryptogamen Flora, 1892. Schroeter in Hngler-Prantl Pflanzenfamilien, 1892. OLPIDIUM. 107 Olpidium brassicae, (Wor.)! (=Chytridium brassicae, Wor.) Cabbage-seedlings die if this fungus finds its way into the tissue at the neck of the root. The spherical sporangia are formed at this place, and their long necks project out of the cells enabling the uniciliate swarm-spores to escape. Lesting-spores with a warty thickened membrane occur in the cells of the epidermis. Fig. 24.—Chytridium brassicae, Wor. Cell containing three sporangia, two of which are discharging zoospores; one sporangium is already empty. Resting- spores inside the cells of a cabbage-plant. (After Woronin.) The disease is favoured by moisture, and restricted by dry surroundings. Ground subject to attack should be planted with crops other than cabbage. Olpidium trifolii, Schroet. (=Synchytrium trifolii, Pass.) Produces deformation of the leaves and petioles of Trifolium repens. The fungus lives in the epidermal cells. Olpidium lemnae, Fisch., in epidermal cells of Lemna. Olpidium simulans, De Bary and Wor. in TZaravacum officinale. A number of other species inhabit algae, spores, fungus- mycelium, pollen-grains, and eggs of Rotatoria. The genera Leessia, Pseudolpidiwm, Olpidiopsis, Pleotrachelus, Ectrogella, Pleolpidivm are parasitic only on lower plants, especially on algae. SYNCHYTRIACEAE. The whole mycelium divides up into a number of sporangia, which remain together as a sorus. The winter resting-spores ' Woronin, Pringsheim’s Jahrbuch f. wiss. Botanik, 1878 (Fig. 31). 108 PHYCOMYCETES. originate from the whole mycelium or parts thereof, and are isolated or united into a sorus. Synchytrium and Pycnochytrium.' Here, as in Olpidium, the mycelial hyphae are wanting, and the vegetative body escapes from the spore as a naked mass of plasma, which is later enclosed in a membrane. This vegetative body may also develop into a sorus of thin-walled sporangia : these separate in water, and each ejects from a pore numerous swarm-spores with a single long cilium. In the event of resting- spores being formed, the membrane of the vegetative body becomes thickened into a brown exospore. The resting-spores on germination liberate their contents as a single mass, or as several zoospores. In the former case the single mass divides at once into zoospores, or Into a sorus of sporangia, which ultimately give off zoospores. These fungi are found in the interior of cells, especially of the epidermis. The one cell inhabited by the fungus grows out as a simple papilla, or several neighbouring cells are also modified, and grow out along with the original one to form a eall-like swelling. The species of Synchytrium generally in- habit the epidermal cells of land plants, yet disease caused by them is commoner in moist than in dry situations. They cause so sheht deformation and damage to cultivated plants that they are of little practical importance. The Pycnochytrium of De Bary is regarded by Fischer as a sub-genus, by Schroeter as a genus. Synchytrium. The sori of zoosporangia are formed by direct division of the mature sporophore, and are enclosed in the colourless membrane of the mother-cell. Synchytrium taraxaci, De Bary and Wor. (U. 8S. America).” This produces, especially on Taraxacum, warty galls composed of a diseased epidermal cell, enlarged and surrounded by a wall of 1Schroeter: Cohn’s Beitrige z. Biol. d. Pflanzen I., 1875, and in Engler- Prantl Pflanzenfamilien, 1892. De Bary and Woronin, Bericht. d. natforsch. Ges. zw Freiburg, 1863. 2 We propose to indicate in this way species recorded in Seymour and Farlow’s ‘« Host-index ” for North America; British species by (Britain). (Edit.) SYNCHYTRIUM. 109 less swollen neighbouring epidermal cells. The sporangia contain reddish-yellow drops of oil, so that the swellings appear yellow. The organs attacked are much distorted and more or less stunted. The same fungus occurs on other Compositae, and is pro- bably identical with S. san- guineum of Schroeter, which produces dark red, crusty swell- ings on Cirsium palustre and Crepis biennis. Along with S. taravaci one often finds Olpidiuwm simulans. S. fulgens, Schroeter (U. 5. America), produces reddish- yellow swellings on the leaves of Ocnothera biennis and 0. - ° Fic. 25.—Synchytrium taraxaci. Leaves of muricata ; When resting-spores farazacum officinale so deformed by the fungus appear they form brown crusts. hit rer ear ras oe i a cae a The sori of zovsporangia are detached from the host-plant as single sporangia, which become scattered over the leaves. S. trifolii, Pass. (=Olpidium trifolii, Schroeter), is as yet little known. Other American species are :— S. papillatum, Farl., on (Geranium. S. decipiens, Farl., on Aimphicarpaca. S. vaccinii, Thomas. on Vaccinium, CGaultheria, Kalmia, Ethododendron, ete. Pycnochytrium. The sori of zoosporangia are not produced directly from the mature sporophore, but the contents of the sporophore pass out by a fine opening and form a thin-walled vesicle, the protoplasm of which breaks up into sporangia. Schroeter divides the genus into two sub-genera. (A) Mesochytrium. The discharge of the original sporophore ‘Halsted, ‘‘Cranberry gall-fungus”; . Jersey Agric. Coll, Bullet. 64, Dee. 1889. With figures. 110 PHYCOMYCETES. and the formation of zoosporangia take place in the cells of the living host-plant. In addition, spores are formed which have a resting period. (2) Only one kind of spore is formed; it has a resting period, and only proceeds to produce sori of zoosporangia after decay of the host-plant. (a) Chrysochytrivm: protoplasm contains a yellow oil. (b) Leucochytrium: protoplasm colourless. Each of these divisions is sub-divided into forms with simple vesicles, and those with compound, Mesochytrium. Synchytrium (Pycnochytrium) succisae, De Bary and Wor.! This parasite forms warty swellings and yellow spots, generally Fic. 26.—Synechytrium succisae. A, A mature sporocarp inside its host-cell. B, In the upper part of the cell a sorus of swarm-sporangia after escaping from its covering, which lies below it. C, Isolated swarm-sporangium. D, Swarm- spores. (After Schroeter.) on the radical leaves and base of stem of Scabiosa succisa. Infection is brought about in damp situations by means of swarm-spores. These have a single cilium, and bore their way into the host-cell. After entrance, they produce a plasma-mass, which becomes enclosed in a delicate membrane. The cell so formed sprouts at its uppermost pole, and gives rise to a new spherical cell, into which the older discharges itself. In the second cell numerous small sporangia are formed, so that it represents a sporangial sorus; beside it is always found the empty membrane of the first cell, The sorus breaks up later into single sporangia, which on opening, set free their contents 1Schroeter, ‘‘ Pflanzenparasiten aus d. Gattung Synchytrium.” Cohn’s Beitr, 2. Biolog. d. Pflanzen, 1875. PYCNOCHYTRIUM. 111] as zoospores swarming by means of a cilium. In addition, resting-spores are developed singly or in groups. The first effect on the host-cell of the entrance of a swarm- spore is that it becomes distinctly larger. At the same time neighbouring cells are so stimulated that they multiply and form a prominent ring-shaped swelling. The sporangia discharge their zoospores on the host-plant itself, and these pass into other cells of the swelling; here they form resting-spores and the host-cells die. Schroeter states that the resting-spores may be found from August onwards. S. stellariae, Fuck. On Stellaria media and S. nemorwm. The reddish-yeliow hemispherical swellings are produced on leaves, stems, flower-stalks, and sepals, either isolated or as a crust. The resting-spores generally form brown crusts. The host-leaves may be somewhat crumpled, but beyond this undergo little de- formation. ' Chrysochytrium. (1) Forming simple vesicles: Synchytrium myosotidis, Kiihn (U.S. America). The epi- dermal cells when attacked swell up to form club-shaped processes, while the cells with no fungus remain unaltered. The normal hairs of the host-plant are fewer on diseased than on healthy parts. This parasite attacks Boragineae, e.g. JZyosotis stricta, Lithospermum arvense. S. cupulatum, Thomas, produces red eruptions on Potentilla argentea and Dryas octopetala; diseased cells of the host-plant contain red sap. S. punctum, Sorokin. On Plantago lanceolata and P. media. S. laetum, Schroet. On (Gagea. (2) Forming compound vesicles : S. aureum, Schroet. Attacks many herbaceous plants as well as leaves of many shrubs and trees. Frequent on Lysimachia Nummularia, Fragaria, ete. The cells attacked are swollen and enclosed in a patch of enlarged neighbouring cells. 8. pilificum, Thomas. On Potentilla Tormentilla. The vesicles are hemispherical, and bear on their summits a tuft of abnormally elongated hairs. Thomas* found this species ' Clendenin (Botanical Gazette, 1894, p. 296) describes and figures a Synchy(rmum on Stellaria media in America (Edit. ). * Ber, d. deutsch, botan, Ges,, 1883, p. 496. We PHYCOMYCETES. on stems, flower-stalks, radical and cauline leaves, and floral envelopes. Leucochytrium. (1) Forming simple vesicles: S. punctatum, Schroet. On Gagea pratensis. S. rubrocinctum, Magnus, forms little red eruptions on Savi- Fraga ygranulata, the cell-sap of the host-plant becoming red. S. alpinum, Thomas. On Viola biflora. S. anomalum, Schroet. (U. 5S. America). On Adoxa Moscha- fellina, less common on Ranunculus Ficaria, Isopyrum thalictroides and Rumex Acetosa. The size and shape of the swellings, as well as of the spores, are very variable. (2) Forming compound vesicles : S. anemones, De Bary and Wor. (U. S. America). On Anemone nemorosa, A. ranuneuloides and Thalictrum purpura- Fic. 27.—Synchytriui anemones. The sporocarps form black points on leaves, petioles and perianth of the Anemone; the laminae are also stunted and distorted. (v. Tubeuf del.) scens, attacking stems, leaves, or flowers, and forming eruptions whose cells contain a red sap. In very bad cases, crumpling and swelling of attacked organs occur. PYCNOCHYTRIUM. LS S. globosum, Schroet. Where the attack is severe, this causes pearly swellings or incrustations; it frequents plants like Viola, Galium, Achillea, Sonchus, Mysotis. S. mercurialis, Fuck., is very common on Mereurialis perennis though seldom injurious to it. One severe case is thus de- scribed by Schroeter: “In spring the stem of the plant was covered by a thick uneven glassy crust, which in course of time became raised into wing-like processes running down the stem and coated on both sides with white granules of the immature parasite ; the leaves were completely rolled together, crumpled, and covered with glistening prominences as with fine silver sand. The plant in this condition developed poorly, scarcely flowered, and soon died, so that by the end of September few diseased examples could be found.” CLADOCHYTRIACEAE. The vegetative body is frequently a branched mycelium. It lives intercellular as a saprophyte, or intracellular as a parasite, and forms intercalary or terminal swellings, in which zoospor- angia or resting-spores are produced, then it disappears. Sexual reproduction does not occur. The parasite lives in and forms swellings on aquatic plants, or land plants in moist situations. The genera Urophlyctis and Physoderma contain species parasitic on higher plants; together with the saprophytic Cladosporangium, these are regarded by Fischer as sub-genera of Cladochytrivm, and as such they are also here regarded. Vrophlyctis has both zoosporangia and resting-spores, Physo- derma has only resting spores, Cladosporangium only zoospor- angia. Urophlyctis. The delicate mycelium is unbranched, or only slightly branched, and lives endophytic, boring through the walls of the host-plant. At the place where a hypha enters a_host- plant it forms a swelling or collecting cell (sammelzell), which generally becomes differentiated into a larger cell rich in contents, and an outer smaller one with few contents, but with fine terminal — bristles. From the collecting cells new hyphae originate and produce other collecting cells in neighbouring host-cells. The zoosporangia are situated outside the host-cells, H 114 PHYCOMYCETES. but send a hyphal process inside, which branches into a tuft of rhizoids. Resting-spores may be found, several in each cell. Cladochytrium (Urophlyctis) pulposum, (Wallr.), causes on leaves, stems, and flowers of Chenopodium and Atriplex glassy swellings, in the undermost cells of which are situated the zoosporangia. The resting-spores have brown shining walls and lie inside the cells. The zoospores are uniciliate. Cl. (Ur.) butomi, Biisgen. On leaves of Butomus wmbellatus. Black spores are produced containing resting-spores. The col- lecting cells have tufts of hair. Physoderma, Zoosporangia are absent. Resting-spores formed, several in each host-cell. Cladochytrium (Phy.) menyanthis, De Bary (U.S. America). On leaves and petioles of Menyanthes trifoliata this forms vesicles containing resting-spores. The collecting cells have terminal hair-tufts. Diseased leaves are generally smaller than healthy. Cl. (Phys.) flammulae, Biisgen, forms little swellings on leaves of Ranunculus Flammula. Cl. (Phys.) Kriegerianum, Magnus, causes transparent swell- ings on Carum Carwi. Cl. (Phys.) iridis, De Bary, on Jris pseudacorus. Fischer mentions other species on Scirpus, Alisma, Ranunculus, Potentilla anserina, Silaus pratensis, Sium latifolium, Phalaris, Glyceria, Symphytum, Mentha, Rumex, Allium, ete. Prunet! describes Cladochytrium viticolum as the cause of the much-discussed Brunisure of vine; also Cl. mori as a new disease of the mulberry.” The same authority * designates as Pyroctonum sphaericum, a parasite on wheat, which has become very abundant in Southern France. (2) ZYGOMYCETES. Unicellular fungi. Sexual reproduction does not take place by the fertilization of an ovum in an oogonium by an antheridium, 1Prunet, Compt. rend., 1894. 2Prunet, Compt. rend., cxx., 1895, p. 222. ’Prunet, Compt. rend., 1894, 11., p. 108. Be ZYGOMYCETES. Ts but by conjugation or union of two cells of the mycelium separated off from the ends of two hyphae by transverse walls. As a result of conjugation, a zygospore is produced, which is a resting-spore and corresponds to the oospore of the Oomycetes. , The zygospore puts forth a germ-tube, which becomes a mycelium bearing sporangia on sporangiophores. From each sporangium, spores, never swarm-spores, are set free, germinate, and produce a mycelium. Sporangia similar in form to the zygospores may be asexually produced on the mycelium. The unicellular and much-branched mycelium grows into its substratum, and is nourished as a rule saprophytically. The Hutomophthoreae cause important insect-diseases on Muscidae, Cabbage Butterflies, and caterpillars of Trachea piniperda (the Pine Beauty). Another common group of the Zygomycetes, the JMucorini, penetrate into bruised places in living fruits, and produce decay {see p. 180). Some other Zygomycetes are parasitic on fungi (Conidiobolus), some on animals. (3) OOMYCETES. These fungi possess a one-celled and much-branched mycelium. In their vegetative structure they most nearly resemble algae like Vaucheria. Reproduction is brought about, asexually by means of swarm-spores formed in sporangia (conidia also occur) ; sexually by oospores derived from oogonia and antheridia. There are three families of Oomycetes: Saprolegniaceae, Mono- blepharideae and Peronosporeae. Two of these groups contain parasitic forms: Saprolegniaceae (eg. Achyla prolifera, dangerous to Fish and Crustaceans); and Peronosporeae. PERONOSPOREAE. The greater number of the Peronosporeae live as parasites in the tissues of higher plants, and obtain nourishment generally by means of haustoria. The mycelium, in earlier life at least, has no dividing septa, and generally grows in the intercellular spaces of the host-plant, and sends haustoria into the cells. Reproduction is effected asexually by formation of swarm-spores in sporangia, and sexually by means of oospores. The latter are produced from the fertilization of an ovum in an oogonium by an antheridium whose contents pass through a fertilization-tube penetrating the 116 PHYCOMYCHETES. oogonium wall.1 No formation of spermatozoids occurs, as is the case in Vaucheria and other groups of algae showing close rela- tionship to these fungi. In certain cases the formation of swarm-spores in sporangia does not take place, but conidia are produced, which germinate directly into a mycelium. Preventive measures against the whole group consist in destruction (by burying or burning) of diseased and dead parts of host-plants which contain the hibernating oospores; by change of crop on infected fields; and by treatment with copper reagents (see Chap. VL). To the Peronosporeae belong the genera Pythium, Phytoph- thora, Cystopus, Basidiophora, Plasmopara, Sclerospora, Bremia and Peronospora. Pythium. The mycelium possesses no haustoria, and grows both between the host-cells and inside them. Cross-septa are not present at first, but later these may be found at irregular intervals. Pythium lives as a parasite in living plants, or as a saprophyte on a dead substratum. The conidia are of various forms, and either germinate directly into hyphal filaments, or discharge their contents into a bladder where zoospores are developed and liberated as free-swimming spores with two lateral cilia. The oogonia contain only one ovum-cell, which is fertilized by means of an antheridial tube applied to the oogonium. The thick- walled oogonia on germination produce hyphae or discharge zoo- spores. Pythium de Baryanum, Hesse? (Britain and U.S. America). This parasite-is injurious to the seedlings of various plants in gardens and fields. Some of its commoner hosts are maize, clover, mangel, millet, and many species of the Cruciferae; it has also been found on the prothalli ef Aguisetum and Lycopodium.’ It may also attack living or dead leaves and tubers of potato. The sporangia have a lateral beak-like outgrowth, into which the plasma passes and divides into biciliate zoospores. The 1In many species the fertilization-tube remains closed e.g. Plasmopara viticola. * Hesse, Pythium de Baryanum, Halle, 1874. Atkinson (Cornell Univ. Agric. Expt. Stat., Bull. 94, 1895), describes and figures this and other fungi causing “) bacteriosis, which will be considered amongst the bacterial diseases of plants. Lagerheim®? has pointed out that Solanum muricatum much cultivated in Ecuador on account of its edible fruit, has been for many years subject to attack from Phytophthora infestans; the fruits sicken and rot off before ripening. The !'This is a well-known point of controversy, for an interesting discussion of which we would refer to ‘‘ Diseases of crops,” Worth. G. Smith, 1884. (Edit.) 2 Boehm, Sitzungsber. d. Zool.-botan. Ges., Vienna, 1892. 3 Rivista Ecuatoriana, 1891. PHYTOPHTHORA. 121 v a Wty . 2 ae a ¢ Fic. 32.—Phytophthora infestans. The Potato disease. A, Potato leaf with brown spots and white patches of fungi on the lower side. 8B, Groups of conidio- phores emerging from a stoma close beside a hair of the potato leaf. C, Conidio pores and conidia, much enlarged. D, Leaf of potato much shrivelled up and srown, a8 in the later stages of the disease. (v. Tubeuf del.) a 122 PHYCOMYCETES. same author also quotes the disease on Solanum caripense at Quito, and on Petunia hybrida at Upsala. The potato disease is above all an associate of moist weather. In such circumstances, the conidia are produced very rapidly and the zoospores readily distribute themselves in the moist soil. There is thus greater risk to the potato crops on wet. soils. For wintering, potatoes as healthy as possible should be chosen. This is particularly the case if the tubers are required as seed : for the fungus-mycelium spreads from the tuber into the shoot. Whole tubers are less lable to infection than those cut or broken. Some varieties (¢g. thick-skinned) are less easily infected than others; such should be selected and _ bred. As a preventive measure the leaves may be sprayed with sordeaux mixture, or with a copper carbonate mixture’ By these means conidia and zoospores which alight on the plants are killed and their germination prevented. The leaves them- selves remain uninjured if the copper compound be used dilute enough. These compounds may even be beneficial to the growth of the host-plant, as was found by Rumm? for the vine, and Frank and Kriiger® for the potato. Frank and Kriiger found on using a two per cent. copper sulphate and lime mixture, in which the copper is known to be the potent constituent, that the potato leaves were stronger, their chlorophyll-contents greater, their power of assimilation and transpiration was increased, the life of the leaf was lengthened, and the yield and starch-contents of the tubers were increased. They regard the effect of the copper on the leaf as the result of a chemotaxic stimulus. Jensen recommends disinfection of seed-potatoes by heating at 40° C. for four hours. Ph. phaseoli, Thaxter, lives in young bean-pods and causes them to shrivel up. The fungus is as yet incompletely known, having only been observed in America where Thaxter+* reports great destruction of Lima bean (Phaseolus lunatus) near New Haven. 'See also § 12. Detailed experiments of this kind are frequently described in the magazines relating to agriculture. (Edit.) 2 Ber. d. deutsch. botan. Ges., 1895, p. 189. 3 Ber. d. deutsch. botan. Ges., 1894, p. 8. 4Thaxter, Botanical Gazette, 1889. CYSTOPUS. 123 Cystopus (Albugo). The mycelium is branched and grows between the cells of living plants, obtaining its nourishment by means of haustoria. The conidial cushions rupture the epidermis of the host. The conidia or sporangia are smooth-coated, and are produced acropetally in chains on short stalks from which they fall off separately when ripe. The sporangia germinate and discharge Fic. 33.—Cystopus candidus on plants of Capsella bursa pastoris. The fungus has caused distortion and thickening; the white porcellanous conidial cushions shew up distinctly on the dark background. (vy. Tubeuf phot.) swarming spores with two unequal lateral cilia. The egg-cells, produced singly in each oogonium, are fertilized by an antheridium. The thick-walled oospores remain enclosed in the intercellular spaces of the host-tissue, and on germinating in spring discharge swarming spores. Cystopus candidus (Pers.) Lev. White Rust. This fungus 124 PHYCOMYCETES. is very frequent on wild and cultivated Cruciferae throughout the whole world, and causes deformation of shoot, leaf, and flower. Fig. 34.—Cystopus candidus. B, Conidiophores isolated from the cushion ; the conidia or sporangia are united by intermediate cells. (C, Sporangia breaking up to form swarm-spores. D, Swarm-spores escaping. 2, Swarm-spores in motile condition. #, Swarm-spores come to rest and germinating. G, Two germ-tubes entering a stoma of Lepidium sativum; the stoma is shown from the inside, so that the spores from which the germ-tube;s arise are on the outer surface and unseen. (After De Bary.) Fic. 35.—Flower of Radish (Raphanus sativus) hypertrophied by Cystopus cundidus. The much-enlarged ovary stands out in the centre. The anthers are leaf-like; the petals are much enlarged and hang downwards; the sepals are somewhat enlarged. (Specimen from Botanical Museum of Erlangen, and photographed by Dr. Bruns.) The conidial cushions form thick white stripes with a porcellaneous appearance, by which they are easily distinguished from the cushions of Peronospora parasitica often present on the same plant. CYSTOPUS 125 Besides conidia, spherical oospores may also be present; these are generally produced on the stems of the host-plant, but also on flower-stalks and ovary-walls. The spherical conidia arise in simple chains on short coni- diophores, and are loosely connected by tiny intermediate cells. The conidial cushions rupture the epidermis and the ripe conidia fall oft to produce biciliate swarming cells (Fie. o4), These give rise to germ-tubes which enter the stomata of seedlings and Fr 6.—Flower of Radish hypertrophi swollen conidial cushions occupy the e Bruns’ phot.) develop to intercellular mycelia, fine short lateral twigs of which pierce the wall of the host-cells and become littl spherical haustoria. The oogonla arise as thick-walled spherical swellings on the mycelium. The antheridium, afte) apply Ing itself to the oo onium widens and projects a fine fertilization-tube through the wall the ege-cell. After fertilization is effected, the ege-cell eC] in a firm uneven membrane, and hibernates inside the « muy In spring the pla ma ol the oospore form Hnumerou 126 PHYCOMYCETES. swarm-spores which escape from the enclosing coats and germinate Fic. 37.—Cystopus portulacae, D. C. m, mycelium; j, basidia; c, spores with intermediate cells. (After Tulasne.) on seedling plants. De Bary! found germ-tubes of Cystopus entering all the stomata of Lepidium sativum and of Capsella, but they only developed further if the part attacked were the cotyledons. Magnus? observed an infection of Raphanus Raphanistrum i which the unopened buds were infected by swarm- spores. Oogonia may be found in the flowers of this same plant, whereas conidia alone only are present in Capsella. White rust is most commonly observed on Capsella, causing slight local swelling or marked hypertrophy. It is also found to injure radish (Raphanus sativus), horse radish (Cochlearia armoracia), cress (Lepidium sativum), species of cabbage and turnip (Brassica Napus, B. nigra, B. Rapa, B. oleracea), wall-flower (Cheir- anthus Cheiri), water cress (Nasturtium amphibium, etc.), caper-plant (Capparis spinosa), and other wild and culti- vated plants belonging to, or closely allied to the Cruciferae. Wakker? investigated the changes brought about on a number of Cruci- ferae by Cystopus. Some plants showed little or no deformation or anatomical alteration, others showed much. While the anatomical changes in the various species examined agreed in general, yet some showed a predominant or exclusive formation of conidia, others of oospores. The changes ' Morphology and Biology of the Fungi. English Edition. 2 Abhand. d. botan. Vereins d. Prov. Brandenburg, xXxxv. ° Pringsheim’s Jahrbuch, 1892. CYSTOPUS. 127 observed on Capsella may be summarized here: the fungus attacks all parts above ground, and causes enlargement of parenchymatous cells; it forms only conidia; formation of chlorophyll is increased; the formation of interfasci- cular cambium isdiminished or altogether suppressed ; the intrafascicular cambium retains its activity longer; accessory vascular bundles make their appearance ; o,/0,,2%;, and P. americana in North America. T. Farlowii (Sad.) on Prunus serotina in North America. T. confusa (Atk.) on Prunus virginiana in North America. T Rostrupiana (Sad.) on Prunus spinosa in Europe. T. communis (Sad.) on Prunus maritima, P. pumila, P. » £7 , americana and P. nigra in North America. T. longipes (Atk.) on Prunus americana in North America. T. rhizipes (Atk.) on Prunus triflora in North America. (ec) On Potentilleae : T. potentillae (Johan.) on P. sylvestris, P. canadensis, and P. geoides in Europe and North America. p. Aeseulus-stem on Euecyelieae: asci plump, cylindrical, with flat or rounded apex. (a) On Sapindaceae : T. aesculi (Ell. et Ever.) on Aesculus californica in California. (6b) On Anacardiaceae : T. purpurascens (Robins.) on Rhus copallina in North America. (¢) On Acerineae : T. acericola (Mass.) on A, campestre and A. Pseudoplatanus in Italy. T. acerina (Eliass.) on A, platanoides in Sweden. T. polyspora (Johan.) on A. tartaricwm in Europe. II. Genus. Magnusiella: asci ovoid or spheroidal. M. githaginis (Sad.) on Agrostemma Grithago in Denmark. M. umlelliferarum (Sad.) on Heraclewm Sphondylium, Peucedanum palustre, and P. Oreoselinum in Europe. & 152 ASCOMYCETES. The Hxoaseeae may be grouped, according to the symptoms of the disease produced, as follows; for this purpose we shall elass all the species as one genus, ‘ Hxvoascus’ (or Taphrina): I. Species which cause deformation of the ovary or other part of the fruit. E. prunt (Fuck.) on Prunus domestica, P. Padus, P. vir- gquniana. EL. Rostrupianus (Sad.) on Prunus spinosa. E. communis (Sad.) on Prunus pumilla, P. maritima, P. nigra, P. ameriwana. E. Farloww (Sad.) (£. varius, Atk.) on Prunus serotina, causing also deformation of twigs. longipes (Atk.) on Prunus americana. confusus (Atk.) on Prunus virginiana. rhizipes (Atk.) on Prunus triflora. cecllomophilus (Atk.) on insect-galls on the fruits of Prunus virginiana, FE. mirabilis (Atk.) on Prunus angustifolia, P. hortulana, P. americana. [Also species on Prunus subcordata, P. Chicasa, and P. pennsylvanica. | E. alni incanae (Kiihn) (2. amentorwm, Sad.) on Alnus incana. E. alni glutinosae (Tubeuf) on Alnus glutinosa. E. Robinsonianus {Giesh.) on Alnus ineanea. E, Johansonii (Sad.) on Populus tremula, P. tremulordes, P. grandidentata. E. rhizophorus (Johan.) on Populus alba. II. Species which (1) produce witches’ brooms, or (2) at least cause deformation of shoots ; asci produced on the leaves. (1) #. epiphyllus (Sad.) (£. borealis, Johan.) on Alnus incana (uniform grey coating of asci on both sides ef leaf.) E. turgidus (Sad.) on Betula verrucosa (coating of asci on under surface accompanied by slight crumpling of leaf). EB. betulinus (Rostr.) on Betula pubescens and £. odorata (coating of asci on under surface). ~ THE PARASITIC EXOASCEAE. 153 £. alpinus (Johan.) on Betula nana (coating on under surface). E., carpini (Rostr.) on Carpinus Betulus (coating on under side, and crumpling of leaf). E. cerasi (Fuck.) on Prunus Cerasus and P. avium (coating, chiefly on under side, and crumpling of leaf). E. insititiae (Sad.) on Prunus Insititia, P. domestica, P. pennsylvanica, (P. spinosa’); (coating on under side, and crumpling of leaf). E. acerinus (Eliass.!) on Acer platanoides; (asci on both surfaces). E. aesculi (EN. et Ever.) on Aesculus californica; (coating on both sides). E. Kruchii (Vuill.) on Quercus Lex. EL. cornu cervi (Giesh.) on Aspidivin aristatum. EL. Laurencia (Giesh.) on Pteris quadriaurita (with detorma- tion of leaves). (2) £. nanus (Johan.) on Betula nana (white coating on upper side). EL. bacteriospermus (Johan.) on Betula nana (coating on both sides). EL. decipiens (Atk.) on Prunus americana (coating on both sides). E. purpurascens (Ell. et Ever.) on Rhus copallina (crun- pling and red-colouration). E. Tosquinetii (West.) on Alnus glutinosa and A. glut.x incana (large blisters and elongation of shoots). E. pruni (Fuck.) on Prunus domestica (blistering and crum- pling). E. minor (Sad.) on Prunus Chamaecerasus. E. deformans Berk. on Persica vulgaris and Amygdalus communis (blistering and crumpling). E. cratacgi (Fuck.) on Crataegus Oxyacantha (spots and blisters on the leaves). E. mirabilis (Atk.) on Prunus angustifolia, P. hortulana, P. americana (on twigs, leaves, and fruits). E. celtis (Sad.) on Celtis australis (brown spots). E. githaginis (Rostr.) on Agrostemma Cithago. ' Svenska Vet.-Akad. Handl, 20, 1895, 154 ASCOMYCETES. III. Species which produce (1) pustule-like outgrowths, (2) leaf-spot, or (3) smooth coatings of asci. E. E. E. aureus (Pers.) on Populus nigra (nel. pyramidalis) and P. monilifera. polysporus (Sor.) on Acer tartaricum and A, Pseudo- platanus. bullatus (Berk. et Br.) on Pyrus communis and Cydonia japonica. carneus (Johan.) on Betula nana, B. odorata, and B. intermedia, coerulescens (Desm. et Mont.) on Quercus pubescens, Q. sessiliflora, Y. Cerris, Q. laurifolia, Q. rubra, Q. tinctoria, Q). aquatica. Sadebeckii (Johan.) on Alnus glutinosa. ulmt (Fuck.) on Ulivus campestris, U. montana, and U. americana (spots and blisters’. virginicus (Sey. et Sad.) on Ostrya virginica. . australis (Atk.) on Carpinus americanus. filicinus (Rostr.) on Aspidium spinulosum. . potentillae (Farl.) on Potentilla geoides, P. canadensis, P. sylvestris. . githaginis (Rostr.) on Agrostemma Githago. lutescens (Rostr.) on Polystichum Thelypteris. umbelliferarum (Rostr.) on Heracleum Sphondyliwm, Peu- cedanum palustre and P. Oreoselinum. ostryae (Mass.) on Ostrya carpinifolia (brown spots). betulae (Fuck.) on Betula verrucosa, B. pubescens, B. turkestanica (whitish spots). . flavus (Farl.) on Betula populifolia, B. papyracea. E. acericolus (Mass.) on Acer campestre and A, Pseudo- platanus. Jasciculatus (Lag. et Sad.) on Nephrodiwm (whitish spots). The following are some of the more important species of Exoasceae : : Exoascus pruni Fuck. (Pocket-plums). This attacks the ovaries of Prunus domestica (plum), P. Padus (bird cherry), and P. virginiana, causing the mesocarp to grow rapidly, whereby the fruits inerease in size and become much changed in form, THE PARASITIC EXOASCEAE. 155 while the stone, including the embryo, remains stunted. (Fig. 49.) The “pocket-plums” (fools or bladder-plums) dry up, and remain hanging on the tree till autumn. De Bary found on the plum a withering of calyx and stamens resulting from the development of the hymenium of this Hvoascus; on the bird cherry, according to Magnus and Wakker, enlargement of the stamens occurs. Sometimes a considerable thickening and twisting of the young shoots takes place, and their leaves curl up. Fic. 48.—Exoascus pruni. Twig of Plum, with four deformed fruits; one normal plum is partially hidden, the other is in the middle. + natural size. (v. Tubeuf del.) The mycelium hibernates in the soft bast of the twigs, and proceeds thence in spring into young shoots and_ ovaries. According to De Bary, the infected ovaries double their size in two days, and are full grown in eight days. The asci form a close layer under the cuticle of the ovary, and _ finally rupture it. 156 ASCOMYCETES. Exoascus Rostrupianus Sad. This fungus causes “ pockets ” on Prunus spinosa (sloe) similar to the preceding species. According to Sadebeck, the asci in this case are more slender. Fia. 49.—Fxrouscus pruni. Malformed Fic. 50.—Ezoascus pruni on twig of Prunus Plums—‘‘ pocket plums”; one which is Padus (at end of July). Four of the ovaries cut shows the rudimentary stone. are malformed. (v. Tubeuf del.) 4 natural size, (v. Tubeuf phot.) Fic. 51.—Exoascus pruni. Young twigs of Plum, showing effects of mycelium. The shoots are swollen and distorted, one diseased leaf remains hypertrophied and much crumpled; on one spur a normal and a ‘‘ pocket” plum are borne. Specimens from the Museum at Geisenheim. } natural size. (v. Tubeuf phot.) ve a rs THE PARASITIC EXOASCEAE. 157 Exoascus communis Sad. This produces pocket-plums on Prunus americana, P. pumila, and P. maritima in America. Similar “ pockets ” also oceur on Prunus subcordata, P. Chicasa, and P. pennsylvanica, in America, as a result of some Hvoascus. Exoascus Farlowii Sad. produces similar deformation of carpels and floral envelopes on Prunus serotina in North America. Exoascus Johansonii Sad. produces carpel- enlargement on the female catkins of Populus tremula, P. tremuloides and P. grandidentata; the contents of the asci are yellow. (Fig. 52.) The anatomy of the deformed ovaries has just been described by Sadebeck.! Exoascus rhizophorus Johan. causes similar — Fis. 52.—Broascus Johansonii Sad. on enlargement of the female catkins of Populus — Populustremula. (v. : Tubeuf del.) alba, Exoascus alni-incanae Kiihn (Hv. wmentorwm Sad.) This species is readily distinguished by the absence of a_ stalk-cell on the ascus. It causes increased growth and enlargement of the seed-scales of alder catkins, the fruit itself being seldom attacked. The fleshy bladder-like outgrowths at first appear as little red processes; later, the asci are developed on the outer surface as a whitish coating. On many of these red processes may still be recognized the trifid apex of the normal scale, (this is really formed from five smaller scales fused into a single large one with a trifid apex). A number of these red outgrowths are generally present on each infected catkin, yet the alders continue to flower vigorously every year. Wakker,? in investigating the anatomy of the deformed scales, found the following alterations :—the scales are increased to many times their original size and contain two cavities; all parenchy- matous cells become regular and iso-ciametric ; lignification of the elements of the wood is more or less interfered with, and fewer wood-fibres are produced; there is an accumulation of transitory starch. Exoascus alni-glutinosae Tubeuf. This is a new species distinguished by v. Tubeuf in 1895. It occurs in the Sudetic mountains, Italy, Denmark, and Sweden, on Alnus glutinosa. Its habit is similar to that of Hz. alni-incanae, but the asci ' Sadebeck (See Literature), 4. p. 144. * Pringsheim’s Jahrbuch, 1892. 158 ASCOMYCETES. contain only conidia, whereas those on ) delicate fungal filaments. Of the latter, those in the depth of the perithecium are known as paraphyses, and among them the asci originate ; others around the sides and opening of the perithecium are the periphyses, which grow inwards so as to close both pore and canal. Perithecia may occur isolated or massed together, and are frequently sunk in a special cushion of fungoid tissue, the stromata. The Pyrenomycetes may also produce chlamydospores and various forms of pycnoconidia and free conidia; these also are frequently developed on special stromata. According to Brefeld’s researches, the structures so well known as spermogonia with their contained spermatia are only pycnidia containing conidia, which have in many cases been artificially caused to germinate. The Pyrenomycetes include a large number of forms par- asitic on all parts of living plants, most of them are capable of existing for some part of their lives as saprophytes, and as ‘This fungus is described as causing a root-rot of Viola odorata in U.S. America (Connect. Agric. Exper. Stat. Report for 1891). (Edit.) “Went, Archief voor de Java-Suikerindustrie. 1893. * Reess and Fisch., ‘‘ Untersuch. tib. Bau u. Lebensgeschichte d. Hirschtriiffel.” Bibliotheca botan. Weft 7. 1887. With lus. 184 ASCOMYCETES. a rule they reach maturity only on the dead remains of host plants. Many of them are enemies of woody plants, and the mycelium of some can live in the elements of the wood itself, hence they constitute a dangerous group of wound parasites. The Pyrenomycetes may be sub-divided thus: 1. The Hypocreaceae having soft coloured perithecia often placed several together on a stroma. 2. The Sphaeriaceae with firm dark-coloured perithecia frequently embedded in a stroma. 5. The Dothideaceae with perithecia so embedded in a stroma that they have no distinct wall of their own. All three divisions include forms parasitic on plants. (1) HYPOCREACEAE. The Hypocreaceae consists of a single family bearing the same name. Of the seventeen genera contained therein only six contain plant parasites, viz.:—Guibberella, Calonectria, Nectria (including Nectriella), Polystigma, Epichloé, Claviceps. The re- mainder are saprophytic only, and do not come within the scope of the present work: they are—ZAelanospora, Selinia, Eleutheromyces, Hypomyces, Sphaerostilbe, Letendraea, Hypocrea, Pleonectria, Barya, Oomyces, and Cordyceps. Gibberella. The perithecia have a transparent blue or violet colour, and form tufts on the stromata. A stroma is not present in all the species. The spores are lhght-coloured, and spindle-shaped or oblong. G. moricola Ces et de Not. Passerini gives this as the cause of a disease of young twigs of mulberry. G. pulicaris (Fr.) is very frequently found on trees. (Britain). Calonectria. The perithecia are yellow or red, and occur isolated or several together. The asci contain spores composed of three or more cells, rarely of one cell. C. pychroa Desm. causes death to young leaves of planes (esp. P. occidentalis); it also multiphes by means of conidia (Fusarium platant). NECTRIA. 185 Nectria. Perithecia yellow or red in colour, and generally produced in close tufts on stromata of the same colour. The asci con- tain eight bicellular spores and few or no paraphyses. Conidia of various kinds and shapes are also produced. Nectrina cinnabarina Fr.! (Britain and U.S. America). The bright-red, button-shaped conidial cushions of this fungus may Fic. 77.—Nectria cinnabarina, with peri- Fic. 78.—Nectria cinnabarina. Portion of thecia on the dead bark of a still-living branch (magnified), Light-coloured cushions stem of Elm. Infection has evidently of conidiophores with conidia are breaking out begun at the wound of a cut branch near towards the upper end, and colonies of hard the middle, and extended outwards, (v. red perithecia towards the lowerend. (After Tubeuf phot.) Tulasne.) be found almost at any time on the dead branches of many deciduous trees, ¢./7., Aesculus, Acer, Tilia, Morus, Ulmus, ete.: also on Lonicera, Sambucus, Robinia, and Pyrus, in America.” ' Tulasne, Select fung., 1865. * Behrens (Zeitsch. f. Pflanzenkrankheiten (1895) ascribes to Nectria the very common tuberous swellings on the twigs of Abies balsamea; these, however, may arise without the agency of the fungus. 186 ASCOMYCETES. The dark-red masses of thick-coated, warty perithecia appear in autumn and winter on the dead branches only; the asci contain eight bicellular hyaline spores which germinate directly to form a mycelium. Infection of a new host-plant is effected by the mycelium, which enters by open wounds into living branches; it is quite unable to penetrate the living bark and is dependent on wounds. The mycelium spreads rapidly through the tissues of the host, especially through the vessels of the wood; the cam- bium and’ rind are not attacked directly, but are killed in conse- quence of the destruction of the wood.! The regions attacked in the wood appear as greenish stripes, and withering of leaves, followed by death of branch after branch, results in conse- quence of the growth of mycelium in the water-conducting elements of the wood. Fic. 79.—WNectria cinnabarina. Enlarged : section of perithecial colony. Germinating For protection against this ascospores. (After Tulasne.) ‘ =) and all other parasites, which find entrance by wounds, it is recommended to prune or dress trees only when necessary, and to paint all wounds with tar or tree-wax. This WNectria is one of the commonest parasites of our parks and fruit gardens, hence all branches already attacked should be removed and burned, likewise all blown timber which might serve as a nursery for production of spores or conidia. Nectria ditissima Tul. (Britain and U.S. America). This is a common parasite and a frequent cause of the canker of beech, apple, and other trees.2 The mycelium lives chiefly in the bark, causing it to die and form cracks. Under ordinary conditions all cracks and fissures are occluded or 1 Mayr in Hartig’s Untersuchungen a. d. forst-botan. Institut zu Miinchen, 111. Berlin, 1882. Brick, Arbeiten d. botan. Museums, Hamburg, 1892. Wehmer (Zeitsch. f. Pflanzenkrankheiten, 1894 and 1895), opposes Mayr’s con- clusions and holds that Nectria can penetrate intact, living bark. * Goethe, ‘‘ Ueber Krebs d. Apfelbiiume.” Rhein. Blatt f. Obst., Weim, u. Gar- tenbau, 1879. R. Hartig, Untersuch. aus d. forst-botan. Institut. za Miinchen, 1. NECTRIA. 187 grown over in course of time by the activity of neighbour- ing living tissues, but the rapid development of the mycelium of this Nectria prevents any such healing, and brings about death of more bark. As a result the so-called “cankers” are Nhe Fic. 80.—Nectvia ditissima. Canker Fic. 81.—Nectria ditissima. Canker on Hazel. The on a stem of Beech. (v, Tubeuf place of infection, a partially broken branch-fork. phot.) produced. The mycelium at first gives off tiny unicellular conidia on the bark, then later white cushions bearing fine conidiophores, from which are abjointed multicellular conidia, shaped like a sickle. Infection is brought about by the germination of spores or conidia on wounded parts of the 188 ASCOMYCETES. bark,! and even on young unfolding leaves. The red lemon- shaped perithecia break through the bark as compact patches. They are distinguished from the perithecia of N. cinnabarina by their smooth exterior and their smaller asci and _ asco- spores. Combative measures to be used are plentiful dressing of wounded places with tar, and the burning of all infected material. Nectria cucurbitula Fr.? (Britain and U.S. America), This parasite on conifers generally, is particularly injurious to spruce (Picea). It enters the host by wounds, such as those caused by the caterpillars of the spruce moth (Grapholitha pactolana), or by hail. The mycelium lives chiefly in the bark and bast; during the active growth of these tissues further extension of the mycelium is almost completely hindered by the formation of a secondary cork, but in the resting periods of these tissues of the host, new hyphae are rapidly produced. Reproduction is brought about by little unicellular, and larger multicellular sickle-shaped conidia produced on conidiophores. The mycelium frequently proceeds as soon as formed to give off the smaller variety of conidia. The dark-red perithecia are produced later on the same stromata as the conidia. The asci contain eight bicellular spores; the paraphyses are very delicate and slightly branched. The fungus sometimes occurs epidemic in spruce plantations, and may be the cause of many deaths. According to Magnus, the larch and cembran pine may also be attacked. As a pre- ventive measure all dead parts should be cut out and burned. Nectria Rousseliana Tul. lives in and kills leaves of the box (Buxus). Nectria pandani Tul.* is said to be the cause of a disease on Pandanus, also ascribed to Melanconium pandani. The Pandanus disease has been reported from the Botanic Gardens 'Young forests in districts subject to hailstorm, (e.g. on the lower Alps of Bavaria), may become completely infested with Nectria through hail-wounds. {Eptr.) “R. Hartig, Untersuch. aus d. forst-botan., Institut. 1., 1888. 3 Schroeter (‘Ueber die Stammfiule d. Pandaneae,” Cohn’s Beitr. z. Biol. d. Pflanzen. Bd. 1., 1895) suggests that this MWelanconium is a conidial form of Nectria. During the investigation of a case of a Pandanus killed in the Palm House at Edinburgh Botanic Garden, J. H. Burrage found both forms present an l agreeing in order of development with Schroeter’s observations. (Kdit.) | ] ’ j ; Pol ie al NECTRIA. 189 of Breslau, Berlin, Paris, Kew, Glasnevin (Dublin), and Edinburgh. Nectria ipomoeae Hals.' Stem-rot of egg-plant and sweet potato. In America this attacks young growing plants, and causes stem-rot. The Fwsarivm-stage developes as a_ white mouldy coating on the withered stem, and is followed later by flesh-coloured clusters of perithecia. Polystigma. On the leaves of species of Prunus, one finds bright-coloured spots, these are the stromata of this family, and in them are embedded pycnidia containing hook- shaped conidia. Perithecia embedded in a similar manner are developed after the fall of the leaves, they contain asci with eight unicellular elliptical spores, which are expelled on reaching maturity in spring. Polystigma rubrum (Vers.) (Britain and U.S. America). This appears as red circular spots on the leaves of plum and sloe. Micropyenidia are developed in summer on the under-surface of the leaf and give rise to curved conidia. The perithecia begin their development in summer, but only reach maturity in the following spring, after the leaves have fallen from the tree and lain on the ground over winter. The asci are club- Fig. 08 — Foleasirwin’ ruben shaped, long-stalked, and contain eight YA Jéf of Plum. The large projecting red spots contain peri- spores, which are set free in succession fi°ci Lath ey lore b mer from April to June. Germination ensues prop he kre el on young leaves, and in six weeks pycnidia reappear. A variety, “Amygdali Desm.,” is found on the leaves of the almond (Amygdalus communis). Frank and Fisch found in connection with P. rubrum, certain hyphae which they designate as_ trichogynes, while they regard the small form of conidia as spermatia which ! Description and illustrations in N. Jersey Agric. Exper. Station Report, 189). Pp YAL 7 } 190 ASCOMYCETES. fertilise the trichogyne and cause it to develop as an ascogonium. P. ochraceum (Wahlenb.) (P. fulvwm D. C.) causes yellowish- red spots on leaves of Prunus Padus. P. obscurum Juel. produces thickened leaf-spots on Astragalus alpinus and A. oroboides; on the under side these are whitish, on the upper side they show the spermogonia as red points. The damage caused by Polystigma is easily kept in check by burning infected leaves in autumn. Epichloé. The stromata form on the stems of grasses yellowish mould- like coatings in which the flask-shaped perithecia are embedded. Fic. 83.—Epichloé typhina, forming numerous white cushions, which completely encircle the grass-stems. (v. Tubeuf phot.) The asci are cylindrical, and contain eight thread-like unicellular EPICHLOE. 19] spores. The formation of perithecia is preceded on the same stroma by that of conidia. Epichloé typhina Tul. (Britain and U.S. America). This may be found on many grasses as a mouldy coating which surrounds the haulms and causes withering of the parts above it. The fungus not unfrequently attacks such fodder-grasses as Dactylis, Poa, and Phleum practense, causing severe loss where these crops are much grown. On the white stromata conidiophores are produced, and from them small, ovate, unicellular conidia are abjointed. These are followed later by perithecia embedded in the same stromata. The asci, of a somewhat yellowish colour, are long with button-shaped apices and contain eight thread- like spores, Ep. Warburgiana, Magn.” is an interesting species found on arrowroot {Maranta) in the Philippines. Claviceps. The sclerotia are black horn-like bodies, and on them the stromata are developed as stalked structures, with spherical heads, in which the flask-shaped perithecia are embedded. The asci contain eight thread-like spores. Claviceps purpurea (Fries*) (Britain and U.S. America). This fungus becomes most apparent, when in the stage of the well-known “Ergot” grains, bluish-black curved sclerotial bodies in which the mycelium perennates over winter. Ergot is found in the ears of our cereals, especially in rye, also in other cultivated and wild Gramineae. The sclerotia fall into the ground direct, or are sown out with the seed, and in spring or early summer produce a large number of stromata, each consisting of a violet stalklet carrying a reddish-yellow head. The ovoid perithecia are completely buried in the head of the stroma, and contain the asci, each with eight thread-like ascospores. The spores, after ejaculation, germinate on flowers of Gramineae, and the septate mycelium developes in the outer of the species of Hpichloé and other species of N. American graminocolous Hypocreaceae. (Kdit.) *Magnus, Internat. Bot. Congress, 1892.' *Tulasne, Annal. d. sci. natur. 3 sér. xx. Kuhn, Mittheilungen d. land- wirth. Institut, Halle, 1863. ‘Atkinson, G. F. (Torrey Club Bulletin, 1894, p. 222), proposes a revision 192 ASCOMYCETES. coats of the ovary, till gradually but completely it fills up the whole cavity. Outside the ovary the mycelium forms an Fic. 84.—Claviceps purpurea. Ergot. Sclerotia or Ergot-grains in ears of Rye. (v. Tubeuf phot.) irregular wrinkled white stroma or sphacelia, from the hollows and folds of which little ovoid conidia are abjointed from short CLAVICEPS. 193 conidiophores. A very sweet fluid, the so-called “honey-dew,” is separated from the sphacelia; this attracts insects, which carry the conidia to other flowers. Since the conidia are capable of immediate germination, and give rise to a mycelium which penetrates through the outer coat of the ovary, the disease can be quickly disseminated during the flowering season of the grasses. After the formation of conidia has ceased, the sclerotia become firmer, with a dark wrinkled cortical layer and an internal firm-walled pseudoparenchymatous Fig. 85.—Claviceps purpurea. A, Sclerotium with seven stromata (c/). B, median longitudinal section through the upper part of a stroma, the flask-shaped perithecia (cp) are embedded in the head. C, Perithecium in longitudinal section (highly magnified)—Ay, hyphal tissue ; sh, cortical tissue of the stroma ; cp, orifice of the perithecium. D, Isolated ascus ruptured, so that the thread-like asco- spores (sp) have begun to escape. (After Tulasne, from Sach’s Lehrbuch.) hyphal tissue. In this condition they are introduced along with grain into bread, which when eaten acts as a powerful poison, producing very serious results (Ergotism). The sclerotia are also used medicinally, and are collected for this purpose (Secale cornutum). Kobert (Fréhner, Lehrbuch der Toxikologie fiir Thieriirtze, 1890) states that Ergot contains three poisonous agents : (1) Cornutin, an alkaloid which produces that particular effect of ergot in causing contraction of the uterus, (2) Sphacelic acid, a non-nitrogenous, resinous, non-crystallizable sub- stance, insoluble in water and dilute acids, but soluble in alcohol, and N 194 ASCOMYCETES. forming, with alkalies, salts soluble in water. This is the real cause of ergot poisoning and gives rise to gangrene. In large doses it produces cramp similar to strychnine, and tetanus of the uterus. (3) Ergotic acid, a nitrogenous, easily decomposed glycoside, which has no effect on the uterus. It is more a narcotic which diminishes reflex excitability and finally stops it. Kobert experimented chiefly with cattle and fowls. He found that an acute course of the poisoning can be distinguished from a chronic; also a gangrenous ergotism from a spasmodic. The symptoms of the disease are: (1) Gastro-enteric, an excessive salivation accompanied with redness, blistering, inflammation, wasting and gangrenization of the mouth-epithelium; similar changes also occur on the epithelium of the gut, producing vomiting, colic, and diarrhoea. (2) Gangrenization and mummifica- tion of extremities, consisting of a drying-up, a dying-off, and a detach- ment of extremities, such as_ nails, ears, tail, wings, claws, toes, and point of tongue, (3) Spasmodic contraction of the uterus and consequent abortion. (4) Nervous phenomena such as in- sensibility, blindness, paralysis, etc. The presence of ergot may be de- tected both microscopically and spec- troscopically. The fungus may be combated by careful separation and de- struction of sclerotia, and by the use of clean seed.! Claviceps microcephala (Wallr.) (Britain). This is found on Phragmites, Molinia, Nardus, ete. It. has smaller sclerotia, which, according to Hartwich,? contain three times as much Ergotin as those of Cl. purpurea. Fic 86.—Sclerotia of Claviceps microcephala on Molinia coerulea. (v. Tubeuf phot.) ‘Smith (Diseases of field and garden crops. 1884. p. 233) describes and figures Claviceps purpurea var. Wilsoni on Glyceria fluitans near Aberdeen. It is distinguished ‘‘in being whitish or yellowish, instead of being pale purple in colour, and in the perithecia or conceptacles being almost free on an elongated club-like growth instead of being immersed in a globular head or stroma.” °Hartwich, ‘‘Sclérote du Molinia coerulea.” Bullet. de la Soc. Mycolog. de France. 1895. CLAVICEPS. 195 Cl. nigricans (Tul.) on Heleocharis and Scirpus, with sclerotia of a dark violet colour (Britain). Cl. setuloso (Quel.) on Poa. Stromata straw-yellow in colour. Cl. pusilla Ces. on Andropogon Ischaemum. (2) SPHAERIACEAE. The group of the Sphaeriaceae includes eighteen families, but only the following contain parasites of interest to us. Families : Trichosphaerieae, Melanomeae, Amphisphaerieae, Cucurbitaricae, Sphaerelloideac, Pleosporeae, Gnomonieae, Valseae, Diatrypeae, and Melanconideae. TRICHOSPHAERIEAE (including Coleroa and Herpotrichia). Coleroa. The perithecia have thin walls with radiating bristles, and sit superficially on the substratum. The asci have thickened apices, and contain eight two-celled, faintly coloured spores. Coleroa chaetomium Kunze, occurs on living leaves of Rubus caesius and Rk. Idaeus. In addition to perithecia, it forms conidia known as EHxosporium rubi Nees. C. alchemillae Grey. (Britain and U.S. America). On leaves of Alchemilla vulgaris. C. andromedae Kehm. On leaves of Andromeda polifolia. C. potentillae Fries (Britain and U.S. America). Leaves of Potentilla anserina. It forms perithecia which are situated on the leaf-ribs; also conidia (Muarsonia potentillae). C. subtilis Fuck. On leaves of Potentilla cinerea. C. circinans Fries. On leaves of Geraniwm rotundifolium and G. molle. C. petasitidis Fuck. On leaves of Petasites officinalis. Trichosphaeria. Perithecia small, spherical or ovoid, and more or less hairy. Paraphyses distinct. Spores with one, two, or four cells, We give this genus a wider scope than Winter, and include species with one, two, and four-celled spores of hyaline or light colour, and whose 196 ASCOMYCETES. other characters coincide; this seems to be all the more justifiable since one finds on the same species asci with spores made up of one, two, or four cells. Trichosphaeria parasitica Hartig.’ (Britain and U.S. America.) Everywhere in young naturally regenerated woods of silver fir, especially in damp places or where the plants are crowded, one finds partially browned needles hanging loosely from the twigs, held only by a fine white mycelium (Fig. 87). In addition to this, one finds in spring young twigs completely enveloped in mycelium, with all their Fic. 87.—Trichosphaeria parasitica on Silver Fir. The withered and dead needles hang loosely downwards, attached to the twig only by a white mycelium. (v. Tubeuf phot.) needles killed, so that the twig itself soon dies. The white mycelium grows especially on the under side of the shoot, and on the lower epidermis of the horizontal needles. Wurzelpilz, Weinstockfaule, Pourridié de la Vigne, Pourriture, Blane des {acines, Blanquet, Champignon blanc, Aubernage, Mal nero, Morbe bianco, ete. (Hartig’s Lehrbuch). wr DEMATOPHORA. 203 Fic. 93.—Vine-root with rows of black sclerotia exposed, and bearing bristle-like conidiophores here and there. (After R. Hartig.) Fic, 92.—Vine-stock with Dematophora necatrix (after a prolonged stay in a moist chamber). a, Fila- mentous myeelium passing over into rhizoctonia- : , i eaus strands (), which anastomose at cc. « and e, Fia, 94,—Portion of Fig. 93 after for- Rhizomorphs growing outwards from the interior. mation of conidiophores. x ¢. (After (After R. Hartig.) R. Hartig.) 204 ASCOMYCETES. Dematophora forms fine rhizoctonia-strands which grow through the soil from root to root. The mycelium kills the fibrous rootlets, and spreads from these into older roots to form rhizomorph-strands, which, however, have a_ structure quite distinct from those of Agaricus melleus. The rhizomorph- strands may pass out of the root into the soil, there to form a filamentous mycelium, or, remaining in the root-cortex, may produce rows of black tuberous sclerotia which on maturity break out to the exterior. On the sclerotia, or other parts of the mycelium, bristle-like conidiophores may be developed as branched panicles from which ovoid colourless conidia are abjointed. This enemy of the vine is rapidly assuming great import- ance. Thus, for instance, in Baden,’ there is no Phylloxera, but whole tracts of vine land are infested with Dematophora. According to Viala,*> Dematophora forms perithecia, which, however, only develop after artificial culture for several years on decayed plants. If this be so, the fungus should be classi- fied between the Tuberaceae and the Elaphomycetes. Berlese,* however, contests this view, and regards it as nearly related to Rosellinia. Hartig* suggests impregnation of the vine poles with creosote as a means of combating this disease. Strickeria. Strickeria Kochii Korb. develops its perithecia on the cortex of living Robinia Pseudacacia; its parasitism is however not yet fully established. CUCURBITARIEAE. Gibbera. The black perithecia, beset with stiff bristles, are developed in large groups on a dark pseudoparenchymatous stroma. Gibbera vaccinii Sow. (Britain). In damp situations amongst moist patches of Hypnwm and other mosses, one often finds the cowberry (Vaccinium Vitis-Jdaea) with its leaves and 1 Beinling, Das Auftreten v. Rebenkrankheiten in Baden, 1891. 2Viala, Monographie du Pourridié d. Vignes, 1891. 3 Berlese, Rivista di patologia vegetale, I. 4R. Hartig, Lehrbuch d. Baumkrankheiten, English edition, 1894, p. 87. GIBBERA. 205 twigs brown and dead (Fig. 95). If more closely examined, the twigs will be found to bear patches of coal-black, Fic. 96.—Gibbera vaccinii. Isolated ascus with eight spores; isolated hair from the outside of a perithecium. (vy. Tubeuf del.) Fic. 95.—Gibbera vaccinii on Cowberry. The perithecia form black patches on the living leafy branch, as well as the dead brown one. (v. Tubeuf del.) Fic. 97.—Gibbera vaceinii. Cross-section of Cowberry showing a patch of perithecia in section; the hairy perithecia contain paraphyses and asci with spores; a mycelium permeates the cortical tissue of the host. Short hooked hairs cover the epidermis of the stem. (v. Tubeuf del.) spherical perithecia, which are coated by short, acute, unicellular, black hairs (Fig. 97). The perithecia contain paraphyses and 206 ASCOMYCETES. asci, the latter with eight or fewer bicellular dark-coloured spores. The mycelium is dark-coloured, very vigorous, and furnished with many lateral bladder-like outgrowths; it permeates the whole cortical tissue as far in as the wood, and under the epidermis forms a brown pseudoparenchymatous stroma, which extends over the cortex, and gives rise to numerous perithecia. The living cells of the cortex turn brown in presence of the fungus-mycelium, and collapse, causing the whole shoot above the place of attack to wither and die. Cucurbitaria. The dark perithecia and pycnidia break through the epidermis in large numbers. The asci contain six to eight brown spores, divided by cross-septa. Cucurbitaria laburni Pers.’ (Britain). The spores of this fungus germinate on wounded parts of laburnum (Cyfisus Laburnum), and, as the branches of attacked plants soon die off, considerable damage to nursery stock may result. The mycelium spreads through the wood, particularly the vessels, in spite of the early stoppage of these by a yellow wood-gum. Diseased parts of the wood of living branches appear as dark strips; reproductive organs are produced in the bark, and there the plant attempts to isolate the diseased parts by con- tinued cork formation. If diseased, but still living spots on stems be examined, they will be found to include many yellow and black pustule- like swellings, some buried in the bark under a periderm eight to ten cells in thickness, others in process of breaking through or altogether exposed. Many of the pustules will attract attention from the presence of red, twisted, elongated tendrils on them. On the lower parts of dead branches the same appearances will be found, but, in addition, the periderm will generally be ruptured, and the openings so produced filled with spherical dark grey or black fructifications. These are variable in form, and amongst them can be distinguished some which are very large, round, smooth-coated, and light-coloured, with a round pore; others, which appear more warty, and have a depressed opening; while still others, generally smaller, have ly, Tubeuf, ‘‘Cucurbitaria laburni.” Botan. Centralblatt, Xxv1., 1886. a just described will be found others: CUCURBITARIA. an acute beaked pore. 207 Where the bark has been lost, a good lens may distinguish the spherical or ovoid dark-coloured perithecia. On the finer twigs the whole bark is often perforated by numerous tiny pyenidia, hardly distinguishable with the naked eye. If these various forms of fructification be submitted to microscopic examination, sections through the yellow pustules will show them to have that colour, because the transparent periderm has become loosened from the rest of the bark; underneath the corky layers will be found a red stroma of pseudo- parenchymatous hyphal tissue. This stroma by its growth causes a gradual rupture and loosening of the corky and other layers of the periderm; wherever this takes place, conidio- phores are developed, and give off numbers of tiny, hyaline, ovoid or cylindrical conidia. The stroma itself is somewhat spongy, and encloses numerous cavities which also become lined with conidiophores. At a later period the tissue enclosing these cavities may become dark coloured, so that structures similar to pyenidia are formed. In such cavities the red colour disappears, and the hyphae, coni- diophores, and conidia appear transparent. The real pycnidia appear later, and consist of a peridium of coarse pseudoparenchyma con- taining conidia similar to those just described (Fig. 99, A). From the openings of these pycnidia the conidia emerge as red tendrils, rising as much as one centimetre above the pore. Adjoining these forms of sporophore unde- veloped perithecia with young asci; dark- brown pycnidia with brownish-grey, multi- septate, compound conidia; or similar pycnidia Fic. 98.—Cucurbitaria laburni. Plant of La- burnum (diagrammatic); the branches 1, 2, 4 are still living, and were in full foliage during the preceding summer; «, b, c, d, ¢, places where the rind is dead and the cork-layer ruptured ; at a and b the perithecia are already developed, and the mycelium has extended into the wood. (After v. Tubeuf.) with unicellular spherical, brownish-grey conidia. Where the disease has made further progress, the pustules 208 ASCOMYCETES. will be found changing from yellow to black on account of the periderm and dead stroma becoming darker. On dead branches the large cushions of fructifications will be found to include: (a) perithecia with a warty exterior and pores set in a depression; (%) large pycnidia, standing out from the cushions, with brown smooth coats, and full of compound multiseptate conidia (Fig. 99, B; see below No. 3, a); (ce) other smaller pycnidia containing the same conidia, but whose pore is situated on a sharp prominence (No. 3, 0). All or any of the three forms may be present. Fic. 99.—Cucurbitaria laburni.. A, Stroma with pycnidia containing minute unicellular conidia. B, One of the large smooth pyenidia. (After v. Tubeuf.) The mature perithecia have a peridium consisting of a loose pseudoparenchyma with a rough warty exterior and a pore set in a distinct depression (Fig. 100.) The paraphyses are long, strong threads, often branched, and between them arise the long cylindrical asci with rounded ends. The normal number of ascospores is eight in each ascus, but fewer is no exception. In addition to the forms already described, pycnidia of still another sort occur (No. 1, ¢). They are spherical, with a dark-coloured coarse peridium, and are smaller than the stroma- pyenidia. These pycnidia contain no conidiophores, but give off CUCURBITARIA. 209 unicellular conidia, at first white, later grey. It is these pycnidia which cause the fine perforations of the periderm of twigs. Yet another form of pycnidia, previously known as Diplodia cytisi (Awd.), (No. 4). This, like the last, breaks through the corky layers of the bark. It has a peridium composed of loose pseudoparenchyma and, without the intervention of conidio- phores, produces two-celled conidia of a dark gvreyish-brown Fic. 100.—Cucurbitaria labuvrni. Perithecium isolated. 4, Ejaculating ascus with the inner membrane as yet unruptured, but emerging beyond the outer ruptured coat. (After v. Tubeuf.) colour. This form, however, I failed to find in the course of my investigation, although I looked through much material. Tabulating these various forms of fructification we have: A. PyYcnipIA. 1. White transparent, small, unicellular conidia on long conidiophores : (a) Free on the stroma. (b) Enclosed in cavities in the stroma : (a) In cavities as yet not resembling pyenidia. (8) In cavities with firm dark-coloured periphery. (ec) Enclosed in dark-coloured free pycnidia, with a peridium of coarse * pseudoparenchyma. 2. Brown, unicellular, round conidia, in little brown pyenidia (Fig. 99, a). 0 210 ASCOMYCETES. 3. Brown, multiseptate conidia : (a) In brown, very large, smooth-coated pyenidia (Fig. 99, b). (6) In darker and smaller pycnidia with pointed aperture. 4. Brown, bicellular conidia, in little dark pycnidia (Diplodia cytis7). B. PERITHECIA. 5. Brown, multiseptate ascospores, in perithecia generally of dark colour, and with depressed pore (Fig. 100). Cucurbitaria sorbi Karsten. This fungus appears to pro- duce disease in a manner similar to C. laburni. It was described by me in 1886! from specimens collected in the Bavarian forest-land from young Pyrus Aucuparia, They were easily distinguished in August by their withered twigs, both bark and wood being killed in tracts by the mycelium. In another locality I found well-developed perithecia, also on P. Aucuparia. Cucurbitaria pityophila Fries. occurs on the living branches of various conifers, eg. Pinus Cembra. SPHAERELLOIDEAE. Stigmatea. The naked perithecia are superficially seated on the sub- stratum. The ascospores, eight in each ascus, are clear and two- celled. The species are parasites. Stigmatea robertiani Fr. (Britain and U.S. America). Occurs on living leaves of Geranium Robertianwm. St. ranunculi Fries. On hving leaves of Ranunculus repens (Britain and U.S. America). St. mespili Sor. (U.S. America). This species appears in spring as reddish-brown spots on the leaves of wild pear-trees. At these places the epidermis becomes ruptured, and cushions are formed from which brown conidia are given off from short conidiophores. This stage was formerly known as Morthiera mespiult. The conidia are at first obovoid, but later seem to consist of four separate cells arranged in a cross, and each furnished with a transparent bristle. Each conidium produces a germ-tube which penetrates the epidermis, and in a month 'T have since found from Saccardo that this fungus was described by Karsten (Mycol. Fenn., tt. ‘‘ad ramos dejectos Sorbi aucupariae in Fennia merid. et media”); it was, however, unknown for Germany to that author. (Auth.) STIGMATEA. 211 new conidial cushions may appear. The mycelium itself is brown. From winter to spring, brown perithecia containing eight-spored asci may be found on the same leaves formerly occupied by the conidia. The colourless spores consist of two unequal cells; they germinate in May, before or after ejaculation from the asci, and bring about new infections. St. polygonorum Fr. occurs on leaves of Polygonum. (Britain and U.S. America). St. andromedae Rehm. On living leaves of Andromeda polifolia. St. alni Fuck. On living leaves of Alnus glutinosa. St. juniperi Desm. On living needles of Juniperus communis. Ascospora. The mycelium forms brown crusts under the host-epidermis, and there the perithecia develop. The asci are small and contain unicellular hyaline spores. The perithecia contain no paraphyses. Ascospora Beyerinckii Vuill.' The conidial form of this fungus (Coryneum Beyerinchii) produces a form of the “ gum- flux” of cherry trees. The mycelium lives in leaves of cherry, peach, plum, apricot, almond, which in consequence become spotted, and die off along with the young fruit. Mature perithecia may be found in spring. The fungus lives to a certain extent as a saprophyte. Sphaerella. The delicate perithecia are embedded in the tissues of the host-plant; they contain asci with two-celled colourless spores, but no paraphyses are present. Sphaerella laricina WHartig.” The needle-cast fungus of Larch. This fungus is the cause of a dangerous larch-disease found everywhere, except in mountainous localities over 1200 metres. The symptoms of disease consist in the needles becoming brown-spotted and falling prematurely in summer. Cushions of conidia are formed in June on the brown spots; these enlarge, and from their surface rod-shaped, four-celled conidia are *Vuillemin, Titres et travaux scientifiques, 1890. *R. Hartig, Forstlich-naturwiss. Zeitschrift, 1895, p. 445. Through the kindness of Prof. Hartig we have been enabled to add an account of this important new disease, with the accompanying figures. (Auth. and Edit.) Pile ASCOMYCETES. abjointed (Fig. 103); in the interior of the spots are produced tiny conidia (Leptostroma laricinum), incapable of germination. Fic. 101.—Tuft of larch needles, the greater number of which are more or less attacked by Sphaerella laricina. (After R. Hartig.) The rod-shaped conidia infect particularly the lower needles of the crown, and three weeks thereafter new conidial cushions Fic. 102.—Sphaerella laricina. Section through a diseased spot on a larch needle. The interior of the leaf is permeated with the intercellular mycelium. Two conidial cushions are shown ; from these numerous long rod-shaped conidia are given off externally (those of the upper cushion have nearly all been carried off by rain), while cavities inside the cushions are filled with micro-conidia. x 199, (After R. Hartig.) appear. Their distribution and germination are facilitated by wet weather. The perithecia (Fig. 104) are matured towards SPHAERELLA. 213 spring in the fallen needles, which lie on the ground over winter. The ascospores are mature and capable of infection at the beginning of June. In _ forests of pure larch, or in mixture with spruce, the ascospores are easily distributed by wind. In larch, underplanted with beech, the spores are kept down towards the ground by the canopy of beech foliage, so that, during the summer, they cannot be carried up to the larch crown. . . » . . . A wy Hartig gives the following interesting facts ais yt) on its distribution. ees oe. : : Fic. 103.—-a, Rod-shapec “As already remarked, the perithecia de- conidia before and after j a9 detachment from the velop in spring on the fallen larch needles, basidia. 4, Micro-conidia 7 = =se from the interior of the and in low-lying localities the spores reach cushions. x 44°. (After maturity at the beginning of June. New ® ts) conidial cushions are not found on the larch in our neighbourhood before July. The parasite has thus four months at its disposal Fic. 104, —Sphaerella laricina. Section through a diseased larch needle in June after it has lain on the ground from the previous year. The mycelium is thick, thick-walled, and of a light-brown colour. The perithecia contain asci and asco- spores. To the extreme right is a pycnidium containing little oblong conidia, alongside a perithecium. x 1$%, (After R. Hartig.) for distribution by means of conidia. As, however, we ascend into the mountains, the snow lies longer, so that the perithecia cannot begin to form so early, the ascospores are correspondingly late in reaching maturity, and the season during which the parasite may spread is still further shortened by the earlier commencement of winter. At an elevation of 1500 metres, 214 ASCOMYCETES. active vegetation begins about two and a half months later than in the plains, ie. at the beginning of June. The season of mature spores of Sphaerella is thus delayed till about the middle of August. On 26th September I found at this elevation only a few spots on the larch needles, and on these hardly any conidial cushions. By 28th September this larch plantation was already under snow.” “Tt will thus be seen, that while at a high elevation the larch can flourish with a vegetative period extending only to three and a quarter or four months, the Sphaerella has not the time necessary for its develop- ment, so that the larch, though much handicapped, remains healthy. Similarly with the larch in Siberia, it grows there, Fic. 105.—Enlarged asci. S$ in the mountains, very slowly, yet this a, Immature asci without paraphyses, as on April 30. parasite Can no longer reach it.” 6, Mature asci, from one of which the spores are escap- Sphaerella fragariae tral: Strawberry ing, ason June 1. x #12, : 1 : m - ‘4: (AlioriRS Hastig) 1 leaf - blight. In summer free conidia (Ramularia Tulasnec Sace.) and pyenidia are produced, while the perithecia ripen in spring. [This destructive disease of the strawberry has been recorded from all parts of the United States. It first appears on the upper surface as small reddish spots, which rapidly enlarge, the centres withering and browning. The growth of the plants and the crop-yield is seriously impaired.] (Kdit.) Sph. gossypina Atks.2 [Cotton leaf-blight is a disease on leaves of the cotton plant caused by the Cercospora-stage of this fungus. Small reddish spots appear on the leaf, enlarge, and become dry whitish spots with a red margin. The conidia are elongated and produced in long chains. The asci contain eight elliptical spores, which are shghtly constricted at the septum when mature, one cell being usually somewhat smaller than the other. This disease frequently accompanies that one known as “yellow leaf-blight,” or mosaic disease.] (Edit.) 1Trelease, Winconsin Haper. Station, 1885. Scribner, F. L., Report U.S. Dept. of Agriculture, 1887. Plate. Other papers by Arthur, Dudley, and Garman. 2 Atkinson, Bulletin Torrey Botan. Club, Vol. xvitt., 1891. SPHAERELLA 215 Sph. mori Fuck. causes a similar disease on leaves of mulberry (U.S, America). Sph. taxi Cke. On the yew.! Sph. longissima Fuck. On living leaves of Bromus asper. Sph. depazeaeformis (Auersw). On living leaves of Ovalis acetosella and Ox. corniculata. Sph. brassicicola (Duby.). On withering leaves of Brassicae. (Britain and U.S. America. Sph. laureolae (Desm.). On living leaves of Daphne laureola. Fic. 106.—Sphaerella fragaviae on leaf of Strawberry. The section through a spot shows formation of conidia. (vy. Tubeuf del.) Sph. hedericola (Desm.). On living leaves of ivy. (Britain). Allescher* describes other fungi on ivy. Sph. Gibelliana (Pass.), On living leaves of Citrus limonum and C. medica, Sph. polypodii (Rabh.). On living fronds of Polypodium vulgare, Aspidium Filiz-mas, Asplenium Trichomanes, Pteris aquilina, Sph. vitis Fuck. On withering vine-leaves. 1Worth. G. Smith, Gardener's Chronicle, xx1., 1884. * Allescher, ‘‘ Blattfleckenkrankheit d. Ephues,” Zeitsch. £. P/l.-krankheiten, 1895. 216 ASCOMYCETES. Sph. sentina (Fr.) (U.S. America). In spring of 1891 this caused at Geisenheim ! a severe spot-disease on the leaves of certain varieties of pear. Other related species occur on pear. There are numerous other species of Sphaerella. Saccardo gives 279 species, many of which are probably more or less prejudicial to plants in orchard or garden. None, however, are recorded as very injurious. Laestadia.” This genus is similar to Sphaerella, but has one-celled conidia; it is distinguished from Physalospora by the absence of paraphyses. L. maculiformis (Bon.) on living leaves of various trees. L. (Physalospora) Bidwellii (Ellis)? (Britain and U.S. America). The Black-rot of the Vine. This parasite attacks all young organs and shoots of the vine. On the leaves the symptoms are spots with dark sharply-defined margins, on which the pyenidia appear later as minute black pustules. The leaves die, but do not fall off, as with Sphaceloma ampelinum. The berries show disease when only the size of peas, and finally fall off singly or in clusters. The grapes are not dusty with a mealy powder, nor do they burst as in attacks of Oidium Tuckeri. Two kinds of pyenidia occur: one sometimes described as spermogonia, has very small rod-like conidia, borne on thread- like conidiophores; these conidia have not as yet been seen to germinate. The other pycnidial form (Phoma wvicola of Berk. and Curt.) contains forked filamentous conidiophores, from which one-celled ovoid conidia (stylospores) are produced and soon germinate by emission of a septate hypha. The latter form of conidia is produced after the spermogonia, and may be 1 Geisenheim Jahrbuch, 1892. * According to the laws of priority this genus must, as shown by Magnus (Oesterreich. botan. Zeit., 1894, p. 201), be called Carlia. Bon. * Bibliography : Cavara, Intorno al dissecamento dei grappoli della vite, 1888. Thiimen, ‘‘ Die Black-rot Krankheit d. Weintrauben.” Alleg. Weinzeitung, Vienna, 1891. Galloway and Scribner, Reports for 1888-89, U.S. Dept. of Agriculture. Viala and Ravaz, Les Progrés agric. et viticole. Montpelier, 1888-89. Rathay (1), Der Black-rot, 1891. With 19 figures (2). Bericht ib. eine nach Frankreich z. Erforschung d. Black-rot Krankheit Reise, 1891. With 7 figures (3). Der White-rot in die Weinlaube, 1892. Viala. Die Krankheiten d. Weinstockes. Linhart u. Mezey, Die Krankheiten d. Weinstockes, 1895 (Hungarian). U.S. Dept. of Agriculture. Numerous references in reports and bulletins, where details of treatment experiments will be found. LAESTADIA. PA by § found right on into autumn, even throughout the winter. Hibernating sclerotia are also produced, the cells of which grow out directly into septate conidiophores with oval conidia. Perithecia, externally resembling pycnidia, are formed in May and June on the fallen berries of the preceding year. The asci have gelatinous walls, which swell and burst so as to ejaculate their spores. Viala and Ravaz successfully infected living grapes by means of the larger conidia, and also by the germinating ascospores. The disease is one of the most dreaded in America. It has been found also in Europe, having been observed in France since 1885, though not as yet in Germany, Switzerland, Italy, or Spain. Moist situations are favourable to it. As with other diseases of the vine, the various varieties have different powers of resistance, and a judicious selection of varieties may prove a good preventive measure. According to Viala, the black-rot is found in the United States on both wild and cultivated vines, but never on the fruit of Vitis rupestris, V. Berlancieri, V. cinerea, V. Liusecomii, V. Monticola, and V. candicans, and very rarely on their leaves. The “vine-stocks” themselves suffer little or nothing from the disease. Rathay says that Vitis riparia, V. rupestris, and V. Solonis, so im- portant as grafting-stocks, are seldom affected; the green shoots of other species, however, may be attacked and the disease be transmitted through the graft-slip. For combating the disease. Galloway, Prillieux, and 1’Ecluse recommend Bordeaux mixture.’ Laestadia buxi (Desm.). The perithecia of this species are found as tiny points on yellow spots on the lower surface of green leaves of box. This fungus, regarded by Desmazieres as saprophytic, is said by Briosi and Cavara to be parasitic. PLEOSPOREAE. Physalospora. The perithecia are formed under the epidermis, but are otherwise devoid of covering; they contain asci and paraphyses ; the spores are one-celled, and ovoid or elliptical. ‘For details see Rathay (/oc. cit.), the American bulletins, ete. 218 ASCOMYCETES. Physalospora laburni (Bonord.) occurs on living twigs of Cytisus Laburnum. Ph. fallaciosa Sace. On withering leaves of Aletris and Musa in Berlin Botanic Garden. Didymosphaeria. Perithecia similar to Physalospora, but with two-celled spores. Didymosphaeria genistae Fuck. occurs on Genista pilosa. D. epidermidis (Fries). On living branches of Berbers, Sambucus, Salix, and Hucalyptus. (Britain and U.S. America). D. albescens Niessl. On living branches of Lonicera Xylosteum and Myricaria germanica. D. dryadis (Spegazz.). On living leaves of Dryas octopetala. (U.S. America). D. populina Vuill. Prillieux and Vuillemin! regard this as a parasite, and the cause of a peculiar dying-off of Populus pyramidalis throughout Germany ; Rostrup, on the other hand, ascribes this to Dothiora sphacriodes Fr. Prillieux regards Napicladiuin tremulae as a conidial form of Didymo- sphaeria ; Vuillemin, however, believes it to be saprophytic. Venturia. The perithecia are. embedded in the stroma, and have stiff bristles round the pore; they contain both paraphyses and eight-spored asci. The spores are two-celled, with or without colour. Venturia geranii Fr. occurs on the living leaves of Geranium pusillum, G. molle, ete. ' V. rumicis (Desm.). On withering leaves of Rumes. (Britain). V. maculaeformis (Desm.). On living leaves of Epilobium. V. vermiculariaeformis Fuck. On withered leaves of Evonymus europaeus and Lonicera Nylosteum. V. Straussii Sace. et Roum. This I have found as a parasite on Erica carnea in Tyrol. It is also said to cause a disease on Ericaceae in France. The various conidial forms at present placed amongst the “Fungi imperfecti” as Fusicladium are probably related to Venturia. Fusicladium dendriticum Wallr. on apple, and F. pirinum Lib. on pear, are at present the subject of an investigation at the hands of Aderhold,? who has, on account of their perithecia, 1 Bullet. de la soc. mycol. de France, 1892; Compt. rend., 1889; Revue mycol., 1892. 2 Aderhold, ‘‘ Die Perithecienform v. Pus. dendriticum” Vorliutig Mittheilung, Ber. d. deutsch. botan. Ges., 1894, p. 338. : , VENTURIA. 219 placed them in the genus Venturia. His investigations are, however, not quite complete. The scab or black spot of apple and pear is a very familiar disease in America and elsewhere. - It attacks leaves, young shoots, and fruits. Dirty greenish spots appear first, then enlarg- ing, they run together, and darken in colour till almost black. If the attack occur on young foliage, it may be dwarfed and killed ; the newly-formed fruit will in such cases be attacked, shrivel up, and fall. If the attack be deferred till the foliage and fruit are well advanced, then spotting results and the fruit remains hard, perhaps cracks. The conidia are oval, unicellular, and yellowish-brown ; they are produced from short conidiophores Fic. 107.—Venturia (Fusicladium) dendriticum forming brown spots on an apple; those still in the earlier stages have a radiate margin and bear conidia. The enlarged section shows two rows of large-celled parenchyma of the apple, covered by a stroma of pseudoparenchyma bearing conidiophores and conidia. (v. Tubeuf del.) with warty prominences which grow on spots of leaf or fruit (Fig. 107). The perithecia (as yet described) are distinguished by black bristles surrounding the pore, and occur on fallen leaves. The asci contain eight greenish ovoid spores with two or three cells. In addition to the injury to leaves and destruction of young fruit, the disfiguration of the apples is a cause of considerable monetary loss. Dilute Bordeaux or copper sulphate mixture applied before the opening of buds, and once or twice after 220 ASCOMYCETES. “setting” of the fruit, is recommended. No fungicide should, however, be applied towards the ripening season.! Gibellina. The spherical perithecia are embedded in the stroma, their necks projecting. The asci contain eight brownish spores, oblong or spindle-shaped, and bicellular. Gibellina cerealis Pass. This parasite of wheat has hitherto been fairly common in Italy; recently it has appeared with disastrous effect in Hungary. According to Cavara, it produces on the under part of the stems, grey plate-like coatings with a brown margin; these may remain as spots, or enlarge till they form a ring round the stem. The perithecia are little black points arranged in rows, and embedded under the epidermis, except the black projecting necks. The asci have thin walls and break up inside the perithecia; they contain eight spores arranged in two rows. The spores are spindle-shaped and bicellular, but their germination has not as yet been observed. The mycelium is found in all the host-tissues, besides forming a stroma-like sheath round the stem. The plants attacked become brown and limp in early summer, and no fruit is produced? Cavara recommends early removal and burning of affected stems, and the cultivation of crops other than cereals on the infected ground. Leptosphaeria. Perithecia black, rarely with bristles; at first they are em- bedded in the host, without a stroma. The spores occur four to eight in each ascus; they are spindle-shaped and _ generally multicellular by means of cross-walls only. Thread-like para- physes are always present. Leptosphaeria herpotrichoides de Not. This species, gene- rally regarded as a saprophyte, was found by Frank? as a parasite on rye. The stalks attacked break over at a node or ‘Further details of treatment-experiments are given in Gall and Scribner’s “* Report on Experiments for 1889,” U.S. Amer. Dept. of Agriculture Bulletin 11. ; also in the Bulletins of various Experimental Stations. The above account has been considerably extended by aid from the American literature. (Edit.) “Cavara (Zeitschrift f. Pflanzenkrankheiten, 111., 1893, p. 16) gives a detailed account of this fungus with illustrations. ‘Frank, Zeitschrift f. Pflanzenkrankheiten, 1895. LEPTOSPHAERIA. to to ww just over the root, thus resembling the symptoms accompanying an attack of Hessian Fly. L. tritici Pass. is said by Frank to be destructive to wheat in Germany. (See also Cladosporium). L. subtecta Wint.!’ In Tyrol the perithecia of this species accompany disease of the leaves of Erica carnea. Simultaneously Hypoderma ericae Tub. and Sphaeria ericina Tub. were found, the former appearing to cause the disease (see p. 234). L. anceps Sacc. On living branches of Ribes nigrum. L. vitigena (Schulzer). On living tendrils of the vine. L. circinans Sacc. (see Rhizoctonia p. 201). L stictoides Sace. on Liriodendron tulipiferu is an American species. Pleospora. The black perithecia are not developed on a stroma, and are at first concealed in the host-tissues only. They contain paraphyses and eight-spored asci. The spores are multiseptate, and generally coloured. Pleospora hyacinthi Sor.” produces black coatings on the bulb-scales of hyacinth. The mycelium inside the tissues is colourless, but outside is dark red, and its presence causes disease of healthy parts. Certain perithecia which appear on the bulb- scales in autumn may perhaps belong to this fungus. P. tropaeoli Hals. is given as a disease of Tropaeolum in U.S. America.” P. hesperidearum Catt. The conidial form (Sporidesminum hesp.) appears as a black coating on the orange. P. ulmi Fr. (var. minor) Allescher, causes a leaf-spot on young elm-seedlings, and the leaves drop off prematurely. P. napi Fuck. is the cause of rape-smut. Leaf-spots carrying conidial cushions (Sporidesmium exitiosum Kiihn) appear on the rape and other allied root-crops. Other “ black smut-diseases” have been ascribed to Polydesmus (Spori- desmium) exitiosum (vv7 DVauev) on carrot; Helminthosporium gramineum Rabh. causing withering of rye and barley leaves; and Sporidesmium putrefaciens Fuck. which attacks and kills the young heart-leaves of beet- root. ty, Tubeuf, Botan. Centralblatt, Xx1., 1885. *Sorauer, Handbuch d. Pflanzenkrankheiten, u. Aufl, p. 8340; and Untersuch. ith. die Ringelkrankheit u. d. Russthau d. Hyacinthen, Leipzig, 1878, 3N. Jersey Agric. Exper. Stat, Report, 13, 1892. bo bo bo ASCOMYCETES. Dilophia. The genus is parasitic and causes swellings. The perithecia remain permanently embedded in the tissues of the host-plant. The asci contain eight transparent, thread-lke, finely-pointed, multicellular spores. Dilophia graminis Sacc. (Britain). This causes deformity of the leaves and inflorescences of wild grasses; also of rye in France, and wheat in England and Switzerland. Fuckel assumes a relationship between this species and Dilophospora graminis Desm., but this we regard as doubtful. Ophiobolus. Perithecia scattered and almost spherical; they contain para- physes and eight-spored asci. The spores are hyaline or yellowish, thread-like, and unicellular or septate. The fungus is minute and inhabits stems and haulms. Ophiobolus graminis Sacc. was indicated by Prilheux, Delacroix, and Schribaux as the cause of a cereal disease in France. The cereals attacked broke over very easily near the eround ; they continued to develop, but produced ears of a poor quality, and often quite withered. The disease was designated “maladie du pied des céréales,” and described in Jour. d’ Agric. practique, 1892; also under the name “la maladie du pied du blé” in Travaux du labor. de pathol. végéale inst. agronom., 1890. The perithecia have a curved lateral beak; the asci contain eight long, spindle-shaped, multiseptate spores. Frank also records this disease as injurious to wheat in Germany in 1894. GNOMONIEAE. Gnomonia. Perithecia without a stroma, and generally remaining embedded in the host-tissues, with only a beaked opening pro- jecting ; they contain no paraphyses. The asci have a thickened apex with a fine central pore. The hyaline spores consist of from one to four cells. Gnomonia erythrostoma Auersw.' This is the cause of an ‘Frank, Ber. d. deutsch. botan. Ges., 1886 and 1887; also Zeitschrift f. Pflanzenkrankheiten, 1891. ee. GNOMONIA. 223 ‘ epidemic disease of the cherry (Prunus avium and P. Cerasus), observed for several years past in North Germany and _ else- where. The fungus attacks the leaves, and there the mycelium grows. The leaves wither prematurely, but remain all winter hanging from the tree by a reddish-brown mycelium. Pyenidia and perithecia are produced in the leaves, the latter reaching maturity in spring, when the two-celled ascospores are ejaculated. The pycnidia contain short conidiophores bearing hook-shaped conidia. The fruit is also attacked, and ripens unequally, so that the cherries are distorted; then they crack and rot. Frank has succeeded in carrying out artificial infection. This takes place in June, and immediately on germination the germ- hypha produces an attachment-dise on the host-epidermis, whence a hyphal filament penetrates the epidermal wall, grows through the cell, and reaches an intercellular space. A thick septate mycelium is formed and spreads, especially amongst the spongy parenchyma. There is no stroma, and the perithecia hibernate on dry leaves. Frank recommends the plucking and burning of dead leaves hanging on the trees. This must of course be done throughout the whole district attacked. In one part of Prussia (Altenlande) this precaution was taken twice each winter for two years, with the result that the disease, which had long completely ruined the cherry crop, disappeared, and the harvest increased to its former amount. Gnomonia quercus-ilicis Berl.’ causes brown spots on leaves of Quercus Llex. VALSEAE. Mamiania. Perithecia produced in a black stroma, from which their long necks project. The asci have a thickened apex, and contain eight oval hyaline spores with one or two cells. Mamiania (Gnomoniella) fimbriata Vers. (Britain and U.S. America). The stromata of this appear in summer as little black cushions on the leaves of the hornbeam (Carpinus). The perithecia are developed in these spots, and their long black beaks projecting distinctly above the surface of the leaf cause ' Berlese, Rivista di Patologia vegetale, 1. 224 ASCOMYCETES. rupture of the epidermis. Numerous leaves may be diseased and each ‘carry many cushions, yet Vuillemin, who described the disease,t does not believe the host-plant is affected to any serious degree. M. (Gnom.) coryli Batsch. (Britain and U.S. America). The black stromata are found in withered spots on the leaves of hazel (Corylus); as a rule, each stroma carries only one peri- thecium with a long beak similar to that of IZ fimbriata. Fic. 108.—Mamiana fimbriata on Carpinus Betulus. Leaf of Hornbeam seen on lower surface. Stroma (enlarged), with the long black necks of the perithecia projecting from the ruptured leaf-epidermis. (v. Tubeuf del.) Valsa. A stroma is generally present, but is of very variable appear- ance ; embedded in it are the perithecia, with only their beak- like mouths projecting. The spores are hyaline or heht-brown, unicellular, and generally bent. No paraphyses are present. Valsa oxystoma Rehm.” This causes disease and death of branches of Alnus viridis in the Alps. The symptoms are withering and drying up of single branches on an otherwise ereen bush. This disease causes severe loss in the Tyrol, 1 Vitres et travaux scientifiques, 1890. 2v. Tubeuf. ‘‘ Zwei Feinde d. Alpenerle,” Yorstlich-naturwiss. Zeitschrift, 1892. VALSA. 225 where leaves of the alder are dried in summer for use as winter-fodder for goats. In the branches attacked, a mycelium is developed in the vessels of the wood, whereby the supply of water is stopped and the bark dries up. Black lens-shaped stromata arise under the epidermis of the twig and rupture it. The perithecia are produced under the stromata in the bark, and communicate Fic. 109.—Valsa oxystoma on Alnus viridis, A, Portion of branch with stroma of Valsa breaking through the periderm at four places. 3B, Enlarged section through a stroma from A. C, Asci and ascospores, isolated from a perithecium. D, Portion of younger branch with periderm ruptured by stromata, in which, however, the perithecia are not yet developed. #, Enlarged section through a stroma of D. (After v. Tubeuf,) with the exterior by means of long projecting necks, The asci contain eight unicellular spores of a slightly bent, rod- like shape. Maturity is reached on the dry dead _ twigs. Externally this disease is identical in appearance with one I ascertained to be due to a beetle (Cryptorhynchus lapathi), p 226 ASCOMYCETES. the larva of which bores canals in the wood of alders, birches, and willows, causing them to die. Most of the other species of Valsa cause only leaf-spot, or occur on dead leaves. Anthostoma. The perithecia are embedded in the substratum or ‘stroma, and have generally long necks. The asci contain eight brown or black, oval, unicellular spores. Paraphyses are always absent. Anthostoma xylostei (Pers.) occurs on living and dead branches of Lonicera Xylosteum. (Britain). Anthostomella pisana Pass. lives on leaves of Chamerops humilis and kills them. DIATRYPEAE. Calosphaeria. No stroma is formed, the perithecia arising singly or in groups in the bark under the periderm; they have often long beaks. The asci are club-shaped, and frequently long-stalked ; they contain eight or more spores, which are little, unicellular, and somewhat curved. Calosphaeria princeps Tul. occurs on living branches of cherry and plum. (Britain and U.S. America). Quaternaria Personii Tul. has black perithecia and, according to Will- komm, causes death of twigs of beech. (Britain and America). MELANCONIDEAE. Aglaospora. The perithecia are beaked and embedded in the stroma. The spores are furnished with appendages. Aglaospora taleola Tul.’ (Diaporthe taleola Fries. and pro- bably nearly related to those MMelanconi with appendages on their spores, e.g. Mel. thelebola, previously known as Aglaospora thel. Tul.). (Britain and U.S. America). This fungus causes a disease of twigs and young stems of oak which have not 1R. Hartig, ‘‘ Hine krebsartige Rindenkrankheit d. Eiche,” Forstlich-naturwiss Zeitschrift; 1893. AGLAOSPORA. Fic, 110.—Examples of Oak-stem attacked by Aglaospora taleola, 1, Portion diseased for two years; a, the portion still healthy (x 4). 2, Portion diseased for four years ( x 4). 38, Section with spots diseased for four, seven, and ten years respectively (x 4). (After R. Hartig.) 228 ASCOMYCETES. as yet formed a bark. Portions of the rind become brown, dry up, and peel off; this on stronger twigs may be followed by a more or less complete occlusion of the wounded part. The browning also extends into the underlying wood. The mycelium is found both in rind and wood, where it probably obtains entrance through small wounds in the bark. In the second year after infection, a circular stroma is formed in the bark under the periderm. Sickle-shaped conidia are Fic. 111.—Aglaospora taleola. Portion of cortex with embedded stromata. a, Corky layer; b, after removal of corky layer; c, section of stroma. (x 7.) (After Hartig.) Fic. 112.—Section of stroma of Aglaospora. a, Boundary Fig. 113.—a, Conidia; b, asco- of stroma formed of dark brown fungus-mycelium ; spore of Massee (British Fungus-flora, tv., 1895). ‘‘There is no evidence of this species having occurred in Britain.” ®Rostrup, Tidsskrift for Landékonomie, 1890. 266 ASCOMYCETES. most from this fungus; red clover was less affected, though the disease often had its origin in that species; while white clover was least often attacked. He recommends keeping out Medicago from clover mixtures, and the addition of a large proportion of grass-seeds. Fields badly affected should be kept out of clover-cultivation for several years. English and French white clovers he found to be very sensitive, but distri- bution of the fungus did not take place by means of seed. Scl. tuberosa (Hedw.) (Britain and U.S. America). This on the rhizomes of Anemone nemorosa causes formation of sclerotia larger than filbert-nuts. The ascospores on germination produce groups of flask-shaped processes from which are given off chains of spherical conidia incapable of germination. Certain pyenidia which appear on the anemone-plants or on the sclerotia belong to a parasite (Pycnis sclerotivora Brefeld). Scl. bulborum Wakk.! (Britain). Wakker observed this form on hyacinth, onion, etc. It is very similar to Sel. trifoliorwm, but the hyacinth-fungus will not infect clover, and vice versa. The leaves attacked become rotten and the plants die.” Eriksson describes, from Wermland (Sweden), a destructive appearance of bulb-rot due to sclerotia, which he attributed to Sel. Fuckeliana De Bary. Scl. candolleana Lev. on oak-leaves. Appendix, Sclerotia of Unknown Affinity. Scl. oryzae Catt. tice plants (Oryza sativa) are often attacked by this Scelerotium, and a disease called “ Brusone ” produced. The sclerotia are found during June in the sheaths and stems. The symptoms are blackening at the base of plants and withering of upper parts. Scl. rhizoides Auersw. occurs on living plants of Phalaris arundinacea, and Calamagrostis; also on dead leaves of Dactylis glomerata. Scl. rhinanthi Magn.® forms sclerotia on the roots and root- l1Wakker, Allgem. Vereenig. voor Bloembollencultur, 1883-84; also Botan. Centralblatt, XxX1x., 1887. 2G. Massee (Gardener's Chronicle, Vol. xvt., 1894) gives description and figures. 3 Verhand. d. botan. Ver. d. Prov. Brandenburg, Xxxv. 1894. ae SCLEROTIA OF UNKNOWN AFFINITY. 267 neck of living Rhinanthus minor: these bodies begin their development in the cambium and bark, which they kill; after- wards the wood itself may be attacked. Sclerotinia with Botrytis-conidia.' Scl. Fuckeliana De Bary. This Sclerotinia is distinguished from all preceding ones by its passing through a Botrytis- conidia stage (Botrytis cinerea). If conidia are sown out on plum-juice gelatine, there appear within fourteen to twenty-one days round groups of sclerotia, which soon give rise to conidia. From such artificially- reared sclerotia I have never succeeded in getting the Peziza-fruit, so easily cultivated from sclerotia gathered in the open-air (¢.4. from vine leaves)” Thus the actual proof that Sel. Fuckeliana and Botrytis cinerea ave pyciiank’ Ripeeut at stages in the life of the same fungus is not Rare Bey reached by this experiment.8 The two forms °? are, however, very frequently met together. The sclerotia of Sel. Fuckeliana are produced in the mesophyll of the leaves, also in the parenchyma and epidermis of the host-plants, but never in the wood. Peziza-fruits with flat apothecia are produced from them. Sclerotia are found in vine leaves and over-ripe grapes (Fig. 140), especially of the Riesling, Orleans, and Sylvaner varieties.*| Other plants and fruits may also be attacked. Diseased parts become brown from the effects, of the parasitic mycelium, and die off. The mycelium can only live parasitic after it has been strengthened by a pre- vious saprophytic existence. Ascospores are thus unable to effect direct infection. The Sotrytis-conidia seem, however, capable of directly infecting a host-plant, at least I have always succeeded in infecting Conifers successfully with the conidial form Botrytis Douglasii. ‘See also Botrytis amongst the ‘* Fungi imperfecti.” *Brefeld, Heft 1v., p. 129, and x., p. 315; Tubeuf, Beitriige x. Kenntniss d. Baumkrankheiten, 188s. *Zopf. (Die Pilze, p. 742) states that Peziza-fruits may be reared from these sclerotia after they have rested a year. *Muller-Thurgau, ‘Die Edelfiiule d. Trauben.” Landwirth. Jahrbuch, 1888 (Ref. in Botan. Centralblatt, xxxv., 1888, p. 94). 268 ASCOMYCETES. Epidemics of great magnitude have been ascribed to attacks by the Botrytis-forms of this Sclerotinia. Thus on lilies in England,’ on yellow gentian,? on male flowers of Conifers, and on the twigs of Conifers and other plants. This is especially the case in houses under glass, where the fungus, favoured by the moist atmosphere, lives as a saprophyte on dead plant-remains, and multiplies till it becomes strong enough to act as a parasite. It is, however, quite possible that conidial forms of other sclerotia (eg. Sel. sclerotiorum) may be confounded with this species. Fic. 141.—Botrytis cinerea (Sel. Fuckeliana). Branch of Prunus triloba with two diseased shoots, withered and dead. (v. Tubeuf phot.) The presence of otrytis and allied forms on the vine is the cause of a disease of great economic importance, because severe loss may be incurred through rotting of the grapes and the injurious after-effects on the “most.” A decay of the potato-plant is said to be caused by sclerotia formed inside the stems, and also by a Botrytis. Smith* has figured similar sclerotia, which he ascribes to Peziza postuma Berk. and Wil. 1H. M. Ward, Annals of Botany, 1888. * Kissling, Hedwiyia, 1889. 3 Ritzema-Bos., Zeitsch. f. Pflanzenkrankheiten, 1894; O. Kirchner, Wurtemburg. Wochenblatt f. Landwirth., 1893. 4 Worthington G. Smith, Diseases of Field and Garden Crops. London, 1884. SCLEROTINIA WITH BOTRYTIS-CONIDIA. 269 Sclerotia, along with Sotrytis-conidia, have been found fre- quently on diseased geraniuins. During the summer of 1894 a withering of twigs of Prunus triloba occurred in several gardens at Munich (Fig. 141). A mycelium was found in the bark, leaf-petioles, and young ovaries, While Sofrytis-conidia were developed on the dead parts. With these I successfully infected young needles and twigs of spruce. Sclerotia were also formed on plum- gelatine in fourteen days. The parasite in this case had killed old twigs of Prunus, and also infected twigs of Conifers. Botrytis Douglasii is a parasite which I studied some time ago on account of its presence along with a disease on the Douglas fir (Psew- dotsuga Douglasii).' I have since had reason to believe that it is allied to some form of sclerotium = like that just considered, and my = view is supported by Behrens.” The disease as seen in various parts of Germany is characterized by withering, curling-up, and death of young shoots towards the sum- mits of young seedlings, and on the lower twigs of older trees up to about five feet above the ground. In autumn, black sclerotia about the size of pin heads, break through fi, a Pye Si apt ae the epidermis under the old bud- Offi, Guts rere Moot are dond. scales, at the base of dead shoots, and on the needles. In addition to these, smaller masses of tangled hyphae are also formed. When sclerotia are placed in a moist chamber, tufts of erect conidiophores arise, and branch, forming numerous whorls of conidiophores, from which 'y, Tubeuf, Beitriige z. Kenntniss d. Bawmkrankheiten, Berlin, 1888. “Behrens, Zeitsch. f. Pflanzenkrankheiten, 1895. 270 ASCOMYCETES. : P oval hyaline conidia are abjointed. These germinate at once in water, and infect young developing shoots or needles of Douglas fir, silver fir, spruce, and larch. Death of these ensues In a few days, and finally the whole plant is killed. On the dead needles a copious development of Botrytis takes place, and the conidia being easily detached, spread _ the disease in damp localities. The mycelium and _ conidiophores are very sensitive to drought. The sclerotia serve to carry the fungus over winter, and may be found in autumn and winter. I have found Juniperis communis with its young shoots dead, and sclerotia similar to the above on the needles. Whether Sclerotinia Kerneri Wettst. found on needles of Abies pectinata is parasitic or not, I do not know. Scl. galanthi Ludw.' Ludwig observed this disease on snowdrops. In place of the flower a shapeless mass was pro- duced, completely covered with conidiophores of Botrytis. The sclerotia develop inside the tuber. Scl. pseudotuberosa (Rehm). (S¢/. Batschiana Zopt or Ciboria pseudotuberosa Rehm) (Britain). The cotyledons of acorns are sometimes found replaced by a firm sclerotium, from which a peziza-fruit (Ciboria) is produced. Nothing is known in regard to mode of infection or the parasitism of this species. EU-PEZIZEAE. The apothecia, at first closed, open out to form saucer- shaped or cup-like discs, with a margin. The discs have usually a thick hypothecium; they are fleshy or waxy in texture, and are often brightly coloured. Dasyscypha. The waxy or membranous ascocarps are sessile or shortly stalked, and beset on the outer surface and margin with hairs of various colours. The asci dehisce by a round apical opening. The spores are ellipsoidal or spindle-shaped, unicellular, and hyaline. The paraphyses are thread-like. Most of the forms are saprophytic on dead plants; the following species alone is known to be parasitic. 1Ludwig, Lehrbuch d. niederen Kryptogamen. DASYSCYPHA. ak Dasyscypha (Peziza) Willkommii, Hartig.! The Larch Canker (Britain and U.S. America). Everywhere in the mountains, the home of the larch, one finds, on young branches and old stems, depressed canker-spots, on which the sporocarps of Dasyscyphu Willkommii are developed. Young twigs, when attacked, are already conspicuous in July and August by their pale and withered needles, and on them small canker-spots will be found; these rapidly enlarge so that on older stems they may reach very great dimensions. Harti easily sueceeded in producing canker-spots on healthy trees by artificial infection. If canker-spots are examined soon after the death of the bark, the stromata will be found as yellowish-white pustules. Conidia are produced either on the free surface or in the internal cavities of a stroma; they are tiny unicellular hyaline bodies, produced from little conidiophores. Hartig never succeeded in getting these spores to germinate. If the atmosphere be moist enough the apothecia make their appearance later on the same places; they are externally yellow, and internally orange- Fie. 143. — Dasyseyphe coloured. The apothecial disc carries long Ric pula eng faeed thread-like paraphyses and cylindrical asci Ei with rounded apices (Fig. 143). The asco- spores are oval, unicellular, and hyaline. They germinate and give off one or two germ-tubes which are unable to penetrate the periderm of a host-plant, and only find entrance through wounded places. Wounds are very common on larch as the result of hail, or injury to twigs by snow or ice, or destruction of needles by insects. For example, the Larch-moth (Coleophora laricella) is well known to cause less damage on the mountains than in the lower regions, and in the same degree Dasyseypha is least injurious to mountain forests. The mycelium is septate and much branched; it spreads chiefly through the soft bast, especially in the sieve-tubes and 'R. Hartig, Untersuch. aus d. forsthbotan. Institut Miinchen, 1., 1880. M. Will- komm, Mikros. Feinde des Waldes, 11., 1868. Die ASCOMYCETES. intercellular spaces, but it may also penetrate the wood as far as the pith. The fungus only spreads during autumn and winter, never during summer, the vegetative period of the larch. The attacked tissues of the bark turn brown and shrivel up, causing the depressed canker-spots. Healthy parts continue their growth normally, and are frequently cut off from diseased areas by formation of layers of secondary cork; this isolation is, however, rarely effective, since fresh invasions of mycelium from the wood into the bast take place annually, and thereby the canker-spots keep enlarging for an indefinite time. The fungus develops reproductive organs only in damp marshy situations. On this account spore-formation is less frequent on mountainous slopes than in moist valleys and ravines. The larch, on its first introduction into the low-lying parts of Germany, Denmark, and England, was much cultivated as a pure forest in close damp localities, and with great success ; but now this parasite has followed its host from the mountains and causes ever increasing damage. As preventive measures may be recommended: larches in low-lying districts should be grown in open, airy situations, and never massed together nor placed in the neighbourhood of diseased larches. Lachnella. The reproductive organs are similar to Dasyscypha, but the apothecia are firmer and generally have no stalk; the spores as a rule become two-celled at maturity. Lachnella pini Brunch.’ occurs in Norway on twigs of Pinus sylvestris, aS a parasite which quickly kills young plants and twigs. It is rare on old plants. The apothecia resemble those of D. Wilkommu, but are larger, externally brown, and covered with brown hairs and scales. The dise is reddish-yellow with a whitish margin. The asci measure about 100u by 9, and contain colourless unicellular spores about 20 long. Rhizina.” This genus contains the single species Rhizina undulata 'Brunchorst, Nogle norske skovsygdomme, Bergens Mus., 1892. *Rhizina has a position somewhere between the Pezizeae and the Helvelleae. Saccardo places the genus under Pezizeae, while Schroeter makes for it the special group of Rhizinacei, included under his Helvellinei. RHIZINA. 273 Fr. (2h. inflata, Schaetf.). Root fungus, or Ring-disease.’ This fungus is found as a saprophyte on the earth, especially where forest fires have occurred; also as a parasite on indigenous and exotic conifers. As such it has been observed in nurseries in various parts of Germany, and in woods of Pinus Pinaster in France. The fungus itself is known in Britain, though not as a parasite. FiG. 144.—Sporophores of Rhizina undulata. Fic. 145.—Section of hymenium. a, Para- «, Upper surface; 6, lower surface; c, small physes; }, secreting-tubes; ¢, asci, each with sporophores. (After Hartig.) eight spores. (After Hartig.) The disease extends from a centre and attacks one plant after another, causing them to lose their needles and die. The sporophores are large (j to 2 inches), chestnut-brown, flattened or undulating structures, which sit directly on the mycelium, without a stalk. On the upper surface is the ascogenous layer which, when moist, is peculiarly sticky and rR. Hartig, Forstl-naturwissen. Zeitschrift., 1892, p. 591; Prillieux, Compt. rend, dela Soc, des Agric., 1880. Ss 274 ASCOMYCETES. glutinous; it consists of small eight-spored asci over which project septate paraphyses, and also non-septate paraphyse-like structures which discharge a brown secretion. The ascospores are unicellular, hyaline, and canoe-shaped; on germination they give off a germ-tube which immediately develops into a_ septate mycelium. The mycelium is found in the intercellular spaces of the rind-parenchyma, but in the bast it grows both inside the cells and between them, so that the sieve-tubes are often completely Fic. 147.—Ascospores of Rhizina. «a, AS Fic. 146.—Rost-system of a Silver Fir taken from the ascus; b, 24 hours after overgrown and killed by the mycelium of sowing; c, 48 hours after sowing; d, the Rhizine undulata. (After Hartig.) spore of c enlarged. (After Hartig.) filled up. Masses of fungoid pseudoparenchyma are frequently formed between the dead and diseased tissues. Strands of the nature of Lhizoctonia emerge from the diseased roots, many of them carrying thread-like processes, at the extremity of which an oil-drop is secreted and escapes on rupture of the apex. According to Hartig, very tiny conidia are abjointed from the mycelium. De la Boulage! and Prillieux have both come to the conclusion that “la maladie du rond” of Pinus sylvestris and P. maritime is the same disease as the “ring-disease ” caused by Rhizina. ' Bull. de la soc. des Agric. de France, 1880. ” THE HELVELLACEAE, 2 Appendix. The Helvellaceae. This family is well known, some as poisonous, others as edible fungi (morel, etc.), and a few are suspected of being para- sites. The ascogenous layer occupies the upper surface of the sporophores, which grow on the earth and assume many various forms. As a rule they are erect and fleshy, and more or less lobed, wrinkled, or folded. USTILAGIN EAE. The Ustilagineae or Smut-fungi are distinguished by their dark-coloured or black chlamydospores, which, on germination, produce some form of promycelium capable of giving rise to an indefinite number of conidia or sporidia.’’ The chlamydospores themselves are produced in large numbers from a mycelium, and serve as resting-spores to carry the fungus through the winter, being often, in fact, the only part which persists. An endogenous formation of spores in sporangia as in the lower fungi, or in asci as in the Ascomycetes, does not occur in the Ustilagineae, Uredineae, or Basidiomycetes. The resting-spores of the Ustilagineae contain only one nucleus, the result of copulation of two nuclei; their formation thus marks the end of one generation, and their germination the beginning of a new. In the case of the Uredineae, Basidio- mycetes, and Ascomycetes, the beginning of the new generation is indicated by the germination of the teleutospore, the formation of basidiospores on the basidium, and the germination of the ascospore respectively. All the Ustilagineae are parasitic on higher plants, the mycelium growing intercellularly and nourished by means of haustoria sunk into the host-cells. The mycelium itself causes neither disease nor deformation of plants, and it is only when ' Brefeld regards the promycelium of the Ustilagineae not, like De Bary, as a mycelial structure, but as a conidiophore or basidial structure. In accordance with this view he has founded his intermediate group, the Hemibasidii corre sponding to the Ustilagineae. Brefeld then subdivides this group into (a) Ustila- neae (Ustilago, Sphacelotheca, Schizonella, Tolyposporium), which as a rule lave a septate promycelium ; and (4) Tilletieae (7'i//etia, Entyloma, Melanotaenium, Schroeteria, Thecaphora, Sorosporium), with non-septate promycelia, (Sehimme/ pile, Heft v., 1883, and Heft x1., 1895.) 276 USTILAGINEAE, the resting-spores are developed that deformation occurs. These spores arise by intercalary growth in the mycelium, which is generally completely used up in their formation; they are produced in large numbers, and scattered after decay of the tissues enclosing them. As a result of the germination of the resting-spores, there is produced either a mycelium capable of immediate infection, or a promycelium from which conidia’ are abjointed. In the latter case, conidia are generally formed in succession, and continue to be given off from the promycelium for a considerable time. They either give out a germ-tube capable of infecting a new host, or give rise to further conidia. The latter process is most frequently observed in artificial nutritive solutions, where the conidia continue to sprout in a yeast-like manner till nourishment is exhausted, when they germinate and form mycelial filaments. In the host-plant, chlamydospores alone are developed, conidia exceptionally (Zuburcinia and Enty- loma). The Ustilagineae are very dangerous and injurious enemies of cultivated plants, especially to the various cereal crops. The species are fairly easy to identify, because each is, as a rule, confined to one or a few species of host. The smut-fungi are best combated by sterilizing the seed of suspected cereals in a copper sulphate solution or in hot water shortly before sowing out; (see General Part, chap. vi.) In this way any adherent smut-spores are killed, and where this preventive measure is regularly carried out, disease is less common and its effects considerably minimized. The Ustilagineae include the following genera: U-stilago, Sphacelotheca, Schizonella, Tolyposporium, Tulletia, Entyloma, Melanotaenium, Uvocystis, Tuburcinia, Daossansia, Schroeteria, Thecaphora, Sorosporium, Graphiola, Schinzia, Tubercularia. Ustilago. The vegetative mycelium makes its way through the tissues of the host-plant without causing any deformation. The spores are developed in certain parts of the host, and form a much- branched, compact, sporogenous mycelium, with membranes 1*Conidia’=the sporidia of De Bary. USTILAGO. 2 which at first swell up in a gelatinous manner. Spores are formed inside the ultimate ramifications of the mycelium, and as they reach maturity, the membrane loses its gelatinous character, the cells break up, and the spores are set free; they are dispersed as a dry dusty powder after rupture of the tissues Fic. 148.—Ustilago maydisa. The head has been « xposed to view by dissecting away the enclosing leaves; it is beset towards the apex by smut-boils (v. Tubeuf phot.) of the host enclosing them. The spores germinate, giving rise to a promycelium (basidium), which becomes divided up by means of cross-septa into several cells, from each of which conidia are laterally abjointed. These conidia sprout yeast- like, and give off new conidia, or they produce a mycelium ; 278 USTILAGINEAE. the former is the case when nutrition is abundant, as when under artificial’ cultivation, the latter under less favourable nutrition; in very unsuitable conditions, the constituent cells Fic. 149.—Ustilago maydis. Diseased Maize-heads after removal of enclosing leaves. The heads are beset with smut-boils of all sizes, some ruptured, others still unbroken. (v. Tubeuf phot.) of the promycelium may each develop directly imto hyphae capable of infecting a new host. USTILAGO. 279 Ustilago maydis (D.C.)! (Britain and U.S. America). This smut of Zea Mais produces large and conspicuous deformations on leaves, leaf-sheaths, stems, roots, and all parts of the male and female flowers. These are whitish, gall-like swellings and _ blisters, containing a mass of gelatinous mycelium, from which spores are produced. The swellings may attain to the size of a fist, or even larger. The spores appear at first as dark olive-green masses seen through the lighter- green outer tissues of the host- plant. When mature the spore masses cause rupture of the enclosing host-tissues, and escape as a dusty powder. The spores are dark-brown in colour, irregu- larly spherical in shape, covered with delicate spines, and measure 9-124 in diameter. They re- main capable of germination for many years. On being sown from the host- plant directly into water, very few spores germinate at once, yet if sown in the following spring they readily do so. In a nutritive solution (¢.g. plum-juice velatine) an abundant germina- tion may be obtained at any : Fic. 150.—Ustilago maydis in head of time. and Wakker* have investigated the anatomical changes produced by this fungus. The latter investigator found that the xylem-elements with unlignified walls remain incompletely developed, and have a_ peculiarly twisted course; that normal sieve-tubes are absent; that the cells of parenchyma undergo secondary division, and give rise to a new tissue provided with little fibrovascular bundles, and rich in starch-contents, in other words, a nutritive tissue to be used up in the spore-formation of the smut. The disease may be found wherever maize is cultivated, and often causes a very serious diminution in the harvest. It may be combated by early removal and destruction of the smut-galls. As a preventive measure, the treatment of seed-corn with copper sulphate solution* is recommended. The avoidance of fresh manure is also advisable, since conidia capable of germi- nation may be lodged in it. The following are the results of an experiment carried out at my instigation by Professor Wollny in his experimental plots at Munich. Three plots were selected distant from each other about 70 metres. On 2nd May, 1893, these were marked out in rows 40 centimetres apart, in which maize was sown at intervals of 50 em. The grain was previously mixed with smut-spores obtained from the Tyrol in autumn, 1892. Plot No. 1 was left without manure, No. 2 was treated with old, No. 3 with fresh cow-manure. Maize had never been grown in the vicinity, so that no infection could result from external sources. The results were: Smutted. Number of Plants. Absol. Per cent. Plot No. 1, unmanured, : - 148 0 0 x 2, old cow-manure, - - 124 2 16 - 3, new cow-manure,- - 132 1] 76 Ustilago Schweinitzii Tul. from Carolina U.S.A. is probably identical with Ust. maydis. Ust. Fischeri Pass.° This smut, observed in upper Italy, ' Knowles, E. J., Amer. Journal of Mycology, Vol. 1v., 1889. *Cugini, ‘‘ Il carbone del grano turco,’’ Boll. dell. stat. Agrar. di Modena, 189). *Wakker, Pringsheim’s Jahrbuch, Bd. 24, 1892. ‘See ‘‘General Part,” chap. vi., and also ‘ Vergleichende Untersuchungen ib. Flugbrandarten.” P. Herzberg in Zop/f’s Beitriigen, 1895. 5 Passerini, Just’s botan. Jahrbuch, 1889, p. 123. 282 USTILAGINEAE. attacks the axis of the maize-heads. Its spores are spherical with slightly granular coats, and measure only 4-6 in diameter. It causes damage through shrivelling up of the grain. Ust. Reiliana Kiihn. This smut frequents Sorghum halepense and S. vulgare (Durra or Indian millet); also maize in various parts of Europe and America,’ as well as in Egypt and India. It is called “Hamari” in the Arabic language. Kiihn? thus deseribes it: ‘“ This species causes the ears of Durra to become large smut-galls of roundish or ovoid shape, with a height of 60-95 mm. and a diameter of 40-60 mm. At first the smut is enclosed in a whitish skin, which is ruptured into shreds to allow the escape of the black spore-powder. After the smut-spores are shed, there remains a stiff skeleton consisting of the fibrovascular bundles of the aborted ear.” The spores are distinguished from those of Ust. maydis by their greater size (9-15), and their almost smooth membrane with very small spines. According to Brefeld, the spores are capable of germination in nutritive solutions after eight years. In the fresh condition they germinate in water to a limited extent, producing multicellular promycelia which give off conidia. In nutritive solutions they germinate and produce thick promy- cela with three or four cells, from which multitudes of conidia (5-124 long and 3-5 broad) are abjointed. The conidia fall off and sprout till the nutritive substratum is exhausted, when they give rise to thread-like conidia which do not coalesce. If kept dry the conidia easily retain their vitality for months. Kiihn distinguishes further Sorosporium Ehrenbergii Kiihn on Sorghuin CETRUUWMN. Ust. cruenta Kiihn.* Another parasite on the ears’ of Sorghum. It is described by Kiihn as follows: “On the spikelets little reddish-brown protuberances of roundish or oblong shape are formed and enclose moderately-sized masses of dark-red smut-powder. If the pustules are very numerous they coalesce with each other, and the branches of the ear become more or less shortened, thickened, and twisted. Where ' Norton, ‘‘ Ustilago Reiliana,” Botanical Gazette, 1895, p. 462. *Kiihn, ‘‘ Die Brandformen der Sorghum-arten,” Mittheil d. Ver. f. Erdkunde zu Halle, 1877. * Kiihn (Joc. cit.) and Hamburger Garten-Zeitung, Bd. 28. Brefeld, Heft v., p. 91. USTILAGO. 283 the pustules are fewer in number the parts of the ear retain their normal position, but all the floral organs contained in the glumes are wholly or partially converted into irregular ereyish smut-masses. Isolated pustules may occur under the inflorescence, on the next internode of the haulm.” Fic. 153.—Ustilago cruenta. Smut of Durra or Sorghum. The head has been divided up and the isolated branches photographed. The ovaries are transformed to long crooked sacs, and pustule-like outgrowths are also present on stalklets and stalks. (v. Tubeuf phot. from material supplied by Prof. Dr. Jul. Kithn.) The spores are yellow to brown in colour, smooth-walled, and of very variable shape, 5-12 long and 5-94 broad. As a rule, germination in water results in the formation of a germ- tube composed of four or five cells, which elongate to long mycelial threads or, exceptionally, produce a single conidium. As a result of germination in nutritive solutions, a_ lively 284 USTILAGINEAE. formation of conidia ensues; the conidia multiply in a yeast- like manner, and only grow out as hyphae on exhaustion of nutritive material. Infection takes place on seedling-plants. Kiihn cultivated this species on Sorghum saccharatum and S. vulgare, and suggests that a common disease of Durra in South Africa may be caused by this parasite. Ust. sorghi (Link.) (Ust. Tulasnei Kiihn) (U.S. America). This is another widely distributed parasite of Sorghum vulgare and S. saccharatum. Its external appearance is described by Kiihn somewhat as follows: “ Diseased plants attain to almost their normal size, and the flower-head is developed as far as the glumes. The ovary, however, is completely metamorphosed into a sac filled with spores, its outer wall forming a delicate Fic. 154.—Ustilago cruenta. Spikelet enlarged from a head of Sorghum. The ‘ ovaries are transformed into long flask- Fig. 155.—Ustilago eruenta. Germin- shaped sacs, from slits of which the ating and sprouting conidia from a spores are emerging as a black powder. cultivation in plum-gelatine. (v. Tubeuf + natural size. (v. Tubeuf del.) del.) whitish coat, which is easily torn, and, when the spores have escaped, a columella will be found to occupy the centre of the smut-mass. The stamens may also become filled with spores, and be externally more or less irrecognizable. As a rule, all the flowers of a head are smutty; if any escape, they remain more or less rudimentary.” The spores, according to Brefeld, germinate only in nutritive solutions. They produce a four-celled promycelium, on which few conidia are formed. Ust. sacchari Rabh. Dust-brand of cane sugar. This fungus injures the stems and heads of Saccharum officinale, S. cylindricum, and S. EHrianthi in Italy, Africa, and Java. Ust. sacchari-ciliaris Bref. occurs on Saccharum ciliare near Calcutta. Ust. avenae (Pers.). The smut or brand of the oat occurs USTILAGO. 285 very frequently on Avena sativa, also on Avena orientalis, A, fatua, and A. strigosa in Europe and North America. So common is it that one seldom sees a field of oats free from the black smutted ears (Fig. 156). All parts of the flower are attacked, the ovary, stamens, glumes, and even the awns. The grains become filled with the black spore-powder, which shows through the transparent eS \ Fic. 156.—Ustilago avenae. The Oat-smut on Avena sativa. (v. Tubeuf phot.) membrane of the ovary wall. The diseased ears emerge from their enclosing leaf-sheaths, and become exposed to wind and rain, under the effects of which the delicate membrane soon becomes ruptured and the spores are blown or washed away, till only the axes of the spikelet are left with a few ragged remains of the flower. As a rule every shoot of a plant and all the grains of an ear are attacked; if single grains do escape, they remain poorly developed. The spores (5-8) have a smooth or slightly granular coat, and 286 USTILAGINEAE. retain their capacity for germination for years. In water they germinate immediately, and produce a single (rarely two) promy- celium consisting of four or five cells, from the ends or partition- walls of which oblong conidia continue to be abjointed for about two days. The cells of promycelia may become connected with one another by lateral branchlets. Delicate germ-tubes are given off by the promycelial cells, by the conidia, or by secondary conidia. In nutritive solutions, on the other hand, the spores germinate much more vigorously, the promycelium is stronger, the conidia are continuously abjointed from little sterigmata, and go on sprouting in a yeast-like manner till, on exhaustion of the nutriment, they germinate to form vigorous mycelial filaments. The fusion of the cells of promycelia never takes place in nutritive solutions. The infection of oat-plants takes place on the soil by means of the germ-tubes produced from the conidia, promycelia, or spores.’ These infect the first leaf-sheath—that one which on germination emerges from the ruptured seed-coats as a whitish or yellowish- green shining shoot, and continues to grow as a sharp-pointed cylinder till, pierced by the first green leaf, it dries up. In 36 to 48 hours after infection, mycelial threads were found to have pierced the epidermal walls, and to have branched freely in the tissues. The mycelium grows from the leaf-sheath into the first green leaf, passes straight through it into the second, and so on till it reaches the haulm or stem.” The young mycelium grows steadily onwards, and the plasma of older hyphae passes over into it. In this way the fungus keeps pace with the host-plant, exhibiting externally no symptom of its presence till the flowers are reached, where the chlamydo- spores are formed. Sterilization of seed-corn by Jensen’s hot-water method is strongly recommended.* In America, steeps containing potassium sulphide, copper sulphate, or lime are also used. As preventive against infection, late sowing is advisable. This is founded on Brefeld’s investigations, in which he found that oat-smut germin- 1Wolf, Der Brand des Getreides, 1874. 2 According to Kiihn, and in Brefeld’s infections (Heft x1., 1895), the majority of the germinating conidia are said to penetrate into the young shoot-axis. 3 Treatment of Smuts of Oats and Wheat,” U.S. Department of Agriculture, Farmers’ Bulletin No. 5, 1892; ‘‘Grain-smuts and their prevention,” Yearbook of U.S. Dept. of Agriculture, 1894. USTILAGO. 287 ated best at 10°C., and not so well above 15°C. This conclusion is supported by experiments of Kellermann and Swingle. Neither these investigators nor Jensen, however, agree Fic. 157.—Ustilago perennans on Arrhenatherum elatius (Oat grass). The grains are transformed into black smut-masses ; the appearance of the infected spikelets is quite distinct from that of the healthy one to the right. (v. Tubeuf phot.) with Brefeld’s view, that the fungus is introduced into fields with fresh farmyard manure. Kellermann and Swingle have found a smut on oats in America which they distinguish as Ust. avenae var. levis. Ust. Kolleri Wille. This is another species of oat-smut recently dis tinguished ; it has smooth spores, and is said to cause even greater damage than Ust. avenae. 288 USTILAGINEAE. Ust. perennans Rostr.' This smut or dust-brand occurs frequently in the flowers of Avrhenatherwm elatius (Fig. 157). The mycelium perennates in the rhizome. An Ustilago nearly allied to the preceding one occurs also on Festuca pratensis, Lolium perenne, and other erasses. The Smut of Barley. There are really two species of Ustilago found on _ barley, Fi0. W8.— Usllage perennans, Speresitore Ose. rordet, and Ust. nuda. Ust. hordei (Pers.) (Ust. Jensent Rostr.) (Britain and U.S. America). This has black spherical spores (6°5 to 75m in diameter), which germinate and give off conidia from a promycelium. The spikelets gene- rally remain enclosed in their coverings. Treatment of seed- corn with a half per cent. copper steep is a certain remedy. Ust. nuda (Jens.) (U.S. America). In ears diseased by this smut the epidermis of the glumes is early lost, so that the spore-powder lies freely exposed when the ears emerge from the leaf-sheath. The spores on germination give off a four- celled promycelium, which however produces no conidia, but develops directly to a septate mycelium. The spores are smooth-coated and oval (5-7m# long and 5-6°5m broad); they are matured and set free at the flowering season of the barley, and probably infect seedlings in spring. The spores of this smut are very resistant against treatment with copper steeps, and it is recommended to soften the barley for several hours in cold water before applying Jenson’s method. Ust. tritici (Pers.) (Britain and U.S. America), Wheat-brand. The spores are developed in the ovary of the wheat, and are black with a tinge of olive-green. On germination they im- mediately form a non-septate mycelium (Fig. 160). Henning? has described spore-cushions on the leaves and leaf-sheaths of Triticum vulgare in Upper Egypt. Ust. bullata Berk. on 7Vriticum orientale in Turkestan. 'Rostrup, Ustilagineae Daniae, 1890. * Henning, Zeztschrift f. Pflanzenkrankheiten, 1894. USTILAGO. 289 Ust. secalis Rabenh. Rye-brand. This occurs but rarely, and destroys only the grain. Ust. panici-miliacei (Pers.) (Ust. destrvens Duby). Smut of Millet. This smut occurs on the flowers of Panicum milia- PIG, 159.—Ustilago hordei. Barley-smut on Hordeum distichum. (v. Tubeuf phot.) coum, LP. chartaginiense, and P. Crus-galli in Italy, France, Germany, and North America. Sometimes it is very abundant and causes great damage. The mycelium makes its way into young plants and grows upwards with them, penetrating every 7 290 USTILAGINEAE. shoot. Spores are developed only in the inflorescence, which in consequence fails to reach its full development as a panicle, and remains more or less spike-like and enclosed in a leaf- sheath. The parts of the inflorescence become completely filled Fic. 160.—Ustilago tritici. Wheat-smut. The central ear is normal and healthy, the others are smutted and most of the spores are already shed. (v. Tubeuf phot.) with a sporogenous mycelium from which arise the spore-masses ; these are at first enclosed in whitish coverings consisting of tissues of the host-plant, but when mature they escape as a black dust or powder. USTILAGO. 291 The spores are smooth-coated and spherical or elliptical, 9-12 u long, and 8-10 broad. According to Brefeld, they germinate in two or three days in water, and produce promycelia with four or five cells; the cells may either bud out directly and become hyphae, or do so after previous fusion. Spores placed in nutritive solutions germinate in about three days, and produce several strong septate promycelia with spindle- shaped conidia. The conidia as a rule germinate directly into branching hyphae; fusion of conidia is not known, and secondary conidia are only rarely formed. The hyphae become septate in their older parts, and produce conidia in two ways, firstly, from hyphae in the solution itself; secondly, from aerial hyphal branches which rise out of the solution and give off conidia in a manner similar to mould-fungi. Brefeld states that infection takes place by means of the germinating conidia. Only resting-spores are produced on the plant itself, and these retain their capacity for germination for years. Ust. Rabenhorstiana Kiihn’ (U.S. America). This is found on Panicum miliaceum, P. glabrum, P. lineare, and P. sanguinale. It destroys flowers, ears, and upper part of haulms. The spores are brown and spiny; they germinate, but do not produce conidia. Ust. sphaerogena Burrill. An American species causing distortion of the spikelets of Paniewm Crus-galli. The malforma- tions resemble those produced on the same host by Zolyposporium bullatum, but differ in having a rough surface with short rigid hairs. The spores are free and germinate easily in water, producing promycelia which give off conidia. The conidia frequently sprout for a time in a yeast-like manner. The following are American species : Ust. diplospora Ell et Ev. On Panicum sanguinale. Ust. trichophora Lk. On Panicum colinum. Ust. setariae Rabh. On Panicum sanguinale ; probably identical with Ust. Rabenhorstiana. Ust. panici-leucophaei Bref. On Panicum leucophaeum in Rio de Janeiro. Ust. digitariae Kze occurs on the flowers of Panicum (Digitaria) sanguinale, P. glabrum, and P. repens. The spores are smooth-walled. 'Kithn, Hedwigia, 1876. 292 USTILAGINEAE. Ust. panici-frumentacei Bref.’ is found on Panicum frumen- taceum, «a cultivated Himalayan millet. | Only isolated grains in an ear are attacked, becoming enlarged to twice their normal size. Germination of spores takes place sparingly in water, but abundantly in nutritive solutions. Two-celled promy- celia are produced bearing numerous sprouting conidia. On exhaustion of nutrition, the conidia give off one or two filaments on the surface of the liquid, and from these other sprouting conidia arise. Ust. Crameri Korn. completely destroys the ovaries of Sefaria italica, S. viridis, and S. ambigua, leaving only the outer wall as an enclosure for the spore-powder. The spores are brown, smooth-walled, and 6-9 broad, 10-12 long. The promycelia consist of four or five cells, which in water as well as nutritive solutions grow out into long threads without producing conidia. Ust. neglecta Niessl fills with its black spore-powder the ovaries of Setaria glauca, S. verticillata, and S. viridis. The cells of the promycelium develop into a mycelium without pro- duction of conidia. Ust. Kolaczekii Kiihn. On Setaria geniculata in Berlin Botanic Garden. Ust. bromivora Fisch. (Britain and U.S. America). This appears in flowers of species of Bromus, so that the ovaries become filled with a dark-brown or black spore-powder, but the glumes or heads undergo no deformation. The spores are smooth, and on germination in water produce only a spindle- shaped one-celled (rarely two-celled) promycelium ; in nutritive solutions, Brefeld found they generally produced two-celled promycelia, bearme conidia from which are produced further promycela with conidia; yeast-like colonies are never formed. Ust. ischaemi Fuck. attacks Andropogon Ischaemum. The inflorescences remain almost completely enclosed in the upper- most leaf-sheath, and are destroyed except their axes. The spores are brown and smooth-walled. Brefeld states that in nutritive solutions they produce conidia which remain adherent to the promycelium and grow out into long hyphae without coalescing. Ust. andropogonis-tuberculati Bref. on Andropogon tuberculatum from Simla. Ust. andropogonis-annulati Bref. on Andropogon annulatum from Culcutta. ! Brefeld, Schimmelpilze, Heft x11., 1895. USTILAGO. 293 Ust. grandis Fries. Reed-smut. (Britain.) This frequents the haulms ot Phragmites communis (also Typha latifolia and T. minor); the internodes of the host in consequence swell out and appear as if the stem carried one or more bulrush-heads. The mycelium permeates the whole host-tissue and _ produces spores, Which escape as a black dust on rupture of the epidermis. According to Kiihn, the spores are capable of immediate ger- mination and retain their vitality for a whole year. A four-celled promycelium is produced and becomes detached from the spore; then follows an abjunction of oblong conidia from the septa of the promycelum. In nutritive solutions, Brefeld found that germination took place in the same way, but more rapidly and vigorously. Numerous conidia are produced, but these only rarely give off secondary conidia, and then only a single one; more commonly they produce promycelia, as the spores did, and conidia again arise from these; yeast-like sprouting does not occur. The resting-spores may continue to give off promycelia in succession for some time. On _ ex- haustion of nutrition the cells of the promycelium, as_ well as the conidia, develop into mycelial threads, to which alone Brefeld ascribes the capacity for infection. Ust. longissima (Sow.) (Britain and U.S. America). This forms elongated brown spore-patches on the leaves of various species of Glyceria. Brefeld states that the smooth spherical spores germinate in water, and give off a short unicellular promycelium which undergoes no further development. In nutritive solutions the spores germinate in like manner, but the promycelium becomes thread-like and septate, and gives off conidia laterally; new promycelia continue to be given off from a cell which remains behind inside the spore, and the conidia ultimately develop into hyphae. Ust. hypodytes (Schlecht). This species forms dark smutty coatings on haulms and leaf-sheaths of Glyceria fluitans, Dip- lachnis fusca, Agropyrum repens, Calamagrostis epigea, Psamma arenaria, Stipa pennata and SN. capillaris, Bromus erectus, Triticwin repens and 7. vulgare, Elymus arenarius, Panicum repens, Phrag- mites communis, Arundinaria, etc. The spores are brown, smooth-walled, and irregularly spherical or quadrangular; they germinate in water or nutritive solutions, producing mycelia direct, without previous formation of conidia. 294 USTILAGINEAE. Ust. grammica B. et B. is reported on haulms of Azra and Glyceria in England. Ust. echinata Schroet. produces smut-strips on leaves of Phaluris arundinacea, (U.S. Amer.) Ust. cynodontis Henn. On Cynodon Dactylon from Simla. Ust. arundinellae Bref. On Arundinellu near Calcutta. Ust. aristidae-cyanthae Bref. On Aristida cyantha from Himalaya. Ust. coicis Bref. On Coiv lacryma from Simla. Ust. esculenta Henn.! causes deformation of plants of Zizania latifolia in Tonquin and Japan. The deformed parts are eaten, while the spores are used for dying of hair and eye-brows, as well as in the manufacture of a varnish. Ust. paspalus-dilatati Henn. On Paspalus dilatatus. Ust. olivacea D. C. frequents species of Carex. The olive- brown spore-masses hang loose and fleecy from the destroyed ovary. The spores, according to Brefeld, are produced from long hyphae which become thickened at intervals and broken up by cross-septa into portions corresponding to the future spores. The hyphae, however, are not completely given up to spore-formation, but parts remain and form fine filaments which give the fleecy appearance to the ruptured ovaries. Germina- tion in water results in the formation of a single conidium, a second being rarely formed. In nutritive solutions similar conidia are produced one after another successively, and sprout off conidia in a yeast-like manner without the formation of pro- mycelia. On failure of nutriment, hyphae are finally produced. Ust. Vuijkii Oudem. et Beyerk. The ovaries of ZLuzula campestris become filled with spores, some colourless, some light-brown. The spores germinate in water, giving four-celled promycelia with ovoid conidia, which do not, however, coalesce or develop further, even in nutritive solutions. Ust. capensis Rees. In fruit of Juncus. Ust. luzulae Sace. In fruit of Luzula. Ust. scabiosae (Sow.)? (Ust. flosculorwm Tul.). (Britain.) The anthers of Anautia and Scabiosa attacked by this fungus become filled with a flesh-coloured to violet spore-powder, and swell to little sacs. The flowers otherwise are but little altered. Brefeld found that spores from Knautia arvensis germinate easily and abundantly in water, and produce promycelia con- 'P. Hennings, Hedwigia, 1895; Miyabe, Tokio Botanical Magazine, 1895. 2 Fischer v. Waldheim, Bot. Zeitung, 1867. USTILAGO. 295 sisting of three or four cells with conidia, and sometimes secondary conidia. Coalescence of conidia may take place, and thereafter production of little mycelial threads. In nutritive solutions everything proceeds more luxuriantly, and conidia are produced in large numbers; they are easily detached and sprout Fic. 161.—Ustilago tragopogonis. Plants of Tragopogon in tlower and fruit 1, normal fruit; 2 and 3, normal flowers; 4, two normal flower-buds. The remaining specimens are attacked by the fungus, and, in consequence, remain in the bud condition, and filled with black spores which escape by the opening of the involucre. (v. Tubeuf phot.) yeast-like, till, on deficiency of nutrition, fusion and subsequent germination takes place. Ust. intermedia Schroet. (sf. Jlosculorum VD. C.) (Britain). The anthers of Scabiosa Columbaria become filled with the dark violet spores of this smut. The spores germinate in water, and, according to Brefeld, produce three-celled promycelia with few conidia; some of. these, as well as the cells of the promy- 296 USTILAGINEAE. celia, may develop to mycelia; coalescence of conidia is unknown. In nutritive solutions conidia are formed in large numbers, and multiply yeast-like till nutriment fails. Ust. succisae Magn.’ frequents the anthers of Scabiosa Suecisa, and forms pure white spores, easily distinguished from those of the two preceding species. The anthers appear to be thickly covered with glassy granules. The spores produce four-celled promycelia from which conidia are formed. (Britain.) Ust. tragopogonis (Pers.) (Britain). This fungus forms its spores in flowers of species of Zragopogon, and in many localities has a wide distribution. The development of the flower is retarded, so that it retains externally the appearance of a flower-bud en- closed in its bracts (Fig. 161). The dark-brown or violet spores escape Fae ae ee ears through intervals between the bracts; Development of spores: successive they are 13-1 Th long, LOST 5x braad, stages of development, in order of the letters. a, Sporogenous branch, with reticulate markings on their just appearing on the surface of ¢ young corolla of Zragopogon pra- coats. They easily produce in water tensis, and beginning to form a tuft es y of branchlets. bande, Formation of four or five-celled promycelia from spores from the mycelium. d, Spore- clump with several ripe spores, the which conidia are given off, often episporium of which is coloured = (eee cel Wisaed oO followed by coalescence. In nutritive De Bary.) solutions development is much more vigorous, secondary conidia may be produced, and coalescence always takes place. Ust. scorzonerae (Alb. et Schwein.) is at first sight very similar to Ust. tragopogonis. Its spores are found in flowers of Scorzonera humilis, Sc. purpurea, and cultivated species, e.g. Sc. hispanica; while its mycelium hibernates in the perennial root-stocks of these. The spores are produced rapidly and - in large numbers; they germinate easily in water, forming a four-celled promycelium, and thereafter conidia which do not pair. Ust. cardui Fisch. v. Waldh. (Britain). This is the cause of a stunting of the flower-heads of Carduus . acanthoides, C. nutans, and Silybum Marianum; while at the same time they beeome filled with a brownish-violet spore-powder. The spores ' Magnus, Hedwigia, 1875. USTILAGO. 297 are about 20m in diameter, and form in water promycelia with conidia. In nutritive solutions Brefeld found conidia produced in large numbers, and multiplying by yeast-budding. The promycelial cells grow out as septate branched twigs, from which conidia are abjointed, and after coalescing in pairs, produce germ-tubes. Ust. violacea (Pers.).. Carnation-smut (Britain and U.S. America). In Silene, Visearia, Saponaria, Dianthus, Stellaria, Malachium, Cerastium, and Lychnis, the pollen sacs of other- wise well-developed flowers become filled with dark-violet spores, which escape and discolour the other floral parts. Pistillate flowers of Zychnis attacked by this fungus develop stamens containing the smut-spores (p. 27). On germination in water, promycelia of three or four cells are formed, and become detached from the spores. Primary and even secondary conidia are produced, while coalescence of promycelial cells and conidia is common; but only a few of them produce germ- tubes. In nutritive solution, according to Brefeld, everything proceeds much more vigorously; from tiny conidiophores on the promycelia numerous conidia are produced in succession, and from these other conidia are budded off like yeast-cells till nutriment fails, when they grow out to form hyphae. The conidia are longer than those formed in the water-cultures, and coalesce in pairs to give rise to longer and_ stronger verm-tubes. Ust. holostei De Bary on Holostewm umbellatum. The host-ovaries become filled with spores which germinate to four-celled promycelia from which pairing sporidia are formed. Ust. Duriaeana Tul. In the ovary of Cerastium. Ust. major Schroet. On Si/ene Otites. The spores germinate only in nutritive solutions. (Britain.) Ust. seminum Juel. In the ovules of Arabis petruea in Scandinavia, The spores on germination produce simple hyphae. Ust. entorrhiza Schroet. In root-cells of Pisum sativum. Ust. pinguicolae Rostr. On Pingwicula vulgaris in Denmark. According to Brefeld, the spores germinate equally in water or nutritive solutions, forming three-celled promycelia, which separate from the spore and bud off conidia from each cell. 'Tulasne, Ann. d. sciences natur., Ser. ui., Vol. vit., 1847. Atkinson (American Carnation Society, 1893), describes this and other smuts frequenting American Carnations. (dit. ) 298 USTILAGINEAE. Ust. betonicae Beck.’ occurs in the anthers of Betonica Alopecurus. Its spores are larger than those of Ust. violacea, and have larger-meshed reticulations on the spore-coat. The spores germinate in water, and as a rule produce a three-celled promycelium from which conidia are abjointed. These at once, or after production of conidia, coalesce in pairs and give off germ- tubes. In nutritive solutions germination takes place much more vigorously, numerous conidia are formed and continue to bud off new conidia till the nutriment is exhausted, when coalescence of conidia and development of hyphae takes place. Ust. bistortarum D. C. frequents leaves of Polygonum and Rumex. (Britain and U.S. America.) Brefeld states that the spores are dark-red and germinate to four-celled promycelia, from which conidia are produced and readily coalesce, especially in presence of abundant nutriment. Ust. marginalis (Lk.) on Polygonum Bistorta. The spore-masses are dark-violet, and occur chiefly on the margins of the leaves. The spores germinate in water and produce a four-celled promy- celium with oval conidia, which do not sprout, oe either pair or grow out as hyphae. Ust. anomala Kunze. On leaves and in ovaries of Polygonum (U.S. America). Ust. utriculosa (Nees). In ovaries and anthers of Polygonum. The ereyish-violet spores, Brefeld says, germinate during the following summer, and give off four-celled promycelia with conidia which do not coalesce in pairs. (Britain and U.S. America.) Ust. Parlatorei Fisch. On twigs and leaves of Rumer maritimus and R. obtusifolius. Ust. Kuhneana Wolf. Inhabits all parts of Rumer Acetosa and R. Aceto- sella (Britain). Ust. Goeppertiana Schroet. On Rumex Acetosa, especially in leaves and leaf-petioles. The spores germinate in water or nutritive solution. The promycelium is unicellular and remains inside the spore, giving off a single conidium, which for a time buds off other conidia (Ust. olivacea alone behaves in this same way). Ust. Molleri Bref. On Polygonum hispidum. Ust. Koordersiana Bref. On Polygonum barbatum in Java. Ust. domestica Bref. On Rumex domesticus in Norway. Ust. vinosa (Berk.). On fruits of Oxyria (Britain and U.S. America). The spores germinate in water or nutritive solutions, and produce a four-celled promycelium from which conidia are given off, especially in nutritive solutions ; the conidia ultimately produce germ-tubes. ! Zoolog.-botan. Gesell., Vienna, 1880. USTILAGO. 299 Ust. Vaillantii Tul.’ appears in the anthers and ovaries of Gagea, Scilla, Muscari, ete. The perianth of diseased flowers remains, but is somewhat enlarged. The ovaries and anthers become filled with spores; the latter organs are, however, fully developed and may even contain pollen-grains mixed with spores. According to Brefeld, the spores germinate easily in water and in nutritive solution. A promycelium is formed which, after detach- ment from the spore, becomes three-celled and develops conidia. These sprout for some time, then produce three-celled promycelia. Ust. ornithogali (Schm. et Kze) forms leaf-swellings on Ornithogalum and Gagea. ° Ust. tulipae (Heufl.) produces swellings on the leaves of the tulip. Ust. plumbea Rostr. occurs on leaves of Arum maculatum in Denmark. Ust. ficuum Reich. In the fruits of Aiews Carica in Asia Minor. Ust. Trabutiana Sacc. In berries of Dracaena Draco in Algeria. Ust. Vrieseana Vuill.2 In the Botanic Garden at Amster- dam, the roots of several species of Eucalyptus exhibited woody tumours from which proceeded outgrowths resembling “ witches’ brooms.” These contained the mycelium of an Ustilago which produced spores in the cortical tissues. Ust. (7?) adoxae Bref. On Adoxa moschatellina in cells of the subterranean stem. The spores produced only simple filaments without conidia. Ust. Lagerheimii Bref. On Rumea from Quito. Ust. Schweinfurthiana Thiim. On /mperata cylindrica from Cairo. Ust. boutelouae-humilis Bref. On Bowteloua humilis from Quito. Ust. Ulei Henn. On Chloris. Ust. spinificis Ludw. On Spinifexr hirsuta from Adelaide, Australia. Ust. Treubii Solms.* This Javanese fungus and the galls produced by it deserve a somewhat lengthened notice on account of their general biological interest. It causes a hypertrophy on Polygonum chinense in Java, which further exemplifies the phenomena already noticed in connection with Caeoma deformans on Thujopsis (p. 30). The stems at attacked places show strong hypertrophy and great change in their anatomical structure. Solms designates the thickenings, in common with those caused by Caeoma 'Tulasne, Ann. d. science natur., Ser. 111., Vol. vut., 1847, with plates of Muscari, Worth G. Smith (Gardener's Chronicle, Xv., 1894, p. 463), gives a figure and note on occurrence of this smut in Britain. (Kdit.) “Vuillemin, Compt. rend., 1894. *Solms, Annal. du jardin botan. de Buitenzory, Vol. vi., 1886-87, p. 79. 300 USTILAGINEAE. deformans and Peridermium elatinum, as “vegetative canker- galls.” On those places are crowded fleshy brittle outgrowths, consisting of an irregular bent club-like stalk, longitudinally furrowed, and expanded at its upper extremity into a broadened head containing the Ustilago spores. Solms calls these out- growths “fruiting galls,’' and he describes them as follows: “if one of these protuberances be divided, the spore deposit will be found as a flattened violet layer, extending to the margins of the head and roofed in by a slight plate of tissue. This last becomes ruptured, shrivelled, and brown. The _ violet spores are thus set free, along with a loose woolly capillitium- tissue, which apparently facilitates distribution of the spores by rendering them difficult to moisten, a contingency very likely to happen in the heavy tropical rains of Java, and with the result that germination would occur before the spores had time to be transported to a new host. After shedding of the spores, the succulent stalk remains. The fruit-galls consist of a hypertrophied tissue developed from the cambium; they first emerge as roundish naked protuberances, covered externally by a smooth epidermis, and containing a meristem from which fibrovascular bundles are developed. The galls are composed of a homogenous parenchyma of large thin-walled cells, elongated in the direction of the long axis of the galls, and containing large cell-nuclei. The epidermis consists of little, polygonal, nucleated cells, and is pierced by a few stomata. The galls are internally permeated by a number of irregularly arranged fibrovascular bundles which show a slightly developed wood and bast region. As the anterior end of the fruit-gall elongates, the bundles keep pace by repeated forkings, and form a system of branches diverging at very acute angles and_ter- minating a short distance from the surface of the gall. The violet-brown sporogenous layer is situated just at the termination of the bundles, and is covered by a slight layer of parenchyma under the epidermis. The sporogenous layer appears as_ if composed of columns arranged beside one another in a palisade manner, and connected above and below with the enclosing tissues. At the margins of a section the columns easily separate, and will be seen to consist of a central strand of elongated cylindrical cells filled with a reddish gum-like mass. The cells 1 Fruchtgallen, USTILAGO. 301 belong to the tissue of the Polygonum and may form simple filaments, or several such filaments may become bound together by lateral connections. Each strand becomes surrounded by spores of the Ustilago which are set free on rupture of the fruit-gall, while the cell-strands laterally bound to each other are loosened from the surrounding tissue as the capillitium. _ “The spores germinate in water, producing short unicellular promycelia and fairly large conidia, which coalesce before they germinate. The mycelium is confined to a small part of the stem, twigs, or inflorescences of the host-plant. The hyper- trophied parts of the stem contain abnormal spongy wood, which easily decomposes and brings about the death of the galls, along with parts of the stem situated beyond them, or even the whole plant. The normal production of cambium is completely destroyed in the galls. The pith and primary rind, however, remain uninfluenced. The cambium produces, both outwards and inwards, such a mass of thin-walled parenchyma that the normal bast is forced asunder and disarranged. In this way rupture of the sclerenchyma-layer ensues, whereby the primary rind is destroyed, and the abnormal tissue formed by the cambium emerges to view. It is from such places that the excrescences described have their origin.” It will be seen we have here the partners of a symbiosis becoming so adapted to each other that the host-plant produces a special tissue for the distribution of the spores. This case goes further than most of those already mentioned in § 5; but the bushes produced by Cacoma deformans for the formation of its spores are again a distinct advance on the “ fruit-galls” of this Ustilago. Cintractia. Spore-masses developed inside a stroma and passing outwards so that the mature black spores lie freely exposed. Magnus! has recently separated Ust//ago caricis Pers. and U. subinclusa Korn., and placed them under this genus, because their spores are developed only in the epidermal cells of the host-ovary. Cintractia caricis (Vers.)' (Britain and U.S. America). The 1Cornu. Annal. d. sciences natur., Ser. v1., Vol. xv., 1883. Plate XV. Magnus, Botan. Verein d. Prov. Branden/urg, XXxvu. Brefeld, Sehimmelpils, Heft x1., 1895. 302 USTILAGINEAE. mycelium forms a stroma on the ovary-wall; there the spores originate and pass out to the periphery as they attain maturity. The spores adhere in black masses, and germinate in water in the following spring. A promycelium is pro- duced, and on emerging into the air becomes divided by means of a cross- septum towards its apex; from both cells so formed conidia are developed and grow out into germ-tubes without previous sprouting. This species occurs on many species of Carex, and the mycelium perennates in the rhizomes. The spores vary somewhat on _ the different hosts. C. subinclusa (Korn.) (U.S. America). The spores form coal-black masses in the ovaries of many species of Carex. They develop on a stroma from within out- wards, and are more easily detached than those of C. caricis; their coat-markings also take the form of thicker and shorter processes. On germination in water after a resting period, the spores produce two-celled promycelia, from the apical cell of which an ovoid conidium is abjointed, while from the lower cell a lateral conidiophore is produced. Numerous conidia are given off from both cells, and erow out without previous sprouting. Fic. 163.—Cintractia caricis. Two ovaries have been replaced by black spherical fungus-fruits ; an isolated normal triangular ovary is shown in longitudinal and cross section. (v. Tubeuf del.) C. (2) sorghi (Endothlaspis sorghi) Sor. The mycelium envelopes the grain of Sorghum cernuum, and fills it with black spore-masses. It has only been observed in Asia. Other species of Cintractia occur outside of Europe, but are of no practical importance. Sphacelotheca. The sporocarp is sharply defined, and consists of a columella round which the loose mass of spores is disposed, the whole being enclosed in a covering formed by non-sporogenous hyphae. Sphacelotheca hydropiperis (Schum.). De Bary describes SPHACELOTHECA. 303 this fungus as follows: “Sphacelotheca forms its compound sporophore in the ovule of its host. When the ovule is normally and fully developed in the young flower, the parasite, which always grows through the flower-stalk into the place of insertion of the ovary, sends its hyphae from the funiculus into the ovule, where they rise higher and higher and surround and penetrate its tissue to such an extent as almost entirely supplant it, and thus an ovoid fungus-body of densely interwoven hyphae takes the place of the ovule. The micropylar end of the integu- ments alone escapes the change, and remains as a conical tip (Fig. 164 C’) on the apex of the fungus-body and gradually turns brown and dries up. The fungus- body is at first colourless and uniformly composed of much- branched hyphae, which are woven together into a compact mass and have the gelatinous walls of the simple sporophore of Ustilago to be described below. FG. 164.—Sphacelotheca hydropiperis in the flower of Polygonum Hydropiper. with terminal conidia. As this process is repeated indefinitely, chains of conidia are formed. Fusion of conidia never occurs. U. Kmetiana Magn. Magnus? describes this as destroying and filling with black spore-powder the ovaries of Viola tricolor (var. arvensis). U. filipendula Fuck. occurs particularly on petioles and leaf-ribs of Spiraea Filipendula. Brefeld found the spores germinating after a year. U. (%) italica (Sacc. et Speg.). In seed of Castanea vesca. U. purpurea Hazsl. Ovaries of Dianthus deltoides and D. prolifera in Hungary. U. (7) coralloides Rostr. In roots of Turritis glabra in Denmark. U. orobanches (Fr.). In roots of Orobanche. U. (7) monotropae (Fr.) In roots and stems of Monotropa in Belgium. U. Johansonii (U. Junci. Lag.) In leaves of Juncus filiformis in Switzerland. Tuburcinia. Spores forming balls as in Uvocystis, but all are equally capable of germination. The spore-aggregations form large or small, slightly thickened spots and crusts, which do not cause very marked deformation of the host. Germination results, as in Tilletia, in the formation of a promycelium bearing a tuft of conidia at one end. White conidia are also produced from the mycelium on the host-plant. Tuburcinia trientalis (Berk. et, Br.)* (Britain and U.S. America). Plants of 7'rientalis ewropaea attacked by this fungus are conspicuous in early summer by their swollen dark-coloured stems and their smaller lighter leaves, which fall prematurely. The conidia appear as a white mould-like coating on the lower 'Prillieux, Bullet. de la Soc. botan. de France, 1880; and Brefeld (/oc. cit.), Heft xu. *Magnus, Naturforsch. Fr. d. Prov. Brandenburg, XXx1. ®*Woronin, Senckenberg. naturforsch. Gesell., 1881. Plates I., U., II. 320 USTILAGINEAE. side of the leaf. The black spore-masses are formed in the rind-parenchyma, and sometimes in the pith; they are set free by rupture of the epidermis. In autumn the symptoms are different. The plants appear normally developed, and have no coating of conidia; dark swollen spots, however, appear on the leaves and _ leaf-petioles, in consequence of the massing of black spore-balls in the par- enchyma under the epidermis. The summer mycelium consists of colourless irregularly branched and slightly septate hyphae occupying the intercellular Fic. 175.—Tuburecinia trientalis. Spore- Fic. 176.—Apex of an isolated promy- mass germinating ; several promycelia have celium from Fig. 175; it carries a whorl of been produced and are proceeding to form branches, some of which have fused in pairs; whorls of branches. (After Woronin.) all are developing conidia. (After Woronin.) spaces of the pith and rind-parenchyma, also the vessels. The hyphae apply themselves closely to the cell-walls, and certain short branched hyphae actually penetrate into the cells. The spore-masses are developed from delicate branched multiseptate filaments of the vegetative mycelium. They begin as two or three little cells round which a coil of hyphae is formed; the central cells, increasing in number and size, become a ball of dark smooth-coated spores, while the enveloping coil of hyphae disappears. The spores germinate during the same autumn, frequently in the position of their formation. A promycelium is first formed, and on its extremity a circlet of conidia arises; there- TUBURCINIA. 321 after the promycelium becomes divided by cross-septa in its upper part, and the conidia too are frequently divided by one or two septa. The two promycelial cells become detached, while the conidia begin to fuse together by means of out- growths near their base; thereafter each conidium gives out a secondary conidium, into which the _ plasma-contents pass over. A similar formation of secondary conidia may take place without previous fusion of the primary conidia. The conidia fall apart, and they, as well as the upper promycelial cells thereby left isolated, grow out as hyphae. It must be these hyphae which infect the rudimentary shoots of T'rientalis when they are already partially formed for next year. The resulting mycelium permeates the shoots in the following spring, and branches of it emerge through the stomata, or pass between the epidermal cells and break the cuticle, to grow up either at once as conidiophores, or to form on the surface of the leaf a web from which conidiophores arise. The pear-shaped conidia are attached by their broader side, and easily fall off, leaving the conidiophores free to produce new conidia. The conidia are capable of immediate germination, and may produce a lateral germ-tube, which grows directly upwards, and gives off secondary conidia; or the conidia themselves grow out into hyphae, capable, as Woronin proved experimentally, of carrying out infection. Such hyphae penetrate between the walls of adjacent epidermal cells, and give rise to a mycelium which spreads in a _ centrifugal direction and forms the spore- masses. This same fungus has also been found on Luphrasia lutea and Paris quadrifolia. On Euphrasia, according to Winter, it causes formation of large swellings, accompanied by consider- able deformation of leaf and stem. T. primulicola (Magn.) Kiihn.’ (Britain). This smut attacks flowers of Primula acaulis, P. officinalis, P. elatior, P. farinosa, In cases described in Germany, the blooms were generally attacked in the filaments or connective of the stamens, but also in the anthers, the ovaries, pistil, stigma, and sometimes in the calyx-tube ; while the whole flower-head was more or less discoloured by the black spore-dust. The mycelium permeates ‘Magnus, Botan. Verein Brandenburg, 1878. Kuhn, ‘die Entwickelungs- gesch. d. Primelbrandes,” Naturforsch. Gesell. 2u Halle, i892. x 322 USTILAGINEAE, the whole host and hibernates in the root-stock. The spores are developed from the ends of hyphae in the host-tissue, and are either isolated or joined into packets, They germinate easily in water, and produce either a fine germ-tube, or a thick promycelium with four oblong conidia on its apex. The conidia are easily detached, and either develop to fine hyphae, or give off secondary conidia. Germination on the whole is similar to that of 7. trientalis, Conidia may be also produced directly on the host-plant ; these were first described by Kiihn, who named them Paipalopsis Irmischiae; later, however, he succeeded in infecting plants of Primula with the conidia, and in proving their relationship to this 7ubureinia. T. Cesatii Sorok. occurs on geraniums in Russia. Here, according to Setchell, the following American genera should be placed : Burillia: BL. pustulata on Sagittaria. Cornuella: C. lemnae on Lemna polyrhiza, Doassansia. Spore-masses consisting of numerous spores capable of ger- mination, enclosed in a layer of sterile cells. The latter are most conspicuous in the species frequenting aquatic plants, and are filled with air,—Brefeld regards them as swimming-organs, The spore-masses le in groups embedded in the host-plant. The species inhabit plants with an aquatic or moist habitat, and produce on them leaf-spots with black pustules. Fisch’ investigated the life-history of Doassansia sagittariae. He found an intercellular mycelium which, inside the stomata, formed sporocarps, consisting of sclerotium-like coils of hyphae enclosing several cells which form spores. The spores on germina- tion give rise to promycelia, which produce sporidia in a manner similar to Hntyloma. The sporidia easily germinate in water, and can immediately infect young leaves. The germ-tubes creep on the surface of leaves, and attaching themselves by an adhesion-disc over the wall between two adjacent epidermal cells, they penetrate this wall. The hypha, while passing 1C, Fisch., Ber. d. deutsch. botan. Gesell., 1884, p. 405. Cornu, Annal. d. set. natur. XV., 1883. Setchell (Botanical Gazette, 1894) records the American species and comments on them. DOASSANSIA. 323 through the wall, remains thin, but on emerging into an inter- cellular space it soon thickens and branches into a mycelium. Infection results in the appearance of yellow spots, due to rapid destruction of the chlorophyll and death of cell-contents, Experiments in germination have been carried out by Setchell and Brefeld. Doassansia sagittariae (West.) (Britain and U.S. America). In leaves of Sagittaria. The spores, according to Brefeld, germinate in water, after hibernation. They produce unicellular promycelia with a terminal tuft of more or less spindle-shaped conidia, which at once begin to sprout and fall off. On the surface of a nutritive solution they continue to sprout yeast- like, and form close mouldy coatings. (Doassansia is the only genus of the Tilletiae in which Brefeld found yeast-like sprout- ing of conidia.) D. alismatis (Nees) (Britain and U.S. America). This inhabits leaves of Alisma Plantago and A. natans, producing knotty swellings. The spores are enclosed in a layer of com- panion-cells containing air, whereby the masses swim on water. On the promycelium the conidia arise from tufts of conidio- phores; they fuse in pairs, and secondary conidia are developed from each pair or even from single conidia. D. Niesslii (de Toni) forms small spots on leaves of Buto- mus umbellatus, The spores are surrounded by companion-cells containing air. They germinate before leaving the spore-patch, and produce ‘conidia, even secondary conidia, before rupture of the host-epidermis takes place. Brefeld describes the spores as germinating in water to form a very short promycelium with short thick conidia which fuse in pairs and give off larger sécondary conidia from their apices. In nutritive solution conidia are developed, which give off septate filaments whence further conidia arise. Aerial conidia are ultimately developed. Magnus found that the spores of D. alismatis, D. Niesslii, and other species germinated at once on reaching maturity. Brefeld, however, found that this took place only after they had lain over winter. It may be that here, as with some higher plants (eg. Pinus Cembra), there is an immediate capability of germination, but also a deferred, the latter requiring 'Setchell, Annals of Botany, v1., 1892. Brefeld, Schimmelpilze, Heft xu1., 1895. 324 USTILAGINEAE. to be preceded by a considerable resting-period, during which germination will not take place. D. Martianoffiana (Thiim.). In leaves of Potamogeton natans and P. gramineus. D. occulta (Hoffm.). In fruits of species of Potamogeton. D. intermedia (Setch.). An American species found on leaves of Sagitturia variabilis. D. comari (Berk.). In leaves of Comarum palustre in Britain. D. limosellae (Kunze.). In flowers of Limosella aquatica. D. hottoniae (Rostr.). In leaves of Hottonia palustris in Denmark. Thecaphora. Spores, large, spherical, and inseparably united into packets of several spores. Germination results in the formation of a Fic. 177.—Thecaphora hyalina. Pluricellu- Fic. 178.—Sorosporium saponariae (var. lar spore, with two cells (spores) germinating. Lychnidis dioicae.) Mature spore-mass, and (After Woronin.) spores germinating. (After Woronin.) promycelium from the apex of which a single conidium is produced. Thecaphora lathyri Kiihn. Spore-balls formed in the seeds of Lathyrus pratensis, and escaping as a brown powder on dehiscence of the pods. The spores germinate in water with formation of a promycelium bearing a single apical conidium, which produces a hypha, but never secondary conidia. In nutritive solutions the spores produce a mycelium from which conidia are continuously given off. Th. hyalina Fingerh. (Britain). This occurs in fruits of species of Convolvulus. _Woronin describes the spores as having germ-pores through which a septate germ-tube is emitted; the individual cells of the germ-tubes develop into hyphae, without formation of conidia. Th. affinis Schneid. In fruits of Astragalus glycyphyllus (U.S. America). Th. Trailii Cooke. In flowers of Carduus heterophyllus in Scotland. THECAPHORA. 325 Th. Westendorpii Fisch. In Loliwm perenne in Belgium. Th. pimpinellae Juel. In fruits of Pimpinella Saxifraga in Sweden. Th. aurantiaca Fingh. In leaves of Urtica dioica. Th. pallescens Fingh. In leaves of Fragaria collina. Sorosporium. Spore-formation takes place in a mass of twisted gelatinous hyphae. Spores at first embedded in a gelatinous investment and united into packets, but later becoming separate. Promy- celium filiform and septate. Sorosporium saponariae Rud. This causes deformation of flowers of Dianthus deltoides, Saponaria officinalis, Silene inflata, and S. velutina, Stellaria Holostewm, Cerastium arvense, Lychnis dioica, and Dianthus prolifer. S. dianthi Rabh, on Dianthus prolifer, is probably identical with the preceding species, We append here as doubtful Ustilagineae, the genera Gvaphiola Schinzia (Entorrhiza), Tuberculina, and Schroeteria. Graphiola. The sporocarps of this genus are formed on the surface of plant-organs containing mycelium; they are litle spherical structures enclosed in a peridium, and contain filamentous septate hyphae. The hyphae may be sterile or fertile; the spores are produced on lateral cells of the fertile hyphae. From the germinating spores, either a thread-like mycelium or spindle-shaped conidia arise. Graphiola phoenicis Pait.! (Britain.) This fungus is a parasite on leaves of palms (eg. Phoenix dactylifera and Chamerops humilis) in the open in Italy and other Mediter- ranean countries, in hot-houses elsewhere. The sporocarps make their appearance as little black protuberances on both sides of the leaf. The mycelium forms a close hyphal tissue, which encloses and kills parenchymatous cells, displaces the bundles of sclerenchyma, and ruptures epidermis and hypoderm. De- formation is, however, localized to these spots. 'Ed. Fischer, ‘‘ Beitrag z. Kenntniss d. Gattung Graphiola,” Botan, Zeitung, 1883, 326 USTILAGINEAE. The sporocarps consist of a two-layered peridium, a sporogenous layer, and tufts of sterile hyphae. The outer layer of the peridium forms the outer layer of the black protuberances on the leaves; the inner layer is delicate. The sporogenous hyphae originate from the centre of the underlying hyphal tissue, and form a palisade-like layer in the bottom of the sporocarp cavity, the remaining space being filled with spores and tufts of barren hyphae. These latter hyphae rise amongst the sporogenous ones, and project as a fine brush-like tuft out of the ruptured peridium. The sporogenous hyphae grow vertically upwards, and become septate, forming chains of loosely united, roundish, hyaline cells or joints. The terminal joints give off several spherical cells laterally, and die away, leaving the cells loose in the sporocarp cavity. From division of the spherical cells yellow spores result, and, on rupture of the peridium, are carried out on the tufts of sterile hyphae to be scattered by wind. The spores germinate in water, and produce either a promycelium or conidia. Gr. congesta Berk. et Ray. occurs on leaves of Chamerops Palmetta. Schinzia (Entorrhiza).’ Spores produced on the ends of lateral branches of a mycelium - in the cortical cells of the root of the host-plant. Germina- tion results in production of a simple or branched sporophore (promycelium), from which kidney-shaped conidia (sporidia) are produced. Schinzia cypericola Magn. This causes deformation of the roots of Cyperus flavescens (Fig. 179). Sch. Aschersoniana Magn. causes swellings on the roots of Juncus bufonius | Britain]. Sch. Casparyana Magn. In roots of Juncus Tenageia. Sch. digitata Lagerh. In roots of Juncus articulatus. Sch. (Naegelia) cellulicola Naeg. In roots of /r’s in Switzerland. Sch. (Entorrhiza) solani Faut.? [This is given as the cause of a disease on potato. The plants droop and ultimately rot at the neck, the leaves become yellow, and neither flowers nor tubers are produced.] (Edit.) 1p, Magnus, at Botan. Verein d. Prov. Brandenburg, 1878; ‘‘ Ueber einige Arten d. Gattung Schinzia,”’ Ber. d. deutsch. botan. Ges., 1888, p. 100; C. Weber, Botan. Zeitung, 1884. 2 Fautrey, Revue mycolog., 1896, p. 11. TUBERCULINA. 327 Tuberculina. Mycelium parasitic on hyphae and spore-patches of Uvedineae. Short rod-like hyphae spring from the spore-patches, and give off from their apices, globose conidia, which on germination produce branched promycelia bearing sickle-shaped conidia. FIG. 179.—Schinzia cypericola on Cyperus flavescens. Several roots show palmately- divided swellings. Isolated spore. (After Magnus.) Tuberculina persicina Ditm. The lilac-coloured spores are found on aecidia of Peridermium pini and other aevidial forms, also on some species of Cacoma.' (Britain and U.S. America.) ' Plowright (British Ustilagineae) gives also Aec, asperifolii, Aec. tussilaginis, and Roestelia lacerata as hosts. 328 USTILAGINEAE. T. maxima Rostr. Occurs on rust-patches on Weymouth pine. It has larger spores than the preceding species. Schroeteria.' Spores joined in pairs, rarely in threes, with their broad faces together. They are developed from single joints of a septate non-gelatinous mycelium, particularly from short curled lateral hyphae. Spherical conidia are produced, like those of Penicillium, by imtercalary growth in chains from the end of a conidiophore which is generally unbranched. Schroeteria Delastrina (Tul.) occurs in seeds of Veronica arvensis, V. hederifolia, V. triphylla, and V. praecox. The spores germinate in water, and produce conidia incapable of further de- velopment, even when transferred to a nutritive solution. In such, however, spore-germination is more vigorous, and an abundant mycelium results, but it seems to be unable to produce conidia. Sch. Decaisneana (Boud.). In seeds of Veronica hederifolia at Paris, UREDINEAE. The Uredineae or Rust-fungi possess several forms of spores, one of which, the teleutospore, is rarely, if ever, absent from the life-cycle of any species. The teleutospores consist of one, two, or more cells enclosed in a thick coat of dark colour, and thereby well adapted to carry the fungus over winter. When germination occurs, each cell of a teleutospore gives off a germ- tube through a pore or thinner place in its wall, and from this a promycelium? is formed, consisting as a rule of four cells. Each teleutospore originates from a sporophore of its own, and in the course of development two nuclei, originally present in each cell of the young teleutospore, fuse together. When germination takes place, and the promycelium is formed, the single cell-nucleus, derived as above, divides into two, then into four, so that a nucleus is produced for each of the cells of the promycelium. From the promycelium four sterigmata are given off, and each produces a single sporidium.2 These : owas the species as forms of higher fungi, not as Ustilagineae (Heft XII., p. 204). * Brefeld considers that the promycelium and sporidium are respectively a basidium and a basidiospore. UREDINEAE. 329 sporidia on germination give infecting mycelial hyphae. In the case of Coleosporium, the promycelium is formed inside the teleutospore in a manner similar to the Protobasidiomycetes. Besides teleutospores, there occur uredospores. These are given off from patches or sori throughout the summer till autumn, when they are followed by teleutospores on the same sori. The uredospores somewhat resemble the teleutospores, but generally consist of one cell only with a thinner coat of lighter colour; they either germinate at once without a resting period, and give rise to a germ-tube capable of direct infection of new hosts; or less frequently they are resting-spores for a time. A third form of spore occurring in the life-history of the Uredineae is the aecidiospore, produced in a special structure, the aecidium. The aecidium is developed inside the leaves or other organs of the host-plant, and when mature ruptures the overlying epidermis; it has as a basis a firm hyphal tissue, the upper surface of which becomes a dise of short erect sporo- phores. From each sporophore there is formed by intercalary growth a chain of cells consisting alternately of spores and smaller intermediate cells, which do not become spores. The youngest cells in an aecidium are those next. the sporophore- disc, and they are forced outwards by intercalation of younger cells between them and the disc. The cells so produced become alternately intermediate cells and spores; the former increase for a time, then decrease and disappear, the spores however continue to increase in size as the chain grows forward and to take on the characters of the mature aecidiospore till they are finally shed from the aecidium. The production and distribution of aecidiospores may thus go on continuously for a considerable time. The sporophores at the periphery of the disc do not however produce spores; chains of cells are also produced from them by interealary growth, but the cells are of equal size, and remain closely connected with their neighbours, so as to form a membranous covering over the spore-sorus, this is the so-called peridium, on rupture of which the aecidio- spores escape. In many Uredineae the peridium is suppressed (Caeoma); in others (Phragmidium) it is replaced by other structures, the paraphyses. The spores of the genus Hadophyllum are produced in series in aecidia enclosed by a peridium, but in 330 UREDINEAE. germination they behave more like typical teleutospores than aecidiospores. Before the relationship of these various forms of spores was known, Accidium and Caeoma were regarded as independent groups, and named as such; even yet many isolated forms of uredospores, teleutospores, and aecidiospores are known, the relationships of which are quite obscure. The aecidia are always preceded or accompanied by a further form of spore produced in a special structure of its own. These spores have hitherto been called spermatia, and their sporocarps spermogonia, on the assumption that they were male organs. Now, however, many of them are known to be capable of germination in artificial nutritive solutions, hence they are more probably a form of asexual bud, and better named conidia, their sporocarps pycnidia. The pyenidia are flask-shaped structures sunk in the tissue of the host, with a pore or mouth emerging through the host-epidermis; they generally occur in leaves, and occupy the upper epidermis, the aecidia occurring on the lower. From the mouth of the pycnidium there frequently emerges a tuft of fine filaments, outgrowths from the inner wall of the flask. The pyenidia possess a lively colour and flowery odour, hence it has been suggested that the conidia may be distributed by insects; but they do not appear to be able to germinate in the open, and infection- experiments with them have never as yet succeeded. On this account they are regarded as degenerate structures.! The various forms of spores are also distinguishable by the manner in which they bring about infection. |Teleutospores on germination produce sporidia, which pierce the membranes of the prospective host at a spot where two adjoining cells are in contact, and thus make their way into the intercellular spaces. Uredospores and aecidiospores, however, first seek a stoma and enter the intercellular spaces of the host through it. The following different forms of Uredineae exist: (1) Those which possess teleutospores alone, e.g. Chrysomyxa abietis; (2) those with teleutospores and uredospores, e.g. Puccinia prunt spinosae ; (3) those with all the forms of spores, e.g. Puccinia graminis ; (4) those without uredospores, e.g. Gymnosporangium. 1Rathay, ‘‘ Untersuchungen iiber die Spermogonien d. Rostpilze,” Denkschrift d. Wiener Akad. d. Wissensch., 1883. UREDINEAE. 331 The different forms of spore may be found on one and the same host-plant (autoecious Uredineae), or the aecidiospores and pycnidial conidia may frequent a different host from the uredo and teleutospore-forms (heteroecious Uredineae).' A mycelium may be produced from the germinating aecidio- spores, uredospores, or sporidia. It spreads throughout the intercellular spaces of attacked organs and causes thickening, distortion of the tissues of its host, or the formation of “ witches’ brooms.” Nutriment is frequently obtained by means of cone- shaped or button-like haustoria in the interior of host-cells. Hibernation of rust-fungus is most commonly attained through the teleutospores, the thick coats of which make them peculiarly suited to pass through a lengthened resting-period. Some forms, however, hibernate by uredospores, by aecidiospores, or by the mycelium remaining on or in living perennating stems, twigs, or underground rootstocks of their host. . Aecidiospores on germination produce, as a rule, a mycelium which gives rise to uredo- or teleutospores, rarely to aecidiospores (eg. Puccinia senecionis and Uromyces ervi).2 Uredospores on germination, produce a mycelium from which ureduspores are first given off, then teleutospores. The sporidia of teleutospores give rise to a mycelium which frequently produces pycnidia and aecidia. In rare cases, the sporidia of species, which normally form aecidia, are said to develop a uredo-mycelium (e.g. Puce. graminis according to Plowright). The Uredineae are for the most part strict parasites, and exhibit marked adaptation to their respective host-plants. Several of the polyxenous members frequenting several species of host- plant have been found to vary according to their habitat, so that one and the same species assumes a slightly different form on each 'The phenomenon of heteroecism was till quite recently known only amongst the Uredineae. Woronin and Nawaschin have, however, recently pointed out that it exists in Sclerotinia ledi, one of the Ascomycetes (p. 277). The conidia of this species are produced only on Vaccinium uliginosum, the apothecia only on Ledum, and alternate with each other, so that the Ledum can be infected only by germinating conidia, the Vaccinium by germinating ascospores. *Dietel (Naturforsch. Verein in Vienna, 1894) pointed out further cases of this kind, in which aecidia were produced the summer through, and no uredospores, while in autumn teleutospores were formed. He has more recently stated the general conclusion (Flora, 1895, p. 394); that with these species of Uromyces and Puecinia, which produce gab and teleutospores, but no uredo- spores, the aecidiospores are capable of reproducing aecidia when no perennating mycelium is present. Similarly with those few species which produce a very small number of uredospores. Bow UREDINEAE. host-species. [I have previously shown,! with regard to the mistletoe (Viscum album), that the different forms on Pinus, Abies, and various broad-leaved trees, which some authors regard as distinct species, might equally well be regarded as forms of one species differing slightly on account of their different substrata. Magnus? designates as “ habitat-races” these forms of heteroecious Uredineae whose aecidial generation has become adapted in some varying degree to each of their respective species of host-plant. Thus the various forms of Ser. III. Aecidia on Rhamnus dahurica (Puce. coronata var. himalensis, Barcl.). Indian forms on Brachypodium sylvaticum, (Piptatherum holeiforme, and Festuca gigantea,) of which nothing more is known. Ser. IV. Aecidia unknown, probably do not exist. 6. Var. melicae on Melica nutans. Amongst our cereal crops the oat alone is attacked by this species, and much damage may result. The uredo-patches have no paraphyses like the preceding species, and they form reddish-yellow spots and stripes; the teleutospore-patches are black. The upper cell of the teleuto- spores is surrounded by a crown of six or seven blunt teeth. The presence of aecidia on Rhamnus is accompanied by thickening and twisting of young shoots, and blister-lke de- formation of leaves, calyces, and ovaries. Wakker! thus summarizes his investigations on the anatomical changes induced by the funeus on Rhamnus Frangula: “It causes the cells of every part to become abnormally enlarged, at the same time giving rise to an orange coloration of the cell-sap and an accumulation of starch; there is no longer any formation of interfascicular cambium, and there is a partial or complete 1Wakker, Pringsheim’s Jahrbuch, 1892. PUCCINIA. 347 suppression of secondary vasa, mucilage canals, and calcium oxalate.” The deformation induced by P. coronata on Rhamnus cathartica was investigated by Fentzling.” The changes were relatively shght: the parenchymatous cells of the rind were enlarged and separated by large intercellular spaces; so also the paren- chyma of the bast; vessels were more numerous in the wood affected; the epidermal cells in some parts of the leaf were broadened and those of the mesophyll enlarged, abnormally shaped, and with large intercellular spaces; in diseased leat- stalks the epidermal cells are thinner-walled and broader, while all parenchymatous cells become enlarged, thinner-walled, and with many intercellular spaces; the fibro-vascular bundles are enlarged, chiefly from an increase of the wood-parenchyma ; this tissue, in normal petioles, occurs as single rows of cells running radially between the vessels, whereas, in diseased places, three parallel layers of cells may separate neighbouring vessels. P. dispersa Eriks.et Henn. Brown-rust. (Britain.) The follow- ing specialized varieties of this species have been distinguished : Ser. I. Aecidium on Anchusa arvensis and A. officinalis (Aee. anchusae). 1. Var. secalis on Secale cereale.” Ser. II. Aecidium unknown. (Whether distinct varieties, somewhat uncertain.) 2. Var. tritici on Triticum vulgare. >}. Var. bromt on Bromus arvensis (and Br. brizaeformis). 4. Var. agropyri on Triticum ‘repens. P. rubigo-vera (D.C.) (P. straminis Fuck., P. striaeformis West.). (Britain and U.S. America.) This, in its uredo- and _ teleuto- spore stages, frequents various grasses, while the aecidia occur on Boragineae. A variety on species of Hordewm has been designated P. simplex. The teleutospore-patches are enveloped in numerous brown paraphyses; the teleutospores have very short stalks. The anatomical changes produced in leaves beset with aecidia have been stated by Wakker as follows: The swelling of the leaf-petioles is due to enlargement of their cells; the large intercellular spaces of the spongy parenchyma are no longer 'Fentzling, Inaugural Dissertation. Freiburg, 1892. *Found along with the Aecidium at Montrose (Scotland) by Prof. J. W. H. Trail. (EKdit.) 348 UREDINEAE. present; the palisade layer is doubled, and rupture of the epidermis takes place; chlorophyll-formation is suppressed, the cell-sap becomes yellow, and starch tends to accumulate. P. dispersa may cause serious damage to wheat and rye; P. rubigo-vera, also on barley and oats. The spore-patches are found on stalks and leaf-sheaths more than on the lamina. The mycelium may hibernate in grasses, so that the fungus is not dependent on the aecidial stage; for this reason the disease is not easily combated. P. glumarum Eriks. et Henn. Golden-rust. This species, hitherto generally included under P. rubigo-vera (D.C.) has been separated by Eriksson and Henning. Experimental infection on Boragineae gave negative results. Eriksson distinguishes the following specialized varieties of this species: A. Definite (and undoubtedly distinct). 1. Var. tritici on Triticum vulgare. 2. Var. hordet on Hordeum vulgare (somewhat uncertain). 3. Var. elymi on Elymus arenarius. agropyrt on Triticum repens. i a =} =) B. Not sharply defined : 5. Var. secalis on Secale cereale. The uredospore-sori are lemou-yellow in colour, and form lines on the leaf-blade which may run together and reach a length of 10mm. The teleutospore-sori form long, fine, brown or black lines: the sori are divided into numerous chambers, each enclosed in a circle of curved brown paraphyses. The spores germinate in the autumn of the same year. The pro- mycelium is yellow till the spores are abjointed; in this way it is distinguished from P. dispersa. P. poarum Niels. (Britain). Uredo- and teleutospores on Poa. According to Nielson, the aecidia occur on Tussilago, Petasites, and Adenostyles. Fentzling (loc. cit.) has described certain anatomical changes which accompany deformations due to the aecidia. P. phlei-pratensis Eriks. et Henn. This has a hibernating mycelium which produces uredospores continuously on Phlewm and probably also on Festuca. Aecidia have not as yet been observed. ' Eriksson and Henning (/oc. cit.). PUCCINIA. 349 P. agrostidis Plowr.' Teleutospores on Agrostis vulgaris ; aecidium = Aec. aquilegiae Pers. (Britain and U.S. America). P. festucae Plowr.! Uredo- and teleutospores on Festuca ovina and F. duriuscula ; aecidium= dec. periclymeni Schum. (Britain). P. phragmitis (Schum.). Uredo- and teleutospores on Phragmites. Aecidium = Aec. rubellum on Rumeax crispus and other species of Rumer, also on Rheum. (Britain and U.S. America.) P. Trailii Plowr. Uredo- and teleutospores on Phragmites communis. ‘Aecidium on Rumev Acetosa. (Britain.) P. Magnusiana Korn. Uredo- and teleutospores on Phragmites communis. Aecidium on Ranunculus repens. (Britain.)* P. moliniae Tul. Uredo- and teleutospores on Molinéa coerulea. Aecidium (according to Rostrup’s out-of-door experiments), on Orchis repens, O. mascula ; probably also on other Orchideae. (Britain.) P. nemoralis Juel. Uredo- and teleutospores on Molinia coerulea ; aecidium (dec. melampyri Kze. et Schm.) on Melampyrum pratense. P. australis Korn. Uredo- and teleutospores on Molinia in Tyrol ; aecidium (Aec. erectum, according to Pazschke) on Sedum reflexve, S. acre, ete. P, perplexans Plowr. Uredo- and teleutospores on Alopecurus pratensis ; aecidium on Ranunculus acris. (Britain.) P. persistens Plowr. On Triticum repens. Aecidium=Aee. thalictri (Britain). P. sesleriae Reich. On WSesleria coerulea. Aecidium on Rhamnus savatilis. P. Winteriana Magn.’ (/. sessi/’s, Schn.). Uredo- and teleutospores on Phalaris arundinacea. Aecidium on Allium ursinum (Aec. alliatum Rbh.). P. sessilis Schn. (including P. digraphidis Soppitt and P. paridis Plowr.) (Britain.) Uvredo- and teleutospores on Phalaris arundinacea. Aecidium, according to Soppitt,t on Convallaria majalis, also on Majanthemum, Paris, Polygonatum, Lilium canadense and Streptopus Smilacina. Klebahn’s experi- ments confirm the relationship of the aecidium on Majanthemum, Convallaria, Polygonatum, and Paris. P. phalaridis Plowr. On Phalaris arundinacea. Aecidium (Aee. ari) on Arum italicum and A. maculatum. (Britain.) P. agropyri Ell. et. Ev. On Agropyrum. Aecidium=Aec. clematidis D. C. on Clematis Vitalba and C. recta, etc., in Europe and America. P. caricis (Schum.) (Britain and U.S. America), Uredospores and teleutospores on species of Carex. | - 4 > a GYMNOSPORANGIUM. 387 may be as long as 10 mm. and are bent like a horn (Fig. 219). A similar case is described by Barclay! in which the peridia of aecidia on Rhamnus dahurica were very long if produced in dry weather, but short if in moist weather.” The aecidiospores are shed during the early part of June, and germinate at once on the bark of young juniper-twigs; the mycelium growing thence into the spurs or branches to spread and hibernate. Teleutospores which germinate on Pomaceae other than species of Crataegus have apparently a normal mycelium, but produce pycnidia only, or aecidia with i ©. OPSOS9) Fic, 220.—Cross-section through a swelling caused by Gymnosporangium on Juniper-stem ; parenchyma with large cells and thin walls is present in abnormal quantity. (After Woernle.) peridia differing from those on Crataegus. My own experiments on the quince and mountain ash regularly produce pycnidia only. Wakker*® summarizes the anatomical changes induced in deformed shoots of hawthorn as follows: cork, collenchyma, sclerenchyma, and chlorophyll are not formed, lignification of the cells of medullary rays no longer takes place, and there are few intercellular spaces. Interfascicular cambium is not formed, while activity of the intrafascicular cambium is suspended at an early period, so that the vessels remain incompletely developed. The epidermis is irregularly formed and liable to rupture, All parenchymatous cells undergo enlargement in a radial direction. Starch is stored up in large quantity, and the formation of calcium oxalate is diminished. 1**On the life-history of Puccinia coronata var. himalensis,” Trans, Linnean Soc., London, 1891, *This probably is the explanation of the long gp obtained by Peyritsch and described by Magnus ( Berichte d. naturwiss. medic, Verein, Innsbruck, 1892-93). * Pringsheim’s Jahrbuch, 1892. 388 UREDINEAE. The anatomical changes induced in diseased plants of Juniperus communis by G. clavariaeforme were investigated by Woernle under my direction. His results were these: in vigorous branches, increased growth took place in the wood, bast, and rind; in weakly and poorly-grown branches, the wood increased less in proportion to the bast and rind. The most marked increase took place in the bast, and to an almost equal extent all round the branches. This abnormal growth absorbs so much il y | { Fic. 221.—Cross-section of a tract of Fig. 222.—Radial longitudinal section parenchyma in a malformed Juniper- through a zone of parenchyma similar to twig. (After Woernle.) Fig. 221. (After Woernle.) water and plastic material that higher parts of the branch gradually die off, and dormant buds break out on the swelling. Increased growth results in increase in the number of medullary rays, while in the tangential section their height is increased from 2-10 cells to 10-20 and more; the wood parenchyma is also more abundant, and together with the rays frequently forms large masses of parenchyma in the wood (Figs. 220-223). The tracheae no longer follow a straight course, and numerous intercellular spaces appear between them; the tracheal walls frequently become thickened and have an increased number of GYMNOSPORANGIUM. 389 fissure-like pores in place of bordered pits. The wood-elements in cross-section are no longer round but polygonal; the bast becomes very irregular, parenchyma grows rapidly, bast fibres remain thin-walled and have no longer a straight course. The mycelium fills the bast and rind, forming masses in the inter- cellular spaces; it is easiest found in the tangential section. On the fall of the club-shaped sporophores, a scar is left and under it will be found a layer of cork many cells thick ; when new sporophores are formed in later years, they seldom break through the cork layer, but emerge through some new _ por- tion of the bark. Gymnosporangium tremel- loides Hartig! on Juniperus com- munis. The sporocarps of this species occur on the branches and needles; its aecidia— Roe- stelia penicillata—on leaves of apple (Pyrus Malus), Pyrus Aria and P. Chamaemespilus. This Roestelia is externally very like Fic. 223.—Tangential longitudinal sec- ‘ mee” tion through the parenchyma-zone of that of G. clavariaeforme on _ Fig. 220. (After Woernle.) Crataegus. The markings on the cells of the peridium consist of somewhat wavy lines, not of short rod-like markings as in &. cornuta; and the cells of the peridium are joimed by a characteristic hinge-joint (Fig. 224, 19 and 20). The mycelium perennates in the rind of Juniperus communis and J. nana, causing thickening of the twigs and a premature death of the distal portion above the swellings. The chocolate- brown velvety spore-cushions break out between the bark-scales on the swollen places, about the middle of April (Fig. 225, 1). The teleutospores are two-celled, the earlier formed ones being short, ovoid, and slightly pointed at each end, while the later ones are thinner-walled and often more elongated (Fig. 225, 6-10). 'Hartig, Diseases of Trees, English edition, 1894. Dietel, Forstlich-natur- wiss, Zeitschrift, 1895, p. 348. KE. Fischer, Hediwigia, 1895, p. 1. 390 UREDINEAE. In May or June the cushions swell up and become large brownish-yellow gelatinous clumps, dotted over with dark points, the teleutospores. Promycelia arise from one or more germ- pores in each spore, and give off basidia with sporidia (basidio- Fic. 224.—Aecidia and Pycnidia of various species of Gymnosporangium : G. tremelloides—1 and 2, aecidia on leaf of Pyrus Aria; 5 and 6, aecidia on leaf of Pyrus Malus; 19 and 20, portions of the peridium of an aecidium from 5, showing the peculiar articulation of the cells. G. juniperinun—3 and 4, aecidia on Pyrus Aucuparia; 7 and 8S, aecidia on Amelanchier vulgaris. G. clavariaeforme—9 and 10, aecidia on Pyius latifolia; 11, 12, and 16, aecidia on Crataegus Oxyacantha, grown out-of-doors; 14, the same aecidia, enlarged ; 13, 15, and 17, aecidia on Crataegus Oxyacantha, indoor culture; 18, deformed twig of Crataegus bearing pycnidia. (After Tubeuf.) spores) capable of immediate germination. The gelatinous mass dries up from time to time, leaving a bright yellow scar on the swollen part of the host-branch. The sporidia germinate most easily on species of Sorbus (Pyrus). Infections with GYMNOSPORANGIUM. 391 Gymnosporangium juniperinum L. and G. tremelloides Hart. from twigs and needles of Juniperus communis produced : On Host-plant. Spore-form. Authority. Pyrus (Sorbus) Aucuparia, - - Rostelia cornuta, - - Tubeuf. Aronia rotundifolia, - - - short aecidia, - - Rathay. Pyrus Malus, - : i - Vovonidia = : - Pyrus (Sorbus) Aria, - ae " oe Z Cydonia vulgaris, - - - - Roestelia (2), - - - a Pyrus (Sorbus) Aucuparia, - - (?) - - - Plowright. Pyrus Malus, - - - - - pyenidia, - . - Thaxter. Amelanchier canadensis, - - - Rostelia cornuta, - - “e Pyrus (Sorbus) Aria, - - - Rk. penicillata, - - Hartig. Pyrus Malus, - - - - - R&R. penicillata, : - Nawaschin. Pyrus (Sorbus) Chamaemespilus, - R, penicillata, - - Mespilus macrocarpa, - - - spots, - - - - Peyritsch. Pyrus communis, — - - - - thick spots, - = - < Pyrus (Sorbus) Aria, - - - pyenidia and aecidia, — - Pyrus (Sorbus) Aria x Chamaemesp., thick spots, - - - me Pyrus Malus, - - - - - pyenidiaand aecidia, — - - Pyrus (Sorbus) Chamaemespilus, - pycnidia only, - - ot Pyrus (Sorbus) Aucuparia, - - pyenidia and aecidia, — - a Aronia rotundifolia, - - - a oe x Pyrus (Sorbus) torminalis, - - pycnidia and spots only, A Crataegus Pyracantha, - = = pe 49 i Cydonia vulgaris, - - - - ss 4 2 Pyrus Malus, - - - - - ostelia penicillata, - Rostrup. Formation of pycnidial spermogonia always precedes that of aecidia. This fungus is of practical import on account of its occurrence on leaves of apple-trees. Its attacks may be very virulent and widely distributed. Eriksson mentions that near Stockholm it is common on apples, and so virulent that many trees have every leaf studded with Roestelia. (American apple-trees suffer from Stoestelia pirata, the aecidia of Gymnosporangium macropus and other species. See p. 402.) Gymnosporangium juniperinum (L.) ((. conicum Hedw.) (Britain and U.S. America). This species, also frequenting Juniperus communis, is distinguished by its shorter spores, which, as Dietel pointed out,’ have a colourless papilla over each germ-pore. The teleutospores are found on both twigs and needles, on the former, however, they are much smaller ' Forstlich-naturwiss. Zeitschrift, 1895, p. 378. 392 UREDINEAE. than those of G. tremelloides. The aecidiospores—Roestelia cornuta—occur on species of Pyrus (Sorbus); they are much smaller than those of Roestelia penicillata, The Roestelia themselves are long, curved, and horn-like, while the walls of the peridial cells are beset with short processes (Fig. 224). Where Pyrus Aucuparia occurs mixed with Pyrus Malus, it has been observed that Roestelia cornuta is confined to the former species exclusively. The oestelia is the cause of a Fia. 225.—Gymnosporangium juniperinum and G. tremelloides. 1, Young spore- cushions breaking through the bark; 2, the same in swollen condition ; 3, gela- tinous cushion arranged to show its lower surface; 4, Juniper-needle with three spore-cushions; 5, young Juniper plant bearing cushions on its needles; 6 to 10, spores of various kinds, to show the variation in size, shape, and thickness of wall; 11, cell of a promycelium with a sporidium attached; 12, germinating sporidium. (After Tubeuf.) marked deformation of leaves, petioles, and even (though rarer) fruits of Pyrus Aucuparia and Aronia rotundifolia, both in the lowlands and mountains. I have produced Roestelia cornuta on Pyrus Aucuparia by artificial infection with portions of spore-cushions from twigs of juniper, and have observed a mountain ash in closed forest, with abundant oestelia, directly beneath an overhanging juniper with diseased needles. GYMNOSPORANGIUM, 393 Woernle investigated the anatomical changes induced by the various Gymnosporangia frequenting the twigs and needles of Juniperus communis. In the needles the mycelium lives inter- cellularly, at first outside the endodermis, but later also penetrating within this. The sporogenous cushions originate on the upper surface of the leaf to right and left of the middle nerve, where the stomata occur and hypoderm is absent. At these places a cushion or stroma of pseudoparenchyma is produced and ruptures the epidermis (Fig. 226). This however is at once healed over by a cork-formation round the margin of the cushion, again to be ruptured as the latter increases in size, once more to be healed by cork-formation, and so on. In Fic. 226.—Comparison of (7) normal Juniper-needle with one (/) bearing teleuto- spores of Gymnosporangium. In a the double outline indicates the hypoderm ; the central vascular bundle and an underlying resin-canal are shown. (After Woernle.) this way a corky layer is formed under the sporogenous cushion and gradually displaces it. If in a following year the cushion be again formed, the scar is ruptured and heals as_ before. Needles frequently remain in position for two, three, or four years, but most of them fall off in the first autumn. Under the sporogenous cushion the cells of the mesophyll increase both in number and size. In considering the twig-deformations, Woernle distinguishes the form assumed by the Gymnosporangium on the needles, as just described, from a form which inhabits the thicker twigs. Both cause deformation of twigs, but their effects differ as follows: “The needle-inhabiting form can only cause a slight swelling extending almost regularly round the whole twig; the twig- inhabiting form, on the other hand, always gives rise to a very 394 UREDINEAE. marked swelling on one side only (Fig. 227). In the needle- form the swelling results from increased growth of the rind, with a simultaneous decrease of growth of the wood; in the twig-form the growth of both wood and rind is much increased. With the twig-inhabiting form the medullary rays and wood- parenchyma increase, and at the same time become filled with mycelium (Fig. 228); whereas with the other form the medullary rays are at most only somewhat broader, and no mycelium can be found in the wood. The greatly swollen rind in the case Fig. 227.—Section of a nine-year twig of Juniper attacked by Gymnosporangium. Fia. 228.—Tangential section through diseased The rind under the spore-cushion is wood beneath a spore-cushion. The wood- much thickened ; the wood towards the elements are much displaced by abnormal tracts same side is much broken up by tracts of parenchyma. (Only one of the latter has been of parenchyma. (After Woernle.) filled in, the others left blank.) (After Woernle.) of the twig-inhabiting form is due more to increased growth of the cortical cells than to increase of bast-parenchyma; in the needle-form, however, the swelling is the result of increase of the bast, especially of the bast-parenchyma. In_ twigs infected by the needle-forin, the mycelium may be found all round, but it has difficulty in making its way radially to the cambium; in the twig-form the mycelium, as early as the spring following infection, will be found to be in close contact with the cambium on the infected side, although it requires several years to pass round to the cambium on the opposite side of the twig. The mycelium and spores of the two forms differ little from each other.” GYMNOSPORANGIUM. 395 The strikingly characteristic cleavage of the wood by the overgrown elements of the medullary rays and the wood- parenchyma, in the case of the twig-inhabiting form, will be seen from the figures (Fig. 229). As already noticed, the sporogenous cushions are generally formed on one side. After Pier * ———— ec OrlC b Fic, 229.—Two sections from a swelling on a Juniper-branch. «a, From the middle of the swelling ; the rind under the spore-cushion is much thickened, and the wood is much broken up by tracts of parenchyma. 6, Section from 2 c.m. under a; abnormal development of parenchyma in the wood has begun in the outer year-rings. (After Woernle.) a > aL, an eel ew the shedding of the cushion, a corky layer arises in the paren- chyma underneath it, and so a bark-scale is produced. Gymnosporangium sabinae (Dicks.). (Britain.) The mycel- ium hibernates chiefly in Juniperus Sabina (Savin), and induces swellings on the twigs. It also occurs on Jun. Oxycedrus,' . 'T found this host-species near Fiume. 396 UREDINEAE. Jun. virginiana, and Jun. phoenicea. (A reported occurrence on Pinus halepensis is probably an error.) The sporogenous cushions are little dark-brown protuberances which break forth in spring from swellings, or on green Fic. 230.—Gymnosporangium sabinae on twigs of Juniperus Sabina, at the time of liberation of spores. (v. Tubeuf phot.) twigs and scale-leaves. These bodies absorb water, swell, and run together, forming transparent gelatinous masses (Figs. 250 and 251). The teleutospores resemble those of G. jwniperinum, but have only four germ-pores; they germinate on the gelatinous masses, and produce promycela and sporidia. The latter germinate at once, chiefly on leaves of Pyrus com- munis. The pyenidia are produced on the upper epidermis as sticky yellow spots bearing darker dot-like pyenidia. The aecidia (Roestelia cancellata) are found in September on the under-surface of the leaves of pear, also on leaf-petioles, young shoots, and even on the fruits. The peridia differ from both GYMNOSPORANGIUM. 397 the species already described in remaining closed at the apex, the spores escaping through trellis-like slits on the lateral walls of the peridia (Fig. 254). This fungus will not germinate on apple-trees, but on pears every leaf may be thickly covered with aecidia and pycnidia, yoo and considerable damage to the crop thereby ensue (Fig. 235). Fic. 232.—Teleutospores of G. sabinae. Fic, 231.—Longitudinal section through The elongated thin-walled ones are a conical teleutospore-sorus of Gymno- lighter in colour than the thick-walled. sporangium Sabinae. (After Woernle.) (After Woernle.) Infections on various hosts with Gymnosporangium sabinae from Juniperus Sabina gave: DoD On Host-plant. Spore-form. Pyrus communis, - - Crataegus Oxyacantha, Mespilus germanica, - Pyrus communis, - - Rostelia cancellata, - Pyrus communis, - : » Michauxii, - » tomentosa, - Pyrus communis, : . zy “ Pyrus communis, . = Bs - Crataegus Oxyacantha, : a + | Basa 4. ; | Authority. Plowright. Oerstedand De Bary. Rathay, Tubeuf, ete. Reess. E. Fischer. Klebahn. = (uncertain). 398 UREDINEAE. The anatomical changes exhibited in diseased parts of pear- leaves have been briefly described by Fentzling." He found a radial elongation of the cells of the spongy parenchyma and an accompanying accumulation of starch. Wakker, about the same time, obtained similar results in the case of Crataegus Oxyacantha deformed by G. clavariaeforme (see p. 387). Wakker observed a diminished formation of calcium oxalate; Fentzling, however, found increased deposit of the same salt, not only in the form of Fic. 233.—Gymnosporangium sabinae in the form of Roestelia cancellata on leaves of Pear. A few twigs showing the abundance of aecidia over the whole tree. (v. Tubeuf phot.) isolated crystals but as masses. Cork-formation was suspended in Wakker’s case, while in Fentzling’s a partial formation of cork was distinguishable beneath the epidermis. The increased thickness of diseased leaves is due principally to multiplication of the spongy parenchyma, the upper layers of which frequently become more or less palisade-hke. When pycnidia (spermogonia) are formed on the upper leaf-surface, the palisade parenchyma 1Fentzling (doc. cit.) and Peglion (Rivista di Patologia Vegetale, 11.), also describe these alterations. 2 GYMNOSPORANGIUM. 399 of the spot in question is either completely destroyed or trans- formed into irregular cells, separated by intercellular spaces. The anatomical changes in swellings (Fig. 235) induced by 7, sibinae on Juniperus Sabina were investigated by Woernle with the following results. Wood, bast, and rind are increased round the whole circumference of the stem. Along with the broadening of the year-rings, however, there occurs a change in the structure of the diseased wood. The same Fic. 234.—A few leaves enlarged from Fig. 233. The leaf to left hand bears pycnidia on red spots on the upper surface of the leaf; the remaining leaves bear aecidia on raised portions of their surface. Several aecidia still further enlarged show the peridia dehiscing by longitudinal slits. (v. Tubeuf del.) tissues occur in the year-rings as already described for 4G. clavariaeforme, viz. thickened twisted tracheids, loosely connected together and with fissure-like pits; medullary rays more numerous and broader; the limits of the year-ring difficult to distinguish ; and a yellow pigment deposited in the walls of all the elements. A tissue of this nature may be found round the whole circumference of a twig even in the first year after infection, and regularly each succeeding year. Woernle only rarely found zones of irregular cell-formation like those 400 UREDINEAE, characteristic of G. clavariaeforme. No myceliuin occurred in the wood. A comparison of normal bast with that of infected twigs revealed changes similar to those already described for G. clavariaeforme. In addition, it is to be noted that the thickened bast-fibres no longer occurred in closed masses, but were often completely absent in the first year after infection, while in all diseased twigs every intermediate stage exists between thin-walled bast-elements and thick-walled bast-fibres, such as never occur in the normal twigs; in fact, many twigs had thin-walled elements only. Fic. 235.—Swelling from Fic. 236.—Sections of a twig of Savin attacked by G. sabinac. a branch of Juniperus a, At thickest part of the swelling ; }, 3¢.m. under a and normal. Sabina attacked by Gymno- In « is shown one of the hollow teleutospore-cushions; five sporangium sabinae. Dia- cushion-scars; in the second-year ring are two shaded zones meter at thinnest part 17 of wood, chiefly composed of parenchyma. (x 25.) (After centimetre, at the thickest Woernle.) 5 6¢c.m. (v. Tubeuf phot.) The sporogenous cushions of (, sabinae are formed in quite a different manner from those of G. clavariaeforme. Beneath each cushion the bast increases very rapidly and forms an out- erowth, which is still further enlarged by the addition to its apex of six or seven rows of radially arranged cells, rounder and smaller than the bast-cells of the cushion. The mycelium penetrates between these outer cells, and forms over the whole cellular outgrowth a pseudoparenchyma from which the sporo- genous tissue arises. GYMNOSPORANGIUM. 401 A sharply detined roundish scar of a light-yellow colour remains after the spores are cast. This is composed of a superficial layer of coloured pseudoparenchyma, with an under- lying scar-tissue of characteristic constitution. The latter con- sists of several layers of cork-cells extending from one edge of the scar to the other, separating the cushion from the twig- tissues. This scar-tissue is not broken through next year, but the new sporogenous cushions break out through other parts of the bark (Fig. 236). G. confusum Plowright. (Britain.) This is found on Juniperus Sabina along with G. sabinae, from which it is difficult to distinguish. Pyenidia and aecidia are produced generally on Crataegus Oxyacantha aud Cydonia vulgaris, rarely on Pyrus communis. The aecidia on Crataegus resemble those of G. clavariaeforme on the same host, and dehisce by the ruptured apex of the peridium. Those produced on Pyrus communis are distinguished? from aecidia of G. sabinae on the same host by dehiscing through the open apex of the peridiuim. Infections of Gym. confusum from Juniperus communis gave the following results: On Host-plant. Authority. Pea wadgoris, : ; { pycnidia and aecidia with | ee | tubular peridia, - -{ Crataegus Oxyacantha, - as * es * Pyrus communis, . - a o Fa Y. Crataegus Oxyacantha, - 3 a “ Plowright. The following American species of Gymnosporangium have been described :* On Arborvitae or white cedar: G. biseptatum Ellis. On twigs and needles of Chamaceyparis thyoides and Libocedrus decurrens. The aecidia on Crataegus tomentosa and Amelanchier canadensis. ' Plowright, Linnean Soc. Journal (Botany), 1887. E. Fischer, Zeitschrift /. Pflanzenkrankheiten, 1., 1891; with summary of literature. Klebahn, Fors?/, naturwiss, Zeitschrift, W., 1893. °K. Fischer (/oe, cit.). *Farlow, The Development of the Gymnosporanyia of the United States, 1886, and other papers. Thaxter in various papers on Gymnosporangia, 1886 to 1891. Halsted (Report on Vegetable Pathology for 1888, U.S. Dept. of Agri culture) gives a résumé, with description and figures of G. macropus and treatment for orchard-rust. Fischer, Zeitschrift f. Pllanzenkrankheiten, 1., 1891. I 402 UREDINEAE. G. Ellisii Berk. On Chamaccyparis thyoides, The aecidial stage on Pyrus Malus and P. arbutifolia. On red cedar (Juniperus virginiana) : G. macropus Lk. The aecidia and pyenidia occur on Pyrus Malus, P. coronaria, P. arbutifolia, Crataegus tomentosa, C. Douglasvi, and Amelanchier canadensis; they are known as Fic. 237.—Spores of Fic. 238.—Spores_ of Fic. 239.—Spores of Gym. biseptatum, (After Gym. — Ellisii. (After Gym. macropus. (After Woernle.) Woernle.) Woernle.) foestelia pirata. This is one of the commonest causes of apple-rust and of the deformation known as “ cedar apples” (Fig. 240). The anatomy of the latter structures has been described by Sanford.! G. clavipes Cooke et Peck, occurs on Juniperus communis, Its aecidia and pycnidia are found on Pyrus Malus, P. arbuti- folia, and Amelanchier canadensis. 1 Sanford, Annals of Botany, 1., 1887. GYMNOSPORANGIUM. 403 G. globosum Farl. Aecidia on Pyrus Malus, P. communis, Cydonia vulgaris, Sorbus americana, and species of Cratuegus. G. nidus-avis Thaxt. Aecidia and pycnidia on Pyrus Malus, Amelanchier canadensis, and Cydonia vulgaris. On the red cedar it causes the “bird’s nest” deformation of the branch-system. G. speciosum Peck. On Juniperus occidentalis. G. Cunninghamianum Barcel. On Cypressus torulosa in the Himalaya. Aecidia on Pyrus Pashia. The following genera do not occur in Europe. Coleopuccinia, Ravenelia, Alveolaria, Trichospora. Ravenelia alone amongst these contains parasitic species of import- auce, ‘They all occur on Leguminosae py Gynncernungiune’ mawonen and Euphorbiaceae in the warmer parts =“ Tent fel) of India, Africa, and America. Ravenelia Volkensii Henn. has teleutospore-sori which appear on “ witches’ broom” deformations of the twigs of an Acacia in Usambara. Rav. pymaea Lager. et Diet. produces its teleutospores on malformed branches of Phyllanthus in Ecuador. Certain forms of Aecidium which cause deformation of species of Acacia should probably be included in this genus (see p. 410). Endophyllum. Teleutospores originate serially on cushions which are enclosed in a peridium similar to aecidia; on germination, a four-celled promycelium is produeed.2 Leaves of Euphorbia, Sedum, or Sempervivum inhabited by mycelium develop abnormally. Endophyllum euphorbiae-silvaticae (1). ©.) (Britain). Accord- ing to Winter, the peridia are regularly distributed over the underside of the leaf of Euphorbia amygdaloides; they have white fissured margins either erect or somewhat turned back. Dietel, ‘* The Genus Ravenelia,” Hedwigia, 1894. * The teleutospores of this genus might be described as aecidiospores which produce promycelia. 404 UREDINEAE. Spores yellow and polygonal. Leaves when attacked remain broad, short, and pale coloured. E. sempervivi (Alb. et Schw.)' (Britain). The aecidium-like patches of teleutospores occur on wild and cultivated species of Sedum and EHscheveria. The spores produce promycelia from which arise sporidia which germinate on the same host-plant. True aecidia are unknown, but orange-red pycnidia (spermo- gonia) may occur. Leaves of attacked plants are pale and abnormally lengthened.” E. sedi (D. C.). Teleutospores occur on species of Sedwin. The genus Pucciniosiva found in Ecuador contains few species, and none of them important parasites. Aecidium-Forms The relationships of which are uncertain. Aecidium elatinum Alb. et Schw. (Britain and U.S. America). The witches’ broom of the silver fir® This Aecidiwm is widely distributed in forests containing silver fir (Abies pectinata), and produces canker of the stem frequently accompanied by that deformation of the branch system known as a_ witches’ broom. In Germany it has also been observed on Abies Nordimanniana, A. cephalonica, A. Pinsapo; in North America on A. balsamea ; and in Siberia on A. Pichia. As a result of the presence of this fungus, globose or barrel- shaped swellings make their appearance on stems and branches of all ages and on all parts of the trees. A single stem may carry one or many of these, and they continue to increase with its growth. If, as is frequently the case, the bark covering the swelling becomes ruptured and partially detached, then the wood left uncovered becomes a wound, and falls an easy 1Leveillé, Bullet. Science. Natur., Xvi., 1825. * Tilustrated in Kerner’s Natural History of Plants, English Edition (Fig. 358). °De Bary, Botan. Zeitung, 1867. Weise, ‘‘Zur Kenntniss d. Weisstannen- krebses,” Miindener Forstliche Hefte, 1891. Heck., ‘* Der Weisstannenkrebs.” Springer, Berlin, 1894; with Illustrations and Bibliography. The canker is common throughout Britain, but witches’ brooms have not been often recorded, (Edit. )} AECIDIUM-FORMS. 405 prey to wood-destroying fungi.’ The presence of such rotting spots renders the tree liable to break over in their neighbour- hood, while they, as well as the swellings on the trunks, cause a considerable depreciation in the value of the timber. The malformations of the branch-system known as witches’ brooms are frequently induced by this fungus. They occur as a rule on the horizontal branches and form a richly branched bush easily distinguished, even at a distance, by a marked Fic. 241.— Witches’ Broom of Silver Fir (winter condition), The needles, with spores of Aecidium elatinum, have fallen off, but the normal foliage remains. (vy. Tubeuf phot.) negative geotropy of its twigs. The brooms not untrequently start from a marked basal swelling. They may be found of all sizes, on young as well as old trees, on any part of the branch- system, and in all localities where the fir occurs (Figs. 241 and 242), The aecidia of Aecidium elatinum are developed only on the deformed needles of the witches’ brooms. These needles are produced anew each spring, live only one season and are cast ' Polyporus Hartigii and Agaricus adiposus in particular accompany this canker and bring about decay of the wood. 406 UREDINEAE. the same autumn; they are small, one-pointed, and pale from an almost complete lack of chlorophyll. In these respects they are quite distinct from the larger double-pointed normal needles with their dark-green colour and a period of growth extending over several years. All the needles on a broom are as a rule stunted in the manner described, yet single branches may be found with needles quite normal; such contain no mycelium, or, if so, it has found its way in too late to have any effect on their growth. Fig. 242.— Witches’ Broom of Silver Fir (summer condition). The markedly negative geotropic broom has its origin in a distinct basal swelling. (v. Tubeuf phot.) The various tissues of the witches’ brooms also undergo considerable modification as compared with normal twigs. A thicker and softer bark is present, due to the parenchymatous cells of both outer rind and bast having enlarged in size and increased in number; the cork layers are also abnormally in- creased. The same changes may be observed in the rind of the swellings, and to this their increased size must be chiefly ascribed. The wood both in twigs and swellings is much increased; the year-rings however are very variable, sometimes they are broader than the normal, again they may be diminished or even altogether wanting; where however the wood decreases, there the bast increases in proportion. This lack of uniformity AECIDIUM-FORMS. 407 in the growth of the wood disturbs the elements, so that they are irregularly developed and more or less twisted.' A mycelium inhabits the tissues of abnormal twigs and cankered swellings. It grows in the intercellular spaces of the rind, between the bast cells and outer parts of the wood, and derives nutriment by means of haustoria; these either bore through the cell-walls, or only press closely against them so as to cause depressions. Spore-formation takes place on the needles of the witches’ brooms. The pyenidia (spermogonia) are produced on the upper side beneath the cuticle and emerge through it as little yellow points. The conidia (sper- matia) are tiny globose colourless bodies. The aecidia come later during June and July in irregular rows on the under side of the leaf. Their peridia break out as low dome-like structures, the apices of which rupture irregu- larly to allow escape of aecidio- spores. In spite of numerous infections, De Bary was unable to observe the penetration of a germ-tube into needles or twigs of silver fir. Weise believes that infection of the fir takes place on twigs which have just emerged from the bud. As a preventive measure, all Fig, 243.—Aecidium atrobilinum on a Spruce witches’ brooms should be cut off cone. (Vv. Tubeuf phot.) before spore-formation begins, and stems with canker-wounds should be removed during forest-thinning. For further details the monograph of Heck may be consulted. Aecidium strobilinum (Alb. et Schw.)® (Britain), Spruce- 'Note.—Further details of the anatomical changes induced in the tissues of these witches’ brooms may be obtained in the German edition of this work (pp. 420-421), or in the original thesis by Hartmann, (Anatom. Vergleichung d. Hexenbesen der Weisstanne. Inaugural Dissertation, 1892.) (Edit.) * Reess, Rostpilxformen d, Coniferen., 1869, Oecerstedt, Naturh. for Vidensk Medd., 1863, 1. 408 UREDINEAE. cone rust. This disease is found on the cones of spruce. The aecidia are brown somewhat flattened spheres, and appear in large numbers on cones distinguished by their scales standing stiffly open even in damp weather (Fig. 243). The germ- tubes of the fungus find entrance in spring into the flowers or young spruce-cones, and the mycelium lives parasitic in the green scales without causing any marked change in their growth, although the ovules are more or less injured. No mycelium has ever been found in the lower cone-axis, nor in the shoots, so that the disease must be the result of in- fection by spores only. 2 3 Fic. 244.—Accidium strobilinuim. 1, Cone-scale of Spruce with aecidia, those to left dehiscing their yellow spores, those to right still closed. (v. Tubeuf del.) 2, Section through an immature aecidium. 3, Part of 2 enlarged—pev, peridium ; sp, Spores; zw, intermediate cells; spti, sporophores; m, mycelium; par, the scale-parenchyma. (After Reess.) The aecidia break out on the inner (rarely the outer) side of the bases of the cone-scales; each is enclosed in a firm brown lignified peridium, which ruptures by a_ cross-fissure and becomes an open disc. The young spores are joined by small intermediate cells, which are gradually absorbed to form a layer of gelatinous lamellae on the _ spore-coats (Fig. 244). Teleutospores of this Accidiwm are wnknown. AECIDIUM-FORMS. 409 Aecidium pseudocolumnare Kiilin.' Occurs on needles of Abies pectinata in Germany; in Britain, however, on this and several other species of Abies. It is distinguished by its large white spores from the Ace. columnare of Calyptospora (p. 372). Aec. Magelhaenicum Berk. ‘This species occurs on various species of barberry. The mycelium hibernates in the shoot- buds and causes them to develop as witches’ brooms, bearing on the lower surface of their leaves aecidia with long, white, sac-like peridia. The allied teleutospore-form is as yet unknown. Aec. clematidis D. C. (Britain and U.S. America). On Clematis Vitalba, (. recta, and other species. It is related to Puccinia agropyri Ell. et Ev.* Aec. Englerianum Henn. et Lind.’ produces a_ peculiar antler-like branching of the twigs and leaves of a Clematis at Eritrea (Lytri) in the Grecian Archipelago. Aec. punctatum Pers. (lec. guadrifidum D.C.) (Britain and U.S. America). This is a common species on Anemone (Fig. 190) and Eranthis. The aecidia have white peridia, which on dehiscence break into four lobes. Aec. leucospermum D.C. (Britain and U.S. America). On Anemone WeMOTrOSa (Fig. 190). Aec. hepaticae Beck. On Anemone Hepatica. Aec. ranunculacearum D. C. (Britain and U.S. America). On species of Ranunculus. A collective name for aecidia of several species of Uromyces (p. 336), and Puccinia (p. 349). Aec. aquilegiae Pers. (Britain and U.S. America) On Aguilegia vulgaris and other species. (See Puceinia agrostidis, p. 349.) Aec. actaeae (Opiz.). On leaves of Actaea spicata in Europe and America. Aec. barbareae D.C. On species of Barbarea (Britain). (See Puce. festucae, p. 349.) Aec. circaeae Ces. On species of Cireaea. Aec. grossulariae Schum. (Britain and U.S. America). On Libes Grossu- laria and R. rubrum. Klebaln believes it is related to a Pucetnia on Carer. Aec. bunii D.C. On Conopodium denudatum in Britain, (See Puce. bistortae, p. 355.) Aec. periclymeni Schum. On species of Lonicera. (Britain.) (See Puce. festucae, p. 349.) Aec. compositarum. A provisional species-name for a large number of aecidia frequenting Compositae, and by no means resembling each other. Aec. leucanthemi D.C. A European species with its Pucernia-form on Carex montana. Aec. cyani D.C. On Centaurea Cyanus. Aec. ligustri Strauss. On Privet. ' Hedwigia, 1884. *Dietel, Ocsterreich botan, Zeitung, 1892. *Engler’s Botan. Jahrbuch, 1893. 410 UREDINEAE. Aec. phillyreae D.C. On species of Phillyrea (Britain ?). Aec. fraxini Schwein. This causes serious damage in America to the foliage of Fravinus viridis and Fr. americana.’ It has also appeared in Europe on the latter species introduced from America. Aec. nymphaeoidis D. C. On leaves of Limnanthemum, Nuphar, and Nymphaea. (Britain.) Aec. pedicularis Lib. On Pedicularis. (Britain.) (See Puec. paludosa, p. 351.) Aec. prunellae Wint. On Prunella vulgaris. (Britain.) Aec. euphorbiae Gmel. is found on many species of Euphorbia, It is probably the Aecidium-form of Uromyces pisi. (Britain and U.S. America.) Aec. convallariae Schum. (Britain and U.S. America). Probably a provisional species-name for aecidial forms found on Convallaria, Polygon- atum, Paris, Lilium, etc. (See under Puccinia.) Aec. ari Desm. (dec. dracontii Schwein.) is found on species of Arwm. (Britain and U.S. America.) (See Pucc. phalaridis, p. 349.) The following species are found on Acacia and seem to have strong affinity with the genus Lavenalia: Aec. esculentum Barcl. produces deformation of twigs of Acacia eburnea in India. Twigs of this kind, likewise shoots deformed by dee. wrticae var. himalayense Barcl., and pine-shoots deformed by certain species of Perider- mium, are eaten in various parts of the world. Aec. acaciae (Henn.) on Acacia etbaica in Abyssinia. This is said by Magnus to cause witches’ broom deformation. Aec. Schweinfurthii Henn. causes malformation of fruits of Acacia Fistula in Africa. Aec. ornamentale Kalch. causes curvature of shoots of Acacia horrida at the Cape. The following gre some of the more important species recorded for North America only : Aecidium dicentrae Trel. Leaves of Dicentra and Corydalis. Aec. monoicum Peck. Leaves of Arabis. Aec. drabae Tr. et Gall. Aec. lepidii Tr. et Gall. Aec. proserpinacae B. et ©. Aec. Mariae-Wilsoni Peck. | (), ppecies oP acl: Aec. Petersii B. et ©. J Aec. cerastii Wint. Aec. pteleae B. et C. On leaves of Ptelea trifoliata. Aec. xanthoxyli Peck. Aec. splendens Wint. In the cotyledons of Croton monanthoqynus. Aec. aesculi Ell. et Kell. Aec. psoraleae Peck, and Aec. onobrychidis Burr. On species of Psoralea. 1Pound, American Naturalist, 1888. — ia | AECIDIUM-FORMS. 411 Aec. Peckii De Toni and Aec. oenotherae Mont. On leaves of species of Oenothera. Aec. sambuci Schwein. On leaves and stems of Sambucus. Aec. ceanothi Ell. et Kell. Aec. abundans Peck. On species of Symphoricarpus. Aec. cephalanthi Seym. On Cephalanthus occidentale. Aec. erigeronatum Schwein. On many species of Erigeron. Aec. asterum Schwein. On species of Aster and Solidago. Aec. polemonii Peck. On Polemonivm and Phlov. Aec. apocyni Schwein. On leaves of Apocynuin. Aec. Jamesianum Peck, and Aec. Brandegei Peck. On leaves of species of Asclepias. Aec. myosotidis Burr. On leaves of MJyosotis rerna, ete. Aec. plantaginis Ces. On leaves of species of Plantago in Europe and America. Aec. pentastemonis Schwein. On species of Pentstemon. Aec. giliae Peck. Aec. lycopi Gerard. On leaves and stems of Lycopus europaeus. Aec. iridis Gerard. Aec. macrosporum Peck, and Aec. smilacis Schwein. On species of Smilar. Peridermium. Peridermium pini (Willd.)! is found on pine-trees in Europe, Britain, and United States. A teleutospore-stage of this has not as yet been identified, although a very similar species (Pe7ri- dermium Cornui Rostr. et Kleb.), also occurring on the bark of pines, has been proved to have as its teleutospore-form Cronartium asclepiadewm.* The mycelium of Peridermium pini lives intercellularly in the rind, bast, and wood of Pinus sylvestris, P. Laricio, P. halepensis, P. maritima, and P. montana. It lives and extends through the stem for years, attacking the living cells and absorbing nutriment from them by little haustoria. The cells of parenchymatous tissues are those most generally attacked, and the mycelium has been found to penetrate along the medullary rays to a depth of 10¢.m. into the wood-mass. The cells of attacked parts lose their normal content including starch, and secrete crude turpentine in such quantity as to completely permeate their walls, and even to form drops, In this way portions of the wood become completely saturated 'R. Hartig, Wichtige Krankheiten d. Waldbdiumem. *Klebahn, Berichte d. deutsch, hotan. Gesellschaft, 1890. 412 UREDINEAE. with resin, and as the same process goes on in bast and rind, the turpentine overflows from fissures or wounds in the bark. During the summer the mycelium grows amongst the dividing cambium-cells and kills them. Where this occurs the year- SN Fic. 246.—Peridermium pini (corticola). Young twig bearing numerous aecidia. (v. Tubeuf phot.) Fic. 245.—Peridermivin pini (corticola). Branch and lateral twigs distinctly swollen where attacked. They also bear aecidia. (v. Tubeuf phot.) ring ceases to thicken, but as the mycelium seldom succeeds during the first year in killing the cambium all round a a; PERIDERMIUM. 413 branch, the living portions of the ring grow on with increased vigour, and even attempt to close over the injured portion. This irregular growth, continued in many cases for years, pro- duces abnormal cross-sections (Fig. 248). The mycelium grows out centrifugally from diseased spots, so that the wounds continue to enlarge, and the disease becomes easily noticeable on account of the deep channels and distorted swellings on the pine branches and stems. As the disease spreads inwards into the stems, the conduction of water is interfered with and the branches above such wounds dry up and die off. Whereas a Vas A ere. So - 42 Biola 0.) (RAO) OPAC? ) SSow: fegly; i KR ( Fic. 247.—Peridermium pini (corticola). «, a, Mycelial stroma developed in the eee Ach metic, Becicts composed! of whoch emallec oells than in the needle-inhabiting aecidia. p, The peridium. (After R. Hartig.) young plants soon succumb to attack, the struggle with old trees may go on for years. Fresh infection of older stems oceurs generally in the higher parts of the tree, where the bark is still thin. Pyenidia (spermogonia) are developed between the rind- parenchyma (periderm) and cork, generally towards the margin of diseased spots. The conidia emerge from the ruptured cork-layers of the bark as a honey-sweet liquid. H. Mayr states that this liquid is given off in such quantity from species of Peridermivm in Japan, that it is collected and eaten by the natives. 414 UREDINEAE. The aecidia appear in June as wrinkled yellow sacs emerging from the bark of swellings. They continue to develop in succession for years on the living parts of attacked branches, but according to Hartig they cease to make their appearance on old stems, even when a mycelium is_ present. This disease is the cause of great damage to pines, especially where planted as pure forest. One case is recorded’ of a forest near Kohlfurt where 90 per cent. of the trees in an old plantation were “stag-headed” on account of a deficient Fic. 248.—Peridermium pini. Section through a diseased stem of Pine showing the gradual killing of the cambium by the fungus. (v. Tubeuf phot.) supply of water in the crown accompanying attacks of this fungus. Until more is known of its life-history, preventive measures cannot be well extended beyond cutting down infected trees. The following species of Peridermium have been observed on species of Pinus: A, On the needles: Peridermium oblongisporium Fuck. (now Coleosporium senecionis) on Pinus sylvestris and P. austriaca (p. 374). P. Klebahni, P. Soraueri, P. Stahlii, P. Plowrightii, and P. Fischeri. On Pinus sylvestris ; related to various species of Coleospordum. P. piriforme Peck. On Pinus speciosa in U.S. America. P. cerebrum Peck. On Pinus rigida in North America. ! Marker at Schlesien. Forstverein, 1893. PERIDERMIUM. 415 P. filamentosum Peck. On Pinus ponderosa, also in America. P. Harknessii Moore. On Pinus ponderosa, P. insignis, P. Sabineana, and P. contorta in California. Fic. 249.— Peridermium giganteum on Pinus Thunbergii from Japan. (vy. Tuber 9 ner, I : i phot.—the specimen presented by Prof. Grasmann of Tokio.) P. brevius Barcel. On Pinus excelsa in India. P. complanatum Barcel. On Pinus longifolia in India; on rind as well as needles. B. On the rind or bark: Peridermium Cornui Rostr. et Kleb. (now Cronartium asclepiadeum, p. 381). On Pinus sylvestris. P. strobi Kleb. (now Cro- nartium ribicola, p. 382). On Pinus Strobus, P. Lambertiana, (and P. Cembra). P, pini( Willd.). On Pinus sylvestris. (Britain and U.S. America. ) P. orientale Cooke. On Pinus rigida and P. virgine- ana in America; also PP. : ' Fic. 250.—DPeridermium giaanté longifolia in India. from Japan. (v. Tube 416 UREDINEAE. P. Ravenelii Thiim. On Pinus australis in North America (probably a variety of P. oblongisporium). P. deformans Mayr. On Pinus mitis in America. P. giganteum (Mayr). On Pinus densiflora and P. Thunbergii in Japan. This causes very conspicuous deformation of its host (Figs. 249 and 250). P. complanatum Barcl. On Pinus longifolia in India. The following species frequent other hosts: Peridermium conorum Thiim.' This aecidium first found by De Bary in Thiiringia, has recently been reported in Denmark, Russia, and America; also in Upper Bavaria by v. Tubeuf in September, 1895.. It takes the form of two large aecidia, which make their appearance on the outer or inner side of the cone-seales of spruce. The white peridia break through the epidermal tissues which then remain as a_ brownish sheath around each ruptured peridium (Fig. 251). The spores are separated by inter- mediate cells, and their outer coats are studded with poly- gonal warts. The cone-scales bearing aecidia contain a very large quantity of starch. Tel- eutospores of the species are unknown. Ba 2 ete TEs CONTI EO ac Peridermium coruscans dium; sp, spore; zw, intermediate cells; spti’, sporophore; m, mycelium; pros, prosenchyma. (after eee) Fries.2, The mycelium of this fungus seems to perennate in twigs and buds of spruce. Twigs unfold from the bud as deformed, shortened, cone-like shoots bearing very short broad needles of a pale colour. The aecidia are produced on the deformed needles as broad lineal cushions with white peridia. They originate under the epidermis which they rupture, and break out on one side of the needle. lReess, Rostpilzformen, 1869. 2 Rostrup, Vidensk. Gelsk. Forhandl., 1884. PERIDERMIUM. 417 The soft hypertrophied shoots are eaten. They occur chiefly in Scandinavia, but recently were observed by Gobi and Tranzschel in the neighbourhood of St. Petersburg.! Fic. 252.—Aecidium coruscans on malformed shoots of Spruce. The compact abnormal shoots thickly covered with white aecidia contrast strongly with the normal portions. (v. Tubeuf phot. from material presented by Prof. Fries, Upsala.) P. Engelmanni ‘Thiim. On cones of Picea Smithiana. (U.S. America. ) P. piceae Barcl. On needles of Picea Smithiana. P. Peckii Thiim. On needles of Tsuga canadensis (U.S. America). P. balsameum Peck. On needles of Abies balsamea (U.S. America). P. ephedrae Cooke. On Ephedra in U.S. America. P. cedri Barcl. On needles of Cedrus Deodara in India. P. Balansae Corn. On leaves of Dammara ovata in New Caledonia. ‘Also reported at Haslemere (Britain), Grevillea, Xx1x., 1890. 2D 418 UREDINEAE. Caeoma. Caeoma abietis-pectinatae [Reess.1 The aecidiospores may be found on the lower surface of young needles of silver fir; the aecidia are yellow elongated cushions situated on either side of the needle mid-rib, and are without peridia. Pycnidia (spermogonia) are pro- duced before the aecidia. The mycelium is septate and intercellular with few haustoria. I have found the fungus fairly abundant Fig. 253.—Cacoma «abictis : pectinatae. Needle of Silver On the Alps and in the Danube valley near Fir showing Cacoma-patches >) on the lower surface. (v. Passau. Teleutospores are unknown. Tubeuf del.) Caeoma deformans (Berk. et Br.) Tubeuf (Uromyces deformans Berk, et Br. or Caecoma Asanuro Shirai).* This induces the formation of “ witches’ brooms” or of antler-lke Fic. 254.—Cacoma dejormans on Thuiopsis dolabrata. (v. Tubeuf phot. from dried material presented by Prof. Grasmann of Tokio.) 1 Reess, Rostpilzformen, 1869. 2Berkeley, ‘‘The fungi collected during the expedition of H.M.S. ‘Chal- lenger.’” Jour. of Linnean Soc., Xvi., 1876. 3 Shirai, Botanical Magazine, Tokio, 1889. CAEOMA. 419 leatiess shoots on 7hujopsis dolabrata in Japan, whence they were sent to me (Figs. 254 and 255). One example (not figured) was as large as a young child’s head. The shoots of the witches’ brooms are furnished with vascular bundles and possess a parenchyma rich in starch-content. Each branch of the deformed shoot termi- nates in a hemispherical saucer-shaped cacoma-cushion, at first covered over by the epidermis, but with no peridium. The caeoma-dises are at tirst brown, but after the epidermis bursts and rolls back, the yellow dusty spores appear. The spores Fic. 255.—Caeoma defor- mans. Portion of the pre- arise serially from very short basidia; they ceding figure enlarged to show the Caeoma-discs on the are yellow and have striped membranes. Bree vee 6 or ee The witches’ brooms also exhibit marked hypertrophy (Fig. 254). In the supporting branch both wood and bark are considerably increased. Large medullary rays occur in the wood, and nests of thin-walled parenchyma are interpolated between the regular tracts of tracheae, so that the general arrangement resembles that shown in juniper by Wornle’s researches on (fymnosporangium. The parenchymatous groups of cells in the wood appear to the naked eye as brown spots. They are permeated by a vigorous intercellular mycelium, which sends off large haustoria into the adjacent cells. Caeoma Iaricis (Westend).! On needles of Larix. (Britain.) C. orchidis A. et S. On orchids. (Britain.) C. chelidonii Magn. On Chelidonium majus (U.S. America). C. fumariae Lk. On Corydalis. C. euonymi (Gmel.). On HLvonymus europaeus (Britain). C. confluens (Pers.). On Ribes alpinum, R. rubrum, ete. C. nitens (C. /uminatum) is the well-known Blackberry-rust so common in the United States. It is probably a form of Pueccinia Peckiana.* C. aegopodii (Rebent.). On Aegopodium Podagraria and Chaerophyllum aromaticum. C. ligustri (Rabh.). On Ligustrum vulgare. C. ari-italici (Duby). On Arum maculatum. C,. alliorum Link. On Allium ursinum, A. oleraceum, ete.* C. saxifragae Strauss. On Sarifraga granulataS C. mercurialis (Mart). On Mereurialis perennis.® ' This and most of the other species are only stages of some Me/ampsora. *Clinton, Botanical Gazette, 1895, p. 116. * These three species are given as British in Plowright’s ‘Uredineae.’ (Edit.) 420 UREDINEAE. Uredo-Forms of uncertain relationship. Uredo agrimoniae (D. C.). On species of Agrimonia (Britain and U.S. America). Dietel regards it as related to Melampsora (Thecospora) agri- monine. U. Muelleri Schroet. On Rubus fruticosus (Britain). U. symphyti D.C. On Symphytum officinale (Britain). U. phillyreae Cooke. On Phillyrea media (Britain). U. macrosora De Toni. On Epilobinimn tetragonum (U.S. America). U. vitis Thiim. This species first attracted notice as a disease- producing fungus in Jamaica in 1879, but it had been found previously in the United States. It causes spots on the upper surface of leaves.! U. fici Cact. On Ficus Carica in Italy and U.S. America. U. quercus (Brond.). On species of Quercus (Britain and U.S. America). U. iridis. On many species of /rzs (Britain). U. glumarum Rob. On Zea Mais in Belgium and England. U. sorghi Fuck. On Sorghum halepense in Greece; (compare with Uromyces and Puceinia on the same host.) U. gossypii Lager.” This has been observed in South America causing a rust on cotton-plants and injuring the yield of cotton. It appears as small purple-brown spots; the spores are oval and yellow. Uredinopsis.”* [This is a new genus found by Magnus to contain several Uredineae parasitic on Ferns* The aecidial stage is unknown. The uredospores are abjointed singly from the endg of sporo- genous hyphae; they are unicellular and without germ-pores. The uredospore-sori are enclosed in a pseudoperidium of elon- gated tubular cells. Unicellular teleutospores (7) are given off from sori similarly to the uredospores. Pluricellular teleuto- spores are developed from the mycelium im the intercellular spaces of the host-plant, never from crust-lke sori. On germination four-celled promycelia with spherical sporidia are produced. Uredinopsis filicina (Niessl.) Magn. On lower surface of fronds of Phegopteris (Polypodium) vulgaris, causing death. Ur. struthiopteridis Stoermer. On sterile fronds of Struthiopteris germanica. Ur. pteridis Diet. et Holw. On Pteris aquilina.| (Kdit.) 1Massee (Grevil/ea XXL, p. 119) states this species to be identical with U. Vialae of Lagerheim (Revue gen. de Botanique, 1890). *Lagerheim, Journal of Mycology, vit. p. 48. *Dietel, ‘‘ Uredo polypodii (Pers.)”’ Oesterreich. hotan. Zeitschrift, 1894; also ‘Der Gattung Uvredinopsis,” Ber. d. deutsch. botan. Gles., 1895, p. 326. +These host-plants do not come strictly within the scope of this work, but a short note on the genus is necessary. (Edit.) BASIDIOMYCETES. 431 BASIDIOMYCETES. The sporophores, known as basidia, are structures with a definite shape, and with lateral branches, the sterigmata, from which a definite number of exospores—basidiospores—are ab- jointed, the basidia then becoming functionless. Basidia and basidiospores are characteristic of all Basidiomycetes, conidia and chlamydospores being produced only exceptionally. The basidia generally arise from an extended layer—the hymenium—which in the higher genera forms part of a con- spicuous complex sporophore. The basidia do not therefore originate from the germination of a spore, as do the promycelia of the Uredineae and Ustilagineae, but from special sporophores (rarely from the mycelium itself), whose surface they occupy, or in which they are enclosed. In the course of development, two nuclei have been found to copulate in the basidial cells. Thereafter they divide and produce four (rarely two) new nuclei (Autobasidiomycetes), or after the division of nuclei, cross-septa are formed, thus making the basidia pluricellular (Protobasidiomycetes). In both cases the nucleus passes through the sterigmata into the developing basidiospores, and on the germination of these spores, it divides into two nuclei, the starting points for further nuclear division. As just indicated two divisions of the group may be dis- tinguished: (1) Protobasidiomycetes, (2) Autobasidiomycetes. PROTOBASIDIOMYCETES. Under this class are included the Auricularieac, Pilacreae, and Tremellinae, the first two possessing basidia divided, as a rule, by cross-septa into four cells, the last with basidia also divided into four cells, which are formed, however, by two longitudinal walls set at right angles to each other. A sterigma grows out from each cell and produces a single spore, after which the basidium dies away. The basidia of the Pi/acreae are produced inside closed sporocarps (angiocarpous), those of the other two groups are exposed (gymnocarpous). Parasites are unknown amongst the Protobasidiomycetes. AUTOBASIDIOMYCETES. Basidia unicellular (autobasidia), the sterigmata formed on the apex of the basidium, and each giving off a single basidio- 422 BASIDIOMYCETES, spore. The basidia originate from basidial layers or from complex hymenia, produced either inside some special structure, or on the surface of special sporophores, or on some definite part of these. The group may be sub-divided into the Dacryomycetes, Hymeno- mycetes, and Gasteromycetes (ncluding Phalloideae). Of these only the Hymenomycetes contain species parasitic on plants, the others include harmless saprophytes, which live in the soil, some of them, however, taking part in the formation of mycorhiza. THE HYMENOMYCETES. The unicellular basidia give off from their apices four (any number from 2 to 6 may occur) sterigmata, from each of which a single basidiospore is abjointed. The basidia arise from free exposed hymenia, which generally occupy the whole or part of large compound sporophores. The greatest develop- ment of the sporophore is attained in the umbrellas of the Agaricineae, and the large dises of the Polyporeae. It is only amongst the lowest genera, like Hxobasidium, that the basidial layers are produced directly on the organs of the host, and the basidia arise directly from the hyphae. teproductive cells, other than basidiospores, are rare. In a few cases amongst the Polyporeae, Brefeld and others have observed conidia and chlamydospores (Oidia, ete.); while some - few Agaricineae have the latter form of spore, but never conidia. The mycelium is of a very varied nature. It frequently inhabits wood, and in many different ways brings about destruction of henified tissues. Other modifications are seen in the forms of mycelium known as rhizomorphs, rhizoctonia, mycorhiza, and other closely felted masses of various shapes, which will be considered in detail as occasion requires. The formation of clamp-connections is also a special feature of the mycelium of the Hymenomycetes. In many cases the mycelium retains its vitality and perennates for several years. The genus Hxobasidium consists of parasites which produce inalformation of their host; many of the Polyporeae and Agaricineae are deadly enemies of forest and fruit-garden, while as wound-parasites many of them are specially dangerous. The general means of combating them consist in cutting out HYMENOMYCETES. 4323 any sporophores and applying tar to the wound, while diseased stems in the forest should be felled. Immediate artiticial closure of wounds in the wood is a very effective preventive measure.’ The Hymenomycetes are divided into Yomentelleae, Exobasi- diaceae, Hypochnaceae (included by Brefeld in the Tomentelleae), Thelephoreae, Clavaricae, Hydneae, Polyporeae, and Agaricineae. All contain parasitic species. EXOBASIDIACEAE. Exobasidium. The basidia are formed on the extremities of branches of the mycelium, which break out through the cuticle of attacked organs. The mycelium lives inside the host-plant, and induces considerable malformation. The basidia emerge on the surface of the host (similarly to the asci of the Hzoasci), and from each of the four sterigmata a single spore is given off. Exobasidium vaccinii Wor.’ (Britain and U.S. America). This is the cause of a very common and conspicuous deformation which affects the leaves, flowers, and shoots of Vaccinium Vitis- Idaea (Fig. 256). Leaves, where affected, become thickened and form irregular blisters vaulted towards the lower surface of the leaf, so that the lower epidermis covers the convex side and the upper epidermis lines the concavity. Chlorophyll is absent in the swollen tissues, but where blisters are exposed to direct light a bright red cell-sap is developed. Parts of the leaf adjoining diseased spots may remain normal and green. Flowers or their parts undergo similar malformation ; twigs become more or less thickened and twisted, their chloro- phyll disappears, and a reddish cell-sap is produced. On such diseased places spores are produced during the summer, after which the poorly developed tissues dry up and wither. When this fungus is present in the young tissues of its hosts, it exerts a very marked influence on their development. The palisade cells of the leaf become enlarged, while their chlorophyll almost wholly disappears, and is replaced by a red ‘Further details on this point have already been given, General part, p. 72. *Woronin, Verhand. d. naturfor. Ges., Freiburg, 1867; with 3 plates, Brefeld, Schimme/pilze, viit., 1889. Wakker, Pringsheim’s Jahrbuch, sve. 424 BASIDIOMYCETES. cell-sap. Cells of the parenchyma in flower and stem enlarge to a still greater degree. Intercellular spaces are as a rule obliterated, but when present are filled with a fine mycelium. Wakker gives us further results of the fungoid attack; crystal-glands, normally numerous, are no longer formed, but are replaced to some extent by indistinctly defined crystals of calcium oxalate. Transitory starch is stored up in large quantity. The fibro-vascular bundles Fig. 256.—Ezobasidium vaccinii inducing outgrowths on leaves of Vaccinium Vitis-Idaca. (v. Tubeuf phot.) present a striking modification, the primary xylem alone is normal, the vessels of the secondary wood remaining rudimentary; other parts are not lignified, and the phloem is only indistinctly laid down. A mycelium is present in all deformed parts, but absent in normal green tissue. It becomes massed to form a hymenial layer beneath the epidermal cells or between their outer walls and the cuticle. The sterigmata do not exceed four in number, and EXOBASIDIUM. 425 from each a spindle-shaped spore is abjointed (Fig. 257). The basidiospores divide in water by formation of cross-septa, and a germ-tube arises from each terminal cell. On a young leaf of Vaccinium the germ-tube penetrates and gives rise to a mycelium (Fig. 258); on other substrata the germ-tube sprouts into several very fine sterigmata, from the extremi- ties of which a series of conidia are abjointed; the conidia may give off secondary coni- dia, perhaps also tertiary. In nutritive solution, Brefeld ob- tained an increased number of re, ap aie ctinitd Dacciués ‘Whe bakidial germ-tubes and a continuous — lser ie shown deseloping from the intercellolar production of conidia; in air, conidia were produced on conidia, but inside the solution the conidia gave off hyphae from which new conidia arose. Fic, 258.— Evobasidium vaccinii, Germinating basidiospores. The septate spores have given off germ-tubes which penetrate into the cowberry leaves, either by stomata or through the epidermis. The lowest spore is forming conidia, (After Woronin.) This Hxobasidivm is very common on the cowberry (Vaccinium Vitis-Idaea) It oceurs less frequently on the bilberry (Vae- I : 'Several American Ericaceae are given as host-plants in the ‘* //ost-Index.” 426 BASIDIOMYCETES. cinium Myrtillus)' causing a premature fall of the leaf and suppression of the flower. The external symptoms of the disease differ somewhat from those on cowberry. Diseased leaves are much larger than the normal, but are neither thickened nor blistered; on the under side they have a whitish or reddish coating, and fall off easily. I have never observed the disease on the stems of bilberry. In spite of these external differences, it is believed that the host-plants are in both cases attacked by the same species of Hzobasidium, but I do not know of any observations on the reciprocal infection of the two hosts. Fic. 259.—Ezxobasidium rhododendri on Rhododendron jerrugineum. (v. Tubeuf phot.) A disease due to an Hrobasidiwm is by no means uncommon on Vaccinium uliginosum (bog whortleberry).2 Shoots of diseased plants are deformed, while their leaves become more or less thickened and assume a beautiful rosy colour. 2 On Vaccinium Oxycoccos (true cranberry) the shoots and leaflets also become thickened and rose-coloured. Rostrup dis- tinguishes this as a separate species (Hxobasidiwm oxycocct). Ex. andromedae Peck. produces on Andromeda polifolia symptoms similar to those just described for the preceding species. (Britain and U.S. America.) ' Sadebeck (Botan. Centralblatt, 1886) records it in large quantity near Harburg- This is the host-species given by Massee (British Fungus- Flora, 1892). 2 Tubeuf, ‘‘ Mittheilungen.” Zeitsch. f. Pflanzenkrankheiten, 1893. EXOBASIDIUM. 427 Ex. rhododendri Cram. (Britain and U.S. America). This causes gall-like outgrowths on the leaves of the Alpine-rose (Rhododendron ferrugineum and kh. hirsutum). The swellings may be small and fairly hard, or, attaining the size of cherries or plums, they may be soft and spongy so that they shrivel up soon after the twig is cut; in colour they are yellowish- white, but on the side exposed to sunlight become rose-red ; the Exobasidium-galis may even be formed on the small rolled- up leaves caused by attacks of mites. Ex. Peckii Hals." [This species occurs in the flowers of Andromeda Mariana in the United States. It is confined almost entirely to the inflorescences, and causes considerable distortion. The bell-shaped corollas are replaced by ones quite polypetalous, and the ovary becomes raised above the re- ceptacle.] (Edit.) The following five species have been recorded on Ericaceae in America : Ex. azaleae Peck. On Rhododendron nudiflorum. Ex. discoideum Ellis. On Rhododendron viscosum. Ex. decolorans Hark. On Rhododendron viscosum and R. occidentale. Ex. arctostaphyli Hark. On Arctostaphylos pungens. Ex. cassandrae Peck. On Cuassundra calyeculuta. Other species to be mentioned are: Exobasidium ledi Karst. On Ledum palustre. Ex. Warmingii Rostr. (U.S. America). This occurs on Suxifraga Aizoon, S. bryoides, S. aspera, etc.; it causes marked hypertrophy of the leaves, and in this way, as well as by its many smaller spores, is distinguished from : Ex. Schinzianum Magn. On the leaves of Suxifraga rotundifolia, causing whitish spots which soon become brown and die. Ex. symploci Ellis. On Syimp/locus tinctoria in North America. Ex. graminicolum Bres. On leaves of various grasses, e.g. Bromus, Arrhenatherum, ete. Ex. lauri Geyl.* is said to produce branched outgrowths of over three feet in length on Laurus nobilis and L. canariensis in the Canary Islands. Urobasidium rostratum Ghgn. occurs on the “ witches’ broom,” out- growths caused by Taphrina cornu-cervi Ghgn. on Aspidium aristatum in India. ' Halsted, Bulletin of the Torrey Club, XxX., 1893, p. 437. *Geyler, Botan. Zeitung, 1874, p. 322, Pl. VIL. 428 BASIDIOMYCETES. HYPNOCHACEAE, Hypnochus. The mycelium forms a cobweb-lke covering on living or dead parts of plants. The sporophores take the form of superficial coatings composed of club-shaped basidia developed on a felted hymenial layer of fungal tissue. Each basidium gives off two to six colourless smooth-coated spores from fine sterigmata. Some species are parasitic, and cause disease. Hypnochus cucumeris Frk.1. In 1882 Frank found at Berlin, on the surface of withering and dying cucumber-plants, greyish coatings of the hymenial layers of this fungus. They occurred principally near the base of the stem, and caused its partial destruction. The symptoms consisted in leaves becoming rapidly yellow from tip to base, and dying off the plant, the lower first. Only cucumbers were attacked, and no further stages could be observed on the killed plants. Hyp. solani Frill. et Del.2 Fine grey crusts, consisting of the hymenial layers of this fungus, were found by these investi- gators on potato-plants; there was, however, no injurious effect on the crop-yield. Aureobasidium. Aureobasidium vitis Viala et Boyer The cause of a vine disease which has done considerable damage in southern France on several occasions since 1882. The grapes when attacked show spots, then shrivel up, their interior becoming completely per- meated by a colourless septate and branched mycelium. On rupture of the epidermis, a firm yellow tissue emerges, and thereon a hymenial layer is developed. The basidia are thick and club-shaped, with a varying number of short sterigmata ; these give off cylindrical unicellular light-yellow spores shehtly curved in shape and with rounded ends. Leaves are also attacked, and fall off after gradually assuming a deep red colour. If this occurs in April, or early in May, the fruit never attains any size. 1Frank, Hedwigia, 1883; and Berichte d. deutsch. botan. Ges., 1883. 2Prillieux and Delacroix, Bulletin de la Soc. mycol. de France, 1891. 3Viala and Boyer, Compt. rend. 1891, p. 1148, and x1x., 1894, p. 248; Aznal. de 1? Ecole nat. @agric. de Montpellier, v1., 1891. THELEPHORA. 429 THELEPHOREAE. Thelephora. The sporophores of this genus assume very varied forms, from simple incrustations to mushroom-like structures. They consist of two layers only, the middle one being absent. The basidia are club-shaped and produce four roundish or oval, hyaline or light-coloured spores. Thelephora laciniata Pers. is not a true parasite, yet it is a dangerous enough enemy to trees. In damp situations, it is ‘common and thrives, growing over young trees and so enveloping them with its sporophores that suffocation ensues. (Britain and U.S. America.) Th. pedicellata Schw. has been reported from America! as a dangerous parasite on apple, Quercus coccinea, and a palm. Th. perdix Hartig, a parasite on oak-wood. (See Stereum frustulosum.) Helicobasidium Mompa. Ichik.* This is injurious to the mulberry tree near Tokyo, Japan. It first attacks the roots, and in consequence the growth of shoots is arrested, the young leaves die off, and gradually death of the tree follows. The mycelium permeates the tissues of the host, and forms an external velvety coating of -basidia. Stereum. Sporophores generally differentiated into three layers, and forming leathery or woody encrustations, or flattened hemi- spherical structures attached by one edge only. Stereum hirsutum (W.) Fr. White-piped or yellow-piped oak. (Britain and U.S. America.) A very common fungus, occurring as a saprophyte on dead branches, on boards, and posts of various kinds of timber, as well as parasitic on living wood, particularly on oak. The sporophores first appear as crusts, later they become cup-shaped ; externally they are brown and roughly hairy with acute yellowish margins. The smooth hymenial layer is orange- red and marked by zones. Between the sterile leathery sporo- phore and the hymenial layer there lies a _ firm white intermediate tissue. ‘Galloway, Journal of Mycology, vi., p. 113. *Nobujiro Ichikawa, ‘‘A new hymenomycetous fungus,” Jour. of College of Science. Imperial University, Japan, 1890. 430 BASIDIOMYCETES. R. Hartig* has investigated in detail the phenomena accom- panying the wood-destruction in the oak. This begins in the branches and extends in white or yellow concentric zones throughout the stem, so producing that appearance which has given rise to the name “ fly-wood.” Portions of the wood appear only white-striped, other parts have a more regular yellowish- white colour. In the white strips the wood has been transformed into cellulose and the middle lamellae of the walls dissolved out ; that of the yellow parts has not undergone this transformation into cellulose, but the destruction has begun from the cell-cavity. A () | i, vi : Fic. 260.—Stereum jfrustulosum. Destruction of Oak-wood. Longitudinal section showing the brown wood with isolated hollow spots containing white mycelium. (v. Tubeuf phot.) Stereum frustulosum Fries. (Thelephora perdix Hartig).” (Britain and U.S. America.) The sporophores form greyish- brown plate-like crusts with concentric markings ; they are small, never exceeding the size of a finger-nail, but generally occur in numbers together, The hymenial layer is composed of club- shaped basidia beset with hair-lhke outgrowths; some of the basidia produce four spores, others are sterile and grow on to form the hymenial layer for the following year. 'R. Hartig, Zersetzungserscheinungen d. Holzes, 1878, Plate XVIII. 2R. Hartig, Zersetzungserscheinungen, Plate XIIT. STEREUM. 431 The very characteristic destruction of oak-wood caused by this fungus was investigated by R. Hartig. The diseased wood has a uniform dark-brown colour, broken at intervals by white rounded spots or hollow cavities ; hence it receives the name _ of “partridge-wood.” In the white spots the wood has by the action of the mycelium become transformed into cellulose, the middle lamellae and starch-grains being dissolved out. In the neighbourhood of old eaten-out cavities the process of decomposition is slightly changed, so that the cell- walls disappear without previous trans- formation into cellulose. CLAVARIEAE. Typhula. Pe zh Sporophores filamentous, and, as t; Mak me a rule, developed from _ sclerotia. Basidia, with four colourless smooth- coated spores. Typhula graminum Karst.' This ee Pag re wee appeared on wheat plants in Sweden, {°" pa Ra iid pp ae killing them and forming yellow eye Peet pies: sclerotia (Selerotium fulvum Fr.). HYDNEAE. Hydnum. Sporophores very variable in form and _ structure. The hymenial layers are spread over teeth-like projections. The basidia bear four white spores. Hydnum diversidens Fr.’ (Britain). The sporophores form yellowish-white crusts or brackets, with spiny outgrowths on the lower side. The hymenial layer consists at first of basidia only, later, however, hyphae grow up through it and build 1 Eriksson, Landthr. Akad. Hand. v. Tidskr., 1879. 7K. Hartig, Zi reelzUndgy se racheinunge n. 432 BASIDIOMYCETES. over it a new hymenium; this is continued for some time so that the sporophore consists of successive layers, and the spiny outgrowths become much thickened. Infection, as was experi- mentally shown by Hartig, takes place on wounds. The wood-destruction, consisting of a white-rot, was studied by Hartig, chiefly on the oak and beech. It begins by the roy) Fic. 262.—Polyporus igniarius. Causing death of a White Alder plantation at Petneu, Stanzer Thal, Tyrol. The stems bear sporophores, and die from above downwards. (v. Tubeuf phot.) appearance of yellowish longitudinal bands (not white as with Stereum hirsutum), and extends gradually till the wood becomes uniformly yellow. The mycelium causes the inner layers of the cell-walls to swell gelatinously without previous transfor- mation into cellulose, and finally to dissolve out leaving the middle lamellae longest intact. oy HYDNUM. 433 Hydnum Schiedermayeri Heufl. (U.S. America).. Sporo- phores fleshy, with a sulphur-yellow colour both outside and inside, and with a smell of anise. They occur on living _ apple-trees, less frequently on other species of Pyrus. Accord- ing to Schroeter, Thiimen, and Ludwig, the mycelium spreads through the stems and kills the trees. Thiimen! thus describes the diseased wood of the apple: “It has a greenish-yellow colour, which passes over gradually to the normal colour of the wood: it becomes soft and _ friable, smelling, like the sporophore, faintly of anise.” Sistotrema fusco-violaceum Schrad. (Britain.) This according to Skiljakow? is parasitic on living pines, entering by wounds, and carrying destruction throughout the wood. POLYPOREAE. Polyporus. Spoerophores large and usually shaped more or less like a hoof or small bracket. The sporogenous layer is com- eet oc seylindrical tubes, 7% 2 oto, iovieries on ake At which generally occupy the Tvevf phot.) lower surface of the sporophore, The substance between the tubes is different from that of the rest of the sporophore. Polyporus (Fomes) igniarius (L.).° (Britain and U.S. America). Sporophores on living stems of oak, alder, apple, willow, and other 'Thumen, ‘* Ein Apfelbaum-Schidling.” Zeitsch. f. Pflancenkrankhe ifen, 1891, *Skiljakow, Scripta botan. horti universitatis Petropolitanae, 1890. *R. Hartig, Zersetzungserscheinungen, Pl. XV. and XVI. Z2E lour, tuber-like or grey in Co BASIDIOMYCETES. 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Zeits incana in Tyrol, which was being Ope Ge (0 f One sisol x QO; one Bh = VSO =| AS. === BB QQ02 Eye es S6 ee BOOBS SS ee, eee ed = RFR Yee WOO) im Ue SS25 Fores |} DY KD )} > — . ~ hoof-shaped, and continue to grow for several years It is f Alnus ation oO ) describes a plant killed out by this fungus (Fig. 262). oy) Oo hrift, 189 « dit. ) y vi (ki species. » common British d POLYPORUS. 435 side is concentrically marked, and has a stone-hard coating which is generally more or less cracked; several zones and layers of tubes will be found when the sporophore is cut in section. This fungus produces a white-rot in the wood, and is one of the most common and dangerous of wound-parasites. The wood attacked by the mycelium is at first dark in colour, then yellowish-white and soft. According to Hartig, a delicate mycelium fills up the elements and eats away the inner layers of the walls; then the middle lamellae are transformed into cellulose and absorbed by it (Fig. 264). Polyporus fomentarius (L.) (Fomes jfomentarivs (L.) Fr.)' (Britain and U.S. America). “ Tinder-fungus.” —Sporophores broad and shaped like reversed brackets or hoofs. Their upper side, at first brownish and velvety, becomes afterwards smooth, grey, and marked with broad concentric zones. The margin is rounded and uniformly grey. The pore-layer is smooth and greyish-brown. A longitudinal section shows a homogenous tinder-like mass, covered on its lower surface by layers or zones of pores. The tinder-fungus is parasitic on beech, elm, and mountain maple. It is particularly common in beech-forests, and was even more so at one time when the infected trees were allowed to remain standing. The sporophores may be found on living stems, on remnants of trees broken by wind, and on felled trees. For some distance above and below the seat of the sporophore runs a furrow on the stem, marking a tract where the mycelium has penetrated to the cambium and killed it, so that growth in thickness ceases (Fig. 266, a). The mycelium causes in the wood a white-rot of a light yellow colour. Where the wood is still firm, though diseased, it will be found to be divided into cubical portions by white tracts of mycelium which run both radially and vertically. A very characteristic feature of the destruction consists of broad white leathery bands of mycelium, formed in a radial direction through the wood; these are best seen on stems shattered by storm, or on wrought timber.” ' Rostrup, Tidsskrift pa Skovburg, 1883. Tubeuf, ‘*Mittheilungen,” A//eg. Forst-. u. Jagd-Zeitung, 1887. A common British species. (Edit. ) * Krull (Schles. Ges. £. vaterland. Kult., 1893) distinguishes a gelatinous mycelium and a cushion-mycelium. 436 BASIDIOMYCETES. Tinder, prepared from the soft central part of the thick sporophores, was at one time used, with the help of steel and flint, for procuring flame. It is very effective in stopping Fic. 265.—Scene in the Bavarian forest near Bischoffsreut. In the foreground, a living 3eech with seven sporophores of Polyporus fomentarius. (v. Tubeuf phot.) haemorrhage from cut blood-vessels, and is still used in surgery. The larger pieces can be manufactured into caps, gloves, vests, POLYPORUS. 437 and hose. The privilege of collecting the tinder-fungi was rented out and regarded as a source of forest-revenue, while the tinder-industry was formerly an important one in many districts, where sporophores were more frequent and larger than now. Measures against this fungus have already been considered in our General Part (§ 12). Fig. 266.—Polyporus Jomentarius on living Beech. «a, A furrow extending above and below the insertion of the sporophore. 4, An injury produced by tearing of the wood in felling. (v. Tubeuf phot.) Polyporus sulphureus (Bull.)' (Britain and U.S. America). The sporophores are flat and soft, the upper side being bright orange-red and the lower sulphur-yellow. They last only for one year, hence are small; they frequently occur in masses, one above another in tiers. After death they lose colour, become brittle, and are easily detached. According to De *R. Hartig, Zersetzungserscheinungen. A very common species in Britain, (Edit. ) BASIDIOMYCETES., near at Hirschau, Polyporus sulphureus on a Willow (Salix alba) Of.— oF phot.) Tubeuf (Vv. Munich. POLYPORUS. 439 Seynes,’ three other kinds of spores are produced in addition - to basidiospores. Willow, poplar, oak, sweet chest- nut, alder, ash, hazel, pear, cherry, robinia, larch, silver fir, ete., are common hosts of this parasite. Wood infested by the mycelium darkens in colour, exhibiting a red- rot. Vessels and all clefts or spaces become filled with white felted masses of mycelium. The wood, in course of destruction, becomes richer in carbo-hydrates, and the walls of the wood-fibres shrink so that fis- sures with an upward right to left direction are formed, but do not reach Fro. 268. — Polyporus sulphureus ial layer, with basidia anc the middle lamellae. Finally the 23me* de netic. wood becomes dry, brittle, and powdery. Polyporus borealis (Wahlenb.) Fr.” (Britain and U.S. America). Sporophores annual, white, and fleshy; the upper Fic. 269.—Polyporus sulphureus. The white mycelium forms concentric zones and radial lines on the cross-section of Oak, (After R. Hartig.) surface is shaggy when fresh, and no internal zones are exhibited. The shape is somewhat cushion or bracket-like, but very variable ; 'De Seynes, Annal. de Sci. nat., Ser. V., Vol. t., 1864, *R. Hartig, Zersetzungserscheinungen, Pl. X. 440 BASIDIOMYCETES. many generally grow near each other. The pores have a torn margin and cystids are frequent between the basidia. The sporophores are common in spruce plantations, and are accompanied by a very characteristic wood-destruction. The wood, in the earlier stages, becomes brownish-yellow and inter- sected by radial and vertical canals filled with a white mycelium (Fig. 270). Gradually, however, it breaks up into small cube- i Sie RS pea re ata Tee SEE eS Fic. 270.—Polyporus borealis. Destruction of Fia. 271.—Polyporus borealis. Later stage Spruce-wood. The white mycelium is present, of destruction. The Spruce-wood is broken dividing the decayed wood into cubical pieces. up into cubical pieces, and the mycelium has (v. Tubeuf phot.) disappeared. (v. Tubeuf phot.) like pieces, particularly evident when the wood is broken (Fig. 271). The cell-walls are dissolved from the cell-cavity out- wards, the lignified wall being first converted into cellulose and disappearing, finally the middle lamella. Polyporus dryadeus Fr.' (P. psewdoigniarius Bull.) (Britain and U.S. America). Sporophores, annual, large, shaped like tubers or hoofs, and generally situated towards the base of the TR. Hartig, Zersetzungserscheinungen, Pl. XVII. A common British species. POLYPORUS. 441 stems of oak-trees. At first they are soft, later hard and brown with grooves on the upper side. The dark heart-wood of the oak exhibits white or yellowish longitudinal stripes of rotten wood converted into cellulose (Fig. 272). In the white portions St Neh mee Ch id Fic, 272.—Polyporus dryadeus. The mycelium forms longitudinal stripes in the Oak-wood. (v. Tubeuf phot.) the destruction is more complete than in the yellow, where dis- solution of the lamellae has not as yet taken place (Fig. 273), A simultaneous destruction of the wood by 2. dryadeus and P. igniarius may occur (Fig. 274); in this case, the medullary 4492 BASIDIOMYCETES., rays appear snowy white at the place where the two forms of rot meet; this is due to an accumulation of starch left after the cell-walls have been almost completely dissolved. Polyporus (Poria) vaporarius (Pers.)' (Britain and U.S. America). The sporophores are white, and have a pungent odour; they form crusts (never brackets) closely adherent to dead substrata, especially to beams and other timber in buildings, Fia. 274.— Polyporus dryadeus and Poly- porus igniarius. Destruction of Oak- Fic. 273.—Polyporus dryadeus. Later wood under the combined agency of stage of decay of Oak-wood. The darker both fungi. The wood is yellowish and places still consist of firm brown wood ; perforated; the medullary rays are the white, however, are soft cellulose. snowy-white, from the accumulation of (v. Tubeuf phot.) : unchanged starch. (v. Tubeuf phot.) where this fungus does great harm. They are also found, how- ever, on bark of living stems of spruce and fir. The destruction takes the form of a red-rot, the wood attacked becoming red- brown, cracked, and soft. The mycelium is found in stems and roots of trees; in cracks in the wood and below the bark, and on the surface of timber in buildings, it forms fan-shaped strands of a permanent white colour. The mycelial strands of the “dry-rot fungus” (Merulius lacrymans) differ from it in being 1 Very common in Britain on dead wood, less so on living trees. (Edit.) POLYPORUS. 443 at first white but becoming grey, and in exhibiting an internal differentiation which those of P. vaporarius do not. The hyphae in the course of their growth do not seek out the pits, but grow straight through the walls and bring about dissolution of the middle lamella for some distance around. At the same time numerous short oblique fissures in the walls are produced vertically one over the other, especially in the elements Fic. 275.—Polyporus squamosus on Acer Negundo. The three upper sporophores are borne on a separate piece of wood, from which a fourth has been cut off. (v. Tubeuf phot.) of the thick-walled autumn wood. (Compare with P. sistotre- moides, Fig. 280). The phenomena accompanying destruction of wood by this fungus are so characteristic that Conwentz* could distinguish it quite clearly in tree-remains enclosed in amber. Brefeld succeeded by artificial culture of the spores, in raising a mycelium on which basidia were formed, at first directly, afterwards from large sporophores. Polyporus squamosus (Huds.). (Britain and U.S. America.) 'R,. Hartig, Der echte Hausschwamm, Berlin (Springer), 1885. *Conwentz, Monographie d. baltischen Bernsteinbiiume, 1890, 444 BASIDIOMYCETES. Sporophores annual, occurring from spring to autumn; at first tender and fleshy, later leathery or almost woody. In form they are short-stalked, flat, semi-circular or kidney-shaped, and attached by one edge; they may also be stalked and circular or cup-shaped. Their upper surface is yellowish, with flat brown scales arranged in concentric nes. The hymenial layer is continued well on to the thick fleshy stalk of the sporophore ; it is yellow in colour, and consists of short angular pores. Fic. 276.—Polyporus hispidus on pieces of living Ash. (v. Tubeuf phot.) The spores are spindle-shaped and colourless. The fungus is especially common on living hazel, ash, species of maple, beech, mountain ash, horse-chesnut, elm, oak, willow, pear, lime, ete. The wood of the specimen in Fig. 275 exhibited extensive white-rot, the inner parts being completely converted into a soft white spongy mass of mycelium. Polyporus hispidus (Bull.).! (Britain and U.S. America.) Sporophores annual, soft and spongy, with a rough brown upper 1A very common form on ash trees in Britain. (Kdit.) POLY PORUS. 445 surface, and a smooth yellowish hymenial surface. They are large and fiat, the thickest part being at their insertion (Fig. 277). Several frequently occur on the same stem, especially if wounds or frost injuries are present. The spores are brown and roundish. Conidia are said, by Schroeter, to be formed on the upper surface of the sporophores. This species is a deadly enemy of fruit-trees, especially apple. In the vicinity of Munich the sporophores are common on ash. Schroeter gives elm and plane as hosts, and Prillieux Fic. 277.—Polyporus hispidus, Longitudinal section through a living stem of Ash, and a sporophore of P. hispidus. The stem shows symptoms of wood- destruction, in that it becomes brown and has short white longitudinal and radial stripes. (v. Tubeuf phot.) and Delacroix state the fungus to be very dangerous to the mulberry in France, It causes! brown discoloration of the wood accompanied by characteristic short white lines in both radial and _ vertical directions, so that the wood becomes marked out in squares. Polyporus (Poria) laevigatus Fr.*. Sporophores dark-brown 1 Prillieux (Bullet. de la Soc. mycolog. de France, 1x., 1893), gives details of the destruction of the wood. *Mayr, Botan. Centralblatt, x1x., 1884. 446 BASIDIOMYCETES. and forming crusts on the bark of birch. Spathulate cystidia occur between the basidia. Spores colourless, and acutely ovate in shape. This is parasitic on birch. The mycelium kills and permeates the wood-parenchyma which forms the greater mass of the later-formed parts of each year-ring, with the result that the various year-rings of the wood separate from each other as concentric hollow cylinders. The mycelium varies according as its pabulum consists of cells just killed, or of wood, or of elements in the last stages of decomposition; in this latter case it suffers from want of food. In woody elements in contact with air, or those destroyed by Polyporus betulinus, the mycelium is brown and forms vesicular tyloses similar to Agaricus melleus. Polyporus betulinus I'v. (Britain and U.S. America). The sporophores are annual, and emerge as_ spherical structures from the unin- jured bark, or from bore- holes of Beetles, or other wounds. When mature Fic. 278.— Polyporus betulinus on Betula verrucosa. F : The sporophore was developed horizontally on a they are hoof-hke or seml- fallen stem; it is here, however, set up vertically PK and photographed from the lower side. (v. Tubeuf elrcular and short-stalked z phot.) € when dead they become soft and break off. The upper side is heht-brown in colour, the pore-layer is white. A section through the sporophore shows it to be white and homogeneous without zones. Lanceolate cystidia occur between the basidia. The spores are rod-like. The pore- layer and the upper brown layer are easily detached, and strips of the remaining tissue are sometimes utilized as razor-strops. This parasite frequents living birches, ultimately causing death. It is known to occur on both Letula verrucosa and B. pubescens in Britain, America, and Europe. Its parasitism and injurious results were first demonstrated by Rostrup.'. Mayr? investigated 1Rostrup, *‘Snyltesvamper Angreb paa Skovtraeerne,” Tidsskrift pa Skov- burg, 1883. 2Mayr, Botan. Centralblatt, x1x., 1884. POLYPORUS. 447 in greater detail the destruction brought about by its mycelium. He found that it penetrates lignified cell-walls, entering the living elements and causing their death; it spreads most rapidly in the vertical direction through wood, bast, and rind, growing through parenchyma and sieve-tubes, and even boring its way into the sclerenchymatous stone-cells; it absorbs the secondary thickening by dissolving out first the ligneous incrustation, next the cellulose, while the middle primary lamella remains behind untouched. Polyporus (Fomes) fulvus (Scop.) (Britain). Sporophores woody and very hard, at first hairy but later smooth, dark, and cracked; in form they are tuberous or triangular. In- ternally they show no stratification. The fungus is very common on living plum where it causes undoubted injury; it also occurs on hornbean and aspen. Polyporus fulvus var. Oleae Scop. In northern Italy may be frequently observed a peculiar splitting of the stems of olive trees into two or more portions; the fissures occur generally on the lower parts of the tree, and may extend so deeply that the stem appears to stand on stilts or props. Hartig! ascribes this phenomenon to the presence in the olive stems of the mycelium of Polyporus fulvus causing rotten places which are cut out by the Italian cultivators ; the disease, how- ever, continuing to make progress, it may be necessary in course of time to cut so deeply into the stem, that tracts extending right through may be removed; this takes place all the more rapidly if several diseased spots are being simultaneously operated on. The destruction of the olive-wood by this parasite is similar to that produced by P. igniarius on oak and other trees. The sporophores appear on rotten spots, but are gener- ally quickly removed by the cultivator. Infection takes place on wounds, hence it is advisable at once to apply tar after cutting out any decayed wood, and also to paint pruning-cuts or other exposed surfaces with tar. Neighbouring fruit-trees, liable to suffer from this same fungus, should be similarly treated, both for their own safety and that of the olive trees. Polyporus (Fomes) Hartigii Allescher” (P. igniarius var. 'R. Hartig, ‘Die Spaltung der Oelbiiume.” Forstlich-naturwiss, Zeitschrift, 1893. *R. Hartig, Zersetzungserscheinungen, Pl. VII. Forstlich-naturwiss. Zeit achrift, 1893, p. 61. 448 BASIDIOMYCETES. pinuum Bresadola or P. fulvus Scop. of R. Hartig). Sporophores on silver fir, less commonly on spruce. Their form varies much, according as they occur on a branch or on the stem. In the former case, the sporophore forms a swelling below and on each side of the more or less horizontal branch. On the stem they are more or less bracket-like. The sporophores are reddish brown with a smooth upper surface on or altogether absent. Internally they are of a brownish or tawny colour, and exhibit concentric strata, which. do not extend into the pore-layer; they are thus distinguished from sporophores of P. igniarius and others. The sporophores are very frequent on cankered stems of fir where the canker-spots afford easy entrance for the spores. The wood-destruction consists in a white-rot. The wood becomes yellowish- white with clear spots and fine dark lines, especially where in contact with healthy parts. The mycelium is yellow- ish, and consists of thick hyphae with lateral. branches forming tangled masses which frequently fill up the cavity of the bordered pits. This mycelium gives off very fine branches which bore through porte 2 — Poluporus Hartigii the cell-walls and dissolve them in such The decayed wood is yellow, but a Way that the middle lamellae disappear Goer estate first and leave the remainder of the wall- thickening for a time isolated before it too is used up. In this way large holes are formed in the elements of the wood. Polyporus sistotremoides (Alb. et Schw.) (P. Schweinitzii Fr. or P. mollis Fr. of R. Hartig)' (Britain). Sporophores almost circular with a short thick central stalk; while young they are light brown and spongy, but when older become dark brown and corky. The upper surface is downy; the hymenial layer extends far down the stalk, when young it is yellowish green, 'R. Hartig, Zersetzungserscheinungen, Pl. IX, which zones are only faintly indicated | TTR) — POLYPORUS. 449 but later becomes brown, and, on being touched, deep red. The spores are white, and various forms of hairs occur among the basidia. Young sporophores appear as little brown cushions on felled timber, also on living stems of pine, and, according to Magnus, on Weymouth pine. The disease generally makes its first appearance in roots and lower parts of the stem, spreading thence into higher parts. Diseased wood has a characteristic odour of turpentine; it has a reddish-brown colour, and, as destruction proceeds, it gradually shrinks and disintegrates till it becomes so soft as to be easily powdered between the fingers. Where broken over, the wood is often covered with a thin white coating of mycelium incrusted in resin so as to appear like chalk. The mycelium penetrates the cell-walls in all directions. A very characteristic feature of this parasite is furnished by shrinkage-fissures in the thick walls of the tracheids of the summer-wood (Fig. 280). These are numerous and run upwards from Bige aol. Araceae. Of Pinusdestroyed by Polyporus jo ‘ Brefeld, Schimmelpilze, Heft 8, 1889. *Rostrup, Afbildning og Beskrivelse af de farligste Snyltesvampe, 1889. : — 'e POLYPORUS. 451 The mycelium penetrates both bast and rind causing a very acute red-rot in the wood, so that death of the tree attacked rapidly follows. The disease makes its appearance on plants of all ages, and in forests of spruce or pine causes gaps which rapidly extend in a centrifugal direction. The roots and lower parts of the stem are generally the parts first attacked. On the roots, the parasite is easily distinguished, even in the absence of sporophores, by the very delicate white mycelial membranes formed between the bark-seales. Destruction of the wood becomes first evident by the appearance of vertical dark lilac- coloured stripes indicating the stage when the parenchyma cells are killed. At a later stage, the wood becomes brown, and shows isolated black spots with white margins (Fig. 282). These last consist of coils of dark mycelium surrounded by wood from which the incrusting _ Fie. 282.—Polyporus annosus. Destrac- 3 = tion of Spruce-wood. Longitudinal sec- substance has been dissolved away, tion showing white (cellulose) spots with : = ° black (mycelium) centres. (v. Tubeuf leaving only cellulose, readily dis- phot.) tinguished by turning blue’ on treatment with chlor-zinc-iodine; here too, the middle lamellae are ultimately dissolved out, so that the elements become isolated. A colourless mycelium may also be found in the other parts of the wood, both inside the elements, and extending in all directions through the cell-wall, leaving holes where it itself has disappeared. Dissolution of the lignifying substance pro- ceeds from the cell-cavity, the middle lamella remaining intact till the last. The resin of the decayed wood passes over into all healthy parts and flows from the bark of diseased stems as a resin-flux. The most effective method for combating the ravages of this parasite is isolation of infected areas. In one case which I investigated in Baden, several spots in the forest formed very evident starting points, and sporophores were everywhere present at the base of stems amongst the moss. Such spots should be enclosed by ditches with vertical sides, and deep 452 BASIDIOMYCETES. enough to cut through all roots, care being taken to leave no diseased stems or roots outside the circle; after remaining open for a time, the ditch must be refilled with soil to prevent development of sporophores on the exposed roots. Diseased stems should be felled, and, along with all root-remains, burned on the spot, where there is no risk of forest fire; failing this, they and their stumps should be deeply covered over with soil, to prevent development, of sporophores. The following species of Polyporus have been observed on living trees, but details in regard to their parasitism and mode of destruction are still wanting: P. officinalis Fr. On larch, chiefly in Russia, but also in France and Switzerland. The sporophores are white irregular masses, and at one time were used in medicine. The mycelium forms bands in the wood similar to those of P. sulphureus. P. albus (Corda), according to Ludwig! is a cause of a disease of Conifers, which extends from the root upwards. (U.S. America.) P, spumeus (Sow.). On apple trees. (Britain and U.S. America.) P. fumosus (Pers.). On willow, ash, maple, and other broad-leaved trees. (Britain and U.S. America.) P. picipes Fr. On willow and other broad-leaved trees. (Britain and U.S. America.) P. (Fomes) cinnamomeus Frog. On cherry trees. (Britain.) P. radiatus (Sow.). On alder (A. incana), birch, and beech. (Britain and U.S. America.) P. (Fomes) ribis (Fr.). On black currant and gooseberry shrubs. (Britain and U.S. America.) P. (Polystictus) hirsutus Fr. (Britain and U.S. America). On living hornbeam, alder, oak, birch, and service. A variety, scrwposus, is common and injurious on cherry. P. ulmarius Fr., is, according to Cavara,” parasitic on living elm near Pavia. (Britain and U.S. America.) P. (Fomes) nigricans. On birch. (Britain and U.S. America.) P. salicinus (Pers.). A dangerous enemy of willow.? (Britain and U.S. America.) Rostrup* gives Corticium comedens as a wound-parasite of oak and alder. Hartig describes Fistulina hepatica, the liver-fungus, as causing a dark- brown colour in oak-wood. lLudwig, Lehrbuch d. niederen Kryptogamen. 2Cavara, Revue Mycol., 1891. ’Tursky, Russian translation of R. Hartig’s ‘‘ Lehrbuch d. Baumkrankheiten.” 4+Rostrup, Fortsatte Undersogelser, 1883. " TRAMETES. 453 Trametes. Sporophores as in Polyporus, except that the substance between the pores does not differ from that of the rest of the sporophore.* Trametes pini (Brot.) Fr.2 Ring-scale of Pine. This is a dangerous forest parasite in Northern Germany; also in Britain and U.S. America. On the pine the sporophores develop from branch-scars, and assume a bracket form. The fungus has also been observed on spruce in Bavaria and elsewhere, but in this case, the sporophores are more frequently found as a coating over the bark on the under side of a branch. Larch, silver fir, and the Douglas fir (in America), have also been mentioned as hosts. The sporophores are brown and woody, and continue to form annual hymenial zones for a number of years. The hymenial layer consists of pore-tubes lined with basidia, between which thick-walled cystidia are formed. The spores are elliptical, and on germination penetrate into wounds or broken branches not protected by an outflow of resin. The older branches of pine and larch have a central heart-wood from which no resin is secreted, and these branches, when broken over, offer the neces- sary access to the germinating spores; for this reason, infection takes place most frequently in old plantations. The mycelium spreads through branch and stem, particularly upwards and downwards in the same year-ring. In this way longitudinal stripes and peripheral zones are formed in the wood, giving rise to the popular name “ring-scale.’ Single hyphae bore through the cell-walls, and a ferment secreted by them dissolves the incrusting substance, so that walls affected show the re- actions for cellulose almost at once. A very characteristic feature is the appearance of isolated white spots or holes, indi- cating where the wood, after becoming cellulose, has been dissolved out entirely. The middle lamellae are dissolved out first in attacks of this fungus, the tertiary lamellae remaining longest intact (Fig. 12). The dark centres of mycelium inside 'The distinction between the genera Polyporus and 7'rametes is badly detined. A reinvestigation of the systematic relationships of the whole group of Polyporeae would in fact be advisable. *R. Hartig, Zersetzungserscheinungen, Pl. V. and VI.; Wichtige Krankheiten, Pl. ILL. ; Lehrbuch d. Bawmkrankheiten, 1894 (English translation by Somerville). 454 BASIDIOMYCETES. white wood-spots, so characteristic of Polyporus annosus (Tram. radiciperda), appear only rarely in this species. The destruction of spruce and fir goes on from pith to bark; in the pine, however, it seems to be confined to the heart-wood, Fic. 283. — Trametes pini on Spruce (Picea Fic. 284.—Trametes pini on Spruce. excelsa). Sporophore on the stem beneath a Board showing the characteristic snag-branch. (v. Tubeuf phot.) white cellulose-spots in the wood. (v. Tubeuf phot.) and is prevented from entering the sap-wood by a firm zone permeated with resin. Remedial measures are the removal of all diseased stems TRAMETES. 455 at thinning; and the prevention of unnecessary injuries to living branches or stems. Trametes suaveolens (L.), common on dead willow, is also reported as parasitic on living stems. (Britain and U.S. America.) AGARICINEAE. Agaricus. Sporophores umbrella-shaped and fleshy, and decaying soon after discharge of the spores.) _Hymenium on the under side of the umbrella, and spread over a series of radiating gills or lamellae, easily divisible in a longitudinal direction. The genus is divided into sections and subgenera distinguished by the colour of the spores; the Coprinarii are black-spored ; the spores of the Prate/li are dark purple, brownish-purple, or dark brown; of the Dermini brown, yellowish-brown, or orange ; ot the Hyporhodii rosy or salmon-coloured; of the Leuwcospori white. Agaricus (Armillaria) melleus Vahl! (Britain and U.S. America.) The honey-fungus or “hallimasch.” The sporophores are present in numbers towards the close of summer on tree- stools of all kinds, and on the bark of dead or living Conifers ; also on timber, and even on earth. The fleshy stalk is somewhat thickened towards its base, and towards the upper part bears the membranous yellowish annulus (Fig. 286). The cap sur- mounting the stalk is honey-coloured or brownish with dark scales. The spores are white and bestrew adjacent objects with a mealy dust. The sporophores are edible. The connection between the sporophores and the rhizomorph- strands was proved by Hartig. These rhizomorphs are very common and vary much in form; they occur as round brown strands running through the earth from root to root of attacked trees; inside hollow stems and in wooden water-pipes, they retain their rounded form, but under the bark of trees they become dark brown flattened bands (Fig. 288). They are not uncommon on. timber; in mines they may be frequently seen hanging from the woodwork as tangled clumps, with 'R, Hartig, Wichtige Krankheiten, Pl. I. and I. ; Zersetzungserscheinungen, Pl. XI. De Bary, Botan. Zeitung, 1859. Brefeld, Schimmelpilze, Heft. 111., 1877, 456 BASIDIOMYCETES. numerous branches like the runners of some hanging plant, e.g. Aaron’s Beard (Saxifraga sarmentosa). The rhizomorphs live as saprophytes and have been long known to emit phosphorescent hight. Sporophores are developed directly on them, and if one sows the spores, a delicate hyphal tissue is produced, which, under suitable conditions, passes gradually over into the rhizo- morph-strand. Brefeld succeeded in raising rhizomorphs from spores in artificial nutritive media. The Agaricus-mycelium forms fan-shaped snowy-white firm membranous expansions under the bark of newly killed or still living trees. They are quite distinct from the much more delicate mycelial expansions of Polyporus annosus, and offer a particularly easy means of distinguishing between the two species. Another indication of Agaricus is the great outflow of resin from the bark at the base of the stem and from roots, whereby hard clumps of earth are formed round the roots. The passage of the rhizomorphs into the white membranous mycelium is easily observed. The AGARICUS. 457 rhizomorphs distribute the fungus in the earth and other dead substrata, as well as bore into the bark of healthy Conifers. This parasite attacks not only the indigenous Conifers (spruce, silver fir, pine, larch, and juniper),’ but also the introduced - forms—Weymouth pine, Douglas fir, Pinus rigida, Abies Pichta, Picea sitchensis, various Cupressineae, etc. It also seems to attack broad-leafed trees, at least as a wound parasite.? In regard to the interesting structure of the rhizomorphs, and the characteristic mode of wood-destruction caused by Fic. 287.— Agaricus iielleus. Section through a lamella. d, The hyphae forming the substance of the lamella are much branched, and send twigs outwards which end in club-shaped basidia, a; on many of Fic. 286.—Agaricus melleus. Sporophore developed from a rhizomorph-strand; the other branch bears arrested sporophores. (After R. Hartig.) these are developed sterigmata with apices swollen into spores, 4; c, isolated thread- like arrested basidia projecting above the hymenial layer. (After R. Hartig.) this fungus, I give directly the account by Hartig in his “ Lehrbuch.” ? “The pathological symptoms can only be ex- plained in the light of the peculiar organization of the mycelial growth that lives in the cortical tissues. The apex of the thizomorphs consists of delicate pseudoparenchyma, which, elongating by the division and growth of the cells, produces delicate hyphae on the inside at a certain distance from the 'T found it on juniper in the pine-forests near Eberswalde. (Auth.) * The a gee does not seem capable of penetrating uninjured broad-leaved trees, but R. Hartig (Forst/.-naturwiss. Zeitschrift, 1894, p. 428) mentions attack and death of cut roots of healthy oaks, *R. Hartig, Lehrbuch d. Baumkrankheiten, 1889. The translation given here is from Prof. Somerville’s English edition of Hartig’s text-book, p., 210. (Kdit.) 458 BASIDIOMYCETES. point, whereby a felted tissue, called the medulla, is produced in the interior. The outer parts of the pseudoparenchyma, on the other hand, coalesce to form the so-called rind, which when young gives off numerous delicate hyphae, and these, taking advantage of the medullary rays, penetrate the wood, and especially the resin-ducts, should such be present. In the wood the growth is upwards. This filamentous mycelium, which pro- gresses much more rapidly in the interior of the wood than the rhizomorphs which grow in the cortex, completely destroys Fic. 288.—Agaricus melleus. Rhizomorphs in the form of dark anastomosing bands, developed between the bark and wood of a tree. (v. Tubeuf phot.) the parenchyma that exists in the neighbourhood of the resin- ducts, and to all appearance this is accompanied by a _ partial conversion of the cell-contents and the cell-walls into turpentine. The turpentine sinks down under its own weight, and in the collar, where the cortex is withered, having been killed by rhizomorphs, it streams outward, pouring partly in between the wood and the cortex, and partly into the surrounding soil at places where the cortex has ruptured owing to drying. On ‘ > this account the disease was formerly called ‘ Resin-flux’ or AGARICUS. 459 ‘Resin-glut.’ In the upper parts of the stem, where the cambium and cortex are still sound, the turpentine also flows laterally, by means of the ducts of the medullary rays, from the injured canals towards the cambium and cortex. In the latter this accumulation induces the formation of large resin-blisters. When, during the summer, the cambium is forming a new ring, the plethora of resin has the effect of causing the production of numerous resin- canals, which are usually large and abnormally constructed, and these impart to the wood-ring formed during the year of sick- ness a very striking and characteristic appearance. “The mycelium gradually spreads from the cells of the medullary rays and from the resin-ducts into the vascular elements of the wood, where it produces a form of decay which may be termed a variety of white-rot. During the progress of the decomposition from the surface of the stem inwards a certain stage is reached, which is highly favourable to the development of the mycelium. While previously it was simply filiform and furnished with numerous lateral hyphae, it now develops large bladder-like swellings, and at the same time the hyphae change into a kind of large-meshed parenchyma, which, like the tyloses in the vessels of many dicotyledonous trees, completely fills up the lumina of the tracheides, On account of the mycelium assuming a brown colour when in this condition, it makes the portion of diseased wood which it infests appear, to the naked eye, like a black line. As this kind of mycelium soon dies off and is dissolved, being replaced by a delicate filamentous mycelium, it seldom happens that the zone which it occupies exceeds the breadth of three to four tracheids. The walls of the elements of the wood afterwards display a_ cellulose reaction, and speedily dissolve from the lumen outwards. “On account of the trees drying up, after the rhizomorphs have spread from the point of infection on the roots into the stem, and again from the stem into the hitherto sound roots, decomposition of the stem usually ceases before the mycelium has advanced from the alburnum into the duramen. It is only in the stool and roots that decay rapidly spreads throughout the whole of the wood.” Methods for exterminating this parasite are unknown, beyond removal of diseased plants and collection of sporophores. — It would certainly be advisable not to plant young conifers on 460 BASIDIOMYCETES. cleared forest-land where the fungus sporophores are numerous on dead stools or roots. Agaricus (Pholiota) adiposus Fr.’ (Britain and USS. America). This is a conspicuous bright yellow or honey-yellow toadstool, with a glistening slimy cap which, as well as the stalk, is beset with concentric darker scales (Fig. 289). The scales and delicate annulus become indistinct or disappear on old sporophores or after much rain. The stalk is thick, fleshy, and stiff, and while growing so changes its direction as to keep the cap always in a horizontal position. The pileus or cap, at Fic. 289.—Aygaricus adiposus. A, Amature and avery young sporophore grown in the forest; the other sporophores were raised on Fir-wood in a cellar. The latter have longer thinner stalks, and a basal swelling beset with white down. B, Mature and germinating spores. (v. Tubeuf del.) first globose, opens out cone-shaped or flat with a diameter of about 9c.m. Remains of the velum adhere to the margin. The underside of the cap is at first yellow, later mouse-grey. The lamellae are of three sizes, the largest extending from margin to stalk. From the lamellae arise the basidia, with four sterigmata each giving off a single spore. The spores fall at maturity, and cover neighbouring objects with a brown dust. They are oval with a length of 7-10 and a breadth of 5-6m. The sporophores spring up rapidly in large numbers on ly. Tubeuf, ‘‘ Eine neue Krankheit d. Weisstanne.” Zeitschr. f. Forst u. Jagd- wesen, 1890. AGARICUS. 461 living stems of silver fir, beech, etc., and on felled wood.’ In the forest, on newly erected piles of firewood, the yellow stools may frequently be found in every stage of development growing from the cut billets, while they are especially numerous on the rotting useless timber left lying. In cellars or other moist Fic. 290.—Agaricus adiposus. Destruc- tion of Fir-wood. The deeply-corroded Fic. 291.—Agaricus adiposus. Destrue- cross-fissures contain white mycelium ; the tion of Fir-wood. Later stage. The remainder of the wood is yellow. (v. corroded fissures no longer contain my- Tubeuf phot.) celium. (v. Tubeuf phot.) : chambers, the sporophores may be abundantly produced till Christmas, but out-of-doors, August is the time of fructification. The mycelium forms felted masses under the bark or in cracks of the wood, and thence the sporophores arise as little pale- yellow buttons, which gradually unfold and become differentiated into cap and stalk. While quite young, they show the darker ‘Previous to the publication of v. Tubeuf’s investigation the fungus had only been observed on living beech and felled wood. 462 BASIDIOMYCETES. scales arranged regularly in concentric lines. The stools break out from living stems through cracks in the bark or from wounds, eg. those made by wood-peckers. The fir-wood, normally white, assumes, when diseased, a yellow or honey-colour, more or less like the sporophore, while here and there, parts may become light brown. The hyphae grow in all direc- tions, but especially as white strands up and down the year-rings, while others in horizontal and vertical direction break up the wood into irregular patches (Fig. 290). Im the final stages of destruction the wood will be found laminated into its separate year-rings and very much broken up into irregular pieces (Fig. 291). The sporophores of this Agaricus are not uncommon protruding from bark- cankers caused by Aecidiwm elatinum, and its mycelium assists in the destruc- Fie. 292.— Agaricus adiposus : and Polyporus Hartigii. Destruc- tion of the stem. tion of Fir-wood. The boundary of the regions affected by the Agaricus (Pholiota) squarrosus (Miill.). On two fungi is formed by a very BSG dark line of demarkation. To living and dead stems of broad-leaved trees. the right the destruction is that ACS produced by Polyporous Hartigii, (Britain. ) to the left by Agaricus adiposus. Ag. (Ph.) destruens (Brond.). On living (v. Tubeuf phot.) oS - and dead stems of poplar. (Britain.) Ag. (Ph.) aurivellus (Batsch.). On living and dead stems of broad-leaved trees. (Britain.) “FUNGI IMPERFECTI.” The fungi placed here have life-histories which as yet have not been completely investigated, most of them being known only in the form of pycnidia or conidia. The number of species was at one time much larger, but it is gradually being re- duced as the forms are proved to be stages in the life of some species of definite systematic position in the other groups already considered. The group may be divided into the provisional sub-groups, the Sphaeropsideae, Melanconieac, and Hyphomycetes. FUNGI IMPERFECTI. 463 I. SPHAEROPSIDEAE. Conidia abjointed from conidiophores contained in dark- coloured pyenidia somewhat spherical in form. The various species are provisionally arranged in genera according to the colour of the conidia and the number of cells contained in them. The families here included are the Sphaerioideae, Nee- trioideae, Leptostromaceae, and Exreipulaceae. 1. FAM. SPHAERIOIDEAE. HYALOSPORAE. Phyllosticta. Fungi with colourless spores, and producing sharply defined spots on living leaves. They occur on all kinds of woody plants, but as a rule the injury caused is too slight to be called a disease. From the long list given by Saccardo (Vols. U1, Ix., and x.) the following have been selected: Phyllosticta persicae Sacc. This produces on leaves of peach brownish-yellow spots, frequently marked by concentric zones. The name “shot-hole fungus” has sometimes been applied to this and other allied forms, because the leaves become more or less perforated by the withered spots falling out. The pyenidia on reaching maturity rupture the lower epidermis of the leaf in a star-like manner. Briosi and Cavara do not regard this parasite as very harmful, because leaves attacked by it remain alive without serious prejudice to their function. Ph. pirina Sacc. has been observed injurious to pear-trees at Geisenheim (Germany). Ph. prunicola Sacc. is the cause of spotting of leaves of apple, plum, cherry, and apricot. (U.S. America.)! Ph. cytisi Desm. On leaves of Cytisus Laburnum in Britain and Europe. Ph. acericola Cook et Ellis. On various species’ of maple (Acer dasycarpum, ete.) It is deseribed by Galloway * 'The chief authorities for the occurrence of the ‘‘ Fungi imperfecti” in Britain and North America are Massee (British Fungus Flora, 1895), Farlow and Seymour (Host-Index for U.S. America, 1891), and Saceardo (Sy/loge eae Professor J. W. H. Trail kindly revised the records for Britain, * Galloway, ‘‘ Report of sect. of Veget. Pathology for 1888.” U.S.A. Dept. of Agriculture, Washington. 464 FUNGI IMPERFECTI. as injurious, especially in nurseries and groves where the trees are grown in number. Ph. sphaeropsoidea FE. et E. is another American species which has become prominent on account of its ravages in nurseries of horse-chestnut. The disease appears about the end of June, and by August the foliage of attacked trees is almost entirely dead. Ph. grossulariae Sacc. On leaves of Ribes Grossularia in Italy and North America. Ph. vulgaris Desm. A common species on leaves of species of Lonicera. (Britain and U.S. America.) Ph. sambuci Desm. On species of Sambucus. (Britain.) Ph. cornicola (D.C.). On leaves of species of Cornus in America. Ph. limbalis Pers. On oblong white spots on leaves of box. (Britain.) Ph. tiliae Sacc. et Speg. On leaves of Zilia. (Britain.) Ph. maculiformis Sacc. is probably a stage of Sphaerella maculiformis Auersw. It is a dangerous parasite causing a leaf-spot on sweet chestnut (Castanea) and other trees. Ph. violae Desm. . ing the constriction characteristic of this disease, Pinus sylvestris in Berlin dotted over with pycnidia. (After R. Hartig.) be fanic varden IU o . 5 Other species of Phoma frequent other conifers and broad- f 1 Bohm (Zeitsch. f. Forst- u. Jagd-wesen, 1896, p. 154) describes and figures an attack of this parasite on Pseudotsuga Douglasii in North Germany. One cannot, however, avoid suggesting some confusion between this and PA. pithya described next. (Edit. ) “E. Rostrup Undersoegelser over Snyltes vampes Angreb paa Skovtraeer, 1883-1888. PHOMA. 467 leaved trees, but details in regard to their parasitism are wanting. Ph. Hennebergii Kiihn.' Brown-spot of wheat-ears. This produces, on the glumes of wheat, brown spots with projecting pyenidia from which unicellular conidia emerge. The grains of attacked ears shrivel up and become spotted, while the value of the chaff as fodder is much diminished. Whole fields may be attacked, showing marked discoloration, and producing but few healthy ears.” The fungus may also appear on the leaves and produce pycnidia. Ph. lophiostomoides Sacc.* — Lopriore revards this as a parasite on cereals, but Cavara looks on it as saprophytic on the dead plants. Ph. ampelinum De Bary (Sphaceloma ampelinum De Bary). Anthracnose of the Vine* The mycelium of this fungus can penetrate into leaves, green bark, or fruit, and kills the tissues. Spots are first pro- duced, then enlargement of the neighbour- ing tissues takes place, causing the spots to appear as if sunk in depressions, and reminding one of hail-wounds. On leaves and grapes, the spots are sharply defined, | Fis. = BR ge at first dark-brown, later with greyish eateries peer ct Rit centres and dark-brown margins. In the later stages the dried-up spots may drop out of the leaves. Anthracnose, or “ birds-eye rot,” constitutes one of the dreaded vine diseases of America and Europe, so that it has received much attention, both as to its life-history and remedial measures ; as yet however with but partial success. Copper sulphate solutions seem to be fairly successful remedies, as shown from results of the many experiments recorded in the Journal of 'Kiihn, Hedwigia, 1877, p. 121; also in Rabenhorst’s Fungi euwrop. No, 2261. Frank, Zeitsch. fiir Pflanzenkrankheiten, 111., 1893, p. 28. Eriksson (Mitthl. der k. Landh, Akad. Stockholm, 1890) records a case of this kind, but the conidia are drawn two-celled as in a Diplodina, whereas Kihn’s original drawings have only one-celled conidia. *Cavara et Eriksson, Zeitsch. (. Pflanzenkrankheiten, 111., p. 23. ‘Goethe, Mitthl. ub. den. schwarzen Brenner d. Reben, Leipzig, 1878. Cornu, Bullet. de la Soc. botan. de France, 1878. Prillieux (idem), 1879. Rathay, ‘* Ver Black-rot,” 1891. Scribner, Report of U.S. Amer. Dept. of Agriculture, 1886. 468 FUNGI IMPERFECTI. Mycology and the bulletins and reports of the American experi- mental stations. (Compare also (Gloeosporium ampelophagum (Pass.) p. 484.) Ph. betae Frank. The younger leaves of well-developed beet-root become black, and the disease extends into the root. Mycelium fills the diseased parts and penetrates into healthy tissues. Pycnidia are developed on the diseased spots. The fungus is no relation of Sporidesmium putrefaciens, a form to which a root-rot is ascribed by Frank. It is however probably identical with Prilleux’s Phyllosticta tabifica. Kriiger found the disease so common, that in many localities as many as 80 per cent. of the plants were destroyed. Sorauer regards the root-rot of beet to be sometimes caused by Phoma, sometimes by Sporilesmium, perhaps in some cases by both together. Ph. sanguinolenta Rostr.? attacks carrot plants in their first year, causing greyish-brown depressed spots on the bulbs with- out however appearing to be very injurious to them. When the seedlings are planted out in spring, the mycelium extends into the stem and causes the umbel to wither at flowering so that no seed is formed. Pyenidia are developed from all attacked spots and give off conidia as red tendril-like bodies—hence the species-name. Certain varieties of carrot appear to resist attack by this parasite better than others. Ph. solani Hals. This causes damage to the egg-plant (Solanum melongena).* Young plants die off on the hot-beds, their stems dying near the earth and shrivelling up. The pycnidia of this Phoma appear on the killed parts. Ph. cydoniae Sacc. has been reported as injurious to quince trees. (U.S. America.) Many species cause leaf-spot diseases. Some of the more important British and American species are: Ph. pinastrella Sacc. On Pinus sylvestris and others. (Britain.) Ph. strobi (B. et Br.). On Pinus Strobus. (Britain.) Ph. taxi (Berk.). On yew. (Britain.) Ph. Candollei (Berk. et Br.). On box. (Britain and U.S. America.) Ph. sorbi (Lasch.). On leaves of Pyrus Aucuparia, (Britain.) 1Rrank, Zeitsch. f. Pflanzenkrankheiten, 11., p. 90, and Deutsche landw. Pr. No. 89, 1893. Kriiger, Zeitsch. f. Pflanzenkrankheiten, tv., 1894, p. 195. * Described and figured by Halsted in Bulletin 91, N. J. Agric. Haper. Station. 1892. PHOMA. 469 Ph. malvacearum West. On mallows and hollyhock in Europe. Ph. longissima (Pers.). In species of Umbelliferae and Chenopodiaceae in Europe and America. Ph. errabunda Desm. In stems of Verbascum. (Britain.) Ph. cucurbitacearum (Fr.). On fruits of various species of Cucur- bitaceae in Europe and America. Dendrophoma. Pyenidia similar to Phoma, conidiophores however bearing several conidia either on branches or little processes Dendrophoma Marconii Cav. attacks Hemp (Cannalis sativa), causing dark oblong spots on the green stem. The pycnidia are embedded and break through the epidermis with a round pore, The conidiophores are branched, with swollen ends varrying little short rod-like unicellular conidia. In case of attack, which generally occurs towards the close of the vegetative period of the hemp, it is suggested to cut the crop somewhat prematurely, and thereby prevent maturing and spreading of the fungus. co D. convallariae Cav. produces dark elongated spots on leaves of Con- vallaria majalis, D. valsispora Penz. is recorded by Penzig on living leaves of Citrus Limonum (Lemon). Sphaeronaema. Pyenidia embedded, membranous, and long-beaked. Conidia ovoid or oblong, unicellular, and almost colourless. Sphaeronaema fimbriatum (Ell. et Hals.), (Ceratocystis Jjimbriata Ell. et Hals.). Black rot or black shank of sweet potato.' The parasite shows itself as black depressed spots on the lower parts of young plants, and these may extend over the whole shoot. The disease is best recognized on the tubers, where it consists of dark, somewhat greenish spots, varying from } to 4 inches in diameter, and extending some distance into the tissue. These spots when once seen cannot be mis- taken, as they are sunk areas with distinct margins, like spots burned into the potato with a piece of metal which has left the skin uninjured. The mycelium consists of thick-walled olive-brown hyphae, which cause death and destruction to the 1 Halsted and Fairchild, Jour. of Mycology, Vol. vu., 1891, with Figures. 470 FUNGI IMPERFECTI. cells of attacked tissues. There are three modes of spore pro- duction: (1) brown macro-conidia inside the tissues; (2) colour- less micro-conidia on the spots; (3) spherical pyenidia with long necks ending in a fringed opening. A sclerotial form is also strongly suspected. Remedial measures recommended are, de- struction of all diseased parts, change of crop on diseased fields, and selection of healthy seed and strong sprouts. Several other species of this genus are recorded from North America, but details in regard to their mode of life are wanting. Asteroma. Fungi forming star-like, dark-grey, mycelial patches on the surface of plants. Pyenidia very small and containing tiny ovoid or short cylindrical spores. Several species frequent living leaves.’ Asteroma impressum Fuck. On Tussilago farfara. A. prunellae Purt. On leaves of Prunella vulgaris. (Britain.) A..ulmi Klotsch. (Britain), and A. maculare Rud. On U/mus. A. padi (D.C.) causes a leaf-fall on Prunus Padus. (Britain.) A. geographicum Desm. is found on the leaves of species of Crataegus. Prunus, and Pyrus in Europe and America. Pyrenochaeta. Pyenidia emergent or sessile, beset with bristles. Conidia oblong, on branched conidiophores. Pyrenochaeta rubi-idaei Cav. forms black spots on leaves of Rubus Idaeus. The pyenidia are spherical with a tuft of bristles projecting from their terminal pore. The conidia are little, oval, and with one or two cells; they are produced from shehtly branched conidiophores. Vermicularia. Conidia unicellular, rarely bicellular, generally spindle-shaped ; they are produced inside pycnidia, and are embedded amongst brown septate hairs. The species are a frequent cause of leaf- spot, but most of them have not yet been sufficiently investigated. 'Cicinobulus Cesatii De Bary, allied to this genus, is a parasite on Oidium Tuckeri, the dreaded vine-parasite. VERMICULARIA. 471 Vermicularia trichella Fr. occurs on living leaves of ivy and other plauts. (Britain.) V. ipomoearum Schw. On species of Zpomoea in America, V. microchaeta Pasc. On living leaves of Camellia japonica in Italy. V. circinans Berk. Onion rot in Britain and U.S. America. Placosphaeria and Cytospora are genera containing forms parasitic on living plants, but of little practical importance. PHAEOSPORAE. Coniothyrium. Pycnidia brown or black. Conidia brown, unicellular, spheroid or ovoid, and borne on short conidiophores. Coniothyrium (Phoma) diplodiella Sace.! White-rot of the vine. This disease has a wide distribution in Hungary, and has also been observed in France, Italy, and America. It has caused considerable damage, especially in Northern Italy, where it was for a long time regarded as the black-rot. According to Mezey, this parasite is distinguished from Laestadia (black-rot) in the following points:—The pyenidia and conidia are larger; the mature pycnidia are greyish or light brown (never black), the mature conidia are brownish. The disease attacks the fruit only, causing it to fall off. Riathay, however, states that it also attacks young shoots, infection taking place from the fruit. Diseased grapes become soft, rotten, and wrinkled; the ridges are beset with pycnidial pustules, as in black-rot, but the grapes never become brittle and hard. Viala and Ravaz? have recently succeeded in rearing perithecia from twigs and fruit-stalks set in sterilized moist sand. None could be found on grapes. The perithecia are globular, enclosed in a black covering several cells thick, and with a large crater- like aperture.’ The asci and paraphyses arise only from the depth of the perithecium, the latter being longer than the former and frequently branched. The asci are club-shaped and short-stalked, and contain eight spindle-shaped colourless or yellowish asco- spores, divided by one to three cross-septa. They germinate and produce one or more geri-tubes. 'Rathay, ‘‘ Der White-Rot,” Die Weinlaube, 1892. General description in Report 9, New York Agric. Exper. Station, 1890, *Viala and Ravaz, Compt. rend., CX1Xx., 1894, p. 443. 472 FUNGI IMPERFECTI. A new genus Charrinia, belonging to the Sphaeriaceae of the Ascomycetes, has been formed to receive this species. Sphaeropsis. Pyenidia black and spherical, with an aperture. Conidia ovoid or oblong, unicellular, dark-coloured, and on stalk-like conidio- phores. Sph. malorum Peck. The cause of a disease in America, known as the black-rot of apple and quince. The mycelium permeates and destroys the skin of the fruit, which, in con- sequence, becomes dried up and mummified. It also occurs in Britain. Other species attack plants of various Rosaceae. PHAEODIDYMAE. Diplodia. Pyenidia, small, spherical, and dark-coloured; the conidia are two-celled when mature. Diplodia gongrogena Temme.’ Temme discovered a mycelium and the pyenidia of this Diplodia in aspen (Populus tremulae) exhibiting hypertrophied outgrowths of wood and rind. As yet it has not been possible to artificially produce these malformations on the aspen, nor other somewhat similar ones which occur on the willow. Other species of this genus attack many trees, eg. holly, lilac, horse-chestnut, mulberry, and various conifers. HYALODIDYMAE. Ascochyta. Conidia ovoid or oblong, bicellular, and hyaline. The pycnidia have a central aperture, and are embedded in discoloured portions of leaves or twigs. The following species are of practical importance : Ascochyta pisi Lib. (Britain). Briosi and Cavara state that 'Temme, Landwirth. Jahrbuch, 1887. Thomas, Verhand. d. botan. Verein d. Prov. Brandenburg, 1874. ASCOCHYTA. 473 this fungus is injurious to Pisum sativum, Phaseolus vulgaris, o ? ‘ Vicia sativa, ete. It causes spots on leaves and pods, followed by , } drying up of the former and deformation of the latter. The pycnidia appear as tiny points on the spots, and give out bicellular eylindrical conidia. As. Boltshauseri Sace.' This species was first observed in Switzerland on bean (Pha- seolus vulgaris). Leaves of all ages become brown-spotted, and premature defoliation may follow. The spots are marked by concentric zones, and bear pyenidia. The conidia are two- to three-celled, being distinguished in this and by their larger size from the preceding species. The following species fre- quent living leaves: Ascochyta tremulae Thiim. On the aspen. Fic. 29§.—Ascochyta pisi on Pea. Enlarged section of pycnidia, and still more enlarged conidia. (v. Tubeuf del.) A. metulispora, B. et Br. On leaves of the ash in Scotland. Armoracia rusticana (Horse-radish). ( Britain.) A. armoraciae Fuck. On (Britain.) A. periclymeni Thiim. On Lonicera Periclymenum. A. maculans Fuck. On Hedera Helir. A. Ellisii Thiim. On Vitis Labrusca in America. A. brassicae Thiim. On Brassica oleracea. A. dianthi (A.8.). On Dianthus. A. pallor Berk. On Rubus Idaeus. (Britain.) A. viciae Trail. On Veta sepium, ete. (Britain.) A. malvicola Sace. On Molva sylvestris. (Britain.) A. graminicola Sace. On various grasses. (Britain.) A. scabiosae Rabh. On Anautia arvensis. A. nicotianae Pass. On Nicotiana Tabaceum. A. digitalis Fuck. On Digitalis purpurea. A. United States, fragariae Sace. has been found injurious to the strawberry crop in the A. aspidistrae given (Gardener's Chronicle, Xvil., 1895) as a parasite on Aspidistra in Britain. ! Boltshauser-Amrisweil, Z itschrift a; Phlanzenkrankheiten, 1.5, D. 135. 474 FUNGI IMPERFECTI. Actinonema. Pyenidia small and situated on a gossamer net of mycelium. Conidia hyaline and divided by one or more cross-septa. Actinonema rosae Lib. (Asteroma radiosum Fr.). This pro- duces black radiating spots on rose-leaves, on which pycnidia with bicellular conidia are developed. A premature defoliation takes place, which in turn causes the upper buds to unfold in autumn before their time. The mycelium is distributed both inside the leaves and superficially. Timely removal of diseased leaves and defoliated shoots might be recommended as remedial measures. A. tiliae Allesch. shows itself in spotting of the leaves and petioles of lime, and may bring about defoliation of the whole tree. A. fagicola Allesch. produces white spots with dark margins on living beech leaves, and causes gradual discoloration of the whole leaf. According to Allescher,' this disease brings about premature defoliation of beech. As yet it has been observed only in Upper Bavaria. A. fraxini Allesch. On living leaves of the ash. A. crataegi Pers. attacks leaves of Pyrus Aria, P. torminalis, and Viburnum Opulus. A. podagrariae Allesch. On living leaves of Aegopodium Podagraria, and Chaerophyllum hirsutum. Darluca. Darluca genistalis (Fr.). On living leaves of Cytisus sagittalis. This may, however, be only a parasite on Uromyces cytisi with which it is frequently observed, just as Dar. filum occurs on several Uredineae. Diplodina. Similar to Diplodia, but having colourless conidia. D. castaneae Prill. et Delac.* produces canker-spots on the stems of chestnut, and brings about death. PHRAGMOSPORAE. Hendersonia. Pyenidia formed under the host-epidermis, which is later ruptured. Conidia brown, two- or more-celled. 1 Allescher, Hediwigia, 1894. 2 Prillieux et Delacroix, Bull. soc. mycol. de France, 1893. HENDERSONIA. 475 Hendersonia foliicola (Berk.) (Britain and Europe). ‘The black globular pycnidia are produced superticially on leaves of Juniperus communis. The conidia are elliptical in shape, three- to five-celled, and abjointed from filamentous conidiophores. (This species is not identical with Podiosoma Juniperi 8 minor Corda, which is more like the needle-frequenting form of Gymno- sporangium juniperinum.) Several species are found on living leaves : H. cydoniae C. et Ell. on quince in America. _H. mali Thiim, on apple. H. rhododendri Thiim, on Ahododendron hirsutum in Northern Italy and Germany. Cryptostictis. Similar to Hendersonia, but having ciliate spores. Cr. cynosbati (Fuck.). Sorauer” regards this as parasitic on Rosa canina, and causing death of portions of the rind. Stagonospora, Couturea, Asteromidium, and Camarosporium contain species said to frequent living leaves of various plants. SCOLECOSPORAE. Septoria. Spores generally multicellular and hyaline; produced from short conidiophores, contained in lens-shaped embedded pyenidia. Septoria parasitica Hartig.* This disease may be frequently observed in young plantations and seed-beds of Spruce. The symptoms are very like those following damage by frost, brown needles appearing in May towards the base or middle parts of young shoots, and followed by a premature needle-cast. The disease is most apparent on lateral shoots, which become sharply bent downwards, the green needles hanging limply till they wither and fall as the whole shoot shrivels up. The pyenidia are little, black, and spherical; they are pro- duced during the summer, particularly towards the lower end 'K. v. Tubeuf, ‘‘ Generations-wechsel Gymnosporangium-Arten,” Cenfralbl. f. Bakteriologie u. Parasitenkunde, 1891. * Handbuch d. Pflanzenkrankheiten, u., 1876, p. 388. 3R, Hartig, Zeitsch. f. Forst. u. Jagd-wesen, 1890; and Forstlich-naturwiss, Zeitschrift, 1893. 476 FUNGI IMPERFECTI. of the shoots, and either rupture the epidermis or grow out from the leaf-scar cushions (Fig. 297). The conidia are abjointed from filamentous conidiophores inside the pyenidia, and emerge as tendril-like structures. They are two-celled, small, cylindrical, and pointed at both ends. Germination takes place easily in water, and the disease spreads rapidly over the young developing shoots during May. The mycelium permeates the twig, living both imside the cells and between them. The disease has _ been observed on Picea excelsa and P. Menziesvi, not only in nurseries and on young trees, but also in pole- forest, where it frequents the upper crown and causes Fic. 296.—Seploria parasiticu. a, Young diseased death. At the beginning shovt of Spruce, with apex still fresh and green. of an attack the pru ning b, Needle diseased towards base. c, Apex of a two- year-old shoot, into which the disease has extended it sec igs j 4 o backwards from the younger shoot; the brown of diseased ties Wy Cue Se an ee is indicated by plantations should be at- tended to. Septoria rubi (Westend.).' Blackberry leaf-spot. This is a parasite of some economic importance in the United States, where it interferes with the blackberry culture. It also occurs in Europe and Britain. S. ribis Desm. produces a somewhat similar disease on leaves of currant and gooseberry.” (Britain and U.S. America.) S. piricola Desm. occurs throughout all Europe, causing little greyish spots on leaves of pear trees. It is probably a pyecnidial form of Sphaerella lucillae Sace. S. crataegi Kich. A common species on leaves of Crataegus in Europe. S. cerasina Peck. On leaves of Prunus serotina in the United States. Many forms of Septoria infest cultivated vegetables : S. petroselini Desm. is the cause of dry spots appearing on leaves of 1 Description in Bulletin, No. 6, Ohio Agric. Exper. Station, 1891. * Description in Bulletin, No. 13, Zowa Agric. Exper. Station, 1891. SEPTORIA. 477 cultivated parsley in Europe and Britain. A variety (ap7/ Br. et Cav.) is an enemy of celery in the United States. S. armoraciae Sacc. On horse-radish in America. S. consimilis Ell. et Mart. frequents lettuce in America. S. lycopersici Speg. This parasite, originally observed in America, has recently been de- scribed by Briosi and Cavara on tomatoes in Italy. It causes spots on leaves, stems, and fruits, inflicting thereby con- siderable loss on cultivators. The following are im- portant forms on _ other cultivated plants : 8. graminum JDesm. causes light spots on leaves of wheat, oats, and grasses. It has been observed to injure the cereal crop in Italy. It is recorded for Britain and U.S. America. S. cannabis (Lasch.). This on leaves of hemp pro- duces spots, which are at first whitish, then yellowish with dark margins. The pycnidia are embedded in the upper side of the leaf. The following species have caused injury to garden plants: S. dianthi Desm. Car- fs ‘ Fic, 297.—a, Leader-shoot of Spruce, partially nation-spot.~ The disease killed and defoliated. 4, Pyenidia emerging from the rind and the leaf-scars(x 5). ¢, Formation appears on the leaves and of conidia inside a pycnidium (x 240). d, Spores germinating in water. ¢, Spores germinating in stems as rounded spots of nutritive gelatine. (After R. Hartig.) 'Cavara (Zeitsch. f. Pflanzenkrankheiten, 11., p. 23) regards this and S. tritici with its varieties, as forms of a single species; also Eriksson (Om Ndgra sjukdomar i odlade Vixter, 1890). * Atkinson, ‘‘ Carnation Diseases,” at American Carnation Society, 1893, 478 FUNGI IMPERFECTI. dirty white or brownish colour with a darker margin. The pycnidia appear as black points on the spots, and rupture the epidermis before giving off their septate spores. . anemones Desm. On Anemone. (Britain.) . lychnidis Desm. On Lychnis diurna. (Britain.) . epilobii West. On Lpilobium. (Britain.) . Stachydis D. et R. On Stachys. (Britain.) . urticae D. et R. On Urtica dioica. (Britain.) cyclaminis Dur. et Mont. This produces roundish spots with concentric markings on the leaves of Cyclamen which then gradually NNNNNYD wither. S. chrysanthemi Cav. causes a leaf-spot on Chrysanthemum japonicum and C. indicum. S. exotica Speg. attacks cultivated New Zealand species of Veronica. S. hydrangeae Bizz. causes injury to cultivated Hydrangea. S. sedi West. injures Sedum under cultivation in the United States and Britain. Other species on many other herbs in Britain and America. Many species of Septoria have been recorded on trees and shrubs, e.g. : rosae Desm. On roses. (Britain.) . hederae West. On ivy. (Britain.) . fraxini Desm. On the ash. (Britain.) nigro-maculans Thum. On green walnuts, stunting their growth. . castaneae Lev. On the sweet chestnut. . aesculi (Lib.). On the horse chestnut. (Britain.) . pseudoplatani Rob. et Desm. On leaves of sycamore. . populi Desm. On leaves of poplar. . didyma Fuck. On Salix triandra and 8. alba. . cornicola Desm. On leaves of Cornus sanguinea. NNNNNHNHHYN Phleospora. True pycnidia are not formed, but the conidia are abjointed from cavities in the stroma; they are hyaline, rod- or spindle- shaped, and consist of two or more cells. Phleospora aceris (Lib.). On living leaves of Acer Pseudoplatanus. (Britain. ) Phil. mori (Lev.). On living mulberry leaves; probably related to Sphaerella mort. (Britain.) Phi. ulmi (Fr.). On living leaves of elm. (Britain and America.) Phl. oxyacanthae (K. et 8.). On living leaves of Crataegus Oxyacantha. (Britain. ) DILOPHOSPORA. 479 Dilophospora. Dilophospora graminis Desm. (Britain.) This attacks rye, wheat, and various grasses. Oblong light spots are produced and bear the pyenidia; when these occur in the flower heads, stunting of the grain takes place. (See also Dilophia, p. 222.) 2. FAM. NECTROIDEAE. The fungi of this family are chiefly pycnidial forms of the Ascomycetes, and as such have already been considered. 3. FAM. LEPTOSTROMACEAE. HYALOSPORAE. Leptothyrium. Pyenidia black and discoid. Spores ovoid or spindle-shaped, unicellular, and hyaline. Leptothyrium periclymeni (Desm.). On living leaves of species of Lonicera. (Britain.) L. alneum (Lév.) produces roundish leaf-spots on species of Alnus. (Britain and America.) L. acerinum (Kunze) causes spotting of the leaves of Acer campestre and A, platanoides. (Britain.) Several other species occur both in Europe and America. Melasmia. The black pyenidia occupy black extended stromata. Conidia simple and unicellular, borne on rod-like conidiophores. Melasmia berberidis Thiim. et. Wint. On living leaves of barberry. Brown spots are produced, bearing the pycnidia as black points; the spots cause total or partial death of the leaves, frequently ending in defoliation of the shrubs. M. empetri Magn. (Britain.) This species was observed by Magnus! causing an epidemic disease on crowberry. The symptoms were abnormal elongation of young twigs, and the leaves remained smaller than usual. The rind of the stem was found to be permeated by a mycelium which produced black | Berichte d. deutsch. botan. Gesell., 1886. With illustrations. 480 FUNGI IMPERFECTI. pustules bearing the pyenidia of this Melasmia. The cells of the cortex dried up, and the rind became detached from the wood in the following year. The leaves were never found attacked. (The species is not a Rhytisma; nor does any species of Rhytisma produce similar hypertrophy of its host.) M. acerina, M. punctata, and M. salicina are now recognized only as pyenidia of the species of Rhytisma bearing these same specific names. Leptostroma. Pyenidia oblong, black, and flattened. Conidia ovoid or oblong, unicellular, and whitish. Leptostroma punctiforme Wallr. Found on the leaves of Salix, Rosa, Buxus, Euphorbia, ete. L. caricinum Fr. frequents leaves of Carex and Eriophorum in Europe and North America. The parasitism of other species is uncertain. Labrella and Discosia are genera whose species have not as yet pro- duced diseases of any serious economic importance. PHRAGMOSPORAE. Entomosporium. Entomosporium maculatum (D.C.) Lev.t This fungus, under the name of leaf-blight of pear and quince, is the cause of considerable loss in the cultivation of these crops. Defoliation takes place early in the season and young seedlings are forced to form a new set of leaves, whereby their reserves of food are exhausted. If this be repeated several times the plants become exhausted and are killed off in winter. Stocks already budded seem to remain immune if not already diseased. The leaves are first attacked, but later the succulent growing apex of the twigs may also succumb. The parasite hibernates on the bark in small depressions containing the pycnidia; thence it spreads in early spring, so that pustules appear on the young leaves before they are fully developed. Spraying with Bordeaux mixture, or solution of copper acetate ‘Galloway and Southworth (Report for 1888 of Section of Vegetable Pathology, Dept. of Agriculture, U.S. America) give a historical account of the fungus and a bibliography. (Kdit.) ENTOMOSPORIUM, 481 (6 oz. in 26 galls. water) have both produced good results in checking the disease.! E. mespili (D.C.). (See Stigmatea mespili, p. 210.) SCOLECOSPORAE. Brunchorstia. Brunchorstia destruens Erikss. (2. pini Allesch.). In Nor- way almost all the plantations of Austrian black pine (Pinus Laricio) from five to thirty years old have become diseased and died out. Similar ravages have also been observed in Germany. Brunchorst ascribes this to a parasitic fungus whose mycelium may be found in all parts of diseased twigs and needles, and whose pyenidia are formed on the killed remains. The disease begins in young first-year twigs, the mycelium growing in the cortex, pith, and wood. The needles are attacked in summer, become brown from the base upwards, and the pyenidia make their appearance under the scale-leaves. Brunchorst ? describes the fungus as follows: Pyenidia par- tially embedded in the tissues of the host-plant; the smaller ones being simple, the larger divided by complete or partial partitions. The inner wall as well as the partitions of the pyenidium are closely beset with straight basidia, from the apices of which stylospores with two to five septa are abjointed. Paraphyses are never present. The perithecia are black, oblong or rounded, slightly grooved, and 1-2 mm. in diameter; they dehisce by one or more irregular pores in the wall. The spores are very minute (30—40=5,), tapering, and rounded at each end. Schwarz considers Brunchorstia as a conidial form of Cenan- guim abietis already described (p. 251). It may be here mentioned that drying-up of pine-twigs may be due to heating by the sun in frosty weather, or to frost itself;* these are, however, quite distinct from the disease just described. ‘Fairchild (Journal of Mycology, Vol. vit.) gives results of treatment with various fungicides on several varieties of pear and quince. (Edit. ) *** Ueber eine neue Krankheit d. Schwarzfohre.” Bergens Museum, 1889. *R, Hartig, ‘‘ Vertrocknen u, Erfrieren d. Kiefernzweige,” Forstlichnaturwiss. Zeitschrift, 1892 and 1895, 9 2H 482 FUNGI IMPERFECTI. 4. FAM. EXCIPULACEAE. The parasitic nature of the species of this family has not as yet been investigated to any extent. Il. MELANCONIEAE. True pyenidia are not formed, but the conidia are developed in clusters or aggregations covered over at first by the epi- dermis of the host-plant, which is ultimately ruptured. HYALOSPORAE. Gloeosporium. Conidial clusters colourless or grey, never black; they rup- ture the overlying epidermis and give off unicellular conidia, one from each conidiophore. Gloeosporium fructigenum Berk.’ (Britain and U.S. America). Apple Rot or Ripe-rot. This is a very serious disease for American cultivators. It not only attacks apple, but also the grape, pears, peaches, and egg-plants.” On the apple it appears first as brown spots which become more conspicuous as the fruit enlarges. The spots on first sight look like decay, but they are quite firm and soon bear pustules of a white or pinkish colour turning to black. The attacked part of the apple has an intensely bitter taste, and should be carefully removed before eating the fruit. On grapes the fungus produces tiny raised pustules, which on the white varieties are situated on spots with a purple centre and a brown margin; the pustules When mature give off flesh-coloured conidia. The grapes gradually shrivel up, but do not become black as in the case of the black-rot, nor do they assume a bitter taste as the apples do. The apple bitter-rot makes rapid progress amongst stored fruit, especially before it has been sorted out. Care should therefore be taken that diseased apples are removed as soon as possible. The spraying of trees bearing young fruit with copper car- 'Southworth, Journal of Mycology, vi., p. 164. ’ Halsted, Bulletin of the Torrey Club, 1893, p. 109. Massee, Gardener's Chronicle, Vol. xtv., 1893. GLOEOSPORIUM. 483 bonate or potassium sulphide solutions has good effects on the yield of the orchards. In vineyards under treatment for black- rot or mildew, there is little chance of the ripe-rot fungus appearing. It is probable that the species known as Gi. phomoides Sace. on tomato, Gl. piperatum E. et E. on peppers (Capsiewm annuum), and Gl. melangeae E. et Hals. on the egg-plant, are identical with Gl. fructigenum. At least they very much resemble each other, even on their widely differing substrata, and cross-infections have been carried out. Gl. venetum Speg. (G/. necator Ell. et Ev.) Anthracnose of raspberry and blackberry.’ This disease appears on both canes and leaves. On the young shoots it produces small reddish- purple spots during early summer; ‘as the season advances the spots run together into irregular blotches of more or less greyish colour with a dark purple margin. ‘The ripening fruit remains small and shrivels up. . Leaves may also bear spots, but they more frequently remain smaller and have an unhealthy look. The conidia are at first enveloped in a thin covering, which becomes gelatinous when wet, so that they escape. The mycelium is believed to perennate in stems or decayed remains, and so to carry the parasite from season to season. Owing to the delicate nature of raspberry foliage, fungicides must be used with great care. Dilute Bordeaux mixture is said to be safe and beneficial. The burning of diseased canes should certainly be carried out each autumn. Gl. ribis (Lib.). This attacks currant bushes throughout Europe and America in much the same way as Gi. venetum. The leaves wither and fall, so that the fruit-crop suffers. ( Britain.) GL amygdalinum Brizi. This has recently been described as destructive to almond cultivation in Italy. The mycelium inhabits twigs and fruits, and gives off tufts of conidiophores bearing conidia; as a result, wounds are produced in the epidermis and stunting of the host-tissues takes place. Gl. rosae Hals. is described as injurious to rose-culture in America. It may be identical with some of the species of Glocosporium already mentioned as frequenting Rosaceae. ‘U.S, America Dept. of Agriculture, Report for 1889, contains a good account, *Brizi, Zeitsch. f. Pflanzenkrankheiten, 1896, p. 65, 484 FUNGI IMPERFECTI. Gl. ampelophagum (Pass.)' Black-rot of the vine. This disease is very injurious and has a wide distribution in Europe. It is known under many names such as “ Pock, Brand, Rost, Jausch, Brussone, and Nebbia nera,” though probably these names include several distinct diseases. The identity of this Glocosporium is somewhat uncertain, and it may really be identical with Phoma ampelinum (p. 467). Rathay ascribes the black rot to Sphaceloma (Phoma) ampelinum, while Thiimen regards Gloeosporium as the cause. Briosi and Cavara consider the two species of fungi as distinct. Thiimen says that the patches of Gloeosporium are for a considerable time disc-like and of a light-grey rose colour; those of Phoma, on the other hand, are always depressed and brown. Rathay, however, de- scribes the spots of Phoma as at first dark brown, and later ashy grey with a brown margin. The spots appear on green parts of the vines during April and May. Those on the leaves frequently fall out, leaving holes. On the grapes the spots are smaller and produce a brown coloration extending deep into the fruit. The conidia are small, hyaline, oval, and unicellular; they are abjointed from very short conidiophores arranged in little clusters. The conidial patches rupture the host-epidermis, and the conidia are liberated. Thiimen suggests that the soil of vineyards should be kept well cleaned, and that the stake mode of culture be used in pre- ference to an overhead trellis; he also recommends the washing of all parts of suspected vines during winter with 10 to 15 per cent. solution of sulphate of iron. This treatment is said to have been very beneficial in keeping many vineyards quite healthy and free from fungi. Gl. nervisequium.” This parasite occurs on species of Platanus in Europe and America. Brown spots appear on the leaves, especially on the veins; these as they extend cause sudden withering and fall of the leaves. Pustules containing a stroma develop on the spots, and unicellular, ovoid, hyaline conidia are abjointed from club-shaped conidiophores. 'Thiimen, Die Pocken an Wein u. Obst. 1885; Die Bekiimpfung d. Pilzkrank- heiten, 1886; De Bary, Annalen d. Oenologie, 1v.; Viala, Les maladies de la Vigne ; Briosi e Cavara, Punghi parasit., u1.; E. Rathay, ‘‘ Der Black-Rot,” 1891. 2 U.S. America Department of Vegetable Pathology, Report for 1888, gives a general account of this disease. GLOEOSPORIUM. 485 Several fungi of very near relationship, if not actually identical, occur on Platanus.' All cause considerable disfiguration of the foliage, so that a systematic destruction of all youn branches is strongly recommended. Gl. cingulatum Atks.” This is the cause of Anthracnose on Privet (Ligustrum vulgare) in the United States. The following is Atkinson’s diagnosis: “ Affected areas light brown, either oblong on one side of the stem or completely girding it. Acer- vuli 100 to 150 in diameter, rupturing the epidermis, in age black from the dark stroma lying in the base or extending irregularly up the sides, frequently forming a pseudopyenidium. Basidia numerous, crowded, simple, hyaline, or when very old perhaps faintly fuligious. Spores oblong, or elliptical, straight or little curved, usually pointed at the base. From pustules on the stem they measure 10-20 by 5-7; in artificial cultures they are frequently much larger, but when crowded in the media, or when the nutrient substances are nearly exhausted, they may be considerably smaller. On stems of Ligustrwm vulgare. “This is quite distinct from Glocosporium ligustrinum Sace.” Many species of Glocosporium frequent broad-leaved trees and cause more or less injury to the foliage. Gl. rhododendri Br. et Cav. attacks the leaves of outdoor cultivated rhododendrons in autumn, or indoor species in winter. Large yellow spots marked with concentric zones are formed, and bear the pycnidia; finally the leaves dry up and fall off. Gl. violae B. et Br. attacks violets in Britain and U.S. America. Gl. vanillae Cke. et Mass. (Calospora vanillae Massee.*) This causes a dangerous disease on Vanilla planifolia and other Orchideae in Mauritius and other parts of the tropics. Death is brought about by the Gloeosporium (Hainsea) form of the fungus, the higher reproductive organs only appearing when the leaves are killed. Other species are known, but their economic importance is not diseased or =] great. 'y, Tavel, Botan. Zeitung, 1886; Leclere du Sablon, Revue gen. de Botanique, 1892. * Atkinson, ‘‘A New Anthracnose of the Privet,” Cornell Univ. Agric. Kape a Station Bulletin, No. 49, 1892. * Massee, Kew Bulletin, 1892, p. 111. 486 FUNGL IMPERFECTI. Myxosporium. Conidia ovoid, hyaline, and abjointed from rod-shaped basidia situated in cavities of the cortical tissues of arboreous plants; a true pyenidium is not formed, and the reproductive mycelium is only covered over by the epidermal layers of the host. Myxosporium devastans [ostr.! is said to attack and kill young twigs of Betula verrucosa. The conidial patches are developed in the killed rind, and give off unicellular colourless conidia. M. carneum Lib. is parasitic on twigs of beech. M. laneola Sacc. et Roum. causes death of oak-twigs. The other known species have as yet been observed only as saprophytes. Colletotrichum. Conidial patches surrounded by setae; characters very like Glocosporium. Colletotrichum Lindemuthianum (Sacc. et Magn.) This disease, first observed by Lindemuth in 1875, has assumed ereat importance as a disease of the kidney bean (Phaseolus vulgaris) both 11 Europe and America. Young pods are most frequently attacked, but neither stems nor leaves are exempt. The pods show brown depressed spots with a distinct margin. The unicellular and oblong conidia are given off from short conidiophores developed on the spots. Germination takes place at once, the germ-tube forming an adhesion-dise on the host- epidermis, and from this a hypha penetrates into the tissues to develop into a brown mycehum. Frank obtained brown spots and mycelium on young beans twenty-four hours after infection. C. Lagenarium (Pass.) (C. oligochaetum Cav.). This parasite is very injurious to seedlings of water melon (Cucumis citrullus), melon (C. Melo), and the gourd (Cucurbita Lagenaria). Leaves and fruits may be attacked, but it is the cotyledons and stems of the seedling plants which most frequently fall a prey. Spots 'Rostrup, Tidsskrift f. Skovvaesen, 1893. *For the relationship of this with the following species, as well as their synonomy, see Halsted in Bulletin of Torrey Botanical Club, 1893, p. 246. Description, treatment, and bibliography by Beach, ‘‘ Bean-spot disease,” Genera N.Y. Exper. Station Bulletin, No. 48. COLLETOTRICHUM. 487 appear on the leaves, and depressions on the stem, sometimes extending so far round that the whole shoot dries up. The conidial patches are very much the same on the different hosts, and consist of short conidiophores from which oval, unicellular, hyaline conidia are abjointed. C. lycopersici Chest. is the cause of a spot-disease on the fruit of tomato in the United States. C. spinaciae Ell. et. Hals. causes a destructive disease on cultivated spinach. C. malvarum Br. et Casp. (C. althacae Southw.') produces a disease of cultivated hollyhock. It is most injurious to the seedling plants, and has caused great loss in America and Sweden. The fungus may attack any organ, and produces spots which enlarge so rapidly that death of the host may result. C. gossypii Southw.2 Anthracnose of Cotton. This disease, although it may be found on stems and leaves, is most frequent and most conspicuous on the fruits or “bolls” of the cotton-plant. The first signs are tiny depressed spots yey o ey eer of a reddish-brown colour, and as these Bey, Enlarged pustule and enlarge they cause blackening of neigh- bouring tissue. When the spores are developed the spots become dirty grey, or perhaps pinkish if the spores are present in large numbers. Fruit attacked in this way does not mature well, and the yield of cotton is greatly prejudiced. Atkinson found the cotyledons easy to infect with the disease. The spores are oblong and tapering, with a shallow constriction in the middle; they are borne either on short colourless basidia or on long, olive-coloured, septate setae, both kinds of conidio- phore being produced in acervuli or patches. Rk RNS Allon C. adustum Ell. is the cause of a leaf-spot on orange in Florida, 1 Southworth, ‘‘ A New Hollyhock Disease,” Journa/ of Mycology, Vi., S890. * Southworth, Journal er Mye ology, Vi., 1890, p. 100. Atkinson, Alabama Agric. Exper. Station Bulletin, No, 41, 1892. 488 FUNGI IMPERFECTI. Faded spots appear on the leaves, becoming later greyish brown dotted over with minute black points, the conidial patches.! C. ampelinum Cav. causes little dry spots on the leaves of vine, fre- quently in such numbers that the whole leaf dries up. C. kentiae Hals. attacks palm-seedlings so that their leaves do not unfold. C. cyclameneae Hals. occurs on Cyclamen. ScoLEco-A LLANTOSPORAE. Cylindrosporium. The white and_ shining conidial cushions are embedded in the host- plants. The conidia are filamentous, frequently somewhat twisted. Cylindrosporium Tubeufianum Alles- cher. This attacks the living green fruit of the bird-cherry, and causes the forma- tion of brown spots from which pustules break out; the premature dropping of diseased fruits follows. In the locality where I observed this disease, numerous trees were attacked and most of the fruiu on each was badly diseased. The my- celium spreads through epicarp and mesocarp, but does not penetrate into Mig... 700 — Cylnarosperium “thes endocarp,, so- thats the gdevelopmend Tubcufianum on fruits of Prunus Hodus. The unshaded parts re —ofithe embryous not direetlysimbterered present parts still green and living, although bearing pustules with. The conidia originate in pycnidial here and there; the remaining 5 parts are completely beset by cavities without any special peridium ; pustules, so that the cells are : : 5 . killed and brown. 4 natural their shape is given in the annexed size. (v. Tubeuf del.) 3 5 S es yee e diagnosis.” The pyenidial cavities arise under the epidermis which is afterwards ruptured and with the cells underlying it becomes brown and dead. 'This note is taken from Underwood, Journal of Mycology, vit., but no mention is made of it in the later paper by Webber and Swingle (‘‘ Diseases of Citrous Fruits in Florida,” U.S.A. Dept. of Agriculture Bulletin, 8, 1896). (Kdit.) * Allescher gives the following diagnosis of this species: Pustulis primum convexis, epicarpio tectis, dein applanatis scutiformibusve, epicarpio ripto cinctis, subcircularibus, saepe caespitosis vel confluentibus, luteo-brunneolis, subfurfuraceis ; acervulis, minutis, innatis, erumpentibus ; conidiis filiformibus, curvatis vel flea- uosis multiguttulatis, hyalinis 40-60=2-3u. Hab. in fructibus immaturis Prunt Padi, quos necat. CYLINDROSPORIUM. 489 As yet the disease has been observed in quantity only in the neighbourhood of Oberammergau (Upper Bavaria). C. padi Karst. Leaf-blight of cherry and plum. This dis- ease is most destructive in the nursery, causing premature defoliation of young trees; it may also cause severe injury to fruit-bearing trees. The leaves become spotted and perforated by holes caused by the falling out of withered spots. Spraying with dilute Bordeaux mixture early in the season is said to have good effects. Fic. 300.—A fruit from Fig. 299 (enlarged). A, Two pustules still further enlarged. B, Pustules before and after rupture of the epidermis. C, Isolated conidia. (v. Tubeuf del.) C. filipendulae Thiim. occurs on leaves of Spiraea Filipendula. C. ficariae Berk. On leaves of Ranunculus Ficaria. (Britain.) C. viridis E. et E., and C. minus E. et E. On leaves of Fraxinus viridis in the United States. C. cercosporoides E. et FE. On living leaves of tulip-tree. C. saccharinum E. et E. On living leaves of Acer saceharinum in the United States. Cryptosporium. Conidial cushions shaped like pyenidia. Conidia rod-like or spindle-shaped. Cryptosporium leptostromiforme Kiiln.” This fungus forms rows of black stromata on the stems of lupines; in the stromata are formed pyenidia-like cavities with several neck-like openings, and in them conidia are given off from conidiophores. The conidia are rods with rounded ends 7-854 lone and about 2 broad; they emerge from the necks of the cavities as long tendril-like chains, and may be continuously given off ‘Fairchild (Journal of Mycology, vu., p. 249) gives results of remedial treatment. *J. Kiithn, Berichte d. landwirth. Inat., Halle, 1880. Fischer, ‘‘ Cryptosporium leptostromiforme.” Breslau, 1893. 490 FUNGI IMPERFECTI. throughout the whole summer. Fischer has proved experi- mentally that the conidia germinate easily in water, that the germ-tubes penetrate into living lupines, and produce a mycelium which spreads through stems and leaves to develop stromata on all the organs of the plant. The formation of both pyenidia and conidia goes on throughout the autumn and following spring on dead plants, the fungus being capable of living as a saprophyte and of hibernating. The disease may occur with great severity. Fischer describes cases where more than the half of the plants in a field were attacked and died before flowering or soon after. There is thus a loss not only in lupine seed, but also in the good effects which the crop has as a “green manure.” Fischer gives the following measures for keeping this pest in check: “ Where the fungus has obtained a footing, lupines should not be planted till at least the year after next, and then only as a catch-crop on stubble; it would be still safer to keep lupines off the land till the third or fourth year. After lupines as a catch-crop, they may safely be sown again in spring as a seed crop, after the lapse of a clear year. No lupines should be cultivated near diseased fields. Instead of ploughing-in a eatch-crop of lupines directly, it should be dried and used as litter for cattle, because the excrement has been found to kill the fungus; the lupines after lying over winter in the manure- heap could then be used as manure in spring. Similarly when the lupines have been grown for seed, they should be closely mowed down so that little stubble is left; the straw may then be used for litter.” This fungus has not as yet been observed on plants other than lupines. DIDYMOSPORAE. Didymosporium. Conidia brown, oval or spindle-shaped, bicellular, and not produced in chains. Didymosporium salicinum Vuill. Vuillemin reports this as very destructive to the Osier cultivation in Bourgogne. Marsonia. Conidia transparent, two-celled, and not produced in chains. The species live on leaves. MARSONIA. 49] Marsonia juglandis (Lib.) produces on leaves of /uglans little greyish yellow spots with brown margins; thereon stromata are formed, which rupture the epidermis and liberate the large sickle- shaped conidia. (Britain.) M. populi (Lib.). On leaves of species of Popu/us in Europe and Britain. M. potentiliae (Desm.). On species of Potentilla. (Britain.) M. campanulae Bres. et All. On Campanula latifolia. The following are North American species : M. toxicodendri (El]. et Mart.). On Rhus Toxicodendron. M. quercus Peck. On Quercus ilicifolia. PHRAGMOSPORAE. Coryneum. The conidial patches are black and disc-like, and rupture the host-epidermis. The conidia are oblong or spindle-shaped, yellowish, and pluriseptate; they are abjointed from short conidiophores. Coryneum Beyerinkii Oud.’ This is stated by Beyerink to be the cause of a “gum-flux” of cherry and allied species of Rosaceae. It is the conidial form of Ascospora (see p. 211). C. camelliae Mass.” occurs on living Camellia leaves at Kew. (Britain.) Pestalozzia. Conidia spindle-shaped, with two or more brown median cells and hyaline terminal cells, the one at the free end carrying several ciliate processes. Pestalozzia Hartigii Tub.” The external effects of this disease have been long known, although the fungus causing it has only been recently detected. It attacks young plants of various trees and shrubs. The symptoms are yellow discoloration of the foliage, and constriction of the stem just above the level of the soil, followed by death of the whole plant. At the constriction of the stem the rind gradually dries up, whereas neighbouring portions continue to grow in thickness till finally the bark is ruptured (Fig. 302). In the living part of the 'Oudemans, //edwigia, 1883. * Cooke, Crevillea, Xx., p. 8, 1891. Sv. Tubeuf, Beitrdge zur Kenntniss d. Baumkrankheiten, 888; and Forstlich naturiwiss, Zeitschrift, 1892. Fic. 301.—Pestalozzia Hartigii. Young Spruce showing constriction just over the surface of the soil. (After v. Tubeuf.) Fic. 302.—Pestalozzia Hartigii. «a, Beech seedling with a diseased constricted part on its stem, *. 0b, Two isolated conidia. (After. Rostrup.) Fic. 303.—Pestalozzia Hartigii. Conidia and coni- diophores on part of stroma. (After v. Tubeuf.) PESTALOZZIA. 493 rind of young plants of spruce and silver fir, I succeeded in finding near the place of constriction, a delicate mycelial stroma enclosing some cavities (pseudopyenidia). | Conidia were formed inside these cavities and emerged to the exterior. They belong to the genus Pestalozzia, and have two brown median cells, a trans- parent stalk-cell to which the long stalk is attached, and a transparent terminal cell carrying two or three transparent thread- like appendages (Fig. 305). Germination results in the emission of a strong germ-tube from one of the three lower cells. If at any time the conidia dry up, the two clear transparent cells Fic. 304,—Pestalozzia funerea on Chamaecyparis Menziesti. At the places marked X cambium and rind have been killed, so that growth in thickness no longer takes place; the higher parts, however, have continued to thicken, but are gradually dying. (v. Tubeuf phot.) collapse and the appendages easily fall off, so that on material of this kind the conidia are only two-celled and brown. The mycelium after cultivation in nutritive gelatine soon produces conidia. This fungus was found by Rostrup on beech, producing much the same effects as just described. On this host it has been found very destructive in young naturally regenerated forest, the loss in Bavaria and Wurtembure within very recent years having been estimated at 30 per cent. It also occurs on ash, sycamore, and other trees. P. funerea Desm. (Britain and U.S. America), The spores of 494 FUNGI IMPERFECTI. this fungus were found by Boehm! on diseased cypress trees, and although investigations are not yet complete, it is believed that this Pestalozzia is the cause of a well-known disease on cypress. The symptoms on Chamaecyparis Menziesii are local constriction of stems and branches, and death of portions beyond. The rind and cambium of constricted places are killed, the bark becomes spht, and the wood dries up. P. funerea is a well-known sapro- phyte on twigs and needles of Cupressus, Juniperus, and other Conifers ; its occurrence as a parasite has been suggested several times. P. gongrogena Temme’ is said to cause the canker of willow. In diseased willows Temme found an intercellular and an intracellular mycelium with pycnidia and conidia of Pestalozzia, but other pyenidia of unknown affinity were also present. P. insidiens Zab. On bark of Ulmus americana. (U.S. America.) P. phoenicis Grev. causes a disease on indoor cultivated palms. The following are some of the more important forms frequenting living leaves : P. Guepini Desm.’ (U.S. America). The conidia of this species are found on large spots with dark margins on living leaves of Camellia japonica, Magnolia, Citrus, Rhododendron, and other plants. Spore-patches appear on the epidermis, and give off conidia embedded in a mucilaginous slime. The conidia have three dark median and two hyaline terminal cells, the distal one bearing the characteristic appendages. The leaves are permeated with mycelium and fall prematurely. P. inquinans ©. et Hark. On Hucalyptus in California, P. stictica B. et C. On Platanus occidentalis and Tilia in United States. P. concentrica b. et Br. On leaves of Crataegus, Pyrus, Castanea, and Quercus in North America. P. suffocata E. et E., and P. discosioides E. et E. On cultivated and wild rose shrubs in America. Pestalozzina. Conidia similar to those of Pestalozzia, but all the cells hyaline. 1 Zeitschrift f. Forst. u. Jagd-wesen, 1894, p. 63. 2Thiel’s landwirth. Jahrbuch, 1887; and Ber. d. deutsch. botan. Ges., 1890. ® Annal. des Science natur., Sér. 11., Vol. xtrt., 1840; Briosi et Cavara, Funghi parasit., VI. ail PESTALOZZINA. 495 Pestalozzina Soraueriana Sacc.’ occurs on foxtail grass (Alopecurus pratensis). The conidial tufts develop on spots which appear on the gradually withering leaves. The bristle-appendages on the terminal cell of the conidia are lateral, only one being terminal. This disease was first observed by Weinzierl at Vienna, Fic. 305.—Septogloeum Hartigianum on Acer canpestye. The dead twigs exhibit black points and lines--the pycnidia of the parasite. (v. Tubeuf phot.) and has not as yet been found out of that neighbourhood ; it attacks the pure-culture seed-beds only. Septogloeum. Like Gloeosporivm, except that it has pluricellular conidia. Septogloeum Hartigianum Sace. Twigs of the common maple (Acer campestre) are subject to a disease, which exhibits ! Sorauer, Zi ischrift ie Phlanze nkrankheite n, ISo4, p. 213. 7K. Hartig, Forstlich-naturwiss. Ze itschrift, 1sev, p. YSU, 496 FUNGI 1MPERFECTI. itself in the drying-up of young twigs before their buds open in spring. The older branches, however, assume their normal fohage. Examination of diseased twigs reveals the mycelium of a parasitic fungus living both inside and between the cells of rind and wood. Conidial patches break through the host’s epidermis about May as long gereyish-green lines. The conidia are hyaline, three-celled, and cylindrical with rounded ends; the conidiophores are short thick rods. In May and June the spores are capable of infecting new hosts, and germinate in a few hours. Infection of twigs takes place in summer, and the mycelium spreads through the first-year shoots, without, however, giving any external indication of its presence till the following spring, when the twigs dry up as already described. S. ulmi (Fr.) may be a form of Phyllachora ulmi. The mycelium lives in parenchymatous cells, and causes the formation of brownish-yellow spots on leaves of the elm. The conidial patches form tiny points onthe lower surface of the leaf; they consist of pycnidia-like structures without a peridium, arising from a stroma developed under the epidermis. The conidia are spindle-shaped and pluricellular. S. mori (Lév.) is stated by Briosi and Cavara to produce yellow spots with brown margins on the leaves of Morus alba and M. nigra. Death and premature defoliation of the host then take place. The conidial patches develop under the epidermis, and rupture it as the conidiophores emerge; they have no real peridium, hence the fungus cannot belong to the group Phleospora, as Saceardo supposed. The conidia are long, cylindrical or fila- mentous, and pluricellular. Amongst the more important North American species are: S. profusum (E. et E.). On living leaves of Corylus americana. S. fraxini Hark. On Fraxinus Oregana. S. apocyni Peck. On Apocynwm cannabinum. Ill. HY PHOMYCETES. Conidia produced neither in pycnidia as in Sphaeropsideae, nor from a special stroma as in Melanconideae, but free on conidiophores given off from the mycelium. HYPHOMYCETES. 497 The group is subdivided into the families of the Mucedineae, Dematieae, Stilbeae, and Tubercularicaes 1. FAM. MUCEDINEAE. 1. Sect. AMEROSPORAE. 1. Subsect. Micronemeae. Oospora. Conidia, transparent or only slightly coloured, globose or ovoid, non-septate, and produced in regular chains from simple short conidiophores ; they thus resemble the genus Zorula in the Dematieae. Oospora scabies Thaxt.? is said to cause the well-known seab or scurf on beet and potato. This consists in portions of the surface of the subterranean tubers swelling out as rough brown excrescences. Other authors ascribe this disease to bacteria. Microstroma. Conidia unicellular, transparent, oval, and shortly stalked. Microstroma album (Desm.). This, although common on living leaves of several species of Quercus, is not a serious disease. The conidial patches on the under side of the leaves are white and very thin. (Britain.) M. juglandis (Béreng.) frequents the leaves of Juglans regia and J. cinerea in Europe and North America. Monilia. Conidia oval or spindle-shaped, and produced in chains from branched conidiophores. Monilia fructigena Pers. (Britain and U.S. America.) This is the cause of certain widespread diseases—the brown-rot of cherry and plum, the peach-rot, and a rot on apples and _ pears. It has been the subject of many papers since Thiimen first described it in 1879.° All parts of the host are attacked, and This is the arrangement followed by Massee, ‘‘ British Fungus Flora,” Vol. 1.; there the characters of the various sub-divisions may be obtained. (Edit.) *Thaxter, Connecticut Agric. Hxper. Station, Report, 1890. “Amongst the more important descriptions are: Thiimen, /ungi Pomicola, 1879; Smith (Worth. G.), Gardener's Chronicle, 1885, p. 52; Arthur, New York Agric. Exper, Station, 1v., 1885. ») I ~ 498 FUNGI IMPERFECTI. exhibit reddish or yellow spots; therein the mycelium spreads rapidly and gives off tufts of conidiophores which rupture the epidermis. The conidiophores are septate, branched, and give off chains of unicellular oval conidia. Meanwhile the affected fruit becomes rotten and gradually shrivels up, it remains, however, hanging on the tree throughout the winter. During Fic. 306.—Monilia fructigena. A, Apple showing the grey conidial patches as more or less concentric lines. B, Young. Peach, shrivelled up in consequence of attack. (v. Tubeuf del.) next spring, when the fruit is again moist, further conidia are given off. Infection takes place by wounds or even through the epidermis of young leaves and blossoms. The conidia have Fic. 307.—Monilia jructigena. Branched conidiophore with chains of conidia. a, Branched hypha of Monilia in the tissue of an Apple. (v. Tubeuf del.) been found to retain their vitality for two years. Smith’ found that twigs were also affected by the disease, so that a gummy degeneration took place in the soft bast and cambium. As remedial measures, the gathering of all diseased fruit left hanging over winter is strongly recommended. ‘This, as well as other diseased parts, should be burned as soon as_ possible. 'Smith (Erwin), Journal of Mycology, vi., p. 36. MONILIA. 499 Washing of stems with a solution of iron sulphate in spring before the buds unfold is suggested, also spraying of young foliage with dilute Bordeaux mixture. _ Oidium. Mycelium epiphytic on living plants. Conidia unicellular and barrel-shaped, produced in chains on erect conidiophores. Many have already been proved to be conidial forms of Erysipheae. Oidium erysiphoides Fr. frequents living leaves of hop, clover, cucumber, ete., and is probably the conidia of species of Hrysiphe on these hosts. (Britain and U.S. America.) O. Tuckeri Berk. On leaves and berries of the vine (see Uncinula, p. 176). O. leucogonium Desm. On roses; probably the conidial form of Sphaerotheca pannosa (see p. 172). O. farinosum Cooke. On living leaves of apple-trees. { Britain.) O. chrysanthemi Rabh. On leaves of cultivated chrysan- themum. (Britain.) O. aceris Rabach. On leaves of Acer Pseudoplatanus. It is probably the conidial stage of Uncinula bicornis, (Britain.) O. mespilinum Thiim. On leaves of medlar. (Britain.) O. destruens Peck. On Amelanchier canadensis and Prunus serotina 11 America. O. tabaci Thiim. On leaves of tobacco. O. monilioides Link, probably the conidial stage of Hrysiphe graminis, occurs on living grasses over the whole world (see p. 175). 2. Sub-sect. Macronemeae. Botrytis. Mycelium grey. Conidia more or less spherical, and pro- duced in aggregations on the ends of branched conidiophores. Many of the species are saprophytes, others are parasitic on plants or insects, and others form sclerotia: the latter have already been considered under Sclerotinia (see p. 267). The following are known to be parasitic on plants: Botrytis cinerea Pers. This enemy of many plants has already been noticed as Sclerotinia Fuckeliana ; so also B. Douglasii Tubeut. 500 FUNGI IMPERFECTI. B. galanthina Sacc. occurs on the bulbs of Galanthus nivalis in Britain. B. parasitica Cav. produces sclerotia and conidia on Tulipa Gesneriana in Italy (Selerotiwm tulipae). B. vulgaris Fr! This is a very common species, and includes. several well-marked varieties. It is ‘said to be parasitic on cultivated lettuce causing a “ leaf-rot.” B. fascicularis Sacc. is reputed to be the cause of a “ fruit- mould” on the ege-plant (Solanum Melongena) in the United States. A Botrytis is figured by Atkinson? as frequent on diseased carnation-plants. Ovularia. Conidiophores simple except for tooth-like projections near the apex on which the conidia are developed. Conidia uni- cellular, colourless, solitary, rarely in chains. “Closely allied to Ramularia, but distinguished by the one- celled conidia” (Massee). Ovularia pulchella (Ces.). Briosi and Cavara distinguish this as a disease of Loliwm italicwum in Italy. The leaves become black-spotted and permeated with an intercellular mycelium, from which arise the erect, branched, septate conidiophores. The more vigorous conidial patches have a delicate rose colour. O. necans Pass. produces large spots on the foliage of quince and medlar, so that the leaves gradually wither and dry up. Conidia appear as a white powder on the dead remains. This fungus is recorded from both Italy and France. The following are British species occurring on leaves; several of them, however, are placed by Saccardo under Ramularia: Ovularia lychnicola (Cke.) Mass. On Lychnis diurna. O. senecionis (Sacc.). On Senecio vulgaris. O. lactea (Desm.). On species of Viola. O. armoraciae (Fuck.). On cultivated horse-radish. It is reported as somewhat destructive in the United States. O. interstitialis (B. et Br.). On under surface of leaves of primrose, forming yellow spots in the angles of the veins. O. primulana Thiim. On leaves of Primula. O. cochleariae (Cke.). On Cochlearia officinalis. 1Wehmer on species of Botrytis, Zeitschrift f. Pflanzenkrankheiten, 1894. 2 Atkinson, ‘‘ Carnation Diseases,” at Amer. Carnation Society, 1893. = OVULARIA. 501 O. alnicola (Cke.). On Alnus glutinosa. O. scelerata (Cke.). On Ranunculus sceleratus. O. rosea (Fuck.) produces irregular brown spots on the leaves of various species of willow. . asperifolii (Sacc.). On Symphytum officinalis. . veronicae (Fuck.). On spots on leaves of Veronica Chamaedrys, etc. . lamii (Fuck.). On Zamium. . syringae (Berk.). On Syringa. . sphaeroidea Sacc. causes spots on leaves of Lotus. carneola Sacc. On spots on leaves of Scrophularia nodosa. . bistortae (Fuck.). On spots on leaves of Polygonum Bistorta. - obliqua (Cke.). On leaves of Rumex. ©CO0000000 2. Sect. DipyMOsPoRAE. Didymaria. Conidia two-celled, colourless, and produced singly at the extremity of simple erect conidiophores. Didymaria prunicola Cav. Cavara states that this causes raised roundish spots on the upper surface of leaves of plum; finally the leaves gradually dry up and fall off. Slender two- celled conidiophores are produced, and give off each a two-celled obovoid conidium. D. Ungeri Cord. On living leaves of Ranunculus repens. (Britain.) D. astragali (Ell. et Hol.). Found on leaves of Astragalus canadensis. D. spissa Hark. On leaves of Solidago occidentalis; both species in North America. Bostrichonema. Conidiophores erect, spirally twisted, unbranched, and non- septate. Conidia elliptic or oblong, two-celled, and hyaline. Bostrichonema alpestre Ces. On living leaves of Polygonwin viviparum and P. Bistorta. (Britain.) B. modestum (1. et B. White). On leaves of Alchemilla alpina. (Britain.) 3. Sect. PHRAGMOSPORAE. Ramularia. Conidiophores emerging in tufts from the stomata; they give off a terminal conidium, then bend over and produce a lateral conidium, and so on they branch in a sympodial manner, pro- 502 FUNGI IMPERFECTI. ducing conidia at the end of each branch. Conidia septate oval or cylindrical, and light-coloured. “The parasitic habit, simple or sparingly branched hyphae, denticulate and bearing the septate conidia at the tips, charac- terize the genus, which differs from Ovularia only in the septate conidia” (Massee). Ramularia cinarae Sacc. is said by Prillieux’ to have caused ereat destruction in the cultivation of artichokes. The leaves became spotted and died, so that no flower-heads were produced. The following are British species: Ramularia hellebori Fuck. On leaves of Helleborus foetidus and H. viridis. R. epilobii (Schn.). On leaves of Epilobium. R. ulmariae Cooke. On leaves of Spiraea Ulmaria. (U.S. America.) R. geranii Fuck. On under surface of leaves of various species of Geranium. R. lampsanae (Desm.). On Lampsana and Hypochoeris. R. pruinosa Speg. On Senecio jacobea. R. plantaginis El. et Mart. On leaves of Plantago major. (U.S. Am.) R. variabilis Fuck. On leaves of Digitalis and Verbascum. (U.S. America.) R. calcea Ces. On leaves of Glechoma hederacea. R. urticae Ces. On leaves of species of Urtica. (U.S. America.) R. pratensis Sace. On Rumer Acetosa. R. rufibasis (B. et Br.). On Myrica Gale. Some of the more important North American species are: Ramularia rufomaculans Peck. On the buckwheat (Fagopyrum esculen- tum), it has proved a somewhat injurious fungus. albomaculata Peck. On leaves of Caryu umericana. viburni E. et E. On leaves of Viburnum Lentago. celtidis E. et K. On leaves Celtis occidentalis. desmodii Cooke. On leaves of various species of Desmodium. Www Dw brunnea Peck. On living Tussilago farfara. R. areola Atks.2. This causes spots on the foliage of cotton. “Spots amphigenous, pale at first, becoming darker in age; irregular in shape, limited by the veins of the leaf, conidia in profusion giving a frosted appearance to the spots. Conidio- phores fasciculate, in small clusters distributed over the spots. Conidia oblong, usually abruptly pointed at the ends” (Atkinson). R. Goeldiana Sacc. is said to kill leaves and twigs of Coffea arabica in Brazil. 1«*Maladie d.. Artichauts,” Bulletin de la soc. mycolog. de France, 1892. * Atkinson, Botanical Gazette, xv., 1890, p. 166. PIRICULARIA. 503 Piricularia. Conidia grey, pluricellular, somewhat pear-shaped, and _pro- duced from the apex of simple erect conidiophores. Piricularia oryzae Br. et Cav. This species is described by Briosi and Cavara as causing a disease of rice in Northern Italy. The plants become spotted and reddish-brown in summer, finally withering. The conidiophores arise on the spots on the lower surface of the leaf, and bear light-grey three-celled Fic. 308.—Mastigosporium album. (v. Tubeuf del.) conidia. Diseased plants may be found bearing this fungus only, frequently however it is in company with other fungi. Cercosporella. Conidia hyaline, similar to those of Cercospora, and produced from simple or branched hyaline conidiophores. Cercosporella persica Sacc. is parasitic on living leaves of peach. In America it has been known since 1890, and receives the name of “frosty mildew.” It causes yellow spots on the lower surface of the leaf. . C. pastinacae Karst. occurs on living leaves of cultivated parsnip, 504 FUNGI IMPERFECTI. Mastigosporium. Conidia hyaline and four-celled, frequently bristled. Mastigosporium album Riess. produces oblong dark spots with light margins on leaves of living grass. The conidia are produced on the margins of the spots (Fig. 308). Fusoma. Sumilar to Fusariwm, but the mycelium is loose and not aggregated into a tuft. Conidia spindle-shaped and _ septate. Fic. 310.—Epidermis of a Pine-seedling with a stoma. Some hyphae of Fusoma have produced partial dissolution of cell- walls. (After R. Hartig.) Fic. 309.—Fusoma parasiticum. Dis- eased Pine-seedlings, with, a, root killed ; Fig. 31].—Fusomea parasiticum. Coni- b, hypocotyl killed; c, first leaves and dia—immature, mature, and germinat- plumule killed. (After R. Hartig.) ing. (After R. Hartig.) Fusoma parasiticum Tub.’ is the cause of a disease of seed- lings, particularly those of Conifers. The first symptoms are dark patches on the seedlings, followed by their collapse. There- IR. Hartig, Forstlich-naturwiss. Zeitschrift, 1893, p. 432. FUSOMA. 505 after in moist weather or under artificial cultivation, a light-grey mycelium appears bearing numerous slightly curved, tapering, pluriseptate conidia (Fig. 311). In Bavaria and Baden this parasite has caused great loss in the seed-beds of conifers. F. inaequale Hoyer. On living leaves of Taraxacum officinale. Septocylindrium. Conidia cylindrical, hyaline or pale-coloured, with two or more septa, and produced in chains. Septocylindrium aromaticum Sace. occurs on living Acorus Calamus, killing leaves and even plants. The mycelium grows ‘intercellularly and produces spots. The conidiophores emerge in tufts from stomata included in the spots, and give off long thread-like, pluriseptate, hyaline conidia. 2. FAM. DEMATIEAE. 1. Sect. AMEROSPORAE. 1. Subsect. Micronemeae. Many of the genera of this subsection contain species found on the living leaves of plants, but none of them are yet of economic importance. 2. Subsect. Macronemeae. Hormodendron. Mycelium grey, epiphytic, and creeping. Conidiophores erect, branched, and septate. Conidia spherical or ovoid, unicellular, and produced in chains. Hormodendron hordei Br.’ This produces a characteristic spotting of the haulms and leaves of barley, accompanied by a stunting of the whole plant and poor development of the ears. This is not a true parasite, but when it appears in quantity it has considerable effect, attacking whole fields and causing great injury. The spots and conidia are found also on wild Hordeum murinum on the margins of roads and fields. !Bruhne in Zopt's Beitrage 2. Physiol. u. Morphol. nied, Organiamen, IV., 1894. 506 FUNGI IMPERFECTI. 2. Sect. DiIpYMOSPORAE. 1. Subsect. Micronemeae. Dicoccum. Conidia oblong, two-celled, and arising from short simple conidiophores. Mycelium subcuticular. Dicoccum (Marsonia) rosae (Bon.) causes brown spots on living leaves of roses, and a premature leaf-cast takes place. Little mycelial stromata develop between the epidermal cells and their cuticle, and give off two-celled hyaline conidia. D. uniseptatum B. et Br. forms dark patches on twigs of Clematis vitalba. (Britain.) D. lathyrinum Ell. et Gall. On living leaves of Lathyrus ochroleucus in America. Cycloconium. Mycehum subcuticular. Conidia one- to three-celled. Cycloconium oleaginum Cast." When this fungus is present, the leaves of the olive show roundish light-brown spots with dark margins, then becoming discoloured, they roll up and drop off. The mycelium grows in the walls of the epidermal cells, branching dichotomously; branches of the hyphae break out through the cuticle as sac-lke cells, which become the conidio- phores. The conidia consist of one to three cells. Kruch states that Cercospora cladosporioides is often present along with this disease of the olive, and may take some part in causing it. Peglion states that this or an allied species occurs on leaves of Quercus Llex. 2. Subsect. Macronemeae. Passalora. Conidia oblong or spindle-shaped, two-celled, and borne on the apex of greenish pluriseptate conidiophores, arising from an olive-green mycelium. Passalora bacilligera M. et Fr. occurs on living leaves of Alnus glutinosa. (Britain.) P. microsperma Fuck. This frequently covers the whole lower surface of the leaves of Alnus incana with little tufts of 'Kruech, Bulletin soc. bot. ital., 1892. Boyer, Recherches sur les maladies de ?Olivicr, Montpellier, 1892. PASSALORA. 507 brown septate conidiophores, bearing long, two-celled, obovate conidia. Fusicladium. Mycelium greenish and sparingly septate. Conidiophores in tufts, short, erect, and bearing terminal conidia. Conidia ovoid or clavate, and one or two-celled. The species are conidial forms of Venturia, and have already been considered. Some of the better-known forms are: Fusicladium dendriticum Wallr. (Britain and U.S. America). This attacks the leaves, shoots, and fruits of the apple (see p. 218). Fie. 312.—Venturia (Fusicladium) dendriticum forming brown spots on an apple; those still in the earlier stages have a radiate margin and bear conidia. The enlarged section shows two rows of large-celled parenchyma of the apple, covered by a stroma of pseudoparenchyma bearing conidiophores and conidia. (v. Tubeuf del.) F. pirinum (Lib.) (U.S. America). This is a cause of “spotting” on leaves and fruits of the pear, also of species of Crataegus and Amelanchier. The conidial patches are brownish in colour. Peglion states that this parasite forms sclerotia in the bark of twigs. It is probably a conidial form of Venturi ditricha var, pyri. F. cerasi (Rabh.) attacks the cherry orchards with such virulence that the crop may be rendered quite unsaleable. F. eriobotryae Cav.' Cavara states that this attacks the 1Cavara, Rivista di Patologia Vegetale, 182. 508 FUNGI IMPERFECTI. leaves of Mespilus (Evrivbotrya) japonica causing them to become spotted and to wither. The hyphae live in the epidermis, and form a stroma from which conidia are given off. F. tremulae Frank. Frank! gives this as the cause of a disease of the aspen (Populus tremula). The leaves turn brown and fall, the shoots in consequence soon drying up. Conidia are developed on the surface of dead leaves and ger- minate on living leaves of aspen, producing a germ-tube which, after forming an adhesion-disc, penetrates into the cavity of the epidermal cells. F. depressum B. et Br. is found on living leaves of Angelica sylvestris. (Britain and U.S. America.) F. praecox Rabh. On living leaves of Tragopogon orientalis. F. sorghi Pass. On living leaves of Sorghum halepense. The following are North American species : F. caryogenum Ell. et Langl. On leaves of Carya olivaeformis. F. effusum Wint. On leaves of Carpinus americana. F. destruens Peck. On living Avena sativa. F. fasciculatum ©. et E. On leaves and stems of Euphorbia. Scolecotrichum. Mycelium greenish. Conidia oblong or oval, produced both terminally and laterally on the conidiophores. Scolecotrichum melophthorum (Prill. et Del.).” This pro- duces a melon disease in France known by the name “ Nuile.” It consists in the fruits and stems becoming spotted, the tissue being completely destroyed. Sc. graminis Fuck. Occurs on grasses, especially on the oat. Pammel*® reports it as also injurious on barley during 1891, in some parts of the United States: the diseased leaves were marked with brown or purplish brown spots. Sc. fraxini Pass. On living leaves of Fraxinus excelsior and F. Ornus. Cladosporium. Mycelium greenish. Conidia globose or ovoid, one to four- celled, and of variable form. The species are mostly sapro- phytes on substrata of all kinds. 1 Ber. d. deutsch. botan. Gesell, 1883, p. 29. 2 Bulletin de la soc. mycolog. de France, 1891. ® Journal of Mycology, vi1., p. 96. CLADOSPORIUM. 509 Cladosporium herbarum (Pers.). This species is found every- where on dead plant remains, but it is also common on living leaves of many plants. The first suggestion that this form might occur as a parasite came from Haberlandt’ and Frank.* It possesses a dirty-grey, thick, septate mycelium, which may be colourless when young or growing inside a substratum; it applies itself closely to the surface of plants and even pene- trates through the stomata or cell-walls into the tissues. The conidiophores are erect, otherwise variable in form; they give off conidia from the apex or from lateral processes. The conidia are oval and contain a variable number of cells. Organs of plants attacked show grey spots, and withered parts if they are still alive. The following are some of the papers describing Cladosporium herbarum as, in certain’ circumstances, a parasite. Prillieux and Delacroix,? on apple-trees and raspberry-bushes; Cavara,* on raspberry, cycads, agave, and other plants; Sorauer,’ on peas. Lopriore® describes this fungus as the cause of a “black” disease on ears of wheat; the results of infection were however somewhat variable. Ritzema Bos reports it as producing disease, and in some cases death, in fields of oats. Kosmahl and Nobbe” found that seedlings of Pinus rigida blackened and died suddenly in the beginning of May, apparently from the attacks of this fungus. Janezewski’ states that this Cladosporium is a conidial form of Sphaerella Tulasnet, a new species of Ascomycete established by him.* Cl. elegans Penz. This causes on the orange a disease or “scab,” which has been injurious both in Southern Europe and the Southern States of America.” It attacks chiefly wild orange 1 Friithling’s landwirth. Zeitung, 1878. 2 Die Krankheiten der Pflanzen, 2nd Edit., 1896, 11., p. 292. * Bulletin de la soc. mycolog. de France, Vi. 4 Revue mycologique, 1891. ® Handbuch d. Pflanzenkrankheiten, 1886. 6 Berichte d. deutsch. botan. Gesell, 1892; Landwirth. Jahrbuch, 1894. 7 Extraits du Bulletin de l’ Academie des sci. de Cracovie, 1892, 1893, 1894. 8 Schostakowitsch (flora, 1895 (ergzbd.) distinguishes Cladosporium from other genera. *Scribner, Bulletin of Torrey Club, xut., 1886, p. 181. Underwood, Journal of Mycology, VIL., p. 34. Swingle and Webber, ‘‘ Diseases of Citrous Fruits,’ U.S.A. Dept. of Agriculture Bulletin 8, 1896. 510 FUNGI IMPERFECTI. trees, more rarely the sweet orange and lemon. The disease first appears as whitish or cream-coloured spots on leaves, young twigs, or fruit. If the spots are numerous the leaves become badly curled or twisted, and covered with wart-like eruptions. Cl. viticolum Ces. is regarded as a dangerous parasite of the vine. Cl. carpophilum Thiim. This species has been found para- sitic on plum and peach in the United States. Its mycelium creeps over the surface of leaves and fruit, causing pale-coloured spots which extend and run together, spoiling the appearance of the fruit. The disease as yet does not appear to have a very wide distribution, nor is it directly very injurious, but as cracking of the ripe fruit occurs when it is present, the way is opened for entrance of fruit-destroying fungi. Cl. condylonema Pass. also occurs on leaves of the plum. It causes leaf-spot and leaf-curl. The mature conidia have fine spines on their coat. Cl. fulvum Cooke. (Britain and U.S. America.) This is the cause of a disease of tomato. It attacks leaves and shoots of plants cultivated indoors, and soon causes their death. Prillieux and Delacroix’ have described a somewhat similar disease in France, found, from artificial infection, to be produced by some species of Cladosporiwm, but whether this particular species, they did not state. Cl. cucumerinum Ell. et Arth* causes a disease of cucumber Frank? describes a disease which he found to be due to a Cladosporium (Cl. cucumeris n. sp.). This attacked the fruit of both cucumbers and melons in cultivation under glass at Berlin, and caused great damage; brown rotten depressions appeared on the fruits, and thereon the tufts of conidiophores. Cl. macrocarpum Preus. causes a “scab” disease of spinach in the United States (WS. Agric. Exper. Station Bulletin, 70, 1890). Other species that may be parasitic are: Cl. pisi Cug. et Macc. On living pods of Pisum sativum in Italy. Cl. epiphyllum Mart. On leaves of Quercus, Platanus, Populus, Hedera, etc. (Britain and U.S. America.) Cl. juglandinum Cooke. On leaves of the walnut. (Britain.) ' Bulletin de la soc. mycology. de France, 1891. ’ Description in Mass. Agric. Kaper, Station Report, 1892. * Zeitschrift f. Pflanzenkrankheiten, 111., 1893. CLADOSPORIUM. 511 Cl. Scribnerianum Cav. On leaves of Betula populifolia in America and Italy. Cl. hypophyllum Fuck. On leaves of Ulmus campestris. Cl. tuberum Cooke. In the tubers of Batatas edulis in Carolina, U.S.A. 3. SecT. PHRAGMOSPORAE. 1. Subsect. Micronemeae. Clasterosporium. Conidia brownish, cylindrical or spindle-shaped, and consisting of three or four cells. Clasterosporium amygdalearum (Vlass.) attacks the leaves of almond, peach, apricot, cherry, and plum. An _ intercellular mycelium has been found, and roundish dry spots with reddish margins are formed. Thereon tufts of short conidiophores are developed, bearing cylindrical, thick-walled, pluricellular conidia. Cl. glomerulosum Sace. (Sporidesmium glom. Sace., 1878, and Pleospora conglutinata Goebel, 1879). Goebel! first described this species as a parasite on Juniperus communis. A colourless intercellular mycelium is present, and in consequence the needles turn brown, die, and fall off prematurely. On the upper side of the needle the mycelium emerges through the stomata, and forms dark-grey coils from which the grey, ovoid, pluricellular conidia are given off. Ceratophorum. Conidia brownish, spindle-shaped or cylindrical, three or more celled, — gyriss trpitaten Meat with diseased the: upper cell with terminal 2.3" Utter) bristles. C. setosum Kirch. Dark spots occurring on the leaves, petioles, and shoots of young plants of Cytisus Laburnum, ete., were found to enlarge and bring about death and defoliation. Kirchner found the leaf-tissue permeated by a colourless septate mycelium, which gives off conidia on both sides of the leaf. The conidia resembled those of Pestalozzia, but their cell-number ! Wurtemburg naturiwiss. Jahreshesle, 1879. Zeitschrift f. Pflanzenkrankheiten, 1892, p. 324. a FUNGI IMPERFECTI. was variable, and the terminal cells, although lighter than the median, were not quite hyaline. The terminal cell bore several very long bristles. C. ulmicolum E. et K. On living leaves of Ulinus fulva in America. Helminthosporium. Conidia brown, cylindrical or spindle-shaped, and pluricellular. Mycelium well-developed and brownish. “Distinguished from Cladosporium by the conidia being more than one-septate at maturity’ (Massee). Helminthosporium gramineum (Rabenh.)' This causes a disease on barley, both in Europe and the United States; as yet, however, it is not very common. It attacks generally the lower leaves, producing long, narrow, dark-brown spots with yellow margins. . The leaves so attacked gradually wither, but do not prejudice the yield of grain seriously. On the spots are developed the black septate conidiophores, each with a large black conidium with from two to eight cross-septa. H. turcicum Pass. causes long spots on the leaves of Zea mais both in Italy and America. The spots are yellow with indistinct dark margins, and from them arise patches of grey septate conidiophores. The conidia resemble those of the species last described, so that some authorities regard the two forms as one. Briosi and Cavara describe the mycelium as consisting of branched septate hyphae, the cells of which frequently become irregularly swollen. The young Indian corn leaves are killed, and the crop may, in consequence, be seriously injured. H. teres Sace. This is a form of H. graminewm which 3riosi and Cavara distinguish as occurring on oats. Infection takes place at the apex of the leaves, and the mycelium spreads through the parenchyma causing elongated dry spots, so that the leaf ultimately dries up and dies. The conidiophores are developed singly, not in tufts, and the conidia are smaller than those of H. graminewm. The conidia are greenish, thick-walled, pluricellular, and produced terminally. H. gracile (Wallr.) causes long marginate spots on the leaves of Iris germanica. ! Eriksson, Botan. Centralblatt, Xx1x., 1887. Kirchner, Zeitschrift f. Pflanzen- krankheiten, 1., 1891, p. 24. CERCOSPORA. 513 Cercospora. Conidia elongated and slender, olive-green, and septate. My- celium greenish. “ Distinguished by the vermiform septate conidia” (Massee). Cercospora circumscissa Sacc.' This is a parasite which occurs on cultivated almond, peach, and nectarine, as well as on wild Prunus serotina in the United States. The leaves are attacked while still young, and exhibit by reflected light a yellowish spot with a dark centre. The conidia arise on the spots as dark-green clusters, thereafter the diseased tissue shrinks, becomes detached, and falls out, leaving “shot-holes” not unlike those produced by species of Phyllosticta. Defoliation may occur in severe cases of attack. As a result of the injury to the foliage, the new wood does not mature well, and second growth may take place during the same season ; shoots of this kind will probably dry up during winter. The fungus may also directly kill the tissue of twigs as far as the cambium. The fruit is never attacked directly, but may be seriously affected through the injury to leaves or twigs. In order to minimize the disease, it is recommended to burn all fallen foliage, and to turn the earth thoroughly below infected trees. Pierce obtained a crown of very healthy foliage on almond trees treated with (1) ammoniacal solution of copper carbonate, and (2) modified eau celeste. C. persicae Sace. On leaves of peach. (U.S. America.) C. acerina Hartig? appears on brown spots on the cotyledons, young leaves, and stalks of young plants of Acer. The conidia are grey, pluricellular, and slightly curved (Fig. 514). The mycelium inhabits the intercellular spaces of the parts attacked, and forms resting sclerotia in the tissues of dead leaves. C. viticola (Ces.).2 This fungus is found in Europe and the United States on Vitis vinifera and V. Labrusca. It causes spots on the leaves, and from these arise close columns of septate conidiophores which give off thick pluricellular conidia. C. beticola Sacc.‘ inflicts considerable injury on cultivated ' Pierce, Journal of Mycology, viui., p. 66 and p, 282. 2R. Hartig, Untersuchungen aus d. forsthotan. Institut, t., Munich. ‘Description and treatment in New York Agric. Exper. Station Report tor 1890, p. 324. *Thiimen, Die Bekiimpfung d. Pilzkrankheiten unserer Kulturge wiichse, 1886, » kK 514 FUNGI IMPERFECTI. sugar beet and beet-root. It is easily recognized by the numerous sharply defined spots produced on the leaves. The conidia are very long and pluriseptate. In the United States this is one of the most serious of beet diseases.’ As preventive treatment, great care should be taken to destroy all infected material. A long rotation should also prove a good remedy. Fic. 314.—Cercospora accrina. 2, Seedling of Acer, with a cotyledon brown and withered, and a leaf partially so. 4, Section through a diseased cotyledon ; the conidiophores (d) emerge from the epidermis, and bear long tapering septate conidia; ¢, sclerotia formed inside the diseased tissues for hibernation. 5, Ger- minating conidia. (After R. Hartig.) C. apii Fres. Common on celery (Apium graveolens) and par- snips (Pastinaca sativa) throughout all Europe and North America. It causes leaf-spots at first yellowish then enlarging and turn- ing brown. The mycelium grows in the intercellular spaces of the leaf, and gives off tufts of conidiophores through the stomata. The conidia are long, tapering, obclavate bodies with an attachment-scar at their larger end.” C. asparagi Sacc. occurs on asparagus in Italy; C. caulicola Wint. frequents the same host in America. C. Bloxami B. et Br. On Brassica in Britain. C. armoraciae Sace. On horse-radish. 'Pammel. Iowa Agric. Exper. Station Bulletin, 15. 1891. 2 Description in New Jersey Agric. Kaper. Station Bulletin 2, 1891. CERCOSPORA. 515 C. resedae Fuck.’ This fungus is the cause of a garden mignonette disease very common in America and Europe. It causes little depressed spots with brownish or yellowish borders, which begin as reddish discolorations of the leaf. The leaves gradually wither and dry up, so that the flowers suffer. The mycelium grows inside the leaves, and gives off tufts of conidio- phores through the stomata. The conidia are elongated, septate, and spindle-like or club-shaped. Spraying with Bordeaux mixture was found to give good results. C. cheiranthi Sacc. produces roundish leaf-spots on wall- flower, and, if severe, causes death of the leaves and premature defoliation of the plants. C. rosaecola Pass. This causes leaf-spot on cultivated and wild roses in the United States. The first indication of disease is the appearance of black spots with reddish margins. The conidiophores emerge from the stomata in tufts, and carry long obelavate conidia. C. angulata Wint. is one of the causes of leaf-spot on currant, and occurs often in company with Septoria ribis. (U.S. America. ) . violae Sacc. occurs on leaves of Viola odorata. . malvarum Sacc. On species of Malva. . althaeina Sace. On hollyhock in the United States. . neriella Sacc. causes leaf-spot on Nerivm Oleander. . Bolleana (Thiim.) produces olive-brown spots on leaves and fruits of the Fig, injuring the crop. C. capparidis Sace. On Capparis spinosa in Italy. C. gossypina Cooke is given by Atkinson as a fungus, frequently present on diseased plants of cotton.* Saccardo records over 230 species of Cercospora, most of which cause spotting of living or fading leaves of many plants, e.g. Phaseolus, Lupinus, Trifolium, Vicia, Gleditschia, Solanum nigrum, Datura, Ricinus, Ampelopsis, Liriodendron, Tilia, Rosa, Potentilla, Rubus, Cydonia, Ptelea, Rhamnus, Euonymus, Ailanthus, Rhus, Sambucus, Viburnum, Olea, Syringa, Morus, Fravinus, Coffea, Ligustrum, Mercurialis, ete. PMOhieliio tle! Heterosporium. Conidiophores simple or branched. Conidia olive, oblong, pluriseptate, and with a spiny or warty outer coat. ‘Fairchild in Report of Section of Vegetable Pathology for 1889, U.S. Dept. of Agriculture. * Botanical Gazette, 1891, p. 61. 516 FUNGI IMPERFECTI. “Resembling Helminthosporiwm in general habit and structure, in fact only distinguished by the minutely warted conidia” (Massee). Heterosporium echinulatum (Berk.).' (Britain and U.S. America.) The “fairy ring spot” of Carnations. This is a serious enemy of cultivated carnations, and causes great damage. It was first described by Berkeley in 1870 as a carnation pest. The symptoms are light-coloured spots on which are concentric rings of dark-coloured conidiophores. These arise from dark- coloured portions of the mycelium inside the leaf and give off conidia with three or more cells. The conidia are at first terminal, but after one has been formed the conidiophore branches laterally and produces another conidium, repeating this process for a considerable time. The spots are produced on leaves, leaf-stalks, and sepals, causing them to wither. In consequence the flowers do not unfold and the plants are rendered unsightly. Cultivation of the carnation in dry airy conditions is said to keep this disease in check. The following are British species occurring generally on fading leaves: H. variabile Cooke. On spinach. H. ornithogali Klotzsch. On Ornithogalum, Convallaria, and other species of Liliaceae. H. typharum ©. et M. On Typha angustifolia. H. laricis C. et M. On larch needles. H. asperatum Massee.? Occurs as a parasite on Smilacina stellata. Napicladium. Conidia oblong, three or more celled, and produced singly on the end of short conidiophores. “Somewhat resembling Helminthosporium and Brachysporiwm, but distinguished by the less rigid fertile hyphae and the large sohtary conidia” (Massee). Napicladium (Helminthosporium) arundinaceum (Cord.). (Britain.) This lives parasitic on the leaves of Phragmites communis, and spreads rapidly from plant to plant. The leaves 1Worth. G. Smith, Gardener's Chronicle, xxvi., 1886, p. 244. Atkinson, ‘‘ Carnation Diseases” at American Carnation Society, 1893. 2?Massee, American Journal of Microscopy, February, 1893. NAPICLADIUM. 517 become coated with conidia and assume a leaden grey colour, so that in many cases only the points remain green. Finally the attacked leaves die and dry up. 4. Secr. Dicrrosrporar. 1. Subsect. Micronemeae, The forms included under genera of this group (e.g. Sporo- desmium and Coniothecium) have as yet been little investigated in regard to their parasitic nature. 2. Subsect. Macronemeae. Macrosporium. Conidia grey, muriform, and borne on the apex of simple or branched conidiophores. Macrosporium sarcinaeforme Cay.' Cavara describes a browning and death of a whole field of red clover (7'rifolium pratense), and ascribes it to this fungus. Minute spots were produced, at first light-coloured, then brown, finally coalescing so as to cause drying-up of the whole leaf. The short thick conidiophores were developed on the lower surface of the leaf, and gave off pluricellular terminal conidia. M. solani Ell. et Mart. This is described? as occurring along with the “black-rot” of the tomato in the United States. It is said to cause a rot in the fruit and a leaf-blight on both tomato and potato. Along with this species there also occur a Fusarium (p. 520) and frequently a Cladosporium; as yet the relationships of the different forms, and the part they take in causing the diseases ascribed to them, is but imperfectly investigated. Sorauer® ascribes a disease on the potato in Germany to this species or to an Alternaria (A. solani). He also believes that it is the cause of the “early blight” of American potato crops, but further investigation is still required. Many other species of Macrosporium have been described on plants of economic importance, yet most of them occur only on parts somewhat faded or languid, so that they cannot be regarded 'Briosi and Cavara, Funghi parasit., v. * Report of the Section of Vegetable Pathology for 1888, U.S. Department of Agriculture. * Zeitschrift f. Pflanzenkrankheiten, 1896, p. 1. 518 FUNGI IMPERFECTI. as important parasites. Amongst these are the following British and North American species: brassicae Berk. On cabbage, generally somewhat decayed. sarcinula Berk. On cucumber. nobile Vize. On Dianthus. alliorum Cke. et Mass. On onion. ramulosum Sacc. On celery. catalpae Ell. et Mart. On Catalpa Bignonioides. nigricantium Atks. is a semi-parasite accompanying other diseases of the cotton plant. SSSSS558 Mystrosporium. “ Allied to Macrosporium, but distinguished by the more rigid and darker-coloured hyphae and conidia” (Massee). Mystrosporium abrodens Neumann.’ This is described as the cause of a disease which destroyed one-tenth of the total wheat-crop in the Haute-Garonne of France. The fungus attacked the nodes and leaves, forming dark patches; the nodes were weakened and frequently broke over, while the ears were badly developed. Alternaria. Conidia grey, muriform-septate, flask-shaped, and borne on short simple conidiophores. “Distinguished by the clavate or flask-shaped muriformly septate olive conidia being united in chains and connected by narrow isthinus-like portions” (Massee). Alternaria brassicae (Berk.) (Britain). This species causes on leaves roundish black spots marked with concentric brown zones. The mycelium lives in the leaf-parenchyma and gives off tufts of conidiophores through the stomata. Briosi and Cavara state that it causes considerable damage to Srassica oleracea, Cochlearia officinalis, and Armoracia, (Probably the same species as Polydesmus exitiosus Kuhn.) Other diseases have been ascribed to species of Alternaria. Septosporium. Conidia brown, and muriform-septate. Conidiophores of two kinds—short and fertile, or elongated and _ sterile. Septosporium heterosporum Ell. et Gall. causes a_leaf- '<*Un nouveau parasite de blée.” Société de Biolog. a Towlouse, 1892. SEPTOSPORIUM. 519 spot on Vitis californica in California. The leaves become quite black on the lower surface, brown on the upper. The fungus has not as yet been reported on cultivated vines. Fumago. Conidia grey and two- or three-celled. The species belong to Capnodium (see p. 181). 3. FAM. STILBEAE. 1. Ser. HyALOSTILBEAE. Sect. Amerosporae. Stysanus. Conidia pale-coloured, more or less spherical, and developed on a dark cylindrical or clavate erect stroma. Stysanus veronicae Pass. This produces irregular spots on the leaves of cultivated Veronica longifolia in Italy, and causes the plant to wither. The columnar stromata are produced on the lower surface of the leaves, and give off unicellular conidia. St. ulmariae M‘W.2) On Spirea Ulmaria in Ireland. Isaria. Stroma erect, clavate, generally branched and bearing conidio- phores all over. The conidia are abjointed from the apex of the conidiophores, and are unicellular, hyaline, and rounded. Isaria fuciformis Berk.® This disease, first observed in Australia, is described by Smith as occurring in England. It attacks grasses, especially Festuca, during summer. The stems and ears are glued together by the fungus-stroma, and conidia are developed on all parts of the plants. 2. Ser. PHAEOSTILBEAE. Sect. Phragmosporae. Isariopsis. Conidia pale-coloured, cylindrical, and pluricellular. Isariopsis griseola Sacc.* produces spots on leaves of living 1 Hedwigia, \877, p. 123. *M‘Weeney, /rish Naturalist, 1895, p. 273. ® Worth. G. Smith, Diseases of Field and Garden Crops, London, 1854, p. 55. *Briosi and Cavara, Funghi parasit. 520 FUNGI IMPERFECTI. cultivated kidney bean. The mycelium lives in the leaf-tissues and forms stromata under the stomata, from which the conidio- phores arise in tufts. The fungus often occurs along with Uromyces phaseolt. Other species of Jsariopsis are recorded on the living leaves of various host-plants, e.g. Cerastium and Stellaria. 4. FAM. TUBERCULARIEAE. Volutella. The conidial patch or sporodochium is disciform, regular, and fringed, or studded over with elongated spine-like hyphae. Conidiophores simple or branched, and bearing elliptical or oblong conidia. The majority of the species of Volutella frequent only dead plant remains. Atkinson,! however, describes and figures a widespread carnation-disease in North America, which is ascribed to a species as yet unnamed. Fresh cuttings are most commonly attacked, and exhibit dirty brown depressed areas, which soon ruin the cutting for purposes of cultivation. Fusarium. Sporodochium more or less effused. Conidia spindle-shaped or sickle-like, pluricellular when mature. The conidiophores are branched, and give off the conidia from their apex. Fusarium heterosporium Nees. Frank? found afield of rye near Kiel completely destroyed, and the ears quite over- grown by this fungus. I have found it on ears of JLoliwm perenne and Molinia coerulea in Bavaria. Species of Fusarium have been frequently described as causing injury to cereal and grass-crops,? in some cases to a_ serious extent. While most of the species of Fusarium are found only on dead or dying plant-remains, a parasitic mode of hfe has been ascribed to some. Fusarium lycopersici Sacc.* The “Sleeping Disease” of '<* Carnation Diseases” in Report of American Carnation Society, 1893. 2 Jahrbuch d, deutsch. landwirth. Gesell., 1892. > Worth. G. Smith, Diseases of Crops, 1884, p. 208. Rostrup (Fusarium avenaceum on Oat) Landboskrifter, v., 1893. 4+Massee, Gardener's Chronicle, xvut., 1895, p. 707. (Kdit.) FUSARIUM. 521 tomatoes. This tomato disease has proved very destructive during recent years in Britain, particularly in the Isle of Wight and the Channel Islands. Plants are attacked when quite young, but the disease seldom manifests itself outwardly till the plant is full grown. The first symptom of disease is drooping of the leaves, with or without discoloration. At this stage the roots of attacked plants will be found to have a yellowish brown colour in the wood region. The mycelium of this fungus will be found in the vessels and other elements of the root. They are believed to originate from resting-spores which have hibernated in the soil and given off germ-tubes by which young rootlets were infected. The mycelium makes its way up the tomato stem, discolouring the vascular bundles as it goes. The conidia are produced on all diseased organs as a whitish bloom on the epidermis. The earlier conidia (Jiplo- cladium) are oval and one- or two-celled, but they are soon replaced by pale orange crescent-shaped conidia of the true Fusarium type. The resting-spores are produced on the hyphae in the tissues of the decaying host-stem; after hibernation, they germinate and produce hyphae which give off the Diplocladium stage. Massee found that only the germ-tubes from resting- spores were able to infect tomato plants The same author does not consider fungicides of much avail on account of the disease beginning from the roots. Careful removal and destrue- tion of all infected material, and a liberal application of lime to the soil are measures recommended. Fus. limonis Briosi (Fusisporium limonis Briosi). This is given by Briosi as the cause of “mal-di-gomma” of orange and lemon trees in Italy and elsewhere ;! Webber and Swingle * ascribe the disease of the orange and lemon in Florida known as “foot-rot” to the same fungus. In Florida the damage done is great and much more serious than that caused by any other disease of the same plants. It may be recognized by the exudation of gum from patches near the base of the tree. The patches enlarge and the disease spreads round the trunk and downwards into the roots, passing inwards from bark to cambium and wood, killing the tissues as it goes. Other symptoms ' Briosi, ‘‘ Mal di gomma,” Memoria della R. Acad. dei Lincei, Kome, 1878, * Webber and Swingle, ‘‘ Diseases of citrous fruits in Florida.” U.S. America Dept. of Agriculture Bulletin, No. 8, 1896, (Edit.) 522 FUNGI IMPERFECTI. are sparse foliage, small yellowish leaves, and death of the smaller branches over the tree. Sweet seedling orange (Citrus aurantium) and lemon (C. limonum) are most subject to this malady, the grape-fruit (C. decwmana) is only slightly liable, and the sour orange (C. bigaradia) is almost wholly exempt. For this reason sour orange stocks should be used on lowlands and flatwoods, and grape-fruit stocks on the higher lands. The most effective treatment is to remove the soil around the crown roots by using a jet of water. Diseased bark should also be cut away and the wounds painted over with carbolic acid or sulphur wash. Good drainage to promote root aeration and the avoidance of excessive use of nitrogenous manures are also recommended. Fus. vasinfectum Atks.' ——— eee ae ree ee ae aaa se 3 . rae" seh -BACTERIOSIS OF BULBS. 539 twenty-four hours to a distance of 5-10 c¢.m. through the leaf- tissue. Infection of leaves and bulbs of common onion with the same Bacillus was also successfully carried out, and the same symptoms of disease followed. Sorauer, in his “ Handbuch,” describes a bulb-rot said to be due to bacteria; but whether it be the same disease as this or not we cannot say. Bacterial Disease of Beans. Halsted! describes a disease on cultivated beans, which caused considerable loss in the United States. Bacteria were present in large numbers in all diseased parts, but to what extent they were responsible for the disease could not be exactly determined. IV. THE PATHOGENIC ALGAE. The Cyanophyceae or Schizophyceae, though generally placed with the Bacteria in the group of the Schizophytes, are here included with the true Algae on account of the great resemblance in their mode of life when they play the part of symbiotes or parasites. The Diatomaceae contain no endophytic species. The Algae differ from the groups of the Fungi, Myxomycetes, and Schizomycetes, in their possession of chlorophyll and their power of assimilation. The relationship of the Algae to other living organisms may be expressed under the following heads: I. Symbiosis of Algae with Fungi. (Lichens.) II. Symbiosis of Algae with animals. Ill. Symbiosis of Algae with chlorophyllous plants. (a) Epiphytes. (b) Endophytes. 1. Inhabitants of free spaces in other plants. 2. Inhabitants of domatia. IV. Parasitism of the endophytic Algae. (2) In relation to animals. (4) In relation to plants. 1. Inhabitants of the cell wall. 2. Inhabitants of the cell cavity. 3. Destroyers of tissues as a whole. 1 New Jersey Agric. Exper. Station, Report, 1892. »_ 540 THE PATHOGENIC ALGAE. The lichen-symbiosis is the most marked example of mutual symbiosis we know. Amongst the partnerships of Algae with animals every form exists from mutual symbiosis to true parasitism or to typical epiphytism. The last condition is, how- ever, more frequently met with amongst Algae or Lichens epiphytic on other chlorophyllous plants. The phenomenon of “shelter-parasitism” is also a frequent one, the Algae inhabiting cavities already present in the host, or “domatia” — places of abode formed with the assistance of the Algae. The full discussion of these and other symbiotic relationships may be had by reference to the works dealing with subject ; some of the more important of these are given: De Bary, Die Erscheinung der Symbiose, 1879. O. Hertwig, Die Symbiose im Thierreich, 1883. Klebs, “Symbiose ungleichartiger Organismen.” Biolog. Centralblatt, 1882. Klebs, “ Beitrage zur Kenntniss niederer Algenformen,” Botan. Zeitung, 1881. Geza Entz, “ Das Konsortialverhaltniss von Algen und Thieren,” Biolog. Centralblatt, 1882. Brandt, “ Die morphologische und physiologische Bedeutung des Chloro- phylls bei Thieren.” Mittheil. d. Zoolog. Station Neapel., 1883. Van Beneden, Animal parasites and Messmates. (English Edition), International Scientific Series. Only these algae which are parasitic on the higher plants come, strictly speaking, within the limits of the present work; we shall, however, also take into consideration the interesting symbiotic adaptations presented by several algae which live endophytic, but not truly parasitic, in higher plants.’ There is a distinct resemblance between the parasitism of algae and that of parasitic fungi. Some parasitic algae live in the intercellular spaces of their host, others inside the host’s cells, and many of them inhabit algae and other aquatic plants. A large number of algae live as endophytes, many of them in cavities occurring naturally in other plants; such we can hardly regard as parasites; nor those which cause the formation of “domatia” on their hosts, since these structures are an indication of a symbiotic rather than of a_ parasitic 1 Altmann (Botan. Zeitung, 1894, p. 207) describes a number of marine algae parasitic on Fucaceae ; Moebius, ‘‘ Endophyte Algen,” Biolog. Centralblatt, 1891 ; also Conspectus algarum endophytarum, etc., 1891, with complete bibliography. -z THE CYANOPHYCEAE. 541 relationship. The manner in which typical chlorophyllous plants gradually become shelter-parasites, and pass from this into the condition of true parasites, is well demonstrated amongst the algae. Few of the parasitic algae can be said to affect their host injuriously by causing death of its tissues; Phyllosiphon is the most marked case of this kind. A. THE CYANOPHYCEAE. These, the blue-green algae, possess a homogenous bluish green plasma, with a colouring matter consisting of phycocyan and chlorophyll. Multiplication takes place only by cell- division; sexual reproduction does not occur. Many of the forms are adapted to a symbiotic life, yet without prejudice to their ability to live as independent organisms. Numerous species form lichens. They are in many cases capable of long resistance to drought. The Cyanophyceae are common both as endophytes and epiphytes on other algae and on higher plants. Amongst them will be found examples of purely shelter-parasitism, of true parasitism, and all intermediate stages, yet no cases are known of real injury or death to host-plants resulting from members of this group. Amongst forms endophytic in Phanerogams may be noticed Nostoc gunnerae in Angiosperms, and Anabaena cycadearum in Gymnosperms; in Pteridophyta only Anabaena azollae is known, and in Bryophyta, Nostoe lichenoides. In every case the Nostoc penetrates as a shelter-parasite into fissures or cavities already existing in the host, and becomes as a rule entangled in a slime occupying the cavity. The Nostoe in Gunnera becomes parasitic at a later stage, and makes its way into the cavity of cells. The other species are never endophytic in the host-cells, though they may affect the cells surrounding a cavity and stimulate them to further growth, apparently, however, without any injurious effect on the host-plant. Nostoc punctiforme (Kutz.) P. Hariot! (Nostoe (Seytonema) gunnerae Reinke).” This occurs in various species of Gunnera, natives of South Africa, New Zealand, and South America; or cultivated frequently in Europe. The occurrence of the Nostoe is in every case the same, its presence being indicated to the 'Hariot, Compt. rend. cxv., 1892. *Reinke, Morphologische Abhandlungen, Leipsic, 1873. 542 THE PATHOGENIC ALGAE. naked eye by bluish-green spots on sections of stems and rhizomes of the host (Fig. 319). Fic. 319.—Nostoc gunnerae. Longi- tudinal section through the apex of a stem of Gunnera manicata. The punc- tated spots indicate the position of Nostoc-colonies. (v. Tubeuf del.) These originate as follows: the species of Gunnera possess charac- teristic mucilage-secreting organs, in the form of fissures of the leaf-tip, collaters on the leaves, and glands on the stems. Merker’ found that these glands originated endogenously in the growing point between each pair of leaves. The mature glands are covered.only by the epidermis, and when activity commences the uppermost cells of the gland-tip, after swelling, become detached and eonverted into slime (Fig. 320). Ultimately the epidermis is ruptured by the pressure of the slime, and the remaining glandular cells are in turn rapidly transformed into the same substance. y 1, X ry) . me aa orre ine Fic. 320.—Nostoe gunnerae. Longitudinal sections through a gland in stem of Gunnera macrophylla. e, Epidermis; s, slime-canal; no, colony of Nostoc. ? (After Merker.) The Nostoc finds its way into the gland as soon as the epidermis is broken, apparently attracted there by some secretion. Merker found that the NVostoe filaments pass down 1 Merker, ‘‘ Gunnera macrophylla,” Inaugural Dissertation, 1888. NOSTOC. 543 the slime-canal into the gland itself, and there occupy the space left by the glandular cells on their conversion into slime. Thence the filaments find their way into the intercellular spaces of the starch-containing parenchyma surrounding the gland, and become closely applied to the cell-walls. The Nostoec then bores through or dissolves the cell-wall, absorbs the starch, and grows vigorously till it fills the whole cell. In this manner the Nostoe spreads through the cortical parenchyma of the Gunnera stem from cell to cell. The stem glands in course of time become inactive and the canal closes up, so that the Wostoe is Fic. 321.—Anabaena cycadearum. A, Coral-like surface-roots of Cycas. B, Trans- verse and longitudinal sections of A; the double line indicates the distribution of the Nostoc. (v. Tubeuf del.) completely shut in. In this condition it is absolutely de- pendent on nourishment derived from the host-cells, and seems to thrive on it. No outward symptoms of disease can be observed on Gunnera with enclosed Nostoc, the local destruction of the cell-contents, the loss of starch, and the filling up of the tissues with filaments of Vosfoc having apparently no effect. The species of Gunnera have a very short stem with a growing point hardly raised above the level of the soil, so that the Nostoc easily finds its way there. No algae have been found in the petiole and lamina of the gigantic leaves. Gunnera may easily be cultivated although it contains no Nostoe. Jonsson! regards Nostoe qunanerae as identical with NV. puneti- g Y / ! Jiénsson, Botan, Notiser, 1894. 544 THE PATHOGENIC ALGAE. forme (the earlier name); he also believes that it exists on damp soil and independently of Gunnera. Anabaena cycadearum (Reinke).' [Nostoc commune (Schneider).”]_ The following account of this species is taken from De Bary.® Seedlings of Cycadeae have a thick tap-root which branches in the soil; from the proximal end of the primary root a few pairs of root-branches grow up _ perpen- dicularly, and, after forking once or twice, their ends swell to form tubercles (Fig. 321). Similar clumps of forked twigs Fic. 322.—Anabaena cycadearum. Section of a Cycad-root, as in Fig. 521 4, and in the condition shown in the upper transverse section B of that figure. The Nostoc-filaments crowd the intercellular spaces and cause alteration in shape and growth of the adjacent cells as shown. (v. Tubeuf del.) arise later on other branches which arise from the tap-root and spread over the surface of the ground. It is into these forked twigs that the ostoc makes its way and causes the following characteristic alteration in their structure. A layer of parenchyma, which in normal roots does not differ from the surrounding compact polygonal tissue, becomes in attacked roots a definite zone round the axile vascular bundle. The zone consists of parenchymatous cells much elongated in one direction, and with their interspaces filled with masses of algal filaments (Fig. 522). In cross-sections of attacked roots the WVostoc 1 Reinke, Botan. Zeitung, 1879, and Abhandlungen, 1873. 2 Schneider, Botanical Gazette, 1894, p. 25. 3De Bary, Die Erscheinung der Symbiose, 1879, p. 14. ANABAENA. 545 zone generally forms a circle; in longitudinal section the cylinder of blue-green algae does not extend quite up to the growing point. According to Reinke, the alga penetrates into the newly-formed intercellular spaces of the developing periblem- cortex, and remains confined to the zone which it has first excited to increased growth. It is not certain whether the Nostoc penetrates only into injured places or into natural fissures. ; The branched aerial masses of tubercles on Cycad-roots are produced independently of the Nostoc but their function is unknown beyond a suggestion that they are organs of respira- tion. They certainly receive no injury from invasion by the Nostoe. Since the Nostoc lives completely cut off from the outer world and frequently in subterranean roots, we must assume that it receives nourishment from the host. Reinke found Anabaena in roots of Cycas, Ceratozamia, Dioon, and Encephalartos. Reinke has also found very fine fungal mycelia in the roots of Cyeads. Schneider observed intracellular bacteria in root- tubercles free from Nostoc. Anabaena azollae Strasb.” This endophyte is never absent from Azolla, neither A. caroliniana so much cultivated in hot- houses, nor the wild species found in America, Africa, Asia, and New Holland. The algal filaments are present even in the neighbourhood of the vegetative point and in the closed indusia of the sporangia. They are, however, most abundant in the cavities formed in the epidermis of the fleshy floating leaves. The Anabaena filaments do not enter the cavity by the opening found in the completed structure, but find their way in during the formation of the cavity, and probably influence its development. As the cavity becomes filled with Anabaena, some cells of its inner walls grow out as segmented branched filaments amongst the coils of the alga, probably in consequence of a stimulus exerted by the Anabaena. No endophytic Schizophyceae are known in the true mosses, ' Moebius (/oc. cit.) states that the roots of Cycads at the Botanic Garden, Heidelberg, never contain Anabaena. *Strasburger, Ueber Azolla, Jena, 1873; also Practical Botany (English Edition by Hillhouse, 1889), *The leaves of Azolla are divided into two parts, the upper fleshy one of which tloats on the water, the under membranous one being submerged. 2M 546 THE PATHOGENIC ALGAE. but several inhabit Hepaticae, chiefly species of Anthoceros, Blasia, Pellia, Aneura, Diplolaena, Sauteria, and Riccia, Nostoc lichenoides Vauch.' is a common endophyte in the mucilage-cavities of Anthoceros laevis. The motile algal filaments gain admission through the stomata or mucilage-fissures on the lower side of the thallus. Only one filament is admitted into each cavity, then the opening is closed by an increased turgescence and growth of the guard-cells; the imprisoned Nostoc multiplies to form a colony. Leitgeb states that after infection has successfully taken place, and frequently before the stoma has quite closed, the guard- Fia. 323.—Anabaena azollae. Longitudinal section through the posterior lobe of a floating leaf of Azolla caroliniana. The cavity is filled with Nostoc-filaments and septate hairs. 1, A septate hair, and a filament of Anabaena. (v. Tubeuf del.) cells divide and ultimately form a three-layered covering of cells over the intercellular space. Simultaneously all the thallus- cells round the infected cavity undergo radial division and grow into the cavity, first as papillae, then as much-branched and septate tubes of various lengths; the space left between them becomes meanwhile filled with the ostoc. In the case of Anthoceros laevis the tubes form a kind of pseudoparenchyma ' Bibliography: Janczewski, Botan. Zeitung, 1872, and Annales d. , sci. natur., Ser. 5, xvi. Milde, Botan. Zeitung, 1851. Leitgeb, Akademie d. Wissen- schaften in Wien, 1878; also Untersuchungen iiber Lebermoose, 1. Goebel, ‘‘ Die Muscineen,” in Schenk’s Handbuch d. Botanik, 1882. NOSTOC. 547 with interspaces filled with Nosfoc. If other algae, e.g. Oscillaria, enter the cavities, the opening is neither closed nor do the walls grow out as processes. Infection by Nostoc only occurs when the mucilage-cavities lie near the apex of the thallus and are secreting mucilage, the substance which evidently stimulates the Nostoe to enter. Leitgeb found many Anthoceroteae (Dendroceros, Nototylas, Anthoceros) with mucilage-cavities containing Nostoc, not sunk in the thallus as with Axnthoceros laevis, but forming warty pro- jections above it. In the case of Dendroceros these occurred on the upper side of the thallus as well as on the lower. The openings of the cavities of Votfothylas do not close after infection, but distinctly open wider. Janczewski observed that chlorophyllous cells of Hepaticae, though at first uninfluenced by the intruding alga, afterwards lose their chlorophyll and plasma; hence he assumed that the imprisoned Nostoc begins in time to live a parasitic life, and to kill the host-cells. This, however, is not supported by other authorities. Goebel, on the other hand, believes that the Vosfoe, like the mucilage amongst which it grows, is useful to the thallus, and that it ultimately completely replaces the mucilage. Prantl held that the alga assimilated free nitrogen, giving up the product to the hairs in the cavity; but this is ex- tremely unlikely, especially when the Vostoe is completely enclosed in its host. Nostoc lichenoides is also very frequent in the leaf-auricles on the under side of the thallus of Blasia pusilla. The auricles contain mucilage, which probably induces the Nosfoe to enter. As a result of the Nosfoc invasion the auricle enlarges and continues to live, whereas without this it would soon have died off. Branched filamentous processes are produced from the inner wall of the auricle and grow amongst the WNostoc.' B. THE TRUE ALGAE. In these Algae the green chlorophyll is limited to certain portions of the plasma, the chromatophores. The true Algae are capable of sexual reproduction. They are all more or less adapted to an aquatic life. Many of them live in symbiosis, some are true parasites. ' Waldner, Akad. d. Wissenschaften in Wien, 1878. D48 THE PATHOGENIC ALGAE. The true Algae may be grouped as follows: (1) Conjugatae ; (2) Chlorophyceac; (3) Rhodophyceae; (4) Phacophyceae; (5) Characeae. Of these the Characeae includes no endophytes, the Conjugatae, Phaeophyceae, and Rhodophyceae only species endophytic in other algae or in animals. The Chlorophyceac, however, include a large number of species which live as “aerial algae” endophytic in Phanerogams, either as shelter-parasites or as true parasites. t. CHLOROPHYCEAE. These are divided into three groups:! 1. Protococcoideae including the families Volvocaceae, Tetra- sporaceae, Chlorosphacraceae, Pleurococcaceac, Protococcaceae (Endos- phaeraceae, Characieae, Sciadiaceae), and Hydrodictyaceae. 2. Confervoideae including the families Ulvaceae, Ulothrichaceae, Chaetophoraceae Mycoideaceae, Cylindrocapsaceae, Oedogoniaceae, Coleochactaceae, Cladophoraceae, Gomontiaceac, and Sphaeropleaceae. 3. Siphoneae including the families Sotrydiaceae, Phyllo- stphonaceae, Derbesiaceae, Vaucheriaceae, Bryopsidaceae, Cauler- paceae, Codiaceae, Valoniaceae and Dasycladaceae. Chlorosphaeraceae. Chlorosphaera endophyta Klebs. This is found between the living epidermal cells of Lemna minor, and produces there spherical cell-masses visible to the naked eye as_ wart-like swellings. According to Frank, this is related to Hndocloniwm polymorphum Frank. Entophysa charae Mob. This lives under the cuticle of the epidermal cell-wall of Chara Hornemanni m Brazil. Endosphaeraceae. Most of the species can penetrate into living organs, but they may also live as saprophytes or vegetate as independent organ- isms. That all the Hndosphacraceae are injurious to their host has not as yet been proved. Chlorochytvium includes eight European species all endophytic in living plants. The arrangement used by Wille in Hngler-Prantl. natur. Pflanzen-familien. gs J CHLOROCHYTRIUM. 549 Chlorochytrium lemnae Cohn.' The zygozoospores have four cilia and swarm for a short time in water. On plants of Lemna trisulea, the ciliated end becomes applied to the epidermis at the place where two cells are in contact, the zoospore becomes spherical, forms a membrane, and comes to rest (Fig. 524). It remains resting for a day or two, and assimilates so that a starch-grain is formed inside it. Next, a transparent process is given off which finds its way between the epidermal cells, widens out, and absorbs the cell-contents, while the portion of the algal Fia. 324.--Chlorochytrium lemnae. a, b, c, Ger- Fic. 325.—Chlorochytrium lemnae minating spores penetrating through the epi- in Lemna trisulea. a, Mature; 3b, dermis and between the cells of the parenchyma. young; c, discharged sporangia. A surface view shows the germinating spores (After Klebs.) from above. (After Cohn.) cell remaining outside becomes filled with cellulose and forms a firm button-like process. The young alga continues to make its way between the cells into the intercellular spaces of the subepidermal layers of parenchyma, preferably taking up its quarters in the thin anterior margin of the thalloid shoot, and avoiding the larger air-spaces. The zoospores are formed by repeated division of the plasma of the original cell; they are enclosed in a gelatinous mass which swells and ruptures the membranes of the alga as well as the tissue of the Lemna. 'Cohn, Beitrage zur Biologie d. nied. Organismen ; Klebs, Botan, Zeitung, 1881. 550 THE PATHOGENIC ALGAE. The zoospores (gametes) copulate in the gelatinous mass which escapes, and break out from it as free swarming zygozoospores. When the Lemna falls to the bottom in autumn, or when it dries up, the cells of the alga become resting-cells capable of sustaining drought. Plants of Lemna seem to be little disturbed by attacks of the endophyte, and develop their flowers normally. Chl. Knyanum Kirchn.- Frequents Lemna minor and L. gibba, Ceratophyllum demersum, EHlodea canadensis, but not Lemna trisulca. It forms zoospores only, and these on penetrating into a host do not produce a cellulose button like the species just described. They appear to be able to enter the host only by the stomata. Chl. pallidum Klebs. Grows in the intercellular spaces of Lemna trisulea. Chl. viride Schroet. Found in the respiratory cavity of Rumex obtusifolius. Other species occur in dead Phanerogams or in Algae. Stomatochytrium limnan- themum Cunningh. Inhabits the respiratory cavity of leaves of Limnanthemum indicum im India. Chlorocystis Cohnii Reinh. Occurs as a “ shelter-parasite ” dit Si pears Merle io Peleneegse in marine algae: Harton ma a few weeks previously. Scotinosphaera paradoxa Klebs. Found between the cells of dying leaves of Hypnum and Lemna triscula. Endosphaera biennis Klebs. The zygozoospores have four cilia, and swarm in water till they reach a living leaf of Potamogeton lucens. They generally come to rest on the under- side of a leaf at the boundary wall between two cells, and become invested in a membrane. A process is next sent in between the epidermal cells, and all the cell-contents pass over into it, the outer portion dying away. The young alga now makes its way into the intercellular spaces of the sub-epidermal tissue and becomes a resting spore. In spring this spore cives off biciliate gametes, which, after copulating, become zygozoospores. This shelter-parasite has not yet been observed ENDOSPHAERA. 551 to have an injurious effect on its host, beyond killing a few isolated leaves. End. rubra Schroet. occurs in leaves of Mentha aquatica and Peplis Portula. Phyllobium dimorphum Klebs. Found in leaves of Lysi- machia Nummularia, Ajuga reptans, Chlora serotina, Erythraea Centaurium. This endophyte may either penetrate into living leaves and there go through its. life-history, or it may do so in dead leaves. The zygozoospores have only two cilia, and enter the leaves chiefly through the stomata of the lower surface. Inside the leaf they form long filaments, which make their way between the elements into the vascular bundles of the leaf-ribs, and follow the course of the spiral vessels. Resting-spores are formed, and give the veins of the leaf a rosary-like appearance. Male and female gametes are pro- duced from the resting cells, and copulate to form zygozoospores. The host-plants are not injured by this endophyte. Chaetophoraceae. Most of the species are aquatic algae which live independent or as epiphytes. Endoclonium polymorphum Frank (see Chlorosphaera endo- phyta Klebs). This form lives Fic. 327.—Entoderma Wittrockii in the : . ° P membrane of a species of Alga (Kelocarpus). endophytic and sometimes intra- 4, A young one-celled plant just after pene . ° tration into the alga. 3B, Plant composed of cellular in living or dead leaves _ several cells in the wall of Eetocarpus. ©, . : Plant which has formed sporangia of swarm- of Lemna. spores, one of which is discharged and ° a r: another is in the act. (After Wille.) Entoderma Wittrockii Wille occurs inside the wall of Eetocarpus (Fig, 327). Periplegmatium and Phaeophila live endophytic in living algae. Trentepohlia endophytica (Ieinsch). In living cells and intercellular spaces of Jungermanniaceae (e.g. Frul/ania dilatata) and kills them. 552 THE PATHOGENIC ALGAE. Mycoidaceae. Cephaleuros Mycoidea, Karsten! (Mycoidea parasitica, Cunning.).2 This alga is epiphytic on the leaves of most trees and‘ shrubs in the tropics. It varies considerably in its appear- Fic. 328.—Cephaleui'os Mycoidea. Section through the epidermis of a leaf of Camellia, showing the thallus-like disc with haustoria. (After Canningham.) ance, but generally forms flattened thalloid discs several layers of cells thick and attached firmly to leaves by means of rhizoids (Fig. 328). Hairs are produced from the thallus-dises, especially Fic. 329.—Cephaleuros Mycoidea. Section through part of an attacked leaf of Camellia. The epidermal layer has been ruptured, and haustoria from the algal dise penetrate to the tissues. The dark-shaded portion is that killed by the alga. (After Cunningham.) the older ones; in addition, sporangial structures are also developed and give off biciliate swarm-spores. The discs form a kind of cuticle which becomes completely fused with that of the leaves. 1 Karsten, Annal. du jardin. botan. de Buitenzorg, Vol. X., 1891. 2Cunningham, Vrans. of Linnean Soc. of London, 1880; H. M. Ward (idem), 1884. MYCOIDACEAE. 553 Where this occurs, black patches are frequently formed so that the leaves become spotted, but the injury to the host-plant is by no means so severe as in the following species. Ceph. parasiticus Karsten. This species is common on the leaves of Calathea and Pandanus at Buitenzorg. It spreads through the whole leaf-tissue blackening and killing it. The epidermis is blistered and its cells filled with the alga; ultimately the cuticle is ruptured and the stalked sporangia are pro- duced. The swarm-spores germinate in the stomatal cavity, or in the adjacent intercellular spaces. Ceph. minimus Karsten is parasitic on leaves of Zizyphus Jujuba at Buitenzorg. It permeates the leaf-parenchyma and kills it, the cells after death becoming completely occupied by the alga. Phyllosiphonaceae. Phyllosiphon arisari Kuhn.’ This is a true parasite as yet observed only on Arisarum vulgare in Italy and the South of France. It causes death of the leaves and is frequently very abundant. The thallus consists of unicellular, non-septate, much branched filaments containing chlorophyll, and filling up the intercellular spaces of the spongy parenchyma of the host. The wall of the filaments gives the reactions for cellulose and consists of an outer and a later-formed inner layer, the latter capable of swelling very much to assist in ejaculating the spores. The chlorophyll corpuscles at first contain no starch, only oil, which, however, decreases during spore-formation, while the starch increases. The spores (aplanospores) are formed inside the algal threads, and are ejected with great force from the extremities of filaments which lie under stomata, and therefore in the position where least resistance is offered to the swelling inner wall. Chlorophyll is not present in the young filaments, but it appears in the older parts, especially about the time of spore-formation, and seems to be stored in the spores. The spores have a nucleus and chlorophyll disc. They germinate to a filament which grows between two epidermal cells into the intercellular spaces of the leaf. 'Kuhn, ‘Eine neue parasitische Alge,” Sitzungsber. d. naturforsch. Ces., Halle, 1878; Just, Botan. Zeitung, 1882 ; Schmitz (idem). 554 THE PATHOGENIC ALGAE. Parts of the leaf and petiole inhabited by this alga appear externally as yellow spots. Only one individual alga inhabits each spot, sending its numerous branches into the intercellular spaces. Attacked leaf-cells lose their chlorophyll and_ starch, © the latter being at first replaced by oil. The cells, however, remain alive and turgescent, even when deprived of almost their whole content ; they die, when spores are produced in the filaments. Cells undisturbed by the alga remain unaffected. A Fic. 330.—Phyllosiphon arisari on Avisarum vulgare. A, Leaf with yellowish spots; B, the branched alga isolated from a spot; C, spore-formation inside a filament; D, spores. (After Just and Engler-Prantl.) single leaf may bear a large number of spots, and all the plants of a locality are generally attacked. The spots were found by Just only from December to April, then they disappeared, indicating that the algal spores must have a_ resting-period outside of the Arisarwm, and return to young plants again in autumn. Phytaphysa Treubii Weber et v. Bosse. Forms characteristic galls on a species of Pilea in Java. 2, PHAEOPHYCEAE. These algae live only in other living algae, and are not en- dophytic in higher plants. PHAEOPHYCEAE. 555 Streblonemopsis irritans Val. forms wart-like galls on Cystosira opuntioides. Entonema grows between and into the cells of Rhodophycea and Melanophyceae. 3. RHODOPHYCEAE. The Rhodophyceae or Florideae occur endophytic only in other algae; e.g. Harveyella mirabilis (Reinsch) vegetates in thalli of Rhodomeleae and Polysiphoneae, but reproduces itself outside its host. Species of Choreocolaxy and other genera have a similar mode of life. I. INDEX OF A abietina, Phoma, - abietinum, Aecidium, Pi; Fusicoccum, abietis, Cenangium, - », Chrysomyxa, - ,, Lophodermium, - abietis-pectinatae, Caeoma, abrodens, Mystrosporium, abundans, Aecidium,-— - acaciae, Aecidium, - acericola, Phyllosticta, », Laphrina, acerina, Cercospora, af Dermaiea, - 3 Melasmia, - a Taphrina, - acerinum, Leptothyrium, 35 Rhytisma, aceris, Oidium, ,, Phleospora, - », Uncinula, acetosae, Depazea, - an Puccinia, - x Uromyces, actaeae, Aecidium, - Actinonema, - - acutatus, Uromyces, adiposus, Agaricus, adoxae, Puccinia, - ,, Ustilago, - - adusta, Sclerotinia, - adustum, Colletotrichum, aecidioides, Melampsora, Aecidium-forms, 377 465 251 379 240 418 518 411 410 463 151, 154 513 253 242, 480 147, 151, 153 479 242 499 478 Mey 465 359 334 PARASITES. aegopodii, Caeoma, - - a Puccinia, aegra, Puccinia,— - aesculi, Aecidium, - : 5 Septoria, - = », Taphrina, - - affinis, Thecaphora, - Agaricineae, - - - Agaricus, - - - aggregata, Scleroderris, - Aglaospora, - - - agrimoniae, Melampsora, =f Uredo, agropyri, Puccinia, - an Urocystis, agrostidis, Puccinia, - albescens, Didymosphaeria, ss Puccinia, - albida, Chrysomyxa, - albidum, Phragmidium, - albomaculata, Ramularia, Albugo (see Cystopus), albulensis, Puccinia, album, Mastigosporium, - ,, Microstroma, albus, Polyporus, - alchemillae, Coleroa, Fs Uromyees, - Algae, - - - alismatis, Doassansia,_ - a Pseudopeziza, alliatum, Aecidium, - allii, Caeoma, - - 2 » Puccinia, - : », Rhizoctonia, PAGE 419 359 340 410 478 147, 151, 153 024 a I. INDEX OF PARASITES. . PAGE alliorum, Caeoma, - - - - 419 ae Macrosporium, - - 518 alneum, Leptothyrium, - - - 479 alni, Frankia, - - = - 10) ;, Microsphaera, - = = = 7G ;, Sclerotinia, - - - - 262 » Stigmatea, - : : 211 alni glutinosae, Taphrina, 150, 152, 157 alni incanae, Taphrina, 147, 150, 152, 157 alnicola, Ovularia, - - - - 601 alpestre, Bostrichonema, - - 501 alpina, Puccinia, - - - - 359 » Taphrina, - 147, 149, 153, 161 alpinum, Synchytrium, - - =e anly- alpinus, Uromyces, - - - 337 alsinearum, Peronospora, - - 134 alta, Peronospora, - - - - 134 Alternaria, - - - - $17, 518 althaeae, Colletotrichum, - - 487 althaeina, Cercospora, - - - 515 ee Phyllosticta, - - - 464 Alveolaria, - - - - - 403 ambiguus, Uromyces,_ - - - 337 amentorum, Exoascus, - : =. ¥LDT ampelinum, Colletotrichum, - - 488 Pe Phoma, - - - 467 za Sphaceloma, = #4 12. AG? ampelophagum, Gloeosporium, - 484 amphigenum, Rhytisma, - - 246 amygdalearum, Clasterosporium, - 511 amygdalinum, Gloeosporium, - - 483 amylovorous, Micrococcus, - - 531 Anabaena, - - - - - 544 anceps, Leptosphaeria, - - - 221 anchusae, Accidium, - - = BAT] Andersoni, Puccinia, — - - - 359 andromedae, Coleroa, - : - 195 - Exobasidium, - - 426 - Rhytisma, - - - 246 “f Stigmatea, - - -* 211 andropogonis-annulati, Ustilago, - 292 “2 -tuberculati, Ustilago, 292 anemones, Septoria, - - - 478 Synchytrium, - - 112 “ Urocystis, - - - 316 anemones-virginianae, Puccinia, - 360 angulata, Cercospora,_ - . - 515 angustata, Puccinia, - . - $54 Anixia, - : = : : . 178 annosus, Polyporus, annularis, Puccinia, anomala, Ustilago, - anomalum, Synchytrium, Antennaria, Anthostoma, - Anthostomella, anthoxanthi, Puccinia, anthyllidis, Uromyces, apli, Cercospora, », Phyllosticta, Apiosporium, - apocyni, Aecidium, * Septogloeum, appendiculatus, Uromyces, aquilegiae, Aecidium, arborescens, Peronospora, arctica, Melampsora, » Tilletia, arctostaphyli, Exobasidium, arenariae, Peronospora, - oP Puccinia, arenariicola, Puccinia, areola, Ramularia, - argentata, Puccinia, ari, Aecidium, ari-italici, Caeoma, ariae, Melampsora, a aristidae-cyanthae, Ustilago, - aristolochiae, Puccinia, - Armillaria, - armoraciae, Ascochyta, - s- Cercospora, - aa Ovularia, bf Septoria, - - aromaticum, Septocylindrium, Arthuri, Peronospora, arundinaceum, Helminthosporium, 516 A Napicladium, - arundinellae, Ustilago, Asanuro, Caeoma, - asarina, Pucecinia, - Aschersoniana, Schinzia, Aschersonii, Entyloma, - asclepiadeum, Cronartium, Ascoboleae, Ascobolus, Ascochyta, | Ascodesmus, 2 349, 409 349, 410 558 I. INDEX OF PARASITES. PAGE Ascoidea, - - - - 138, 141 | balsamitae, Puccinia, Ascomyces, - = enc Ae - 146 | bambusae, Neovossia, - - Ascomycetes, - - - - - 136 | barbareae, Aecidium, = - Ascospora, — - - - - - 211 | Barclayana, Neovossia, - - asparagi, Cercospora, — - = - 514 | Barclayella, - - - - AS Puccinia, - - - 341 | Barclayi, Phragmidium, - asperatum, Heterosporium, - - 516 | Bardanae, Aecidium, — - Aspergillus, - - - - - 179 | Bargellinia, - - - - asperifolii, Ovularia, 2 = = 501 | Barya, - - - aspidistrae, Ascochyta, - z - 473 | Baryanum, Pee de, - - Asterina, - - - - - 179 | Baryi, Puccinia, - - - asteris, Puccinia, - - - - 361 | basicola, Thielavia, Asteroma,— - - - - 470, 474 | Basidiomycetes, Asteromidium, = 4 : - 475 Basidiophora, = asterum, Aecidium, - - - 411 | batatas, Rhizoctonia, - astragali, Didymaria, — - - - 501 | bataticola, Phyllosticta, 56 Microsphaera, - - 176 | Batschiana, Sclerotinia, - e Uromyces, — - : - 337 | Behenis, Uromyces, - - astrantiae, Fabraea, - : - 255 | bellidiastri, Puccinia, - astroidea, Piggotia, : - - 230 Beloniella, — - 5 atragenis, Puccinia, - - - 358 | berberidis, Aecidium, atriplicis, Phyllosticta, - - - 465 % Melasmia, - aucupariae, Sclerotinia, - - - 260 As Microsphaera, aurantiaca, Thecaphora, - 325 | Berkeleyi, Puccinia, f aurea, Taphrina, - 148, 150, 154, 166 betae, Phoma, 3 : Aureobasidium, — - - 428 », Phyllosticta, - = aureum, Synchytrium, - - > blil ,, Rhizoctonia, 3 Auricularieae, - - - = G7 », Uromyces, - - aurivellus, Agaricus, S - - 462 | beticola, Cercospora, australe, Rhytisma, - = - 246 | betonicae, Puccinia, z australis, Peronospora, - - - 134 i Ustilago, = = » Puccinia, : : - 349 | betulae, Hormomyia, — - - . Taphrina, - - 150, 154 5, Sclerotinia, = - : Auteupuccinia, - - - - 340 ,, Taphrina, = W148 Autobasidiomycetes, — - - - 421 | betulina, Dothidella, — - - autumnale, Rhytisma, - - - 246 s Melampsora, - 2 avenae, Ustilago, - - - . 64, 284 ay Taphrina, 147, 149, azaleae, Hxobasidium, - - - 427 | betulinus, Polyporus, — - = azollae, Anabaena, - - - - 545 Beyerinckii, Ascospora, - ys Coryneum, - 2 B bicolor, Entyloma, - - baccarum, Sclerotinia, - - - 260 | Bidwellii, Laestadia, — - - bacilligera, Passalora, - - 506 | biennis, Endosphaera, - Bacillus, - - - 532-538 | biseptatum, Gymnosporangium, bacteriospermum, Taphina 147, 149, 153 | bistortae, Ovularia, - - Bacteria, - - - 530 | 55 Pseudopeziza, Bacterium, - = 100 143,534. 537 | a Pseudorhytisma, Balansae, Peridermium, - 2 - 417 49 Puccinia, - - balsameum, Peridermium, — - =) TAT a Rhytisma, PAGE - 355 ae fi 409 Bere il 373 - 7368 a 35t 138 arise 4, 116 - 354 - 182 42] 127 - 202 - 464 270) 337 359 255 341 479 S ATG - 356 - 468 - 464 202 334 251s - 359 - 298 - 262 - 261 149, 154 - 230 366, 367 152, 159 - 446 - 201 211, 491 - 312 - 216 - 555 401 - 5Ol - 256 - 255 - 355 246 I. bistortarum, Ustilago, - Bivonae, Uncinula,- bliti, Cystopus, — - Bloxami, Cercospora, Bolleana, Cercospora, = - Boltshauseri, Ascochyta, borealis, Exoascus, - a Polyporus, Bostrichonema, Botrytis, = = . 4, 267, boutelouae-humilis, Ustilago, brachysporum, Lophodermium, Brachypuccinia,- Brandegei, Aecidium, brassicae, Alternaria, ie Asochyta, = + Chytridium, Pe Macrosporium, oH Olpidium, Se Plasmodiophora, brassicicola, Sphaerella, - Bremia, - - - - brevipes, Uromyces, brevius, Peridermium, bromivora, Ustilago, - Brunchorstia, - - Brunchorstii, Frankia, brunnea, Ramularia, bulborum, Sclerotina, bulbosum, oacummaaamm Bulgaria, - - bullata, Puccinia, », Taphrina, - = Ustilago, - - - bullatum, Tolyposporium, bunii, Aecidium, - - ,, Puccinia, Burillia, - . butomi, Ty ee - buxi, Laestadia, - 5, Puccinia, . - Byssothecium circinans, - C caealine, aaa as Caeoma, - . calamagrostidis, Tilletia, calcea, Ramularia, - calendulae, Entyloma, 148, 150, INDEX OF PARASITES. 154, 355, 364, PAGE 298 178 127 514 515 | 473 158 439 501 499 299 233 353 411 518 473 107 518 107 524 215 131 337 415 292 481 101 502 266 363 253 353 168 288 306 409 356 322 114 217 361 201 337 418 310 502 californica, Plasmodiophora, Calonectria, Calosphaeria, - | ealospora, Tilletia, - 312 | calthae, Pseudopeziza, 33 Puccinia, Calyptospora, - Camarosporium, camelliae, Coryneum, a Meliola, - campanulae, Coleosporium, Se Marsonia, A Puccinia, cancellata, Roestelia, candida, Peronospora, candidus, Cystopus, Candolleana, Sclerotinia, Candollei, Phoma, - canescens, Entyloma, cannabis, Septoria, - capensis, Ustilago, - Capnodium, - capparidis, Cercospora, carbonarium, Phragmidium, - caprearum, Melampsora, cardui, Puccinia, », Ustilago, earicicola, Puccinia, caricis, Cintractia, - », Puccinia, » Ustilago, caricinum, Leptostroma, - caries, Tilletia, : - : carnea, Taphrina, - carneola, Ovularia, - carneum, Myxosporium, - carpinea, Dermatea, carpini, Melampsora, — - », ‘Taphrina, Carpoasci, - - carpophilum, Cladosporium, carthami, Puccinia, caryogenum, Fusicladium, caryophyllinus, Uromyces, Casparyana, Schinzia, cassandrae, Exobasidium, castagnei, Melampsora, - Podosphaera, - ” Puccinia, ” 148, 149, 154, 147, 150, 153, 370 162 168 510 355 50S 336 326 427 SOS 178 B55 560 castagnei, Sphaerotheca, castaneae, Diplodina, oe Septoria, - - catalpae, Macrosporium,- - catenulatum, Entyloma, - “ caulicola, Cercospora, — - - caulium, Melanotaenium, - ceanothi, Aecidium, - = cecidomophilus, Taphrina, cedri, Peridermium, - - cellulicola, Schinzia (Naegelia), celtidis, Ramularia, - = celtis, Taphrina, - - Cenangium, - = - cenchri, Tolyposporium, centaureae, Aecidium, Puccinia, cephalanthi, Aecidium, - - Cephaleuros, - - - Cephalotheca, cepulae, Urocystis, - cerasi, Fusicladium, - - 5, Euccinia, - - - 5. Laphrina, | 295 147, cerasina, Septoria, - - - cerastii, Aecidium, - - - ,, Melampsorella, - = cerastiorum, Fabraea, - - Ceratocystis fimbriata, Ceratophorum, - - cerealis, Gibellina, - = E cerebrum, Peridermium, Cercospora, - Cercosporella, - : cercosporoides, Gumaree osporium, Cesatii, Tuburcinia, - - chaetomium, Coleroa, - - Chaetophoraceae, - - - Charrinia, - - - g cheiranthi, Cercospora, - chelidonii, Caeoma, - chenopodii, Phyllosticta, 5 Uromyces, - Chlorochytrium, Chlorocystis Cohnii, Chlorophyceae, - - Chlorosphaera sraene - Chlorosphaeraceae, - - Choreocolax, - 148, 151, 149, 153, PAGE 173 474 478 518 313 514 314 411 152 417 326 502 153 251 306 351 353 411 552 178 316 507 355 163 476 410 370 255 469 511 220 414 513 503 489 S22, 195 551 472 515 419 465 337 549 550 548 548 548 555 I. INDEX OF PARASITES. chrysanthemi, Oidium, - a Septoria, - Chrysochytrium, - - Chrysomyxa, - - - chrysosplenii, Entyloma, * Puccinia, - Chrysospora, - - Chytridiaceae, - - Ciboria, - - cichoriacearum, Erysiphe, Cicinobolus Cesatii, cicutae, Puccinia, - cinarae, Ramularia, - cinerariae, Aecidium, § - cinerea, Botrytis, - - a Monilia, - cingens, Melanotaenium, cingulatum, Gloeosporium, cinnabarina, Nectria, — - cinnamomeus, Polyporus, cinnamonea, Dermatea, - Cintractia, - - - circaeae, Aecidium, - 5 Melampsora, e Puceinia, - - circinans, Coleroa, - - i Leptosphaeria, ah Vermicularia, - circinata, Uncinula, - circumscissa, Cercospora, cirsii, Phyllosticta, - », Puccinia, - - citri, Capnodium, ;, Meliola, - - - Cladochytriaceae, - - Cladochytrium, = - - Cladosporium, - - cladosporoides, Cercospora, Clasterosporium, - - 180, 267, Sls clavariaeforme, Gymnosporangium, Clavarieae, - - - clavellosum, Triphragmium, - Claviceps, - - - clavipes, Gymnosporangium, - claytoniae, Peronospora, clematidis, Aecidium, - Clithris, - = - = Cocconii, Tolyposporium, cochleariae, Ovularia, he sf —_ = I. INDEX OF PAGE | coerulescens, Taphrina, 148, 150, 154, 167 coicis, Ustilago, - - - - 294 colchici, Urocystis, - = - - 316 | », Uromyces, - - - 338 | Coleopuccinia, - - - - 403 | Coleosporium, - - - 4374 | Coleroa, - - - - = - 195 Colletotrichum, — - - - - 486 | Colpoma, - - - - - 248 columnare, Aecidium, - - 372, 409 | comari, Doassansia, : - - 324 | commune, Nostoc, - - - - 544 communis, Erysiphe, — - - - 175 | a Taphrina, 147, 151, 152, 157 complanatum, Peridermium, - 415, 416 | compositarum, Aecidium, - - 409 concentrica, Pestalozzia, - - 494 condylonema, Cladosporium, - - 510 confluens, Caeoma, : - - 419 confusa, Taphrina, - Sy Sy on ey BR confusum, Gymnosporangium, - 401 congesta, Graphiola, - - - 326 conglomerata, Puccinia, - - - 359 | conglutinata, Pleospora, - - - DL | conicum, Gymnosporangium, - - 391 Coniothecium, - = - erty Coniothyrium, - - - - 471 conorum, Aecidium (Peridermium), 416 consimilis, Septoria, - - - 477 controversa, Tilletia, = - - - 309 convallariae, Aecidium, - - - 410 ae Dendrophoma, - - 469 convolvulacearum, Cystopus, - FF | convolvuli, Puccinia, — - - - 341 coralloides, Urocystis, - - - 319 Cordyceps,_ - - - - - 184 cornicola, Phyllosticta, - - - 464 ae Septoria, - - - - 478 cornu cervi, Taphrina, 29, 147, 149, 153 Cornuella, - - - - = $22 Cornui, Peridermium, — - - $881, 415 cornuta, Roestelia, - - 385, 391, 392 coronata, Puccinia, - - 346 coronifera, Puccinia, : : - 346 | Corticium comedens, — - . - 452 coruscans, Peridermium, . - 416 corydalis, Entyloma, — - - - 312 - Peronospora, - . - 134 coryli, Gnomoniella, - . - 224 | PARASITES. | coryli, Mamiania, - Coryneum,— - - - Couturea, - - - Crameri, Ustilago, - crastophilum, Entyloma, - crataegi, Actinonema, - - oe Septoria, - - - 561 PAGE " 224 211, 491 475 292 312 474 476 £ Taphrina, 147, 150, 153, 166 cristatus, Uromyces,_— - crocorum, Rhizoctonia, - Cronartium, - - - cruenta, Ustilago, - - Cryptomyces, - - : Cryptosporium, Cryptostictis, - Ctenomyces, - - cubensis, Peronospora, - cucumerinum, Cladosporium, - cucumeris, Cladosporium, — - og Hypnochus, - - cucurbitacearum, Phoma, - Cucurbitaria, - - : . Cucurbitarieae, —- - - cucurbitula, Nectria, = - = 337 202 380 282 246 489 475 138, 170 134 510 510 425 469 206 204 188 Cunninghamianum,Gymnosporangium403 cupulatum, Synchytrium, — - cyani, Aecidium, - - Cyanophyceae, - - - cycadearum, Anabaena, - cyclameneae, Colletotrichum, - cyclaminis, Septoria, Cycloconium, - - cydoniae, Hendersonia, - 2 Phoma, - Cylindrosporium, - - cynodontis, Phyllachora, - Ustilago, - cynoglossi, Peronospora,- eynosbati, Cryptostictis, cypericola, Schinzia, —- _ Cystopus, > haustoria, ” oogonia, a spore-germination, cystosiphon, Pythium, cytisi, Diplodia, », Peronospora, », Phyllosticta, Cytospora, . 2N 111 409 541 O44 488 478 12, 506 475 468 488 229 204 134 475 326 28, 123 13 47 47, 60 117 vou 133 463 471 D Dacromycetes, - dactylidis, Uromyces, Darluca, - - Dasyscypha, - - Decaisneana, Schroeteria, decipiens, Endomyces, - SS Synchytrium, - i, Taphrina, - ‘5 Tilletia, - ~ decolorans, Exobasidium, deformans, Barclayella, - A Caeoma, - aa Peridermium, 3 Taphrina, sis Uromyces, - Dehnii, Beloniella, - - Delastrina, Schroeteria, - Dematieae, - - Dematophora, - > dendriticum, Fusicladium, A Venturia, - Dendrophoma, - densa, Plasmopara, - densissima, Microsphaera, dentariae, Puccinia, Depazea, - - - depazeaeformis, Sphaerella, depressum, Fusicladium, Dermatea, - - Dermateaceae, - Dermatella, - - - desmodii, Ramularia, destruens, Agaricus, - ae Brunchorstia, - Fusicladium, - 3 Oidium, ‘3 Ustilago, - devastans, Myxosporium, devastatrix, Phragmidium, Diachora, - - - dianthi, Ascochyta, - . Septoria, - - A Sorosporium, - Diatrypeae, - - - dicentrae, Aecidium, - Dicoccum, - - - dictyospermum, Pythium, didyma, Septoria, - - I. INDEX OF PARASITES. 147, 15 147, 151, Didymaria, - - Didymosphaeria, - - Didymosporium, digitalis, Ascochyta, — - digitariae, Ustilago, : digitata, Schinzia, - - digraphidis, Puccinia, - Dilophia, - - - Dilophospora, - - Dimerosporium, — - - dioicae, Puccinia, - - Diorchidium, - - - Diplodia, - - - diplodiella, Coniothyrium, Diplodina, — - - - diplospora, Ustilago, — - Dipodascus, - = dipsaci, Peronospora, = - discoideum, Exobasidium, Discomycetes, - - Discosia, - - . discosioides, Pestalozzia, dispersa, Puccinia, - - ditissima, Nectria, - - diversidens, Hydnum, - Doassansia,_— - - domestica, Ustilago, Dothidea, - - - Dothideaceae, - - Dothidella, — - - - Dothiora, - - - Douglasii, Botrytis, - drabae, Aecidium, - - ae eucciniaas - dracontii, Aecidium, dryadeus, Polyporus, - dryadis, Didymosphaeria, Duriaeana, Ustilago, Dyscomycopsis rhytismoides, E | echinata, Ustilago, - echinatum, Triphragmium, echinulatum, Heterosporium,- Kcetrogella, - - - effusa, Peronospora, - effusum, Fusicladium, Ehrenbergii, Sorosporium, Elaphomyces, - - - ~ PAGE - 501 - 218 - 490 - 473 - 291 - 326 - 349 - 222 - 479 178 - 3d] - 361 - 472 471 - 474 - 291 - 138 - 1382 - 427 240 - 480 - 494 347, 548 - 186 - 431 - 322 - 298 - 230 184, 229 230 - 249 269, 499 410 - 309 - 410 - 440 = 218 = 2 Or - 245 - 294 - 362 - 516 Bey. * age - 508 - 282 97, 183 elatinum, Aecidium, = elegans, Cladosporium, - Eleutheromyces, - - elliptica, Peronospora, - Ellisii, Ascochyta, - = ,, Entyloma, - - ;, Gymnosporangium, elymi, Puccinia (Rostrupia), empetri, Caeoma, - - i Chrysomyxa, - - Melasmia, - aa Rhytisma, - endiviae, Puccinia, - - Endoclonium polymorphum, endogenum, Melanotaenium, Endomyces, - - - Endophyllum, - Endosphaera, - - - Endosphaeraceae, - - Engelmanni, Peridermium, Englerianum, Aecidium, enormis, Puccinia, - - Entoderma Wittrockii, - Entomophthoreae, - - Entomosporium, - - Entonema,~ - - - Entophysa charae, - - Entorrhiza, - = - entorrhiza, Ustilago, — - entospora, Basidiophora, Entyloma,_— - - - ephedrae, Peridermium, Epichloé, - - - E epidermidis, Didymosphaeria, epilobii, Plasmopara, = - a Ramularia, ia Septoria, - »» Sphaerotheca, - epilobii-tetragonii, Puccinia, epiphylla, Tapbrina, 20, 147, 150, 152, ep Tilletia, - - epiphyllum, Cladosporium, epitea, Melampsora, erectum, Aecidium, Eremascus, - - Eremothecium, - - ericae, Hypoderma, erigeronatum, Aecidium, eriobotryae, Fusicladium, INDEX OF PARASITES. 563 PAGE PAGE 18, 72, 404 | eriophori, Puccinia, - - 352 - 509 | errabunda, Phoma, 469 -, 184 | ervi, Uromyces,_ - - - - 333 - 135 | eryngii, Entyloma, - - - - 312 - 473 | Erysiphe, - - - - 175 - 313 | Erysipheae, - - 7, 170 - 402 re haustoria, - 9, 10 - 354 ye remedies, - - - 68 - 380 | erysiphoides, Oidium, 499 - 389 | erythronii, Uromyces, - - - 338 - 479 | erythrostoma, Gnomonia, 222 - 246 | esculenta, Ustilago, ~ 2 -. 294 - 3595 | esculentum, Aecidium, - 410 548, 551 | euonymi, Caeoma, - : 368, 419 - 314 uy Microsphaera, - - 176 - 141 | Eupezizeae, - - - - 270 - 403 | Euphacidieae,- -~— - - 241 - 990 | euphorbiae, gecidium, 24, 410 - 548 Pe Peronospora, 135 - 417 ~ Uromyees, - - - 334 - 409 | euphorbiae-dulcis, Melampsora, 370 - 359 | euphorbiae-sylvaticae, Endophyllum, 403 551 | euphrasiae, Coleosporium, - - 376 - 115 | Eu-puecinia, - 340 - 480 | Eurotium, — - : - - 178 - 559 | excavatus, Uromyces, 337 - 548 | Excipulaceae, - : : - - 482 - 326 | exitiosum, Polydesmus, - 291, 518 - 297 e Sporidesmium, - 22] - 127 | Exoasceae, - - - ll, 41, 52, 144 =o re galls, - - - 25, 29 - 417 | Exoascus (see Taphrina), 147, 152 190 | Exobasidiaceae, = - - 423 218 | Exobasidium, - - - - 7, 13, 423 131 oP galls, - 21, 25, 28 502 | exotica, Septoria, - 475 478 | expansa, Puccinia, - . - - 359 - 174 | extensa, Taphrina, - 148 - #41 | extensicola, Puccinia, - - 351 158 310 F - 510 | fabae, Uromyces, 333 368 | Fabraea, - 255 349 | fagi, Phytophthora, 117 138 | fagicola, Actinonema,— - - 474 138 | fallaciosa, Physalospora, 218 234 | farinosa, Melampsora, - 36S 411 | farinosum, Oidium, 173, 499 - 507 | Farlowii, Taphrina, - 147, 151, 152, 157 564 fascicularis, Botrytis, fasciculata, Taphrina, fasciculatum, Fusicladium, Fenestella, — - - Fergussoniu, Puccinia, festucae, Puccinia, » Urocystis, - ficariae, Cylindrosporium, ,, | Peronospora, - » Uromyces, - fici, Uredo, - = ficuum, Ustilago, - - I, INDEX OF PARASITES. PAGE 500 148, 149, 154 filamentosum, Peridermium, - 148, 149, 154 filicina, Taphrina, - - a Uredinopsis, filipendulae, Cylindrosporium * Triphragmium, re Urocystis, - filum, Darluca, - fimbriata, Ceratocystis, - he Gnomoniella, - - Mamiania, fimbriatum, Sphaeronema, firma, Puccinia, - Fischeri, Entyloma, - Pe Peridermium, - ro Tilletia, 55 Ustilago, - Fistulina hepatica, - flaccidum, Cronartium, - flammulae, Cladochytrium, flava, Taphrina,-— - Florideae, - - flosculorum, Ustilago, foeda, Chaetophoma, foedum, Capnodium, foliicola, Hendersonia, fomentarius, Polyporus, - Fomes (see Polyporus). fragariae, Ascochyta, ‘3 Sphaerella, fragariastri, Phragmidium, Frankia, fraxini, Actinonema, - x Aecidium, - aA Hysterographium, x Scolecotrichum, - » Septogloeum, s, Septoria, > 508 e229 2 Bhi) 349 316 489 134 = 33iy/ - 420 - 299 - 415 141, 420 - 489 362 319 = 47a 469 fructigena, Monilia, - fructigenum, Gloeosporium, frustulosum, Stereum, - fuciformis, Isaria, - Fuckeliana, Sclerotinia, fulgens, Synchytrium, fuliginosa, Scleroderris, - fulvum, Cladosporium, - Fe Polystigma, - PP Sclerotium, fulvus, Polyporus, - - Fumago, - - - fumariae, Caeoma, - fumosus, Polyporus, funerea, Pestalozzia, - Fungi, classification of, - Fungi imperfecti, - - Fusarium, - - fusca, Puccinia, = - - fusco-violaceum, Sistotrema, fuscum, Entyloma, - fuscus, Protomyces, - Fusicladium, - - - Fusicoccum abietinum, - fusiforme, Phragmidium, Fusisporium, - - Fusoma, - - - galanthi, Sclerotinia, galanthina, Botrytis, galeopsidis, Erysiphe, s5 Phyllosticta, galii, Melampsora, ,, Puccinia, - - 261, 184, 189, ganglioniformis, Peronospora, Gasteromycetes, geicola, Depazea, - - genistae, Didymosphaeria, 3 Uromyces, genistalis, Darluca, gentianae, Puccinia, - geographicum, Asteroma, geranii, Plasmopara, ss Puccinia, as Ramularia, ,, Uromyees, - Fe Venturia, Gibbera, - - PAGE 497 482 430 519 267 109 251 510 190 431 448 519 419 452 493 105 462 520 356. 433 312 141 507 465 362 521 504 I. INDEX OF PARASITES. 565 PAGE PAGE Gibberella, - - = - - 184 | grandis, Ustilago, - - 293 gibberosa, Puccinia, - - 346,354 | Graphiola, — - - 325 Gibelliana, Sphaerella, - = - 215 | grisea, Peronospora, - 134 Gibellina, - - - - - 220 | griseola, Isariopsis, - : - 519 giganteum, Peridermium, - - 416 | grossulariae, Aecidium, - - - 409 giliae, Aecidium, - - - - 4i11 a Microsphaera, - - 176 », Puccinia, - - - - 3d5 3 Phyllosticta, : - 464 gilvum, Lophodermium, - - 240 | Guarinonii, Microsphaera, — - - 176 githaginis, Magnusiella, 148, 151, 154 | Guepini, Pestalozzia, — - = - 494 gladioli, Urocystis, - - - 316 | gunnerae, Nostoc, - . - §4] glaucum, Penicillium, - - 3, 180 | guttata, Phyllactinia, - - - BS glechomatis, Puccinia, - - - 361 | Gymnoasci,— - - - - AT globosum, Gymnosporangium, - 403 | Gymnoascus, - - - - 138, 170 Bs Synchytrium, - - ‘113 | Gymnosporangium, 48, 51, 74, 332, 383 Gloeosporium, - - - - 482 ~ deformations, 18, 43 glomerulata, Tilletia, = - - - 310 glomerulosum, Clasterosporium, - 511 H glumarum, Puccinia, ~— - - - 348 | Halstedii, Plasmopara, - - - 131 a Uredo, - - - - 420 | Harknessii, Peridermium, — - - 415 glycyrrhizae, Uromyces, - - 337 | Hartigianum, Septogloeuin, - - 495 Gnomonia,~— - = - - 17, 222 | Hartigii, Melampsora, - . - 368 Gnomoniella, - - - - - 223 3 Pestalozzia,_ - - §3, 491 Goeldiana, Ramularia, - - - 502 of Polyporus, - = - 447 Goeppertiana, Calyptospora, - - 370 | Harveyella mirabilis, — - - - 555 > Melampsora, - - 370 | hederae, Septoria, - - ~ - 478 ea Ustilago, - - 298 | hedericola, Sphaerella, - - - 215 gongrogenua, Diplodia, - - - 472 | hedysari-obscuri, Uromyces, - - 338 “e Pestalozzia, - - 494 | helianthi, Aecidium, - - - 340 gossypii, Colletotrichum, - - 487 Be Puccinia, - - - 340 os Uredo, - - - - 420 | Helicobasidium, — - = - 429 gossypina, Cercospora, - - - 515 | helioscopiae, Melampsora, — - - 370 5 Sphaerella, - - - 214 | hellebori, Ramularia, = - : = 602 gracile, Helminthosporium, — - - 512 | Helminthosporium, - - 512, 516 A Phragmidium, - : - 363 | helosciadii, Entyloma, - - he » Pythium, - - - - 117 | Helotieae, - - - - - 256 gramineum, Helminthosporium, 221, 512 | Helvellaceae, - - - : - 275 graminicola, Ascochyta, - - 473 | helvetica, Puccinia, - - - 333 i> Sclerospora, - - 131 | Hemibasidii, - - : - - 275 graminicolum, Exobasidium, - - 427 | Hemileia, - - = : 32, 361 graminis, Dilophia, - - - 222 | Hemipuccinia, - - - - 353 ~ Dilophospora, - - - 479 | Hendersonia, - - - - - 474 Fr Erysiphe, - = - 175 | Hennebergii, Phoma, © - : - 467 Fe Ophiobolus, — - - - 222 | hepatica, Fistulina, - . - 452 ¥e Phyllachora, - - - 229 | hepaticae, Aecidium, — - - 409 a Puccinia, - - 75, 341 | herbarum, Cladosporium, . 4, 509 5 Scolecotrichum, . - 508 | Herpotrichia, - : : 61, 83, 198 graminum, Septoria, — - . - 477 | herpotrichoides, Leptosphaeria, - 220 a8 Typhula, : : - 431 | hesperidearum, Pleospora, — - - 22) grammica, Ustilago, - - - 294 | Heteropuccinia, — - . - B41 566 I. INDEX OF PAGE heterogenea, Puccinia, - 360 heteroica, Sclerotinia, - 263 Heterosphaeria, = - - - 249 Heterosporium, = - : - 515 heterosporium, Fusarium, 520 heterosporum, Septosporium, 518 hieracii, Puccinia, - 353 himalayensis, Chrysomyxa, 379 hippuridis, Aecidium, 336 hirsutum, Stereum, - 429 hirsutus, Polyporus, 452 hispidus, Polyporus, — - : 444 Holboelli, Puccinia, 359 holostei, Ustilago, - 297 hordei, Hormodendron, - 505 i. Lilletias = 310 » Ustilago, - 288 Hormodendron,~ - - 505 hottoniae, Doassansia, 324 hyacinthi, Bacillus, 538 af Pleospora, - - 221 hyalina, Thecaphora,— - - 324 Hydneae, - - 431 Hydnum, - - - - 431 hydrangeae, Septoria, - - 478 hydropiperis, Sphacelotheca, - 302 Hymenomycetes, - = 422 hyoseyami, Peronospora, 134 hypericorum, Melampsora, 370 Hyphomycetes, - 496 Hypnochaceae, - - 428 Hypnochus, - - - - 428 Hypocreaceae, = = = 184 Hypoderma, - - - - 233 Hypodermieae, - - - 235 Hypodermella, 234 hypodytes, Ustilago, 293 hypogaea, Niptera, 254 Hypomyces, - - - 184 hypophyllum, Cladosporium, - 511 Hysteriaceae, - - - 232 Hysterineae, - - 232 Hysterium, - - - - 237 Hysterographium, - - 232 I igniarius, Polyporus, oe “3 var. pinuum, 35, 433, 441 447 PARASITES. impatientis, Depazea,_ - - impressum, Asteroma, - - inaequale, Fusoma, - infestans, eeheeaet - inflata, Rhizina, - - inquinans, Bulgaria, - - ae Pestalozzia, - - insidiens, Pestalozzia,_ - = insititiae, Taphrina, Aja olemlioss intermedia, Doassansia, - - +5 Ustilago,- E intermedium, Pythium, - = interstitialis, Ovularia, - - ~ Puccinia, - - inulae, Coleosporium, — - - ipomoeae, Coleosporium, - = Nectria, = 2 56 Puccinia, - - ipomoearum, Vermicularia, - ipomoeae-panduranae, Cystopus, iridis, Aecidium, - - - ,, Cladochytrium, - - Pe uccinia eae - . 5) Uredo,- = - - Irmischiae, Paipalopsis, - - sariae 2 = 2 - - : Isariopsis, - - - ischaemi, Ustilago, - - italica, Urocystis, - - - italicum, Penicillium, = - = dJ Jamesianum, Aecidium, - = Jenseni, Ustilago, - = = Johansonii, 4s Urocystis, - = juglandinum, Cladosporium, - juglandis, Marsonia, : : 3 Microstroma, - - junci, Puccinia, - - ,, Tolyposporium, - ., Urocystis, - = ,, Uromyces, - = = juncicolum, Rhytisma, - - juniperi, Clithris, - - RS Stigmatea, - - juniperinum, Gymnosporangium, f Lophodermium, - Taphrina, 147, 150, 152, I. INDEX OF K PAGE Kaufmanniana, Peziza, - - - 265 kentiae, Colletotrichum, - - - 488 Kerneri, Sclerotinia, — - - - 270 Klebahni, Peridermium, - 376, 414 Kmetiana, Urocystis, - - - 319 knautiae, Peronospora, - - - 132 Knyanum, Chlorochytrium, - - 550 Kochii, Strickeria, - = - - 205 Kolaczekii, Ustilago, — - - - 292 Kolleri, Ustilago, - - = - 287 Koordersiana, Ustilago, - - - 298 Kriegerianum, Cladochytrium, - 1l4 Kruchii, Taphrina, - - #47, 150, 153 Kiihneana, Ustilago, — - - - 298 L Labrella,- = - - - - 480 laburni, Cucurbitaria, - = 6, 206 s Physalospora, - = ae DAL: laceianum, Phragmidium, - - 363 lacerata, Roestelia, - - - 385 Lachnella, — - - - - = 272 laciniata, Thelephora, - - - 429 lactea, Ovularia,_ - = - 500 lactucae, Bremia, - - - Sais Laestadia, : - Se OAS laetum, Synchytrium, — - - = AL) laevigatus, Polyporus, - - - 445 laevis, Tilletia, - . - 309 Lagenarium, Golletateiahum, - - 486 Lagerheimii, Leuconostoc, — - - 143 f Ustilago, - - - 299 lamii, Ovularia, - - : - 601 lampsanae, Puccinia, — - - - 340 Ee Ramularia, - - - 602 Janeola, Myxosporium, - - - 456 lapponicus, Uromyces, - - - 337 laricina, Sphaerella, - - ee) laricinum, Leptostroma, - : 2 212 um Lophodermium, — - - 240 laricis, Caeoma,~— - - 366, 367, 419 | » Heterosporium, - - - 616° | alban - - - 234 Lasiobotrys, - - - - 182 lathyri, Sten - - 3824 lathyrinum, Dicoccum, - - - 506 Laurencia, Taphrina, 29, 148, 149, 153 laureolae, Sphaerella, — - . - 215 PARASITES. 567 PAGE | lauri, Exobasidium, 31, 427 ledi, Chrysomyxa, - - 378 », Exobasidium, - 31, 427 »» Sclerotinia, - : - 263 leguminosarum, Rhizobium, - - 101 Leimbachii, Urocystis, 317 lemnae, Chlorochytrium, 549 a Cornuella, - =. Boe ue Olpidium, - - 107 lepidii, Aecidium, - 410 lepigoni, Cystopus, - - - 127 leproides, Oedomyces, - 313 leproidum, Entyloma, - - 313 Leptopuccinia, - - - 359 leptosperma, Peronospora, = - - 134 Leptosphaeria, - - - - 220 Leptostroma, - - - - - 480 Leptostromaceae, - - - 479 leptostromiforme, Cryptosporium, - 489 Leptothyrium, - - - 479 Letendraea, - - - - - 184 leucanthemi, Aecidium, - - 851, 409 +s Puccinia, - < - Sol Leucochytrium, — - - - 112 leucogonium, Oidium, - - 499 Leuconostoc, - - 143 leucospermum, Aecidium, - 409 lichenoides, Nostoc, - - 546 ligustri, Aecidium, - 409 ss Caeoma, - 419 liiacearum, Puccinia, - 356 limbalis, Phyllosticta, 464 limonii, Uromyces, - 334 limonis, Fusarium, - - §2) limosae, Puccinia, - 351 limosellae, Doassansia, 324 _ linariae, Entyloma,- 312 ~ Peronospora, : 18¢ Lindemuthianum, Colletotrichum,- 486 lineolatus, Uromyces, 336 Linhartiana, Monilia, 261 lini, Melampsora, 369 », Peronospora, 134 lolii, Tilletia, - . - $10 longipes, Taphrina,- 147, 151, 152 longissima, Phoma,- . - 469 “- Puccinia, 354 a Sphaerella, 215 7 Ustilago, 2038 568 ie INDEX OF PARASITES. PAGE lonicerae, Lasiobotrys, - 182 Ss Microsphaera, - - 176 lophanti, Peronospora, - - 154 lophiostomoides, Phoma, 467 Lophodermium, — - - - 235 lucillae, Sphaerella, - - - 476 Ludwigii, Saccharomyces, 141, 143 luminatum, Caeoma, - - - 419 lupini, Uromyces, - - - = Bay lutescens, Taphrina (Maguusiella), 148, 149, 154 luzulae, Urocystis, - - - - 316 55 Ustilago, - - 294 lychnicola, Ovularia, - - 500 lychnidis, Septoria, - - - 478 lycii, Microsphaera, - 176 lycopersici, Colletotrichum, - - 487 a Fusarium, 520 55 Septoria, - 477 lycopi, Aecidium, - - 41] M macrocarpum, Cladosporium, 510 macropus, Gymnosporangium, 391, 402 macrosora, Uredo, - - 420 Macrosporium, - - - = Oly macrosporum, Aecidium, - 411 > Hysterium, - 237 ne Lophodermium, 34, 237 macrosporus, Protomyces, — - 31, 138 maculaeformis, Venturia, = iis} maculans, Ascochyta, - - = Ailes maculare, Asteroma, = - - 47 maculatum, Entomosporium, 480 maculiformis, Laestadia, = 216 mr Phyllosticta, 464 Magelhaenicum, Aecidium, - 409 Magnusia,-—- - - 178 Magnusiana, Puccinia, - - 349 Magnusiella (see Taphrina), 146, 148, 151 Magnusii, Endomyces, - - - 148 nF Entyloma, 312 - Puccinia, - 350 major, Ustilago, 297 mali, Hendersonia, - 475 malorum, Sphaeropsis, 72 malvacearum, Phoma, - 469 ae Puccinia, 359 malvarum, Cercospora, - - 515 malvarum, Colletotrichum, - malvicola, Ascochyta, - - Mamiania, mammillata, Puccinia, - - Marconii, Dendrophoma, - marginalis, Ustilago, — - - marginatus, Polyporus, - - Mariae-Wilsoni, Aecidium, maritimus, Uromyces, - - Marsonia, - - - - Martianoffiana, Doassansia, - martii, Erysiphe, - - - Masseela, - - - - Mastigosporium, - maxima, Tuberculina, - . maximus, Cryptomyces, - maydis, Puccinia, - > - ;, Ustilago, - - megalospora, Sclerotinia, Melampsora, - Melaimpsorella, — - - melampyri, Aecidium, - 2p Coleosporium, = - Melanconideae, = = Melanconieae, - - - melangeae, Gloeosporium, — - melanogramma, Schizonella, - Melanomeae, - - - - Melanospora, - - - : Melanotaenium,-— - - - Melasmia, - - - - Meliola, - - - - melleus, Agaricus (Armillaria), melophthorum, Scolecotrichum, menthae, Puccinia, - - menyanthis, Cladochytrium, - mercurialis, Caeoma, - 56 Synchytrium, — - Merulius lacrymans, Mesochytrium, - - - mespili, Entomosporium, A Morthiera, », Stigmatea, - - mespilinum, Oidium, — - - metulispora, Ascochyta, - - microcephala, Claviceps, - microchaeta, Vermicularia, - Micrococcus, - - - Micropuccinia, - - = - 48, 53, 74, 143, 366, 367, 40, 46, 531, PAGE 487 473 223 355 469 298 449 410 336 506 324. 175 361 504 328 246 303 279 260 363 370 349 376 226 482 483 305 200, 184 3138 479 181 455 508 341 114 419 113 442 110 481 210 210 499 473 194 471 039 356 microsora, Puccinia, - microsperma, Passalora, Microsphaera, - - microsporum, Entyloma, Microstroma, - - Microthyrium, - - millefolii, Puccinia, - minimus, Cephaleuros, minor, Taphrina, - ,, Uromyces, - - minus, Cylindrosporium, mirabilis, Taphrina, - mirabilissima, Puccinia, - mixta, Melampsora, modestum, Bostrichonema, moliniae, Neovossia, - = Puccinia, Molleri, Ustilago, - - mollis, Polyporus, - Mollisia, : - Mollisieae, = - - = Mompa, Helicobasidium, Monilia, - - monilioides, Oidium, monoicum, Aecidium, - Monospora, - - - monotropae, Urocystis, - montana, Puccinia, - morbosa, Plowrightia, - mori, Bacterium, - - ,, Cladochytrium, - ,, Phleospora, - : », Septogloeum, ,, Sphaerella, - : moricola, Gibberella, mors-uvae, Sphaerotheca, Morthieri, Mollisia, ae Puccinia, “ Mucedineae, - - - Mucor, - . . Muelleri, Uredo, - mutila, Dothiora, Mycoidaceae, - Mycoidea, - Mycoidea, Cephaleuros, - Mycomycetes, - - myosotidis, Aecidium, x Peronospora, ” Synchytrium, INDEX OF PARASITES. PAGE 355 506 -7G 312 497 179 361 553 147, 150, 153, 164 338 489 147, 151-153 340 368 501 311 349 298 448 254 253 429 497 499 410 138 319 340 231 53 114 478 496 478 184 173 254 359 497 180 420 249 552 552 552 135 411 134 lil myrtillina, Podosphaera, Mystrosporium,~ - - Myxomycetes, - - - Myxosporium, - - Naegelia, - : < Naevia piniperda, - - - nana, Taphrina, napi, Pleospora, — - - - Napicladium, - necans, Ovularia, necator, Gloeosporium, necatrix, Dematophora, - Nectria, - - - : Nectriella, = - Nectroideae, - . - neglecta, Ustilago, - nelans, Ovularia, - - nemoralis, Puccinia, - Neovossia, - - - nepalense, Phragmidium, neriella, Cercospora, - nervisequium, Gloeosporium, - ¥ Lophodermium, nicotianae, Ascochyta, - nidus-avis, Gymnosporangium, Niesslii, Doassansia, - ;, Sphaerotheca, - nigra, Herpotrichia, - nigricans, Claviceps, = - ,, Polyporus, nigricantium, Macrosporium, - nigro-maculans, Septoria, - Niptera, - - - nitens, Caeoma, nivea, Plasmopara, nobile, Macrosporium, Nostoc, - nuda, Ustilago, nymphaeae, Entyloma, - nymphaeoides, Aecidium, \) obducens, Plasmopara, obliqua, Ovularia, oblongata, Puccinia, oblongisporium, Peridermium, obscura, Puccinia, - 147, 149, 153, 229, 375, 569 PAGE 175 518 520 vse 486 326 238 161 221 516 500 483 202 185 1s4 479 292 261 349 311 363 515 484 239 473 403 323 174 199 195 452 518 475 254 419 128 518 DAG USS 3138 410 131 501 3O4 414 352 570 I. INDEX OF PARASITES. PAGE obscurum, Polystigma, - - - 190 obtusa, Puccinia, - - - - 341 occulta, Doassansia, : - - 324 », Urocystis, - - - - 315 ochraceum, Polystigma, - - 190 Ochrospora, - - - - - 369 octoloculare, Phragmidium, - = 38i{h3) oenotherae, Aecidium, - - - 411 FA Puccinia, - - - 355 officinalis, Polyporus, — - - - 452 Oidium, - - - - 144, 175, 499 oleae, Bacillus, - - - - §32 oleaginum, Cycloconium, - - 506 Oleina, - = - - - = Bhs) olida, Tilletia, - - - - 310 oligochaetum, Colletotrichum, - 486 olivacea, Ustilago, - - - - 294 olivaceum, Penicillium, - - - 180 Olpidiaceae, - . - - - 106 Olpidiopsis, - - - - =e alOn Olpidium, - - - - - 106 omnivora, Phytophthora, 71, 83, 117 onobrychidis, Aecidium, : - 410 se Diachora, - - = 250) Oomyces, - - - - = lltey! Oomycetes, - - - - - 115 Oospora, - - - - - 497 Ophiobolus, - - - - - 222 orchidis, Caeoma, - - - 368, 419 oreophila, Sclerotinia, - - - 259 oreoselini, Puccinia, - - Pes )3) orientale, Peridermium, - - - A415 ornamentale, Aecidium, - - 410 ornithogali, Heterosporium, - - 516 ne Urocystis, - - - 316 5 Uromyces, - - - 338 35 Ustilago, - 5 - 299 orobanches, Urocystis, - - - 319 orobi, Uromyces, - - - - 333 oryzae, Piricularia, - - 5 105} ,, Sclerotium, - - - 266 yy. Jhlieie - - - 310 ,, Ustilaginoidea, - - = Sill ossifragi, Entyloma, - - - 313 ostryae, Taphrina, - 148, 150, 154 Ovularia, - - - = 2615000) oxyacanthae, Phleospora, - - 478 3 Podosphaera, — - - 174 oxybaphi, Peronospora, - - - 135 oxycocci, Exobasidium, - A Sclerotinia, oxyriae, Puccinia, - oxystoma, Valsa, - - iP pachydermus, Protomyces, padi, Asteroma,— - - ,, Cylindrosporium, - ,, Melampsora, - - », sclerotinia, - - Paipalopsis, — - - - paliformis, Puccinia, pallescens, Thecaphora, - pallidum, Chlorochytrium, pallidus, Uromyces, _ - pallor, Ascochyta, - - paludosa, Puccinia, - pandani, Melanconium, - Se Nectria, - - panici-frumentacei, Ustilago, - », -leucophaei, 3 ;, -miliacei, a pannosa, Sphaerotheca, - papillatum, Phragmidium, eS Synchytrium, parasitica, Botrytis, - rr Mycoidea, - ry Peronospora, - ae Septoria, = A Tetramyxa, AA Trichosphaeria, parasiticum, Fusoma, = - parasiticus, Cephaleuros, paridis, Puccinia, Parlatorei, Ustilago, parnassiae, Aecidium, - 35 Uromyces, - paspalus-dilatati, Ustilago, Passalora, - - - pastinacae, Cercosporella, Patella, Heterosphaeria, - Pazschkei, Puccinia, - Peckiana, Puccinia, Peckii, Aecidium, - : ,, EHxobasidium, = - ,, Peridermium, = - pedicillata, Thelephora, - pedicularis, Aecidium, - PAGE 426 259 355 224 141 470 489 370 261 322 354 325 550 338 473 351 188 188 292 291 289 499 363 109 500 552 133 475 529 196 504 552 349 298 352 334 294. 506 503 249 359 419 411 427 417 429 410 oe Pellicularia, - - - penicillariae, Tolyposporium, - - penicillata, Roestelia, - Penicillium, - - - pentastemonis, Aecidium, Penzigi, Meliola, - - perdix, Thelephora, - perennans, Ustilago, — - periclymeni, Aecidium, - 35 Ascochyta, - os Leptothyrium, Peridermium, - - - Periphlegmatium, - - Perisporiaceae, Perisporieae, - - - Perisporium, - - - Peronospora, - - Peronosporeae, - “e remedies, perplexans, Puccinia, - persicae, Cercospora, — - + Cercosporella, - ae Phyllosticta, - persicina, Tuberculina, - persistens, Puccinia, Personii, Quaternaria, Pestalozzia, - Pestalozzina, - - = petasitidis, Coleroa, Petersii, Aecidium, petroselini, Septoria, Pezicula (see Dermatea),- Peziza, - = : - Pezizeae, - - - Phacidiaceae, - - - phacidioides, Sphaeronema, Phacidium, - - - Phaeophila, - - Phaeophyceae, - Phakospora, - - - phalaridis, Puccinia, phaseoli, Phytophthora, - a Uromyces, phaseolina, Phyllosticta, phillyreae, Aecidium, a Uredo, - phlei-pratensis, Puccinia, Phleospora, phoenicis, Graphiola, - 3,4, 35, 429, 241, PAGE 181 306 385, 389, 391 180 411 181 430 288 409 473 479 41] 551 170 178 179 132 115 68 349 513 503 463 327 349 226 491 494 195 410 476 253 271 253 241 255 255 551 554 361 349 122 334 404 410 420 348 478 325 I. INDEX OF PARASITES. phoenicis, Pestalozzia, Pholiota = Agaricus. Phoma, - - phomoides, Gloeosporium, Phragmidium, phragmitis, Puccinia, Phycomycetes, Phyllachora, - Phyllactinia, - - Phyllobium dimorphum,- Phyllosiphon arisari, Phyllosticta, - Physalospora, - Physoderma, - Phytaphysa Treubii, phyteumatum, Uromyces, Phytomyxa, Phytophthora, piceae, Chrysomyxa, ,, Peridermium, - picipes, Polyporus, - picridis, Entyloma, a Puccinia, - Piggotia astroidea, - Pilacreae, - - - pilificum, Synchytrium, - pimpinellae, Puccinia, P Thecaphora, pinastrella, Phoma, pinastri, Lophodermium, pinguicolae, Ustilago, pini, Brunchorstia, », Lachnella, - ,, Peridermium, - ;, Trametes, pinicola, Hypoderma, », Polyporus, pinitorquum, Caeoma, pinophilum, Apiosporium, piperatum, Gloeosporium, Piptocephalis, pirata, Roestelia, piricola, Septoria, - Piricularia, piriforme, Peridermium, piriformis, Mucor, - pirina, Phyllosticta, pirinum, Fusicladium, a Venturia, 101, 411, 358, 391, 218, I, INDEX PAGE pirolae, Chrysomyxa,— - - 380 », Melampsora,_ - - - 370 pisana, Anthostomella, - - 226 pisi, Ascochyta, — - - - - 472 ,, Cladosporium, - - = alo ,, Uromyces, - 334 pithya, Phoma, : - - - 466 pityophila, Cucurbitaria, - = 210 Placosphaeria, - - - - 471 plantaginis, Aecidium, - - - 411 a Ramularia, - - - 502 Plasmodiophora, — - - - - 524 Plasmopara, - - - - = 127 platani, Fenestella, : = - 229 Be Fusarium, - = 2 - 184 Pleolpidium, - - - - - 107 Pleonectria, - 184 Pleospora, - - 221, 511 Pleosporeae, - - - 27) Pleotrachelus, = Wy Pay, isis II(07/ Plowrightia, - - - - - 230 Plowrightii, Peridermium, 376, 414 plumbea, Ustilago, - - - 299 poae, Uromyces, - - - - 336 poarum, Puccinia, - - - - 348 podagrariae, Actinonema, - 474 5 Phyllachora, - 229 Podiosoma juniperi, - - - 465 Podocapsa, — - - - - - 138 podophylli, Phyllosticta, - - 465 Podosphaera, - - - - - 174 polemonii, Aecidium, 411 polygoni, Peronospora, - - 135 ae Puccinia, = 355 7 Uromyces, = - = 304: polygonorum, Stigmatea, - a Db polymorpha, Bulgaria, - - = 53} polypodii, Sphaerella, — - - - 215 . Uredo, - - - - 420 Polyporeae, - 2 5, 6, 17, 4383 Re action on starch, - = ay ae remedies, - - 70, 72 Polyporas,6-)) 0 = = 9 433 polyspora, Taphrina, 148, 151, 154, 168 Polystictis, - - ; = - 402 Polystigma, - roads - 7, 189 Polythrincium, 229 populi, Marsonia, - - - - 491 55 Septoria, 478 OF PARASITES. populina, Didymosphaeria, — - as Melampsora, - - Poria = Polyporus. porri, Puccinia, = - - - portulacae, Cystopus, - postuma, Peziza, - - - potentillae, Coleroa, - - a Magnusiella, - 35 Marsonia, - - As Peronospora, a Phragmidium, — - 3 Taphrina, praecox, Fusicladium, - - pratensis, Ramularia, - E prenanthis, Puccinia, — - - primulae, Puccinia, - - Pe Uromyees,_ - < primulana, Ovularia, — - - primulicola, Phyllosticta, - or Tuburcinia, princeps, Calosphaeria, - Pringsheimiana, Puccinia, — - proeminens, Uromyces, - profusa, Aglaospora, — - - profusum, Septogloeum, - proserpinacae, Aecidium, - Prosti, Pueccinia, - - Protobasidiomycetes, — - - Protomyces, - - - pruinosa, Ramularia, = - - #3 Sphaerotheca, - prunastri, Dermatella, - 5 Uncinula, - - prunellae, Aecidium, < a Asteroma, - - pruni, Puccinia, - - - 3) laphrimeas —- - prunicola, Didymaria, es Phyllosticta, - : pseudocerasus, Exoascus, pseudocolumnare, Aecidium, - Pseudocommis, - - - pseudoigniarius, Polyporus, - Pseudolpidium, — - - Pseudopeziza, Pseudophacidieae, - pseudoplatani, Septoria, - Pseudorhytisma, - pseudotuberosa, Ciboria, 218, 148, 151, 147, 151- PAGE 249 367 341 127 268 195 148 491 . 134 363 154 508 502 340 341 334 500 465 321 226 350 337 229 496 410 359 421 138 502 174 252 178 410 470 35d 154 501 463 164 409 529 440 107 254 246 478 255 27 = I. INDEX OF PARASITES. PAGE pseudotuberosa, Sclerotinia, - - 270 psoraleae, Aecidium,~— - - - 410 ptarmicae, Leptothyrium, — - - 242 4. Schizothyrium, - - 242 pteleae, Aecidium, - . - - 410 | pteridis, Cryptomyces, - - - 248 re Fusidium, - - - - 248 ie Uredinopsis, - - - 420 Puccinia, - - - - - 339 Pucciniosira, - - - 2 - 404 Pucciniopsis, - - - - - 3896 pulchella, Ovularia, - - - 500 pulicaris, Gibberella, — - - - 184 pulposum, Cladochytrium, -— - - li4 pulsatillae, Coleosporium, — - - 377 pulverulenta, Puccinia, - - - 341 punctata, Melasmia, - - - 480 punctatum, Aecidium, - = ~ + 23:.409 - Rhytisma, - - - 244 a Synchytrium,— - - 112 punctiforme, Leptostroma, - - 480 ae Nostoc,—- - - 641 punctiformis, Phyllachora, — - - 241 punctum, Synchytrium, - - », Puccinia, - Soraueri, Peridermium, - Soraueriana, Pestalozzina, sorbi, Cucurbitaria, - - - 5, Melampsora, - - - . », Phoma,- - - - - sordida, Peronospora, - - : sorghi, Bacillus, - - - : », Cintractia, - ,, Endothlaspis, > Fusicladium, s Puccinia, », Uredo, - » Ustilago, Sorosphaera, - sorosporioides, Urocystis, Sorosporium, - - - sparsa, Melampsora, : : - ,, | Peronospora, sparsus, Uromyces,- speciosum, Gymnosporangium, i Phragmidium, 376, | spergulae, Puccinia, Sphaceloma, - - - - Sphacelotheca, Sphaerella, Sphaerelloideae, Sphaeriaceae, - sphaerogena, Ustilago, sphaeroidea, Dothidea, Ovularia, ” sphaeroides, Dothiora, - 218, 1s4, | Sphaerioideae, . : : 7 495 210 369 468 134 534 302 302 508 353 420 284 530 317 325 S70 133 337 403 362 361 467 302 211 210 195 463 291 230 501 wy 576 IE Sphaeronema, - - - Sphaerophragmium, - Sphaeropsideae, —- - - Sphaeropsis, - - - - sphaeropsoidea, Phyllosticta, - Sphaerostilbe, - - - Sphaerotheca, - - - sphagni, Tilletia, spinaciae, Colletotrichum, spinificis, Ustilago, - - - spinulosus, Cystopus, = - - spiralis, Uncinula, - - spissa, Didymaria, - - splendens, Aecidium, — - Sporidesmium, - - Sporodesmium, - : spumeus, Polyporus, spurium, Sphaeronema, - squamosus, Polyporus, squarrosus, Agaricus, stachydis, Puccinia, - e Septoria, Stagonospora, - - Stahlii, Peridermium, stellariae, Synchytrium, - Stemphylium, - - ois Stereum, - = - : : stictica, Pestalozzia, stictoides, Leptosphaeria, Stigmatea, — - - - Stilbeae, - = B C stolonifer, Mucor, - Stomatochytrium, - straminis, Puccinia, Straussii, Venturia, - Streblonemopsis irritans, striaeformis, Puccinia, - striatus, Uromyces, - - - Strickeria, striiformis, Tilletia, strobi, Peridermium,~— - - is Phoma, : - strobicola, Hypoderma, - strobilinum, Aecidium, - struthiopteris, Uredinopsis, Stysanus, - - 3 suaveolens, Puccinia, — - = Trametes, - subcorticium, Phragmidium, INDEX OF PARASITES. PAGE 469 361 463 472 464 184 171 310 487 299 127 176 501 410 511 517 452 253 443 462 356 478 475 , 414 111 182 429 494 22) 210 519 180 550 347 218 5d0 347 336 204 310 , 415 468 233 407 420 519 353 455 362 subinclusa, Cintractia, xs Ustilago, subtecta, Leptosphaeria, subtilis, Coleroa, - succisae, Synchytrium, a Ustilago, suffocata, Pestalozzia, suffulta, Phyllactinia, sulcigena, Hypodermella, sulphureus, Polyporus, sylvatica, Puccinia, symmetricum, Rhytisma symphyti, Uredo, - symploci, Exobasidium, synantherearum, Coleosporium, — - Syncephalis, - - - - Synchytriaceae, Synchytrium, - syringae, Ovularia, ap tabaci, Oidium, », - Phyllosticta, tabifica, Phyllosticta eee: - taleola, Aglaospora, », - Diaporthe, - tanaceti, Puccinia, - Taphria = pees Taphrina, - Taphrinopsis, - taraxaci, Puccinia, ss Synchytrium, taxi, Capnodium, ~ ehomass = » Sphaerella, - tenuistipes, Puccinia, Tepperianus, Uromyces, terebinthi, Uromyces, teres, Helminthosporium, Tetramyxa, - - thalictri, Aecidium, Ss Entyloma, ang Puccinia, Thecaphora, - - Thecopsora, - - thelebola, Aglaospora, rs Melanconium, Thelephora, thesii, Puccinia, a4 “ alll Ustilago, - - - - a ris) utriculosa, Ustilago, — - - - 298 uvicola, Phoma, - - - - 216 V vaccinii, Exobasidium, - - - 423 i Gibbera, - - - - 204 ‘e Melampsora, - - - 370 Be Sclerotinia, : - - 256 » Synehytrium, - - 3 KO) vagans, Fumago, - - - + ltsiil Vaillantii, Ustilago, - - - 299 valantiae, Puccinia, - - ool valerianae, Puccinia,~— - - - 356 55 Uromyces, - - - 334 Valsa, - - - - - - 224 Valseae, - - - - - = OR; valsispora, Dendrophoma, — - - 469 vanillae, Calospora, - - - 485 is Gloeosporium, - - - 485 vaporarius, Polyporus, 6, 442 variabile, Heterosporium, — - - 516 I. INDEX OF PARASITES. variabilis, Ramularia, - varius, Exoascus, - - E vasinfectum, Fusarium, - - - vastatrix, Hemileia, - = venetum, Gloeosporium, - - Venturia, - - - veratri, Puccinia, - - 5 Uromyces, - Vermicularia, - - - - vermiculariaeformis, Venturia, vernalis, Endomyces,_— - - - 4 Melampsora, - veronicae, Ovularia, - - mA Puccinia, - - An Sorosphaera, - - - 55 Stysanus, - - - veronicarum, Puccinia, - - - verruculosum, Entyloma, Vialae, Uredo, - - viburni, Plasmopara, — - : 2 aA Ramularia, - - viciae, Ascochyta, - ,, Peronospora, - ,, Phyllosticta, - vineae, Puccinia, - - - - vinosa, Ustilago, - - - violacea, Rhizoctonia, - a Ustilago, - . - 27, violaceum, Phragmidium, — - violae, Cercospora, - - - - ,, Gloeosporium, - - - ,, Peronospora, - », Phyllosticta, ,, . Puccinia, - - - - », Urocystis, SG. 2 veole virgaureae, Puccinia, - - virginica, Taphrina, - virgultorum, Dothidea, - viride, Chlorochytrium, - viridis, Cylindrosporium, vitellinae, Melampsora, - viticola, Cercospora, —- 5 Plasmopara,— - viticolum, Cladochytrium, 5 Cladosporium, vitigena, Leptosphaeria, vitis, Aureobasidium, - ,, Plasmodiophora, - », Sphaerella, - 148, 150, Tae lem I. INDEX OF PARASITES. PAGE vitis, Uredo, - - = - - 420 Willkommii, Peziza, Volkensii, Ravenelia, - - - 403 Winteriana, Puccinia, - Volutella, - - - - - 520 Wittrockii, Entoderma, - Vrieseana, Ustilago, - - - 299 Vuijkii, Ustilago, - - - ~- 294| x vulgaris, Botrytis, - : : - 500 Xanthoxyli, Aecidium, - », Phyllosticta, - - - 464 Xenodochus, - - vulpinae, Puccinia, : ; - 351 | xXylostei, Anthostoma, 2 Z Ww zonata, Tilletia, — - Warburgiana, Epichloé, - - - 191 | Zopfia, Warmingii, Exobasidium, - - 427 | Zopfiella, - Westendorpii, Thecaphora, - - 325 Zopfii, Puccinia, Willkommii, Dasyscypha, — - - 271 | Zygomycetes, - ] lf GENERAL INDEX OF HOST-PLANTS, COMMON NAMES, Eice. PAGE A Abies, Aecidium, 404, 409 Agaricus, 457, 461 | Apiosporium, - 181 | Caeoma, = - 418| Calyptospora, =) 372)| Lophodermium, - 239 Nectria, - - - 185 Peridermium, - 417 Pestalozzia, - - 493 Phoma, - - - 465 Polyporus, 439, 442, 448, 449, 450 Rhizina, - - 274 Sclerotinia, - - 270 Trametes, - - 453 Trichosphaeria, - 196 Abietineae, Mycorhiza, 96 Acacia, Aecidium, - 410 Ravenelia, —- - 403 Uromyces, - \ - 338 Acer, Cercospora, ols Cylindrosporium, - 489 Dermatea,- - = a8} Dyscomycopsis, - 245 Leptothyrium, - 479 Melasmia, - 242, 480 Nectria, - - = 15} Oidium, - - - 499 Pestalozzia, - 493 Pezicula, - - 253 Phleospora, - - 478 Phyllosticta, - 463 Polyporus, 435, 444, 452 PAGE Rhytisma, - 242, 244 Septogloeum, - 495 Septoria, - - 478 Taphrina, 151-154, 168 Uncinula, sire ize! Achillea, Cystopus, - 127 Leptothyrium, - 249 Protomyces, - - 141 Puccinia, - 361 Schizothyrium, - 242 Synchytrium, - 113 Aconitum, Puccinia, - 356 Urocystis, - 316 Acorus, Septocylindrium, - 505. Actaea, Aecidium, - 409 Urocystis, - - 316 Adenostyles, Aecidium, 348 Coleosporium, - 377 Niptera, - - - 254 Puccinia, - 348, 359 Uromyces, 337 Adhesion-dises = appres- sorla. Adonis, Urocystis, - 317 Adoxa, Puccinia, 341, 359 Synchytrium, - 112 Ustilago, - - 299 | Aecidium, Tuberculina, 327 Aegopodium, Actinonema, - - 474 Caeoma, - 419 Phyllachora, - 229 Protomyces, - 138 PAGE Puccinia, = = 39 Aesculus, Aecidium, - 410 Diplodia, - - 472 Nectria, - - - 185 Phyllosticta, - - 464 Polyporus, — - - 444 Septoria, - - 478 Slime-flux, - - 14 Taphrina,. - 151, 153 Aethusa, Puccinia, - 353 Agaricus, Endomyces, 141 Agave, Cladosporium, 509 Agrimonia, Melampsora,370 Uredo, - - - 420 Agropyrum (see also Triticum) Puccinia, - 345, 349 Ustilago, - - 293 Agrostemma, Taphrina, - 151, 154 Agrostis, Puccinia, 345-349 Tilletia, - - - 310 Ailanthus, Cercospora, 515 Aira, Entyloma, Spills} Puccinia, - - 345 Ustilago, - 294 Ajuga, Phyllobium, 551 Albizzia, Uromyces, - 338 Alchemilla, Bostrichonema, - 501 Coleroa, - - - 195 Uromyces, - - 337 Alder, black = Alnus glutinosa. PAGE Alder, white=Alnus incana. Aletris, Physalospora, 218 Algae, Chlorocystis, - 550 Entoderma, - = Dol Entonema, - = DDD Harveyella, - - 5d5 Olpidium, = - - 107 Periphlegmatium, - 551 Phaeophila, - - 551 Pythium, - ae Lee Streblonemopsis, - 555 », blue-green, - 541 », pathogenic, 539, 547 Alisma, Cladochytrium, 114 Doassansia,_ - ys: Pseudopeziza, - - 255 Allium, Aecidium, - 349 Bacteriosis, - - 539 Caeoma, - - 367, 419 Cladochytrium, - il4 Macrosporium, - 518 Peronospora, - - 132 Puccinia, sortie Rhizoctonia, - - 202 Sclerotinia, - - 266 Urocystis, - - 316 Uromyces, 337, 338 Vermicularia, - 471 Almond (see Amygdalus) Alnus, Corticium, - 452 Frankia, - - 101 Leptothyrium, - 479 Microsphaera, - 176 Mycodomatia, - 99 Mycorhiza,— - - 99 Ovularia, - ~- & DOL Passalora, - 506 Polyporus, 433, 439, 452 Sclerotinia = - - 262 Stigmatea, - - 211 Taphrina, 150, 157, 158, 166, 168 Valsa, - . - 224 Alopecurus, Pestalozzina, - - 495 Pueccinia, 345, 346, 349 Tilletia, - - 310 Alpine-rose (see Kho- dodendron) Il, GENERAL INDEX, PAGE Alsineae, Puccinia, - 361] Althaea, Cercospora,- 515 Colletotrichum, - 487 Phoma, - - 469 Phyllosticta, - 464 Puccinia, - 360 | Amarantaceae, Cystopus, - - 127 Amelanchier, Fusicladium, - =, ‘507 Gymnosporangium, 385, 391, 401, 402, 403 Oidium, - : - 499 Podosphaera, - - 173 Roestelia, 385, 391, 402 Ampelopsis, Cercospora, 515 Amphicarpaea, Synchytrium, - 109 Amygdalus (see also Prunus) Ascospora, - - 21) Cercospora, - - 513 Gloeosporium, - 483 Polystigma, - - 189 Taphrina, 150, 153, 165 Anbury, - - - 524 Anchusa, Aecidium, - 347 Andromeda, Coleroa, 195 Exobasidium, 426, 427 Rhytisma, - - 246 Stigmatea, - 211 Andropogon, Claviceps, 195 Puecinia, - 345, 354 Tilletia, - - - 310 Ustilago, - - 292 Androsace, Peronospora, - - 134 Anemone, Aecidium, 23,409 | Coleosporium, 377 Protomyces, 141 Puccinia, . 356, 358, 360 Sclerotinia, 266 | Septoria, 478 Synchytrium, - 112 | Urocystis, 316 Anethum, Puccinia, 3538 Heterosphaeria, 249 | Angelica, Fusicladium, 508 Puecinia, 340 581 PAGE Anthoceros, Nostoc, - 546 Anthoxanthum, | Puccinia, - 345, 354 | Tilletia, - - 310 Anthracnose, Blackberry, - - 483 Cotton, 487 Privet, - - - 485 Raspberry, 483 Vine, - - 467 Anthriscus, Plasmopara, 128 Puecinia, 340 Anthyliis, Tiehieweene 337 Apium, Cercospora, 514 Entyloma, 312 Macrosporium, - 518 Phyllosticta, 464 Puccinia, 353, 355 Septoria, - - 477 Apocynum, Aecidium, 411 Septogloeum, - - 496 Apple (see Pyrus Malus) -blight, - - - 531 -rot, - - 482 powdery mildew, 174 Appressoria, = - - 9 Apricot (see Prunus) Aquilegia, Aecidium, 349, 409 Arabis, Aecidium, 410 Puccinia, 359, 361 Ustilago, 297 Aralia, Triphragmium, 362 Arctium, Aecidium, - 351 Arctostaphylos, Exobasidium, - - 427 Melampsora, - 370 Arisarum, Phyllosiphon,553 | Aristida, Ustilago, 294 | Aristolochia, Puccinia, 341 Armeria, Uromyces, 334 Armoracia, Ascochyta, 473 | Alternaria, 518 Cercospora, 5l4 Cystopus, 126 Ovularia, 500 Septoria, 477 | Arnica, Entyloma, 312 _Arnoseris, Entyloma,- 312 582 PAGE Aronia, Gymnosporangium, 391 Roestelia, - - 392 Arrhenatherum, Exobasidium, 427 Puccinia, 345 Tilletia, - 310 Urocystis, 316 Ustilago, - - 288 Arrowroot (see Maranta) Artemisia, Peronospora, 134 Puccinia, - = 3{pll Artichoke (see Cynara) 7 Jerusalem (see Helianthus) Arum, Aecidium, 349, 410 Caeoina, - 419 Ustilago, = 299 Arundinaria, Ustilago, 293 Arundinella, Ustilago, 294 Asarum, Puccinia, 32, 359 Asclepias, Aecidium,- 411] Ash (see Fraxinus ex- celsior) Ash-canker, - > -6p8} Asparagus, Cercospora, 514 Puccinia, - - 341 Rhizoctonia, - a B40 Aspen (see Populus tremula) Asperula, Phacidium, 241 Puecinia, - 341, 353 Aspidistra, Ascochyta, 473 Aspidium ) Asplenium { (see Ferns) Aster, Aecidium, 351, 411 Basidiophora, = 127 Puccinia, - - 361 Astragalus, Didymaria, 501 Microsphaera, eG Polystigma, 190 Thecaphora, 324 Uromyces, - - 337 Astrantia, Fabraea, - 255 Puccinia, - - 359 Athamantha, Puccinia, 340 Atragene, Puccinia, - 358 Urocystis, - - 316 Atriplex,Cladochytrium,114 II. GENERAL INDEX. PAGE Phyllosticta, - - 465 Atrophy, - - 22, 26 Aubernage, 202, 534 Autoecism, - « 45 Avena, Cladosporium, 509 Erysiphe, - Se} Fusarium, - = 512 Fusicladium, - - 508 Helminthosporium, 512 Phoma, - - - 467 Puccinia, 345, 346, 348 Scolecotrichum, - 508 Septoria, - - 477 Uromyces, - 336 Ustilago, - 284, 287 Azolla, Anabaena, - 545 B Bacterial diseases or Bacteriosis, - - 530 Bacteroids; - - 102 Bambusa, Neovossia, 311 Puccinia, - - 354 Barbarea, Aecidium, 409 Barberry (see Berberis) Barley (see Hordeum) Bartsia, Coleosporium, 376 Batatas, Ceratocystis, 469 Cladosporium, = oll Cystopus, = - 27 Nectria, - - 189 Phyllosticta, - - 464 Rhizoctonia, - - 202 (see also Ipomoea) Bean (see Vicia) ,, Kidney(see Phaseolus) Beech (see Fagus) Beet-root and Sugar Beet (see Beta) Beet, heart rot, - - 464 a eo, ee ay Bellidiastrum, Aecidium,~— - = Bf Entyloma, - - 312 Puccinia, - - 399 Bellis, Aecidium, noose | Berberis, Aecidium, 341, 409 | Didymosphaeria, - 218 Melasmia,-~ - - 479 PAGE Microsphaera, = 76 Puccinia, 340 Beta, -Bacteria, - 537 Cercospora, 514 Entyloma,- - - 313 Oospora, - - 497 Peronospora, - 132 Phoma, - 468 Phyllosticta, - 464 Pythium, 116 Rhizoctonia, - - 202 Sclerotinia, - - 264 Sporidesmium, 221 Uromyces,_ - - 334 Betonica, Puccinia, - 359 Ustilago, - - 298 Betula, Cladosporium, 511 Dothidea, : - 230 Dothidella, - - 230 Hormomyia, - - 262 Melampsora, - 366, 367 Microsphaera, - 176 Myxosporium, - 486 Phyllactinia, - - 178 Polyporus, 446, 449-452 Sclerotinia, - = 261 Slime-flux, - - 1438 Taphrina, 149-154, 159-161, 167 Twig-galls, - - 532 Uncinula, = - 178 Bilberry (see Vaccinium Myrtillus) Birch (see Betula alba, etc.) Bitter-rot, - - - 482 Black-knot, - = », “rot, Vine, 216, 484 +) -Lusts - 341, 419 a Shanks 469 Bladder-plums, - 155 Blane des Racines, - 202 Blight, a common name for diseases Boragineae, Erysiphe, 175 Puccinia, - 347, 348 Borago, Entyloma, - 312 Bordeaux mixture, 69, 172 Bouillie-Bordelaise, - 69 Bouteloua, Ustilago, 299 PAGE Brachypodium, Puccinia, - 346, 354 Tilletia, - - 310 Uromyces, - - 336 Bramble (see Rubus) Brand, Vine, - - 484 Brands (see Ustilagineae) Brassica, Ascochyta, 473 Alternaria, - - 518 Cercospora, - - §&14 Cystopus, . 126 Macrosporium, - 518 Olpidium, - - 107 Peronospora, - - 133 Plasmodiophora, 524 Pleospora, - ~ 22] Sclerotinia, - - 264 Sphaerella, - - 215 Sporidesmium, 221 Briza, Puccinia, - 345 Tilletia, - - 310 Bromus, Exobasidium, 427 Puccinia, - 345, 347 Sphaerella, - - 215 Tilletia, - - 310 Urocystis, - - 316 Ustilago, 292, 293 Brown-rot, - - 497 Brown-rust, - - 347 Brunissure, - 114, 528 Brusone, Rice, - - 266 Brussone, Vine, - 484 Bryzopyrum, Puccinia, 345 Buckwheat (see Fagopyrum) Buds, premature, - 20 Bulb-bacteriosis, - 538 Bunt (see Tilletia) Butomus, Cladochytrium, 113 Doassansia, - - 323 Buxus, Laestadia, - 217 Leptostroma, - - 480 Nectria, - - 188 Phoma, - - - 468 Phyllosticta, - - 464 Puccinia, - 361 C Cabbage (see Brassica) II. GENERAL INDEX, PAGE Cacalia, Coleosporium, 377 Uromyces, - 337 Cactus, Phytophthora, 118 Caeoma, Tuberculina, 327 Calamagrostis, Puccinia, - - 346 Sclerotium, - - 266 Tilletia, - - 310 Ustilago, - - 293 Calathea, Cephaleuros, 553 583 PAGE Tilletia, = 310 Tolyposporium, - 306 Ustilago, . 27, 294 Carlina, Puccinia, - 353 Carnation (see Dianthus) Calcium oxalate and fungi, 35 Calendula, Entyloma, 312. Caltha, Fabraea, = 255 Pseudopeziza, - 255 Puccinia, - - 341 Camellia, Cephaleuros, 552 Coryneum, - 491 Meliola, : - 182 Pestalozzia, - - 494 Vermicularia, 471 Campanula, Coleosporium, - 377 Marsonia, - 491 Puccinia,; - - 359 Cankers (see under Hosts) Nectria, Aecidium, ete. Cannabis, Dendrophoma, - 469 Peziza, - - - 265 Septoria, - - 477 Caper = Capparis spinosa Cercospora, - - 515 Cystopus, — - - 126] Capsella, Cystopus, - 126) Peronospora, - - 133) Plasmodiophora, - 524, Capsicum, | Gloeosporium, - 483) Caragana, Uromyces, 337 | Carduus, Puccinia, 340, 353, 359 Thecaphora, - - 324) Ustilago, - - 296 Carex, Cintractia, 302 | Leptostroma, 480 Phyllachora, - - 229 | Puccinia, 349-351, 355 _ Schizonella, - 305 bacteriosis,— - - 532 fairy ring spot, 516 rosette, - 522 rust, 336 spot, - 477 Carpinus, Dermatea, 253 Fusicladium, - 508 Gnomoniella, - 22% Mamiania, 223 Melampsora, - 370 Pezicula, 253 Phyllactinia, - - 178 Polyporus, - 447, 452 Slime-flux, - 143 Taphrina, 150-154, 162 Carrot (see Daucus) Carthamus, Puccinia, 355 Carum, Cladochytrium, 114 Puccinia, - - 356 Carya, Fusicladium, - 508 Ramularia, - - 502 Cassandra, Exobasidium, 427 Castanea, Diplodina,- 474 Pestalozzia, - 494 Phyllosticta, - 464 Polyporus, 439 Septoria, 478 Slime-flux, 143 Urocystis, 319 Castration of flowers, 27 Catalpa, Macrospo- rium, - - §18 Ceanothus, Aecidium, 411 Cedar-apples, 402 Cedrus, Peridermium, 417 Celery (see Apium) Cell-contents and fungi, 31 Cell-sap and fungi, 33 Cell-walls and fungi, 36 Cellulose-destroying fungi, 35, 38 502 149, 153 Celtis, Ramularia, Taphrina, 584 PAGE Cenchrus, Tolyposporium, - 306 Centaurea, Aecidium, - 351, 409 Puccinia, - 340, 353 Centranthus, Puccinia, 356 Cephalanthus, Aecidium, - 411 Cerastium, Aecidium, 410 Fabraea, - —eEOD Isariopsis, — - - 520 Melampsorella, - 370 Peronospora, - - 134 Sorosporium, - - 325 Ustilago, - 3 D7 Ceratophyllum, Chlorochytrium, - 550 Cereals (under Avena, Hor- deum, Secale, Triticum) Cereal-rust, 82, 84 Chaerophyllum, Actinonema, - - 474 Caeoma, - - 419 Protomyces, - 138 Puccinia, 340, 359 Chamaecyparis, Gymnosporangium, 401, 402 Pestalozzia, - - 4094 Chamerops, Anthostomella, - 226 Graphiola, 325, 326 Chamomilla, Cystopus, 127 Champignon blanc, - 202 Chara, Entophysa, 548 Cheiranthus, Cercospora, - 515 Cystopus, 126 Peronospora, - - 133 Chelidonum, Caeoma, 419 Chenopodiaceae, Peronospora, - 132 Phoma, - - 469 Chenopodium, Cladochytrium, 114 Phyllosticta, - - 465 Uromyces, 337 Cherry (see Prunus avium and Prunus Cerasus) II. GENERAL INDEX. PAGE Cherry, Bird (see Prunus Padus) Chervil (see Anthriscus) Chestnut, horse (see Aesculus) Chestnut, sweet (see Castanea) Chives (see Allium) Chlora, Phyllobium, - 551 Chloranthy, - 33, 90 Chloris, Ustilago, 299 Chlorosis, - - = Bw Chrysanthemum, Aecidium, - 351, 409 Oidium, - - - 499 Septoria, - - 478 Chrysosplenium, Entyloma, — - 312 Puccinia, - - 361 Cichorium, Puccinia, = 3358}, 315s Sclerotinia, - - 264 Cicuta, Puccinia, 5 35) Cineraria, Aecidium,- 352 Bremia, - - - 132 Circaea, Aecidium, - 409 Melampsora, - - 370 Puccinia, - = 361 Cirsium, Aecidium, - 351 Cystopus, 127 Phyllosticta, - 464 Puccinia, =) 3905, ob) Synchytrium, - - 109 Citron (see Citrus medica) Citrus, Capnodium, - 182 Cladosporium, - 509 Colletotrichum, 487 Dendrophoma, 469 Fusarium, - =) yell Fusisporium, - oo! Meliola, - = = Jiehl Penicillium, - - 180 Pestalozzia, - 494 Pleospora, - 221 Sphaerella, - - 215 Sporidesmium, - 22) Claytonia, Peronospora, 134 Clematis, Aecidium, 349, 409 Dicoccum, - Clover (see Trifolium) Clover-rust, - Club-root, - - | Cochlearia, Alternaria, - Cystopus, - Ovularia, - Coffea, Cercospora, Hemileia, - Pellicularia, - Ramularia, Coix, Ustilago, - Colchicum, Urocystis, Uromyces,_ - Colutea, Uromyces, - Comarum, Doassansia, Compositae, Aecidium, 409 Bremia, - - Cystopus, Erysiphe, Peronospora, - Plasmopara, Protomyces, - Puccinia, - Sphaerotheca, Synchytrium, Conifers, Agaricus, Botrytis, - Diplodia, Fusoma, - Pestalozzia, - Phytophthora, Polyporus, Rhizina, - Conium, Puccinia, 338 337 324 132 127 175 134 131 141 353 173 109 457 268 472 504 494 117 450, 452 273 353 Conopodium, Aecidium, 409 Puccinia, - Conservation of Host, Convallaria, Aecidium, Dendrophoma, Heterosporium, Convolvulaceae, Cystopus, - 355 PA| 349, 410 469 516 127 Convolvulus, Puccinia, 341 Thecaphora, - Copper salts as Fungi- cides, 66, 69, 122, 171, 467 324 PAGE Coralliorhiza, Mycorhiza, 97 Cork (see Wound-cork) Cornus, Cryptomyces, 248 Erysiphe,— - - 175 Phyllosticta, - - 464 Septoria, - - 478 Corydalis, Aecidium,- 410 Caeoma, - - 419 Entyloma, - = 312 Peronospora, - - 134 Corylus, Gnomoniella, 224 Mamiania,_ - - 224 Phyllactinia, - - 178 Polyporus, - 439, 444 Septogloeum, - - 496 Cotoneaster, Sclerotinia, - - 261 Cotton (see Gossypium) », frenching, - §22 Cottonwoods (see Populus) Couch-grass (see Triticum) Cowberry | (heeV accinium) Cranberry | Crataegus, Asteroma, 470 Fusicladium, - - 507 Gymunosporangium, 385, 391, 397, 401-403 Pestalozzia, - 494 Phleospora, - - 478 Podosphaera, - 174 Polyporus, - 450 Roestelia, 385, 397, 402 Septoria, - 476 Taphrina, 150, 153, 166 Crepis, Puccinia, - 353 Synchytrium, - 109 Cress (see Lepidium) », water(see Nasturtium) Croton, Aecidium, - 410 Crowberry (see Empetrum) Cruciferae, Cystopus, 124 Erysiphe, - - 175 Peronospora, - 133 Plasmodiophora, - 524 Pythium, 116 Cucumber (see Cucumis) Cucumis, Bacteria, - 536 Cladosporium, - 510 Colletotrichum, 486) Il. GENERAL INDEX. PAGE Erysiphe, - 175, 499 Hypnochus, - - 428 Macrosporium, - 518 Peronospora, - - 134 Pythium, - = 217 Scolecotrichum, - 508 Cucurbita, Bacteria, - 836 Colletotrichum, - 486 Peronospora, - - 134 Cucurbitaceae, Phoma, 469 Sclerotinia, - - 265 Sphaerotheca, - 173 Cupressus, Agaricus, 457 Gymnosporangium, 403 Pestalozzia, - - 494 Cupuliferae, Mycorhiza,_ - 93, 94 Cycads, Anabaena, - 544 Cladosporium, - 509 Cyclamen, Colletotrichum, - 488 Septoria, - - 478 Thielavia, - - 183 Cydonia, Bacteria, - 531 Cercospora, - 515 Entomosporium, 480 Gymnosporangium, 385, 391, 401, 403 Hendersonia, - - 475 Ovularia, - - 500 Phoma, - - - .468 Roestelia, - - 391 Sphaeropsis, - - 472 Taphrina, 150, 154, 168 Cynanchum, Cronartium, - 381 Cynara, Ramularia, 502 Cynodon, Phyllachora, 229 Tilletia, - - - $10 Ustilago, - - 294 Cynoglossum, Peronospora, - 134 Cyperus, Schinzia, - 326 Cytisus, Ceratophorum, 511 Cucurbitaria, - 206 Darluea, - - 474 Diplodia, - 209 Microsphaera, 176 Peronospora, - 133 PAGE Phyllosticta, - 463 Physalospora, - 218 Uromyces, 337, 338 D Dactylis, Entyloma, - 312 Epichloé, - - 191 Puccinia, - 345, 346 Sclerotium, - - 266 Tilletia, - - - 310 Uromyces, - - 336 Dahlia, Sclerotinia, - 264 Dammara, Peridermium,417 Damping-off, 116 Dandelion (see Taraxacum) Daphne, Sphaerella, - 215 Datura, Cercospora, - 515 Daucus, Heterosphaeria,249 Phoma, - 468 Plasmopara, - 128 Polydesmus, - 221 Protomyces, - 138 Rhizoctonia, - - 202 Sclerotinia, - 264 Sporidesmium, 22) Desmodium, Microsphaera, 176 Ramularia, 502 Dentaria, Puccinia, - 359 Dianthus, Ascochyta, 473 Bacteriosis, - - 5382 Botrytis, - - 500 Fusarium, 522 Heterosporium, 516 Macrosporium, 518 Puccinia, 361 Septoria, 477 Sorosporium, - 325 Urocystis, 319 Uromyces, 334, 336, 337 Ustilago, 297 Volutella, 520 Dicentra, Aecidium, - 410 Peronospora, - 134 Digitalis, Ascochyta, 473 tamularia, 502 Digitaria (see Panicum) Dill (see Anethum) Diplachne, Ustilago,- 203 586 PAGE Dipsacus, Peronospora, 132 Disposition : Chap. V. Domatia, - 93, 99, 540 Doronicum, Puccinia, 361 Doubling of Flowers, 28 Douglas Fir (see Pseudo- tsuga) Draba, Aecidium, - 410 Puccinia, - - 359 Dracaena, Ustilago, - 299 Dry-rot = Merulius. Dryas, Didymosphaeria, 218 Synchytrium, emmlel et Durra (see Sorghum) E Eau celeste, - - 69 Echeveria, Endophyllum, - 404 Echinocystis, Peronospora, - - 134 Echinospermum, Peronospora, - 134 Ectotrophic mycorhiza, 94 Egg-plant (see Solanum Melongena) Elder (see Sambucus) Eleagnaceae, Mycorhiza, 99 Elm (see Ulmus) Elodea, Chlorochytrium, 550 Elymus, Puccinia, 345, 348, 354 Ustilago, - - 293 Empetraceae, Mycorhiza, 98 Empetrum, Caeoma, - 380 Chrysomyxa, - - 380 Melasmia, — - - 479 Podosphaera, - 175 Rhytisma, — - - 246 Sclerotinia, - - 260 Endophytes, — - - 8, 11 Endotrophic, Mycorhiza, 93 Enzymes, - - 35, 37 Epacrideae, Mycorhiza, 98 phedra, Peridermium, 417 Epilobium, Melampsora, - - 370 Plasmopara, - - 13] Puccinia, - - 341 Ramularia, 502 II. GENERAL INDEX. PAGE Septoria, - - 478 Sphaerotheca, - 174 Uredo, - - - 420 Venturia, - 218 ipiphytes, - - 8, 10 Epipogon, Mycorhiza, 97 Equisetum, Pythium, 116 Eranthis, Aecidium, 409 Urocystis, - 316 Ergot (see Claviceps) Ergotism, symptoms, 193 Erica, Hypoderma, - 234 Leptosphaeria, - 22) Venturia, - - 218 Ericaceae, Exobasidium, 423, 427 Mycorhiza, - - 98 (see also Vaccinium) Erigeron, Aecidium,- 41] Basidiophora, == Py Entyloma,- - = Billy Eriksson on Cereal-rusts, 82, 332, 345 Eriophorum, Leptostroma, - - 480 Puccinia, - 352, 354 Eryngium, Entyloma, 312 Puccinia, - - 340 Erythraea, Phyllobium, 551 Eucalyptus, Didymosphaeria, - 218 Pestalozzia, - - 494 Ustilago, - - 299 Euonymus, Caeoma, 368, 419 Cercospora, - - 515 Microsphaera, = 176 Venturia, - - 218 Euphorbia, Aecidium, = - 24, 410 Endophyllum, - 403 Fusicladium, - - 508 Leptostroma, - - 480 Melampsora, - - 370 Peronospora, - = lis5 Ravenelia, - 403 Uromyces, 334, 336, 337 Euphrasia, Coleosporium, S oyils) Tuburcinia, - 5) oyll Exoasceae, parasitic, 144 PAGE F Fagopyrum, Phytophthora, a lal Ramularia, - - 502 Fagus, Actinonema, - 474 Agaricus, - - 461 Hydnum, - - 432 Myxosporium, - 486 Nectria, - - - 186 Pestalozzia, - 493 Phyllactinia, - 178 Phytophthora, a ally Polyporus, 435, 444, 450 452 Quaternaria, - - 226 Slime-flux, 143, 144 Fennel, Rhizoctonia, - 202 Ferments, - 2, 16, 35, 37 Ferns, Cryptomyces, 248 Sphaerella, - - 215 Taphrina, 29, 149, 153 Uredinopsis, - 141, 420 Urobasidium, Sty Festuca, Isaria, - - 519 Puccinia, 345-349, 354 Tilletia, - - 310 Urocystis, - 316 Ustilago, - - 288 Ficus, Cercospora, - 515 Uredo, - - - 420 Ustilago, - - 299 ‘“Pinger and Toe,” - 524 Fir, silver (see Abies) Flax (see Linum) Flower-hypertrophy, 26 Flowering, premature, 20 Flux of Trees, - 142 Fly-wood, - - 430 Forests, close and mixed, 80 Fragaria, Ascochyta, 473 Peronospora, - 134 Ramularia, = - - 214 Sphaerella, - = Bile Synchytrium, = “100 Thecaphora, - - 325 Fraxinus, Actinonema, 474 Aecidium,- - 410 Ascochyta, - - 473 Bacteria, - > GBR PAGE Cercospora, - - 515 Cylindrosporium, - 459 Hysterographium, 233 Pestalozzia, - - 453 Phyllactinia, - - 178 Polyporus, 439, 444, 445, 452 Scolecotrichum, - 508 Septogloeum, - 496 Septoria, - - 478 Fritillaria, Uromyces, 338 Fruit-rot, - 58, 115, 179 Fungi, pathogenic, - 104 » Classification, - 105 s, heteroecious, 45, 74, 331 », infodder, 85, 306 », sexuality, 104, 135 Fungicides, - - 68 Fungus-digesting plants, 92, 97 3» ~ -galls, 15, 25, 32, 33, 40, 300 » roots, - - 93 » -traps, - 92, 97 G Gagea, Puccinia, - 356 Synchytrium, sli Uromyces, - 338 Ustilago, - - 299 Galanthus, Botrytis, - 500 Caeoma, - - 368 Sclerotinia, - - 270 Galega, Uromyces, - 337 Galeopsis, Phyllosticta, 465 Galium, Melampsora, 370 Melanotaenium, 314 Phacidium, - 24) Puccinia, 341, 359, 361 Synchytrium, - 113 Gaultheria, Synchytrium, - 109 Genista, Didymosphaeria, - 218 Rhizobium, 101 Uromyces, - - 337 Gentiana, Botrytis, 268 Cronartium, - 351 II. GENERAL INDEX. PAGE Heterosphaeria, - 249 Mycorhiza, - - 97 Puccinia, - 341 Sclerotinia, - - 268 Geraniaceae, Sphaerotheca, =e 53 Geranium, Botrytis, - 269 Coleroa, - =. 195 Plasmopara, - 130, 131 Puccinia, - - 359 Ramularia, - 502 Stigmatea, - - 210 Synchytrium, - 109 Tuburcinia, - 322 Uromyces, - - 334 Venturia, - - 218 Geum, Depazea, - 465 Peronospora, - - 134 Gilia, Aecidium, - 411 Puccinia, - = 0D Gladiolus, Urocystis,- 316 Glaux, Aecidium, =! Lao Glechoma, Puccinia, - 361 Ramularia, - - 502 Gleditschia, Cercospora, Glyceria, Cladochytrium, Claviceps, - Ustilago, 293, Glycyrhiza, Uromyces, Gnaphalium, Entyloma, Golden-rust, Goodyera, Mycorhiza, Gooseberry (see Ribes) -mildew, | Gossypium, Cercospora, Colletotrichum, Fusarium, Macrosporium, Ramularia, Sphaerella, Uredo, . Gourd (see Cucurbita) Grain-smuts, 65, (see also Ustilagine: Gramineae, Ascochyta, Claviceps, Dilophia, Dilophospora, 515 ll4 194 294 337 312 348 97 173 515 487 522 518 502 214} 420 | 479 PAGE Epichloé, - 191 Erysiphe, 175 Exobasidium, 427 Fusarium, - 520 Tsaria, - 519 Mastigosporium, 504 Oidium, - 75, 499 Ophiobolus, 222 Phyllachora, - 229 Puccinia, 341-349 Scolecotrichum, - 508 Septoria, 477 Ustilago, 288, 306 Grape (see Vitis) Grape-fruit (see Citrus) Grasses (see Gramineae) Grass-mildew, - =/) 375 Gumming, 211, 491, 538 Gunnera, Nostoc, =» BAT H Habitat-races, 332 Hail-wounds, 61, 78 Hallimasch, - - 455 Hartig’s, R., chief works,37 Haustoria, 8, 10, 12 Hawthorn (see Crataegus) Hazel (see Corylus) Heart-wood, antiseptic, 5, 76 Hedera, Ascochyta, - 473 Bacteria, 533 Cladosporium, 510 Septoria, 478 Sphaerella, - - 216 Vermicularia, 471 Hedysarum, Uromyces, 338 Heleocharis, Claviceps, 195 Helianthus, Plasmopara, 131 Puccinia, 340 Ramularia, 502 Sclerotinia, 4 Helichrysum, intyloma, 312 Helleborus, Ramularia, 502 Urocystis, 316 Helosciadium (see Apium) Entyloma, - 312 | Hemi-parasite, - 3, 4, 6 Hemi-saprophyte, 3, 4 Hemp (see Cannabis) 588 PAGE Hemp-canker, - 265 Hepaticae, Nostoc, - 547 Schizophyceae, - 546 Heracleum, Heterosphaeria, - 249 Protomyces, - - 138 Puccinia, - - 340 Taphrina, - 151, 154 Heteroecism, - 406, 331 Hieracium, Entyloma, 312 Puccinia, : - 35D3 Hippuris, Uromyces, 336 Holcus, Puccinia, 346 Tilletia, - - 310 Hollyhock (see Althaea) Holosaprophytes, —- 3 Holosteum, Ustilago, 297 Homogyne, Puccinia, 359 Honey-dew, 181, 193 Honey-fungus, - 455 Honeysuckle (see Loni- cera) Hop (see Humulus) Hop-mildew, - anal 7, Hordeum, Erysiphe, 175 Helminthosporium, 22 oz Hormodendron, - 505 Phoma, - = - 467 Puccinia, - 345, 348 Tilletia, - - 310 Ustilago, - - 288 Hornbeam (see Carpinus) Horse-radish (see Armor- acia) Hottonia, Doassausia, 324 Hot-water sterilization, 66 Humulus, Capnodium, 181 Oidium, : - 499 Sphaerotheca, 173 Hyacinth (see Scilla) Hydrangea, Septoria, 478 Hypericum, Melampsora, - - 370 Hypnum, Scotinosphaeria, 550 Hypochoeris, Ramularia, - 502 Hypopitys, Mycorhiza, 93 ! II. GENERAL INDEX. I PAGE Iberis, Plasmodiophora, 524 PAGE Clithris, - - 248 Dothidea, - - 230 Nex, Diplodia, - - 472| Gymnosporangium, 384, Impatiens, Depazea, 460 389, 391, 395, 401-403 Plasmopara, - - 131] Hendersonia, - - 475 Puccinia, = - 356] Herpotrichia, - - 199 Imperata, Ustilago, - 299} Lophodermium, - 240 Indian Corn (see Zea) Pestalozzia, - - 494 Individuation, - 87| Pleospora, - == Sill Infection methods, 53,56) Polyporus, - - 450 Insect diseases, - 8,115] Stigmatea, - 2) OAT Inula, Coleosporium, 376 K Ipomoea, Coleosporium, 377 : : Baceiniat ; 356 SEER Synchytrium, 109 Vermicularia, 471 ee SIGUE INE ae Iris, Aecidium, - 5 ail Ee) CuO SD Ora: ee ee Cladochytrium, 114 Ustilago, ane = ; i s Koeleria, Puccinia, - 354 Helminthosporium, 512 E a Preciran f _ 355 Kohlhernie, or Kropf, 524 Schinzia, - > 326 1p, Uredo, - - ~- 420)|Labiatae, Erysiphe, - 175 Isopyrum, Puccinia, : - 341 Synchytrium, - 112) Laburnum (see Cytisus) Ivy (see Hedera) Lactuca, Botrytis, - 500 Ivy-canker, - - 533] Bremia, - Z = 139 Puccinia, - - 340 J Septoria, - - 477 Jasione, Puccinia, - 359|Lamium, Ovularia, - 501 Jausch, - : - 484|Lampsana, Puccinia,- 340 Jensen’s method, - 66] Ramularia, - =) 502 Juglans, Botrytis, 180 | Larch (see Larix) Cladosporium, - 510] -canker, - - = BY/l Entorrhiza, - - 326 | Larix, Agaricus, - 457 Marsonia, - 491 Caeoma,- 366, 367, 419 Microstroma, - - 497| Dasycypha, - = 27a: Penicillium, - - 180} Heterosporium, - 516 Septoria, - - 478} Hypodermella, - 234 Juncus, Puccinia, - 354] Leptostroma, - - 212 Rhytisma, — - - 246] lLophodermium, - 240 Schinzia, - - 326| Nectria, - - 188 Tolyposporium, - 305] Peziza, - - 20, 271 Urocystis, - 319} Polyporus, - 489, 452 Uromyces,_ - - 336] Sphaerella, - - 211 Ustilago, - =) 204: Trametes, 2 - 453 Jungermanniaceae, Lathyrus, Diachora, - 230 Trentepohlia, = 551 Dicoccum, - - 506 Juniperus, Agaricus, 357| Peronospora, - - 132 Botrytis, - - 270} Thecaphora, - - 324 Clasterosporium, 511} Uromyces, - 333, 334 PAGE Laurus, Exobasidium, 31, 427 Leaf-cast, - - - 236 Ledum, Chrysomyxa, 379 Exobasidium, - 427 Sclerotinia, - - 263 Leguminosae, Erysiphe,- - - 175 Mycodomatia, 101, 524 Mycorhiza, - =."2:99 Ravenelia,-— - - 403 Lemna, Chlorochytrium, 549, 550 Chlorosphaera, - 548 Cornuella, = - - 822 Endoclonium, - 551 Olpidium,— - - 107 Pythium, - ap aay Scotinosphaera, - 550 Lemon (see Citrus) Lepidium, Aecidium, 410 Cystopus, - - 126 Peronospora, - - 133 Lettuce (see Lactuca) Libocedrus, Gymnosporangium, 401 Lichen-symbiosis, - 86 Lignification of cell-walls, 37 Lignin-destroying fungi, 38 Ligustrum, Aecidium, 409 Caeoma, ~ - - 419 Cercospora, - - 515 Gloeosporium, - 485 Lilac (see Syringa) Lilium, Aecidium, 349, 410 Botrytis, - - 268 Peronospora, - - 135 Sclerotinia, - 268 Uromyces, - - 338 Lima bean (see Phaseolus) Lime or Linden (see Tilia) Limnanthemum, Aecidium, 352, 410 Stomatochytrium, - 550 Limosella, Doassansia, 324 Linaria, Mntyloma, 312 Melanotaenium, 314 Peronospora, - 134 Linum, Melumpsora, - 369 II. GENERAL INDEX. PAGE | Peronospora, - - 134 Liriodendron, Cercospora, - - 515) Cylindrosporium, - 489 Leptosphaeria, - 22] Lithospermum, Synchytrium, - iil Lolium, Fusarium, - 520 Ovularia, - - 500 Puccinia, - 345, 346 Thecaphora, - - 325) Tilletia, - - - 310 Ustilago, - - 288 Lonicera, Aecidium, 349, 409 Anthostoma, - - 226 Ascochyta, - - 473 Didymosphaeria, - 218 Lasiobotrys, - - 182 Leptothyrium, - 479 Microsphaera, - 176 Nectria, - - Bet Se Phyllosticta, - - 464 Venturia, - - 218 Lophanthus, Peronospora, - - 134 Lotus, Ovularia, - 501 Uromyces, - - 336 Lucerne (see Medicago) Lupinus, Cercospora, 515 Cryptosporium, - 489 Erysiphe, = - - 175 Pythium, - mo NT Thielavia, - = 182 Uromyces, - - 337 Luzula, Phyllachora,- 229 Puccinia, - 852, 354 Urocystis, - - 316 Ustilago, - - 294 Lychnis, Ovularia, - 500 Puccinia, - - 340 Septoria, - - 478 Sorosporium, - 825 Uromyces, 337 Ustilago, 27, 297 | Lycium, Microsphaera, 176 Lycopodium, Pythium, 116) Lycopus, Aecidium, 411) Lysimachia, Aecidium, 351 Phyllobium, 5O1 | Synchytrium, 11h) PAGE M Magnolia, Pestalozzia, 494 Mahonia, Aecidium, - 341 Maize (see Zea) Majanthemum, Aecidium, 349 |Malachium, Ustilago, 297 Maladie digitoire, 524 Mal-di-gomma, - 521 Mallow (see Malva) Mal nero, - 202, 534 Malva, Ascochyta, 473 Cercospora, - - 515 Phoma, - 469 Puccinia, - 359, 360 Mandarin (see Citrus) Mangel Wurzel (see Beta) Mangold (see Beta) Manure and Fungi, 67, 279 5, Plasmodiophora, 528 Maple (see Acer) Maranta, Epichloé, 191 Matthiola, Plasmodiophora, - 524 Medicago, Peronospora, - 133 Pseudopeziza, 255 Rhizoctonia, - 201 Sclerotinia, - 265 Tilletia, - - - 310 Uromyces, - - 336 Medlar (see Mespilus and Amelanchier) Melampyrum, Aecidium, 349 Coleosporium, - 3876 Melica, Puccinia, 346 Melon (see Cucumis) », Water (see Cucumis) Mentha, Cladochytrium, 114 Endosphaera, 551 Puccinia, : 341 Menyanthes, Cladochytrium, 14 Mercurialis, Caeoma, - 367, 419 Cercospora, DLS Synchytrium, 113 Mespilus, Fusicladium, - 50s 590 PAGE Mespilus, Gymnosporangium, 391, 397 Mucor, - - 180 Oidium, - : - 499 Ovularia, - 261, 500 Penicillium, - - 180 Podosphaera, - - 174 Meum, Triphragmium, 362 Mignonette (see Reseda) Mildew, powdery (see Ery- sipheae) re false (see Perono- sporeae) Milium, Puccinia, 345 Tilletia, - - - 310 Millardet’s vines, =) 1 o2 Millet (see Panicum and Sorghum) Mint (see Mentha) Mistletoe, - - 18, 64 Molinia, Claviceps, - 194 Fusarium, - - §20 Neovossia, - 311 Puccinia, - - 349 Monotropa, Mycorhiza, - Sis Urocystis, - - 319 Monoxeny, - 45 Morbe bianco, - e202, Morus, Bacteria, 534 Cercospora, 515 Cladochytrium, 114 Diplodia, E = 2, Gibberella, — - - 184 Helicobasidium, - 429 Nectria, - - - 185 Phleospora, - - 478 Polyporus, - 445 Septogloeum, - - 496 Sphaerella, - 215, 478 Mountain Ash (see Pyrus [Sorbus] Aucuparia) Mucilage flux, - - 142 Mulberry (see Morus) Mulgedium, Puccinia, 340 Mummification of fruits, 29 Musa, Physalospora, - 218 Muscari, Urocystis, 316 | II. GENERAL INDEX. PAGE Uromyces,_ - 338 Ustilago, - 299 Mycocecidia, 25 ; Mycodomatia, - 93, 99 Mycorhiza, - 32, 93 Myosotis, Aecidium, - 411 Entyloma,_ - 312 Peronospora, - - 134 Synchytrium, 111, 113 Myosurus, Peronospora, 134 Myrica, Ramularia, 502 Frankia, — - 101 Myricaceae, Mycodomatia, 99 Myricaria, Didymosphaeria, 218 Myrrhis, Puccinia, 340 N Narcissus, Puccinia, - 359 Nardus, Claviceps, 194 Narthecium, Entyloma, 313 Nasturtium, Cystopus, 126 Nebbia nera, - - 484 Needle-cast, 211, 236, 481 Neottia, Mycorhiza, - 97 Nepeta (see Glechoma) Nephrodium (see Ferns) Nerium, Capnodium,- 181 Cercospora, - Seeroile Nicotiana, Ascochyta, 473 Bacteria, : - 535 Erysiphe, 175 Oidium, - = 499 Peronospora, - - 134 Phyllosticta, - 464 Nucleus-parasites, 32 Nuile, - - - 508 Nuphar, Aecidium, 352, 410 Nutricism, - 88, 92 Nyctaginaceae, Peronospora, - 135 Nymphaea, Aecidium, - 352, 410 Entyloma, - 313 O Oak (see Quercus) Oak-root fungus, 200 PAGE Oat (see Avena) Oenothera, Aecidium, 411 Peronospora, - - 134 Puccinia, - 355 Synchytrium, - =e LOS Olea, Cercospora, - 515 Cycloconium, - - 506 Polyporus, - - 447 Twig-galls, - - 532 Oleaceae, Hysterographium, - 232 Oleander (see Nerium) Olive (see Olea) Rogna or Loupe, - 532 Onion (see Allium) Onion-rust, - 341, 3395 Onobrychis, Diachora, 230 Thielavia, - - 183 Uromyces, - = 8Biy/ Orange (see Citrus) foot-rot, - - = byal mal-di-gomma, 521 sooty mould, - - 182 Orchideae, Aecidium, 349 Caeoma, - - 3868, 419 Gloeosporium, - 485 Mycorhiza, - Mo SY Ornithogalum, Heterosporium, - 516 Puccinia, - = hie Urocystis, - - 316 Uromyces, - - 338 Ustilago, - - 299 Orobanche, Urocystis, 319 Oryza, Piricularia, - 503 Sclerotium, - - 266 Tilletia, - - 310 Ustilaginoidea, - 311 Ostrya, Taphrina, 150, 154 Oxalis, Sphaerella, - 215 Oxyria, Puccinia, - 355 Ustilago, - - 298 P Paeonia, Cronartium, 382 Palms, Colletotrichum, 488 Graphiola, — - - 325 Pestalozzia, - - 494 Thelephora, - 429 PAGE Pandanus, Cephaleuros, 553 Nectria, - - - 188 Panicum, Pythium, - 116 Tolyposporium, - 306 Ustilago, 289, 291, 293 Papaver, Entyloma, - 312 Peronospora, - - 133 Papilionaceae, Erysiphe,175 Peronospora, - - 132 Parasites, - - 1-3, 7, 14 classification, - - 3 mode of life, - - te Paris, Aecidium, 349, 410 Mycorhiza, - 32 Tuburcinia, - - 32) Urocystis, - - 316 Parnassia, Aecidium, 352 Uromyces, - - 334 Parsley (see Petroselinum) Parsnip (see Pastinaca) Partridge-wood, - 431 Paspalus, Ustilago, 294 Pastinaca, Cercospora, 514 Cercosporella, 503 Heterosphaeria, - 249 Pea (see Pisum) Pea-rust, - - - 334 Peach (see Persica) -curl, - - - 165 frosty mildew, - 503 -rot, 497 Pear (see Pyrus communis) | -blight, - - - 531 Pedicularis, Aecidium, 351, 410 Penicillaria, Tolyposporium, - 306 Pennisetum, Neovossia, 311 Pentstemon, Aecidium, 411 Peplis, Endosphaera, 551 Peridermium, Tuberculina, - 327 Persica (see also Prunus) Cercospora, - - 533 Cercosporella, 503 Cladosporium, 510 Clasterosporium, 511 Gloeosporium, 482 Monilia, 497 II, GENERAL INDEX, 591 PAGE PAGE Phyllosticta, - - 463 | Picea, Aecidium, 377, Puccinia, - - 355 379, 407, 416 Sphaerotheca, - 172) Agaricus, - - 457 Taphrina, 150, 153, 165) Barclayella, - 373 Petasites, Aecidium,- 348) Chrysomyxa,- 379, 380 Coleosporium, = OLE Herpotrichia, - - 199 Coleroa, - - 195 Lophodermium, 237, 240 Petroselinum, Mycorhiza, - - 95 Heterosphaeria, 249/ Naevia, - : 238 Plasmopara, - 128| Nectria,- .- - 188 Puccinia, 353; Peridermium, 416, 417 Septoria, - 477'| Pestalozzia, - - 493 Petunia, Polyporus, 440-450 Phytophthora, - 122) Septoria, - + 475 Sclerotinia, - - 264 Trametes, ° 453 Peucedanum, Trichosphaeria, 197 Puccinia, - 393, 359) Picris, Entyloma, 312 Taphrina, - 151,154) Puccinia, - - 355 Phalaris, Pilea, Phytaphysa, - 554 Cladochytrium, - 114 Pimpinella, Puccinia, - 346, 349 Puccinia, 340, 356 Sclerotium, - - 266! Thecaphora, - «- $25 Ustilago, - 294 Pine (see Pinus) Phaseolus, Ascochyta, 473) |, Cembran (see Pinus Bacteria, - - 539 Cembra) Cercospora, - - 515) ,, Mountain (see Pinus Colletotrichum, - 486 montana) Isariopsis, . - 520 ,, scots (see Pinus Phyllosticta, - - 464 sylvestris) Phytophthora, - 122 ,, Weymouth (see Pinus Sclerotinia, - 264 Strobus) Uromyces, - - 334) Pine-twister, § - - 364 Phegopteris (see Ferns) Pinguicula, Ustilago,- 297 Phillyrea, Aecidium,- 410 | Pinus, Agaricus, - 457 Uredo, - - 420 Brunchorstia, - - 481 Phleum, Epichloé, 191} Caeoma,- 364 Puccinia, - 345, 348} Cenangium, 251, 481 Phlox, Aecidium, 411 Cladosporium, - 509 Puccinia, - - 855} Cucurbitaria, - - 210 Phoenix (see Palms) Elaphomyces, 183 Phragmites, Claviceps, 194; Gymnosporangium, 396 Helminthosporium, 516) Lophodermium, 240 Napicladium, - 516} Melampsora, - 364 Puccinia, - - 349} Mycorhiza, - - 96 Ustilago, - - 293) Peridermium, 376, Phyllanthus, Ravenelia, 403 382, 411-416 Phyteuma, Phoma, - 466, 468 Coleosporium, 377 Polyporus, 449, 450 Uromyces, - - 338 Rhizina, - - 273 592 PAGE Pinus, Sistotrema, - 433 Trametes, - - 453 Twig-galls, - 532, 533 drying up of twigs, 481 Pinus Cembra, Cucurbitaria, - - 210 Nectria, - - - 188 Peridermium,- 382, 415 Pinus montana, Herpotrichia, - = IR) Hypodermella, - 234 Melampsora, - - 364 Peridermium, - - 411 Pinus sylvestris, Agaricus, - 457 Caeoma, - - - 364 Hypoderma, - - 234 Hypodermeila, - 234 Lachnella, - = ODE Lophodermium, - 235 Melampsora, - - 364 Peridermium,- 374, 376, 377, 381, 411, 414, 415 Phoma, - - 466, 468 Polyporus, - 450 Trametes, - - 453 Pinus Strobus, Agaricus,457 Hypoderma, - - 233 Peridermium, 382, 415 Phoma, - - 468 Polyporus, - 449, 450 Tuberculina, - = Bye Piptatherum, Puccinia, 346 Pistacia, Uromyces, - 337 Pisum, Ascochyta, - 473 Cladosporium, 509, 510 Erysiphe, 175 Peronospora, - 132 Pythium, - 117 Thielavia, = - - 1838 Uromyces, 334 Ustilago, 297 Plane (see Platanus) Plantago, Aecidium,- 411] Peronospora, - - 134 Ramularia, - - 502 Synchytrium, - = eee Tilletia, - - - 310 Platanus, Calonectria, 184 Il. GENERAL INDEX. PAGE Cladosporium, - 510 Fenestella, - 4!) Gloeosporium, - 484 Pestalozzia, - - 494 Polyporus, - - 445 Plum (see Prunus domestica) Plums, pocket, - 154, 156 Plum-rust, = Sans Poa, Claviceps, - - 195 Entyloma,~ - - 312 Epichloé, - > - UG) Puccinia,° - 345, 348 Tilletia, - - - 310 Urocystis, - - 316 Uromyces, - - 336 Pock of Vine, - 484 Podophyllum, Phyllosticta, - - 465 Podospermum, Puccinia,356 PAGE Mycorhiza, - = oe Polyporus, - 489, 447 Septoria, - - 478 Slime-flux, - ela y Taphrina, 150-154, 157, 166 Uncinula, - = 7S Portulaca, Cystopus,- 127 Potamogeton, Doassansia, - - 324 Endosphaera, - =O) Potassium sulphide, 173, 483 Potato (sce Solanum tuberosum) -disease, - - - 119 early blight, - - 517 -rot, - - > bet “scab, = - Dai, Potato, sweet (see Batatas) Polemonium, Aecidium, 411 | Potentilla, Beloniella, 256 Polygonatum, Cercospora, - ata) Aecidium, - 349, 41 Cladochytrium, - 114 Polygoneae, Puccinia, 355 Coleroa, - - =" 195 Polygonum, Marsonia, - - 491 Bostrichonema, - 501] Peronospora, - - 1384 Ovularia, - = 501 Phragmidium, - 363 Peronospora, - - 135) Synchytrium, = 01 Pseudopeziza, - 255| Taphrina, = eos: Pseudorhytisma, - 255) Pourriture of Vine, - 202 Puccinia, - 352, 355| Powdery mildew (see Rbytisma,- - 246 Erysipheae) Sphacelotheca, - 302) Prenanthes, Puccinia, 340 Stigmatea, - - 211) Primula, Ovularia, - 500 Uromyces, - - 334] Phyllosticta, - - 465 Ustilago, - 298, 299 Puccinia, - - 341 Polypodium (see Ferns) Tubureinia, - - 321 Polyxeny, - - - 45 Uromyces, - 21, 334 Poplar (see Populus) Primulaceae, Poppies (see Papaver) Peronospora, - - 134 Populus, Agaricus, - 462/ Privet (see Ligustrum) Ascochyta, - 473 | Proserpinaca, Aecidium, 410 Capnodium, - 181 | Prunella, Aecidium, - 410 Cladosporium, - 510] Asteroma, = - - 470 Didymosphaeria, 218| Pruning, - - =) Sri Diplodia, - - 472) Prunus, Ascospora, - 211 Dothiora, - 218, 249 Asteroma, - - 470 Fusicladium, - - 508 Botrytis, : - 269 Marsonia, —- - 491} Cercospora, - - 513 Melampsora, - 364, 367 Clasterosporium, - 511 Il. GENERAL INDEX. PAGE PAGE Coryneum, 211, 491 Melampsora, - = a0 Dermatella, - - 252) Monilia, - 261 Gloeosporium, 482/| Polystigina, 190 Monilia, - - 497| Sclerotinia, - - 261 Oidium, - = - 499 Taphrina, 151, 152, 154 Phyllosticta, - 463 Psamma, Ustilago, 293 Podosphaera, - - 174| Pseudotsuga, Agaricus, 457 Puccinia, - - 355 | Botrytis, 269 | Septoria, - - 476) Phoma, - - 466) Sphaeronema, - 253] Polyporus, 450 | Sphaerotheca, 172| Trametes, - - 453 Taphrina, 150-157, 164| Psoralea, Aecidium, - 410 Uncinula, 178 | Ptelea, Aecidium, 410 Prunus avium and P. Cercospora, 515 Cerasus, Ascospora, 211 | Pteris (see Ferns) Calosphaeria, - - 226) Pulicaria, Uromyces,- 236 Clasterosporium, 511 | Pulsatilla(see also Anemone) Coryneum, - 211, 491 | Puccinia, - - 356 Cylindrosporium, - 489) Urocystis, 316, 317 Fusicladium, - - 507) Pyrola, Chrysomyxa,- 380 Gnomonia, —- - 222) Melampsora, - 370 Monilia, - 261, 497 | Pyrus, Actinonema, 474 Penicillium, - - 180) Asteroma, 470 Phyllosticta, - 463| Bacteria, - - 531 Plowrightia, - 231) Gymnosporangium, Podosphaera, - - 174 385, 391, 402 Polyporus, 439, 449,452, Hydnum, - - 433 Puccinia, - - 355! Melampsora, - 369, 370 Taphrina, 151, 153, 163 Nectria, - : 185 Prunus domestica, Pestalozzia, - 494 Calosphaeria, - 226) Polyporus, 445, 452 Cladosporium, 510 Roestelia, 385, 391, Clasterosporium, 511 | 392, 397, 402 | Cylindrosporium, 489| Sphaerotheca, 174 Dermatella, 252 Pyrus (Sorbus) Aucuparia, Didymaria, 501) Cucurbitaria, - 210 Monilia, - 497, Gymnosporangium, Mueor, - . 180 | 385, 391 Penicillium, - - 180) Melampsora, - 369 Phyllosticta, - 463 | Phoma, - - 468 Plowrightia, - - 231) Podosphaera, - 174 Podosphaera, - 174| Polyporus, ~ 444 Polyporus, 447 Roestelia, 385, 389,391,392 Polystigma, 189 Sclerotinia, 260 Puccinia, - 855 Pyrus communis, Tapbrina, = 151-154, 164 Bacteria, 53] Uncinula, 178) Entomosporium, 480 Prunus Padus, Asteroma,470) Fusicladium, - 218, 507 Cylindrosporium, 485) Gloeosporium, - 482 2P Gymnosporangiun, 385, 391, 396, 401, Monilia, Morthiera,— - - Mucor, - Penicillium, Phyllosticta, - Plasmodiophora, - Plowrightia, - - Podosphaera, - Polyporus, Roestelia, Septoria, Sphaerella, Stigmatea, - - Taphrina, Venturia, - Pyrus Malus, Bacteria, Cladosporium, Fusicladium, - Gloeosporium, Gymnosporangium, 385, 389, 391, 402, Hendersonia, - Hydnum, - - Monilia, - Mucor, Nectria, - Oidium, - Penicillium, Phy llosticta, Podosphaera, - Polyporus, 433, 445, 385, 391, Roestelia, Slime-flux, Sphaeropsis, - Sphaerotheca, Thelephora, Venturia, Q Quercus, Agaricus, Aglaospora, Bulgaria, Capnodium, Ciboria, - Cladosporium, Clithris, - Colpoma, 439, 385, 216, 150, 154, 218, 593 PAGE 403 497 210 180 180 463 529 231 174 444 396 476 476 210 168 218 531 590 507 482 403 475 433 497 180 186 . 499 180 463 174 452 402 143 472 173 429 218 457 26 253 18] or ~i 510 248 248 594 PAGE Quercus, Corticium, - 452 Cycloconium, - - 506 Dermatea, 25383 Diaporthe, 226 Fistulina, 452 Gnomonia, 223 Hydnum, 432 Marsonia, 491 Microsphaera, 176 Microstroma, - 497 Myxosporium, 486 Pestalozzia, 494 Pezicula, 2 =) 2983 Phyllactinia, - - 178 Polyporus, 433, 439, 440, 444, 450, 452 Rosellinia, =e) Sclerotinia, - 266, 270 Slime-fux, 142, 143 Stereum, 429, 430 Taphrina, 150, 153, 167 Thelephora, 429, 430 Uredo, - 420 Quince (see Cydonia) -rust (see Roestelia) leaf-blight, - - 480 black rot, 472 R Radish (see Raphanus) Radish, horse (see Armoracia) Ranunculaceae, Erysiphe, 175 Plasmopara, - - 130 Ranunculus, Aecidium, 349, 409 Cladochytrium, 114 Cylindrosporium, - 489 Didymaria, - - 501 Entyloma, 312 Fabraea, 255 Ovularia, 501 Peronospora, - 134 Stigmatea, 210 Synchytrium, 112 Urocystis, - - 316 Uromyces, 336, 337 Rape (see Brassica) Il. GENERAL INDEX. Raphanus, Cystopus,- Peronospora, - Raspberry (see Rubus) Red-rot (due to Polyporeae) Reed (see Phragmites) Reseda, Cercospora, - Peronospora, - Resin, - » collecting, no allo, Rhamnus, Aecidium, Cercospora, Microsphaera, Rheum, Aecidium, Rhinanthaceae, Scleroderris, - Rhinanthus, Coleosporium, Sclerotium, Rhizoctonia, Rhizomorphs, Rhododendron, Apiosporium, - Chrysomyxa, - Exobasidium, Gloeosporium, Hendersonia, - Pestalozzia, Sclerotinia, Synchytrium, Rhodophyceae, Entonema, Rhus, Cercospora, Marsonia, Sphaerotheca, Taphrina, Uromyces, Ribes, Aecidium, Caeoma, - Cercospora, Cronartium, Gloeosporium, Leptosphaeria, Microsphaera, Phyllosticta, - Plasmopara, - Polyporus, Puccinia, 44, 151, 350, 363, PAGE 126 133 458 346, 349, 387 ols 176 349 337 409 419 515 382 483 221 176 464 131 452 359 Scleroderris, - Septoria, Sphaerotheca, Rice (see Oryza) Ricinus, Cercospora, - Ring-scale, Robinia, Aglaospora, - Dothiorella, - Nectria, - Polyporus, Rhizobium, Strickeria, Root-hypertrophy, PAGE 251 476 173 old 453 229 229 185 439 102 204 26 ,, tubercles, 99, 101,528,544 Rosa, Actinonema, Cercospora, - Cryptostictis, - Dicoccum, Gloeosporium, Leptostroma, - Marsonia, - Peronospora, - Pestalozzia, Phragmidium, Septoria, Sphaerotheca, Rosaceae, Oidium, Peronospora, - Phragmidium, Sphaeropsis, - Sphaerotheca, tose- Mildew, Rost of Vine, Rowan (see Aucuparia) Rubiaceae, Phacidium, Rubus, Ascochyta, Caeoma, Cercospora, Chrysomyxa, - Cladosporium, Coleroa, - Gloeosporium, Mollisia, Phragmidium, Puccinia, - Pyrenochaeta, Septoria, Uredo, Pyrus 474 515 475 506 483 480 906 138 494 362 475 172 499 134 362 472 173 172 484 241 473 419 515 380 509 195 483 254 363 341 470. 476 420 PAGE Rumea, Ustilago, - 299 Rumex, Aecidium, - 349 Chlorochytrium, - 550 Cladochytrium, - 114 Depazea, - - 465 Ovularia, - - 501 Puccinia, : - 355 Ramularia,_ - - 502 Synchytrium, a iS Uromyces, - 334, 337 Ustilago, 298 Venturia, - 218 Ruppia, Tetramyxa, - 529 Ruscus, Phyllosticta,- 465 Rust or Uredineae, 328 Rust in Australia, ete., 85 Rust, white, - 123 | Rye (see Secale) NS) Saccharum, Thielaviopsis, - 183 Trichosphaeria, .- 198 Ustilago, - 284 Saffron, Rhizoctonia,- 202 Sagittaria, Burillia, - 322 Doassania, - 323, 324 Sainfoin (see Onobrychis) Scrophularia, Ovularia, 501 Salicornia, Uromyces, 334 Salix, Capnodium, - 181 Cryptomyces, - 246 Didymosphaeria, 218 Didymosporium, - 490! Leptostroma, - 480 Melampsora, - 367, 368 Melasmia,_~ - - 480 Ovularia, : - 501 Pestalozzia, - - 494 Polyporus, 433, 439, 444, 452 Rhytisma, 245, 246 | Scleroderris, - 251 Septoria, 478 Trametes, 455 Twig-galls, 582 Uncinula, 178 | ‘Salvia, Puccinia, 341 | Sambucus, Aecidium, 411 Cercospora, 515 / Il. GENERAL Didymosphaeria, Nectria, - Phyllosticta, - Sanguisorba, Phragmidium, Sanicula, Fabraea, Puccinia, : Saponaria, Sorosporium,325_ Ustilago, - Saprophytes, - Saxifraga, Caeoma, Exobasidium, - Melampsora, - Puccinia, Synchytrium, - INDEX. d95 PAGE PAGE 218 Sedum, Aecidium, 349 185 Endophyllum, 404 464 Puccinia, 359 Septoria, 478 363 | Seed-control stations, 65 255 | Seed-sterilization, 65 341 Seedling-diseases, Fusoma, - - 540 28,297 Peronosporeae, 116, 117 1-3 | Selection against dis- - 419) ease, Sl 427 | Selinum, Puccinia, 353 370 Sempervivum, 359, Endophyllum, 403 112 Phytophthora, 118 Scabiosa, Peronospora, 132 Senecio, Aecidium, 350, 351 Puccinia, Synchytrium, - Ustilago, Scilla, Bacteriosis, Pleospora, Puccinia, Sclerotinia, Urocystis, Uromyces, Ustilago, Scirpus, Cladochytrium,114 Claviceps, Puccinia, Uromyces, Sclerotium-diseases, 29, 256 Scorzonera, Cystopus, 353, 356 Puccinia, - Ustilago, Peronospora, - Uromyces, Scrophularineae, Plasmopara, - Secale, Claviceps, Dilophia, Dilophospora, Fusarium, Helminthosporium, Leptosphaeria, Pucecinia, 345, Tilletia, - Urocystis, Ustilago, 352, 354 347, 348 361, Coleosporium, 374, 377 - 110, Ovularia, 500 294-296 Puccinia, 356, 359 538! Ramularia, 502 221| Thielavia, - - 183 356, 359 | Seseli, Puccinia, 353 266 | Sesleria, Puccinia, 349 316| Tilletia, - 310 - 338) Setaria, Sclerospora,- 131 - 299) Ustilago, - 292 Ustilaginoidea, 311 195 | Shelter-parasitism, 540 336 Capsella) 127 | Sicyos, Peronospora, - - 296 | Silene, Peronospora, - Puccinia, Sorosporium, - Uromyces, Ustilago, Silver Fir (see Abies pectinata) Silybum, Ustilago, Sium, Aecidium, Cladochytrium, Slime-flux of trees, Slime-fungi, Sloe (see Prunus) Smilacina, Heterosporium, Smilax, Aecidium, 134 338 131 19] 222 479 520 221 220 310 B15 289 Shepherd’s Purse (see Shot-hole fungi, 463, 467,513 134 Silaus, Cladochytrium, 114 134 340, 361 325 334, 337 297 296 336 ll4 142 “hh o 411 596 PAGE Smuts or Ustilagineae 275 Smuts, stinking (see Tilletia) Smyrnium, Puccinia,- 356 Snag-pruning, - 2 ie Solanum, Cercospora, 515 Phytophthora, 120 Solanum Lycopersicum, Bacteria, 536 Cladosporium, 510 Colletotrichum, - 487 Fusarium, = 520 Gloeosporium, 483 Macrosporium, - 517 Phytophthora, 119 Septoria, - 477 Solanum Melongena, Botrytis, - - 500 Gloeosporium, 482, 483 Nectria, - - - 189 Phoma, - - - 468 Solanum tuberosum, Alternaria, 517 Bacteria, = pi, Seyi Botrytis, = 268 Entorrhiza, 326 Hypnochus, - - 428 Macrosporium, 517 Oospora, 497 Peziza, - - - 268 Phytophthora, 119 Pythium, - 116 Rhizoctonia, - - 202 Schinzia, 326 Sclerotinia, - - 264 Soldanella, Puccinia,- 341 Solidago, Aecidium, - 41] Basidiophora, - - 127 Didymaria,_ - 501 Puccinia, 359 Uromyces,_ - 338 Sonchus, Bremia, 132 Coleosporium, 377 Puccinia, - 309 Synchytrium, - 113 Sorbus (see Pyrus) Sorghum, Bacteria, 534 Cintractia, - - 302 Endothlaspis, - - 302 Il. GENERAL INDEX. PAGE Fusicladium, - = 508 Puccinia, 353 Uredo, 420 Ustilago, -. 282, 284 Spergula, Puccinia, - 361 Spergularia, Cystopus, 127 Uromyces, - - 337 Spermatia and Spermo- gonia, - 55, 137, 328 Sphagnum, Tilletia, - 310 Spinach (see Spinacia) Spinacia, Cladosporium,510 Colletotrichum, 487 Entyloma, 313 Heterosporium, 516 Peronospora, - 132 Phyllosticta, - - 465 Spinifex, Ustilago, 299 Spiraea, Cylindrosporium, 489 Podosphaera, - - 174 Ramularia,_— - 502 Sphaerotheca, 173 Stysanus, - - 519 Triphragmium, 361, 362 Urocystis, - - 319 Spore, distribution, - 53 » germination, - 46 Sporobolus, Tilletia, - 310 Spraying of Plants, - 69 Spruce (see Picea) Spurge (see Euphorbia) Stachys, Puccinia, 356, 359 Septoria, - - 478 Starch and fungi, - 33 Statice, Uromyces, - 334 Steeps for fungi, = (835) Stellaria, Isariopsis, - 520 Melampsorella, 370 Sorosporium, - 325 Synchytrium, =a Uromyces, - ~ 8By/ Ustilago, 297 Stenactis (see Erigeron) Stiftia, Protomyces, - 141 Stipa, Ustilago, 293 Strawberry (see Fragaria) Streptopus, Aecidium, 349 Struthiopteris, (see Ferns) PAGE Sugar-cane (see Saccharum) Sulphur for Mildew, 68, 170 Sulphur-puff, - ail Summer-rust, - - 341 Sunflower (see Helianthus) Sunflower-rust, - - 340 Sycamore (see Acer) Symphoricarpus, Aecidium, - 411 Symphytum, Cladochytrium, 114 Entyloma, 312 Ovularia, 501 Uredo, - - 420 Symplocos, Exobasidum,427 Syringa, Bacteria, 533 Cercospora, - - 515 Diplodia, - - 472 Ovularia, 501 T Tanacetum, Puccinia, 355 Taraxacum, Aecidium, 39 Fusoma, - - - 505 Olpidium, — - a ly Protomyces, - - 141 Puccinia, - - 303 Synchytrium,- 22, 108 Tare (see Vicia) Tarring of Wounds, 77, 201 Taxus, Capnodium, - 181 Phoma, - - - 468 Sphaerella, - - 215 Teucrium, Puccinia, - 361 Thalictrum, Aecidium, 349, 352 Entyloma, - - 312 Puccinia, 356, 358 Synchytrium, - 112 Urocystis, —- 317 Thesium, Puccinia, 341 Thlaspi, Puccinia, 361 Tilletia, - - 310 Thuja, Polyporus, 450 Thujopsis, Caeoma, 30, 419 Thymus, Puccinia, - 359 Tilia, Actinonema, - 474 Cercospora, - - 515 Nectria, - - 185 Il. GENERAL INDEX. : PAGE PAGE Pestalozzia, - - 494 Tropaeolum, Pleospora, 221 Phyllosticta, - - 464 Tsuga, Peridermium, 417 Polyporus, - - 444) Trichosphaeria, 197 Tinder-fungus, - - 435) Tulipa, Botrytis, 500 | Tobacco (see Nicotiana) | Puccinia, 359 | Tomato (see Solanum Lyco-| Sclerotium, - 500 | persicum) | Ustilago, - - 299 Sleeping disease, - 520 Turnip (see Brassica) Topinambur, Sclerotinia, 264 | Turritis (see also Arabis) Tragopogon, | Urocystis, - - 319 Fusicladium, - - 508 Tussilago, Asteroma, 470 Puccinia, - - 3856} Coleosporium, 376 Ustilago, - - 296 Puccinia (Aecidium), 348 Trientalis, Tuburcinia, 319 Ramularia, - =" 502 Trifolium, Cercospora, 515 | ‘l'wig-galls, - - 532 Erysiphe, - 175, 499|Tyloses, - 76 Macrosporium, - 8517 | Typha, tae aie 516 Peronospora, - 133| Ustilago, - - 293 Phacidium, - - 255 Phyllachora,- - 229 U Polythrincium, - 229|Ulmaceae, Taphrina,- 149 Pseudopeziza, - 255|Ulmus, Apiosporium, 181 Pythium, - - 116] Asteroma, - - 470 Rhizoctonia, - 201 Ceratophorum, LD Sclerotinia, - 265) Cladosporium, - 511 Sphaeronema, - 255 Dothidella, 230 Synchytrium, 107,109} Nectria,- 185 Uromyces, 333, 336, 338 Pestalozzia, 494 Trigonella, Thielavia, 183 Phleospora, - 478 Uromyces,_ - = OO7 Phyllachora, - 496 Triticum, Bacteria, - 535 Piggotia, 230 Cladosporium, 509 Pleospora, 221 Dilophia, - - 222) Polyporus, 435, 444, 445, Dilophospora, 479 452 Erysiphe, 175| Septogloeum, - 496 Gibellina, 220] Slime-flux, 143 Leptosphaeria, 221 Taphrina, - 149, 154 Mystrosporium, 518| Uncinula, 178 Ophiobolus, - - 222] Umbelliferae, Erysiphe, 175 Phoma, - - 467| Heterosphaeria, 249 Puccinia, 345-349 Phoma, 469 Pyroctonum, - 114) Protomyces, - 138 Sclerotium, - - 431) Puecinia, 353, 359 Septoria, - 477|Umbilicus, Puccinia,- 361 Tilletia,- 306, 309, 310| Uredineae, Darluca, - 474 Typhula, - - 431) Tuberculina, - 327 Urocystis, 316} Uromyces, Darluca, - 474 Ustilago, 288, 293 | Urtica, Aecidium, 349 Trollius, Puccinia, 356 Ramularia, 502 Rhytisma, Septoria, Thecaphora, - Urticaceae, Peronospora, 135 Vv Vaccinium, Calyptospora, Exobasidium,- Gibbera, Melampsora, - Podosphaera, - Sclerotinia, 256-260, Synchytrium,- Valeriana, seit Uromyces, 423, Vanilla, Gloeosporium, Veratrum, Heterosphaeria, Puccinia, Uromyces, Verbascum, Phoma, Ramularia, Uromyces, Veronica, Ovularia, Peronospora, - Puccinia, Schroeteria, Septoria, Sorosphaera, - Stysanus, Vetch-rust, Viburnum, Actinonema Cercospora, Microsphaera, Plasmopara, Ramularia, Vicia, Ascochyta, Cercospora, Erysiphe, Peronospora, - Phy llosticta, Uromyces, Vinca, Puccinia, Vine (see Vitis) bird’s eye rot, black rot, false mildew, 333, 334, 216, 370 426 204 370 175 263 109 356 334 485 249 355 337 469 502 338 501 134 361 328 478 530 519 333 , 474 515 176 131 502 473 515 175 132 464 336 356 467 454 129 PAGE Vine, mildew, - 176 root-fungus, - 202 vine-epidemics, 81, 84 white rot, 471 Vines, American hybrid, 81 Vingerziekte, - - 524 Viola, Aecidium, - 410 Cercospora, - 515 Gloeosporium, 485 Ovularia, = - 500 Peronospora, - - 134 Phyllosticta, - 464 Puccinia, - 340, 359 Synchytrium,- 112, 113 Urocystis, 16, 21, 31, 317 Violet-rust, - - 340 Viscaria (see Lychnis) Vitis, Ascochyta, - 473 Aureobasidium, 428 Bacteria, - = pt! Botrytis, - 180, 267 Cercospora, - 513 Cladochytrium, - 114 Cladosporium, 510 Colletotrichum, - 488 Coniothyrium, 471 Dematophora, - 202 Gloeosporium, 482, 484 Laestadia, - 216 Leptosphaeria, - 22] Oidium, - - 177, 499 Penicillium, 180 II. GENERAL: INDEX. PAGE Phoma, - 216, 467 Plasmodiophora, 528, 529 Plasmopara, - 81, 128 Pseudocommis, - §29 Sclerotinia, - -, 267 Septosporium, - 519 Sphaceloma, - - 467 Sphaerella, — - - 215 Uncinula, 176 Uredo, - : - 420 W Wallflower (see Cheiranthus) Walnut (see Juglans) Weinstockfaule, - Weymouth Pine (see Pinus Strobus) Wheat (see Triticum) White-rot of timber (due to Polyporeae) Willow (see Salix) Witches’ Broom, due to Aecidium, 18, 24, 72, 88, 202 404, 410 Caeoma, - 2 - 418 Exoasceae, 19, 24,52, 145, 158, etc. Ravenelia, = - 403 Wood-destroying fungi, 5; 34, 36, 62, 72 (see also Polyporeae, Agaricus, Nectria, etc. ) THE END. PAGE Wound-cork, - 42, 76 » -duramen, 3 7KS », -Infection, ey (5° », -parasites,5, 17, 72, To, V42 (see also Polyporeae, Nectria, etc. ) ;, -treatment, = ee Wounds from animals, 78 xX Xanthoxylum, Aecidium, - = A Y Yeast (see Saccharomyces) Yew (see Taxus) Z Zea, Bacteria, - = bys Helminthosporium, 512 Puccinia, 353 Pythium, 116 Tilletia, - 310 Uredo, - - - 420 Ustilago, 279, 281, 282 Zinnia, Sclerotinia, 264 Zizania, Ustilago, 294 Zizyphus, Cephaleuros, 553 Zoocecidia, - = 25) Py “Anis eo : , e ats: et Pine HOR AN SARA IV ORE RIE (Ch OREEROROTR CRIER ORES Buta). Vip } ; eH hay TOS AN At a Ne ety pi de A , \ . putt a a a ee | 6h! ; SB Tubeuf, Karl Freiherr von 731 Diseases of plants induced T793 cryptogamic parasites of, Bot ee ticag “a fs SOLE of bf erie ra pyre ea i ae wESy a ee 3] ay Aol ee nest gue eh £ bas aah Sgt hoes i hal (aad Wtf ney en Me i meet wd ; ie ee nD og A ithn a3 vise ‘ grr. Case) ~ fststs esi? Teese * WO Werte ML AN i OU we oor '