beth Se ti at Ses dno eS Seca ee Taree eee RN Eee eee eet eevee tete e's ete Reo the atte, ated aretha, oe Y easel tet tenes ch cneaes Bate tiat ie te RD i SCA ganas ct reat Ae cS eA Le Aes eRe Ot aod ee A tata dt eas eee ce eS EERSTE Ce nS = 2960: ALBERT R. MANN LIBRARY NEW YorK STATE COLLEGES OF AGRICULTURE AND HoMeE ECONOMICS AT CorRNELL UNIVERSITY Tai 626 345 mann A MONOGRAPH OF THE BRITISH UREDINEAZ AND USTILAGINEE A MONOGRAPH OF THE BRITISH UREDINEZE AND USTILAGINEE WITH AN ACCOUNT OF THEIR BIOLOGY INCLUDING THE METHODS OF OBSERVING THE GERMINATION OF THEIR SPORES AND OF THEIR EXPERIMENTAL CULTURE BY CHARLES ai ownIGHT. F.L.S., M.R.CS. SENIOR SURGEON TO THE WEST NORFOLK AND LYNN HOSPITAL HONORARY MEMBER OF THE NORFOLK AND NORWICH NATURALISTS’ SOCIETY AND OF THE WOOLHOPE NATURALISTS’ FIELD CLUB CORRESPONDING MEMBER OF THE SOCIETA CRITTOGAMOLOGICA ITALIANA socleTe MYCOLOGIQUE DE FRANCE, OF THE SCOTTISH CRYPTOGAMIC SOCIETY OF THE BIRMINGHAM NATURAL HISTORY AND MICROSCOPICAL SOCIETY AND MEMBER OF THE SCIENTIFIC COMMITTEE OF THE ROYAL HORTICULTURAL SOCIETY ILLUSTRATED WITH WOODCU7TS AND EIGHT PLATES LONDON KEGAN PAUL, TRENCH & CO., 1, PATERNOSTER SQUARE 1889 (The rights of translation and of reproduction are reserved.) PREFACE. OF the many interesting problems involved in the elu- cidation of the life-history of the various members of the fungus kingdom, there are none more important to the agriculturist than those connected with the development of the Uredinez and Ustilagineze. Not alone to the culti- vator of plants are these phenomena of interest, but also to the student of botany and microscopy. The work of Tulasne in France, and of De Bary in Germany, during the past thirty years, laid the foundation of our knowledge of the biology of these fungi. More recently the continued researches of De Bary, of Schroter, the late Dr. Winter, Wolff, Kiithn, Magnus, Woronin, and Hartig in Germany, of Oersted, Rostrup, and Nielsen in Denmark, of Fischer Von Waldheim in Russia, and of Farlow and Thaxter in America, have so increased our information as to render necessary a re-classification of our British species. The object of the present work is to supply the British student, not only with improved descriptions of these para- sites, but also with an account of their life-history as far as this is at present known. vi Preface. With this object, the materials from which the following pages are compiled have been collected from the various handbooks, transactions, pamphlets, and periodicals in which .the several authors have published their work. This has been a task of some little difficulty, and has taken several years to accomplish. It has been my en- deavour in all cases to acknowledge the individual work of each author, but has not always been possible for me to do so. The descriptions of the species are in the main those of the late Dr. Winter in the new edition of Rabenhorst’s “Kryptogamen-Flora,” although I have not scrupled to amend them in various ways, partly from my own observa- tions, and partly from the writings of others. The arrange- ment of the species is that first proposed by Dr. J. Schréter and employed by Winter in the above-named work, to which I must refer the student for the descriptions of those European species which are not known to be British, as well as for the full details of the synonymy of those described in the present work. As the present generation of British students owe their knowledge almost exclusively to the writings of Dr. M. C. Cooke, the synonymy of his two works, “The Handbook of British Fungi,” and “ Micro- scopic Fungi,” is given in full. The synonymy of the older British botanists has, however, not been omitted. The biology of the Uredinez is, in the main, the work of the late Professor De Bary, and of the Ustilaginez that of Von Waldheim, Brefeld, and Schréter. During the past seven years I have devoted much time to these two groups of fungi, and have made between nine hundred and a thousand experimental cultures, with the object of working out the life-history of those species of which it was unknown, Preface. vii as well as of testing the accuracy of the statements of other botanists. The results of my cultures have been from time to time published in various botanical periodicals of this and other countries, and in the Transactions of the Linnean and Royal Societies. It was at one time hoped, when the zcidiospores and uredospores were shown not to be distinct species, that con- tinued biological investigation would materially lessen the number of species of the Uredinee. I am convinced, however, that modern botanists have erred in grouping together forms on account of the similarity of the teleuto- spores; for instance, the Pucciniz on the Composite, Labiate, Umbellifere, etc. The proper limitation of species of these parasites cannot be effected on purely morphological grounds ; it can only be accomplished when the morphological characters are supplemented by a know- ledge of the life-history of each individual species. I am greatly indebted to my friend, Professor J. W. H. Trail, of Aberdeen, for his kind assistance in. the revision of the proof-sheets. I have also to thank numerous friends and corre- spondents, both in this country and abroad, for information and material for experiment, amongst whom I must mention Professor Farlow, of Harvard University; Mr. W. B. Grove; Rev. Dr. Keith; Dr. P. Magnus, of Berlin ; Mr. W. Phillips, F.L.S.; Mr. H. T. Soppitt; Professor P. Sorauer, of Proskau; Rev. Dr. Stevenson; Rev. J. E. Vize ; the late Dr, Winter, and many others. CONTENTS. CHAPTER I II. III. IV. Vv. VI. VII. VIII. IX, xX. XI. XII. XIII. BIOLOGY OF THE UREDINE#-—INTRODUCTORY REMARKS MYCELIUM OF THE UREDINEE aii ls SPERMOGONIA AND THE SO-CALLED SPERMATIA ALCIDIOSPORES a wd ssi UREDOSPORES ... ve oot TELEUTOSPORES at et tals st HETER@CISM MYCELIUM OF THE USTILAGINEH ... FORMATION OF THE TELEUTOSPORES OF THE USTILAGINES GERMINATION OF THE TELEUTOSPORES OF THE USTILAGINEZ INFECTION OF THE HosT-PLANTS BY THE USTILAGINEE SPorE CULTURE be oe sits aie THE ARTIFICIAL INFECTION OF PLANTS ee DESCRIPTIONS OF THE BRITISH UREDINEE APPENDIX—IMPERFECT FoRMS, OF WHICH THE FULL Lirr-History IS UNKNOWN ane see vee ane ove ne DESCRIPTIONS OF THE BRITISH USTILAGINEE ee ols SUPPLEMENT—ALLIED AND ASSOCIATED SPECIES THE BARBERRY LAW OF MASSACHUSETTS GLOSSARY sie “Gi ae as wat AUTHORS QUOTED ai i DESCRIPTION OF PLATES INDEX OF HosT-PLANTS... eas ote “a sit BIOLOGICAL INDEX sles ae sie tii a ei INDEX OF SPECIES an ba sit sit aig PAGE 2I 28 36 46 58 61 72 99 105 114 119 254 272 297 302 305 309 317 325 331 339 A MONOGRAPH OF THE BRITISH UREDINEA! AND USTILAGINEA?. CHAPTER I. BIOLQGY OF THE UREDINEAZi—INTRODUCTORY REMARKS. THE Uredinez constitute a large and important group of parasitic fungi, affecting fone plants and ferns, representatives of which are familiar to every student of nature, practical agriculturist, and working microscopist. In whatever part of the world phanerogams appear, these parasitic cryptogams accompany them. Our present object is to obtain an insight into. the life- history and the structure of those species which are known to occur in Great Britain, Much has been elucidated con- cerning these points during the past twenty years, but still much remains to be discovered. They all grow as parasites upon some living plant, independently of which they cannot exist, and consist of two essential elements—spores and mycelium. The spores every one sees; the mycelium, on account of its inconspicuous nature, and because it cannot be seen without some little effort, no one takes the trouble to look B a British Uvredinee and Ustilaginee. for. The vegetative mycelium of most of the Uredinez is very similar, although the spores are very diverse. The mycelium, being present in the appropriate host-plant, may give origin to several different kinds of spores, according to the nature of the Uredine under examination, each kind of which was regarded by the older botanists as being a distinct genus (cidium, Uredo). While it is true that the relationship between the various spore-forms was suspected as early as the beginning of the present century, yet its actual demonstration has been accomplished only within the last thirty years. From the investigations of Tulasne and De Bary, we now know that each spore-form has a life-history of its own, that they are often products of the same mycelium grown among different environments, and that they all arise from some antecedent spore-form. The ultimate condition in which all the Uredineze are encountered is the teleutospore, which, after a longer or shorter period of quiescence, manifests its vitality by ger- minating and producing a body—the promycelial spore. The promycelial spore may then be regarded as the beginning of the series of spore-forms of which the teleu- tospore is the end. The actual number of spore-forms intervening between the promycelial spore and the teleutospore is subject to considerable variation in different species, 1. The promycelial spore, after emitting a germ-tube, which enters the tissues of its appropriate host-plant, may give rise to a mycelium, which produces teleutospores exactly similar to the teleutospore from which it originated (Leptopuccinia, Micropuccinia). 2. The promycelial spore in a second group of the Uredinez, in like manner, gives rise to a mycelium which produces uredospores, and subsequently teleutospores (Hemipuccinia). « Biology of the Uredinea. 3 3. The promycelial spore may give rise to a mycelium which produces, first, spermogonia, then zecidiospores, then uredospores, and finally teleutospores (Auteupuccinia). Certain minor variations in the life-cycle occur; for example, the uredospores may be suppressed, so that we have only zcidiospores and teleutospores (Pucciniopsis), or the spermogonia and zcidiospores may occur on one plant, and the uredospores and teleutospores upon another of a totally distinct kind (Heteropuccinia). Each of the above spore-forms has its own peculiarity of structure, as well as origin, so that it will be better, for clearness, to describe them separately. The spore relations may be thus tabulated :— PROMYCELIAL SPORE. Teleutospore Uredospore Spermogonium Spermogonium Teleutospore | Uredospore 4Ecidiospore — Teleutospore Uredospore - Teleutospore - 4 British Uredinee and Ustilaginee. CHAPTER. II. MYCELIUM OF THE UREDINEA’. THIS is, of course, common to all the different spore- forms, inasmuch as it is that part of the fungus which develops them. It consists of a number of hyaline tubes that extend themselves principally between the cells of the host-plant. In some instances these tubes send short branches into the cells (haustoria), but the haustoria are not so common nor so well developed as in some other parasitic fungi (Peronosporee, Ustilaginee). In the tropical genus Hemileia* the haustoria are unbranched, thin-stemmed vesicles, very like those of Cystopus. Bagn ist has figured the haustoria of Puccinia malvacearum, but his figure is of a doubtful character ; and Barclay} has depicted an arborescent haustorium on the mycelium of Acidium urtice, var. himalayense. The mycelial tubes (Plate I. Figs. 1 and 2) consist of hyaline, membranous walls, containing usually a colourless watery fluid. They are rendered more distinct by the * Marshall Ward, ‘‘On Hemileia vastatrix,” Zinn. Soc. Four. Bot., vol. xix., and Quart. Four. Micr. Science, new series, vol. xxi. + Bagnis, ‘‘ Obs. Vita et Morphol. Funghi Uredinei,” t. i. fig. 11. t Barclay, ‘‘ Scientific Memoirs by Medical Officers of the Indian Army” (1887), t. iv. fig. 4. Mycelium of the Uredinee. 5 action of caustic potash (KHO). The tubes themselves are rather irregular in their outline, and branch at frequent intervals, These branches unite with other mycelial tubes, so as to form an anastomosing irregular network, which pervades more or less widely the tissues of the affected plant. At rare intervals transverse septa are seen. The function of this mycelial network is to utilize the elabo- rated material which the host has prepared for its own use, and to turn it into a suitable pabulum for the sustenance of the fungus. The extent to which the mycelium permeates the tissues of the host-plant varies; in the majority of cases it is localized and confined toa limited area. The germ- tube from a single spore having once entered the tissues of a leaf, the tendency of the mycelium thereby pro- duced is to spread equally in all directions in a centrifugal manner. Many causes, however, come into operation which tend to prevent the ultimate spore products being equidistant from the centre. This may be due, in part, to the mycelial hyphe growing more luxuriantly in one direction than in another; but to a great extent it arises from a want of uniformity in the tissues of the host-plant. Still, however, we are often able to observe that the fructification of those Uredines which have a limited mycelial growth, is arranged either in a circle or in more or less circular manner (Puceinia lychnidearum, Uroutyces scillarum, Caoma orchidis, Acidium zonale). In those cases in which the mycelium is developed in a leaf with strongly marked venation, this tends to exert a directive influence upon its extension; for example, the primary uredospores of Triphragmium ulmarie and Uromyces alchemille. The same directive influence of the tissues of the host-plant is seen in the linear arrangement of the sori of Puccinia magnusiana, graminis, rubigo-vera, etc., 6 British Uredinee and Ustilaginee. especially when they occur upon the stems or cauline sheaths of their graminaceous host-plants. The presence of the mycelial hyphe generally acts as a local stimulant to the tissues in which they are present, as is evinced by the increased thickness which is often associated with a concave arching or vaulting of the affected places (Zcidium grossularia, berberidis, Restelia cancellata). When the stems are affected, this is usually shown very markedly by the development of swellings and distortions; as a rule these are more or less fusiform, and often induce considerable bending of the attacked stem (P. difformis, Kze.; Ur. trifolit), Even upon these cauline tumefactions may often be traced the concentric arrange- -ment of the sori. The stimulation of the affected part may be carried to such an extent as to kill the invaded tissues; thus we often find the older leaves of Malva sylvestris and Althea rosea with numerous circular holes, punched as it were out of them. Each of these has been the seat of the very localized mycelium of Puccinia malva- cearum, which has by its presence killed a circumscribed portion of the entire thickness of the leaf tissue, so that, as the leaf itself expands, the dead area above described becomes separated round its circumference and falls out, leaving a circular hole. The above-described perforated foliage is most observable after a period of drought; in rainy weather the reproduction of the parasite is so rapid that the entire leaf tissue is quickly invaded by the fungus and totally destroyed. The same dropping out of mycelial areas occurs upon the stem; here, however, the hyphz only penetrate the external parts, so that when the affected spots drop out an elongated or fusiform wound is left, at the bottom of which the central woody part of the stem is exposed. Schroter* has pointed out that a somewhat * Schroter, ‘‘Cohn’s Beitrage,” vol. ii. p. 88, Mycelium of the Uredinee. 9 similar circumscribed dead area surrounds those sori of P. annularis which are formed late in the year, and that it is by means of teleutospores contained in these sori that the fungus is reproduced in the following spring. The localized mycelium of the zcidiospores especially, has a powerful influence upon the chlorophyll, causing it to lose its green colour, and, as De Bary * has pointed out with &. cancellata, making it disappear entirely. The affected tissue is found to contain an immense number of minute starch granules, To such an extent does the development of starch take place, that in the Himalayas the natives eat the hypertrophied stems of Urtica parvi- folia, which are affected with 2c. urtice, on account of the abundant nutritive starchy material they contain. The hypertrophies are eaten just before the acidia open, and are said to resemble cucumber in flavour.t The affected places generally assume some tint of yellow or reddish yellow, more or less bright. In the Polygonee the spots are bright red or purplish (4c. rumicis,U redo bifrons). In other cases they are yellow, surrounded by a purplish or reddish border (4. onale, behenis), Ina few instances all colour is more or less discharged, and the spots appear whitish (4c. albescens, leucospermum). The mycelium of all species is by no means thus localized. With some Uredines, on the contrary, it per- vades the whole plant with the exception of its roots— stem, leaves, petioles, peduncles, and the upper part of the root-stock. De Bary{ has shown that when the mycelium of a Uredine can be traced into the perennial parts of the host-plant, it is itself perennial; thus with * De Bary, ‘‘ Brandpilze,” p. 73. + Surgeon-Major A. Barclay, ‘‘ On Akcidium Usgice,” ‘Scientific Memoirs by Medical Officers of the Indian Army ” (1887), p. 2. t De Bary, ‘‘ Neue Untersuchungen iiber Uredineen ” (1865), pp. 20, 21. 8 British Uredinee and Ustilaginea. Endophyllum euphorbie it is found more particularly in the pith and in the inner parenchyma of the bark of the affected plants. In plants of Axemone nemorosa, it may be found in the leaves, stems, and vascular bundles of the upper part of the rhizome, as is likewise the case with the host-plants bearing Uvedo suaveolens and Ac. tragopogonis. ( 9) CHAPTER III. SPERMOGONIA AND THE SO-CALLED SPERMATIA. THESE bodies are produced by the mycelium, which arises directly from the entrance of the germ-tube of a promy- celial spore, or from a perennial mycelium permeating the tissues of the host-plant. The hyphz which are destined to form a spermogonium becéme interwoven into an inextricable network in the subepidermal tissues of the host-plant. They pass for the most part between the cells, and are found more frequently septate where a spore-bed is about to be formed than when they are encountered elsewhere in the tissues of the host- plant. Their contents are watery and transparent. From this tangled mesh, immediately beneath the epidermis, are given off a great number of branches of much smaller diameter, as a rule about 2 or 3u. The general direction of these finer branches is towards the epidermis. They incline towards a central point, however, and so come together, forming a pyriform or subglobose body, the upper part of which is covered only by the cuticle of the host- plant. This body soon assumes a pyriform or flask-shaped contour (Plate I. Fig.3). The neck of the flask bursts through the epidermis as a minute conical point, which can be seen to consist of a vast number of straight hyphe, parallel to one another in the main, but all sloping upwards, converging 10 British Uredinee and Ustilaginee. to the apex of the cone. The outer walls of this body are embedded between the cells of the host-plant, and are like- wise composed of similar fine hyphae, placed side by side. A section of the spermogonium at this time shows its interior to consist of similar converging parallel hyphz. Very soon those hyphz which constitute the emerging apex separate from one another, so as to appear as a brush of stiff hair like bodies (paraphyses) ; in the centre of this brush is a minute canal, which passes downward to the interior of the body of the spermogonium (Plate I. Fig. 4). The apices of those converging hyphze which occupy the lower part of the spermogonial interior, are now seen to be surmounted by very minute irregularly oval or rounded bodies (Plate I. Fig. 5)—the so-called spermatia. These spermatia vary in size, not only in the different species, but also in the same spermogonium. In those I have examined, they were from 5 to 8u long, and 3 or 4 or even 6u wide. Tulasne * gives the measurement of the spermatia of Triphragmium ulmarie and Puccinia fusca as from § to 6u long, while those of most the A®cidia and of Ceoma pingue and C. ridesii are rarely more than 4u long. The spermatia are produced in linear series from the apices of the hyphz (sterigmata), which fill the interior of the sper- mogonium. These are held together by a viscid, gelatinous substance, which at first fills the bottom of the canal; but as more spermatia are produced, gradually the whole canal becomes full, and eventually the mass oozes out at its upper end in the form of a globule. The cause of the expulsion of the spermogonial mass is, as De Bary ¢ has shown, the imbibition of moisture, which causes the investing gela- tinous material to swell. As the spermogonium advances in maturity, its flask-like neck opens out, so that, instead * Tulasne, ‘‘2e Mémoire,” p, 118, + De Bary, ‘‘ Brandpilze,” p. 60, Spermogonia and the so-called Spermatia. 11 of resembling a flask with a bristly mouth, it comes to be a cup-shaped depression on the surface of the leaf, sur- rounded by a hedge of stiff bristles (Plate I. Fig. 4). The function of these bristly paraphyses appears to be that of preventing the exuded mass of spermatia and jelly from being bodily washed off the surface of the leaf by rain. This is the more necessary, because it is during wet weather, as we have seen, that the spermatia, from the imbibition of moisture by their investing jelly, are brought to the surface of the leaf at all. The paraphyses occur with all spermo- gonia except those of the Phragmidia. Whatever their functions may be, whether as a spore- form or as spermatia properly so called, it is obvious that, by being held together by a viscid substance, their chance of dissemination by currents of air, etc, is but small. To a certain extent their diffusion over a limited area might take place in very wet weather. Rdthay* has, however, shown that the spermatial mass contains a certain amount of some saccharine material mixed with it—a substance which has the power of reducing Fehling’s solution; and further, that, as a matter of fact, insects do visit spermo- gonia for the sake of this saccharine matter, and are thereby unwittingly the agents for the distribution of the spermatia. Of this latter point, he obtained actual demonstration in the following manner. He had standing upon his window- sill some plants of Euphorbia amygdaloides, upon which a large number of spermogonia of Exdophyllum euphorbie were in the act of exuding their contents. One day, as he approached the window in question, he noticed some flies, which were busy upon these leaves, fly away from the leaves and alight upon the window pane. Closer scrutiny showed that the flies had left their wet footmarks upon the * Rathay, ‘‘ Untersuchungen iiber die Spermogonien der Rostpilze.” Wien: 1882, 12 British Uredinee and Ustilaginee. glass. These footmarks were found, upon microscopic examination, to contain the spermatia of the Endophyllum. Further investigation of the subject led him to the con- clusion that this sugary matter in the spermogonial contents acts as a bait to attract insects, and he has observed some 135 species, of which 31 were Coleoptera, 32 Hymenoptera, 64 Diptera, and 8 Hemiptera, thus visiting various spermogonia. The examination of the. spermo- gonia of Uromyces pist, Puccinia suaveolens, fusca, trago- pogonis, pimpinelle, Endophyllum euphorbia, Gymnosporan- gium sabing, and G. juniperinum, gave rise to copious deposits of suboxide of copper, when the washings of the affected leaves in distilled water were treated with Fehling’s solution ; less was obtained from Uvomyces dactylidis, Puccinia graminis, covonata, rubigo-vera, sylvatica, viola, Gymnospo- vangium clavariaforme, and Atcidium magelhenicum, and least of all with Puccinia poarum and Aicidium clematidis. Some of the spermogonial contents actually taste sweet when the tongue is applied to them, as those of G. sabine and junipertnum. ‘The spermogonial contents of G. sabine were further found to contain dextrose and levulose, the latter predominating. Insects are not only attracted to the spermogonia by their saccharine contents, but also by the powerful odours which many of them possess. Persoon* long ago noticed the penetrating odour of the spermogonia of Puccinia suaveolens, which precede its uredospores; hence he called the Uredo, suaveolens. In a paper upon “Mimicry in Fungi,” ¢ I pointed out the probability of this. odour being for the purpose of attracting insect visitors, mimicking as it does the perfume of the flowers of Anothera biennis, Sowerby had already observed the fact that flies * Persoon, ‘‘ Synopsis Fungorum,” p, 221. t Plowright, ‘‘ Mimicry in Fungi,” ‘ Grevillea,” vol, x. pp. 1-14, Spermogonia and the so-called Spermatia. 13 visited this plant. Tulasne* remarked the odour of the spermogonia of &cidium pini and of Uredo serratula, which he compared to that of the pollen of willows. Léveillé-+ noticed the odour of &cidium tragopogonis, which he said resembled the perfume of some flower. Rathay indicates that the possession of the pleasant odours is a special characteristic of those Uredines which have a perennial mycelium. The plants thus invaded are further rendered more conspicuous to insects by the manner in which the affected leaves or shoots grow. As he says, these have a strong negative geotropism. Not only are they more erect, standing above the other leaves or shoots, but they are further characterized by an alteration in their foliage ; while they are, as a rule, not only paler in colour, but also modified in form, being either more attenuated or the reverse. The Uredines with a short-lived mycelium, on the contrary, have their spermogonia produced upon the upper surface of the leaves on brilliantly coloured spots, which contrast more or less strikingly with the green colour of the healthy foliage. These spots are generally bright yellow or orange, often with a tinge of red. Sometimes they are white (4c. fabe) or purple-red (4c. rumicis). The negative geotropism is not, however, confined to the species with the perennial mycelium ; I have noticed it very markedly with some leaves of Senecio jacobea, upon which I had produced the zcidium of Puccinia scheleriana ; as soon as the ecidial spots became sufficiently developed to produce spermogonia, the leaves which bore them became almost erect, while the unaffected leaves remained horizontal. In either case the darker paraphyses surmounting each conceptacle finally attract the insect to the sugary globule. * Tulasne, ‘2° Mémoire,” p. 118. t Léveillé, in D’Orbigny’s ‘“‘ Dict. Univ. d’Hist. Nat.,” tome xii. p. 175, sub. Uredines, 14 British Uredinee and Ustilaginee. Although the paraphyses themselves, when seen singly as transparent objects, appear hyaline or yellowish, yet ex masse they are almost black in many species, and in all distinctly darker than the surrounding tissues. Their function, however, is not confined to the attraction of the insect visitors ; they also (as has been already stated) main- tain the spermatial globule in its place upon the leaf, and prevent it being bodily washed away. With regard to the presence of some saccharine matter in the spermo- gonial masses, I may add that I have found, in confirmation of Rathay’s statements, that the spermogonial contents of Puccinia obscura both reduced Fehling’s solution and also gave the reaction with the indigo-carmine test. The so-called spermatia were long regarded as not possessing the faculty of germination. Cornu,* however, found that when they were placed in water, in which a little white cane sugar had been dissolved, and exposed to the free action of the air, they were capable of germination. During the past five years I have repeated M. Cornu’s experiments, at first employing white sugar as the nutrient material. Under these conditions the individual spermatia at first assume a more regular outline than one commonly observes them to have as they are naturally exuded from the spermogonial receptacles (Plate I. Figs. 6, 7). They at one extremity give out a minute prolongation. This pro- longation does not develop into a germ-tube, but gradually increases in size, until it acquires the size and form of its parent spermatium ; at the end of twenty-four hours many of these twin bodies, joined end to end, will be observed in the culture (Plate I. Figs. 8, 9). Resembling Saccharomyces spores as these bodies do, I at first thought some stray yeast-spores had accidentally gained admission * Cornu, Belletin de la Société Bot, de France (1876), tome xxiii. pp. 120, 121. Compt. rendus, January 21, 1875. Spermogonia and the socalled Spermatia. 15 into my culture; in fact, De Vauréal* has suggested that yeast-spores are simply the spermatia of Uredines. By frequent repetition the result was always the same—the so-called spermatia budded in the same manner as Saccha- romyces spores. Mr. A. Lister has observed the same budding with the spermatia of Acidium albescens, of which process he sent me an excellent figure made by him in 1873 (Plate I. Fig. 10). In 1883, I made a number of cultures in which honey instead of sugar was employed ; with this substance the germination is much more active and prolific, the budding spores often remaining in yeast colonies, attached by their ends in chains of half a dozen or more. I have germinated the spermatia of the ecidio- spores on Bellis perennis, Ranunculus bulbosus, R. ficaria, Anemone coronaria, Lapsana communis, and some others. The older germinating spores present two nuclei of notice- able size. Not only are the spores budded in linear series, but also from these chains young spores are developed laterally, from the point of union where two older ones meet (Plate I. Figs. 12-16). This budding goes on con- tinuously, but not with the same rapidity as with Saccha- romyces, nor, as far as my observations go, to the complete exhaustion of the saccharine material. Probably these changes in the fungus are accompanied by the production of alcohol, but I was unable to detect it in any of my cultures. When the Uredine yeast-spore falls to the bottom of the fluid, it is excluded from the air and does not further change; whereas if a small number of Saccharomyces spores be placed in a saccharine fluid, they rapidly multiply themselves until all the sugar disappears, while the Uredine spermatial cultures, at the end of several days, retain their sweet taste. Attempts to produce alcohol from brewers’ * De Vauréal, “ Schutzenberger on Fermentation ” (1876), p. 61. Inter- national Scientific Series, vol. xx. 16 British Uredinee and Ustilaginee. wort ina Pasteur’s flask, with Uredine yeast-spores, were with me unsuccessful. The spermatia do not germinate in pure water, but only in the presence of sugar. There is, however, often enough saccharine matter in the investing jelly of the exuded globule for a few of them to bud, as Mr. Lister’s figure shows, and as I have several times observed. Rebentisch * noticed the small black points which the spermogonia present in the spots upon pear leaves, on which Restelia cancellata are produced. Ungerft figures these structures, and describes them as a distinct species under the name Zcidiolum exanthematicum ; he regarded them as a peculiar exanthem of the affected plant. Meyenf{ thought they played the part of the male element in the _ reproduction of the Uredinez, with which they were asso- ciated. Tulasne§ considered they were spore-forms developed from the same mycelium as their accompanying spore-forms, but, like De Bary, was unable to observe their germination. The spermatia of the Uredinee occur with all the spore-forms. They almost invariably precede and accom- pany the zcidiospores, being produced from the same mycelium, but generally occupying the upper surface of the leaf, while the acidial cups occur on the lower. The first formed spermogonia are produced in the centre. of a spot; the next more externally, and so on centri- fugally. In many cases, however, they occur on the same surface and between the ecidia. When the acidiospores are cauline, the accompanying spermogonia are often * Rebentisch, ‘‘ Prodrom. florze Neomarch.” me Unger, ‘Die Exantheme der Pflanzen,” t. iii, figs. 18, 19. Wien; t Meyen, ‘‘ Pflanzenpathologie” (1841), pp. 143-147. § Tulasne, Comptes rendus, tome xxxii, p. 472; and tome xxxvi. Pp. 1093. Spermogonia and the so-called Spermatia. 17 arranged circumferentially. Morphologically, they present a great similarity to the spermogonia of the lichen-fungi, especially to those of Collema. Since our knowledge of the life-history of the lichen- fungi has been increased, from the facts added to it by Schwendener, De Bary, Stahl, and others, many botanists have come to look upon the Uredine spermatia as fertiliz- ing bodies, and to consider the AEcidiomycetes as being nearly allied to the Ascomycetes. There are, however, certain facts which cannot be overlooked. In the first place, the faculty which the Uredine spermatia have of multiplying themselves by budding in saccharine solutions in exactly the same manner as the spores of Saccharomyces, and also as Brefeld has shown the conidia of the Ustila- gineze do in his ndhrlésung, points rather to their being conidia than spermatia. Then, again, it is not asserted that all spore-forms of the Uredinez are sexual—this is claimed as probable for the zecidiospores alone ; true it is that the zecidia are almost always accompanied by spermogonia, but this is not in- variably the case. De Bary* cultivated a single plant of Sempervivum affected with Exdophyllum sempervivi which bore no spermogonia, but the zcidial cups were perfectly developed, and their spores germinated in the normal manner. In the autumn of 1883, I transplanted into my garden a wild plant of Zvragopogon pratensis affected with the Aécidium. During the spring and summer of 1884, this plant continued to produce a suc- cession of zcidia, the spores from which were used for infecting seedlings of Tragopogon. This they successfully did, causing the development of the teleutospores, with their scanty accompaniment of uredospores. I was never able to find any spermogonia upon this Tragopogon, * De Bary, ‘Morphol. und Physiol.,” 1st edit. p. 169. Cc 18 British Uvedinee and Ustilaginee. although I repeatedly examined it for them. These two cases, the Sempervivum and the Tragopogon, in both of which perfectly normal, and in the latter instance function- ally active, zcidiospores were produced without the presence of spermogonia, show, at any rate, that each individual zcidial cup was not the result of a spermatial fecundation. It may have been that in both instances the plant bore spermogonia before they came under observation, and that they both bore within them an already fecundated mycelium ; but this is only a supposition, and, even if it be admitted as possible, yet it does away entirely with the analogy of the sexually produced spore-beds of the lichen-fungi and Polystigma, in which each spore-bed is the result of a separate spermatial impregnation of the trichogyne. Nor is the case of the Uredinez with short- lived mycelia more tenable. If with them each ecidial cup is a sexual product, then it cannot arise in the first instance without a spermatial fecundation. But Schréter * has shown that when he produced the zxcidium of P. porrt (Sow.) by infecting the Adium schenoprasum with the teleutospores, the resulting zcidia were always unaccom- panied by spermogonia, as was also the case when they occurred on onion (A. cepa); and, further, he found while the vernal specimens of the ecidiospores of Uromyces ervi on Ervum hirsutum and of Puccinia galii on G. aparine were accompanied by spermogonia, yet those produced later in the year were invariably without this accompani- ment. If the spermatia were necessary to the development of the zcidia in spring, how could they be dispensed with by the same formation in autumn? The fact, however, stands that spermogonia almost always accompany zcidio- spores ; but not only is this the case with the true Acidia, but also with the analogous Caomata—analogous in as * Schroter, ‘*Cohn’s Beitrage,”’ vol. ii, p. 83. Spermogonia and the so-called Spermatia. 19 far that they originate from promycelial infection of their respective host-plants, and that their spores are produced in linear basipetal series. In fact, the Ceomata only differ from the Aécidia in the want of a peridium. Yet we do not find spermogonia exclusively associated with these two spore-forms; on the contrary, they occur not very rarely with the uredospores—a spore-form which does not present any morphologically biological parallel with the two above named, inasmuch as its spores are produced singly on separate and distinct spore-forming hyphe ; nor has any sexuality ever yet been claimed for uredospores. As instances in which spermogonia accompany uredospores may be quoted Zrifhragmium ulmarig and the Brachy- puccinie, Puccina hieracit (Schum), P. suaveolens (Pers.), P. bullata (Pers.), P. oreoselini (Strauss), and Uromyces terebinthi (D.C.). Nor is this all, for spermogonia also occur with the teleutospores, as in P. selphiz (Schw.),* P. Salcarie (Pers.), P. liliacearum (Duby), and P. fusca (Relh.).t It has been suggested that the association of the spermogonia with the teleutospores in the last-named species might be only an exceptional case in which the corresponding female spore occurred upon some other plant—in other words, that it is an hetercecious species; but while admitting the full weight of this argument in the abstract, yet of the Puccini in question, if P. fusca be a Pucciniopsis, and have for its zcidiospores Aicidium leucospermum, an zcidium so markedly accompanied by spermogonia that their presence was recognized by the older botanists, we must consider it to be an exceptionally well-provided species. The escaped spermatia are very commonly found scat- tered amongst the ezcidiospores, and often adhering to their outsides. Mr. W.G. Smith f has represented this condition, * Trelase, ‘‘ Parasitic Fungi of Wisconsin.” 1884. + Tulasne, ‘2° Mémoire,” p. 116, note. t W. G. Smith, ‘‘ Diseases of Field and rae Crops,” p. 166, fig. 86. Z San Saran Seat a8 ayer ame Yip’ Bue by Paul” 20 British Uvedinee and Ustilagineae. and suggested that this may in itself be the fecundative act, but the same objection obtains to this latter suggestion as to the former. As far as I can judge, the balance of evidence is against the supposition that the spermatia are sexual organs. The other supposition, that they are conidia, is more plausible ; but one would have thought, had such been the case, it would have been capable of demonstration. I have at- tempted various experimental cultures with these bodies, but uniformly without result. These consisted of the application of the spermogonia in active germination in honey and water— 1. To the foliage of the plant upon which the spermo- gonia occur. Thus the spermatia of Acidium ranunculi repentis were applied to the healthy foliage of Ranunculus repens (Exp. 123) ;* of Restelia cornutato Sorbus aucuparia (Exp. 124); of cidium bellidis to Bellis perennis (Exp. 244, 247). 2. To the corresponding host-plant bearing the uredo- spores and teleutospores, The spermatia of Zcidium ber- beridis were applied to wheat (Exp. 392). 3. Remembering the fact that the spermatia are carried by insects, and that they germinate so freely in honey, they were applied to the stigmata of certain flowers, I was further induced to try these experiments from the frequency with which the fruit of so many plants in this country are attacked by the zcidiospores of some of the Uredinex ; for instance, the Mahonia berries, the barberry fruit, the goose- berry, and the fruit of the hawthorn, The spermatia of Ac. bellidis were applied to the stigmata of Bellis perennis (Exp. 267), and those of 4c. ficarie to Ranunculus jicaria (Exp. 274), but no result was obtained ; the infected plants produced in due course perfectly normal ripe seeds, * The numbers of these experiments refer to my private note-book of experimental cultures, CHAPTER IV. AECIDIOSPORES, THE mycelium originated in the tissues of the host-plant by the entrance of the germ-tube from a promycelial spore, after it has produced a certain number of spermogonia, proceeds in the course of its development to give rise to the zcidiospores, This spore-form is of considerable interest, not only on account of the manner in which it arises, but also by reason of its attractive appearance in the mature state. The first appearance of the zcidiospores consists in the formation of the receptacle (pseudoperidium) in which they are contained. The mycelium destined to give rise to an zecidium becomes more frequently branched and interwoven. At certain points, not immediately beneath the epidermis, but pretty deeply placed in the parenchyma of the plant, the mycelial hyphz compact themselves into spherical bodies, which at first are not larger than the parenchymatous cells of the host-plant, The bodies (the primordia of De Bary) gradually increase in size by the addition of fresh hyphe from the mycelium, It can soon be observed that the interior and bulk of these bodies consist of a cellular structure, and that they bear the greatest possible resemblance to perithecia (Plate IT. Fig. 1). They are globose, but rather flattened upon their upper surface—that is, the surface nearest to the epidermis. 22 British Uvedinee and Ustilaginee. A section through one of them shows it to consist of an external envelope of cells, which encloses the body on all sides, above, below, and laterally. The base of the body is a circular disc (hymenium), which bears upon its upper surface a number of closely packed, erect hyphze (basidia), each of which supports a linear series of spores. The hymenium increases in width as the ezcidium develops, but whether from new basidia arising between those at first formed, or circumferentially, is not known. Sur- rounding the circumference of the hymenium is a circle of sterile cells (the peridial cells; Plate II. Fig. 2). Spore- formation in the ecidium proceeds from above downwards, in the following manner. The cells destined to become spores are enclosed in a hyaline tube (the mother-cell) ; they are at first colourless, but‘soon within each, from above downwards, a number of granules appear, which, becoming invested with a delicate membrane, rapidly augment in size, from an increase of the enclosed protoplasm, until they touch the walls of the mother-cell. ‘he granular contents are by this time orange (cidium), or brown (Restelia), or remain white in a few species, such as Ac. rumicis, vince, etc.* The growth of the cell continues until it is indistinguish- able from the mother-cell. By the mutual pressure of the neighbouring cells the young spores become polygonal. As this process begins above and continues downwards, or, as it is now termed, in a basipetal manner, it follows that the ripe spores are uppermost. As soon as they have attained their full maturity, they separate from one another and are blown away. In some species (Restelia, xcidio- * Mr, G. Massee considers the whole A®cidium to be a sexual product resulting from the conjugation of two dilated mycelial hyphee in the tissues of the host-plant (Anmals of Botany, June, 1888, vol. ii. Pp. 47-51, plate iv. figs. 1-7). : NY : a 6 sees SE aaa eo Cos Ye ) Fig. 3.—Chrysomyxa rhododendri. Section of spore-bed showing the compound teleu- from appet Conserv of ieamsperioe cosiavinsde daa ot tie nner sce uean arin lee (be Bay e, The epidermal cells of the host-plant (A. Azrsutum) ; », mycelial hyphae. at that end of the cell which is nearest the septum (Plate III. Figs, 22, 23; and Plate IV. Figs: 11, 13, and 14). Germination.—The period at which germination takes place in teleutospores is subject to variation with the different species of Uredines. In the greater number of cases this process ensues only after the teleutospore has remained for some considerable time in a state of qui- escence or rest. Generally this resting period extends from the summer or autumn of one year until the ensuing spring, corresponding, in fact, with the conditions of the host-plant, which of course, in the case of deciduous plants, is without suitable foliage during the winter months into which the fungus can gain an entrance. On the other hand, there are several species in which teleutospores germinate as soon as they are mature, without undergoing any resting period (Leptopuccinia, Lepturomyces); while there are others which have only a very short resting period (P. rubigo- Leleutospores. 43 vera, poarum). In either case the germinative process is the same; This consists in the protrusion of a germ-tube from the teleutospore through the germ-pore above men- tioned, which tube is really an extension of the endospore through the germ-pore. This tube consists of a hyaline, homogeneous membrane, into which the protoplasm con- tained within the spore passes. This tube (the promy- celium) does not, like the germ-tubes of the uredospores and zcidiospores, elongate indefinitely, but, after attaining a certain length, ceases to grow onwards and terminates in a blind, blunt extremity. Into the promycelial tube the protoplasm passes from the spore, and accumulates towards the peripheral extremity, so that the end nearest the spore becomes empty. The distal end now exhibits one or more septa, which are formed from above downwards. At the same time, from each compartment thus divided off there arises a single, short, pointed branch ; these at their points dilate. The dilated end becomes the receptacle for the protoplasm originally contained in each compartment, and rapidly assumes an oval or subreniform outline (the pro- mycelial spore), which in the course of a few hours becomes abstricted and falls off (Plate IV. Figs. 2-10). The pro- mycelial spores, when placed in water or upon any damp surface, forthwith germinate (Plate III. Fig. 24). This process consists in the outgrowth, from some point of their surface, of a short but acutely pointed tube. Placed upon the cuticle of a living leaf of the proper host-plant, this germ-tube turns its point downwards, and, piercing the epidermal cell-walls, enters the tissue of the leaf (Plate IV. Fig. 1). Having thus gained an entry into its host, the protoplasm contained in the promycelial spore passes down- wards into the germ-tube, leaving the spore empty. The empty spore-cell falls off, and the minute opening through which the germ-tube has entered the epidermal cell ceases 44 British Uvedinee and OUstilaginee. to be visible; this is probably due to the elasticity of the epidermal cell-wall, The germ-tube itself continues its onward growth; soon branching, it insinuates itself between the cells of the host, where it gives rise to a mycelial development, similar to that which arises from the zcidio- spore or uredospore germ-tubes. With the Leptopuccinia and Lepturomyces, the germ- tube of the promycelial spore does not bore its way through the epidermal cells, but enters, as De Bary* first showed with P. dzanthi, through the stomata, or, as Rathay f points out with P. malvacearum, it enters, like the germ-tube of LTubercinia trientalis, between the epidermal cells, and then pierces laterally into the side of the adjacent cell. If the germination of the promycelial spores takes place in water, and not upon their proper host-plant, the germ- tubes sometimes become swollen at their extremities, so as to form reserve spores in the same manner as has been previously described with the uredospores (Plate III. Fig. 24). The mycelium produced from a promycelial spore in the tissues of its proper host-plant in due time gives rise to a fresh spore-development. This spore-development, however, varies in different cases. 1. In certain species the mycelium produces spores. which are exact counterparts of the original teleutospores (Micropuccinia). 2. It may give rise to teleutospores similar in general appearance as to its progenitors, but endowed with the faculty of immediate germination (Leptopuccinia, etc.). 3. It may give rise to a crop of uredospores, which may * De Bary, “Champ. paras.,” Ann. Se, Nat., 4° sér. tome xx, p. 84 (reprint). + Rathay, “‘ Ueber d. Eindringen. Puccinia Malvacearum,” Verhandi. d. Kk, Zool, bot, Ges., band xxxi. (1881), t. 1. Teleutospores. 45 be typical of the species in question. There are instances, however, in which, instead of being truly typical, these uredospores depart in some degree from those subsequently found in the life-history of the Uredine. For example, in Triphragmium ulmarie the primary uredospores do not differ in their individual form and size from the secondary, but the former occur in large dusty sori principally on the petioles and midribs of the host-plant, while the latter are much smaller and scattered over the under-surface of the leaves. A more common deviation, however, consists in the association of the primary uredospores with spermo- gonia (Brachypuccinia). 4. In other instances the mycelium produces spermo- gonia and ecidiospores, The zcidiospores may be upon the same species of host-plant upon which the teleuto- spores occur (Auteupuccinia), or upon some plant of a totally diverse nature (Heteropuccinia) The teleutospores of Endophyllum, although they are produced in the same way and closely resemble zcidio- spores, yet in germination they behave like the teleutospores of Puccinia (Plate IV. Fig. 7). In Coleosporium (Plate IV. Fig. 9) each cell produces a single promycelial spore, while in Melampsora (Plate IV. Fig. 8) and Chrysomyxa (Plate IV. Fig..10) three or four are developed from each promycelium, as is also the case with Cronartium. 46 British Uredinea and Ustilaginee. CHAPTER VII. HETERGCISM. flistory—The fact that a certain number of Uredines possess the faculty of passing a part of their lives upon one plant, and the remainder of it upon another and a totally different one, is so remarkable that until quite recently there were persons who declined point-blank to believe it. It is quite unnecessary now to undertake serzatim to answer the theoretical objections which have been raised against the fact that hetercecism does occur, for the simple reason that these objections are only theoretical. The process of simply placing the spores of the fungi in question upon their various host-plants is so easy, that any one wishing to appeal to Nature herself can do so with very little trouble. The first Uredine in which this peculiarity of develop- ment was observed was Puccinia graminis, the wheat mildew. The mildew of wheat has, as a blight, probably been known from remote antiquity. The Romans held a festival on April 25—the Robigalia, or Rubigalia—with the object of protecting their fields from mildew. The sacrifices offered on this occasion consisted of the entrails of a dog and a sheep, accompanied with frankincense and wine.* In Wickliffe’s Bible, the suggestion is made that * Smith, ‘Smaller Dictionary of Greek and Roman Antiquities,” sth edit. (1863), p. 322. Heterecism. 47 the seven years of famine with which the Egyptians were afflicted were caused by mildew. This, of course, does not prove that the ancient Egyptians had any acquaint- ance with mildew, but it does show that in early English times the disease was not only known as an affection of cereals, but also that it was regarded as an agency suff- ciently powerful to cause famine. In Shakspeare’s time, mildew must have existed, for we read of how “ The foul_ fiend Flibbertigibbet mildews the white wheat.”* The fungoid nature of the mildew was not known, however, until the latter half of the last century, for Tull,t writing in 1733, attributes it to the attack of small insects, “brought (some think) by the east wind,” which feed upon the wheat, leaving their excreta as black spots upon the straw, “as is shown by the microscope”! Felice Fontana,{ some thirty years later, published an account of the fungus, with figures. Persoon,§ in 1797, gave it the name it still bears (Puccinia graminis), and also figured it, as did Sowerby || in 1799, under the name of Uvredo frumenti. With regard to the fact that the barberry in some way favours the growth of mildew upon wheat, there is no’ doubt that it was well known to practical farmers during the eighteenth century, for we find in America, as early as 1760, in the state of Massachusetts, an Act was passed by the legislature, compelling the inhabitants to extirpate all barberry bushes. In New England a similar law existed, which is referred to by Schopf** In our own country, * Shakspeare, ‘‘ King Lear,” act iii. sc. iv. + Jethro Tull, “‘ Horse Howeing Husbandry,” 3rd edit. (1751,) p. 151. + Felice Fontana, ‘‘ Osservazioni sopra la Ruggine del Grano.” Lucca: 1767. : § Persoon, ‘‘Tentamen Dispos. Method. Fungorum ” (1797), p. 39, t. iii. fig. 3. | Sowerby, “ English Fungi,” vol. ii. (1799), t. 140. {| For ‘‘ Barberry Law of Massachusetts,” see p. 302. ** J.D. Schépf, ‘‘ Reise durch die mittleren und siidlichen vereinigten Nordamerikanischen Staaten,” theili. p. 56, Erlangen: 1788. 48 British Uvedinee and Ustilaginee. especially in Norfolk, this belief also existed, for we find Marshall,* writing in 1781, says, “It has long been con- sidered as one of the first vulgar errors among husband- men that the barberry plant has a pernicious quality (or rather a mysterious power) of blighting wheat which grows near it. “ This idea, whether it be erroneous or founded on fact, ‘is nowhere more strongly rooted than among the Norfolk farmers; one of whom mentioning, with a serious counte- nance, an instance of this malady, I very fashionably laughed at him. He, however, stood firm, and persisted in his being in the’right, intimating that, so far from being led from the cause to the effect, he was, in the reverse, led from the effect to the cause ; for, observing a stripe of blasted wheat across his close, he traced it back to the hedge, thinking there to have found the enemy; but being disappointed, he crossed the lane into a garden on the opposite side of it, where he found a large barberry bush in the direction in which he had looked for it. The mischief, according to his description, stretched away from this point across the field of wheat, growing broader and fainter (like the tail of a comet) the further it proceeded from its source. The effect was carried to a greater distance than he had ever observed it before, owing, as he believed, to an opening in the orchard behind it to the south-west, forming a gut or channel for the wind. * * * * * * “Being desirous of ascertaining the fact, be it what it may, I have inquired further among intelligent farmers concerning the subject. They are, to a man, decided in their opinion as to the fact, which appears to have been so long established in the minds of the principal farmers that * Marshall, ‘‘ Rural Economy of Norfolk,” 2nd edit. vol ii, p. ¥9. Lon- don: 1795. fleteracism. 49 it is now difficult to ascertain it from observation, barberry plants having (of late years more particularly) been extir- pated from the farm hedges with the utmost care and assiduity. One instance, however, of mischief this year I had related to me, and another I was myself eye-witness to. Mr. William Barnard, of Bradfield, says, that this year, seeing a patch of his wheat very much blighted, he looked round for a barberry bush, but seeing none conspicuous in the hedge, which was thick, he with some difficulty got into it, and there found the enemy. He is clearly decided as to the fact. Mr. William Gibbs, of Rowton, telling me that a patch of his wheat was blighted in the same manner, and that he believed it to proceed from some sprigs of barberry which remained in the neighbouring hedge (which a few years ago was weeded from it), I went to inspect the place, and true it is that near it we found three small plants of barberry, one of which was particularly full of berries. The straw of the wheat is black, while the rest of the piece is of a much superior quality. “These circumstances are undoubtedly strong evidence, but do not by any means amount to proof.” On October 16, 1782, Marshall writes *— “To endeavour to ascertain the truth of this opinion, I had a small bush of the barberry plant set in February or March last, in the middle of a large piece of wheat. “T neglected to make any observations upon it until a little before harvest, when a neighbour (Mr. John Baker, of Southrepps) came to tell me of the effect it had pro- duced. “The wheat was then changing, and the rest of the piece (about twenty acres) had acquired a considerable degree of whiteness (white wheat), while about the barberry bush there appeared a long but somewhat oval-shaped * Marshall, Joc. cet., p. 359. E 50 British Uredinee and Ustilaginec. stripe of a dark livid colour, obvious to a person riding on the road at a considerable distance. “The part affected resembled the tail of a comet, the bush itself representing the nucleus, on one side of which the sensible effect reached about twelve yards, the tail pointing towards the south-west, so that probably the effect took place during a north-east wind. “ At harvest, the ears near the bush stood erect, hand- ling soft and chaffy; the grains slender, shrivelled, and light. As the distance from the bush increased the effect was less discernible, until it vanished imperceptibly. “The rest of the piece was a tolerable crop, and the straw ‘clean, except on a part which was lodged, where the straw nearly resembled that round the barberry; but the grain on that part, though lodged, was much heavier than it was on this, where the crop stood erect. “The grain of the crop, in general, was thin-bodied ; nevertheless ten grains, chosen impartially out of the ordi- nary corn of the piece, took twenty-four of the barberried - grains, chosen equally impartially, to balance them.” In 1784, Marshall repeated his experiment at Statfold, in Staffordshire, with the same result. He says *— “Upon the whole, although I have not from this year’s experience been able to form any probable conjecture as to the cause of the injury, it nevertheless serves to fix me still more firmly in my opinion that the barberry is injurious to wheat.” _— Withering, writing in 1787 of Berberis vulgaris, says,t “This shrub should never be permitted to grow in corn- lands, for the ears of wheat near it never fill, and its influ- ence in this respect has been known to extend as far as three hundred or four hundred yards across a field.” * Marshall, ‘‘ Rural Economy of the Midland Counties” (1790), vol. ii, p. 11. + Withering, ‘‘ Botanical Arrangement” (1787), 2nd edit., p. 366. fTeteracism. 51 In 1804, this country suffered severely from an outbreak of wheat mildew, in consequence of which Mr. Arthur Young, the Secretary to the Board of Agriculture, issued a circular of questions, so as to obtain a consensus of opinion, from farmers, landowners, and others interested in the sub- ject, as to various points connected with causation of mildew. The ninth question ran thus: “ Have you made any observation on the barberry as locally affecting wheat?” The replies to these questions were published,* and from them the following are selected :— Isaac King, Esq., Wycombe, Bucks, in answer to the question about the barberry, says, “In 1795, a field of about twenty acres had two large barberry bushes growing within twenty yards of it. These appeared to be the focus of destruction to several acres ; in front, close to the hedge, the wheat was as black as ink, and further off it was affected to a less degree. ... In short, I had fifteen acres very good, and five of very little value. You may conclude the bar- berries were destroyed.” Mr. S. Johnson, Thurning, Norfolk, says,t “My ob- servations on the barberry have been for several years. I have seen the blast from a small stem blown on the wheat in one direction upwards of two furlongs, like smoke from a chimney.” __ Mr. W. Maxey, Knotting, Bucks, says,t “When pass- ing a wheat-field a few years ago on the eve of harvest, I noticed some streaks of a different darker hue across a furlong of wheat from the hedge directly opposite ; at the end of each streak was a barberry bush.” __ Mr. James Sheppard, Chippendale, Newmarket, says,§ “T have never seen an instance of wheat growing near a barberry not being injured more or less.” * A, Young, ‘‘ Annals of Agriculture ” (1805), vol. xliii. p. 457. + Marshall, Joc. céz., p. 469. $ Loc. cit., p. 505. § Loe. cit., p. 510. 52 British Uvedinee and Ustilaginee. It is quite unnecessary to quote any further from Mr. Young’s correspondents upon this point. The injurious influence of the barberry was, however, a matter of observation only at this time. Various sugges- tions were made as to the cause of this: some affirmed the barberry bush exhaled a noxious effluvium ; others, that the pollen of its flowers poisoned the wheat ; others, again, that it appropriated to itself all the nourishment from the soil in its- vicinity. In 1805, however, Sir Joseph Banks, in his paper on “ Wheat Mildew,” alluding to the subject before us, men- tions the belief as being prevalent amongst farmers, but scarcely credited by botanists, and points out the resem- blance the yellow fungus on barberry has to rust, although it is larger. He says,* “Is it not more than possible that the parasitic fungus of the barberry and that of wheat are one and the same species, and that the seed is trans- ferred from the barberry to the corn?” The suggestion of our eminent countryman was soon put to the test of experiment by a totally independent observer. The honour of being the first to demonstrate the con- nection between the barberry A®cidium and the wheat mildew belongs to a Danish schoolmaster, who lived in the village of Hammel, near Aarhus, in Jutland, at the begin- ning of the present century. In 1818, the Royal Agricul- tural Society of Denmark published a paper by Schoeler, “On the Pernicious Influence which the Barberry Bush exercises on Cereals.”t This paper was almost over- looked until Mr. Nielsen brought it under notice in 1874, in his capacity as Consulting Botanist to the Royal Agri- * Banks, ‘‘ Annals of Agriculture,” vol. xliii. p. 521. +t Om Schoeler, ‘‘ Berberissens skudelige Indflydelse paa Sceden,” Lazda:- homminske Tidender (1818), part viii. p. 289; Nielsen, Ugeskrift for Land- meda, 1884. Hleterceecism. 53 cultural Society of Denmark. Schoeler began the study of the subject in 1807, when, by closely observing the yellow spots on the under side of barberry leaves, he came to the conclusion that they were due to a microscopical fungus. In 1810, he noticed that the barberry bushes were nearly free from this fungus, and that the rye was that year almost free from rust.* “I then thought,” he says, ‘that there might possibly be a close relationship between the rust on the rye and that upon the barberry ; and when, in the fol- lowing year (1811), I noticed that the rust upon the bar- berry appeared much earlier in the spring than the rust did upon the other plants—grasses and cereals—I thought I had found out the true origin of the rust in rye. Still, however, this question again and again presents itself to my mind, ‘Where does the rust on rye come from in those places in which no barberries are to be found?’ In the summer of 1812 I convinced myself that the barberry bushes are indeed able to communicate the rust to the rye, by means of the wind, even to a considerable distance.” For several years prior to 1813 he experimented in his garden by planting different kinds of corn around barberry bushes, and found that rye and oats were especially liable to be destroyed almost every year by the rust, which always appeared first nearest the barberries. From 1813 to 1817 he planted large and small barberry bushes in his rye-field. He found that the larger bushes did not give rise to the rust in rye, when they lost their foliage in the process of transplanting ; but, on the con- trary, the smaller bushes, which did not lose their leaves so readily, did give rise to the rust in the rye to a very marked degree. One of his experiments he thus describes: “I planted * By the word here translated ‘‘rust” is evidently meant, not only the Uredo, but also the mildew. 54 British Uvredinee and Ustilaginee. in May, 1816, in a rye-field, three small barberries ; one of them did not thrive, but the other two developed nineteen leaves before the end of May, nearly all of which had the yellow spots (of Acidium berberidis) upon them. When, in the middle of the following June, the rye immediately surrounding these two bushes became rusty, I invited, by a notice in the newspapers, every one interested in the question to come and convince themselves of the pernicious influence which the barberry exerts. On account of this invitation many people came to visit me, and all of them, the learned as well as the unlearned, declared that they could accept no other cause for the rust on the rye than _ the small barberry plants.. They were astonished that so small a cause should have produced so great an effect. On June 22, most of the rye plants for thirty to forty feet (square feet) around these small barberries were more or less rusty, mostly so to the north and north-west, but for a long time afterwards not even a single rust spot could be found elsewhere in the field.” In the same year (1816) Schoeler performed the follow- ing experiment :—Some fresh branches of the barberry bush having rusty leaves upon them were cut off, put into a box, and carried to a rye-field, where the rye was still moist with dew. The rusty barberry leaves were applied to some of the rye plants—to the straw as well as to the leaves—by rubbing them with the underside of the affected barberry leaves, until he could see some of the “yellow dust” (spores) of the fungus adhered to the rye plants. The infected rye plants were then marked by tying them to sticks driven into the ground. In five days’ time these plants were badly affected with rust, “while at the same time,” says Schoeler, “not one rusty plant could be found anywhere else in the whole rye-field.” The question, however, was not even now fully decided ; fletereecism, 55 for although the above facts were in themselves unanswered, yet the so-called scientific botanists urged that the fungus upon the barberry leaves belonged to a totally distinct genus (AEcidium) to that upon the wheat (Puccinia). There is evidence to show that many careful observers, even at this time, suspected that the Puccinia was connected in some way with the uredospores which occur as its pre- cursors, This remained a suspicion only until Tulasne_ demonstrated that the connection between the uredospore and the teleutospore existed not only in the species in. question, but was the general rule amongst the Uredinez. In 1861, De Bary showed that many of the Uredinee not only had uredospores and teleutospores, but also that the latter gave rise in many cases (but not in all) to zcidio- spores, and conversely the zcidiospores to uredospores. De Bary also pointed out that in certain cases the sowing of germinating teleutospores upon the same species of host-plant which bore them was not followed by any result. Amongst these were Puccinia graminis. It further occurred to him that, as there were several zcidia unac- companied on their host-plants by any other spore-form, these might belong to Uredines which passed a part of their life upon one plant and the remainder upon another. Familiar with the facts already known to the practical agriculturist concerning the barberry and wheat mildew, he put the matter to the test of actual experiment. In 1864, he sowed Puccinia graminis on barberry and produced the Aicidium, and in 1865 he did the converse culture, by sowing the zcidiospores upon rye. The results obtained by his experiments with P. gvaminis led De Bary to investigate the life-histories of other zcidia, which, like 4c. berberidis, are unaccompanied by any other spore-form on the same host-plant. Thus he found that P. rubigo-vera has its ecidiospores upon Lycopsis 56 British Uredinee and Ustilaginee. arvensis, and P. coronata upon Rhamnus frangula* In the following year, 1866, Oersted showed that Gymnosporangium sabine and juniperinum were similarly connected with Restelia cancellata and cornuta; and in 1867, that G. cla- varieforme was connected with . lacerata. In 1869, Fuckel indicated the connection of Uvomyces junc with ic. zonale. In 1873, Magnus worked out the life-history of P. caricis in its relationship to 4c. urtice ; and Schréter, in the same year, the hetercecism of Zc. ranunculd bulbosi and Uromyces dactylidis. Since this time these facts have been repeatedly verified by numerous workers, and our knowledge of the subject has continuously increased, as the subjoined tabular statement will show. * Puccinia graminis fEcidium berberidis De Bary | 1864 * Puccinia rubigo-vera Aécidium asperfolii De Bary | 1865 * Puccinia coronata Aécidium rhamni De Bary | 1865 * Gymnosporangium sabine | Reestelia cancellata Oersted | 1866 * Gymnosporangium junipe- | Reestelia cornuta Oersted 1866 rinum 3 * Gymnosporangium clava- | Reestelia lacerata Oersted | 1867 riceforme * Uromyces junci fEcidium zonale Fuckel 1869 * Puccinia caricis fEcidium urticee Magnus 1873 * Uromyces dactylidis 4écidium ranunculi-bulbosi | Schroter | 1873 * Coleosporium senecionis Peridermium pini Wolff 1874 t Puccinia moliniz AEcidium orchidearum Rostrup | 1874 * Puccinia sessilis Aécidium alli Winter 1874 * Puccinia phragmitis 4Ecidium rumicis Winter 1874 Uromyces pisi Ecidium cyparissize Schroter | 1875 * Puccinia poarum AEcidium tussilaginis Nielsen 1876 Puccinia limosze Ecidium lysimachize Magnus | 1877 Puccinia seslerize Ecidium rhamni-saxatilis Reichardt | 1877 Puccinia sylvatica Aicidium taraxaci Schroter | 1879 * Uromyces pox AKcidium ficarice Schréter | 1879 Chrysomyxa rhododendri | Aicidium abietinum De Bary | 1879 Chrysomyxa ledi Aacidium abietinum De Bary | 1879 Calyptospora goeppertiana | Aécidium columnare Hartig 1880 t Melampsora populina “Ecidium clematidis Rathay 1881 * Puccinia magnusiana fEcidium ranunculi-repentis | Cornu 1882 t+ Melampsora caprearum Ceeoma euonymi Rostrup 1883 + Melampsora hartigii Czeoma ribesii Rostrup 1883 t Melampsora tremulze Czeoma mercurialis Rostrup 1883 Puccinia dioicze /&cidium cirsii Rostrup 1883 * De Bary, ‘‘ Neue Untersuch. iiber Uredineen,” 2nd paper (1886), pp- 208—214. Fletevccism. Bo Puccinia eriophori AEcidium cineraria Rostrup | 1883 * Puccinia obscura Ecidium bellidis Plowright | 1884 * Puccinia schdleriana Akcidium jacobzeze Plowright | 1884 * Puccinia perplexans Ecidium ranunculi-acridis | Plowright | 1884 Puccinia vulpine Ecidium tanaceti Schroter | 1884 * Puccinia arenariicola Ecidium centaureze Plowright | 1885 * Puccinia phalaridis ABcidium ari Plowright | 1885 + Melampsora tremulze Czeoma laricis Hartig 1885 * Gymnosporangium bisep- | Reestelia botryapites Farlow 1885 tatum * Gymnosporangium confu- | Acidium mespili Plowright | 1886 sum Puccinia pollinise 4&cidium strobilanthis Barclay 1886 Gymnosporangium calvipes | Roestelia aurantiaca Thaxter | 1886 Gymnosporangium macro- | Roestelia pyrata Thaxter | 1886 us Crmartiita asclepiadeum | Peridermium acicola Cornu 1887 * Puccinia extensicola AKcidium asteris Plowright} 1888 * Melampsora zcidioides Czoma mercurialis Plowright} 1888 * Puccinia paludosa Aicidium pedicularis Plowright | 1888 * Puccinia persistens fEcidium thalictri-flavi Plowright | 1888 * Puccinia trailii AEcidium acetosze Plowright | 1888 * is affixed to those species which I have personally investigated. + to those of which I have repeated the cultures, but am not able to confirm the above statements. The question naturally presents itself to us, Why are some species hetercecious and others not? One reason is pretty obvious, namely, that those Puccinize and Uromyces which are hetercecious occur upon host-plants whose cuticle is, if not silicous, at least very hard and difficult for the germ- tube of the promycelial spore to pierce—namely, on grasses, Carices and Junci. This, however, can hardly be the only reason, since Schréter has produced the A‘cidium on Euphorbia cyparissias from Uromyces pist ; in this case both the host-plants have soft epidermal cells. The Coleosporia and Melampsore afford similar instances. Whatever may have been the cause or causes in bygone ages, the fact is that at the present time so completely have these parasites become hetercecismal in habit, that the most profuse appli- cation of their promycelial spores to the graminaceous host is always without result. 58 British Uredinee and Ustilaginee. CHAPTER VIII. MYCELIUM OF THE USTILAGINEZ, THE vegetative mycelium of the Ustilaginee is to the parasite one of its most important organs, for by it, and by it alone, does the fungus derive its nutriment from the host-plant upon which it subsists. Yet the mycelium is of all parts that which is least frequently observed ; nor can this be wondered at when one remembers that, like the mycelium of the Uredinez, it can be seen only by careful search —by cutting and teazing out numerous thin sections of the host-plant. We owe most of the information we possess upon the mycelium of the Usti- laginee to Fischer von Waldheim,* although, among others, both Kiihnt and Hoffmann} had previously figured it. It exists most abundantly in the tissues of the host-plant, in the immediate vicinity of those places in which the spores are developed; but it can also be found in other parts of the affected plant—in the monocotyledons particularly in the stem, but especially in the nodes and in the root-stock. In the dicotyledons it is not so easily found at a distance from the spore-beds ; still, in them too it has been seen in the root-stock and * Fischer von Waldheim, ‘‘Pringsheim Jahrbiicher” (1869), vol. vii. pp. I, 2. + Kithn, “ Krankheiten der Kulturgewichse,” 2 aufl. 1859. t Hoffmann, “ Ueber der Flugbrand.” 1866, Mycelium of the Ustilaginee. 59 stem, especially in the nodes. The mycelium consists of hyaline tubes, frequently septate, and enclosing watery or pellucid, frequently vacuolated contents (Plate V. Figs. 1, 2). Its walls vary in thickness, but very often they have a distinctly double contour. The number and frequency of the septa are subject to much variation ; in some instances they are close together, at others they are only found at distant intervals. This is also the case with the mycelial ramifications ; sometimes the hyphe do not extend for more than 2u without branching, at others they extend for 20u or more without dividing. These long unbranched hyphz are found mostly in the internodes ; in the nodes themselves not only are the branches more abundant and convoluted, but here too are encountered, more abundantly than elsewhere, the little intercellular haustoria, or suckers, which characterize the mycelia of many of the Ustilaginez. The mycelium ramifies not only between the cells of the host-plant, but, frequently piercing their walls, grows through them. Its diameter varies from 2 to 5u. The addition of caustic potash to a section of the host-plant containing mycelium renders the latter more distinct, and otherwise clears up the preparation ; so does prolonged treatment in glycerine. Its walls are not composed of cellulose, as they do not show any blue reaction when treated with sulphuric acid and iodine; but in some cases, as with U. maydis and Sorosporium saponarig, they do get an external coating of cellulose from the tissues of the host- plant, which, completely investing them, hides them from view (Plate VI. Fig. 3). The mycelium of almost all the Ustilagineze permeates more or less the whole of the affected plant, and although in the advanced state we can find it only near the spore-beds, yet originally it could be found in all parts of the axis of the young plant. In this it differs from the localized mycelia of most of the 60 British Uredinee and Ustilaginee. Uredinee. These permeating mycelia, whether of the Uredinee or Ustilaginee, are perennial. When the plant, if perennial, dies down in winter, the mycelium, of course, dies down with it, but remains alive, although quiescent, in the upper part of the root-stock ; and when fresh shoots are sent up in spring, the mycelium is sent up in them. One peculiarity of most of the Ustilagineous mycelia is that, although it pervades more or less the whole plant, it produces its spore-formation at certain favoured places only ; these are, for the most part, in the flowers or seeds of the plants, but not always, sometimes on the stems or in the leaves. The place of spore-formation, however, is constant with each species. If a plant affected with one of the Ustilaginee be transplanted into a garden, it will, year after year, be affected with the parasite. In my own garden at King’s Lynn I have had growing for the past six years, plants of Colchicum autumnale affected with Urocystis colchicd, Triticum repens with U. hypodytes, and Avena elatior with U. segetum. De Bary mentions that a plant of Saponaria offcinalés, in the Freiburg Botanic Garden, was for more than ten successive years affected with U. violacea. ( 61 ) CHAPTER IX. FORMATION OF THE TELEUTOSPORES OF THE USTILAGINEA. WHILE it is true that the spores of Ustilaginee are formed from the mycelium, yet the process does not take place directly from the vegetative mycelium which has just been described. On the contrary, at those favoured parts of the affected, host-plant at which the spores are developed the vegetative mycelium often quite suddenly changes its character. The double-contoured hyphe with pellucid vacuolate contents lose their double contour, become swollen or distended, and contain, instead of a clear watery fluid, a gelatinous, granular protoplasm. in which numerous oleaginous particles may often be seen (Plate V. Figs. 3-6). The gelatinization of these spore-forming hyphz is a great character of the Ustilaginez ; it does not, however, occur in all species. The first change observable in the mycelium before it becomes a spore-forming hypha, is that its walls increase in thickness at the expense of its calibre, which becomes proportionately diminished ; soon, however, the whole hypha becomes dilated, so that its lumen is increased. Its contents can now, by the action of reagents, be shown to consist of protoplasm. Spore-formation takes place, after these changes in the mycelium, so differently in the different genera that it will be necessary to describe the process, as it takes place in each one, separately. 62 British Uvedinee and Ustilaginee. Ustilago—The spore-forming hyphe enlarge and branch in various ways. The gelatinization of their in- terior takes place to such an extent as to almost obliterate their lumen, which, however, may frequently be seen as a narrow shining line in the middle of the hyphz (Plate V. Fig. 7). At certain points the surface of these hyphe en- large, so that they appear nodose. The increase in size - of the hypha continues, so that adjacent hyphze become variously tangled and intertwined together, and eventually many of the hyphe appear glued together, or to have coalesced. The irregularities of the hyphe become more marked, and it is obvious that each tumefaction will even- tually become a spore, inasmuch as they gradually get more and more rounded (Plate V. Fig. 8). It is in the interior of these distended hyphe that spore-formation takes place. It is, however, always found that the external spores are the most developed, the formation being, in fact, centripetal. The commencement of the differentiation of the protoplasmic contents is at the exterior of the mass, and it gradually proceeds inward towards the centre. The spores when first formed have gelatinous envelopes, and gradually become more or less polygonal from mutual compression. The interior of the spore is now seen to have a distinct contour, and to contain fatty granules. The outer edge of this contour darkens, and even while it is still surrounded by a thin gelatinous envelope the irregularities of the epispore begin to be apparent. As the spores ripen this gelatinous membrane disappears, so that at their maturity they have no remnant of it; nor are any remains of the mycelial hyphe attached to them, as is often seen in Tilletia. Sphacelotheca.—This genus differs from Ustilago in the spore-mass being developed in a receptacle. De Bary thus describes its development: “The vegetative mycelium, Formation of Teleutospores of Ustilaginee. 63 entering the ovary through the flower-stalk, sends its hyphz through the funiculus into the ovule, which becomes permeated by densely interwoven hyphe. The micro- : pylar end of the integuments alone escapes and remains, as a cap on the top of the diseased ovule, for some time, Fig. 4.—Sphacelotheca hydropiperis (Schum.) A, Ovary and perianth of Polygonum hydropiper affected with S. hydropiperis. B, The same more advanced, showing the micro- pylar cap (c). C, Section of ovary and perianth at an early stage 5 & the style; o, the imtegument of the micropylar end of the ovule; / wall of the ovary; Z, the perianth. The spore-formation is seen to be commencing above, and the rudimentary columella is visible. D, Section of a more advanced ovary, showing the walls of the receptacle and the columella (c). Slightly magnified. (De Bary.) but eventually falls off. The hyphz develop partly into spores and partly into the receptacle. The latter consists of a thick external case with a central columella. The cells of which it is composed are but loosely compacted, colourless, and about the size of the spores, The least injury fractures this case and allows the escape of the 64 british Uredinee and Ustilaginee. spores.” The spore-formation itself is similar to that of Ustilago,* and, commencing above, proceeds downwards, Sorosportum,— The spore-formation in Sorosporium differs considerably from that which has just been described in Ustilago, although it is obviously of the same type. In Ustilago the mature spores are separate and distinct, form- ing usually a pulverulent mass. In Sorosporium, on the other hand, they are in their perfect state aggregated into spore-balls, which individually often contain fifty or a hundred separate spores. The process of spore-formation has been studied by Von Waldheim with S. sagonaria, and is as follows :—The mycelium, which is very abundant in the blossom and ovary, rapidly changes into spore-form- ing hyphe, from 4 to 74 in diameter, which are gelatinous and full of shining protoplasm. The free ends of these hyphe have a tendency to curve inwards and roll them- selves up (Plate V. Fig. 9). The spore-forming hyphe from several contiguous mycelial branches, incline together, and twist themselves into a ball, as happens in the forma- tion of a lichen thallus. These convoluted and contorted spore-forming hyphe, being gelatinous, soon become so intertwined and entangled that they cease to be indi- vidually recognizable; to all appearances they coalesce together in part, if not entirely, and on the exterior of this gelatinous ball other hyphe are now seen encircling it (Plate V. Figs. 10,11). These latter, also being gelatinous, soon lose their individuality, although at times traces of their concentric arrangement can be made out. Spore- formation takes place only in the central gelatinous ball, in the middle of which it commences by the central part darkening in colour and becoming differentiated into spore-like bodies, which vary in number from four to sixteen. Apparently these bodies again subdivide, so * De Bary, “‘ Vergleichung,” p. 187, ‘Formation of Teleutospores of Ustilaginee. 65 that when the spores arrive at their maturity the spore- balls contain sixty to a hundred or more spores. In the young state these developing spores are polygonal from mutual pressure, and they are to be found in spore-balls not more than Sou in diameter. In the subsequent de- velopment of the spores the balls increase in size, and the gelatinous zone swells also. When, however, the spores assume their characteristic deep brown colour this gela- tinous zone begins to be absorbed, having been utilized in the development of the spores. In spore-balls of 7ou in diameter the gelatinous zone is only from 4 to 6u thick. It entirely disappears when the spores have attained their full maturity. In a certain sense it may be said that the spore-formation is centrifugal, inasmuch as it commences in the centre of the gelatinous ball; but the peripheral spores are as in Ustilago, the oldest, having been pushed outward by the continued formation of the younger spores in the centre of the mass. These externally placed sporcs either continue their development independently, or, what is more probable, their spore-forming hyphze have become greatly elongated, but still remain in connection with the host-plant (Plate V. Fig. 12). In Sorosporium, however, certain solitary spores occur independent of those aggre- gated into spore-balls. The development of these isolated spores takes place in single hyphe. The end of the hypha becomes gelatinized, swells up, and a spore is developed inside. They are at first surrounded by the gelatinized hyphae, which generally, however, disappear entirely by the time the spores are mature. _ Tubercinia—The spore-formation of Tubercinia has been worked out by Woronin* in Tudbercinia trientalis. * Woronin, “ Beitrag. sur Kenntniss der Ustilagineen” (1882), pp. 4-16, t. ii. figs. 3-10. De Bary and Woronin, ‘‘ Beitrage zur Morphol. und Physiol. der Pilze,” 5 reihe. 1882. F 66 British Uvredinee and Ustilaginee. -Subjoined is a summary of his observations. The mycelium pervades all parts of the affected plant, excepting the roots. It consists of intercellular, branched, sparsely septated, hyaline hyphz, having a diameter of from 2 to 3u. It is most abundantly distributed in the cortical tissues, and sends botryform haustoria (Plate VI. Fig. 1) into the adjacent cells. At certain places it gives off smaller, more richly septate branches, unprovided with haustoria, which are destined to form the spore-beds. This is accomplished in the following manner :—A number of straight and rather larger branches are given off, which soon become curved and interwoven in various ways, gene- rally more or less spirally, so as to form an entangled knot. The spore-bed which this entanglement forms develops spores from within outwards ; each spore-ball contains from fifty to a hundred spores, The spores measure from 55 to 75u in diameter, and consist of an endospore, containing granular protoplasm, and an exospore. No gelatinization of the spore-forming hyphe, such as takes place in the spore-formation of Sorosporium saponari@, was observed. Tilletia.—The spore-formation in this genus was first indicated by Tulasne,* afterwards by Kiihn,t who gave a figure of the process; but it is to Von Waldheimt that we are indebted for the most detailed account. He thus describes the process: The vegetative mycelium of Tilletia tritict is nearly 2u in diameter. The swelling and gelatinization of the spore-forming hyphe is not so marked as in Ustilago, so that the lumen of the hyphe is never so contracted. Spore-formation begins by the hyphe giving off pyriform buds, I°5u across (Plate VI, Fig. 4), in succession, from their sides ; these outgrowths in- * Tulasne, ‘17 Mémoire sur les Ustilaginées comparées aux Uredinées,” Pp: 27, e¢ seq. + Kiihn, ‘‘ Krank. der Kulturgewachse,” p. 56, t. iv. fig. 5. t F. von Waldheim, Joc. cet. Formation of Teleutospores of Ustilaginee. 67 crease in length and breadth, so that the pyriform swellings become spheres, from 2 to 3 across, attached to the hypha by thin stems about Iu thick. These become differentiated ; their contents vacuolated and oleaginous, and the con- necting branches speedily wither (Plate VI. Figs. 5, 6). The epispore subsequently darkens and becomes uneven, while to many ripe spores the remains of the spore-forming hyphez continue attached. In 7. strvieformis the process is very similar, but the spore-forming hyphe are larger and more gelatinized, and invest the spores to their maturity, after which they dis- appear without leaving any trace. Doassansia.*—The mycelial hyphe in the tissues of the host-plant give off branches, which at certain points become interlaced into tangled knots. From these knots are formed the spores; the central portions forming the true spores, while the external develop into a layer of oblong or wedge-shaped sterile cells, which constitute an investing peridium. The peridial cells are darker in colour than the spores (Plate VIII. Fig. 4). Entyloma—tThe spore-formation in Enty- loma is very similar to that of Tilletia. De Bary ft investigated it in E. mzcrosporum and calendulg. The much-branched mycelium is principally intracellular. At certain places blister-like swellings appear along the spore- forming hyphz, and also at their ends. The ,,F8: 573" Hers : 1 f contents of these swellings become differen- ine’ scbutospores tiated into spores, so that they are intercalated foming.bpha 5 ; e Bary. in the hyphe (Fig. 5). Often a series of r spores, formed one behind the other, may be seen still con- * Cornu, Ann. des Science Nat., 6° sér., Bot., tome xv. p. 280, ef seg. “+ De Bary, Bot. Zeitung (1874), pp. 81-93; pp. 97-108, t. ii, 68 British Uredinee and Ustilaginee. nected with the remains of the spore-forming hyphe. At maturity these spores do not break up into dusty masses like Ustilago, but remain as compact colonies embedded in the tissues of the host-plant. Each spore has two coats, and contains coarse protoplasm. The outer coat is some- times gelatinous at its maturity. Urocystis.—-The process has been studied by Kiihn,* De Bary,t Wolff,f and Winter§. The spore-forming hyphz become swollen at their ends ; two or more of these branch and wind themselves together, generally in a spiral manner, so as to form a glomerulus, and then, becoming highly gelatinized, they are quite indistinguishable from each other. In JW. occulta, Wolff considers that they have a common investing membrane. At the same time, other branches are given off from the spore-forming hyphe, which apply themselves to the outside of the glomerulus (Plate V. Figs. 13, 14, 15). As Winter points out, this might be looked upon as a sexual act, the central spiral branches of the glomerulus being the carpogonium, and the external enveloping branches the pollinodium ; but there is no proof that this is really a sexual act, especially as similar conditions occur in the spore-formation of various Ustilaginez, and in Geminella, in which no sexuality occurs. The spores are formed entirely from the gelatinized central glomerulus, and, as De Bary first showed, the pseudo- spores are formed by the outer branches. Prillieux || con- siders that the spores of Urocystis are formed in a similar manner to those of Sorosporium. The details of his obser- vations are given on the plate which accompanies his paper. * Kiihn, loc. cit., pp. 78, 79. + De Bary, ‘‘ Morph, und Physiol.,” p. 125. t Wolff, Bot. Zedtung, 1873 ; “ Der Brand des Getreides,” 1874. § Winter, Flora, 1876, ‘‘ Ustilagineen.” || Prillieux, ‘‘Sur la formation et la germination des Spores des Urocystis,” Ann. des Science Nat., 6° sér., Bot., vol. x. pl. i, Formation of Teleutospores of Ustilaginee. 69 Localization of Spore-beds.—The bulk of the Ustilagineze are characterized by the constancy by which they repro- duce their spore-formation, in the same tissues of their respective host-plants. In the majority of the species the spores are localized somewhere in the reproductive organs of the host-plant ; very frequently in the ovary (T77dletia tritici, decipiens, U. caricis, Thecaphora hyalina), or in the anthers (U. violacea, scabios@) ; often the blossoming and fructifying organs are attacked and destroyed (U. segetum, bromivora, kiihneana, Sphacelotheca); sometimes the leaves are affected (U. longissima, T, strieformis, Uroc. occulta), or the stems (U. hypodytes, grandis), or even the subterranean organs (U. hypogaea, Uroc. gladiol). With Entyloma and its allies, however, this selection of tissue by the parasite does not obtain; with these species the spore-formation seems to occur in almost any part of the plant. Structure of the Ti eleutospores.—The teleutospores of the Ustilaginez consist of two membranes: an outer, which is thick and generally dark-coloured—the exospore; and an inner, which is thin and hyaline—the endospore. The exospore is subject to considerable variety; it may be quite smooth, or covered with extensive reticulations. Between these various intermediate conditions occur ; thus in U. segetum it has generally been regarded as smooth, but it is rather to be described as granular, although Winter speaks of it as being “generally very minutely verrucose, and seldom quite smooth.” The degree of roughness of the epispore varies from extremely minute elevated points to marked tubercles. These elevations may be evenly distributed over the whole spore, or they may be confluent in lines or ridges. Sometimes these ridges anastomose, and so form a reticulate or alveolate spore (7. ¢rzticé, Sphacelotheca). The colour of the spore depends upon that of the exospore; as a general rule the spores, as seen x 70 British Uredinee and Ustilaginee. en masse, are blackish, but frequently, however, with an olive-brown or yellowish lustre, especially when viewed in an oblique light. UW. segetum, when it occurs in wheat, has a distinctly golden lustre, but when on Avena elatior it is sooty black. Physiological research will possibly show that these two forms are specifically distinct. With U. scabtose the spores in bulk are flesh-coloured, and in U. succise they are quite white. Individually, the spores of the various species, as seen under the microscope, afford a considerable range of colour—black, dark violet, brown, olive-brown, and yellowish, or quite colourless. In some species a germ-pore is said to exist, through which the promycelium is protruded in germination ; but these germ- pores of the Ustilaginez are by no means so marked a formation as in the Uredinee. In U. tragopog? the germ- pore is said to occupy from one-quarter to one-half the epispore.** Much more commonly do we find, as in T. trttic?,} a small opening which splits into a rift as the promycelium grows out. In Thecaphora hyalina the germ- pore is round and paler in colour than the rest of the epispore ; moreover, it is smooth, while the epispore is verrucose. Upon the whole, although germ-pores probably exist in all species, they are inconspicuous, and are very easily overlooked in the smaller spores. The reticulations on the epispore of Sphacelotheca are shown by the action of sulphuric acid, when examined: by a high magnifying power, to consist of a series of distinct palisades, placed vertically.f As a general rule, the spores are globose, but in most species this is subject to a certain amount of variation ; they often have one diameter rather longer than the other, but more frequently they show the result * De Bary, ‘‘ Morph. und Physiol.,” p. 128. + Brefeld, ‘‘ Hefenpilze,” p. 48. { F. von Waldheim, ‘Sur la structure des spores des Ustilagingées ” (1867), pp. 243-245. Formation of Teleutospores of Ustilaginee. 71 of mutual compression by being flattened in one or more directions. In JU. caricis they are often flattened on one side, so as to be subhemispherical. In size they vary from 4u in U. hypodytes to 30u or more in Uvocystzs fischert. They may be simple or collected into spore-balls (Soro- sporium), in which case they appear, when seen separately, asin Z. hyalina, to be segments of a sphere, being convex externally, but internally more or less wedge-shaped. In Urocystis the spore-balls are surrounded by a variable number of barren spores or pseudospores, which are paler in colour, often almost hyaline ; these do not germinate. Hartsen * found that the spores of the U. maydis would not yield their colouring matter to any of the ordinary solver.ts, but that strong sulphuric and nitric acids de- colourized the epispore, rendering it more transparent without at once destroying it, so that the contour of the exterior remains unaffected. Although the spores some- times burst, yet by these reagents the structure of the epispore can be conveniently examined. In nitric acid the spores of the U. maydis swell up, and after a time they dissolve, giving off an odour of bitter almonds. Sul- phuric acid is the better reagent to employ in the exami- nation of the spores, as, although it decolourizes and renders the epispore transparent, yet it does not so rapidly destroy the latter. * Hartsen, ‘Compt. rendus ” (1874), pp. 441, 442. 72 British Uredinee and Ustilaginee. CHAPTER X. GERMINATION OF THE TELEUTOSPORES OF THE USTILAGINEE, ArT the beginning of the present century, Prevost * discovered the fact that the spores of U. segetum and T. tritici, when placed in water, would germinate. He observed the pro- cess in Tilletia to consist of the protrusion of a germ-tube and the development upon it of primary spores, which became united in pairs below, and which bore above the secondary spores. His observations were confirmed by De Candolle,t by Caron and Vandenhecke,t and by Mr, Berkeley,§ and they have been accurately described and delineated by almost all the more recent observers. With U. segetum Prevost observed the germ-tube, and that it gave off small secondary branches. His observations were confirmed by Tulasne,|| Bonorden,{ Kiihn,** and others. Little additional light had been thrown upon the subject, * Prevost, B., ‘‘ Mém. sur la cause immédiate de la Carie.” Montauban; 1807. g + De Candolle, ‘‘ Physiol. Veget.” (1832), vol. iii. p. 1436. + In 1835 referred to by Tulasne, ‘‘ 1° Mémoire,” p. 38. § Berkeley, ‘‘ Propagation of Bunt,” Zrans, Hort. Soc. London (1847), vol. ii. p. 113. | Tulasne, ‘‘Mém. sur les Ured, et Ust.,” dan. de Sct. Nat., 4° sér., tome ii. p. 113. { Bonorden, ‘‘ Handb, d. Allg. Mykologie” (1851), p. 39. ** Kiihn, ‘Krank, Kult.,” 2 aufl, 1859. Germination of Teleutospores of Ustilaginee. 73 until Brefeld,* in 1882, published his investigations, by which it appears that the Ustilagineze are capable of per- petuating themselves for long periods, outside and inde- pendently of their host-plant, in the excreta of herbivorous animals. It will be more convenient to describe the ger- mination of the spores of the various Ustilaginez in detail, as was done with their spore-formation. Much confusion exists in the works of various authors who have written upon the subject, from the diverse appli- cations to which the word “spore” has been made. For instance, Cooke} speaks of the teleutospores of the Ustila- ginez as pseudospores, which term he applies to the zcidio- spores, uredospores, and teleutospores of the Uredinez. Again, these bodies are sometimes called conidia, which term is applied by Brefeld to the secondary spores of the Ustilaginez. In order to avoid confusion, it may as well be stated at once that while the term “spore” may be correctly applied to all the reproductive bodies possessed by these fungi, in a general sense, yet it becomes necessary to affix to it some qualifying word, such as teleuto-spore, resting-spore, uredo-spore, promycelial spore, zecidio-spore, and so forth. In speaking of the Ustilaginez, the word “spore” has generally been applied to the perfect, last- ‘formed bodies—the teleutospores, analogous to the teleuto- spores of the Uredinez ; those bodies, in fact, with which we are all familiar as the black dust of bunt and smut. These black teleutospores, when they germinate, protrude a germ-tube—the promycelium. This promycelium bears certain very small hyaline bodies, which are spores, and may be fairly enough designated promycelial spores, inas- much as they have been produced by the promycelium ;- as, however, the term “ sporidia” is very commonly applied * Brefeld, ‘‘ Hefenpilze.” 1883. t Cooke and Berkeley, ‘‘ Fungi, x: 1875, » Tnternational Scientific Series, vol, 74 British Uredinee and Ustilaginee. to them by Continental botanists, we may use it when writing or speaking in English. The promycelial spores very commonly produce second- ary promycelial spores by budding. Hence we may speak of primary and secondary promycelial spores. When these promycelium spores continue many times to multiply them- selves by budding, after the manner of Saccharomyces, Uredine spermatia, etc. they may very well be called, as Brefeld suggests, “ yeast-spores” and “ yeast-colonies.” The term “ conidia” will be confined to that form of fruit in Tubercinia and Entyloma which is produced in the air from the mycelium in the living host-plant. Ustilago.—The germination of the spores of Ustilago varies somewhat in different species. The commonest type is that of U. segetum, violacea, maydis, kiihneana, scabios@, etc. U. segetum.—If a few spores be placed in a drop of water, they will begin to evince signs of vitality in six or eight hours. Germination occurs more rapidly in summer than in winter, and in fresh spores than in those which have been kept some months. At one point of its surface the spore emits a germ-tube, which grows straight out- wards, until it is from three to four times as long as the spore is wide; and under certain circumstances this tube may, according to Kiihn, have the functions of a germ- tube (Plate VII. Fig. 6), entering by its pointed extremity the tissues of the host-plant. Normally, however, it becomes divided by septa into from three to five compartments, generally into four. This germ-tube is an outgrowth of the endospore, which is pushed upward through the exo- spore. It is from 30 to 4ou long, and from 4 to 5u broad at its maturity. At first it is in direct communication with the endospore, and the protoplasm therein contained passes. into the germ-tube and fills it. Adopting the phraseologt, Germination of Teleutospores of Ustilaginee. 75 of De Bary, this germ-tube constitutes the promycelium. In the course of a few hours the promycelium has received into itself all the protoplasm originally contained in the spore. Transverse septa now make their appearance in it, and it thus becomes divided into four or five equal compart- ments. From the outer walls of the now septate promy- celial tube little offshoots or buds arise, into which the _ protoplasm of the tube passes (Plate VII. Fig. 1). These buds continue to increase in size until they become elon- gated, ovate, or elliptical promycelial spores; they then fall off. Generally they are produced from the side wall of the promycelium, near the septa, and almost always one is produced from the apex of this structure. If the proto- plasm in each segment be exhausted by the production of promycelial spores, then spore-formation from it ceases ; but if all the protoplasm be not used up in the formation of the first promycelial spore, a second but usually a smaller one is budded off. In fact, spore-development from the promycelium goes on until its protoplasmic contents are exhausted. It is not at all uncommon for one of these primary promycelial spores to remain attached to the pro- mycelium instead of falling off, and at its free end to give off a small bud, which gradually grows into a secondary promycelial spore, the latter being of smaller dimensions than the one from which it sprang (Plate VII. Fig. 2). These primary and secondary promycelial spores, after they have fallen away from the promycelium, show still further developmental changes in water. (1) They may, as Tulasne has figured, emit a germ-tube (Fig. 12)—a very narrow tube pointed at its extremity, into which the contents of the promycelial spore are passed.* (2) Two promycelial spores, being near one another, may become joined by a transverse branch, through which the contents of one of * Tulasne, ‘2° Mémoire,” pl. 12, figs. 22-24, 76 British Uvedinee and Ustilaginee. them passes into the other (Fig. 11). When the first pro- mycelial spore has become emptied of its contents, the second emits a tube which may remain as a germ-tube, or, as it becomes full of protoplasm, its end may swell out and form a third spore. (3) At a variable distance from the spore from which it arose a detached spore may form a connection with one of the segments of another promyce- lium ; sometimes as many as three spores may thus become united.* Germination, however, does not always occur in the above typical manner, namely, by the development of pro- mycelium and promycelial spores. From some of the largest teleutospores two promycelia are occasionally given off (Fig. 7). More commonly we find that, instead of pro- mycelial spores being produced in the regular manner above indicated, only one or two segments give rise to them. The others send off branches, into which their con- tents are emptied in the same manner as occurred when spores were formed. The free ends of the promycelial branches often come into contact with one another, When this happens they fuse together and become one continuous tube. (1) Thus a tube given off from one of the upper segments may form a connection with one of the lower segments of the same promycelium in the form of a bow (Fig. 4). (2) Two continuous segments may become united ° by forming what Brefeld calls a buckle-joint. This con- sists of the unequal growth of one side of the promycelium at the level of one of the septa; as this growing-out con- tinues the promycelium itself becomes bent at an angle, at first obtuse, but eventually acute. A reference to the figures (Plate VII. Figs. 3 and 9) will render this obvious. (3) Promycelia from two different spores may unite by branches in various ways, either by their ends or at any * Brefeld, loc. czt., pp. 54-67, t. ii, iii., figs, 1-17. Germination of Teleutospores of Ustilaginee. 77 part of their length ; or the end of one may become united with the central segment of another, and so forth (Fig. 5). These various fusions take place only in spores grown in water. Brefeld has shown, however, that when cultivated in a medium which is capable of supplying suitable nutri- ment to the spores, these various fusions do not take place. He discovered that an aqueous extract of the excreta of herbivorous animals, sterilized by discontinuous boiling, afforded such a medium (ndhrlisung). Spores of U. sege- tum placed in Brefeld’s nutrient fluid germinate sooner than in water. Not only so, but the promycelia and the promycelial spores are larger than those produced in water. No fusion of spores or buckling of promycelia occur. The most remarkable fact is that the promycelial spores mul- tiply themselves by budding, very much after the manner in which yeast-cells multiply themselves in saccharine fluids (Plate VII. Figs. 8-10). Brefeld has kept them thus repro- ducing themselves for more than a year, by replacing the nutrient fluid as it became exhausted. The promycelial spores continue to reproduce themselves by budding, as long as the nutrient fluid remains unexhausted ; when this occurs they cease to bud, and fuse in various ways, as is seen when germination takes place in water. The yeast- spores of this species produced in his nutrient fluid are rather larger than the primary spores produced from the promycelium in water; they are, however, of the same shape, and measure from 9 to 30m in length, and from 3 to 5in breadth.* Kept moist, they retain their vitality for about two months, but if allowed to dry, none germi- nated after the sixth week. The teleutospores, on the contrary, if kept dry, retain their germinative faculties for a very long period. Brefeld found that at the end of two years they germinated as freely as when fresh; other * Brefeld, loc. cz#., p. 13. 78 British Uredinee and Ustilaginec. observers have found that they even germinate after seven and a half years.* U. Cardui, F. v. W.—Kiihn found that teleutospores in water produced promycelia and small ovate promycelial spores.f In nahrldsung yeast-colonies are produced very abundantly, being from 5 to 8u long, and from 3 to 5 wide; they very much resemble the cells of beer-yeast, but of course they do not bud in saccharine media. U. flosculorum, D.C. (U. intermedia, Schroter).— The germination has been studied by Schréter,§ who found the spores, when placed in water, germinated very quickly, for in twelve hours all had produced promycelia. These attained a length of from 16 to 20u, and a width of from 5 to 64; they generally become triseptate, and bear sporidia both laterally and terminally. These are shortly ovate in form, and measure 6u in length, and from 4 to 5 in width. In the course of thirty-six hours the sporidia tend to become spherical. The promycelia have a great tendency to fall away from the teleutospores and . subsequently to produce promycelial spores, so that they become sporophores (/ruchttréger). The budding in nahrlésung Brefeld || found to be so profuse as to be quite phenomenal. The yeast-spores are from 4 to Su long, and 4u wide. U. violaca, Pers. (U. antherarum, Fries)—The teleuto- spores of this species are even better suited for observa- tion than those of the one just described, inasmuch as, being rather larger, the promycelia are proportionately bigger, and the septation and spore-formation more easily observed. The teleutospores germinate after a very short * Von Liebenberg, ‘‘ Oesterr landw. Wochenblatt” (1879), Nos. 43, 44. t Kiihn in Rabenhorst’s ‘‘ Fungi Europzi,” No, 1798, t Brefeld, Zoc. ci¢., pp. 86-88, t. vi. figs. 1-16. § Schroter, “‘ Beitrage zur Biol.,” bd. ii. heft. iii, pp. 352, 353. || Brefeld, /oc. cz?., pp. 89, 90, t. vi. figs. 17-27. Germination of Teleutospores of Ustilaginee. 79 immersion in water. The process has been described and figured by Tulasne * and Von Waldheim.t Brefeld ¢ found the production of the yeast-cell colonies was very prolific in nahrlésung, where they multiplied themselves through endless generations, whereas in water no budding took place after the second day. These yeast-spores are ovate, but somewhat elongated; from 5 to 7u long, and from 3 to 44 wide. The germinative faculty lasts for about six weeks. The spores in water or in exhausted nahrlésung frequently unite in the same manner as those of U. segetum. When this fungus attacks the anthers of Lychuzs diurna, a plant which is usually unisexual, the styles which would normally be short, acute, and erect, become long and recurved, as they are in the female flower. This has been pointed out by M. Cornu, as well as by other observers. U. maydis—The germination of these spores was observed by Kiihn § and Wolff, and does not materially differ from the above: the yeast-spores,? however, are elongated and fusiform, being from 10 to 36 long, and from 3 to 5u wide. U. scabiose (Sow.) (U. flosculorum).—The spores germi- nate very freely and very soon in water. According to Schroter ** the promycelia are three or four-partite ; about from 20 to 224 long, and du wide; the promycelial spores are about 4u long. In nahrlosung Brefeld tt found the yeast-spores to be from 4 to 8u long, and from 1°5 to 2 wide. They were produced continuously and * Tulasne, ‘‘ 17 Mémoire,” t. iv. fig. 18. + F. von Waldheim, Joc. czt., t. xii. { Brefeld, Joc. ci#., pp. 36-54, t. i. figs. 1-27. § Kiihn, “‘ Krankh d. Kulturgew.” p. 260, t. iii. figs. 22, 23. || Wolff, ‘Brand des Getreides,” p. 11 t. i. fig. c. q{ Brefeld, oc, czt., pp. 67-75, t. iv. figs. 1-17. ** Schréter, “ Cohn Beitrage,” vol. ii. 1877 tt Brefeld, oc. cit., pp. 78-81, t. v. figs. 1-6. 80 British Uredinee and Ustilaginee. abundantly. As the teleutospores are of pretty large size, their germination is easily observed. I have found them germinate much more freely in summer than in late autumn (Plate VII. Fig. 18). U. tragopogi (Pers.) (U. veceptaculorum, Fries).—The germination was observed by Tulasne,* and by Von Wald- heim,t to consist in the protrusion of a promycelium through a very marked germ-pore in the epispore. It becomes three or four-septate, and pro- duces subcylindrical promycelial spores, which are rounded, especially at their distal ends. These often grow nearly parallel to the promycelium, as figured by Von Waldheim; sometimes they : are produced terminally upon branches pore taetinge wast given off from the segments of the pro- nating ,and promyceiaispores mycelium. Conjugation often takes place between them after they have fallen off. In n&hrlésung Brefeld t{ found them to be abun- dantly reproduced by budding, but they were larger, mea- suring from 5 to 20u in length, and from 5 to 7u in breadth. U. kithneana, Wolff.—The promycelium is three to four- septate, the lowermost compartment being the longest and empty. It produces numerous promycelial spores in whorls at each septum (Plate VII. Fig. 17). In nahrlo- sung § they budded very profusely, forming very character- istic small yeast-spores, from 3 to 54 long, and from 3 to su wide. Wolff|| also investigated the germination in water. U. hypodytes (Schlecht.).—The germination of this plant * Tulasne, ‘‘2° Mémoire,” pp. 159-160, t. xii. figs. 34-40. t F. von Waldheim, loc. cét., t. xi. figs. 27-37. t Brefeld, oc, czt., pp. 81, 82, t. v. figs. 7-11. § Brefeld, Joc. cit., pp. 83-88, t. v. figs. 12-20, || Wolff, Bot, Zeitung (1874), pp. 814, 815. Germination of Teleutospores of Ustilaginee. 81 is markedly different from that of any of the former species. Winter * figures an elongated promycelium with a pro- mycelial spore, borne laterally upon a long pedicel. He found that the spores germinate freely in water, and that the endospore sends out a process—the promycelium—from the interior of the spore, which grows to 30 or 50m in length, but is only about 3u wide. It becomes septate, and gives off short lateral branches, which become spores. They are slightly clavate, and from 6 to 7 long.t Brefeldt found that in nahrlésung they only produced mycelial hyphz without spores. Although I have tried many times, I have never succeeded in getting the teleuto- spores of this species to germinate. CU, longissima (Sow.).—When the spores of this species are placed in water they very soon begin to germinate. The process, as carried on in this species, differs very materially from that which obtains with the previously mentioned species. This, as was first pointed out by Von Waldheim,§ consists in the protrusion of a very narrow straight tube through a small opening in the epispore. This acquires a length of about 10 or 12, when it becomes divided below by a septum into two un- equal parts (Plate VII. Fig. 14), the upper of which is about 6 or 8, and the lower 3 or 4u long. The lower portion is, moreover, narrower than the upper, and is the true promycelium, the upper being the pro- mycelial spore. The promycelial spore soon falls off, and the promycelium produces, in about an hour, Von Waldheim says (but I have personally made no observation as to time), a second promycelial spore, which in like manner falls off, and is followed by a third (Plate VII. Figs. 15, 16). * Winter in Rabenhorst, ‘‘ Kryptogam, Flora,” vol. i. p. 81, fig. 4. + ‘Flora” (1876), Nos. 10, II. t Brefeld, Zoc. c¢¢., p. 103. § F. von Waldheim, Zoc, cit., t. v. figs. 42-46. G 82 British Uvredinea and Ustilaginea. According to Brefeld,* it is seldom that more than three promycelial spores are produced from one teleutospore before it becomes emptied of protoplasm and exhausted. Be this as it may, if a few teleutospores be placed in a drop of water on a glass slide, and examined at intervals for two or three days, one can see with the naked eye that there has fallen to the bottom of the drop a whitish cloud. Upon microscopic observation, this cloud is found to con- sist of an immense assemblage of promycelial spores. They are cylindrical bodies, with somewhat attenuated’ extremities, and often measure from 8 to 10m in length, and from 1°§ to 2u in breadth. Hence it appears that they have increased in size since they fell off the pro- mycelium. After a time this increase in size ceases, but not before some few odd ones here and there have attained a length of from 20 to 30pm. Brefeld found, by the culture of isolated promycelial spores in nahrldsung, that after these bodies had fallen away from the teleutospore which produced them they not only multiplied themselves, but increased enormously in length and thickness. This they did with great rapidity. They more resembled hyphe than promycelial spores, and each soon became more or less septate. They multiplied by giving off a small bud-like projection laterally, and at a short distance from one or other of their extremities. This bud rapidly grew into a second spore, but before it attained the dimensions of its parent the latter had given off a similar bud towards its opposite extremity; and so the process of multiplication goes on until the nahrlésung is exhausted. When this takes place, instead of multiplying in the manner above described, the promycelial spores give off hyphz of considerable length, which become septate at intervals from below upwards, and the protoplasm is passed * Brefeld, /oc, cé¢., pp. 104-116, t. viii., ix., figs. 1-16. Germination of Teleutospores of Ustilaginee. 83 along to its growing end. Frequently two promycelial spores become united by a transverse bridge (conjugation) ; one of them then gives off a germ-tube, as in the case of single promycelial spores, only it is longer. U. grandis——In 1876, Kiihn* investigated the germina- tion of this species. He found that the promycelia, which are about from 50 to 6ou long, and from 5 to 84 wide, had a great tendency to fall off from the teleutospores before they produced promycelial spores, although this was not by any means always the case. Brefeldt found in nahrlé- sung that the promycelia were not only larger, but produced more and also larger promycelial spores. These sporidia not only reproduced themselves, but also grew into sporo- phores (Fruchitréger), which were indistinguishable from the ‘original promycelia, inasmuch as they were cylindrical septate tubes, which in their turn budded off spores. Thus colonies of yeast-cells do not occur in the life-cycle of U. grandis any more than they do in that of U. longéssima. With U. grandis the promycelium produces spores which grow out into sporophores, and they in their turn produce spores again. These sporophores are multicellular. U. bromivora, Tul.{—In water each spore produces a small promycelium through a minute opening in the epispore, very much after the manner of U. longissima. This bears terminally a spore which soon falls away. Between the fallen-off spores conjugations are frequently to be seen ; sometimes they become uniseptate and buckle- jointed. In nahrlésung the typical germination takes place in the production of single promycelial spores from a short promycelium. These spores increase in size and become uniseptate, or, as Brefeld terms them,§ bicellular sporo- * Kiihn in Rabenhorst ‘‘ Fungi Europei,” cent. xxiii. No. 2299, fig. + Brefeld, Zoc. cét., pp. 116~123, t. ix. figs. 17-26. t Cf Kiihn, ‘‘ Vortrages iiber Getreidebrand.” 1874. § Brefeld, Joc. cé#., pp. 123-129, t. x. figs. 2-8. 84 British Uredinee and Ustilaginee. phores. There is no observable difference between the sporophores produced from the teleutospore and those pro- duced from a promycelial spore. These developments go on until the nahrlésung becomes exhausted, when the pro- mycelial spores and sporophores alike give off mycelium- like tubes and fuse in various ways. In other .words, U. bromivora is characterized by its promycelial spores growing into bicellular sporophores, which sprout directly into new promycelial spores. True yeast-cell colonies do not occur. In the teleutospores of this species which Mr. Soppitt sent me I found that promycelia were freely produced in water, and that they developed elongate, elliptical promy- celial spores (10O—-I12 X 3—4y), which tended to become vacuolate after they had fallen off, and afterwards emitted pointed germ-tubes. I found that teleutospores gathered in June germinated freely in September. U. olivacea (D.C.)—The teleutospores germinate,* after a few hours in water, very much like those of U. longzssima. The promycelium is, however, so curtailed as practically not to exist, and the promycelial spores are really produced at once out of the teleutospores without any promycelium. These promycelial spores are variable in size; each is sub- fusiform, and measures from 5 to 20u in length, and from 2 to 3u in breadth. In nahrlésung they form yeast- colonies. U. major, Schréter—I gathered some specimens of this fungus near Paris in the middle of October, 1887. The spores germinated very readily when placed in water. In twenty-four hours they had developed cylindrico-fusiform promycelia (10—12 X 2), which fell off from the teleuto- spores (Plate VII. Figs. 19g—22) much after the manner of U. longissima. In forty hours these had attained a * Brefeld, Joc. cit, pp. 129-133, t. x. figs. 9-26, Germination of Teleutospores of Ustilaginee. 85 size of I5—18 X 3°5—4'5u. They formed an opalescent cloud at the bottom of the culture-drop. Each promy- celium was cylindrical in form, vacuolate, and eventually became triseptate, and produced promycelial spores both laterally and also at either end (Plate VII. Figs. 22, 24, 25). The latter were elliptical in form, and measured from 3 to 4u in length by 2u in width. Many of the promycelia were observed to be slightly curved. U. utriculosa (Nees).—The teleutospores of this species are by no means easy to germinate, and although I have made a great many attempts to do so, I have always failed. Schréter* states that they emit a cylindrical promy- celium, which becomes triseptate, and produces elliptical promycelial spores in pairs, which conjugate in couples, Sphacelotheca.—The spores do not germinate at all readily, and I have been unsuccessful in observing the process. Schréter ft states that the teleutospores emit a cylindrical promycelium, which becomes triseptate, an 1 bears elliptical promycelial spores laterally, and that these conjugate in pairs at their bases.f. Sorosporium.—W oronin § succeeded in getting the spores of S. saponarig, Rud. on Lychnis dioica and Saponaria officinalis to emit a germ-tube, but no spore-formation was * Schroter, Cohn’s ‘* Krypt. Flora von Schlesien,” vol. iii. Pp. 273. t Loc. cit., p. 275. } The two following species of Ustilago can in no sense be regarded as being British, yet they occur in this country sufficiently commonly to render them objects of interest. U. ficuum, Richdt., is often met with on the cheaper kinds of figs, known in the trade as ‘natural figs.” The spores are formed in the interior of the fruit, and are black or dark violet, smooth, globose, from 6 to 8u across. I found no germination took place below 10° C., but between 10° and 13° C., when placed in water, they emitted promycelia 20—150 x 4—5u, but I was unable to observe any further development. U. phenicis, Corda, is a. closely allied species, which is frequently to be met with on cheap dates. The spores are globose, smooth, dark violet, from 4 to 5 in diameter. § Woronin, /oc. c#t., pp. 18, 19, t. lii. figs. 13-18. 86 British Uredinee and Ustilaginee. observed. The spores collected in June germinated in December, after being placed in water from three to five days. Thecaphora—The germination of 7. hyalina has been investigated by Woronin.* He found that spores collected in August germinated in October and November, after ~fourteen to eighteen days’ maceration in water, but older spores did not germinate at all. Each spore is provided with a germ-pore in the epispore, which is pale in colour and free from any of the verrucosities which occur upon the other parts of the epispore. Every germ-tube becomes filled with protoplasm, and generally contains four nuclei. It becomes septate, and each compartment contains one of the nuclei. From each segment of the promycclium narrower lateral branches are given off. Those from the upper compartments tend to grow downwards, while those from the lower, on the contrary, grow upwards. If one of the upper branches comes in contact with one of the lower, they unite at their ends and form a bow-like conju- gation. From this a long germ-tube is given off, into the end of which the protoplasm is passed. No spore-formation was observed. I have made many attempts, but have always been unsuccessful in getting the teleutospores of this species to germinate. Brefeld,f in an allied species (7. /athyrz, Kiihn), found promycelia, at the end of which spherical promycelial spores were formed. These promycelial spores in nahrlésung germinated and produced a mycelial mass, which in turn also produced spores upon those of its branches which came in contact with the air. Tilletia—The germination of Z7lletta triticd has been * Woronin, /oc. cét., pp. 21, 22, t. iii. figs. 19-28. + Brefeld, /oc. cit., pp. 134-138, t. xi. figs. 2-12. Germination of Teleutospores of Ustilaginee. 87 known since 1807, when Prevost* figured not only the promycelium, but the primary and secondary promycelial spores. Mr, Berkeley,f in 1847, discovered the conjugation of the primary spores, which was again more fully investi- gated by Tulasne.{ Kiihn§ gives a full account of the process. Since then nothing has been added to our know- ledge of the subject, till Wolff || showed the method by which the germ-tube enters the host-plant, and Brefeld investigated the further development of the spores in nahrlésung. The spores do not germinate until they have been placed in water for some considerable time, not before forty-eight or fifty hours ; but often I have found them to take a much longer period. They retain their germinative power for two or three years, and one author says as long as eight and a half years.** The process differs materially from that previously described in the other genera. The promycelial tube is emitted from a small germ-pore, but very soon, as the tube increases in diameter, it causes the epispore to split. Its length varies according to circum- stances, its diameter being about 8u. If it be given out from a spore under water, at the bottom of the culture-drop, it grows upwards until its apex reaches the air. As soon as the promycelium has reached the air several tubercula- tions appear upon its‘ssummit. The protoplasmic contents of the spore are passed along the promycelium to its extremity. If the promycelium happen to be a very long one, then numerous septa occur from below upwards ; but, * Prevost, ‘* Mémoire sur la cause immédiate de la Carie.” 1807. + Berkeley, ‘‘ Propag. of Bunt,” Zrans. Koy. Hort. Soc. (1847), *vol. ii. p. 113. t Tulasne, “1° Mém. sur les Ured. et les Ustilag.” 1854. § Kiihn, “‘ Krank. der Kulturgew.” 1859. | Wolff, ‘Der Brand des Getreides.” 1874. { Brefeld, Joc. cit., pp. 146-163, t. xii., xiii. figs. 25- oe ** Liebenburg, Joc. cit. 83 British Uredinee and Ustilaginee. however short it may be, there is always one septum developed near its upper end. The tubercles above men- tioned increase rapidly in length, and become the primary spores (Plate VI. Fig. 7). They are filiform bodies, curved in various ways, and measure from 80 to 100 in length; in number they vary from four to twelve or more, according to the size of the spores from which they are developed. When all the protoplasm from the promycelium has been absorbed into these primary spores, they become cut off from it by septa at their attached ends. If the promycelium be so situated that it cannot reach the air, no primary spores are produced. Shortly after their maturity these primary spores conjugate (Plate VI. Fig. 8), or become united by transverse bridges, usually in pairs. The primary spore is possibly a Fig. 7.—Germinating teleutospore of Tilletia tritici, producing a cluster of wind-carried spore, but, as in primary promycelial spores, A conjugated pair of promycelial spores producing two artificial cultures they are not secondary spores—an unusual circumstance é (ss). A secondary promycelial spore which E€Xposed to this force, they fre- has produced a tertiary (s¢). (Tulasne.) . i ‘ 3 quently germinate zz sztu. This they do by a repetition of the oft-described process of protoplasmic migration, with septation of the emptied parts ; the protrusion of a bud-like process into which the protoplasm is emptied, and which becomes a secondary spore. The end of the promycelium, after the primary spores’ have fallen off, remains tuberculated, showing the points of their attachment. The conjugation of two primary spores cannot be considered a sexual act, in- asmuch as the single spores, which have not been sub- jected to it in any way, germinate as freely, and produce Germination of Teleutospores of Ustilaginee 89 secondary spores as effectually, as when two or three have become connected, the only difference being that with fused or conjugated spores larger germ-tubes and larger secondary spores are produced. The secondary spores (Plate VI. Figs. 9, 10) are at first cylindrical, but they soon become reniform, and at length, by the attenuation of their ends, more or less crescent-shaped. They may be pro- duced from any part of the primary spores, even from the connecting bridge, and at almost any distance; very seldom is more than one produced from each fused pair of primary spores. Three or more primary spores have been seen connected,* and double fusion between two has been also observed.f Not only do the primary produce second- ary spores, but they may emit a pointed germ-tube, for direct penetration of the host-plant. In like manner, between the secondary spores all sorts of connections and conjugations or fusions occur, with the same emptying of the contents from. one spore into the other (Figs. 11, 12). The secondary spores are, however, essentially the spores the germ-tubes of which enter the host-plant. In nahrldsung, according to Brefeld, the primary and aa are larger, and are produced in greater abundance. No conjugations or fusions occur, except when it becomes exhausted. The primary spores, how- ever, comport themselves very differently in nahrlosung ; they send out germ-tubes which are narrower than those given off by them: in water. These tubes are unseptate, but branch and inosculate with each other so as to form a mycelium (Plate VI. Fig. 14), which, as it grows out into the air; forms a white floccose mass. For five or six days no secondary spores are found in well-nourished mycelia, whereas spore-formation occurs soon in badly ‘nourished ones. The spores are produced only on those * Brefeld, /oc. cit., t. xiii. fig. 39. t Ibid., fig. 38. 90 Lritish Uredinee and Ustilaginee. branches of the mycelium which are given off into tl air. Upon these numerous short, lateral branches are giv: off, which swell up at the ends and become crescent spores (Fig. 15). No conjugation takes place betwe« these bodies. The branches of the mycelium which a given off in the fluid do not produce spores, but gro outwards until at length they reach the air, when the produce terminal spores or they remain sterile. In tl latter case the hyphe are empty and septate. Brefe has further observed that by long-continued culture tt hyphe, under certain circun stances, become nodose, and a parently develop certain globo: bodies which closely resemble tk original teleutospore.* Entyloma—tThe germinatio of Entyloma, though similar t that of Tilletia, is far less con plex. E. microsporum.—De Bary found that if the spores wer wholly immersed in water, they in the course of twenty-four hour. would give out a germ-tube fror Fig. 8.—Entyloma microsporum, four to ten times the length c a, ‘Teleutospore germinating (the promycelial spores have conjugated the spore, At its rounded sum at their upper ends); 4, two secondary spores produced from the conjugated mit this promycelium gave o pairs of primary promycelial spores ; ¢, teleutospore of £. calendule, which Six or seven branches each c a preeveas oes Pee eel Spares, m our Of whic. lave conjugate elow. 1 1 . eee jug: which was dilated upwards; an when they attained a lengt measuring 30 or 40u, each became cut off by a base septum. They conjugate in pairs, by a transverse bridg: * Brefeld, doc. cét., t, xiii. figs. 46-52, t De Bary, Bot. Zeitung (1874), pp. 81-92, 97-108, t. ii, Germination of Teleutospores of Ustilaginee. 91 either at their lower or upper ends; after which one of the branches grows onwards in its original direction, and at its end develops a long, fusiform, secondary spore (Fig. 8). This falls off, and in its turn germinates by the pro- trusion of a long, very narrow germ-tube. If the number of the primary branches be odd, the odd one does not con- jugate, but grows more slowly, and ultimately shows no further change. Various deviations from the typical ger- mination take place; thus the promycelium itself can grow into a germ-tube, or the primary whorl of branches may send a-branch downwards. In £. calendule the process is similar. E. vanunculii—In this species Brefeld* found that conidia were produced upon the host-plant, and that in the interior of the leaves a richly developed mycelium existed, sending up conidiophores through the stomata. The fresh conidia in naéhrldsung produced a mycelium less extended, but little different from that produced by the spores of Tilletia under similar conditions. This very soon becomes covered with conidia, which in their turn produce mycelia and conidia again. Marshall Wardf found that the conidia which are produced in spring are clavate or elongate-oval in form ; that they germinate in from twenty- four to thirty hours by emitting a delicate germ-tube from both extremities, one of which grows, the other becoming empty of protoplasm and septate. The growing germ-tube generally becomes swollen into a secondary conidium when the culture is madé in water. From this secondary coni- dium a branched germ-tube is emitted. If the conidia germinate on a leaf, the formation of the secondary conidia is rare. The germ-tubes enter the stomata, and * Brefeld, oc. c2t., pp. 163, 164. + Marshall Ward, PAZ. Trans. of the Roy. Soc., vol, 178 (1887), B., pp. 173-185, plates 10-13. 92 British Uredinee and Ustilaginee. the conidia reproduce themselves in from fifteen to.twen days. The teleutospores. are developed in spherical dil tations, in the continuity of the mycelial tubes, rare at the ends of branches. Conjugation between: conid sometimes takes place. The conidiophores emerge eith through the stomata or between the epidermal cel: The conidia germinate more rapidly and throw out larg: germ-tubes when this process takes place on the livir leaves of the host-plant than. when it does so in wate This may be in part due to: the more abundant supply « oxygen which they would receive in the former situation. £. canescens —Schroter * finds the spores germinate : soon as they are ripe by protruding a germ-tube, from 2 to 30u long, and 4 wide, on the end of which a tuft « cylindrico-fusiform spores are produced. They measui from 25 to 40y in length, and from 2's to 3 in thickness. Doassansia.—The germination of this genus is ident cal with that of its ally, Entyloma, consisting in the prc trusion of a promycelium of limited growth, which develo apically a tuft of promycelial spores.t In D. alésmatzs th promycelial spores are long and cylindrical, and they ar produced in great numbers (Plate VIII. Fig. 5). The process is also similar in D. sagiltart@, as observe by Fisch,} who observed conjugation to take place betwee the fallen-off promycelial spores. He found that the germ tubes of these spores entered the sides of the cells of th host-plant, having insinuated themselves between th epidermal cells. The teleutospores of D. alismatis germ: nated as soon as they were ripe, but those of D. sagittart did not do so until the ensuing spring. * Schroter, “‘ Cohn Beitriage,” vol. ii, (1877), p. 372. t Commu, Ann. des Scienc. Nat. Bot., 6° sér., tome xv. p. 281. } Fisch, ‘‘ Entwickelungsgeschichte von Doassansia Sagittariae,” Berich der deutschen botan. Gesellschaft,” September, 1882, bd. ii. t. x. Germination of Teleutospores of Ustilagince. 93 Urocystis—The germination of Urocystis was first observed by Kiihn in U.,-occulta,* and is also described by Wolff.t U. colchicd was studied by Winter ;t U. ane- mones, by Von Waldheim ;§ U. viola, by Prillieux ; || and U. primulicola, by Pirotta.1 The process consists in the protrusion of a promycelium from the inner coloured ‘spores (the paler peripheral pseudo- spores do not germinate), into which the protoplasm passes to the upper end, where it gives rise to a variable number of primary spores. If the promycelium be produced under water, no spore-formation occurs until its point comes into the air. The primary spores fall off, and occasionally con- jugate in various ways, but not so constantly as in Tilletia ; they also frequently germinate whilst attached to the pro- mycelium. U. gcculta—The central, dark-coloured spores (as first described by Kiihn **) emit a promycelium, at the apex of which from two to six primary spores are borne. These sometimes conjugate by a transverse bridge at their upper ends, and often germinate—as Wolff has more recently shown—while still attached to the promycelium, from their -lower, ends, sending out a long, narrow germ-tube, which receives the protoplasm from the interior of the spore, so that the upper part of the spore is first emptied of its contents.fTt U. fischeri—The spores of this species, which Mr. Soppitt was kind enough to send me, germinated only after a con- * Kiihn, doc. cit., pp. 78-80, t. ii. figs. 13-34. + Wolff, ‘‘ Der Brand des Getreides,” pp. 16, 17, t. ii. figs. I-10. t Winter, ‘“ Ustilagineen Flora” (1876), Nos. 10, 11. § F. von. Waldheim, Joc. cét., t. vi. figs. 38-43. || Prillieux, Azz. des Scienc. Nat. Bot., 6° sér., tome x, (1880), p. 49, t. i. { Pirotta, ‘*R. Nuovo Giornale Bot. Ital.,” vol. xiii, 12 Luglio (1881), No. 3. ** Kiihn, loc. cét., t. ii, fig. 202. tt Wolff, Zoc. cit., t. ii. B. figs. 7, 8. 94 British Uredinee and Ustilaginee. siderable period of soaking in water. The germ-tube we larger than in any of the other species of Urocystis tt germination of which I have watched; the promycelii spores were also not only larger, but more numerous. counted as many as eight on some of the promycelia (Pla' VII. Figs. 34, 35). U. anemones——Von Waldheim * points out that unle: the promycelium grows in the air no spore-formatic takes place. At its end it divides into three or fou branches, which become spores. They are elongated, ova and generally wider at their upper end. In length, the measure from 10 to 14"; and in breadth, from 3 to 3°5, They become vacuolate, and enlarge in size till they ofte measure 22 by 4u. After several hours’ (forty-eight an more) immersion in water, in November and December, found the teleutospores germinated. The promycelial spore were of the same size and form as described by Von Walc heim (Plate VII. Fig. 31); I also observed they becam vacuolate when old (Figs. 32, 33). The promycelial spore were applied to the foliage of Ranunculus repens, in tw experimental cultures, on December 12, 1884. No chang was observed in the plants until February, when it wa noted that they showed signs of the formation of spore beds. On February 11 in ome experiment, and on th 22nd in the second, spores were developed. This is on of the few species in which mycelium is localized, and th infection of the host-plant occurs at the same place < which the teleutospores are subsequently formed. U. viole—The spore-balls generally produce only on promycelium, which bears at the end a cluster of five c six fusiform spores, If the promycelium remain shor spores are produced ; but if it grow to any great lengtl either no spores at all are formed or only small ones. C * Waldheim, /oc. cit. Germination of Teleutospores of Ustilaginee. 95 the six spores generally only three germinate, and pro- duce at their distal extremities secondary spores similar in size and shape to themselves.* U. primulicola, Magnus.—Pirotta { found that the fresh ripe spores germinated in water in about ten hours, by emitting a short cylindrical promycelium, which at its extremity gave off three or four branches that became spores, measuring from 9 to 18 in length, and from 4 to gu in width. These, while still attached, produced secondary spores from their ends. The secondary spores germinated by the protrusion of a germ-tube (about 3u wide, and from 10 to 20 times as long as the spore), into which the protoplasm migrated. Lateral conjugation was occasionally observed. This species occurred in 1884, in Rev. C. Wolley Dod’s garden, on P. farinosa. In August of that year I received some specimens from Mr. Dod. The spores germinated readily in water, and emitted short promycelia, which bore a cluster of promycelial spores as figured by Pirotta. I found that no spore-formation took place unless the end of the promycelium grew in the air. If a spore germinated at the bottom of a drop of water, the promycelium grew upwards through the water until it reached the air. In these cases the lower part of the promycelium became emptied of its protoplasm and septate, just as one sees in Tilletia (Plate VII. Fig. 26, 27). The promycelial spores varied from 12 to 20u in length, and were 4 or 5 in width (Figs. 28, 29, 30). After keeping the promycelial spores in nahrlésung for two hundred and sixty-four hours, no further spore-formation was observed; but they became septate and nucleate (Fig. 29). Melanotenium—The mycelium is principally inter- * Prillieux, Zoc. cit, + Pirotta, ‘*Nuovo Giornale Bot. Ital.,” vol. xii. (1881), pp. 235-239, t. vi. 96 British Uredinee and Ustilaginee. cellular, and pervades all parts of the affected plant especially the cortical parts of the stem, and also to sor extent the pith. It also occurs in the upper part of tk root-stock. It is hyaline, about 4°5u wide, richly branche: and septate. Its contents are colourless and vacuolate: The spores are formed inside. the mycelial hyphe, whe: it becomes coarsely granular, much after the manner « Entyloma.* Woronint finds that the mycelium is abu dantly provided with very marked haustoria, which ente the cells. These botryform prolongations enter and occup a third or a half of their interior. Germination takes plac in autumn. Specimens gathered in June germinated i October and November. The epispore splits, and th endospore grows out as a blunt cylindrical promyceliun At its extremity it emits a cluster of from four to seve apical branches. The outgrowth of the endospore is < first often in the form of two equal branches, one which develops into the promycelium, while the othe ceases to grow and has become emptied of its protoplasn which passes into the developed branch. Towards th upper half of the promycelium a septum appears, cuttin off the protoplasm above from the empty tube below; bt true spore-formation was not observed. Tubercinia—The germination of the spores of 7. trée talis has been worked out by Woronin.t Teleutospore collected at the end of September and the beginning October were found often to have already germinated upo the plant. Placed in a damp atmosphere, they germinate freely after the manner of Tilletia, each spore producing promycelium surmounted by a cluster of spores. All th teleutospores of one spore-ball do not germinate at th * De Bary, loc. cz. t Woronin, /oc. cit., pp. 27, 28, t. iv. figs. 27-35. $ Woronin, ‘‘ De Bary und Woronin Beitrage,” 5 reihe (1882), pp. 4-1 t. i, ii, iii, figs, I-12. Germination of Teleutospores of Ustilaginee. 97 same time. The promycelium emerges through a small ~ round opening in the epispore, its length corresponding to the size of the spore. On the upper, free, blunt end of the promycelium from four to eight protuberances appear, which elongate themselves into branches and become the cylindrico-fusiform promycelial spores. After all the pro- toplasm from the interior of the teleutospores has been passed into the upper end of the promycelium and into the developing promycelial spores, a septum is formed close to its upper end. If the promycelium happen to be a very Jong one, two or more septa occur. The promycelial spores, while still attached to the promycelium, become united in pairs by a bridge-like connection. This conjuga- tion takes place at the bases of the promycelial spores, and but rarely at their summits. One of the conjugated spores then buds out a secondary spore, which in its turn sometimes produces a tertiary ; sometimes all these may be observed in achain. If there be an odd spore on the promycelium which has not conjugated, it does not bud. Conzdia.*—These are produced from a mycelial mesh that exists for the most part just beneath the epidermal structures, and is provided with very numerous botry- form haustorie (Plate VI. Fig. 1). The conidiophores emerge through the stomata, or between the epidermal cells (Plate VIII. Fig. 1). The conidia are borne almost horizontally; they are from 11 to 154 long, and consist of subpyriform cells attached by their larger end. A thin hyaline membrane encloses the granular protoplasm, in which a small vacuole may be observed. When placed in a damp atmosphere the vacuole enlarges and a germ-tube is produced, generally from the larger end of the conidium ; into this germ-tube the protoplasmic contents of the coni- dium are received and passed onwards as it elongates * Woronin, oc. cit. H 98 British Uvedinee and Ustilaginee. (Plate VIII. Fig. 2). If the conidium germinate upon a leaf, the germ-tube squeezes its point between the two epidermal .cells (Fig. 3), and soon produces in the leaf a mycelium with haustoria. In from twelve to twenty days after infection this mycelium produces the black teleutospores, but not the conidia. The life-history of this species is peculiar: the teleuto- spores germinating in autumn produce promycelial spores, which, entering the young subterranean shoots of the host- plant, develop a mycelium, which remains quiescent during the winter, and in the spring produces, first the conidia on the leaves, and afterwards teleutospores mostly in the stem. The entrance of the germ-tube in this species is (as already stated above) between the epidermal cells. It grows downwards in the partition wall, splitting it into two lamine, and so makes its way through the epidermis. ( 99 ) CHAPTER XI. INFECTION OF THE HOST-PLANTS BY THE USTILAGINE#. THE manner in which the Ustilaginez gain admission into their respective host-plants has been studied very carefully by many botanists, but is not yet fully understood. With those species which affect the flowering parts of annual graminaceous plants, such as 77/etia tviticd and U. segetum on wheat, it is noteworthy that not only are all the blossoms or fruits upon an ear affected, but also all the ears which arise from one plant. It is very exceptional ever to find one sound ear upon a plant of which the others are diseased ; in like manner, it is very unusual to find a sound kernel upon an ear in which the other kernels are affected. Coupled with the fact that in diseased plants the mycelium of the fungus can be found in all parts of the axis, it is obvious that the parasite gained admission into the plant at an early stage of its growth. Kiihn * specially investigated this point with I. triticé, and found, in very young wheat seedlings, that the mycelium was present in them. Hoffmannt came to the conclusion that the spores entered between the split in the young sheath and the rootlet. He also figures the spores . forming a mycelium which enters the stomata of the young * Kiihn, oc. cit., pp. 48, 49. t Hoffman, ‘ Flugbrand,” pp. 202-206, t. xiv. figs. 14-18. 100 British Uvedinee and Ustilaginee. plant, but this is probably incorrect. To Wolff,* however, we owe the first accurate explanation of this process. He investigated it with U. segetum and maydis, Urocystis occulta and 7. tritic?. The outcome of his observations is that the germ-tube of the promycelial spores of the species is capable of piercing the embryonic plant at any time before the primary enveloping sheath of the young plant is ruptured. The germ-tubes of 7. ¢rztzc? squeeze their points through the epidermal cells of the young plant, at first piercing through the outer epidermis of the primary sheath ; they then grow through the cells of the sheath itself, then through the inner epidermal cells of the sheath, across the interspace to the outer epidermal cells of the embryo, and so into the embryo itself (Plate VI. Fig. 2). With certain species the entering germ-tube acquires for itself an invest- ing sheath from the cells through which it passes (Plate VI. Fig. 3)—a sort of invagination of the outer wall of the outer epidermal cell, which is continued over the young mycelium as it grows through one cell after another. With Uvrocystds occulta the investing sheath exists only where the mycelium passes through the first epidermal cell. Kiihnf subse- quently repeated and confirmed Wolff's observations as far as they went, but he also found that the germ-tubes could enter, not only through the primary sheath-leaf, but also into the true root-node at the base of the sheath, and, in point of fact, into almost any part of the embryo. While it has long been known that, by merely dusting wheat with the teleutospores of 7. ¢retzcz and planting it, it became affected with bunt, yet with U. segetum such dusting rarely, if ever, succeeds in producing the disease. Hoffmann was able to produce only a few smutted plants in many hundred * Wolf, ‘‘ Roggenstengelbrand,” ot. Zeitung (1873), t. viii; “ Der Brand des Getreides” (1874), pp. 18-24, t. iii., iv. + Kiihn, Bot, Zeitung (1874), pp. 121-124; Fahlingsladw, Zeitunz (1879) p. 84. L[nfection of Host-Plants by Ustilaginee. 101 experiments in which he applied the teleutospores to the young plant. F. von Waldheim was equally unsuccessful. Kiihn asserts that he found that, if too many germ-tubes entered an embryo plant, they developed into spore-forming ‘hyphee and formed a “ brand-knot” in the sheath-node, and killed the young plant. All the experiments which I have conducted with a view of infecting the young plants of wheat, barley, and oats with J. segetum have uniformly failed. I have attempted the infection in various ways: dusted the spores on the dry grain and planted it; soaked the grain in water, and then dusted it with the dry spores ; planted the grain in flower-pots, and dusted the spores thickly on the surface of the soil; the grain allowed to germinate, and applied the dry spores to the embryos just as they emerged from the seed-corn ; placed the teleuto- spores of U. segetum in water for twelve hours, and, when an abundant development of promycelial spores had taken place, applied the spore-charged water to the emerging embryos ; germinated the teleutospores of U. segetum in nahrlésung and dipped the young embryos in it; watered the grain, after it was planted, and before it came up, with nahrlésung, containing spores, but the result was uniform failure. Wolff has stated that the infection will not be successful if the infected plants be kept too moist at first. This point was attended to, but the result was the same. Mr. A. S. Wilson, however, has been more successful, for he showed me some oat-plants which he had artificially infected by removing the glumelle and applying the spores to that part of the kernel from which the embryo emerges; but he also informed me that he had many failures, But more than this remains to be considered. 7. trétic¢ matures its teleutospores at the same time that the wheat- plant matures its fruit; but with U. segétum the case is 102 British Uredinee and Ustilaginee. altogether different, for the smut is formed at the time, or soon after, the cereals are in blossom, and long before harvest it has been scattered by the winds, so that in the harvest field one never finds a smutted ear. We know, moreover, that when once the teleutospores fall on the ground, or in any way become damp, they forthwith ger- minate, and although they are capable of retaining their power of germination for some years, it is only when they are perfectly dry—a condition which never obtains with them in a state of nature. There must, therefore, be some means by which the interval is bridged over between the ripening of the teleutospores of WU. segetum, which takes place in early summer, and the time when the grain itself germinates, for this, under any circumstances, can only be one or two months later. This may be by a metcecism, but there is no proof whatever that any such occurs; or it may be by the continued reproduction of yeast-spores, as Brefeld suggests taking place in manure heaps. My own experiments, however, with nahrlésung containing U. Ssege- zum spores have all been negative. There is a certain point in connection with the repro- duction of smut (U. segetum) wherein it differs essentially from bunt (7. ¢rétzc¢); it is this—that however carefully wheat may be dressed with cupric sulphate, arsenic, brine, lime, etc., while such dressing almost absolutely protects the crop from bunt, yet it has no appreciable affect upon smut. This fact is obvious to any one residing in an agricultural district. The wheats are dressed for bunt on every well-managed farm, but they are as much affected with smut as the barley and oat crops, which latter, never being affected with bunt, are never subjected to protective dressing. In 1883, I made a series of experiments by applying the teleutospores of U. segetum to the wheat and oat plants L[nfection of Host-Plants by Ustilaginee. 103 d while they were in flower, but the plants were subsequently destroyed by an accident. Recently Mr. J. L. Jensen has published the results of his experiments and observations conducted on the ex- perimental farm of the Royal Agricultural School near Copenhagen. He believes the spores of U. segetum effect their entrance into the host-plant at the time it is flowering, and either infect the ovum or remain quiescent, enclosed within the grain until the ensuing spring, when they germinate when the grain does, and so cause its infection. He found that barley grown for twenty-five years con- secutively upon one experimental plot was not more affected with UW. segetum than when grown in the ordinary rotation with other crops, which clearly shows the teleutospores do not remain effective in the soil. He further found that manuring with ‘farmyard manure does not produce moré U. segetum in the crop than occurred when artificial manures were employed. But he did find that seed obtained from a field in which the fungus had been abundant produced a more severely diseased crop than when the seed was taken from a healthy field ; but that, if the seed oats were dipped in water at a temperature of 57°C. (134° F.), and allowed to remain there for five minutes, the disease was prevented, and, moreover, the vitality of the seed was unimpaired.* With regard to 7iletia tritici, the important question of the protective dressing of the seed corn has long ago engaged the attention of agriculturists. Many have been employed, but that most generally used (and probably the best) is a ‘5 % solution of cupric sulphate in water. Alum, ferrous sulphate, and even sulphuric acid, unless used sufficiently concentrated to injure the seed corn, were * Some additional observations of Mr. Jensen on this subject, made during the year 1888, will be found under Usti/ago segetum, See Descriptions. British Uvredinee and Usttlaginee. red by Wolff to be quite useless. With regard to perature, a series of experiments made by Schindler * vs that the teleutospores of J. ¢v7tic¢ will withstand a heat of 65° C. before altogether losing their power of iination ; but with moist heat they were sterilized reen 45° and 50°C. Cold, on the other hand, had little t upon them, even after exposure to —20° C. for a onged period. ichindler, ‘‘ Ueber den Einfluss verschiedenea Temperaturen auf die ahigkeit den Steinbrandsporen,” “ Forschungen auf Geb. der Agrikultur- :. bd. iii.” (1880), pp. 288-293. ( 105 ) CHAPTER XII. SPORE-CULTURE., THE microscopic examination of the Uredineze and Usti- laginee is a very simple matter. The acidia viewed as opaque objects with a low power are always very attractive objects. To examine the various spore-forms, all that is necessary is to remove a small quantity with the point of a penknife, place them in a drop of water on a glass slide, and, having covered them with a circle of thin glass, view the preparation as a transparent object with a quarter-inch objective. The various markings on the exterior of the zecidiospores and uredospores are more readily seen if they be examined dry. In order to obtain more accurate in- formation of the structure of the spore-beds, thin sections must be cut with a sharp knife, including both the spore- bed and a small portion of the host-plant. With a little patience, and by teazing out the cells of the host-plant, the mycelium can be observed. This is often rendered more conspicuous by the application of a drop of caustic potash. To observe the germination of the spores is not difficult, and can be accomplished without the aid of expensive or elaborate apparatus. All that is necessary is to place the spores in asufficiently humid atmosphere, or in a sufficiently moist place, This may be conveniently accomplished by placing a drop of pure water upon an ordinary glass slide, 106 British Uredinee and Ustilaginee. and putting into this drop of water the spores whose germination it is desired to watch ; as, however, the spores do not germinate for several hours, the drop of water would evaporate, unless means be taken to prevent it, before the germination takes place. This is easily done by placing the glass slide under a bell-glass, inverted over a plate of water, so that it is kept continuously in a saturated atmo- sphere. Any suitable object will do to place the slide on; but the most convenient appliance will be found to be a simple stand or rack (Fig. 9), which can be constructed el, i IS Hl Fig. 9.—Stand with slides for the culture of Uredine spores, placed on a plate of water, and covered by a bell-glass. in a few minutes out of two flat pieces of gutta-percha and four glass rods. The gutta-percha which is used for soling boots answers well enough. If two oblong pieces of equal size, say three and a half by four and a half inches (9 X 12 cm.), be taken, and a hole bored at each of the four corners, and through each pair of holes let a glass rod be passed, a very simple stand is made at the cost of a few pence. For the glass rods an old barometer tube, cut into suitable lengths with a file, does well enough. The great advantage of such a stand is, that when the slides are laid across the rods, they touch only at two Spore-Culture. 107 points, so that when placed under the microscope there is no necessity to wipe the lower side of the slide. Of course, the length and height of the stand must be propor- tionate to the size of the bell-glass. The best form of bell- glass is that known by gardeners as a propagating glass, which is rather flatter than the ordinary bell-glasses ; the advantage being that the slides are not too far above the surface of the water in the plate, so that the drops of water on them do not evaporate so rapidly. Supposing we wish to observe the germination of the ecidiospores of Puccinia graminis, having obtained a per- fectly fresh-gathered leaf of barberry with the Aécidium on it, we proceed as follows. The spores can either be brushed upon the slide with a camel-hair pencil, or what will often be found more convenient, the zcidium can be gently struck upon the dry slide, and a drop of water let. fall upon the tiny heap of golden spores that have been displaced. The ripe spores will most of them float on the top of the water and can be readily observed with a quarter-inch objective. The preparation must, of course, not be covered with a cover-glass, and it takes a little patience to examine these uncovered objects, because the front of the object-glass is apt to become'bedewed and misty. The only plan is to raise it by the coarse adjustment, wipe it dry, and try again. Germination will be well advanced in the course of ten or twelve hours, and the migration of the yellow endochrome along the germ-tube will by that time have taken place. This will be followed by their circumnutatory movements and ultimate branching. Earlier examination of the preparation will show the germ-tubes emerging from the germ-pores. It is useless to attempt to get «cidio- spores to germinate unless they are perfectly fresh and perfectly ripe. For instance, the spores dug out from the bottom of an zcidial cup with a needle will not germinate ; 108 British Uvedinee and Ustilaginee. nor will they if they have once become thoroughly dry. The same method is to be adopted with the uredospores, and the same precautions observed. With regard to the teleutospores, certain modifications of the above are neces- sary. If one of the Leptopucciniz is to be examined, all that is requisite is to cut up one of the sori and place the fragments in a drop of water, and in a few hours the pro- mycelia will be developed. Those species which have a prolonged period of rest in their life-history, it is, of course, useless to attempt to germinate except at the proper season of the year. Suppose it is desired to observe the germina- tion of P. graminis, in the autumn some specimens of mil- dewed straw must be procured, and preserved through the winter, I have always found the best plan is to tie them up in a bundle and keep them out-of-doors, so that they are exposed to the same vicissitudes of temperature and moisture as would happen to them in a state of nature ; for it is obvious that if they be kept throughout the winter indoors, they will not only be maintained at a higher temperature, but also will become more completely dried than is natural to them. Under such circumstances they neither germinate so freely nor so uniformly as they do when they have passed the winter in the open air. Specimens of P. graminis may be obtained on Triticum repens, in February or March, from the immediate vicinity of any barberry bush, and these will be found to germinate very readily. Having obtained the material, in March or April, however it may have been preserved, in order to get it to germinate all that is required is to place it in water. If some of the spore-beds be cut into pieces about one- eighth of an inch (2 or 3 mm.) across, and placed in water in a watch-glass, under the bell-glass, the process of germi- nation soon commences, perhaps in twelve hours, perhaps longer, according to the manner in which the material has Spore-Culture, 109 been preserved, and according to the temperature of the atmosphere at the time the experiment is made. I never remember having seen any teleutospore germinate if the temperature within the bell-glass was below 5° C. Germina- tion is very partial and very slow at 8° C., but at from 10° to 15° C. it is both vigorous and rapid. The germination can be recognized to have taken place, if it be atall free, by the naked eye; the clusters of spores will then be seen to be surrounded by an opalescent, hazy cloud, which, when placed under the microscope, will be found to consist of myriads of promycelia. A very convenient method of preserving material on grasses—such, for instance, as Uro- myces po@, which occurs on the leaves of the grass that in the ordinary course of events become disintegrated by decay during the winter—is to gather a small bundle of affected leaves, attached to the stems, place this bundle in a flower-pot just as if it were a living plant, to cover the flower-pot with a bell-glass, and keep it out-of-doors in a shady place all winter. There will be enough moisture in the atmosphere to prevent the material from being injured by desiccation, but not enough to allow the teleutospores to germinate until they are purposely placed in water. The bell-glass will protect the grass from injury by wind and weather; so that when spring comes you will have abundance of material ready to hand, in excellent condi- tion for germination, which you can induce at pleasure, by merely soaking it in water. The same method may be conveniently adopted with those species which occur on leaves, and in which the spores are lost by their decay in the ordinary course of events. The Melampsore on willow and poplar may thus be kept out-of-doors under a bell- glass with great facility. With those species which occur on leaves of plants which have a very perishable foliage, it is necessary to IIO British Uvredinee and Ustilaginee. collect specimens which are perfectly mature, if possible, upon leaves that are beginning to fade from age, and dry them in the ordinary way between blotting-paper. When the spring comes, the affected leaves must be soaked for twenty-four hours in water, and the spores examined to see if any attempt at germination is observable. If not, the soaked leaves may be wrapped in an old, well-washed piece of calico, and buried for a day or two in the ground ; after which treatment, a few spores must be tried in a drop of water on a glass slide. Should they fail to evince any signs of vitality, the leaves must again be buried for a day or two longer, and re-examined. The entrance of the germ-tube into the host-plant can be observed in various ways. The promycelial spores can be applied to the surface of a leaf, and sections made a few hours afterwards. This is, however, an exceedingly delicate process, and requires not only patience, but considerable manipulative skill. A piece of the epidermis may be stripped off and laid flat upon a moistened slide, and the spores placed on it ; if the preparation be kept in a moist atmosphere for a few hours, the germ-tubes can be seen boring through the cells. Another plan is to place a mass of teleutospores, which has first been seen by the micro- scope to be in active germination, on a leaf, and to keep it for a few hours under a bell-glass in a moist atmosphere. The teleutospore mass can be seen by the naked eye, and is a guide to the exact part of the leaf to be examined. By a little deft manceuvring, pieces of the epidermis at this spot can be ripped off with the point of a penknife, and examined either on their external or internal surfaces. By similar methods, the entrance of the uredospore and zcidio- spore germ-tubes can be observed, only, being larger and containing yellow endochrome, the process is less difficult. With most of the Ustilagine, all that is necessary is to Spore-Culture. III place the spores in water, and they will germinate at once— for example, U. segetum, longissima, etc.; but some require a longer period of immersion, eg. T. ¢riticz, which will not germinate till after being several days in water. The germination of the Tilletia spores can also be conducted in a hanging-drop culture in the following manner :—A piece of glass tube, about half an inch (12 mm.) in diameter and about three-quarters of an iach (18 mm.) long, is cemented on an ordinary glass slide, so as to form a deep cell (Fig. 10). Fig. 10.—Deep cell for hanging-drop cultures, made by cementing a piece of glass or lead ‘tube upon an ordinary glass slide. Into this is placed a small quantity of water. The drop containing the spores to be germinated is placed on the centre of a circular cover-glass, which will fit the top of the cell (Fig. 11). If the upper edge of the tube which forms Fig. 11.—Hanging-drop culture, seen in section. the cell be oiled, no air can get into the culture, and the germination can be watched for months, as the hanging- drop will not evaporate, because the water placed in the bottom of the cell keeps the atmosphere saturated, and any loss which it may sustain from evaporation is made up from the water at the bottom of the cell. There is no need to keep these sealed hanging-drop cultures in a moist atmosphere, because the necessary 112 British Uredinee and Ustilaginee. moisture is contained in the cell itself; all that is require: is to place them under a bell-glass to keep them free fron dust. The disadvantage of this mode of culture is, tha all spores will not germinate normally unless they get ; supply of free air. To obviate this hindrance, the glas. cover may be fixed to the cell with three tiny fragments o wax. These cells may be readily constructed with leac tubing; an ordinary piece of gas-piping, cut into suit able lengths with a knife, and the ends smoothed on < whetstone, will answer all the requirements of the case. The germination of the spores in xa&hrlisung, however requires more care. The nahrlésung is prepared by boiling fresh horse-dung in pure water, and filtering first througl Fig. 12.—One of Brefeld’s ‘‘ésmmer” for nahrlésung cultures. It consists of a glass daceralby seadige mmtlotars peoaletimeaciatner auuearig ie Mikes ia uae oe ends of the tube are closed with cotton wool. coarse filtering-paper, then through fine ; then boiling again for a short time in a flask, the mouth of which is closed, while the steam is issuing from it, by a firm plug of cotton wool. After three or four hours the flask is again boiled for a short time. To sterilize the fluid this boiling requires to be repeated several times, carefully closing the mouth ot the flask each time with a plug of cotton wool. The spores to be germinated are put in a small quantity of sterilized nahrlésung, and the process watched zz camerd. The little apparatus necessary consists of a glass tube about eight or ten inches long, open at both ends; in the middle of this tube a bulb has been blown, the sides of which have.been compressed laterally (Fig. 12), so that they are flat and parallel to each other. Before using, the apparatus (Aammer) Spore-Culture. 113 must be sterilized by heat; and after the nahrlésung has been introduced, the two ends of the tube are closed by cotton wool; this allows free access of air, but filters out any germs that may be floating in it. Or the culture may be made in a hanging-drop cell made of lead tubing, in the sides of which two holes have been cut (Fig. 13). I have found it most convenient to _ Fig. 13.—Hanging-drop culture cell, made of lead tubing, with two holes for the admis- sion of air; when in use with nahrlésung these openings are closed with cotton wool. wrap cotton wool round the cell, and to hold it in its place by a small elastic band. These cells require to be sterilized by washing with a weak solution of corrosive sublimate. (14) CHAPTER XIII. { 1 THE ARTIFICIAL INFECTION OF PLANTS. IN order to ensure success in the artificial infection oi plants, attention to several little details is absolutely neces- sary. Of course, if you simply wish to produce 4edium urtice on a cluster of nettles, you may throw a handful o! Carex hirta affected with Puccinia upon the ground where the nettles grow in autumn, and, trusting to chance, you will probably find them bearing the A¢cidium the following spring. But such a procedure is open to many objections - the wind may blow away your Carex during the long winter and spring months either before the Puccinia has germinated or before.the nettles have appeared above ground. A still greater objection is, that even if a few clusters of zcidia happen to be produced on the nettles you have no proof that they arose from the Puccinia you threw down. Still more important is it to avoid this clumsy method of “laying on,” if you are investigating the life. history of any particular species of Uredine, for it ofter happens that more than one species attacks the same host- plant; P. magnusiana, trailii, and phragmitis, on the reed for instance. The first thing to be done is to provide suitable plant: for infection. These should, it is hardly necessary to state be healthy, and have had time to become established before The Artificial Infection of Plants. 115 they are infected. It is a good plan to establish a number of plants, say half a dozen, in the autumn; they will then be ready for use in the following spring. It is often con- venient to infect every alternate plant, so that the remaining plants may be kept as control specimens. The reason for using established plants is that the young foliage is so much more easily infected by the Uredinee than the older ; in fact, it is by no means uncommon for an old leaf to die off before the Uredine has had time to complete its development. Let us suppose we wish to perform the classical in- fection of the barberry with P. graminis. In the autumn, six young barberries, small enough to be covered with a bell-glass, having been planted, as soon as their leaves are fully developed in the spring they may be infected in the following manner. A quantity of P. graménds having also been provided in the autumn and kept during the winter in the mode before explained, as soon as the barberry foliage is ready, test the germinative power of the P. gramznis by placing a few fragments in water in a watch-glass. If it germinate freely and produce a good crop of promycelial spores, as proved by microscopic examination, the contents of the watch-glass may be at once employed. It is best to do your infection experiments in the evening. Water one of the barberries freely through the rose of a watering-can and then cover it with a bell-glass ; then water the outside of the bell-glass. By so doing the temperature of the enclosed air is reduced, and the inside of the bell-glass as well as the leaves of the barberry become bedewed with condensed vapour. After leaving it a few minutes, remove the bell-glass, and apply the germinating spores with a camel-hair pencil. As the promycelial spores easily become diffused in the water in the watch-glass, by stirring it with the camel-hair pencil the water becomes equally charged 116 British Uredinee and Ustilaginee. with them; then, by simply brushing the water on the leaves, you may be pretty sure of successfully infecting the plant. Replace the bell-glass, and give it another douching outside with the watering-can. If sufficient material has been prepared, each alternate barberry may be infected in the same manner. The bell-glass need not be kept over the infected plants more than two or three days. If the weather be very bright, the bell-glasses should be shaded by putting a piece of matting or carpet over them to prevent the foliage being scorched by the sun. In the course of eight or ten days the yellow spots on which the spermogonia are produced will appear, and in two or three weeks the perfect xcidiospores will be de- veloped. It will then be seen that only those barberries to which the spores: were applied have the zcidiospores on them, while the alternate plants remain free. If an attempt be made to infect a plant in the day-time, when the sun’s rays are full upon it, it will be found that the water all runs off the leaves ; but by operating in the evening in the manner directed, the leaves are bedewed with a thin layer of moisture, and no difficulty will be found in applying the spore-charged water. Should it be desired to perform the converse culture— the infection of wheat plants with the barberry zcidiospores —greater care is necessary to prevent the control plants from becoming infected, because the zcidiospores will not diffuse themselves in water, and are very readily carried away by currents of air. The simplest way is to plant some wheat in two flower-pots, and at once to place the pots on two plates of water and cover each with a bell-glass. As soon as the young wheat plants have made a green leaf, those in one of the flower-pots may be infected, using the same precautions as are given above. The acidiospores may be collected by brushing them into a watch-glass of water The Artificial Infection of Plants. 117 with a dry camel-hair pencil, taking care to use only those which will readily brush off. The contents of the watch- glass may then be applied to the wheat plants. By this method the accidental infection of the control plants is avoided, because the bell-glass is never removed from them. Personally, I have found infection with promycelial spores more certain than with the zcidiospores, because we can see that they are actually germinating at the time they are used, while with the ecidiospores this cannot be done so certainly ; moreover, the zcidiospores being generally brought from a distance, they are apt to lose this germinative power, unless used immediately, from becoming too dry on the one hand, or on the other, if kept in too moist an atmosphere during their transit, from many of them having already germinated before they are em- ployed. It is only by attention to these minute details that we can ensure uniform success. The Gymnosporangia are very easy to cultivate. A few seedling hawthorns can be obtained anywhere, and it is necessary only to soak the Gymunosporangium clavarie- JSorme for twelve hours, when the golden promycelial spores will be visible to the naked eye. In producing R. cancellata on pear, it is necessary to infect two-year old plants, because if seedlings be infected the spermogonja alone will be produced, because seedling pears lose their foliage before the Reestelia has had time to develop ; with these plants it is essential to success that they should be thoroughly established before they are made the subiect of experiment. In working out the life-history of the allied species duplicated cultures are very valuable. Suppose we wish to produce the acidia of P. magnusiana and P. phragmitis. Having provided the proper material and a number of 118 British Uredinee and Ustilaginee. growing plants of Rumex obtusifoliius and Ranunculus repens, germinate a quantity of the P. maguusiana in a watch-glass, and then put half on a Rumex and the other half on a Ranunculus ; in a week or ten days we shall find the Ranunculus affected and the Rumex free. Care, of course, must be taken that there is no mixture of teleuto- spores in the watch-glass. In like manner the P. phrag- mitis may, on a subsequent occasion, be applied to the other two plants, when we shall find the Rumex become affected and the Ranunculus will remain free. The main points to be attended to in order to ensure success in performing these cultures are, first and foremost, to have ocular demonstration that your infecting material is actually germinating at the time you use it; and, secondly, to infect the young growing foliage of established plants. DESCRIPTIONS OF THE BRITISH UREDINE#:. UREDINEA. Tulasne. MYcELIuM parasitic in living plants. Spores formed from the ends of erect, crowded hyphe, usually of more than one-kind. Teleutospores germinating by a short promycelium. VROMYCES. Link. Teleutospores separate, unicellular, pedicellate, produced in flat sori (spore-beds), apex perforated by a single germ-pore. I, EUUROMYCES. Schroter. Having spermogonia, zcidiospores, uredospores, and teleutospores, the latter germinating only after a period of rest. A. AUTEUUROMYCES. Having all spore-forms on the same host-plant. Uromyces fabe. (Pers.) Atcidiospores—Pseudoperidia generally crowded upon whitish spots, which are more or less circular, short, slightly pro- minent, flat, with torn white edges. Spores subglobose, orange-yellow, finely echinulate, 15-25 in diameter. Uredospores—Sori chestnut-brown, roundish, amphigenous, scat- tered, often confluent, soon naked. Spores subglobose or ovate, with three germ-pores, shortly echinulate, yellowish brown, 20-30 X 17-20. Teleutospores—Sori rounded .on the leaves, more abundant and elongate on the stems, persistent, black. Spores variable in form, ovate or broadly clavate, dark brown, smooth, apex much thickened (8-10), rounded or conical with one in- 120 British Uredinee and Ustilaginea. fundibuliform germ-pore, 25-40 X 20-254. Pedicels long, persistent, pale brown. Synonyms. Uromyces orobi, Pers. Winter in Rabh., ‘Krypt. Flor.,” 2nd edit., vol. 1. p. 158, in part. Uromyces fabe. Cooke, ‘‘ Grevillea,” vol. vii. p. 135. Uromyces appendiculata, Lév. Cooke, “ Hdbk.,” p. 518; ‘Micro. Fungi,” 4th edit., p. 212. Credo faba, Pers. Romer, “New Mag.,” vol. i. p. 92. Grev., **Scot. Crypt. Flor.,” t. 95; ‘‘ Flor. Edin.,” p. 436, in part. Credo fusca. Purton, ‘‘ Mid]. Flor.,” vols. ii. and iii., No. 1130. Uredo leguminosarum, Link. Berk., “ Eng. Flor.,” p. 383. Credo appendiculosa, Berk., ‘‘Eng. Flor.,” p. 383. Trichobasis faba, Lév. Cooke, “Micro. Fungi,” 4th edit., p. 225. Puccinia globosa. Grev., “ Flor. Edin.,” p. 368; ‘Scot. Crypt. Flor.,” t. 29. Puccinia faba, Link. Berk., “Eng. Flor.,” p. 434. Cooke, “ Hdbk.,” p. 508; “Micro. Fungi,” 4th edit., p. 211. Exsiceati. Cooke, i. 71; ii. 52. Vize, “Micro. Fungi,” 44; “Micro. Fungi Brit.,” 63, 223. On Faba vulgaris, Vicia cracca, sepium, sativa, Lathyrus pra- tensis, Pisum sativum. Ecidiospores, April and May; uredospores, May to July; teleutospores, July to November, and lasting through the winter on the dead stems. BioLocy.—There are several species of Uromyces parasitical upon the Leguminose. By most botanists the above is considered identical with U/. ervz. As early in the year as February 8 (1884), I produced the 4Zcidium upon a bean plant (/_faé@) from teleutospores on bean straw which had been grown for agricultural purposes. The latent period between placing the teleutospores on the host-plant and the appearance of the spermogonia was twenty-three days. The cul- ture was repeated on March 20 in two experiments, both of which were successful, the spermogonia in each case showing on April 16. In 1886, four cultures were made with U. evvi by placing the germi- Auteuuromyces. 12 nating teleutospores on bean, pea (Pzsum sativum), and two vetch seedlings. Although the teleutospores were germinating very freely, yet they produced no result. In 1888 two further series of cultures were made with U. fabe on bean, pea, Vic¢a sativa, cracca, Lathyrus pratensis, and Ervum hirsutum. No zcidiospores were produced except upon the bean and the pea. The ecidium on the bean occurs on white spots, which are thickened and very conspicuous. On pea the spots are pale dirty yellow, and the pseudoperidia few and scattered, while on bean they are numerous and crowded together. “There can’ be no doubt that U. fade and ervé are biologically distinct, but the 1888 cultures show that continued investigation will probably lead to further subdivision of the forms now grouped under U. fase. De Bary says that in some instances the same mycelium which produces the ecidia of U. fade gives rise to a few isolated uredo- spores. Uromyces orobi. (Pers.) cidiospores—Spots scattered or circinate. Pseudoperidia flat or slightly prominent, with torn whitish edges. Spores sub- globose or polygonal from mutual pressure, orange, echinulate, 16-27 in diameter. Uredospores—Sori small, crowded or scattered, soon naked, elon- gated on the stems. Spores subglobose or ovate, echinulate, yellowish, 15-28 x 16-224. Teleutospores—Sori roundish or elongate, very dark brown, at first covered with epidermis. Spores oblong or subpyriform, api- culate above, often obliquely, from a thickening of the epispore, dark brown, smooth, 25-4o xX 18-28. Pedicels very long, persistent, pale brown. Synonyms. Uromyces orobi (Pers.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 158, in part. citium orobi, Pers. Romer, “‘ New Mag.,” vol. i. p. 92. “cidium orobt, D. C. Cooke, “ Hdbk.,” p. 542; “ Micro, Fungi,” 4th edit., p. 197. Berk., ‘‘ Eng. Flor.,” vol. v. p. 378. Pucinia fabe. Johnst., ‘ Flor. Berw.,” vol. it, p. 197. Exsiccati. Vize, “ Micro, Fungi Brit.,” p. 327. On Lathyrus macrorrhizus (Orobus tuberosus). July to October. 122 British Uredince and Ustilaginee. Uromyces phaseoli. (Pers.) 4 cidiospores—On circular spots about 2 mm. across. Pseudoperidia cup shaped, crowded, with everted, whitish, deeply toothed edges. Spores polygonal, finely verrucose, colourless, 20-25 X 16-184. Uredospores—Sori_ scattered, pale cinnamon brown. Spores rounded or shortly elliptical, pale brown, echinulate, 25-34 x 15-18. Leleutospores—Sori black-brown, soon naked, Spores spherical or shortly elliptical, apex thickened, with a wide germ-pore and often a colourless papilla, smooth, dark brown, 26-35 x 22-364. Pedicels short, deciduous. Synonyms. Uromyces phaseoli (Pers.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 157. Uredo appendiculata, var. phaseoli, Pers. Observat. in “ Usteri ann. d. Botan.,” vol. xv. p. 17. Uromyces phaseolorum, De Bary. Cooke, “ Grevillea,” vol. vii. P. 135. On Phaseolus vulgaris. May and October. BioLocy.—-The spots on which the zecidia occur are at first pale, becoming yellowish. I have never seen British specimens of this, which is inserted on the faith of Dr. Cooke’s paper upon Uromyces in “ Grevillea.’ Uromyces limonii. (D. C.) Ecidiospores—-On purplish spots. Pseudoperidia hypophyllous, white, cylindrical, with much-torn white edges. Spores sub. globose or ovate, yellow, minutely verrucose, 16-25 X 15—20p, Uvredospores—Sori brown, roundish, bullate, then naked, scattered, Spores globose or ovate, pale brown, finely verrucose, 30-35 X 25—30p. Teleutospores—Sori small, rounded, black. Spores ovate or sub: pyriform, darker and thickened at the apex, often attenuatec below, smooth, rich brown, 25-50 X 15-254. Pedicels very long. A uteuuromyces. 123 Synonyms. Uromyces imonit (D. C.). Winter in Rabh., “ Krypt. Flor.,” vol. i, p. 156. Puccinia limonit. D.C., “ Flore frang.,” vol. ii. p. 595. Ecidium statices, Desm. Cooke, “ Micro. Fungi,” 4th edit., Pp. 197. Uredo statices, Desm. Cooke, “ Hdbk.,” p. 528; ‘Micro. Fungi,” 4th edit., p. 217. Credo armeria, Duby. Berk., Eng. Flor.,” p. 377. x Uromyces limonit, Lév. Cooke, “Hdbk.,” p. 5183; “ Micro. Fungi,” 4th edit., p. 212. LE xstccatt, Cooke, i. 632, 444, 591; ii. 83, 324. Vize, “ Fungi Brit.,” 60, 71; “Micro. Fungi Brit,” 128. On Statice limonium, Armeria vulgaris. AEcidiospores, May and June; uredospores, June and July; teleutospores, July to October. BIOLOGY.—The ecidia occur on pallid spots, which are usually tinged with purple round their circumference; on the stems they cause considerable distortion. As the fungus occurs in situations which are often covered by the spring tides, it is obvious that the presence of salt is not inimical to it. Uromyces polygoni. Pers.) Acidiospores—Spots generally hypophyllous, rarely cauline. Pseudoperidia in small irregular clusters, rather flat, with broad, whitish torn edges. Spores subglobose, finely verru- cose, pale yellow, 15-23 in diameter. Uvedospores—Sori brown, scattered, rarely circinating, amphi- genous, pulverulent. Spores globose or ovate, pale brown, finely echinulate, 20-25 X 15-z20p. Teleutospores—Sori blackish, on the leaves roundish, on the stems elongated. Spores globose or elliptical, smooth, chestnut brown, apices thickened, rounded or conical, 23-35 X 15-204. Pedicels yellowish, long, -persistent, 124 British Uvedinee and Ustilagince. Synonyms. Uromyces polygont (Pers.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 154. Puccinia polygont. Pers., ‘ Disp. Meth.,” p. 39. Acidium avicularia, Kze. Cooke, “ Micro. Fungi,” 4th edit., p. 199; ‘“ Hdbk.,” p. 545. Uredo polygonorum. Grev., “Scot. Crypt. Flor,” t. 80; “Flor. Edin.,” p. 434. Johnst., “Flor. Berw.,” vol. li, p. 201. Berk., “Eng. Flor.,” p. 377. Trichobasis polygonorum, B. Cooke, “Micro, Fungi,” 4th edit., p. 226. Puccinia vaginalium, Link. Cooke, “ Hdbk.,” p. 519. Berk., “Eng. Flor.,” vol. v. p. 363. Cooke, ‘Micro. Fungi,” 4th edit., p- 204. Puccinia avicularia. Grev., “Flor. Edin.,” p. 429. Johnst., “Flor. Berw.,” vol. iL p. 195. Uromyces polygoni, Fckl. Cooke, “ Hdbk.,” p. 519; “ Micro. Fungi,” 4th edit., p. 213. Uromyces avicularie, Schrot. Cooke, “Grevillea,” vol. vit p- 136. Exstecati. _ Cooke, i. 123; ii. 144, 312. Vize, “ Micro. Fungi Brit.,” go; “ Micro. Fungi,” 170. On Polygonum aviculare. Ecidiospores, May ; uredospores, May to July; teleutospores, July to November. BIoLoGy.—The ecidium usually occurs on yellowish spots, which are often surrounded by a reddish margin. On the stems it causes considerable distortion. Uromyces trifolii, (Alb. and Schw.) :cidiospores in circular clusters, on pallid spots. Pseudoperidia shortly cylindrical, flattish, on the stems in elongated groups; edges whitish, torn. Spores subglobose or irregular, finely verrucose, pale orange, 14-23 in diameter. Autéuuromyces. 125 Uredospores—Sori pale brown, rounded, scattered, surrounded by the torn epidermis. Spores round or ovate, with three or four germ-pores, echinulate, brown, 20-26 X 18-20. Teleutospores—Sori small, rounded, almost black, long covered by the epidermis. Spores globose, elliptical or subpyriform, with wart-like incrassations on their summits, smooth, dark brown, 22-30 X 15-204. Pedicels long, deciduous, Synonyms. Cromyces trifolid (Alb. and Schw.). Winter in Rabh., “ Krypt. Flor.,” vol. i, p. 159. Credo faba, B. trifolit, Alb. and Schw., “Consp.,” p. 127. Puccinia fallens. Cooke, “ Hdbk.,” p. 508; “ Micro. Fungi,” 4th edit., p. 212, in part. Trichobasis fallens. Cooke, “ Micro. Fungi,” 4th edit., p. 226. Uromyces apiculatus, Lévy. Cooke, ‘Grevillea,” vol. vii. p. 136, Lxsiccate, Cooke, i. 116; “L. F.,” p. 40. Vize, “ Fungi Brit.,” 18. , » P g On Trifolium pratense, and repens. fécidiospores, May; uredospores, May; and teleutospores, May to November. BioLoGy.—The Uredo and Uromyces frequently attack the petioles, where they cause elongated swellings and distortions. Schréter has found the zcidiospores only on Trifolium repens, and states that this stage is of very short duration. A plant of 7. vepens, with the Uromyces upon it, was in October brought indoors and kept there until the following summer. During all this time it produced only teleutospores, no zecidiospores. In the open air the foliage would probably have been destroyed by the cold, so that the fungus would, therefore, have been unable to have kept itself alive, and would of necessity have been compelled to develop zecidiospores in spring from the last year’s teleutospores (Schroter, ‘‘ Cohn’s Beitrage,” vol. ii p. 78). Dr. Cooke has observed bicellular teleutospores (Seem. Jour., vol. iv., 1866) on Vicza sepium ,; they were few in number, and mixed with the uredospores. 126 British Uvredinee and Ustilaginea. Uromyces geranii. (D. C.) Z:cidiospores—Pseudoperidia crowded. in irregular or roundish patches on reddish spots, shortly cylindrical, edges white, at first adpressed, afterwards recurved, toothed. Spores roundish, finely verrucose, orange, 20-30 X 15-20p. Uredospores—Sori dark chestnut-brown, in rounded groups, small rounded, soon naked. Spores spherical, shortly elliptical, echinulate, pale brown, 20-25 X 19-23m Teleutospores—Sori blackish, rounded, often circinate. Spores elliptical or pyriform, with a colourless wart-like papilla on the summit, smooth, brown, 20-25 xX 17-234. Pedicels short, deciduous. Synonyms. Uromyces geranit (D. C.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 160. Uredo geranii. D.C., ‘‘Synop. Plant.,” p. 47. Berk., “ Eng, Flor.,” vol. v. p. 380. Sow., t. 398, fig. 5. Grev., “Scot, Crypt. Flor.,” t. 8; “Flor. Edin.,” p. 434. Johnst., “Flor. Berw.,” vol. ii. p. 201. : Trichobasis geranit, Berk. Cooke, “ Hdbk.,” p. 530. i Acidium geranit, D. C. Cooke, “ Micro. Fungi,” 4th edit. p. 199; “Hdbk.,” p. 543. Berk., “Eng. Flor.,” vol. v. p. 371. Johnst., “Flor. Berw.,” vol. ii. p. 205. Uromyces geranit. Cooke, “ Micro. Fungi,” 4th edit., p. 213 “Grevillea,” vol. vii. p. 134. Exsicati. Cooke, i. 107, 440; ii. 50. On Geranium sylvaticum, pratense, dissectum, and molle. May to October. BioLoGy.—At the Mycological Conference in Paris, in 1887, Dr. Richon exhibited a figure of the uredospores accompanied by large clavate, hyaline paraphyses, but I have not observed these in any British specimen I have examined. The zcidiospores often cause great distortion when they occur on the stems. On the leaves they usually occur on reddish spots. Professor Trail finds this fungus near Aberdeen, on the two first-named host-plants, but not upon G, dissectum or molle. A uteuuromyces. 127 Uromyces betes. (Pers.) Atcidiospores—On yellowish rounded or elongated spots. Pseudo- peridia irregularly scattered or circinate, cup-shaped, with whitish torn edges. Spores polygonal, isodiametric, orange- yellow, smooth, 15-25 in diameter. Uredospores—Sori brown, irregularly roundish, surrounded by the ruptured epidermis, scattered or circinate. Spores ovate or elliptical, pale yellowish brown, echinulate, 25-30 X 16-25). Teleutospores—Sori black brown. Spores roundish, elliptical, ovate or obovate, dark brown, smooth, with a colourless papilla on their summits, 25-35 < 20-254. Pedicels long, deciduous. Synonyms. Uromyces bete (Pers.). Winter in Rabh., “Krypt. Fior.,’ vol. i. p. 155. Uredo bete. Pers., ‘‘Syn.,” p. 220. Berk., “Eng. Flor.,” vol. v. p. 377. Uromyces bet, Kithn. Cooke, “Grevillea,” vol. vit. p. 136; ** Micro. Fungi,” 4th edit., p. 213. Trichobasis beta, Lév. Cooke, “ Hdbk.,” No. 1587; ‘“ Micro. Fungi,” 4th edit., p. 225. E-xsiccatt. Cooke, i 70; ii. 31; “L. F.,” 39. Vize, “ Micro. Fungi Brit.,” 553. On Beta vulgaris and maritima. Ecidiospores, April and May; uredospores, June and July ; teleutospofes, August to October. BIOLOGY.—The zcidiospores are very rarely found in this country in a state of nature. In 14885, I produced the zcidia on two plants of mangold from the teleutospores from wild plants of Beta maritima grown on the banks of the River Ouse at West Lynn. Some dead stems of the Beta were laid on the mangold plants on March 20, and on April 21 the zcidiospores were found, which in due course were followed by the uredospores. The zecidium was found in April, on the wild Beta maritima. 128 British Uredinee and Ustilaginee. Uromyces valeriane. (Schum.) Acidiospores—Pseudoperidia hypophyllous, rarely cauline, circi- nate, scattered, cup-shaped, on the stem often elongate, slightly prominent, with torn, erect, white edges. Spores polygonal, finely echinulate, orange-yellow, 17-24 in diameter. Uredospores—Sori small, reddish brown, rounded, scattered or clustered, amphigenous. Spores spherical or elliptical, echi- nulate, pale brown, 20-30 XK 18—20n. Teleutospores—Spots irregular, dark brown, sometimes forming dendritic figures. Sori long covered by the epidermis, amphi- genous, slightly elevated. Spores elliptical or ovate, summits thickened, smooth, chestnut brown, 20-25 X 15-20mu. Pedi cels short, deciduous. Synonyms. Uromyces valeriane (Schum.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 157. Cooke, “ Grevillea,” vol. vii. p. 137. Credo valeriane, Schum. ‘Enum. Plant. Sell.,” vol. il. p. 233. Lecythea valeriane. Berk., ‘ Outl.,” p. 334. Cooke, “ Hdbk.,’”’ p- 523; ‘‘ Micro. Fungi,” 4th edit., p. 222. Ecidium valerianacearum, Duby. Cooke, ‘‘Hdbk.,” p. 540; ““ Micro. Fungi,” 4th edit., p. 196; “ Eng. Flor.,” vol. v. p. 370. Johnst., ‘‘ Flor. Berw.,” vol. ii. p. 206. LE xsiccatt, Berk., 349. Cooke, i. 63, 103; il. 64, 88; “L. F,” 32, 56. Vize, “‘ Micro. Fungi Brit.” 132, 448 ; “ Fungi Brit.,” 68. On Valeriana officinalis and dioica. Ecidiospores, May and June; uredospores, June and July ; teleutospores, July to September. B1oLoGy.—The presence of the mycelium in the leaves and stems causes the acidiospore-sori to be seated on the thickened spots. Uromyces parnassiz. (D. C.) A:cidiospores—Hypophyllous on pallid spots, in rounded patches. Pseudoperidia, tawny yellow, between urceolate and concave, with thick edges. Spores pallid. Uredospores—Spores ‘spherical, rough, 20-2544 in diameter. A uteuuromyces. 129 Teleutospores—Sori amphigenous, at first bullate, then rupturing the epidermis, scattered, often confluent. Spores subglobose, ovoid, brown, smooth, 25-30 X 20-224, Synonyms. Credo parnassiz. OD. C., * Flore frang,” vol. vi. p. 68. LEcidium parnassia, Grev. Cooke, “ Micro. Fungi, 4th edit., p- 198. Trichobasis parnassie. Cooke, Seenr, Jour. Bot, vol. ii. p. 3443; “Hdbk.,” p. 531. Cromyces parnassie, Schrot. Cooke, “ Grevillea,” vol. vit. Pp. 134. Lxsiccate, Cooke, i. 74. Vize, “‘ Micro. Fungi,” 4, 226. On Farnassia palustris. The ecidiospores were found by Dr. Greville near Glasgow, and by Professor Trail near Aberdeen ; the teleutospores by Dr. Cooke at Irstead, Norfolk, in 1864. Uvromyces salicorniz. (D. C.) icidiospores—Pseudoperidia scattered or in small clusters, at first hemispherical, then shortly cylindrical, with erect, white torn edges. Spores polygonal, isodiametric, finely verrucose, orange-yellow, 17-35 in diameter. Uvedospores—Sori rounded, small, long surrounded by the rup- tured epidermis. Spores oblong or subpyriform, echinulate 20-35 X 18-20. Teleutospores—Sori generally larger than those of the uredospores, pulverulent, dark brown, soon naked. Spores rounded, sub- pyriform, apex often thickened, smooth, dark brown, 24-36 x 15-26u. Pedicels long, persistent. Synonyms. Uromyces salicornie (D. C.). Winter in- Rabh., “ Krypt. Flor.,” vol. i. p. 156. Cooke, * Grevillea,” vol. vii. p. 137. cidium salicornie. YD. C., “ Flore frang.,” vol. vi. p. 92. K 130 British Uredinee and Ustilaginee. Exsiccatt, Cooke, i. 538; ii. 143. Vize, “ Micro, Fungi,” 139. On Salicornia herbacea. BioLoGy.—The ecidiospores often occur on the young cotyle- donary leaves, on yellowish spots. B. HeTERUROMYCcES. Schrot. Having the spermogonia and ecidiospores on one host-plant, and the uredospores and teleutospores upon another of a different genus. Uromyces dactylidis. Otth. Aicidiospores—Pseudoperidia on rounded or elongated spots, often in confluent clusters, cup-shaped, with everted torn white edges. Spores polygonal, subglobose, or isodiametric, 15—25p. in diameter. Uredospores—Sori small, elliptical or oblong, scattered, long covered by the epidermis. - Spores almost spherical, rarely ovate, echinulate, orange-yellow, 18-30 x 15-20p, without paraphyses. Teleutospores—Sori small, elongated or roundish, long covered by the epidermis. Spores irregularly rounded or oblong, some- what thickened and darker above, smooth, brown, 18-20 x 14-174. Pedicels short, persistent. Synonyms. AEcidium ranunculacearum, Y). C. in part. Cooke, ‘“ Hdbk.,” P- 539; “ Micro. Fungi,” 4th edit., p. 196. Uromyces dactylidis. Otth., “Nat. Ges. in Bern.” (1861), p. 85. Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 161. Cromyces graminum. Cooke, “ Hdbk.,” p. 520; “ Grevillea,” vol. vii. p. 138; “Micro. Fungi,” 4th edit., p. 214. Lixsiccati. Cooke, i. 537. Vize, “Fungi Brit,” 37; ‘Micro. Fungi Brit.,” 139. Ecidiospores on Ranunculus bulbosus, May and June. Teleutospores on Dactylis glomerata, July to October, and continuing on the dead stems until the following spring. fleteruromyces. ge BIOLOGY.—This species has been the subject of many cultures by me. It has been stated that the uredospores are provided with para- physes ; in this country they certainly are not. It has been affirmed, too, that Ranunculus repens and acris bear the ecidiospores ; but in numerous cultures, many of which were serial (ze. the same infecting material was simultaneously applied to a series of plants), no result was obtained on the above-named plants (2. acris and repens). The series included &. duZbosus, and on it, and on it alone, the ecidium developed, the other species named above (R. repens and acris), as well as &. ficaria and auricomus remaining free from the parasite ; conversely, the zecidiospores from &. du/bosus applied to Poa pratensis and ¢ivialis produced no result. For a detailed account of these cultures, see Quart. Four. of Micro, Science, vol. xxv., new series, pp. 152-156. Uromyces poz. Rabh. Zcidiospores—Similar to the preceding. Spores 15-20uin diameter. Uredospores—Sori orange, rounded, elliptical, or linear ; at first covered by the epidermis, which splits longitudinally. Spores rounded, elliptical, or ovate, finely echinulate, orange-yellow 16-26 in diameter, without paraphyses. Teleutospores—Sori brown, small, punctiform or elongate, covered by the epidermis. Spores generally irregular in form, often elliptical or ovate, pale brown, with a smooth epispore, 17-25 x 25-4opu. Pedicels long, narrow, rather persistent. Synonyms. Atcidium ficariea. Pers., “Obs. Myc.,” vol. ii. p. 23. Purton, “ Midl. Flor.,” vol. iil. p. 333. Sow, t. 397, fig. 4. Acidium ranunculacearum, D. C. in part. Cooke, “ Hdbk.,” p. 539; “Micro. Fungi,” 4th edit, p. 196, plate ii. figs. 12-14. Johnst., “ Flor. Berw.,” vol. ii. p. 206. Berk., “ Eng. Flor.,” vol. v. p- 370. ecidium confertum, Grev., “ Flor. Edin.,” p. 446. Johnst., “ Flor. Berw.,” vol. ii. p. 205. Uromyces poe. Rabh., “Unio. Itin.” (1866), No. xxxviii. Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 162. Exstecati. i Cooke, i. 8; ii 87; “L. F.,” 55. Vize, “ Fungi Brit.,” 72. E93 British Uvredinee and Ustilaginee. «Ecidiospores on Ranunculus ficaria, repens, and bulbosus, March to May. Teleutospores on Poa ¢rivialis, pratensis, and annua, April to July. BioLocy.—This species is said to occur on P. nemoralis as well as on P. pratensis and ¢rivialzs, but I have been unable to produce it on the first mentioned, although I have done so several times on the two latter. In one serial culture in which the Uromyces from P. trivialis was applied to R. ficaria, (2) repens, and (3) bulbosa, the zecidium was produced on the two latter, but not on the first named. No result was obtained on &. auricomus and acris. At present it is safer to say that Ur. foe has its xcidiospores on R. ficaria, repens, and apparently on &. ddbosus (because I do not like to be too con- fident about one culture), and that it has its teleutospores on Poa trivialis and pratensis. I have failed more than once in producing the zecidium on &. ficaria from the Uromyces on P. ¢rzvdalis, although I have always succeeded with teleutospores from P. “ivialés on R. repens, and I think it quite possible that there may be two species, the one having its teleutospores on P. ¢vévéalzs, and its zcidiospores on R. repens, the other with its teleutospores on P. Zratensis, and its wcidiospores on #&. ficayza, Further cultures can alone determine the truth of this surmise. Uromyces junci. (Desm.) Aicidiospores—Pseudoperidia circinating, cup-shaped, with whitish torn edges. Spores polygonal, irregular, globose or elongate, pale orange, smooth, 15-23 in diameter. Uredospores—Sori on brown or yellowish elongated spots, scattered, irregular, rounded or elongated, confluent. Spores rounded or elliptical, echinulate, pale brown, 17-28 X 15-17. Teleutospores—Sori round or elongate. Spores dark brown, usually elliptical or cuneiform, with much thickened and often attenuated summits, deep brown, smooth, 20-40 x I5- sia Pedicels long, thick, pale brown. Synonyms. Uromyces junci (Desm.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 162. ; Puctinia junct, Desm. “Plant. Crypt. Edin.,” 2nd edit. No. 170. cidium zonale, Duby. “Bot. Gall.,” vol. ii. p. 906. fTeteruromyces. ise Uromyces junci, Tul. Cooke, “Grevillea,” vol. vii. p. 139; “ Micro. Fungi,” 4th edit., p. 213. Exsiccatt, Vize, “ Micro. Fungi Brit.,” 445. fEcidiospores on Jnula dysenterica, May to July. Teleutospores on /uncus obtusifiorus, July to October, and lasting through the winter on the dead stems. BIoLoGy.—The presence of the mycelium of the zcidiospores in the leaves causes round spots, which are yellowish in colour, and surrounded by a purple line. Fuckel suggested the probability of the connection between the ecidiospores and the teleutospores, but I believe the actual demonstration of their relationship by culture was never made until I did so in 1882. Uromyces pisi. (Pers.) Aicidiospores—Scattered over the whole leaf surface. Pseudope- ridia cup-shaped, with whitish edges. Spores subglobose or polygonal, orange, finely verrucose, 17-26 in diameter. Uredospores-—Sori roundish, scattered or crowded, cinnamon- brown. Spores subglobose or elongate, yellowish brown, echinulate, 17-20 X 20-25. Teleutospores—Sori roundish or elliptical, blackish. Spores sub- globose or shortly elliptical, finely but closely punctate when recent, apex only slightly thickened, 20-30 x 17-20" Pedicels long, colourless, fragile. Synonyms. Uromyces pist (Pers.). Winter in Rabh., “Krypt. Flor.,” vol, i p. 163. COredo appendiculata, B. prst. Pers., ‘Observ. myc.” in Usteri, Annal. d, Botan., vol. xv. p. 7. Uromyces pist, De Bary. Cooke, “Grevillea,” vol. vii. p. 135. Aicidium cyparissia. WD. C., “Flore frang.,” vol. ii. p. 240. A&cidiospores on Luphorbia cyparissias. Uredospores and teleutospores, on Fesum sativum. 134 British Uredinee and Ustilaginea. BIOLOGY.—I1 have no acquaintance with this as a British species, and the host-plant of its zecidiospores is not native in this country. Its life-history was worked out by Schréter, who states that the teleuto- spores occur on Vicia cracca, Pisum sativum, Lathyrus pratensis and sylvestris, “He further finds that another acidium on £. cyparissias produces another Uromyces—U. striatus, with brown teleutospores ; which, when recent, are delicately striate with wavy lines, and which have a flat conchiform brown cap over the germ-pore. The latter occurs on Lotus corniculatus, Trifolium arvense, Medicago sativa, etc. II]. BRACHYUROMYCES. Schrot. Having spermogonia, uredospores, and teleutospores. Il]. HEMIUROMYCES. Schrot. Having only uredospores and teleutospores, which occur on the same host- plant. Uromyces scutellatus. (Schrank.) Uredospores—Sori small, roundish, at first covered by the epi- dermis. Spores scanty, mixed with the teleutospores, roundish or subpyriform, with a thick colourless or yellowish-brown spore-membrane, smooth, verrucose or finely echinulate, 20- 35 X 15-25. Teleutospores—Irregular, rounded, ovate, or oblong, apices of the spores often surmounted with a broad, flat, pale or colourless papilla. Spores brown, smooth, tuberculate or reticulate, 20-40 X 15-25. YVedicels rather long, deciduous. Synonyms. Lycoperdon scutellatum, Schrank. “ Baiersch Flor.,” vol. ii. p. 631. Uredo excavata, D.C. “Synop. Plant.,” p. 47. Uromyces scutellatus, Lév. Cooke, ‘ Grevillea,” vol. vii. p. 137, in part. Uromyces excavatus, D. C. Cooke, ‘Grevillea,” vol. vii. p. 138; “Micro. Fungi,” 4th edit., p. 213. Oromyces scutellatus (Schrank). Winter in Rabh., “Crypt. Flor.,” vol. i. p. 144. On various Euphorbieze (?). Hlemiuromyces. 135 BIOLOGY.—The mycelium of the teleutospores permeates the whole plant. The foliage of the affected plants is altered by its presence, being shorter, broader, and thicker ; moreover, they seldom blossom. The sori are scattered over the whole plant. Winter con- siders this species to belong to Hemiuromyces, Schréter that it is without uredospores. It is doubtfully British. Uromyces anthyllidis. (Grev.) Uredospores—Sori roundish, chestnut-brown. Spores subglobose, 22-24u in diameter, echinulate,.chestnut-brown, with four or five germ-pores, contents orange-red. . Leleutospores—Sori brownish black. Spores short, elliptical or globose, 19-22 xX 17-20, dark chestnut-brown, markedly verrucose, apex rounded. Pedicels short, deciduous. Synonyms. Uredo anthyllidis. Grev.in Hook. Herb. Berk., “ Eng. Flor.,”’ vol. v. p. 383. Uromyces anthyllidis, Schrot., “ Krypt. Flor. Schl.,” vol. iii. p. 308. On Anthyllis vulneraria. June to October. Uromyces rumicis. (Schum.) Uredospores—Sori amphigenous, brown, small, round, scattered. Spores elliptical or subrotund, echinulate, pale brown, 20-25 X 20-30p. Teleutospores—Sori dark brown, roundish, scattered. Spores roundish, elliptical or subpyriform, chestnut-brown, smooth, 25-35 X 15-25m, apex of the spores having a pale, rounded, wart-like point. Pedicels short, deciduous. Synonyms. Uromyces rumicis (Schum.). Winter in Rabh., “ Krypt. Flor.,” vol. 1. p. 145. Uredo rumicis, Schum. ‘Enum, Plant. Seell.,” vol. ti. p. 231. Purton, “ Midl. Flor.,” vol. iii. No. 1544. Uredo bifrons. Grev., “Flor. Edin.,” p. 435. Berk., “Eng. Flor.,” vol. v. p. 382. Johnst., “Flor. Berw.,” vol. ii. p. 201, Cooke, “Hdbk.,” p. 528; “Micro. Fungi,” 4th edit. p. 217, t. vil. figs. 137-139. 136 British Uvredinee and Ustilaginee. Uredo apiculosa, Link. Berk., “Eng. Flor.,” vol. v. p. 382. Purton, “Midl. Flor,” vol. ili, p. 297. Grev., ‘‘ Flor. Edin.,” Pp. 436. Trichobasis rumicum, D.C. Cooke, ‘ Micro. Fungi,” 4th edit., p. 225. Uromyces apiculosa, L.év. Cooke, “*Hdbk.,” p. 518; ‘‘ Micro. Fungi,” 4th edit., p. 212, t. vil. figs. 154-155. Uromyces rumicum, Lév. Cooke, “Grevillea,” vol. vil. p. 136. Exsiccatt. Cooke, i. 318, 322; “L. F.,” p. 26. Vize, “ Fungi Brit.,” 63, 553 ‘Micro. Fungi Brit.,” 225. On Rumex conglomeratus, obtusifolius, crispus, hydrolapathum, acetosa. May to September. BioLocy.—The sori are often accompanied by very little dis- coloration of the foliage, but the presence of the mycelium often causes those parts of the leaf which are adjacent to the sori to retain their original green colour long after the unaffected portions of the leaves have become yellow from age. Sometimes, however, on R. acetosa there is considerable red discoloration. Uromyces sparsus. (Kze. and Schm.) Oredospores—Sori on pale spots, often convex, round or elliptical, amphigenous or cauline, long covered, at length surrounded by the ruptured epidermis. Spores round or oblong, 20-23 in diameter, to 30m in length. Zeleutospores—Round or ovoid, frequently attenuated below, brown, smooth, epispore thickened above, sometimes hooded, 30-40 X 15-244. Pedicels long and persistent. Synonyms. Uromyces sparsus (Kze. and Schm.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 148. Uredo sparsus, Kze.and Schm. “ Deutsch, Schwamme,” 170. Uromyces sparsa, Lév. Cooke, ‘‘ Hdbk.,” p. 519 ; “ Grevillea,” vol. vii. p. 137; ‘‘ Micro. Fungi,” 4th edit., p. 214. On Spergularia rubra. May to July. Flemiuromyces. 137 Uromyces alchemille. (Pers.) Uredospores—Sori golden-scarlet, hypogenous, rounded or ellipti- cal, frequently arranged radially, following venation of the leaves, becoming confluent. Spores globose, elliptical, or oval, epispore thickly covered with short sharp points, orange-yellow, 15-25 X 1520p, Teleutospores—Sori chestnut-brown, hypogenous, scattered, round- ish, discrete, rarely confluent. Spores elliptical or ovate, verrucose, brown, 30-40 X 20-35. Pedicels rather long, deciduous, ; Synonyms. Uvromyces alchemille (Pers.), Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 146. Uredo alchemilla. Pers. “Syn.” p. 215. Grev., “ Flor. Edin.,” p. 439. Johnst., “ Flor. Berw.,” vol. ii. p. 199. Uredo intrusa, Grev., “ Flor. Edin.,” p. 436. Berk., “ Eng. Flor.,” vol. ii. p. 382. Johnst., “ Flor. Berw.,” vol. ii, p. 207. Uromyces intrusa, Lévy. Cooke, ‘‘ Hdbk.,” p. 519; “ Micro. Fungi,” 4th edit., p. 213. Uromyces alchemilla, Fck\. Cooke, “ Grevillea,” vol. vii. p. 136. Trachyspora alchemille, Fckl. Bot. Zeit, 1861, p. 250. Schroter, “ Krypt. Flor. Schlesien,” vol. iii. p. 350. Lixsiccatt. Cooke, i. 121; “L. F.,” 27. Vize, “ Fungi Brit.,” 40; ‘ Micro, Fungi Brit.,” 43. On Alchemilla vulgaris, May to September. BioLocy.—Those leaves which are attacked by the mycelium of the uredospores do not properly develop, and are usually more elongated in the stalk than the healthy ones. The teleutospores are much less conspicuous than the uredospores. Uromyces alliorum. (D. C.) Uredospores—-Subglobose, pale, 22 X 254 Epispore thin. Teleutospores—Elliptical, brown, 30-35 X 15-184. Pedicels very evanescent. 138 British Uvedinee and Ustilaginee. Synonyms. Uromyces alliorum, D.C. Cooke, “ Grevillea,” vol. vii. p. 138; “Micro. Fungi,” 4th edit., p. 212. Uredo alliorum, T). C. Cooke, “Micro. Fungi,” 4th edit., p. 217, in part. I am unacquainted with this species. IV. UROMYCOPSIS. Schrot. Having spermogonia, zcidiospores, and teleutospores, Uredospores absent. Uromyces behenis. (D. C.) Ecidiospores—Pseudoperidia in round, often confluent, circinate clusters, sometimes almost covering the affected leaf, slightly prominent, rather short, with broad, torn, whitish-yellow edges. Spores round or elongate, finely echinulate, orange-yellow, 15-20 in diameter. Teleutospores—Sori dark brown, often occurring with the ecidia in small, roundish clusters, sometimes confluent, long covered by the epidermis. Spores elliptical or ovate, smooth, with rather markedly thickened summits, 25-40 X 17-25p- Pedicels very long, stout, persistent, hyaline or yellowish. Synonyms. Uromyces behenis, D.C. Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 153. Ecdium behents, D.C. “ Encycl.,” vol. viii. p. 239. Berk., “Eng. Flor.,” vol. v. p. 372. Cooke, “‘Hdbk.,” p. 541; “ Micro. Fungi,” 4th edit., p. 197. Uromyces behenis, Lév. Cooke, “ Grevillea,” vol. vii. p. 134; ‘* Micro. Fungi,” 4th edit., p. 213. LExstecatt. Baxt.,go. Cooke, i. 442. Vize, “ Fungi Brit.,” 167 ; “ Micro. Fungi Brit.,” 134. On Silene inflata, maritima. July to September. B1ioLoGY.—The mycelium causes considerable discoloration of the host-plant, the affected spots being sometimes yellow, sometimes Uromycopsts. 139 brown, and generally having a purple margin. Both zcidiospores and teleutospores are produced from the same mycelium, as is the case with the Chilian species, Puccinia berberidis. Uromyces scrophularie. (D. C.) cidiospores—Pseudoperidia on yellowish spots, in roundish cir- cinate clusters, generally hypophyllous, with rather prominent, yellowish-white, erect or sometimes inverted, entire edges. Spores rounded, polygonal, finely verrucose, 17-30 in diameter. Teleutospores—Sori frequently intermingled with the accompanying zecidia, circinate, confluent or following the venation, small, round or elliptical. Spores round, obovate or oblong, or subpyriform, with much-thickened, often conically attenuated summits, chestnut-brown, smooth, 20-35 X 10-20. Pedicels rather long, deciduous. Synonyms. Uromyces scrophulane (D. C.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 15. Abcidium scrophularia, WD. C., “ Flore frang.,” vol. vi. p. 91. Cooke, ‘‘ Hdbk.,” p. 544; “ Micro. Fungi,” 4th edit., p. 199. Uromyces concomitans, B. and Br. Cooke, “ Micro. Fungi,” 4th edit., p. 213. Uromyces scrophularia, Lév. Cooke, “ Grevillea,” vol. vii. p. 136; ‘Micro. Fungi,” 4th edit., p. 213. Puccinia scrophularia, Lév. Cooke, “ Hdbk.,” p. 497. Exsiccatt, Cooke, i. 209 ; il. 82, 637. Vize, ‘‘ Micro. Fungi Brit.,” 41. On Scrophularia nodosa. May to October. BioLoGy.—The mycelium causes considerable distortions of the stem and petioles ; the spots are yellowish, usually surrounded by a purplish margin. In this species the same mycelium which produces the ewcidiospores also gives rise to the teleutospores. 140 British Uvedinee and Ustilaginez. Uromyces ervi. (Wallr.) Ecidiospores—Scattered over both surfaces of the leaves and on the stems. Pseudoperidia cylindrical or abbreviated, with torn white edges. Spores subglobose, orange, often polygonal, 10-12 in diameter. Teleutospores--Sori oval, elliptical or linear, shining black, per- sistent, erumpent, mostly cauline, Spores ovate or subglobose, smooth, brown, apex thickened and darker, rounded or flat- tened, often with a blunt conical incrassation, ro-12 high 30-35 X 15-254. Pedicels brown, long, firmly attached, stout. Synonym. Acidium ervi, Wallr., “ Flor. Crypt. Germ.,” vol. ii. p. 247. On Vicia hirsuta (Ervum hirsutum, L.). Ecidiospores, May to June and August to October ; teleuto- spores, from July, lasting on the dead stems through the winter. BIOLOGY.—Intermixed with the teleutospores occasionally are. found a few ovate, pale brown uredospores, 20 X 104, This species morphologically closely resembles U. orobz, faba, and pisz; but I found, in 1886, that the germinating teleutospores, when placed on young plants of Prsum sativum, Faba vulgaris, and a vetch seedling, produced no effect (Exp. 623, 645, 646, 647). In 1888, no result was obtained on Victa cracca, sativa, and Lathyrus Drgtenes; but only upon Ervum hirsutum (Exp. 813 to 827). V. MICRUROMYCES. Schrot. Having only teleutospores, which germinate after a period of rest. Rarely there are found hidden between the teleutospores a few solitary uredospores. Uromyces ficariz. (Schum.) Teleutospores— Sori numerous, amphigenous, in more or less rounded clusters. Spores mostly ovoid, obovate, or pyri- form, smooth, brown; the apex of each has usually a wart- like, pale brown papilla, 25-44 x 16-26. Pedicels short, colourless, Micruromyces. 141 Synonyms. Uromyces fcarie (Schum.). Winter in Rabh., “ Krypt. Flor.,” vol. i p. 141. Uredo ficaria, Schum, “Enum. Plant. Seell.,” pt. ii p. 232. Grev., “ Flor. Edin.,” p. 434. Johnst., “ Flor. Berw.,” vol. ii. p. 203. Berk., “ Eng, Flor.,” vol. v. p. 380. Uromyces ficaria, Lév. Cooke, “ Hdbk.,” p. 518; ‘Micro. Fungi,” 4th edit., p. 212, t. vii. figs. 156, 157; “ Grevillea,” vol. vil. p. 134. Lexsiccat, Cooke, i. 122; li. 145; “L. F.,” 24. Vize, “ Fungi Brit.,” 41. On Ranunculus ficaria, L. April to June. BioLocy.—The sori occur upon pale spots on the leaves and stems ; on the latter, they cause elongated distortions. This species was at one time considered to be the teleutospores of £céadium ficaria, until Schroter (Cohn’s “ Beitrage,” vol. ii. pt. iii. p. 63) worked out the life- history of the Ranunculus Acidia. Uromyces scillarum. (Grev.) Teleutospores—Sori amphigenous, brown, arranged more or less concentrically, sometimes irregularly, becoming confluent, especially in the centre of the larger groups, at first covered with the cuticle. Spores roundish, ovate, or elliptical, with a uniformly thick, smooth, brown epispore, apex rounded or flattened, 20-30 X 15-20,4.. Pedicels short, slender, deciduous. Synonyms. Uromyces scillarum (Grev.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 142. Uredo scillarum, Grev., “Flor. Edin.,” p. 376. Berk., “Eng. Flor.,” vol. v. p. 376. Uromyces concentrica, Léy. Cooke, “‘ Hdbk.,” p. 519. Oromyces concentricus, Lév. Cooke, “Grevillea,” vol. vii. p. 138; “Micro. Fungi,” 4th edit., p. 213. Puccinia scillarum, Baxt., Exs. 40. 142 British Uredinee and Ustilaginee. On Scilla bifolia and nutans. May and June. BIOLOGY.—The presence of the mycelium causes the sori to be produced on paler discoloured spots on the leaves and stems. Uromyces ornithogali. (Wallr.) Teleutospores—Sori dark brown, scattered, elliptical or oblong, on the smaller leaves causing various distortions, often confluent, at first covered by the epidermis, then pulverulent. Spores elliptical or subpyriform, sometimes smooth, sometimes rough, pale or chestnut-brown, having a colourless wart-like point above, rounded or attenuated below, 25-40 x 17-224. Pedi- cels long, slender, deciduous. Synonyms. Uromyces ornithogali (Wallr.). Winter in Rabh., “ Krypt. Flor.,” vol. i p. 141. Erysibe rostellata, var. orntthogalé. Wallr., “Flor. Crypt. Germ.,” vol. il. p. 209. Uromyces ornithogali, Lévy. Cooke, ‘‘Grevillea,” vol. vii. p. 138. On Gagea lutea, L. B1oLoGy.—The presence of the mycelium causes variously shaped pale spots on the affected leaves. Uromyces urticz. Cooke. Teleutospores—Subpyriform, apiculate, pale, 30 x 184. Epispore thickened, on hyaline pedicels. Synonym. Uromyces urtice. Cooke, “ Grevillea,” vol. vii. p. 137. On Ortica dioica, Shere, Surrey. This species has never been found but once. It is remarkable that no one except Dr. Cooke should have met with it, seeing how common the host-plant is. A uteupuccinia. 143 VI. LEPTUROMYCES. | Schrot. Ilaving only teleutospores, which germinate as soon as they are mature upon the living host-plant. PUCCINIA. Pers. Teleutospores separate, pedicellate, produced in flat sori, con- sisting of two superimposed cells, each of which is provided with a germ-pore. The superior cell has its germ-pore, as a rule, piercing its apex; in the inferior the germ-pore is placed laterally, immediately below the septum. I, EUPUCCINIA. Schrot. Having spermogonia, zcidiospores, uredospores, and teleutospores; the latter germinating only after a period of rest. A. AUTEUPUCCINIA, De Bary. Spermogonia, ecidiospores, uredospores, and teleutospores on the same host-plant. Puccinia galii. (Pers.) icidiospores—Pseudoperidia scattered or clustered in irregular groups, edges torn, whitish. Spores roundish or shortly ellipti- cal, orange-yellow, smooth, 16-23 in diameter. Uredospores—Sori reddish brown, round or oval, often confluent. Spores globose, oval, or ovate, echinulate, pale brown, 20-30 xX 17-22h. Teleutospores—Sori black, roundish, persistent. Spores elliptical, oblong, or clavate, base attenuated, apex much thickened (9-10), often obliquely conical, constriction slight, brown, smooth, 30-55 X 15-254. Pedicels rather long, brown. Synonyms. Puccinia gali(Pers.). Winter in Rabh., “ Krypt. Flor.,” vol. i. Pp. 210. Ecidium galit, Pers. “Syn.,” p. 207. Cooke, “ Hdbk.,” p- 540; “Micro. Fungi,” 4th edit., p. 196, t. ii. figs. 15-17. Lrichobasis galt, Y.év. Berk., “ Outl.,” p. 332. Cooke, “ Hdbk.,” p. 501; “ Micro. Fungi,” 4th edit., p. 226. 144 British Uredinee and Ustilaginee. _ Puccinia galiorum, Link. Berk., ‘Eng. Flor.,” vol. v. p. 366. Cooke, “‘Hdbk.,” p. 501; “Micro. Fungi,” 4th edit, p. 208, t. viii. figs. 172, 173. Puccinia difformis, Fckl. Cooke, “ Hdbk.,” p. 501; “ Micro. Fungi,” 4th edit., p. 208. E-xstccatt. Cooke, i. 9, 72, 1133; i. 318, 325, 575. Vize, “ Fungi Brit.,” 82, 229; ‘ Micro. Fungi Brit.,” 110. On Asperula odorata, Galium cruciata, aparine, uliginosum, palustre, verum, mollugo. BroLocy.—The presence of the mycelium in the stems, especially in G. afarine, causes considerable swellings and distortions. Puccinia asparagi. D. C. A%cidiospores—Pseudoperidia in elongated patches upon the stems and larger branches, short, edges erect, toothed. Spores orange- yellow, round, very finely echinulate, 15—26y in diameter. Uredospcres—Sori brown, flat, small, long covered by the epider- mis. Spores irregularly round or oval, clear brown, echinu- late, 17-25 X 20-30. Teleutospores—Sori black-brown, compact, pulvinate, elongate or rounded, scattered. Spores oblong or clavate, base rounded, apex thickened, darker, central constriction slight or absent, deep chestnut-brown, 35-50 X 15-254. Pedicels persistent, colourless or brownish, as long as or longer than the spores. Synonym. Puccinia asparagi. D. C., ‘‘ Flore frang.,” vol. ii. p. 595. Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 201. Grev., “ Flor. Edin.,” p. 429. Berk., “Eng. Flor,” vol. v. p. 363. Cooke, “ Hdbk.,” p. 494; “ Micro. Fungi,” 4th edit. p. 203. Lixsiccati. Cooke, i. rrr. Vize, “ Fungi Brit.,” 113. On Asparagus officinalis. April to October. Auteupuccinia. 145 Puceinia thesii. (Desv.) cidiospores—Pseudoperidia scattered and thickly crowded over the whole plant, cylindrical, whitish, with torn edges. Spores nearly spherical, orange-yellow, smooth, 17-26 X 12-17. Uredospores—Sori reddish brown, round, subconfluent. Spores round or shortly elliptical, brown, echinulate, 20-284 in diameter. Teleutospores—Sori compact, round or elongate, brownish black. Spores elliptical or ovate, central constriction slight or absent, base rounded, apex thickened and darker, smooth, brown, 30-40 X 14-284. Pedicels brown, persistent, very long. Synonyms. Puccinia thesit (Desv.). Winter in Rabh., “Krypt. Flor.,” vol. 1. p. 202. Atcidium thesit, Desv., Jour. de Bot., vol. ii. p. 311. Cooke, “ Hdbk.,” p. 537; ‘‘ Micro. Fungi,” 4th edit., p. 195, t. iii. figs. 50-51. B. and Br., Aun. Nat. Hist, No. 1048. Puccinia thesit, Chail. Cooke, “ Hdbk.,” p. 495; “ Micro. Fungi,” 4th edit., p. 204. Lxsicati. Berk., 318. Cooke, ii, 311. Vize, “Fungi Brit.,” 12, 81; “Micro. Fungi Brit.,” 214, 457. On Thesium humifusum. April to October. Puccinia calthe. Link. cidiospores—Pseudoperidia on roundish spots, or when cauline on long swellings, flat, with white torn edges. Spores sub- globose, finely verrucose, orange, 20-30 in diameter. Uredospores—Sori small, round, chestnut-brown, soon scattered. Spores globose or elliptical, echinulate, brown, 22-30 x 20-2 5p. Teleutospores—Sori small, black, pulverulent, but persistent. Spores oblong, attenuated towards both extremities, rarely rounded, central constriction little or none, apex conical or with a paler wart-like papilla, smooth, brown, 30-44 X 13-22¢. Pedicels persistent, rather long, L 146 British Uredinee and Ustilaginea. Synonyms. Puccinia calthe, Link., “Sp. Plant,” vol. vi. pt. ii. p. 79. Winter in Rabh., “Krypt. Flor.,” vol. i. p. 216. Berk., “Eng. Flor.,” vol. v. p. 367. Cooke, “ Hdbk.,” p. 504; “ Micro, Fungi,” 4th edit, p. 210, Grev., “ Flor. Edin.,” p. 367. Johnst., “ Flor. Berw.,” vol. ii. p. 196. Acidium calthe. Grev., “Flor. Edin,” p. 446. Berk., “Eng. Flor.,” vol. v. p. 371. Cooke, “ Hdbk.,” p. 539; “ Micro. Fungi,” 4th edit., p. 196. Lixsiccate, Cooke, i. 114. Vize, “ Micro, Fungi,” 159; “ Micro. Fungi Brit.,” 219. On Caltha palustris. May to September. Puccinia convolvuli. (Pers.) Li cidtospores—In circular clusters on the leaves and on elongated swellings on the stems. Pseudoperidia cup-shaped, with broad, recurved, torn white edges. Spores polygonal, finely verrucose, pale yellow, 17-26 X 25-30m. Uvedospores—Sori scattered or circinate, often confluent, brown, soon naked. Spores subglobose, rarely ovate, echinulate, pale brown, 22-26 xX. 25-30. Teleutospores —Sori scattered, or cireinate, dark brown, long covered by the epidermis, sometimes confluent. Spores of two kinds: (1) Teleutospores proper, oblong, oval, or sub- clavate, apex truncate, slightly thickened, or attenuated, or much, thickened, base rounded or slightly attenuated, deep brown, 38-66 X 20-304. Pedicels stout, brownish. (2) Mesospores, generally ovate, apex much thickened, truncate, or attenuated, brown, 25-35 xX 20-25p. Pedicels pale brown, persistent. Synonyms. Puccinia convolvuli (Pers.), Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 204. Credo beta, vax. convolvuli. Pers., “‘Syn.,” p, 221. On Convolvulus sepium, Miss Jelly. June to October Auteupuccinia. 147 Puccinia gentianz. (Strauss.) A:cidiospores—Pseudoperidia on circular or elongated brownish spots, flat, with white torn edges. Spores globose, orange, finely verrucose, 16-23 X I4-I7p. Uredospores — Sori roundish, scattered or circinating, at first covered by the epidermis, chestnut-brown. Spores elliptical, ovate, or obovate, brown, echinulate, 20-30 X I9-24m. Teleutospores—Sori small, blackish, pulverulent. Spores elliptical or ovate, rounded at both ends, median constriction slight or absent,. smooth, thickened above, 28-38 x 20-254. Pedicels. colourless, delicate, very deciduous. Synonyms. Puctinia gentiane (Strauss). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 205. Uredo gentiane, Strauss. “ Wetter Ann.,” vol. ii. p.. Loz. W. G. Smith, Gard. Chron., Sept. 19, 1885, p. 372, fig. 82. On Gentiana acaults. Kew Gardens. Puccinia silenes. Schrot. icidiospores—Pseudoperidia on pale yellowish spots, in small clusters,. small,. white, shortly cylindrical, edges torn. Spores subglobose, granular, orange-yellow, 17-26 X 1420p. Uvedospores—Sori brown, roundish, scattered or circinating, often confluent. Spores roundish or elliptical, pale brown, echinu- late, 20-26 X 17-20pm. Teleutospores—Sori small, pulverulent.. Spores elliptical or oblong, very slightly constricted, rounded at both ends, apex slightly thickened, smooth, chestnut-brown, 25-40 X 16-264. Pedicels short, deciduous. Synonyms. Puccinia silenes, Schrot. Winter in Rabh., “ Krypt.. Flor.,” vol. i. p. 215. Puccinia lychnidearum. ¥ckl.,.“Symb.,” p. 50, p70 parte. Puccinia silenes, Rabh.. Cooke, “Micro. Fungi,” 4th edit., Pp. 211. On. Silene inflata. 148 British Uvredinee and Ustilaginee. Puccinia porri. (Sow.) Acidiospores—Pseudoperidia in linear or circinate clusters, shortly cylindrical, with everted torn edges. Spores polygonal, finely verrucose, membrane colourless, contents orange, 19-28m in diameter. Uredospores—Sori reddish brown, scattered or in elongate clusters, linear or oblong. Spores roundish, shortly elliptical, very finely echinulate, orange-yellow, 20-27 X 25-30. Teleutospores—Sori small, bluish grey from the dark spores being covered by the semitransparent epidermis. Spores clavate or oblong, central constriction slight, generally attenuated towards the stem, apex rounded or truncate, smooth, brown, 30-45 X 20-26, Pedicels long, but deciduous. Mesospores numerous, unicellular on long deciduous pedicels, often irregular in form, sometimes thickened above, 22-36 x 17-2 3p. Synonyms. Puccinia porri (Sow.). Winter in Rabh., ‘Krypt. Flor.,” vol. i. p. 200. Credo porrit. Sow., “ Eng. Fungi,” t. 411. Uredo alliorum, D.C. Berk., “ Eng. Flor.,” vol. v. p. 376, in part. Cooke, “Micro. Fungi,” 4th edit., p. 217, in’ part; “ Hdbk.,” p. 528. Uromyces alliorum. Cooke, ‘‘ Hdbk.,” p. 518 (?). Puccinia mixta, Fckl. Vize, Exs. Exstccatt, Cooke, ii, 425. Vize, “Fungi Brit.,” 38; “Micro. Fungi Brit.,” 430. ‘ On Allium cepa, sthenoprasum. May to August. Puccinia prenanthis. (Pers.) A cidiospores—Pseudoperidia in circular or elongate patches, hemispherical or shortly conical, opening above ‘by small A uteupuccinia. 149 irregular clefts without typical pseudoperidium. Spores rounded, pale orange, densely verrucose, 15-25 X 12-20p. Uredospores—Sori reddish brown, scattered or circinate, small, rounded. Spores spherical, pale yellow, finely echinulate, with very remarkably thickened margins to the three germ- pores, as seen in the moist state, 16-23 in diameter. Teleutospores—Sori round, blackish, dusty. Spores elliptical or ovate, central constriction slight, extremities rounded, apex scarcely thickened, finely punctate, brown, 25-35 x 20-25m. Pedicels deciduous, colourless, short. Synonyms. Puccinia prenanthis (Pers.) Winter in Rabh., “ Krypt. Flor.,” vol, i, p. 208. Schrot., Krypt. Flor. Schl.,” vol. iil. p. 318. 4icidium prenanthis. Pers., “Symb.,” p. 208 Cooke, “Hdbk.,” p. 542; “Micro. Fungi,” 4th edit, p. 198. Grev., Flor. Edin.,” p. 444. Johnst., “ Flor. Berw.,” vol. il. p. 205. Acidium compositarum, Mart. in part. Berk., ‘Eng. Flor.,” vol. v. p. 370. Puccinia chondrille, Corda. Vize, Exs. Exstccatt. Vize, ‘ Micro. Fungi Brit.,” 211. On Lactuca muralis. April to September. Some of the teleutospores are as much as 4op in length. Puccinia lapsane. (Schultz.) Ai cidiospores—Pseudoperidia flat, amphigenous, with torn, white reflexed edges, on purple, irregular spots. Spores ovate or subrotund, smooth, yellow, about 18 in diameter. Uvedospores—Sori orbicular, pulverulent, very numerous, often confluent, cinnamon-brown. Spores subglobose or ovate, brown, finely echinulate, 18-21 x 15-17p. Teleutospores—Sori small, flattish, black, pulverulent, numerous. Spores ovate or roundish, obtuse at both extremities, slightly (if at all) constricted, punctate, dark brown, 23-28 x 18-21u Pedicels hyaline, short, often oblique. 150 British Uvedinee and Ustilaginee. Synonyms. Ecidium lapsane. Schultz, “ Prod. Flor. Starg.,” p. 454. Puccinia lapsane, Fckl. Schrot., “Krypt. Flor. Schl.,” p. 318. Cooke, ‘ Micro. Fungi,” 4th edit., p. 207. cidium lapsane, Purton. Cooke, ‘“ Hdbk.,” p. 543; “* Micro. Fungi,” 4th edit., p. 198. Aicidium prenanthis, Pers. Cooke, “ Hdbk.,” p. 542; ‘ Micro. Fungi,” 4th edit., p. 198. Trithobasis lapsane, Fckl. Cooke, “Micro. Fungi,” 4th edit., p. 224. Exsiccati. Cooke, i. 13; ii. 91, 92. Vize, “ Micro. Fungi,” 162, 169. On Lapsana communis, Crepis paludosa. 4icidiospores, March to April; uredospores, April to June; teleutospores, June to August. BIOLOGY.—This species is said by Schréter to occur on Crepzs paludosa. It is certainly distinct from Puccinia variabil’s and taraxact on Taraxacum, as neither the zcidiospores, uredospores, nor teleuto- spores of P. /afsan@, when applied to Taraxacum in duplicated ex- periments made in 1885, had any effect (Exp. 481, 497, 499, 500), although they readily enough infected plants of Lapsana communis. The species which occurs so frequently on the Composite is without zecidiospores. Puccinia variabilis. (Grev.) cidiospores—Spots very small, purple, scattered over the under surface of the leaves, from 2 to 5 mm. across. Pseudoperidia often single, cup-shaped, with torn whitish edges. Spores sub- globose or oval, hyaline, with orange contents, echinulate, 20-25 X 15-20, Uredospores \ Sori small, round or elongate, dark brown, soon ee naked, containing both spore-forms. Uredospores scanty, subglobose, often irregular, brown, echinulate, 20-25 in diameter. Teleutospores oval, oblong, often sub- globose, frequently distorted in various ways, -constriction almost none, dark brown, finely verrucose, 28-30 x 18-20p, Pedicels hyaline, deciduous, of variable length. A uteupuccinia. 151 Synonyms. Puccinia variabilis. Grev., “Scot. Crypt. Flor.,” t. 75; ‘“ Flor. Edin.,” p. 431. Johnst., “Flor. Berw.,” vol. ii p. 196. Berk., “Eng. Flor.,” vol. v. p. 365. Cooke, “ Hdbk.,” p. 500; ‘ Micro. Fungi,” 4th edit., p. 207, t. 4, figs. 82, 83. Ecidium taraxact. Grev., “ Flor. Edin.,” p. 444. cidium grevillei. Grove, Jour. of Bot., May, 1886. On Zaraxacum officinale, July to October. BIOLOGY.—Mr. H. T. Soppitt met with this AZcidium at Grassing- ton, in Yorkshire, and found, by applying the zcidiospores to healthy plants of Taraxacum officinale, the uredospores and teleutospores were produced in about fourteen days. Greville’s description of the Aicidium is clear : “ Spreading over the whole leaf, and generally collected into numerous little clusters, with single ones scattered between them.” The Acidium to Puccinia sylvatica, which also occurs on Taraxacum, has its pseudoperidia not scattered but collected into large clusters. In July, 1888, I found the zcidiospores of P. vardadbziis at Watlington, near King’s Lynn, and successfully repeated Mr. Soppitt’s experiment. Puccinia pulverulenta. Grev. Aicidiospores—Pseudoperidia flat, scattered over the whole surface of the leaves. Spores subglobose, finely verrucose, orange- yellow, 16-26 in diameter. Uredospores—Sori scattered or circinating, often confluent, pul- verulent. Spores elliptical or ovate, echinulate, brown, 20-28 X 15-25p¢- Teleutospores—Sori dark brown, often concentric, pulverulent. Spores elliptical or oblong, centrally constricted, apex thick- ened, hooded, base generally rounded, smooth, brown, 24-35 x 16-204. Pedicels colourless, deciduous. Synonyms, Puccinia epilobii tetragoni (D. C.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 214. Uredo vagans, var. epilobit. D. C., “Flore frang.,” vol. ii, p. 228. 152 British Uvredinee and Ustilaginee. cidium epilobii, D. C: Berk., “Eng. Flor.,” vol. v. p. 372. Cooke, “Hdbk.,” p. 536; ‘Micro. Fungi,” 4th edit. p. 195. Grev., “Flor. Edin.,” p. 444. Johnst., “ Flor. Berw.,” vol. i. p. 204. Uredo epilobii, D.C. Purton, “ Midl. Flor.,” vol. iii., No. 1604. Johnst., ‘Flor. Berw.,” vol. ii. p. 200. Berk., “Eng. Flor.,” vol. v. p. 381. Trichobasis epilobit. Berk., “Outl.,” p. 333. Cooke, “‘ Micro. Fungi,” 4th edit., p. 226. Puccinia pulverulenta, Grev., “Flor. Edin.,” p. 432. Berk., “‘Eng. Flor.,” vol. v. p. 368. Cooke, ‘‘ Hdbk.,” p. 507; “‘ Micro. Fungi,” 4th edit., p. 211, t. iv. figs. 78, 79. E-xsiccatt. Berk., 108, 349. Cooke, 1. 4, 49; ii. 80; “L. F.,” 52. Vize, “ Fungi Brit.” 19, 79; “Micro. Fungi Brit.,” 36, 215. On L£pilobium hirsutum, montanum, tetragonum. BroLoGy.—The mycelium of the zcidiospores is diffused through the greater part of the affected plant, but whether it be truly perennial I cannot say. The mycelium of the uredospores and teleutospores is strictly localized. I found in June, 1882, that the ecidiospores sown on seedlings of ,Z. hirsutum gave rise to ecidiospores in seventeen days. Puceinia viole. (Schum.) Acidiospores—Pseudoperidia on the leaves in circular concave patches, often causing much distortion on the stems, flat, with white torn edges. Spores subglobose, finely verrucose, orange- yellow, 16-24 X to-18p. Uvedospores—Sori brown, small, roundish, scattered, soon naked. Spores roundish or elliptical, brown, echinulate, 20-26 in diameter. Teleutospores—Sori black, roundish, small, pulverulent. Spores elliptical or oblong, slightly attenuated at the base, with an apical thickening, constriction almost absent, brown, 20-35 X 15~20u. Pedicels long, deciduous. A uteupuccinia. 153 Synonyms. fuccinia viole (Schum.). Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 215. Grev., ‘Flor, Edin.,” p. 432. Johnst., ‘ Flor. Berw.,” vol. vi. p. 196. cidium viole, Schum. ‘ Enum. Plant Szll.,” vol. ii. p. 224. Grev., “Edin. Flor.,” p. 444. Berk., “Eng. Flor.,” vol. v. p. 372. Johnst., “Flor. Berw.,” vol. ii. p. 205. Cooke, “ Hdbk.,” p. 543; “Micro. Fungi,” 4th edit., p. 198. Credo violarum, D.C. Berk., “Eng. Flor.,” vol. v. p. 380. Sow., t. 440. Johnst., ‘‘ Flor. Berw.,” vol. ii. p. 202. Trichobasis violarum, Berk. Cooke, “‘ Micro. Fungi,” 4th edit., p. 226. Puccinia violarum, Link. Berk., “Eng. Flor.,” vol. v. p. 367. Cooke, “ Hdbk.,” p. 505; “Micro. Fungi,” 4th edit., p. 210. Lxsiccats. Berk., 223, 228. Cooke, i. 46, 49, 104; ii. 135; “L. F.,” 43. Wize, “ Micro, Fungi Brit.,” 57-112; ‘‘ Fungi Brit.,” 77. On Viola canina, odorata, sylvatica, hirta, tricolor. May to October. Puccinia albescens. (Grev.) cidiospores—Pseudoperidia short, whitish, with toothed edges, on bleached places on the stems, petioles, leaves, and blossoms. Spores roundish, colourless, 15-224 in diameter. Mycelium diffused, perennial. Oredospores—Sori brown, small, round or oblong, long covered by the epidermis. Spores oval or elliptical or subglobose, nucleate, echinulate, clear brown, 20-25 X 15-20. Teleutospores—Sori small, round, scattered. Spores often in the same sori as the uredospores, rarely confluent, long covered by the epidermis, fusiform or elliptical, attenuated at both ends, often surmounted by a colourless papilla, constriction slight, smooth, brown, 34-40 X 18-25. Pedicels short, hyaline, deciduous. Synonym, cidium albescens. Grev., “Flor. Edin.,” p. 444. Berk., “Eng. Flor.,” vol. v. p. 372. Johnst., “Flor. Berw.,” vol. ii 154 British Uredinee and Ustilaginee. p. 205. Cooke, “ Hdbk.,” p. 536; ‘Micro, Fungi,” 4th edit., Pp. 194. Exsiccatt, Cooke, i. 636; ii. 78. Vize, “Fungi Brit,” 164. “ Micro. Fungi Brit.,” 561. On Adoxa moschatellina. March to May. B1oLocy.—The mycelium of the zcidiospores is perennial, that of the uredospores and teleutospores annual. Schriter found that when the zecidiospores were sown on healthy plants, a small quantity of the uredospores were produced. I have grown plants affected with the zecidiospore mycelium in a flower-pot, which for three successive years came up affected with the A:cidium. Mr. Soppitt produced the uredo- spores and teleutospores in June, 1888, from the eecidiospores. He states that the teleutospore sori remains longer covered by the epidermis than those of P. adoxe do. Puccinia bupleuri. (D. C.) A cidiospores—Pseudoperidia uniformly scattered over the whole leaf-surface, flat, with torn white edges. Spores polygonal, nearly smooth, 14-21 in diameter. Uredospores—Sori scattered or circinating, small, roundish or irregular. Spores very few, subglobose, verrucose, yellowish brown, 17-23 in diameter. Teleutospores—Sori numerous, scattered, dark brown, oblong, surrounded by the ruptured epidermis, rather small. Spores broadly elliptical or oblong, apical thickening slight or absent, rounded at both ends, broadly constricted, smooth, dark brown, 26-42 X 17-304. Pedicels long, deciduous. Synonyms. icidium falcaria, var. Bupleurt falcati. D.C., “ Flore frang.,” vol. vi. p. 91. Puccinia Bupleuri falcati (D. C.). Winter in Rabh., “‘ Krypt. Flor.,” vol. i. p. 212, Cooke, ‘‘Grevillea,” vol. xvi. p. 47. On Bupleurum tenuissimum. Walton-on-the-Naze. August, 1887. Auteupuccinia. 155 Puccinia pimpinelle. (Strauss.) cidiospores—Pseudoperidia mostly on distortions of the stems, at first hemispherical, then rounded or elongate, edges slightly torn, Spores roundish, yellow, finely verrucose, 18-35 x 16-21. Uredospores—Sori chestnut or cinnamon brown, rounded or elon- gated, pulverulent. Spores rounded, shortly elliptical or pyriform, pale brown, echinulate, 23-32 X 20-24. Teleutospores—Sori blackish, rounded or elongated, pulverulent, often on the stems, which are then much distorted. Spores elliptical or ovate, scarcely constricted, rounded at both extremities, membrane reticulated with broad net-like ridges, chestnut-brown, 25-35 X 17-26u. Pedicels long, deciduous, colourless. Synonyms. Puccinia pimpinelle, Strauss. Winter in Rabh., ‘‘ Krypt. Flor.,” vol. i.’ p. 212. Credo pimpinella, Strauss. “ Wett. Ann.,” vol. il. p. 102. cidium bunti, D.C. Grev., “ Flor. Edin.,” p. 445. Berk., “Eng. Flor.,” p. 370, in part. Acidium pimpinelle, Kirch. Cooke, “Micro. Fungi,” 4th edit., p. 196. Uredo heraclei, Grev. Berk., “ Eng. Flor.,” vol. v. p. 380. Trichobasis heracte. Cooke, “Hdbk.,” p. 502; “Micro. Fungi,” 4th edit., p. 225. Trichobasis angelica, Schum. Cooke, “ Micro. Fungi,” 4th edit., p. 224. Trichobasis pimpinella, Strauss. Cooke, “‘ Micro. Fungi,” 4th edit., p. 224. Puccinia umbelliferarum, D.C. Grev., “Flor. Edin.,” p. 431. Berk., “Eng. Flor.,” vol. v. p. 316. Johnst., “ Flor. Berw.,” vol. ii. p. 196. Cooke, “‘ Hdbk.,” p. 501, in part. Puccinia cherophyllt. Purton, ‘ Midl. Flor.,” vol. iii., No. 155 3. Puccinia heracled, Grev., “Scot. Crypt. Flor.,” t. 42. Cooke, “ Hidbk.,” p. 502; “ Micro. Fungi,” 4th edit., p. 208. Puccinia angelice, Fckl, Cooke, “Micro. Fungi,” 4th edit., p. 208. 156 British Uredinee and Ustilaginee. Puccinia pimpinelle, Link. Cooke, ‘“ Micro. Fungi,” 4th edit., Pp. 209. LExstccatt. Vize, “ Micro. Fungi Brit.,” 124, 425. On Pimpinella saxtfraga, Heracleum sphondylium, Anthriscus sylvestris, Myrrhis odorata. May to October. Puceinia apii. (Wallr.) Acidiospores—Spots rounded or irregular, yellow above, paler below, causing elongated yellow swellings on the stems. Pseudoperidia subcylindrical, with torn, white, everted edges. Spores subglobose, orange-yellow, finely echinulate, about 25. in diameter. Uredospores—Sori rather large, cinnamon-brown, pulverulent, round or elongate, scattered, sometimes circinating. Spores globose, oval, or subpyriform, pale brown, finely verrucose, 30-40 X 20-30, Teleutospores—Sori rounded, elongated, or irregular, long covered by the epidermis, blackish brown. Spores oval or clavate, smooth, dark brown, scarcely constricted, apex not thickened, 30-50 X 15-204. Pedicels hyaline, rather persistent, not very short. Synonyms. Oredo apii. Wallr., “Flor. Crypt. Germ.,” vol. ii. p. 203. Puccinia apit. Corda, “Icones,” vol. vi. p. 30, fig. 11. Cooke, “ Hdbk.,” p. 502; “ Micro. Fungi,” 4th edit., p. 208. YOR3 LExsiccatt. Cooke, i. goa. Vize, “ Fungi Brit.,” 54, 127 ; “ Micro. Fungi Brit,” 554. On Apium graveolens. May to September. BioLocy.—The aecidiospores occur at the latter part of May, and are by no means profuse, although the yellow spots on which they occur are very conspicuous. On May 27, 1888, I applied the ecidio- spores and uredospores freely to a plant of Prmpinella Saxtfraga in my A uteupuccinia. 157 garden, but without effect. This is the only Uredine affecting the Umbelliferze with which I have made any experimental cultures, and I have therefore followed Winter and Schréter in grouping the various spore-forms together under Puccinia pimpinell@ and bullata; but it is probable that biological research will show several of them to be good species, Puccinia menthe. Pers. cidiospores—Pseudoperidia immersed, flat, opening irregularly, edges torn; principally on the stems, which are much swollen, more ‘rarely on concave spots on the leaves. Spores sub- globose or polygonal, coarsely granular, pale yellowish, 17-26 x 26-35p. Uredospores—Sori small, roundish, soon pulverulent and confluent, cinnamon-brown. Spores irregularly rounded or ovate, echinu- late, pale brown, 17-28 X r4—19p. Teleutospores—Sori black-brown, roundish, pulverulent, Spores elliptical, oval, or subglobose, central constriction slight or absent, apex with a hyaline or pale brown papilla, verrucose, deep brown, 26-35 X 19-234. Pedicels long, delicate, colourless. Synonyms. Puccinia menthe. Pers., “Syn.,” p. 227. Winter in Rabh., “ Krypt. Flor.,” vol. i. p. 204. cidium mentha, D.C. Berk., “ Eng. Flor.,” vol. v. p. 369. Cooke, “ Hdbk.,” p. 544; “ Micro. Fungi,” 4th edit., p. 199. Trichobasis labiatarum,Lév. Cooke, “ Hdbk.,” p. 496 ; “Micro. Fungi,” 4th edit., p. 224. Trichobasis clinopodii, D. C. Cooke, ‘‘Micro. Fungi,” 4th edit., p. 224. Puccinia clinopodii, D. C. Cooke, ‘ Micro. Fungi,” 4th edit., p. 205. Puccinia menthe, Pers. Berk., ‘‘ Eng. Flor.,” vol. v. p 364. Grev., “Flor. Edin.,” p. 430. Johnst., “Flor. Berw.,” vol. ii. p- 195. Cooke, “ Hdbk.,” p. 496; “Micro. Fungi,” 4th edit., p. 204, t. iv. figs. 69, 70. Uredo menthe. Purton, “Midi. Flor.,” vol. ii, No. 1550. Sow., t. 398, fig. 3, sub Aicidium, 58 British Uredinee and Ustilaginee. Uredo labiatarum, D. C. Grev., “Flor. Edin.” p. 436. erk., “Eng, Flor.” vol. v. p. 378. Johnst., ‘Flor. Berw.,” ol. i, p. 203. LExsiccatt. Berk., 217, 232, 233. Cooke, i. 395 ii: 33,60, 444. Vize, Fungi Brit,” 14, 15, 75; “ Micro. Fungi Brit.” On Mentha rotundifolia, sylvestris, aquatica, arvensis, viridis, wiganum vulgare, Calamintha clinopodium. May to October. BIoLoGy.—The ecidiospore mycelium: is diffused. and probably. srennial —at any rate, lasting for more than one year ;, at least, this ypears to be the case with AZ, wirddis, which I. have cultivated for a. 2riod of three years. The mycelium of the uredospores and teleuto- dores is purely local. Johnoston records this species on Ajuga reptans. Pucecinia zgra. Grove Ecidtospores—Pseudoperidia on all: green. parts. of the plants, scattered, not collected on. swollen roundish or elliptical spots, with torn, white, somewhat recurved margins. Spores roundish or oblong, angular, smooth, orange-yellow, 17-21 X 14-16p. 7redospores—Sori numerous, amphigenous, on yellow spots, not very small, scattered or collected in groups, roundish, flatly onvex, covered with the silvery, shining, persistent epidermis. Spores elliptical or obovate, finely echinulate, brown, 28-30p, ‘eleutospores—Sori resembling those of the uredospores. Spores elliptical, oblong, or roundish, very irregular, rounded or taper-~ ing at the base or apex,.sometimes truncated, smooth, not constricted, dark brown, 22-30 X 18-24. Pedicels short,. hyaline. Synonyms. Puccinia egra. Grove, Jour. of Bot., vol. xxi. (1883), p. 274, Ecidium depauperans, Vize. Cooke, “ Micro. Fungi,” 4th dit., p. 195, : Exsiccati, Vize, ‘Micro. Fungi Brit.,” 56. On Viola cornuta, lutea var. amena, May to September. A uteupuccinia. 159 BIoLocy.—The zcidiospore mycelium is perennial, and the affected plants are altered in habit, being paler in colour and more elongated in their stems and petioles. The mycelium of the uredospores and teleutospores is purely local. Puccinia primule. (D.°C.) , Czeomata, 18 3» uredospores, 19 Spore-culture, 105 formation in Doassansia, 67 35 Entyloma, 67 7 Sorosporium, 64 is Sphacelotheca, 62 4 Tilletia, 66 4 Tubercinia, 65 30 Urocystis, 68 55 Ustilago, 62 nomenclature, 74 Sporenlager, 28 Staffordshire, 50 Stahl, Dr. E., 17 Starch, 7 Stellaria holostea, 211 Sterigmata, 10, 31 Stroma, 28 Taraxacum officinale, 151, 186, 187 Teleutospores, 36 abnormal septation of, 40 absence of, 36 “development of, 37 Biological Index. gar Teleutospores— germination of, 42 mother-cell, 39 of Chrysomyxa, 42 »» Coleosporium, 41 >, Gymnosporangium, 42 5» Phragmidium, 40 »» Puccinia, 38 3, Uromyces, 37 », Xenodochus, 40 Temperature, effect on— Puccinia spores, 109 Uredo spores, 35 Ustilago spores, 104 Thalictrum flavum, 181 Thecaphora hyalina, 86 trailii, 296 Thiimen, F., 258 Tilletia decipiens, 284 lathyri, 86 strizeformis, 285 tritici, 86 Tragopogon porrifolius, 198 pratensis, 17, 198 Trail, Professor J. W. H., 126, 127, 172, 203, 208, 215, 241, 262, 284, 291, 296, 299, 300, 301 Trichobasis, 31 Trichogyne, 18 Trifolium arvense, 134 Triphragmium ulmarize, 19, 219 Triticum repens, 181 Tubercinia trientalis, 96, 294 Tuberculina, 299 Tulasne, L. R. and C., 13, 16, 72 Tull, Jethro, 47 Tussilago alpina, 217 Unger, F., 16 Uredinex, biology of, 1 haustoria, 4 mycelium, 4 localized, 5 = perennial, 7 spore-forms, 3 spore-relationships, 3 ” Uredo fabze, 30 frumenti, 47 Uredo iridis, 258 linearis, 30 miilleri, 257 phaseoli, 30 serratulz, 13 suaveolens, 12, 30, 103 symphyti, 36 Uredospores, 28 colouring matter, 31 development of, 28 germ-pores, 30 paraphyses, 33 primary, 45 quantity, 34 -resistance of cold, 35 structure, 29 Urocystis anemones, 94 colchici, 68 fischeri, 93 occulta, 93 primulicola, 95 viole, 94 Uromyces, teleutospores of, 37 Urtica dioica, 170 parvifolia, 7, 170 Ustilaginese, infection of host-plants, 99 Hoffmann’s views, 99 Jensen’s views, 103 Kiihn’s views, 99 . Wolff’s views, 100 Ustilaginez, localization of spore-beds, 64 mycelium, 58 spore-formation, 61 teleutospores, colour of, 69 x5 formation of, 61 ” germ-pores, 70 germination of, 72 resistance of tempera- ture, 104 35) structure of, 69 Ustilago bromivora, 83 cardui, 78, 296 ficuum, 85 flosculorum, 79 grandis, 83 oe” hypodytes, 80 kiihneana, 80 longissima, 81, 273 major, 84 38 Biological Index. Ustilago maydis, 79 olivacea, 84, 277 pheenicis, 85 scabiosee, 79 segetum, 74, 101, 274 » var, nuda, 274 ” var, tecta, 274 tragopogi, 80 utriculosa, 85 violacea, 78 Vicia sepium, 125 Vincetoxicum asclepiadeum, 250 Vize, Rev. J. E., 40 Waldheim, F. von, 58 Ward, Professor M., 297 Wheat-mildew, 46 Wickliffe’s Bible, 46 Wilson, Mr. A. S., 101 Withering, W., 50 Wolff, R., 249 Vaccinium vitis-idzea, 271 Vandenhecke, 72 Vicia cracca, 121, 134, 140 faba, 121, 140 Young, A., 51 sativa, I2I, 140 Xanthophyll, 31 INDEX OF SPECIES. PAGE ECIDIUM, Pers. . ‘ : . 262 albescens, Grev. . i » 153 allid, Pers. . - i . 165 aquilegiz, Pers. . » 263 argentatum, Schultz : » 193 ari, Desm. . * < - 166 asperifodit, Pers. . 2 . 167 avicularia, Kze. . , » 124 barbaree, D.C... , » 265 behenis, D. C. . , . 138 bellidis, D.C. : : » 175 berberidis, Gmel. . i . 163 bunii, D, C. . 155, 270 calthe, Grev. ‘i 5 . 146 cancellatum, Pers. . : . 231 cichoracearum, Johnst. . - 198 cirsiz, D.C. . , ‘ » 173 clematidis, D. C. . ; . 265 columnare, Alb. and Schw. . 271 compositarum, Mart. 149, 169, 171, 172, 175 confertum, Grev. . ‘ 2 32 convallarize, Schum. i‘ . 264 cornutum, Pers. . 7 » 235 crassum, Pers. 2 F » 164 cyparissia,D.C. . : « 133 depauperans, Vize . ‘ » 158 dracontii, Schw. . ‘ « 266 elatinum, Alb, and Schw. . 270 epilobit, D. C. . ° » 152 ervi, Wallr. . i e » 140 euonymi, Gmel. . j . 260 euphorbie, D.C. . ° . . 228 | BCIDIUM—continued. euphorbiz, Gmel. . #6 Pers. . Jalcaria, D. C. Jicaria, Pers. Juscum, Rehl. gali, Pers. 5 5 geranit, D. C. : 5 glaucis, Dozy. and Molk. grevillei, Grove. . grossulariz, Gmel. incarceratum, B. and Br. jacobe@, Grev. laceratum, Sow. lapsane, Purt. . >> schultz leucospermum, D. C. mentha, D. C. ‘ a mespilt, D. C. : orchidearum, Field . orobi, Pers. parnassia, Grev. . E pedicularis, Libosch, . penicillatum, Mill. periclymeni, Schum. phylliree, D.C. . ‘ pimpinelle, Kirch. . pind, Pers. poterii, Cooke prenanthis, Pers. . primule, D.C. . prunelle, Wint. . pseudo-columnare, Kiihn 149, PAGE 270 228 154 131 205 134 126 268 151 263 267 171 237 150 150 269 157 232 179 121 129 174 232 264 267 155 249 268 150 159 264 271 340 Index of Species. PAGE ECIDIUM—continued. punctatum, Pers. 268 pyrole, Gmel. 247 » * Cc. 253 quadrifidum, D. ©, . 268 ranunculacearum, D. C. 130, 131 178, 180, 266 i var. thalictri- fiavi, D. C. 181 rhamni, Gmel. - 164 rubellum, Pers. ~ . 176, 177 rubt, Sow... 3 : . 224 rumicis, Pers.. 176 >» Grey. 177 salicis, Sow. : ee 239 salicornia, D. C. 129 saniculi,Carm. . 160 scrophularia, D. C. 139 soldanellg, Hornsch. 160: sonchi, Johnst. 266 staticis, Desm. 123 strobilinum (Alb. and Stier ) 266 taraxaci, Grev. 151 thalictri, D. C. 181 thesii, Desv. : » 145 tragopog?, Pers. . + 198 tusstlaginis, Pers. 169 urtice, D. C. i 170 valerianacearum, Duby . 128 viole, Schum. 153 zonale, Duby . 132 Aregma acuminatum, Fries . 221 bulbosum, Fries 224 gracile, Grev. 227 mucronatum, Fries 225 obtusatum, Fries 225 Ascomyces trientalis, Berk. 293 AUTEUPUCCINIA 143 AUTEUUROMYCES 11g BRACHYPUCCINIA 182 BRACHYUROMYCES . 134 CHOMA, Link. 259 egirtnum, Schlecht. 242 alliatum, Link, 261 alliorum, Link. . 261 colchict, Schlecht. . 287 euonymi (Gmel.) 260 hydrocotyles, Link. 195 CHOMA—continued. hypodytes, Schlecht. laricis (West.) mercurialis, Pers. mixtum, Schiecht. blongatum, Link. : orchidis, Alb. and Schw. poteriz, Schlecht. saxifragze (Strauss. ) utriculosum, Nees . CHRYSOMYXA, Unger empetri (Pers.) pyrole (D.C.) . COLEOSPORIUM, Lév. . cacaliag (D. C.) campanulee (Pers.) euphrasiz (Schum.) miniatum (Pers.) . ochraceum, Fckl. petasitis (Lév.) pingue, Lév. . ; rhinanthacearum, Lévy. . senecionis (Pers.) . sonchi (Pers. ) sonchi-arvensis, Lév.. symphyti, Fckl. tusstlaginis . . CRONARTIUM, Fries . a flaccidum (Alb. and Schw.) paonia, Cast. ‘ DOASSANSIA, Cornu . alismatis (Nees) sagittarize (Fckl.) . ENDOPHYLLUM, Lév. euphorbiz (D. C.). sempervivi (Alb. and Schw.) . ENTORRHIZA, Weber. - cypericola, Magnus ENTYLOMA, De Bary . bicolor, Zopf. calendulze (Oud.) canescens, Schrot. . . chrysosplenii (B. and Br.) fergussoni (B. and Br.) . ficarie, F. v. Waldh. matricariz, Trail . microsporum (Unger) ranunculi (Bon.) » .254 PAGE 237 262 260 240 190 261 221 259 280 252 253 248 251 251 252 225 255: 251 225, 252 248 250 25k - 255. 251 254 254 294 294 295 228 228 229 298 299 289 296 292 290 291 289 290 291 291 290 L[ndex of Species. 341 PAGE PAGE ENTYLOMA—continued. Lycoperdon scutellatum, Schrank. . 134 ungerianum, De Bary . » 292 subcorticatum, Schrank. . 225, Epitea baryi, Be and Br. . . 192 tritict, Bjerk,. 283, Erysibe occulta, Wallr. . . 285 | MELAMPSORA, Castagne 236 rosellata, var. ornithogali, ecidioides (D. C.) . 241 Wallr. 7 ‘ ‘ . 142 betulina (Pers.) 243 subterranea, Wallr. ; - 204 cerastii (Pers. ) 247 EUCOLEOSPORIUM . 5 . 248 circaeze (Schum.) 245 EUVPUCCINIA. ‘ : . 143 epitea (Kze.. and Schum.) 239 EUUROMYCES . . % . II9 farinosa (Pers.) 236 Farinaria carbonaria, Sow. . . 276 helioscopize (Pers.) 238. scabiose, Sow. met 2 279 hypericorum (D. C.) 243 stellaria, Sow. . ‘i . 280 lini (Pers.) .. 237 Gleosporium ficaria, Berk. . . 290 mixta (Schlecht.) ‘ 239 Granularia viole, Sow. ; . 288 padi (Kze. and Schum.) . 246 GRAPHIOLA, Poit. é ‘ » 297 populina (Jacq.) 242 pheenicis (Moug.) . 7 . 298 pustulata (Pers. ) 244 GYMNOSPORANGIUM, Hedw. . 230 pyrole (Gmel.) 247 clavarizforme (Jacq). 233 salicina, Lév. . 238 confusum, Plow. . 232 salicis-caprea, Pers. 238 juniperinum (Linn.) : » 235 tremule, Tul. 240 sabine (Dicks.) . é . 230 vacciniorum, Link. 246 tremelloides, Hartig, ‘ . 236 vitellinze (D. C.) . 240 HEMICHRYSOMYXA . : . 253 | MELAMPSORELLA . 247 HEMICOLEOSPORIUM 5 . 250 | MELANOTZNIUM, De Bary 292 HEMIPUCCINIA . 5 . 188 endogenum, De Bary . 292 HEMIUROMYCES ‘ . 134 | MICROPUCCINIA A : 199 HETEROPUCCINIA . ; . 162 | MICRUROMYCES 3 és 140 HETERUROMYCES. ‘ . 130 | Ailesta polypodii, B, White . 256 Lecythea baryi, Be and Br... . 192 | Mycxosporium colliculosum, Berk. . 231 betulina, Lév. i : . 244 | Perichena strobilina, Fries 267 caprearum, Berk. . ‘ . 238 | Peridermium acicolum, Link. 249 epitea, Lév. . ; : . 239 columnare, Alb. and Schw. 271 euphorbia, Cast. . ‘ . 237 elatinum, Alb. and Schw. 271 gyrosa, Berk. . : : 227 pini, Chevall. 249 lint, Berk. . ‘ 5 . 237 | Phelonitis strobilina, Pers. 267 mixta, Lév. . ; . 240 | PHRAGMIDIUM, Link.. 220 populina, Lév. ; ‘ » 242 acuminatum, Fries 221 poterit, Lév. . 5 ‘ . 228 bulbosum, Fries 224 rose, Lév. ‘ . 225 bullatum, West. . 225 ruborum, Lévy, 4 . 223, 224 carbonarium (Schlecht.) . 227 saliceti, Lév. . : . » 240 fragariastri (D. C.) 220 valeriane, Berk. . : . 128 Susiforme, Schrot. . 226 LEPTOPUCCINIA ‘ : . 210 gracile, Berk. 227 LEPTUROMYCES - - 143 mucronatum, Fries 225 Licea strobilina, Alb. and Schw. . 267 obtusatum, Fries. . 222 Lycoperdon innatum, With. . . 269 potentillee (Pers.) . . 221, 222 populinum, Jacq. « ‘ « 242 poterti, Fekl. . . 221 Z 3 342 Index of Species. PAGE PHRAGMIDIUM—continued. rosz-alpinze (D, C.) . . 226 rubi (Pers.) . : 3 + 224 rubi-ideei (Pers.) . 3 . 226 sanguisorbze (D. C.) : oc SE subcorticatum (Schrank. ) . 224 tormentille, Fckl. . » 222 violaceum (Schultz.) - . 223 Podisoma juniperini, Fries. » 243 sabina, Fries . ; 5 & 231 PROTOMYCES, Unger . ‘< + 300 ari, Cooke. ‘ : » 301 calendule, Oud. . ‘ . 292 chrysosplenit, B, and Br. - 291 Jergussoni, B, and Br... » 290 Jicarie, Cornu and Roze. » 290 hieracit, Berk. ‘i i + 292 macrosporus, Unger. + 300 menyanthis, De Bary. =~ 301 microsporus, Unger a + 292 pachydermus, Thiim. . + 300 rhizobius, Trail . : ~ 300 Sagittarig, Fckl. . ! » 295 tuberi-socani (Mart.) 4 + 204 PUCCINIA, Pers. : ‘ » 143 acuminata, Fckl. . é » 212 adoxe, D. C. a . » 207 zegopodii (Schum.) ‘ + 201 zegra, Grove . : 4 158 @thuse, Link. ‘ 3 » 184 albescens (Grev.) . . » 153 amphibii, Fckl. . : - 188 andersoni, B. and Br. . + 204 anemones, Pers. : + 205 53 var. betonica, Alb. and Schw. a 2 + 200 angelica, Fckl, . : » 155 annularis (Strauss. ) - . 217 anomala, Rost. ‘ . » 167 anthoxanthi, Fckl. . ‘ » 194 apii (Wallr.) ‘ i 156 arenariee (Schum.) ; . 210 arenariicola, Plow. : - 170 argentata (Schultz) . + 193 arundinacea, Hedw. A - 176 asarina, Kze. .. 4 » 202 asparagi, D.C... : . 144 asteris, Duby . . < . 215 PAGE PUCCINIA—continued. avicularig, Grev. . 124 baryi (B. and Br.) . : 19 betonicze (Alb. and Schw.) 199 va D. C. 200 bistorte, D.C. .. 192 bulbocastani, Fckl. 206 bullaria, Link. 184 bullata, Pers. 183 bunii, D.C. . 206 bupleuri, D. C. 154 buxi, D.C... 217 -calthee, Link. 145 -campanulz, Carm. 200 ‘cardui, Plow. 216 caricina, Grev. 169 caricis (Schum.) . . 169, 174 caulincola (Schum.) : 201 centaurese, Mart. . . . 186 cherophylli, Purt. . 155 chondrilla, Corda . 149 chrysosplenii, Grev. 211 circaeze, Pers. . + 213, 245 carsi?, Fckl. . - 216 clandestina, Carm. 185 clinopod#, D.C. . = + 157 compositarum, Schlecht. 185. 4 Mart. . 182, 186 conit, Fckl. . z si « 184 convolvuli (Pers.) . 146 coronata, Corda 163 dianthi, D, C. 210 aifformis, Fckl. 144 dioicee, Magnus 173 discoidearum, Link. 189 epilobii, D. C. L 202 epilobit-tetragont (D. C.) 151 extensicola, Plow. 181 Jabe, Link. 120 »» Johnst. 121 Sallens, Cooke 125 fergussoni, B. and Br. 207 Sragaria, Purt. : F 221 Sragariastri, D. C. 220 Jlosculosorum (Alb. — and Schw. ) - : . 185, 187 fusca (Relh.), : + 205, 269 galii (Pers. ) . +143 Index of Species. 343 PAGE ~ PAGE PUCCINIA—continued. PUCCINIA—continued. galti-cructati, Johnst. . » 212 potentille, Grev. . : a 222: galiorum, Link. . a » 144 45 Pers... f » 222 gentianze (Strauss. ) Z - 147 prenanthis (Pers.) . ‘ . 148 glechomatis, D. C. : » 214 primule(D.C.) . : . 159 globosa, Grev. : » ¥20 pruni, Pers. . ‘ . . 192 glomerata, Grev. . + 209, 249 pruni-spinose, Pers. : . 193 gracilis, Grev. f é we 227, prunorum, Link. . , » 193 graminis, Pers. - 162 |, pulverulenta, Grev. ‘ » 151 » var. arundinacea, rhodiole, B. and Br. . . 207 Cooke ‘ 5 - . 178 vos@, D.C. . ; ‘ . 225 heraclei, Grev. ‘ ‘ - 155 rubi, Sow... . ‘ + 224 hieracii (Schum.) . ‘ - 184 rubigo-vera (D. C.) ‘ . 167 >» Mart. ‘ ‘ » 185 ruborum, D.C. . : » 224 hydrocotyles (Link.) . » 195 sagine, Fckl. ‘ : . 210 iridis (D. C.) js 2 . 189 sanguisorbe, D. C. ‘ . 221 Junct, Desm. - 3 132 sanicule, Grev. . 5 . 160 lapsanee (Schultz) . ‘ . 149 saxifragee, Schlecht. ‘ . 208 liliacearum, Duby . ‘ « 197 saxifragarum, Schlecht. . » 208 Umonii, D. C. : : » 123 schneideri, Schrot. : . 201 linearis, Rob. : - 165 scheeleriana, Plow. , « AZT « luzula, Libk ., . ‘ » IQI scillarum, Baxt. . ‘ . 142 lychnidearum, Link. 147, 196, 210 scirpi, D. C. .. ‘i ; - IQ1 magnusiana, Korn. » 177 scorodonie, Link. . 5 . 218 malvacearum, Mont. . . 212 scrophularie, Lév. . 5 . 139 menthe, Pers. : : - 157 senecionis, Lib. . , + 209 millefolii, Fckl. . 2 » 215 sessilis, Schneid. . 2 . 165 mixta, Fckl. . ‘: . - 148 stlai, Fckl. . ‘ 3 . 184 mehringia, Fckl. . ‘ . 210 silenes, Schrot. . -* . 147 moliniz, Tul. , : - 179 smyrnii, Corda. : - 199 mucronata, Pers. .. a 224 soldanelle (D. C.). A » 159 noli-tangeris, Corda 5 - 194 sonchi, Rob. . 5 . 196 oblongata (Link.) . 3 - 190 sparsa, Cooke . , . 198 obscura, Schrot. . - 174 spergule, D.C. . : . 210 obtegens, Fckl. : : » 182 spirea, Purt. - : » 219 oxyriz, Fckl. , : . 194 stellarig, Duby . 5 . 210 paliformis, Fckl. . ; + 203 straminis, De Bary ‘ - 167 paludosa, Plow. . a - 174 striola, Link. 7 F » 170 perplexans, Plow. . . - 179 suaveolens (Pers. ) . “ . 182 persistens, Plow. . : . 180 sylvatica, Schrét. . : » 172 phalaridis, Plow. : . 166 syngenesiarum, Link. . 198, 216 phragmitis (Schum.) « F76.377 tanaceti(D. C.) . : » 189 pimpinella (Strauss.) . » 155 taraxaci, Plow. . ‘ » 186 53) Link. . 5 . 156 thalictri, Chevall. . : . 206 poarum, Niel. . @ - 168 thesii (Desv.) é ‘ . 145 polygoni, Pers. . . 124, 188 tragopogi (Pers.) . s . 197 polygonorum, Link. : - 188 trailii, Plow. . ; é . 176 porri (Sow.) . , ; . 148 tripolii, Wallr. 5 a 285 344 Lndex of Species. PUCCINIA—continued. truncata, Berk. tumida, Grev. ‘ é ulmaria,D.C. . . umbelliferarum, D.C. . umbilici, Guép. vaginalium, Link. . valanti, Pers. variabilis, Grev. . veronice (Schum.) ‘ var. persistens, Korn. veronicarum, D. C. verrucosa (Schultz) vince (D. C.) violacea, Schult. violee (Schum.) violarum, Link. virgaurez (D. C.) . ” Lib. PUCCINIASTRUM PUCCINIOPSIS . Reticularia segetum, Bull. Restelia cancellata, Reb. . cornuta, Tul. . ‘ lacerata, Tul. . . SOROSPORIUM, Rudol. paridis, Unger . saponariz, Rudol . triertalis, Woron. . SPHACELOTHECA, De Bay hydropiperis (Schum. ) , Spheria flaccida, Alb. and Schw. THECAPHORA, Fing. . cirsit, Boud. . hyalina, Fing. trailii, Cooke THECOSPORA . TILLETIA, Tul. . bullata, Fckl. cartes, Tul. decipiens (Pers.) . Spherococca, F. v. Waldh. 184, 155; . 150, strizeformis (Westd.) tritici (Bjerk.) Juniperina, Linn. . sabine, Dicks, . * Trachyspora alchemilla, Fekl, Tremella clavariaformis, Jacq. 293 293 282 282 254 295 296 295 296 245 283 277 283 284 284 284 283 137 234 235 231 Trichobasis angelica, Schum. artemisia, Berk. . beta, Lévy... 3 caricina, Berk. clinopodiz, D. C. conit, Strauss. . . cynapit, D. C. epilobit, Berk, Jabe, Lév. . Sallens, Cooke galt, Lév. geranit, Berk, glumarum, Lévy. heraciei, Berk. hieracit, Schum. hydrocotyles, Cooke impatiens, Rabh, tridis, Cooke . labiatarum, Lév. lapsane, Fckl. linearis, Lév.. lychnidearum, Lév. lynchit, B. and Br. oblongata, Berk. parnassia, Cooke . petroselini, Berk. pimpinella, Strauss. polygonorum, Berk. primule, Cooke pyrole, Berk. . rhamni, Cooke rubigo-vera, Lév. rumicum, D.C. senecionis, Berk. suaveolens, Lév. . symphytt, Lév. umbelliferarum, Lév. vince, Berk. . violarum, Berk. TRIPHRAGMIUM, Link. filipendulze (Lasch.) ulmarie (Schum.) . TUBERCINIA, Fries scabies, Berk. trientalis (B. and Br.) TUBERCULINA, Sacc. . persicina (Ditm.) vinosa, Sacc. . PAGE 155 189 127 170 157 184 184 152 120 125 ~ 143 126 167 155 185 195 194 190 157 150: 163 196 259 191 129 184 155 124, 188 159 247, 253 193 167 136 249 181 255 184 161 153 218 219 218 293 294 293 299 299 299 Index of Spectes. 345 UREDO, Pers. eciditformis, Grev. . e@cidioides, D.C, a Mill. egopodit, Schum. agrimoniz, D.C. . agropyri, Preuss. alchemille, Pers. alliorum, D.C. anemones, Pers. annularis, Strauss. antherarum, D. C. anthyllidis, Grev. . apiculosa, Link. api, Wallr. appendiculata, Pers. x var. Pers. 5 var. fist, Pers. . appendiculosa, Berk. arenarig, Schum. armeria, Duby artemisia, Berk. aurea, Purt. bete, Pers. »» var. convolvuli, Pee bifrons, Grev. bistortarum, D. C. bullata, Pers. campanule, Pers. . caprearum, Berk. . caricis, Pers. . » Schum . caries, D. C. ‘ ‘ie Weatacens tis Jannst, cichoracearum, D. C. circe@ Schum. : compransor, Schlecht. confluens, D. C. a 3.9 Schw. crustacea, Berk, cylindrica, Strauss. . effusa, Berk. . >, Strauss. empetri, Pers. epilobit, T). C. 138, 148, phaseoli, var. mercurialis, Pers. var. orchidis, Alb. and PAGE 254 183 242 257 201 255 286 137 261 288 218 280 135 136 156 122 133 120 210 123 189 225 127 146 135 277 183 251 238 276 169 283 248 185 245 250 260 260 261 251 244 219 225 253 152 UREDO—continued. epitea, Kze. and Schum. . euonymi, Mart. euphorbia, Reb. euphrasia, Schum. . excavata, D. C. Jaba, Pers. y «Var. Schw. Jarinosa, Pers. é a «Var. Semectonis, oo ‘ fearig, Schum. filicum, Vesm. fillipendula, Lasch. frosculorum; D. C.. Sragaria, Putt. Srumenti, Sow. fusca, Port. . gentiang, Strauss. . geranit, D. C. gladioli, Requien. . gyrosa, Rebent. helioscopia, Pers. heraclei, Grev. hieracti, Schum. hydropiperis, Schum. hypericorum, D. C. intrusa, Grev. iridis, Thiim. . 5 Di:Ge e labiatarum, D.C. . laricis, Westd. leguminosarum, Link. linearis, Pers. trifoli, Alb. and > «Var, oa Peis, lini, D.C. . longissima, Sow. . lynchii (B. and Br.) mentha, Purt. miniata, Pers. 7 » var. zz, Pers... miilleri, Schrot. oblongata, Grev. obtusa, Strauss. olivacea, D. C. orchidis, Mart. ovata, Grev. . § padi, Kze. and Schum. . PAGE 239 260 237 252 134 120 125 238 249 141 256 220 279 221 162 120 147 126 287 226 237 155 185 283 243 137 257 190 158 262 120 162 256 237 273 259 157 228 237 256 190 222 277 262 244 246 346 Lndex of Spectres. UREDO— continued. parallela, Sow. parnassia, D. C. petasitis, Grev. phragmitis, Schum. phyllireze, Cooke pimpinelle, Strauss. pinguis, D.C. : . plantaginis, B. and Br. . polygonorum, D. C. - 124, polymorpha, var. saxifrage, Strauss. . . polypodii, Pers. . populina, Grev. 5 var. betulina, Pers. . Lorphyrogenita, Kze. porrt, Sow. . potentilla, D. C. > ~Grev. potentillarum, De C. =. 2:22, 5 var. agrimonie, DC . poterii, Spreng... : primule, D. C. pustulata, Pers. ee var. cerastit, Pers. . pyrole, Grev. quercus, Brond. rhinanthacearum, D. C. rose, D.C. . rubigo-vera, D. C.. rubi-tdei, Pers, é rumiis, Schum. saliceti, Lév. . : s» Schlecht. . saxifragarum, D. C. scillarum, Grev. scolopendri, Fckl. . segetum, Pers. : ; 3» var. decipiens, Pers. sempervivt, Alb. and Schw.. senecionis, Schlecht. . soldanelle, D.C. sonchi, Pers. . sonchi-arvensis, Pers. sparsa, Kze, and Schum. statices, Desm. ‘ strieformis, Westd. PAGE 285 129 251 176 258 155 226 259 188 260 256 242 243 246 148 221 221 255 255 220 159 245 248 253 257 252 225 167 226 135 238 240 260 141 256 274 284 230 249 160 250 250 136 123 284 PAGE UREDO—continued. suaveolens, Pers. 182 symphyti, D. C. 255 tanaceti, D. C. A 189 tragopogi, Pers. 282 tropzoli, Desm. . 258 tusstlaginis, Pers. . 251 ulmari@, Schum. 219 umbellatum, Johnst. 184 urceolorum, D.C. . 276 utriculosa, D. C. 280 vaccintorum, Link. 246 zagans, D. C. » var. eptlobiz, D. C. 151 valeriane, Schum. . 128 veronica, Schum. 211 vince, D.C... : 161 vinosa, Berk. . . 279 violacea, Pers. 280 violarum, D.C. 153 vitelline, D. C. ‘ 240 UROCYSTIS, Rabh. . 285 agropyri (Preuss.) . 285 anemones (Pers.) 288 colchici (Schecht.) . 286 fischeri, Korn. ; 286 gladioli (Req.) . . . 287 occulta (Wallr.) . 3 . 285 >> Preuss. ‘ < . 285 parallela, B. and Br. . 285, 286 pompholygodes, Lévy. ¢ . 288 primulicola, Magnus 289 sorosporioides, Korn. 287 violze (Sow.) . 288 UROMYCES, Link. . 11g alchemillze (Pers.) . 139 alliorum (D. C.) . . 137, 148 anthyllidis (Grev.) . 135 apiculatus, Lév. . 125 apiculosa, Lév. 126 appendiculata, Cooke 120 avicularia, Schrot. . 124 behenis (D. C.) . - . 138 betze (Pers.) . : 127 concentricus, Lév. . 14! concomitans, B. and Br. . 139 dactylidis, Otth. . 130 ervi (Wallr.) . 140 Index of Spectes. UROMYCES— continued. excavatus (D. C.) . fabee (Pers.) . ficarize (Schum.) geranii (D. C.) graminum, Cooke . intrusa, Lévy. tridis (Lév.) . junci (Desm.) limonii (D. C.) . ornithogali (Wallr.) orobi (Pers.) . parnassiz (D.C.) . phaseoli (Pers.) pisi (Pers.) . poe, Rabh. polygoni (Pers.) rumicis (Schum. ) rumicum, Lév. salicornize (D. C.) . scillarum (Grev.) scrophularize (D. C.) scutellatus (Schrank.) sparsus (Kze and Schum.) trifolii (Alb. and Schw.).. ulmaria, Lév. urticze, Cooke valerianz (Schum. ) UROMYCOPSIS . USTILAGINEE USTILAGO, Pers. . antherarum, Fries .. bistortarum (D. C.) bromivora (Tul.) candollet, Tul. PAGE 134 119 140 126 130 137 257 132 122 142 12a 128 122 133 134 123 135 136 129 141 139 134 136 124 219 142 128 138 272 272 280 277 278 283 USTILAGO— continued. carbo, Tul. . ‘ 3 cardui, F. v. Waldh. caricis (Pers.) . flosculorum (D. C.) » Tul. grammica, B. and Br. grandis, Fries hypodytes (Schlecht. ) hypogeea, Tul. . intermedia, Schrot. kiihneana, Wolff longissima (Sow.) . major, Schrot. marina, Durieu maydis (D. C.) montagnet, B. and Br. olivacea (D. 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