ct ALBERT R. MANN LIBRARY NEw York STATE COLLEGES OF AGRICULTURE AND HoME ECONOMICS CORNELL UNIVERSITY ‘ornell Universit ce Cornell University Library The original of this book is in the Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www.archive.org/details/cu31924000590541 Dtate of Gonnecticut PUBLIC DOCUMENT No. 47 State Geological and Natural History Survey COMMISSIONERS HENRY ROBERTS, Governor of Connecticut (Chairman) ARTHUR TWINING HADLEY, President of Yale University BRADFORD PAUL RAYMOND, President of Wesleyan University FLAVEL SWEETEN LUTHER, President of Trinity College (Secretary) RUFUS WHITTAKER STIMSON, President of Connecticut Agricultural College SUPERINTENDENT WILLIAM NORTH RICE * BULLETIN No. 5 HARTFORD PRESS The Case, Lockwood & Brainard Company 1905, Qk bag A | COHU 379276 THE USTILAGINEA, OR SMUTS, OF CONNECTICUT By GEORGE PERKINS CLINTON, S.D. Botanist of Connecticut Agricultural Experiment Station Hartrorp Press The Case, Lockwood & Brainard Company May, 1905 The Ustilaginee, or Smuts, of Connecticut. GENERAL CHARACTERS OF THE SMUTS. The Ustilaginez are an order of parasitic fungi commonly known as the smuts, of which corn, oat, and onion smuts are familiar examples. See Figs. 55, 43, 42. Thus the more evi- dent characters of these fungi are the black, dusty masses that break out on the surface of the infected plants (hosts). In these macroscopic characters the smuts are often very similar to the rusts, but with a little experience one can readily dis- tinguish them from these even with the naked eye. The dusty uredo stage of the rusts, for instance, is of a lighter reddish color, and the darker teleuto stage is usually more firmly em- bedded in the plant tissues, than are the smuts. Not all of the smuts, however, produce dark dusty outbreaks, since the white smuts are light colored and usually permanently embedded in the tissues of the host. Generally they occur in the leaves, and often are distinguished by light colored spots produced without distortion of tissues. More experience is needed to distinguish these forms. See Figs. 29-34. The smuts occur on a great variety of flowering plants, and at least 35 different families of plants in North America and 17 in Connecticut are subject to their attack. The grasses, however, are by far the most frequent hosts. Be- tween 500 and 600 species of smuts have been described from different parts of the world, and over 200 of these occur in North America. The number of species known from this state * is 50, and these represent 12 of the 19 genera reported from North America. While time no doubt will reveal a few more species and a greater number of hosts not reported here, we may still consider the group rather thoroughly worked up, since, as yet, only one other state has reported a greater num- ber of species. * The list of smuts reported in this paper is based chiefly on the collections made by the writer during the past three seasons. Specimens of these are to be found in the herbarium of the Conn. Agr. Exp. Station at New Haven and in the writer’s herbarium. Collections made by others are indicated in each case by the name of the collector. 6 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull. | LIFE HISTORY OF THE SMUTS. In the study of these fungi, there are three parts that we need to consider, namely, the mycelium, the spores, and the sori. Mycelium is the technical name for the vegetative part of the smut, by means of which it spreads through its host and gathers nourishment. This consists of microscopic threads confined to the interior of the host, so it is necessary to exam- ine very thin cross sections of the tissues under the micro- scope to see this stage. See Fig. 26. These threads vary but little with the different species, and therefore are of no value in classifying the different species. They are simple, hyaline, more’ or less branched, and at first filled with protoplasmic contents, but gradually lose most of their contents and be- come septate. They push their way between the cells of the host or sometimes through them. Usually, however, they enter the cells only by short special branches called haustoria, whose special function is to gather nourishment from the cell. The mycelium may be localized or, rather generally spread . through the host. In those cases where it gains entrance through the germinating seed, it usually remains evident fi- nally only at the nodes or where it breaks forth to the exterior in its fruiting stage. In perennial plants the mycelium often becomes established in the perennial parts, from which each year it sends threads into the new growth. Eventually the mycelium becomes prominent in certain regions of the host, and there undergoes special modification to give rise to the reproductive or spore stage. The walls of the mycelial threads that form the spores are generally indis- tinct through more or less complete gelatinization. The spores are usually formed from contents in the interior of these fertile threads. This gelatinization apparently serves to nourish the developing spores, as often all signs of the threads disappear on maturity of the spores. In some cases, as in the genus Neovossia, Fig. 13, and often in Entyloma, however, these threads remain as more or less evident envel- opes to the spores. Spores are the bodies by which the smuts reproduce them- selves. See Figs. 1-20. They are the evident, usually ex- No. 5.] USTILAGINEE OF CONNECTICUT. 7 ternal part of the fungus, making up the smutty masses visible to the naked eye, and are the characteristic and variable parts upon which species are based. The spores are usually so formed that at maturity they break out to the exterior of the host in a dusty mass that is easily scattered, thus securing their dispersal. In some species, however, they are embedded in the host so that they are liberated only on the rotting of the tissues, and sometimes they germinate in situ, and secure their dispersal by the formation of secondary spores of.a different nature. While easily seen in mass, the spores are really microscopic in size, varying according to species from 4p to 35u-* They are usually subspherical or spherical, but vary to ovoid, ellipsoidal, or even oblong. Pressure often makes them more or less polyhedral, or irregularly so. They are generally simple, consisting of single separate cells; but are sometimes bound more or less permanently into spore balls, Figs. 7a, 10a. These spore balls may consist entirely of fer- tile cells (spores) or they may have an external coating (cor- tex) of sterile cells, Fig. 19, or an internal matrix of sterile cells or even of threads, Fig. 20. The spore walls may be smooth, or marked with minute spines (echinulations if sharp, verruculations if dull), or reticulated with a net-work of ridges or wings. In color they are hyaline, yellowish, red- dish or olive brown, violet, or purplish. Often the color is so deep that the spores are opaque or nearly so. Upon these variations the genera and species are chiefly classified, though on the whole the variations are not so great as with many of the other fungi. This is partly due to the fact that the spore-producing threads are not diversified*or permanent. In some few species, most frequently in the genus Enty- loma, the mycelium also gives rise to secondary spores called conidia. These are usually hyaline, elongated, ephemeral, and are produced originally on the exterior of the host, gener- ally from mycelial tufts protruding through the stomates and forming these conidia at their tips. Their object is to spread the smut over the host or to new hosts immediately. Sori are the evident masses of the spores that break out singly or in clusters on the various parts of the hosts. See * Au, or micron, is one twenty-five thousandth of an inch. 8 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull. Figs. 27-55. These are protected more or less permanently by thin coverings of plant tissue, or in some genera, as Spha- celotheca, and often in Cintractia, by false membranes com- posed chiefly of fungous cells or threads, Fig. 5a. The sori vary greatly in size, sometimes forming outbreaks smaller than a pin-head, and sometimes, as often in corn smut, reach- ing several inches in diameter. Usually each species has a definite place on its host where the sori appear. This may be on the leaves, Fig. 35, on the stems, Fig. 28, in the blossoms as a whole, Fig. 54, or confined to the anthers, ovaries, Fig. 52, or seeds, Fig. 48. Occasionally a smut may break out in any of these places. In the grasses, when infecting the inflorescence, the sorus may be confined to the ovary, infest the spikelet as a whole, or even involve the entire inflorescence. Sometimes all of the ovaries or spikelets are infested and sometimes only part of them. The characters of the sori, of course, are modi- fied somewhat by the part of the host in which they develop, and also by the kind of host on which the smut occurs. Usually the sori are dusty and easily dissipated. With a few species they are hard, and the agglutinated spore mass is dis- seminated by gradual disintegration, often through the ab- sorption and loss of water. In these cases there is sometimes a gradual ripening of the spores from the outside of the sorus inward. With some genera (Entyloma, Doassansia, etc.) the sorus is rather permanently embedded in the tissues of the host, usually in the leaves, and often produces only a discolora- tion of these. The species of one of the European genera form gall-like growths in the roots. Germination of Spores. There are two chief types of germination of the Ustilaginee upon which the two families are based. In the Ustilaginacee, Fig. 21, the spores, when placed in a drop of water, send out a single, hyaline thread, several times their length, which usually divides into about four cells by cross partitions, or septa. This is called the promycelium. Usually at the apex of each cell one or more elongated thin-walled spores, or sporidia, are budded out, the apical cell bearing its sporidium terminally and the others lat- erally. The sporidia when full-sized are pinched off at the base, and others are sent out until the protoplasm of the cell No. 5.] USTILAGINEE OF CONNECTICUT. 9 is exhausted. These sporidia germinate by a short slender thread (infection thread) that usually pushes out from near one end, Fig. 24. In some species the cells of the promyce- lium, instead of forming the sporidia, give rise directly to the infection threads, Fig. 23. Sometimes the adjacent cells of the promycelium become connected by closely applied short threads, forming buckle or knee joints, and from these may develop infection threads. Some species germinate directly into elongated threads which scarcely partake of the nature of a promycelium. When nutrient is added to the drop of water, the spores germinate much more luxuriantly, the spor- idia usually sprouting out other sporidia while still attached to the promycelium. Especially in a solid cultural medium these chains develop a more or less complicated system of branching, Fig. 25. In a liquid solution, the sporidia usually soon fall off from the promyceliurn, but continue to multiply by the yeast fashions of budding new sporidia, which soon sep- arate and develop others, until the nourishment is exhausted. Then the sporidia may develop infection threads. The second chief type of germination is shown by the spe- cies of Tilletiacee, Fig. 22. The simple or septate promyce- lium in this case bears all of the elongated sporidia in a terminal cluster. Sometimes these primary sporidia develop terminal secondary sporidia. With some species the sporidia, before or after falling off from the promycelium, become yoked in pairs by short connecting threads. In nutrient solutions some of the species develop finally a complicated mycelium that bears numerous aérial sporidia or conidia quite different from the normal type produced in water. The conidia develop in- fection threads on germination. Infection of Host. The host is entered by the fine infec-. tion threads of the promycelia or the sporidia boring through its tissues into the interior. Once inside, these develop the mycelium of which we have already spoken. Many of the smuts infect their host only through the young hypocotyl or epicotyl of the seedling. In these cases the smut usually can not gain successful entrance after the plants appear above ground. Once inside, the mycelium rapidly penetrates the young tissues, seeking to gain access to the growing tip, where 10 CONNECTICUT GEOL, AND NAT. HIST. suRVEY. [Bull. it then follows the upward growth of the plant, often not giv- - ing evidence of its presence until it breaks out in its spore stage in the fruiting organs of the host two or three months later. This is the case with the oat smuts, wheat and barley smuts, and the grain smut of broom-corn. In this latter case the mycelium has traveled upward through six to ten feet of the cane. In other species infection may take place through - any very young tissue of the host upon which the germs may be blown or washed into contact. This is the case with corn, and the corn smut appears soon after infection in its smutty outbreaks, the mycelium usually remaining localized. With the Entylomas the infection is largely confined to the leaves, and apparently the mycelium is limited to the vicinity of the sorus. ECONOMIC IMPORTANCE. Injury. While most of the smuts occur on plants of no economic importance, there are at least twenty-five species in America that do more or less injury to cultivated plants. Some of these cause such serious injury that they are counted among the worst species of parasitic fungi. It is with the cereals that the smuts do most damage. Wheat, oats, barley, rye, and corn are all subject to attacks from one or more species. With the first three hosts the smuts break out in the inflorescence, en- tirely preventing the formation of the seed. It thus becomes an easy matter to determine rather accurately the per cent. of loss in fields of these grains by counting the number of smutted and of free heads in a given area. Oat smut, for instance, in our. central states generally claims from one to fifteen per cent. of the grain, and in some few fields as high as thirty or forty per cent. has been destroyed. In Illinois there is perhaps an average loss per'year of one million dollars worth of oats, caused by the two smuts of this plant. In Indiana Arthur found one field of wheat where fifty per cent. of the grain was destroyed by the stinking smut. In the northwestern states, in the wheat districts, the loss caused by this latter smut is very great, as it is one of their worst fungous pests. When abundant it renders the grain inferior for milling purposes, as the spores may be so numerous in the flour that they darken No. 5.] USTILAGINEZ OF CONNECTICUT. II it. In South Carolina during recent years a smut of rice has been introduced from Japan, and causes some loss to this crop. In the eastern states where onions are raised extensively, the onion smut often does considerable injury. In this case the smut becomes established in the soil and often prevents profit- able onion culture on this land. Corn smut is common every- where, and especially on sweet corn causes considerable injury. In Connecticut the most injurious smuts, from an economic standpoint, are those of onion, corn, barley, and oats. Prevention. With those plants where infection takes place through the germinating seed, the danger usually comes from the spores that mechanically adhere to the seed. It has been found that, if these are killed, the crop from this seed will be free from smut. Investigations have shown that certain treat- ment of the grain with chemical solutions or with hot water will kill the spores with little or no injury to the seed. For instance, it has long been known that soaking the seed of wheat in a solution of copper sulphate of a certain strength for a cer- tain time serves to prevent or Jessen the amount of stinking smut in the crop. Later it was found that ‘soaking the seed for ten or fifteen minutes in hot water at a temperature of 132° to 135° F. was a more efficient remedy, since there was less likelihood of injury to the seed. More recently still came the less cumbersome practice of sprinkling piles of the grain with formalin, one pint to fifty gallons of water, stirring the grain to thoroughly wet it, and leaving it in piles or in bags over night for the fumes to act on the spores. The smuts of oats, the covered smut of barley, the stinking smut of wheat, the grain smut of sorghum and broom-corn, and the grain smut of millet, have all yielded to seed treatment. The loose smuts of wheat and barley can be prevented or lessened by a severer method of the hot water treatment (in which a preliminary soaking of several hours in cold water precedes the hot water treatment), but this also injures the seed more or less, so that a greater quantity per acre must be sown. Corn smut can not be prevented by seed treatment, as the smut gains entrance through any young tissue of the host. Some experimenters have advocated the removal of the smut-balls as soon as they appear. It is known that the fungus develops aérial conidia 12 CONNECTICUT GEOL. AND NAT. HIST. suRVEY. [Bull. in manure, and so fresh manure applied to corn land is apt to increase the amount of smut. With onion smut we have a case where the seedlings while coming through the soil are in- fected by the smut established there through a previous smutted crop. . Onions raised from sets in this land are little. infected. Some success has been had in preventing this smut by treating the soil with a mixture of lime and sulphur, drilled into it at time of seeding. SYSTEMATIC TREATMENT OF CONNECTICUT SPECIES. The order Ustilaginee is divided into two families, based chiefly on the method of spore germination. They are the Ustilaginacee (represented in this state by the genera Cintrac- tia, Schizonella, Sorosporium, Sphacelotheca, Tolyposporium, Ustilago), and the Tilletiacee (represented by Doassansia, Entyloma, Neovossia, Tilletia, Tracya, Urocystis). Key to Genera. I. Spores simple. A. Usually forming a dusty sorus at maturity (see also IB and IJJAza). 1. Large, usually 16-35. a. With an elongated hyaline ees sauce Neovossia. b. Without a conspicuous appendage. Tilletia. 2. Small to medium, usually 5-18. a. Sorus covered with a false membrane of definite fungous cells..... Sphacelotheca. b. Protecting membrane of plant tissue only (sometimes with gelatinized hyphe)...... Ustilago. B. More or less firmly agglutinated at maturity....... Cintractia. C. Permanently imbedded in leaves, praducing dis- COlMG. SNES aacsudbedueusesnaueeens Entyloma, II. Spores chiefly in pairs, forming an agglutinated sorus. Ss rhizomella, No. 5.] USTILAGINEZ OF CONNECTICUT. 13 ITI. Spores in more or less permanent balls. A. Forming a dusty or granular sorus at maturity. I. Spore balls consisting only of spores. a. Often evanescent (Ustilago-like spores)... Sorosporium. b. Quite permanent, adhering by folds of spore Oats se csvseeensew sewn Tolyposporium. 2. Spore balls with a cortex of sterile cells...... Urocystis. B. Permanently embedded in the plant tissues. 1. Spore balls with a definite cortex of sterile cells Doassansia. 2. Spore balls without cortex but with sterile threads in center ................ Tracya. USTILAGINACEZ §Schrot. The sori usually form exposed, dusty, or agglutinated spore masses. Germination is by means of a septate pro- mycelium, which usually gives rise to terminal and lateral sporidia (capable of yeast-like multiplication in nutrient solu- tions), or else to infection threads. Figs. 21, 23-25. Ustilago Rouss. The sori occur on various parts of the host, according to the species, and at maturity form dusty, usually dark colored spore masses. The spores are single, separate, of small to medium size (5-18), and are produced irregularly in fertile threads that entirely disappear through gelatinization at ma- turity. Figs. 1-4, 43-55. : This is the most common and the typical genus of the smuts. Saccardo, in his Sylloge Fungorum, describes about 250 species for the world: 72 species occur in North America and 18 in Connecticut. A number of the species possess eco- nomic importance as parasites of the cereals. Key to Species of Ustilago. I. Spores olive or reddish brown. A. Spores smooth. 1. Sori in leaves forming linear strie.......... ‘ U. longissima. 14 CONNECTICUT GEOL. AND NAT. HIST. suRVEY. [Bull. 2. Sori in spikelets, ovoid. a. Spores usually lighter colored on one side, 5-Op. * Spikelets entirely destroyed. ..U. Hordei. %* Spikelets smutted only at base. .U. levis. b. Spores uniformly colored, 8-IIp.....-.++- U. Crameri. B. Spores echinulate or verruculose. 1. Sorus in each spikelet, distinct. a. Spores usually lighter colored on one side, 5-9r- * Host Arrhenatherum avenaceum (Oat (GEASS). geacesgr sic: sioveraiainsae aise U. perennans. ** Host Avena sativa (Oats)...U. Avene. “ek Host Hordeum vulgare (Barley)..... U. nuda. b. Spores uniformly colored, 10-14p.......+- U. Panici-glauci. 2. Sorus involving the entire panicle............ U. Rabenhorstiana. 3. Sori in the ovaries. a. Smooth.and inconspicuous, 1-2 mm. * Spores echinulate ...... U. spermophora. *k Spores verruculose ....... U. Eriocault, b. Hispid and conspicuous, 4-Io mm........ U. spherogena. 4. Sori in leaves (rarely on stem). a. Forming nodular swellings, often at nodes U. Crus-galli. b. Forming linear striz in the blades........ U. strieformis. 5. Sori breaking out on any part of host, often NATE: ssisriccs terrasse eratorsteuie sages U. Zee. II. Spores golden brown, verrucose; sori in seeds......... U. Oxalidis. III. Spores violet or purplish, reticulate, sori in flowers. 1. Reticulations rather fine, 1-3n....U. anomala. 2. Reticulations rather coarse, 2-4u.U. utriculosa. No. 5.] USTILAGINEZ OF CONNECTICUT. 15 / Ustilago longissima (Sow.) Tul. Fig. 1. The sori form more or less distinct, linear strie from a féw mm. to length of leaf; the epidermal covering soon ruptures, and the dark reddish brown spore mass becomes scattered from the more or less shredded tissues. The spores are light brown, oblong, or more commonly ellipsoidal to spherical, smooth (or sometimes scarcely granular under an immersion lens), 4-8» in length. Host and Distr.: Glyceria grandis, Shaker Station, June 29, 1903. This species undoubtedly has a wider Wistribution than in- dicated above. Ustilago Hordei (Pers.) Kell. & Sw. Fig. 46. The sori occur in every spikelet, forming an ovate, adhering, purple- black spore mass, 6-10 mm. in length, and rather permanently covered by the thin, usually transparent plant tissue. The spores are reddish brown, slightly lighter colored on one side; chiefly subspherical or spherical, smooth, and 5-ou, rarely (most elongated) 9-11p, in length. Host and Distr.: Hordeum vulgare, Whitneyville, Sept. 24, 1902. This covered smut of barley is not nearly so common in this state as is the loose smut of the same host. Ustilago levis (Kell. & Sw.) Magn. Fig. 45. The sori usually occur in all of the spikelets, are more or less hidden by the enveloping glumes (which may be infected only at their base or in the interior), and form rather permanent, black- brown, semi-agglutinated, ovate spore masses 6-10 mm. in length. The spores are reddish brown, slightly lighter col- ored on one side, chiefly subspherical or spherical, smooth, 5-9m, rarely (most elongated) 11, in length. Host and Distr.: Avena sativa, New Haven, July Io, July 28, 1902. This hidden smut of oats is not so common in our oat fields as is the loose smut, U. Avene. Ustilago Crameri Korn. Fig. 44. The sori occur in all of the spikelets of the spike, destroying especially the inner and basal parts, and form ovate dusty brown spore masses 2-4 mm. in length. The spores are reddish brown, ovoid to 16 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull. subspherical, or occasionally more elongated or irregular,’ smooth, often with pitted contents, and 8-11p in length. Host and Distr.: Setaria Italica, New Haven, Sept. 11,. 1903. So far this species has been found in this state only once, | in a small plot of Hungarian grass grown at the Experiment : Station. In some of the central and western states it some-* times causes considerable injury to its host. It can be pre- vented by seed treatment with hot water, formalin, etc. Ustilago perennans Rostr. The sori occupy all of the. spikelets, usually destroying the basal and inner parts (some/ times even running down slightly on the pedicles) ; they are oblong, 3-8 mm. in length, and have a somewhat protected | semi-dusty olive brown spore mass. The spores are chiefly subspherical or spherical, occasionally ellipsoidal to ovate, ‘usually lighter colored on one side, minutely echinulate, espe- : cially on lighter side, and 5-8 in length. Host and Distr.: Arrhenatherum avenaceum, South Man- chester (Thaxter). This is closely related to the oat smuts. Its mycelium be-- comes perennial in the underground parts of the host, and so smuts the infested plants year after year. Ustilago Avene (Pers.) Jens. Figs. 23, 43. The sori appear in all of the spikelets, completely destroying all the floral parts, except a very temporary transparent covering membrane; eventually the olive brown dusty spore mass be- comes dissipated, leaving behind only the naked pedicles. The spores are reddish brown, lighter colored on one side, sub- spherical to spherical, or occasionally more elongated, mi- nutely echinulate, especially on the lighter side, and chiefly 5-9 in length. Host and Distr.: Avena sativa, New Haven, July 8, July 28, Aug. 17, 1902, Aug. 9, 1903; West Cornwall, July 18, 1902; Westville, July 7, 1903; West Haven, July 22, 1903. This is a common pest in the oat fields of the state, though the per cent. of infected plants is smaller than in the central and western states. It yields to seed treatment. Ustilago nuda (Jens.) Kell. & Sw. Fig. 47. The sori infest all of the spikelets, changing each into an olive brown,: No. 5.] USTILAGINEH OF CONNECTICUT, 17 oblong, dusty spore mass about 6-10 mm. in length; this is very temporarily protected by a thin transparent membrane, and upon dispersal of the spores nothing remains but the naked rhachis. The spores are lighter colored on one side, sub- spherical to spherical, or occasionally more elongated, minutely echinulate, especially on the lighter side, and chiefly 5-9» in length. Host and Distr.: Hordeum vulgare, Storrs, 1901; New Haven, July 8, 1902, June 15, 1904; Whitneyville, Sept. 24, 1902; Westville, July 7, 1903. This is the loose smut of barley, and is rather common in the fields of this crop. The smuts of barley, oats, and oat grass, as will be seen by the descriptions, are very similar, in fact were not long ago considered one species. It requires the modified form of the hot-water treatment to prevent the loose smut of barley. Ustilago Panici-glauci (Wallr.) Wint. Fig. 49. The sori occur in all of the spikelets, are ovate, 2-3 mm. in length, and at first protected by thin transparent glumes, but soon rup- ture these, and scatter the dusty black-brown spore mass. The spores are dark reddish brown, ovoid, spherical or sometimes more elongated, prominently and abundantly echinulate, and 10-14 in length. Host and Distr.: Setaria glauca, Southington, Aug. 8,’ 1902; Bridgeport, Sept. 15, 1902; Berlin, Oct. 3, 1902; Glas- tonbury, Oct. 23, 1902; Andover, Sept. 15, 1903; New Canaan, Sept. 29, 1903; Manchester, Oct. 2, 1903. Throughout the United States this is a common smut on the yellow fox-tail grass, though it does not occur on the green fox-tail. ‘ . Ustilago Rabenhorstiana Kithn. Fig. 50. The sorus involves the entire inflorescence, changing it into a linear or oblong body, 3-5 cm. in length, which is usually hidden by the enveloping leaf sheath; it is covered for a short time by a very fragile transparent plant membrane, and within the dusty brown-black spore mass are often elongated remains of the plant tissues. The spores are reddish or olive brown, ovoid to spherical, or occasionally slightly angled, echinulate to ver- ruculose, and 10-14m in length. 2 18 CONNECTICUT GEOL. AND NAT. HIST. suRVEY. [Bull. Host and Distr.: Panicum sanguinale, Centreville, Aug. 10, Sept. 1, 1902; Unionville, Aug. 26, 1902; New Haven, Sept., 1902, Oct. 5, 1904; Westville, Aug. 25, 1903; Monto- wese, Sept. 14, 1903; New Canaan, Sept. 29, 1903. This is one of the common grass smuts here as elsewhere, and is to be looked for in the fall of the year. Ustilago spermophora B. & C. Fig. 52. The brown- black dusty sori occur in the ovaries, infesting one here and there; they show as small ovate bodies, about 2 mm. in length, extending between the spreading glumes, and are at first pro- tected by a thin ovary membrane at the apex of which are the remains of the styles. The spores are light brown, ovoid to subspherical, usually prominently echinulate, and 8-11, or occasionally 134 in length. Host and Distr.: Eragrostis major, New Haven, Oct. 31, 1902; Westville, Oct. 17, 1903. The sori usually occur in only a few of the ovaries, and are so inconspicuous that the smut is easily overlooked. Fig. 52 shows isolated spikelets of the grass with a single sorus in each. Ustilago Eriocauli (Mass.) Clint. The sori occur in the ovaries, scarcely showing between the glumes as slightly swollen ovoid bodies about 1 mm. in length; a thin membrane protects the rather firm, at first semi-agglutinated, but finally dusty, olive-black spore mass. The spores are polyhedral, sub- spherical, or occasionally more elongated, rather prominently verruculose, and 9-15 in length. Host and Distr.: Eriocaulon septangulare, Whitneyville, Sept. 21, 1902. It requires very close examination to detect the flower heads containing this inconspicuous smut, since they resemble the normal ones. The smut should be searched for in the fall. Ustilago sphzrogena Burr. Fig. 53. The sori occupy the ovaries, forming ovate bodies, 4-10 mm. in length, which are covered by a tough hispid plant membrane that ruptures irregularly from the apex, disclosing at first an agglutinated, but finally a dusty, olive brown spore mass. The spores are ovoid to subspherical, prominently and sharply echinulate, and 9-12 in length. No. 5.] USTILAGINEH OF CONNECTICUT. 19: Host and Distr.: Panicum Crus-galli, Conn. (Setchell) ; Savin Rock, Sept. 14, 1902; Westville, Sept. 12, 1902. This species and the next are very closely related, and oc- cur on the same host, but are rarely, if ever, found together. Fig. 53 shows a portion of the flower panicle with a single un- usually large sorus. Ustilago Crus-galli Tr. & Earle. The sori form nodules often encircling the leaf node, or more rarely occur in place of inflorescence, infecting both leaves and stem; they are one to several cm. in diameter and protected by a tough hispid plant membrane, which on rupture discloses an olive brown spore mass. The spores are reddish brown, ovoid to spherical, occasionally more elongated, rather bluntly echinulate or even verruculose, and 10-14 in length. Host and Distr.: Panicum Crus-galli, New Haven, Sept. II, 1903. Only a single specimen of this smut was found on a culti- vated variety of this barn-yard grass grown at the Experiment Station. Ustilago strizformis (West.) Niessl. Fig. 51. The sori occur in the leaves, rarely in the inflorescence, forming short linear striz, or by terminal fusion reaching several cm. in length, and laterally are often so crowded as to cover most of the leaf; at first they are covered by the epidermis, but this soon ruptures, and the dusty brown-black spore masses be- come scattered from the shredded tissues. The spores are red- dish brown, vary from ellipsoidal to spherical, or occasionally irregular, are prominently echinulate, and 9-14» in length. Hosts and Distr.: Agrostis alba var. vulgaris, Whitney- ville, July 20, 1902, May 9, 1903; Centreville, June 12, 1904; Phleum pratense, Whitneyville, Bae 9, 1903; New Haven, May 16, 1903. Both of these hosts are economic eae grown for pasture or hay, but so far the smut has been found on them in this state only in door yards. Fig. 51 shows the leaves of Agrostis alba var. vulgaris shredded by this fungus. Ustilago Zez (Beckm.) Ung. Figs. 2, 55. The sori break out on any part of the host, often forming prominent smut balls, though these vary from a few mm. to over a dem. 20 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull. in diameter, and also vary in shape according to the part at- tacked; the brownish black spore mass is at first covered with a whitish membrane, composed largely of semi-gelatinized fun- gous threads. The spores are ellipsoidal to spherical, occa- sionally irregular, prominently echinulate, and 8-11, rarely even I5y, in length. Hosts and Distr.: Euchlena luxurians, New Haven, Sept. II, 1903; Zea Mays, New Haven (Sturgis), Aug. 18, 1901 (Rorer) ; Southington, July 17, 1902; Westville, Aug. 14, Sept. . 2, 1902, June 28, 1904; Hartford, Oct. 20, 1902; New Canaan, Sept. 29, 1903. Corn smut is common on both the sweet and field varieties, though in this state the former is more subject to its attacks. Seed treatment will not prevent the smut, as it can gain en- trance to its host through any exposed young tissue. The first host given is teosinte, a plant that is closely related to corn. Fig. 55 shows smutted staminate blossoms of corn re- duced to one-half size. Ustilago Oxalidis Ell. & Tr. Figs. 3, 48. The incon- spictious sori are found in the seeds, all or part of these being changed into reddish brown dusty spore masses, that show to the exterior only on the dehiscence of the otherwise little modi- fied ovaries. The spores are golden yellow, ovoid to spherical, or rarely more elongated or irregular, coarsely verrucose, and 13-20 in length. Host and Distr.: Ovalis stricta, Yalesville, July, 1902; West Cornwall, July 18, 1902; New Haven, July, 1902; Whit- neyville, Oct. 18, 1902, Sept. 9, 1903; Manchester, Oct. 2, 1903. An inconspicuous conidial stage is also produced on the surface of the anthers of the infected flowers. These temporary spores are thin-walled and ovoid to subspherical. They are so placed that they are probably carried from the flowers by in- sects, as are pollen grains. Fig. 48 shows two ovaries in which all of the seeds have been changed into smutty bodies. Ustilago anomala Kze. The sori occur in the essential organs of all the flowers, the floral envelopes forming a cov- ering to the dusty, purplish spore mass. The spores are light violet, ovoid to spherical, occasionally somewhat irregular, provided with rather fine winged reticulations (1-34 wide by Iu deep), and 10-15, rarely 17p, in length. No- 5.] USTILAGINEH OF CONNECTICUT. 21 Hosts and Distr.: Polygonum Convolvulus (?), Mt. Car- mel, May 8, 1904 (last year’s flowers); Polygonum dume- torum var. scandens; Montowese, Sept. 14, 1903; New Canaan, Sept. 29, 1903. This species is closely related to the next, but differs in having usually lighter colored and somewhat larger spores, with finer reticulations. Ustilago utriculosa (Nees.) Tul. Figs. 4, 21, 54. The sori destroy the essential organs of all of the flowers, and form ovate, dusty, purplish spore masses 3-4 mm. in length, pro- tected at first by the floral envelopes. The spores are violet or purplish, chiefly subspherical or spherical, provided with rather coarse, winged reticulations (2-4 wide by about 1.5 deep), and chiefly 9-14 in diameter. Hosts and Distr.: Polygonum Hydropiper, Montowese, Sept. 20, 1902; Polygonum hydropiperoides, Whitneyville, July 24, 1902; Polygonum lapathifolium, Westville, Sept. 2, 1902; Polygonum Pennsylvanicum, Westville, Sept. 2, 1902; Ham- den, Sept. 11, 1902; Hartford, Oct. 20, 1902; Glastonbury, Oct. 23, 1902; Yalesville, Sept. 11, 1903, Oct. 14, 1904; Monto- wese, Sept. 14, 1903; New Canaan, Sept. 29, 1903; Manches- ter, Oct.-2, 1903; Green’s Farms, Sept. 30, 1904. This is one of the most common smuts of the state, espe- cially on the last host. Sphacelotheca DeBy. The sori are usually found in the inflorescence (often con- fined to the ovaries), are provided with a false membrane of fungous cells that soon ruptures, disclosing a dusty spore mass and a central columella composed chiefly of plant tissues. The false membrane is formed largely or entirely of definite sterile fungous cells which are hyaline or slightly tinted, and vary in shape from linear to subspherical or cuboidal, and in size from less than to larger than the spores. The spores are like those of Ustilago, simple, free, usually reddish brown, and of small to medium size. Figs. 5, 6, 37, 38. Very often groups of the sterile subspherical cells are scat- tered through the spore mass. The columella is usually the remains of the woody plant tissues, and often protrudes above 22 CONNECTICUT GEOL. AND NAT. HIST. survEY. [Bull the spore mass as the latter wears away. See Fig. 37. This genus has not been thoroughly worked up for the whole world, and so a number of the species belonging under it are now in- cluded under Ustilago. Only two of the 16 species occurring in North America are of economic importance. So far only two species have been found in Connecticut. Key to Species of Sphacelotheca. I, Spores olive brown, smooth............+++.. S. Sorghi. II. Spores purplish, verruculose.......... S. Hydropiperis. : Sphacelotheca Sorghi (Lk.) Clint. Figs. 5, 24, 26, 38. The sori occur in the ovaries, forming oblong or ovate bodies usually 3-8 mm. in length, or rarely fusing the aborted spike- lets into longer forms. The brownish false membrane wears away from the apex, revealing the olive brown spore mass and finally the evident slender columella. The sterile cells of the membrane easily break up into groups, and are hyaline, oblong to subspherical, and chiefly 7-18 in length. The olive brown. spores are subspherical or spherical, smooth, often have pitted contents, and are 5.5-8.5 in diameter. Hosts and Distr.: Sorghum vulgare var. sorghum, New Haven, Sept. 30, 1903; Sorghum vulgare var. technicum, New Haven, Sept. 20, 1901 (Rorer), Sept. 11, 1903. This species was found on both sorghum and broom-corn grown at the Experiment Station. So far as I can learn, neither of these plants is grown commercially in the state, so the smut is of no economic importance here, though further west it often does considerable injury to these plants. Sphacelotheca Hydropiperis (Schum.) DeBy. Figs. 6, 37. The sori are found in the ovaries, forming oblong or ovate bodies 3-5 mm. in length; with the false membrane de- hiscing at the apex, and revealing the purple-black spore mass and finally the slender columella. The sterile cells, besides forming the false membrane, constitute part of the columella; they easily separate, are hyaline or slightly violet tinted, chiefly subspherical, and 6-17 in length. The spores are purplish, Broadly oblong or ovate to (chiefly) subspherical, very mi- nutely but abundantly verruculose and 10-1 7» in length. Hosts and Distr.: Polygonum acre, Green’s Farms, Sept. No. 5.] USTILAGINE® OF CONNECTICUT. 23 30, 1904; Polygonum sagittatum, Westville, Sept. 2, 1902, Sept. 14, 1903 (Britton); Oct. 4, 1904; Montowese, Sept. 20, 1902; Whitneyville, Sept. 21, 1902; Centreville, Sept. 27, 1902; New Canaan, Oct. 9, 1902 (Britton), Sept. 11, 1903; Yales- ville, Sept. 11, 1903; Andover, Sept. 15, 1903; Cheshire, Oct. 25, 1903. This is another of the common smuts of the state, especially on the arrow-leaf Polygonum, and is found in the fall. It is the only purple-spored species of the genus, of which it is the type, however. Fig. 37 shows the smut on Polygonum acre, several of the sori exposing the slender central columella. Cintractia Cornu. The sori occur on various parts of the host, but most com- monly in the ovaries; at maturity they form a firmly aggluti- nated, or more rarely a dusty, spore mass, which usually is pro- tected at first by a false membrane of sterile threads or indefi- nite cells. The single spores develop centripetally around a central axis, the outermost wearing off as they ripen; they are usually of medium to large size, and of a dark reddish black : color, often opaque. Figs. 8, 27, 28. The species of this genus occur on the Cyperacez, or occa- sionally on related families. So far 17 species have been de- scribed, 13 occurring in North America, and 4 in Connecticut. None of the species are of economic importance. Key to Species of Cintractia, I. Sori dusty at maturity, concealed by the glumes. A. Spores usually irregular polyhedral...... C. Cyperi. B. Spores usually ovoid to spherical (often with hya- liné Wines.) wap anid ian re nee ea C. Montagnei. II. Sori firmly agglutinated at maturity. A. Subspherical, in ovaries................ C. Caricis. B. Oblong to linear, surrounding base of pedicles..... CC. Junci. Cintractia Cyperi Clint. The sori occur in the interior of the spikelets, infecting all of the head, and are hidden by the enveloping glumes until finally the dusty spore mass sheds 24 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull. out on their exterior. The spores are reddish brown, oblong to polyhedral, chiefly irregular, smooth, but often showing darker lines on surface due to pressure of spore mass, and 12- 18u, or (most elongated) even 22, in length. Host and Distr.: Cyperus filiculmis, North Haven, July 26, 1902; Montowese, Sept. 14, 1903. This species was described originally from this state by the writer, having been found not uncommon on the sand plains near North Haven and Montowese. The affected plants look very much like those free from the smut, but can usually be detected by the darker aspect of the spikelets, which in this sedge are clustered into heads. Cintractia Montagnei (Tul.) Magn. The sori occur hidden in the ovaries, forming inconspicuous, oblong to sub- spherical, usually dusty spore masses. The spores are brown or brownish black, often compressed laterally, and so appear- ing ovoid to subspherical or occasionally more irregular and angled, smooth but minutely pitted, and 12-19, chiefly 13-16y, in length; they are very often provided with conspicuous, hya- line, wing-like bladders on either side. Host and Distr.: Rhynchospora alba, Berlin, Sept. 3, 1902; Cheshire, Oct. 25, 1903. This is another smut readily overlooked, because the in- conspicuous sori are hidden by the floral bracts. Cintractia Caricis (Pers.) Magn. Figs. 8,27. The sori form subspherical bodies about 3-4 mm. in diameter in the ovaries; at first the sorus is protected by a white membrane of sterile fungous tissue, but this soon wears off, revealing the black, firmly agglutinated spore mass, the spores of which gradually ripen and wear off toward the interior. ~ The spores are black-brown, subopaque, chiefly irregular polyhedral, or occasionally ovoid to subspherical, smooth or pitted to gran- ular or even papillate, and 16-27y, chiefly 18-22y, in length. Host and Distr.: Carex Penysylvanica, Rainbow, June 6, 1903; East Hartford, June 1, 1904 (Weatherby). This variable smut has been reported on a large number of Carex species in North America, but the above is the one upon which it commonly occurs. The illustration shows it in the ovaries at the base of the staminate spikelet of this host. No. 5.] USTILAGINE® OF CONNECTICUT. 25 Cintractia Junci (Schw.) Trel. Fig. 28. The sori are linear, usually surrounding the pedicles or peduncles for half or more of their lower length, occasionally developing in the basal parts of the flowers; they form a black, rather firmly ag- glutinated spore mass. The spores are black-brown, sub- opaque, oblorig to irregular polyhedral or subspherical, very minutely pitted, and 14-22» in length. . Host and Distr.: Juncus tenuis, Westville, June, 1891 (Thaxter) ; Milford, June 26, 1894 (Sturgis), July 27, 1902; Whitneyville, June 20, 1902. This was one of the first smuts reported from North Amer- ica, having ‘been described by Schweinitz from Carolina in 1834. The figure shows one of the peduncles smutted for half its length. * Schizonella Schrot. The sori form black agglutinated spore masses in the leaves. The spores are united in pairs (formed by internal division of a mother cell), and often become laxly connected by their bulging contiguous surfaces, or even entirely separ- ated; they are of reddish brown color and of small or medium size. Figs. 9, 35. Only one species, with a variety, is known for this genus, but it has a wide distribution on various species of Care-x. In North America it has been found chiefly on Carex Pennsyl- vanica, its host in this state. Schizonella melanogramma (DC.) Schrét. Figs. 9, 35. The sori form black, agglutinated, linear, elevated striz, I or 2 mm. long, or, by terminal fusion, of considerable length, chiefly on the upper sides of the leaves. The spores are dark reddish brown, often with the cells entirely or partially sep- arated by the bulging out of their contiguous surfaces, chiefly ellipsoidal to hemispherical, or, when separated entirely, poly- hedral or subspherical, and 8-12» in length. Host and Distr.: Carex Pennsylvanica, Westville, July 12, 1902; Whitneyville, May 4, 1903. This is a species found chiefly in the spring; it probably has a much wider distribution than indicated here. 26 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull. Sorosporium Rud. The sori occur in various parts of the host, forming dusty dark colored spore masses like Ustilago. The medium-sized spore balls are composed of numerous spores, often so loosely , held together that in time they separate entirely. The spores are like those of Ustilago, simple, olive to reddish brown, and of medium size. Figs. 7, 36. When the spore balls become separated into the individual » spores, it is difficult to distinguish the species from Ustilago. The number of species described by Saccardo is over 30, but some of these probably belong under other genera. For | North America 9 species are now known, and 2 or 3 of these occur in Connecticut. Key to Species of Sorosporium. I. Sori in ovaries, 1-2 cm. in length; spores 8-12p........ S. Everhart. II. Sori involving the entire infloresence usually. A. Sorus 1-5 cm.; spores 12-IQp.......... S. Ellisu. B. Sorus usually 3-7 cm.; spores Q-I 3M... eee eee S. Syntherisme. Sorosporium Everhartii Ell. & Gall. The sori develop in the ovaries, forming linear bodies 1-2 cm. in length, and are covered with a prominent whitish false membrane that de- hisces at the apex into several lobes, disclosing the black- brown, semi-agglutinated spore mass and the flattened colu- mella of plant tissue. The spore balls are oblong to subspher- ical, composed of many firmly agglutinated spores, and vary from 55 to 125u in length. The spores are reddish brown, or the interior ones often nearly hyaline, ovoid to subspherical or polyhedral, smooth (outermost rarely granular), and 8-12, in length. Host and Distr.: Andropogon scoparius, Westville, Oct. 22, 1903; [Southington, July 16, 1902; North Haven, July 26, 1902; Montowese, Sept. 20, 1902; New Haven, Oct. 18, 1903]. The first specimen mentioned above is typical of the species as described here. Those included in the brackets were origi- No. 5.] USTILAGINEZ OF CONNECTICUT. . a7 nally considered by the writer as belonging under S. Ellisii, since their sori involve the entire inflorescence, instead of being limited to the ovaries. The spores, however, are smaller than those of the typical members of that species, and both the spores and spore balls are like S. Everhartii, so they may possibly be only vigorous specimens of this species, in which the sorus has involved the whole inflorescence. Sorosporium Syntherismz (Pk.) Farl. Figs. 7, 36. The elongated sori usually involve the entire inflorescence, 3-7 cm. in length, or more rarely are limited to the individual spikelets, when they are shorter; they are provided with a prominent false membrane which ruptures irregularly, disclosing the black- brown spore mass, within which are often shredded filaments of plant tissues. The sterile cells of the membrane are hya- line, oblong to cubical or subspherical, and tend to adhere in filaments when crushed apart. The spore balls, often evanes- cent when old, are irregular oblong to subspherical, and 40- 1oou in length. The reddish brown spores are minutely ver- rucose (the inner often smooth and more lightly colored), subspherical, polyhedrat or occasionally more elongated, chiefly 9-13 in length. Host and Distr.: Cenchrus tribuloides, Savin Rock, Aug. 30, 1902; Montowese, Sept. 20, 1902. This species should also be looked for on Panicum pro- liferum, which is a common host for it elsewhere. Tolyposporium Wor. The sori are usually found in the ovaries, forming a gran- ular spore mass at maturity. The spore balls are of medium size, dark colored, and composed of numerous permanently united spores. The spores are bound together by ridged folds or thickenings of their outer walls, and are of small to me- dium size. Figs. 10, 40. The spore balls can be ruptured by pressure, when the outer darker colored covering often breaks apart as ridges or spine-like projections on the light colored or hyaline spores. So far 13 species have been described, though it is doubtful if all belong to this genus; 2 of these occur in North America, 28 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull. and 1 in Connecticut. None of the species apparently are of economic importance. Tolyposporium bullatum (Schrét) Schrét. Figs. 10, 40. The ovate sori are found in the ovaries, are about 3-5 mm. in length, and are covered by a smooth greenish plant membrane, which upon rupturing discloses the granular, black spore mass. The spore balls are opaque, black, oblong to spherical or polyhedral, contain numerous (over 100) firmly aggluti- nated spores, and are 50-160 in length. The spores are light reddish brown, or the inner ones semi-hyaline, and are covered with a thin, tinted outer coat thrown more or less into ridges or folds that bind the spores together; they vary from ovoid to spherical or polyhedral, and are 7-12y, rarely 14, in length. Host and Distr.: Panicum Crus-galli, Woodmont (Setch- ell) ; Unionville, Aug. 26, 1902; New Haven, Sept. 11, 1903; New Canaan, Sept. 29, 1903. Sometimes Ustilago spherogena occurs in the ovaries of the same plant with this. It can easily be distinguished from this by its larger sori, which are covered by a hispid mem- brane. Figure 40 shows the uppermost ovary only infected by the fungus. TILLETIACEZE $= Schrot. The sori form dusty erumpent spore masses, or are per- manently embedded in the plant tissues, often without evident distortion of these. The germination is by means of a pro- mycelium, which usually gives rise to a terminal cluster of elongated sporidia, which sometimes bear whorls of similar secondary sporidia; or the primary sporidia, with or without fusing in pairs, may give rise to infection threads, or in a nutrient medium to a mycelium bearing dissimilar secondary sporidia (aerial conidia). Fig. 22. Tilletia Tul. The sori occur in various parts of the host, usually in the ovaries, forming a dusty, dark spore mass. The spores are simple, separate, and originate singly in the ends of special: mycelial threads that generally disappear rather completely No. 5.] USTILAGINEE OF CONNECTICUT. 29 through gelatinization; they are of large size, 16-35y. Figs. II, 12, 39. This is the type of the family, and is one of the larger genera. The sorus is quite like that of Ustilago, but the spores are usually considerably larger. Saccardo records 53 species; 15 have been found in North America, but only 2 in Connecticut. The important stinking smuts of wheat, which cause so much damage in the western wheat districts, belong in this genus; but, as wheat is little grown in this state, neither species has been reported here. The smooth-spored species, Tilletia feetens (B. & C.) Trel., however, has been found, on microscopic examination, in ground cattle food offered for sale in the state, apparently made from smutted western wheat. The cattle did not relish this food. Key to Species of Tilletia. I. Spores reticulate.......... ccc eee ee eee T. Anthoxanthi. II. Spores apparently verruculose........... T. Maclagani. Tilletia Anthoxanthi Blytt. Figs. 12, 39. The sori are ovate, about 3 mm. in length, and usually occur in all of the ovaries of the spike; they are somewhat hidden by the envel- oping glumes, and at first are covered by a thin plant mem- brane, which eventually ruptures, disclosing the dusty, reddish black spore mass. Hyaline cells, or immature spores, are mixed with the spores, and are smaller than those, and have thin to medium-thick walls. The mature spores are reddish brown, ovoid to spherical, reticulate (3-64 wide and 1-3p deep), and 24-30p, occasionally even 34, in length. Host and Distr.: