Balansia and the Balansiae in America

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V BALANSIA AND THE BALANSIAE IN AMERICA

WILLIAM W. DIEHL Mycologist

Bureau of Plant Industry, Soils, and Agricultural Engineering

Agriculture Monograph No. 4

United States Department of Agriculture, Washington, D. C. December 1950

For sale by the Superintendent of Documents, U. S. Government Printing Office Washington 25, D. C. — Price 30 cents

I ACKNOWLEDGMENTS

Indebtedness, especially for the privilege of examining herbarium materials, is acknowledged to the following: The late Roland Thaxter and D. H.

Linder, of Harvard University; F. J. Seaver, of the New York Botanical Garden; Miss E. M. Wakefield and the late Sir A. W. HiU, of the Royal Botanic Gardens, Kew; H. D. House, of the New York State Museum; E. B. Mains, of the University of Michigan; C. W. Dodge, of the Missouri Botanical Garden; L. W. Pennell, of the Philadelphia Academy of Sciences;

J. H. Miller, of the University of Georgia; R. M. Harper, of the Alabama Geological Survey; E. West, of the University of Florida; B. C. Tharp, of the University of Texas; and R. E. D. Baker, of the Imperial College of Agriculture, Trinidad. Thanks are due to the many persons who have sent me living materials or personal herbarium specimens: To H. A. Allard,

I. H. Crowell, R. W. Davidson, G. D. Darker, G. B. Sartoris, J. L. Seal, G. F. Weber, E. West, and A. S. MuUer. I especially grateful F. am to Mrs. Agnes Chase, J. R. Swallen, J. Her- mann, and the late A. S. Hitchcock, for their generosity in identifying grass host plants and for the gift of numerous specimens that have come to their hands. It is desired also to express a sincere sense of obligation to the late A. B. Seymour for the privilege of examining the unpublished catalog records of fungi of the Farlow Library. To Mr. J. A. Stevenson I am sincerely appreciative of his long-time interest in furthering this work and for the use of the facilities of the Mycological Collections of the United States Department of Agriculture and the fungus collection of the United States National Herbarium, both now housed at the Plant Industry Station, Beltsville, Md. The work involved was undertaken under the spon- sorship of Prof. W. H. Weston, of Harvard University, to whom I am deeply grateful for helpful suggestions as well as sustained interest and inspira- tion. It is, furthermore, only proper to acknowledge very real gratitude to Mrs. Rose Beam for patience and special care in typing this manuscript. F. L. F. Foubert, L. I grateful To J. Brewer, M. and R. Taylor am very for their painstaking and sympathetic preparation of certain key photo- graphs used in the plates, as well as for many incidental courtesies: To Mr. Brewer for plates 1; 2; 3, D and E; 5, C, D, F, I, and /; 8, A, B, C, and D; 10, A and B; and 11, B; to Mr. Foubert for plates 3, B; 4, A, B, C; and 7, D; to Mr. Taylor for plates 3, C; 5, E and G; 6, C and E; 7, B, C, and E; 8, E, F, and G; and 9, A, B and C. I am indebted for plate 10, C, to M. M. L. Jaeger, and for plate 5, A, B, and H, to the C. G. Lloyd Myco- logical Collection of the Smithsonian Institution. The rest of the illustrations are my own. W. W. D. UNITED STATES DEPARTMENT OF AGRICULTURE Agriculture Monograph No. 4 Issued December 1950

Washington, D. C.

BALANSIA AND THE BALANSIAE^ IN AMERICA'

By William W. Diehl, jnycologist, Division of Mycology and Disease Survey, Bureau of Plant Industry, Soils, and Agricultural Engineering, Agricultural Research Administration

CONTENTS

Page Page

Introduction 1 Systematic account—Continued Historical background 2 The genus Balansiopsis 54 Morphological basis for the taxonomy Doubtful species or species not of the genus Balansia 3 eluded in the Balansiae 59

Criteria 3 Host relations in the Balansiae. . . . 62

t The mycelium and the hypothallus . . 3 Host specificity 62 The conidial fructiJScation, the ephe- Teratology 63 lidial stage 5 Mode of infection and dissemina- The ascostroma 7 tion Sequence of fructifications and the Incidence, epidemiology, and control 65

concept . . . of the genus Balansia. Susceptibility and resistance 66 Taxonomic position of Balansia and Recovery from infection 66 related genera Climate and geographic distribution . . 61 The Balansiae and the Clavicipita- Temperature relations 61 ceae Moisture relations 68 A classification of the genera of the Clavicipitaceae Phenology 68 Endemism Host relations in the Clavicipi- 68 toideae Economic considerations 69 Systematic account Bibliography 71 The genus Balansia Host index 77 The genus Atkinsonella 48 Index of fungus names 81

INTRODUCTION

H Balansia, Dothichloe, arid Ophiodothis general there is a distinct group or tribe are generic names that have been applied of species in America that have not been rather indiscriminately to certain species found in Africa or in Eurasia. of Clavicipitaceous fungi thus far known This monograph is restricted to a con- only as parasites of grasses and sedges. sideration of American^ species of these These species are found in various parts genera in the tribe here recognized as the of the world, most frequently in the Balansiae. It is based on observations of Tropics and Subtropics, and the evidence these fungi, where possible on their liv- of herbarium material suggests that in ing host plants in the field and in experimental culture, as well as the examination of herbarium ^ Exclusive of the genus Epichloe. materials, where available, 2 Submitted for publication May 11, 1950. from various institutions, *- and through the ^ The word "American" " here' used--^ for both North and South America. courtesies of many individuals. AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

In this account special consideration is cations and their sequence should be re- given to the peculiar pleomorphism of quisite in the interpretation and classifica- species of the Balansiae in recognizing tion of the Clavicipitaceae; hence, in this the several genera and in relating them study special effort has been made to take to the family Clavicipitaceae. Except for account of them. It is always desirable in the genus Claviceps the genera of this studies of fungi to observe them in pure family have not been clearly distin- culture (in vitro) where that is possible, guished, partly because many species, but various attempts at cultivating the even generic types, have been described species of Balansm on agar media have from limited material, usually ascomyce- thus far been unsuccessful, so that culture tous fructifications only. Conidial fructi- studies noted here have been limited to fications, sometimes present in herbarium plantings of infected grasses kept under specimens of Clavicipitaceae, are usually observation in the greenhouse and lacking, partly due to their ephemeral na- garden. Although it would have been de- ture but chiefly because of the practice of sirable for all the species considered to collectors and of herbarium technicians have been observed in culture, this has in preserving only the more conspicuous been possible for but eight of them. structures. As previously pointed out These eight species may, however, be (Diehl, 1930b)* these conidial fructifi- considered representative.

HISTORICAL BACKGROUND

In his original description of the genus studies by Moller (1901) and others of Balansia Spegazzini (1885) did not dif- the period antecedent who had described ferentiate the genus from Claviceps by species of Balansia for which no "pseu- any criteria other than the black color of dosclerotium" was noted. the ascostroma and the position upon the Atkinson's genus Dothichloe was not, inflorescence of the host plant of the however, described in sufficient detail to sclerotic structure from which it arises. have been accepted by Moller or by most In Claviceps, however, the ascostroma is other students of these fungi. These said to be colored and the sclerotic struc- workers have, in general, without- ex- ture from which it comes is developed planation eliminated the genus from con- within the ovary wall of the living host sideration and have at the same time con- plant. sidered species of Dothichloe under the Atkinson (1905), in revising the genus Ophiodothis of Saccardo (1883) genus Balansia, drew special attention to in the Dothideales. Hohnel (1910) has the fact that the pseudomorph from recognized the inclusion of species of which the ascostroma arises contained Dothichloe in the Hypocreaceae, al- both host and fungus tissues; he termed though he also considered the genus in- this pseudomorph the "pseudosclero- valid, and on what seems ill-defined tium" and distinguished the genus by premises he proposed the genera Balan- means of it. He noted here that in the siopsis and Linearistroma. Theiszen and two species of Dothichloe previously de- Sydow (1915) have chiefly reiterated the scribed by him (1894) the homologous conclusions of Von Hohnel. sclerotic structure was less developed, Considerable confusion has resulted merely forming an effuse layer on limited from reference of certain Clavicipitaceous surfaces .tissue. curious- of host Most species to the genus Ophiodothis Sacc, ly, Atkinson seems to- have been unac- although Atkinson (1905) would seem the extensive and detailed quainted with to have shown that this generic name is therefore unrec- ' The bibliography (p. 71) contains all references a nomen dubium and to literature given in the text. The year following ognizable. Ophiodothis was proposed by author's name is the key to the citation. BALANSIA AND THE BALANSIAE IN AMERICA

Saccardo (1883), based on Dothidea for Balansia trinitensh and Atkinson vorax Berk. & Curt. While it is possible, (1905) had recognized a similar fructifi- even likely, that this Asiatic species is cation in B. hypoxylon (Pk.) Atk. A congeneric with Balansia of Spegazzini conidial fructification was originally (1885), which it antedates, the type described for B. cyperi by Edgerton specimen of D. vorax at Kew is inade- (1919), while Rehm (1889) and Weber quate evidence because of its poor con- (1924) have reported ascostromata for dition. Ephelis, and the present writer (1930b) None of these accounts considered has called attention to the possible ad- seriously any morphological structures vantage in considering conidial fructifi- other than the pseudomorphs and asco- cations in the taxonomy of both Balansia mata to be of taxonomic value, although and Doth'ichloe. It seems desirable to the scolecosporous ephelidial stage was include all observed structures in the described by Cooke and Massee (1889) taxonomy of these species. MORPHOLOGICAL BASIS FOR THE TAXONOMY OF THE GENUS BALANSIA Criteria

The traditional criteria for distin- recognized as a synonym of B. claviceps guishing Balansia, here considered the Speg., had previously been pointed out type of the Balansiae, from related by Cooke and Massee (1889), and the

genera as expressed by ' the Saccardian sphaceiial stage as the accessory fructifica- heritage in the Clements and Shear tion of Claviceps had long been appreci- (1931) key or by the somewhat similar ated as of special taxonomic significance. treatment of Lindau (1897) have the In the present treatment the conidial virtue of simplicity in being restricted stage referable to the form-genus Ephelis,

to characters afforded by the ascostroma here termed the "ephelidial" stage, is

and the structure from which it arises. considered more important in defining

Despite a mistaken statement in Lindau's the genus than is the ascostroma alone, key, he is in agreement with Clements since in certain other genera, as Clavi- and Shear (loc. cit.) that the sclerotium ceps, for instance, the ascostromata are includes tissues of both host and fungus. practically identical with those of some

They do not take cognizance of the role species of Balansia. Consideration is also of accessory fructifications, although the given to the taxonomic value of the relation of Ephelis trinitensis, the coni- sclerotium or pseudoscierotium, here dial stage of Balansia trinitensis here termed the "hypothallus." The Mycelium and the Hypothallus For those species of Balansia observed in living condition, the affected host normally has failed to produce any natural inflorescence and new shoots from the same plant have continued to bear the fungus. This evidence of systemic infection is supported by the presence of characteristic hyphae (fig. 1) in nodes and to some extent in internodes and even in leaves of plants bearing the fructifications of the particular species concerned. These hyphae are most readi- ' ly detected in meristematic tissues and in /o7* ' Figure 7. — Endophyfic mycelium of Balansia intercellular spaces. Although difficult to obfecta within node of Cenchrus echinatus. AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE find, they can frequently be seen within The hypothallus of one species of the middle lamellae between cells. The Balansia may differ markedly from that vegetative hyphae in all species are of of another: In some it is persistent and very small diameter, usually 1/x to 1.3 fi, extensive; in others ephemeral and slight. ordinarily not exceeding 2/x, but where It is always present previous to the they are massed to form a superficial formation of any fructification, either as layer their diameters may be as much as a film of hyphae or as a robust stromatic

3fx. Many seeds of Cenchrus echinatus. mass. It is found either in florets as in suspected of being infected by Balansia, two species, superficial upon host tissues when sectioned and examined critically, in some species, or highly developed as have been found to bear vegetative my- the fungus component along with the celium like that of infected hosts. aborted or mummified inflorescence in The hyphae aggregated at certain- the relatively large pseudomorphs char- points on the surface of the hosts, as on acteristic of Balansia clariceps, B. cyperi, leaves or surrounding the culm or the and B. hypoxylon. The surface and con- inflorescence, become the basal layer text are white or yellowish at first but from which the fructifications arise. This may become yellow or brown to black. basal layer, when it includes host tissues In all cases the tissue of the hypothallus intermingled, is a pseudomorph that has is a compact stroma. sometimes been termed a "sclerotium." When a bit of the hypothallus of any Atkinson (1905) used the term "pseu- of these species is examined in sterile dosclerotium" for this sclerotic basal water in a microscopic preparation, stroma containing fragments of host clouds of minute bacterioid bodies are to tissue and utilized this feature in distin- be seen streaming out into the surround- guishing the genus. His term is not ap- ing medium. These minute bodies are plicable to the fungus equivalent in his not motile but are particularly evident genus Dothichloe, which does not con- because of Brownian movement. They tain host tissues intermingled, although are subspherical and do not exceed 0.5/x it does apply to the structure in the type in diameter. In those species of Balansia species of the genus Balansia. where the black rind of the hypothallus

The sclerotic basal stroma in some of is particularly well developed, these the species here under discussion, in- minute bodies appear to make up a great cluding those referred by Atkinson to share of the bulk of the rind, where they

Dothichloe, is a simple prosenchymatous, are first attached to the peripheral hyphae rarely pseudoparenchymatous, layer over- (fig. 2). No attempt is made here to lying the surface, usually of but a small explain the occurrence of these minute area, of host tissue. To call this basal bodies, and it is convenient to regard layer a "sclerotium" or "pseudosclero- them as minute granules or concretions. tium" seems inadvisable, as these terms imply a resting stage and other concepts not normally applicable to species of this genus. Despite a certain etymological disadvantage, the term "hypothallus," applied hitherto to this type of structure, will be used in the present discussion of the species. Although seemingly not previously applied with reference to Balan- siae, it was accepted by Von Hohnel ' '^ ' (1909) for the homologous structures 10 Figure 2.— Vertical section through rind on peri- in Konradia and Oomyces following the phery of hypothallus of Balansia obtecfa, to usage of Raciborski and Moller (1900) show structure of closely appressecJ hyphae (1901). and detachable minute spherical bodies. BALANSIA AND THE BALANSIAE IN AMERICA

The Conidial Fructifications, the Epheiidial Stage

Ellis and Everhart (1886) reported Ephelis borealis as a stage of Balansia hypoxylon, and Cooke and Massee (1889) described E. trinitensis as the conidial fructification for B. trinitensis here considered the same as B. claviceps, the type species of the genus. Since that time some consideration has been given to these conidial fructifications of Balan- sia — in America chiefly by Atkinson (1905), Edgerton (1919), Weber (1924), and Diehl (1930b, 1939); in Europe by Von Hohnel (1913, 1925); and in India by Sundararaman (1921). Of genera with similar ascostrom.ata only Balansia, Dothichloe included, has been known to possess this type of fructification. ;-; i: f-; \:- The ephelidial fructification is an %wm0^:-^ apically expanded pycnidium or acervu-

lus that is erumpent from beneath the surface of the hypothallus. The typical ephelidial fructification of Balansia claviceps emerges from the hypothallus as a protuberance that expands rapidly, open- ing above into a cupulate and ultimately S l.v patellate structure with the opened, glis-

1"; '^: tening, inner surface lined with an hy- ^:i 'i4i menial palisade of scolecosporous conid-

ia. The periphery of one of these struc-

tures is slightly hairy to smooth and resembles superficially the exciple of a :^1':!!:.M--I '}^t ^^r^ discomycetous apothecium. The excipular hairs when present may be conspicuous in some species, and in certain others ' '" none have ever been observed. In '-•'' '•.; some ' 1 '/I jk^" species the fructification appears as a mere crack, usually elongated-hysteriform, sometimes furcate, with the thin, folded-back surface of the hypothallus 1 im Tn,| ^

as an exciple (fig. 3) . In one species the pycnidium is reduced to a mere pit in Figure 3. — Ephelidial fructifications of Balansia erumpent the hypothallus. The hysteriform fructi- ambiens as irregular whitish hymenia from and covering hypothallus. fications are usually very ephemeral and, being less conspicuous than the more ence of septa, which, although usually emergent types, easily escape notice. not discernible even with the higher The conidia are acicular, hyaline, powers of the compound microscope, are waxy-glutinous, and characterized by indicated by one or two slight constric- peculiar indentations that are difficult to tions and a general tendency to break explain unless they are due to the pres- apart at those points. But in Ophiodothis

I AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE linearis, here referred to Balansia, defi- each branch identical with the simple nite septa are demonstrable. It should be type. At each of these apices a single noted that the acicular conidia of six conidium is first formed, followed by a species of Balansia have been germinated succession of additional conidia pushed on numerous occasions in tap water, in off in turn. The mass of conidia under rain water, and on the surface of sterile moderately dry conditions will eventual- agar observations media, and frequent ly form a cirrus (see pi. 4, A) , which, have been made of -these germinating however, is dissipated on contact with conidia. In all cases the germination, as water. In certain species, where herb- well as the appearance of the conidia, arium specimens only have been seen, was so much of one type that it was not simple conidiophores have been noted; ordinarily possible to distinguish any but in those species observed in living difference between the conidia of these condition the conidiophores were often species. Generally a single apical germ swollen and irregularly branched. In tube is produced (fig. 4), although Linearistroma linearis they were noted lateral germ tubes have been observed in an herbarium specimen as branched also. but not swollen. Diehl (1930b) has pointed out the homology of the ephelis- like conidia of Dotbichloe with those of Balansia, but failed to report the presence of the very ephemeral peridia in the O conidial fructifications of species referred to Dotbichloe that are here regarded as homologous with the exciple of the ephelidial fructifications of Balan- sia. In the present treatment these thin peridia are thus considered to be excipular and the scolecosporous conidial

stage of Dotbichloe is regarded as an

ephelidial stage of Balansia, treating it as a genus with considerable variation in the form of the ephelidial structure. In Diehl's (1930b, 1939) discussions some emphasis was given to observations upon three species of Dotbichloe, which when kept moist immoderately long, produced masses of ovate conidia upon bottle- shaped conidiophores similar to those of Epichloe typhina. Later observations have brought out the fact that in the species then referred to Dotbichloe these ovate conidia were produced only under extraordinary conditions and are not ordinarily present. On the other hand in

Figure 4.— Ephelldial conidia of Balansia obfecfa Balansia bypoxylon, here segregated germinating in tap water. from the genus, they are always present first in the sequence of fructifications. The conidiophores of the Balansia on Cenchrus echinatus vary from a simple The function of ephelidial conidia is hyphal tip to a complex, much swollen, presumably the same as that of ascospores and irregularly branched structure of in disseminating and propagating the considerable size, but with apices of species. They are waxy-gelatinous to BALANSIA AND THE BALANSIAE IN AMERICA

glutinous, easily adhering to various ob- in systemically infected plants. It is also jects on contact. They could adhere to possible that spattering rain could effect insects and thus be transported to stig- conidial dispersal, since the conidial mas of florets of susceptible hosts where masses readily separate in water. As initial infection evidently takes place. As ephelidial fructifications materially out- noted in the following discussion of one number the ascostromata in certain spe- species of Balansia (p. 25), germinating cies of Balansia it would seem that conidia when placed upon stigmas of a ephelidial conidia might be of greater susceptible host brought about infected significance in dissemination than are seeds, which when germinated resulted ascospores.

The Ascostroma

Balansia claviceps Speg., the generic features the life histories of species of type, B. cyperi, and B. hypoxylon are Dothichloe and Balansia are similar, characterized by capitate ascostromata justification for so distinguishing these that follow a well-defined ephelidial genera is lacking. Although a particular

stage, sometimes arising within it. These species of Balansia, as here treated, may capitate ascostromata are arranged upon be characterized by ascostromata of a

the hypothailus, that here covers and is specific form or orientation with respect intermingled with the tissues of an un- to the hypothailus, these features are developed inflorescence of the grass host. not regarded as of generic significance. These ascostromata in the type specimen These ascostromata have certain fea- of B. claviceps are definitely long-stipi- tures in common: In most of the species tate (pi. 1), but in many instances, as there is a tendency for them to become in part of the type of B. trinitensis (pi. brown to black when mature, although 2, C), the stipes are very short. If one in one South American species they are examines a goodly number of the speci- yellow. In all species the ascostromata mens here referred to B. claviceps, it is contain an indefinite number of ostiolate obvious that in this species the length of perithecia with aparaphysate asci arising the stipe varies considerably. In other from a fertile area at the base, the asci species, as B. discoidea P. Henn., for in- characterized by a slime-cap and filiform stance, the stromata vary from capitate ascospores. Except for the tendency of to flattened-pulvinate, and they some- stromata to blacken with maturity, these times coalesce. When capitate the sterile are all features characteristic of Claviceps base of each ascostroma is a short stipe, and other related genera as well. but when pulvinate and coalescent the The blackening of the ascostroma in combined ascostromata are clearly an most species is limited to the outermost effuse layer — the kind of stroma that is cells, the prosenchymatous to pseudo- said to distinguish the genus Dothichloe. parenchymatous context being whitish in Actually every gradation between the some species, yellowish in others. The capitate and even short-stipitate stromata perithecia of one species may be com- of B. discoidea and the effuse stromata pressed by crowding and of another of D. nigricans can be found, so that the relatively free and well-rounded. Al- two species are merely extremes of one; though with the development of the i.e., B. discoidea is an extreme condition, perithecia the walls show the effect of a form or variety of the other. To dis- pressure as compared with the tissues of tinguish the genus Dothichloe from the stroma proper, they retain their in- Balansia solely because the stroma in the dividuality. When an ascostroma is still former may be effuse rather than broken tender and turgid before the maturity up into several heads as in Balansia of the asci the several perithecia may seems questionable; and since in all other readily be squeezed out with walls intact AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE if the stroma is crushed in water under the characteristic periphyses. In the base a cover glass (pi. 10, E) . Actually the of the perithecium there is usually a v/ails of these fungi, as shown by Balan- raised region or basal coUiculus of par- sia claviceps, B. hypoxylon, and Dothi- tially gelatinized prosenchyma from chloe epichloe, are formed from the which the asci have arisen. Interthecial stromatic cells by compression and hyphae (paraphyses) gelatinize early and stretching into a layer surrounding the have disappeared by the time the asci are developing asci and paraphyses that re- evident. The asci extend from the basal sem.bles a true perithecial wall. This wall colliculus to the base of the ostiolar is usually less distinct at the base of the canal. When living ascostromata are perithecium than at the sides and in the kept in a moist chamber the tips of asci region of the ostioie. The condition is may be seen to extrude from the ostioie. comparable to that shown by Vincens The asci are cylindrical-clavate, nor- (1917) for Claviceps and Epichloe and mally with eight filiform ascospores, by Killian (1919) for Claviceps. Re- although in some instances a reduced gardless of the origin of the walls, they number, usually two or four, have been so simulate those of true perithecia that observed. The spores are somewhat they are so termed in the present dis- coiled within the ascus and are ejected cussion. through an apical pore surrounded by a hyaline apparatus known as the slime- cap (fig. 5). The ascospores develop septa before or at germination. At germination each cell may form one or more germ tubes perpendicular to the axis of the spore.

The manner of spore ejection is very likely the same in all species of this genus and of those genera closely related. In Balansia hypoxylon and Dothichloe epichloe the ascospores have been observed to "squirt" from ostioles when matured stromata in a moist chamber were exposed suddenly to a beam of light directed from an electric arc. The ascospores of all the species of Balansia are delicate, so gelatinous and glutinous that they adhere readily to any object

upon which they have been ejected. It is likely that they, as well as ephelidial conidia, which are of like texture, can adhere to insects that visit grass florets Figure 5.—An ascospore, an ascus containing and thus be transported to stigmas of eight ascospores, and swollen (germinating) susceptible grasses where infection is segments of an ascospore of Balansia linearis. thought to take place. In this respect The upper part of the perithecial wall, there would seem to be an exact parallel including the ostiolar canal, is lined with with the condition in Claviceps.

Sequence of Fructifications and the Concept of the Genus Balansia

Normally in Balansia claviceps the marked, but in certain other species the sequence of hypothallus, ephelidial fruc- hypothallus is scant and the ephelidial tifications, and ascostromata is well fructification so ephemeral that both are BALANSIA AND THE BALANSIAE IN AMERICA

early obliterated and covered by the terial delay. ascostroma. Nevertheless, there are in- The genus Balansia is thus defined by stances where, possibly due to moisture the production from an intramatrical relations, ephelidial fructifications con- m.ycelium of a superficial hypothallus tinue to be produced even after maturity from which arise the conidial fructifica- of ascostromata. The concept of the tions characteristic of the form genus genus as here considered would require Ephelis, followed by or synchronous with ephelidial fructifications regardless of clavicipitaceous ascostromata that may their superficial appearance, orientation, show great variation in form or color. or sequence. The hypothallus in some Insofar as the species have been studied species may function as a resting sclero- in living condition, they show every in-

tium or not, without affecting its taxon- dication of obligate parasitism with sys- omic validity. Certainly in the type temic, endophytic, vegetative mycelia. species of the genus the hypothallus on The recorded hosts are all grasses or

one host {Setaria) produces its char- sedges, although other host relations are acteristic fructifications without any ma- entirely conceivable. TAXONOMIC POSITION OF BALANSIA AND RELATED GENERA The Balansiae and the Clavicipltaceae

Spegazzini's (1885) description of like ascostromata, although, like Epich- the genus Balansia recognized its close loe, possessing asci each with a "hyaline relation to the genus Clavkeps TuL; and apex" or slime-cap. the affinity of the two genera has never Atkinson (1894) noted the possibility since been questioned. The genus was of the relation of his genus Dothichloe accepted by Saccardo (1891) in the to Ophiodothis of Saccardo (1883) and Sylloge, and a few years later Lindau later (1905) demonstrated the likeli- (1897) classified it along with ten other hood that Ophiodothis and Balansia of genera in his subfamily Clavicipiteae of Spegazzini (1885) were synonymous, the family Hypocreaceae. The subfamily although he erroneously credited Spegaz- was recognized primarily by the presence zini's publication to the year 1880 as of of filiform ascospores in fruit bodies prior date. He concluded that, since the (perithecia) not necessarily sharply de- type specimen of Ophiodothis vorax limited within a hypocreaceous stroma. (Berk. & Curt.) Sacc. on which the By definition the mature stromata in the genus was based was in poor condition family Hypocreaceae and the order and inadequate for descriptive purposes, Hypocreales were bright-colored and of the genus Ophiodothis was invalid, a a fleshy or waxy consistency, features decision that has since been acceptable lacking in most species of Balansia thus to most taxonomists of these genera re- far known and not necessarily present in gardless of Atkinson's confusion as to some related genera. Certain other genera their publication dates. Atkinson (1905) included by Lindau have since been took account of the "cap-cell" of the shown to be unrelated, but the sub- ascus and of the conidial stage referable family has continued to be recognized, to the form-genus Ephelis as taxonomic nevertheless, as a unit typified by the features in Balansia, although he stressed well-known genus Claviceps. Lindau the worth of the "pseudosclerotium" placed Ophiodothis of Saccardo (1883) (pseudomorph consisting of both host in the Dothideales, yet failed to include and fungus) in delimiting the genus. in the Clavicipiteae Atkinson's (1894) This latter feature is not considered in genus Dothichloe, which had been de- the present discussion as of so great a scribed as a close affinity of Epichloe but significance taxonomically. separated from it by its black, Dothidea- InSeaver's (1910a,b, 1911) treatment 10 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE of the Hypocreales the four genera lated to each other and so distinct from Oomyces, Barya, Typhodium {Epich- the genera usually considered in the loe), and Hypocrella are placed in his Hypocreaceae that the Clavicipiteae may tribe "Hypocreae" and Ech'modothis in properly be considered a family. "Clavi- his tribe "Nectriae," leaving in his tribe cipitaceae" has been proposed by Earle "Cordicipiteae" but three genera: Cordy- (1901) and by Nannfeldt (1932), an(^ ceps, Spermoedia {Clavkeps), and Ba- recognized in Bessey's (1935) textbocK lans'ia. Von Hohnel (1910), however, as well as in Miller's (1949) revision of recognized that the Clavicipiteae were Ascomycete classification. characterized by scolecosporous asci with The characters of the family do not a terminal "slime-cap" in perithecia in commend its retention with the Nectri- stromata that might be black in color, aceae and Hypocreaceae in the dubious since he included his new genera Balan- order Hypocreales; but whether the siopsis and Linearistroma, which have origin of the perithecial wall in the black ascostromata, although he con- Clavicipitaceae would exclude the family sidered the "tribe" to be in the Hypo- from other Pyrenomycetes requiring ac- creaceae. ceptance of Nannfeldt's (1932) order Gaumann (1926), like Spegazzini, Clavicipitales is problematical. Gau- Atkinson, and Von Hohnel, recognized mann's (1949) recent treatment recog- the Clavicipiteae in the Hypocreaceae by nizing Nannfeldt's order Clavicipitales the slime-cap of the scolecosporous ascus, retains his previous concept of the three although he followed the tradition of phylogenetic groups, presumably within distinguishing the order Hypocreales by the single family Clavicipitaceae. For the the criteria of color and texture, never- present it is convenient to aline it with theless including Balansia with its black the Sphaeriales and it is likewise con- ascostromata. He, however, proposed a venient to accept Gaumann's subdivision revolutionary concept of three groups of into three groups which may be regarded a subfamily, each group supposedly rep- as subfamilies, termed as follows: (1) resenting a phylogenetic unit: (1) The Oomycetoideae, (2) Clavicipitoideae, Oomyces-Ascopolyporus group; (2) the and (3) Cordycipitoideae. Epichloe-Clavkeps group; (3) the Cor- It is questionable whether any natural dyceps group, comprising Cordyceps and system of classifying the genera in the by implication segregates from that Clavicipitaceae can be determined by genus. utilizing as criteria only ascostromatic

Although there may be some difficulty features. It is, therefore, to be expected in accepting Gaumann's expressed justi- that recourse be had to characters pro- fication for this grouping and his in- vided by conidial fructifications, which clusion of certain genera in group (1) may well be as advantageous, even as rather than in group (2), it is obvious requisite here as in the classification by to anyone having an intimate and long- Nannfeldt (1932) of those many oper- time acquaintance with species of these culate D'ncornycetes, \vhere comparable genera that Gaumann's arrangement is difliculties in segregating and defining indicative of three divergent evolution- genera have been solved by the use of ary trends in the development of the the characters secondarily provided by ascostroma. As such, it may foreshadow conidial structures. It is to be expected an arrangement of these genera in an also that conidial structures are of simi- ultimate natural system of classification lar significance in the taxonomy of many that will reflect their phylogeny. All the Pyrenomycetes. Indeed, in the Clavicipi- genera in these three groups, together by toideae the genera Epichloe, Claviceps, virtue of the slime-cap character of the and Balansia have conidial fructifications scolecosporous ascus, are so obviously re- especially convenient for this purpose. BALANSIA AND THE BALANSIAE IN AMERICA 11

Far too little is known as yet about the clavate to linear phialides bearing glut- stages of the various genera of the inous or "wet" conidia. The conidial Oomycetoideae to make a comparable stage of Balansia as discussed previously

assertion, but from the evidence now is recognized as the very distinctive but available this would seem most likely variable scolecosporous form-genus also. Considerable knowledge has been Ephelis, here termed the "ephelidial accumulated relative to the life histories stage." of species in the Cordycipitoideae, and As has been noted by Diehl (1930b) it is to be expected that the future classi- and considered previously in the present fication of that subfamily will make discussion, certain species of Dothichloe, ample use of conidial structures. To con- here referred to Balansia, under some sider these seeming secondary features conditions of moisture develop ameros- in the classification of the Clavicipitoi- porous conidia as in Typhodium, but deae would be in line with present trends seemingly only under unusual condi- in the treatment of the other two sub- tions. Since typhodial stages are normal- families. ly lacking in those species, they are here In the Clavicipitoideae as here recog- considered as of negligible significance nized there are certain species that do in the taxonomy of the genus Balansia. not appear to produce any conidial fruc- On the other hand Balansia hypoxylon, tifications; they can readily be allocated as was reported by Diehl (1930), regu- with the genus Balansiopsis, of which larly develops the typhodial stage as a the type species B. gaduae has no recog- primary fructification covering the hypo- nizable conidial structures. The little- thallus. This sequence of fructifications, known genus Balansiella has been rele- typhodial stage, ephelidial stage, and gated by Clements and Shear (1931) to ascostroma, is here recognized as justi- synonymy with Claviceps, a disposition fication for the concept of a new genus justified by features reported in the fol- Atkinsonella, proposed in honor of G. F. lowing treatment. Atkinson, whose contributions have Claviceps is recognized by all investi- marked an epoch m the study of species gators as having conidial fructifications of Balansiae. referable to the tuberculariaceous form- it is noteworthy that, de- genus Sphacelia with glutinous conidia, Incidentally, justified while the conidial fructification of Epi- spite Atkinson's (1897, 1905) disposition of his Dothichloe as a close chloe is referable to the distinctly dif- and ferent form-genus Typhodium Lk.^ relative of Balansia and Claviceps of Ophiodothis as in- There are numerous references to this his elimination valid, certain students of these fungi ovoid-amerosporous stage where it is the names called the "sphacelial stage," following have continued to include Ophiodothis in the the usage of Saccardo (1882) in naming Dothichloe and of the this stage Sphacelia typhina. Actually Dothideales, probably because of the this "typhodial" fructification of Epi- connotations of the black color of the names chloe is a smooth, eff^use layer or palisade stromata and the etymology of clavate-lageniform conidiophores rather than because of the nature or Theiszen (phialides), each with an apiculate origin of the perithecial wall. sterigma bearing "dry" conidia, whereas and Sydow (1915), in their monograph nevertheless the sphacelial fructification of Claviceps of the Dothideales, have allo- is a much-wrinkled, effuse palisade of accepted Atkinson's (1894, 1905) cation of Ophiodothis to Balansia and ' This use of Link's (1826) generic name Typho- Hypocreaceae. They dium as a form-genus name is a present convention of Dothichloe to the of convenience. Link's meager description along with recognized also the close relationship in his illustration actually suggests a stage in stromatic development where immature perithecia are present the same family of the genera Balan- but in which the conidial stage would still be chiefly in evidence. siopsis and Lin earistroma. Poroniopsis 12 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE described with one species, P. bruchi, by spores justifies its recognition as the Spegazzini (1921) has been considered monotypic representative of the tribe a synonym of Clavkeps by Clements and Clavicipiteae, thus restricting the tribe Shear (1931), although Spegazzini's name at present to this single genus, photograph of the type specimen brings The rest of the genera in the subfamily that disposition into question. It is here distinguished by species with different suggested that Poroniopsh may more kinds of conidia, or none, are referred properly be alined with the Cordycipi- .to the tribes Balansiae and Ustilaginoi- toideae. deae. The latter tribe is distinguished by The subfamily Clavicipitoideae may the peculiar smutlike conidia (radula- be divided into three tribes, Clavicipiteae, spores) borne pleurogenously on closely Balansiae, and Ustilaginoideae, based on packed parallel hyphae, whereas in the the fundamental differences in conidia. Balansiae the conidia if present are dr}' The distinctive sphacelial state of Clavi- phialospores. ceps with the "wet" or glutinous amero-

A Classification of the Genera of the Clavicipitaceae

SUBFAMILY 1. OOMYCETOIDEAE Oomyces ? Barya {Globulina) Konradia ? AUtosporium {Aciculosporium) Hypocrella M.ycomalus ? Hypocreophh Ascopolyporus SUBFAMILY 2. CLAVICIPITOIDEAE Tribe 1. Clavicipiteae Clavkeps (Balansiella) Tribe 2. Balansiae Balansia {Dothtchloe, Linearistroma, ? Ophiodothis) Atkinsonella Epichloe Balansiopsis Tribe 3. Ustilaginoideae Munkia {? Shropshiria, ? Neomunkia) Ustilaginoidea SUBFAMILY 3. CORDYCIPITOIDEAE Cordyceps Torrubiella Dussiella {Echinodothis) Ophiocordyceps Poroniopsis ? Coscinaria

Key to the Clavicipitoideae and the genera o] the ^alansiae

(Based primarily on conidial fructifications)

a. Conidial fructifications not developed . Balansiopsis. aa. Conidial fructifications normally present previous to development of ascostromata. b. Conidial fructifications sphacelial CLAViaPiTEAE. bb. Conidial fructifications otherwise. c. Conidial fructifications both typhodial and ephelidial Atkinsonella. cc. Conidial fructifications either typhodial or ephelidial, but not both. d. Conidial fructifications typhodial Epichloe. dd. Conidial fructifications ephelidial Balansia. cii. Conidial fructifications ustilaginoid UsTlLAGINoroEAE.

Host Relations in the Clavicipitoideae

It is singular that with one possible genera in this subfamily are parasites of

exception all the species of the several grasses or sedges. That exception is Epi- .

BALANSIA AND THE BALANSIAE IN AMERICA 13

chloe hertonil, described and illustrated and shortening of internodes. Further- by Spegazzini (1922) from Paraguay more, all the evidence suggests the like- as an endoparasite with ascostromata lihood of seed transmission of these covering areas on stems of the composite, parasites, most likely initiated by ovule Mikanm scandens. The fungus has yellow infection by way of the stigma and style. ascostromata similar to that of E. typhina Investigations of species of Claviceps on grasses. But since its conidial stages, by various workers have shown initial if any, are unknown thus far, additional infection through the stigma, resulting information is needed to confirm this in ultimate localized infection and de- allocation. struction of the ovule. In species of Epichloe typhina has been studied Epichloe and Balansia thus far investi- critically by Sampson (1933) in Great gated the parasite normally hibernates or Britain. She has shown it to be a parasite aestivates over unfavorable seasons as an with intramatrical mycelium transmitted endophytic mycelium either in seeds or through the seed, causing a systemic in- in the living plant itself, but in Claviceps fection of the grass plant that emerges unfavorable seasons are survived by the from that seed. She has noted extreme localized fungus in the sclerotial stage variability in effects of the fungus on (hypothallus) within the dead and the host: In certain cases it causes com- transformed ovule wall. As will be noted plete sterility and in others only partial for the monotypic species of Atkinson- sterility of the host; and in some in- ella, ascospores within the perithecia may stances the tissues of normal-appearing sometimes overwinter intact, a condition grasses can be shown to contain mycelium that might also be expected for some of the endophyte. As will be noted in species of Balansia, Balansiopsis, and detail further on, in those species of Epichloe but less likely for species of

Balansia and Atkinsonella where they Claviceps. It is of special interest that have been observed critically, the effects species of Balansia and the other genera on the hosts parallel somewhat the effects here included in the tribe Balansiae are caused by E. typhina. All the hosts in- systemic endophytes, whereas the species fected with Balansia, Atkinsonella, and of the monotypic tribe Clavicipiteae are Balansiopsis, as well as those infected distinguished as endophytes with a with E. typhina, tend to be dwarfed, and limited mycelium infecting localized some are fasciated. They tend to have tissues only. This is a correlation that specific effects such as deformation of appears to be sheer coincidence without ligules, usually of the terminal leaf only, fundamental taxonomic bearing. dwarfing of terminal leaves of the shoot.

SYSTEMATIC ACCOUNT (Exclusive of the genus Epichloe) The Genus Balansia

Balansia Speg., An. Soc. Cient. Argen- Dothichloe Atk., Torrey Bot. Club tina 19: 45. 1885 (emend.) Bui. 21: 224. 1894. Type spe- Type species: B. claviceps Speg. cies: D. aristidae Atk. (lectotype) } Ophiodothis Sacc, Sylloge Fun- ^^Ephelis^ Fr., Summa Veg. Scand. gorum 2: 652. 1883. Type spe- 9: 370. 1849 et ap. Berk., Linn. cies: Dothidea vorax Berk & Curt. Soc. London, Bot. Jour. 10: 353. Hyalodothis Pat. & Hariot, Soc. 1869. Type species: E. mexicana Mycol. de France Bui. 9: 210. Fr. ap. Berk. 1893. Type species: Hyalodothis •'Names marked with an asterisk (*) refer to claviis Pat. & Hariot. form-genera applying to conidial stages. 14 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

"^Sphaceliopsis Speg., Buenos Aires doseptate, irregularly acicular conidia Mus. Nac. de Hist. Nat. An. 20: (scolecospores) that break up into two 457. (1911) 1910. Type species: to three segments, conidia borne upon S. cyperkola Speg. simple conidiophores often branched and irregularly swollen. Ascostromata arising hinearhtroma Hohn., Akad. der from the hypothallus, effuse to pulvinate Wiss. Wien. Math.-Nat. KI. Sitz- or capitate, sessile to stipitate, context ber. 119 (Abt. 1): 938-939. prosenchymatous to pseudoparenchymat- 1910. Type species: Ophiodothis ous, whitish to yellow, surface gray or linearis Rehm. yellow and generally becoming black Hypothallus or basal stroma scant or with maturity, punctate from, the numer- well-developed, prosenchymatous-sclero- ous ostioles; perithecia ovate-lageniform, tic to pseudoprosenchymatous, context immersed in one layer within the stroma white to gray or yellow, surface white or except for some ostiolar protrusion; peri- yellow to black, sometimes laccate, sur- thecial walls well defined, composed of rounding tissues of undeveloped inflores- compressed stromatic cells; ostioles usu- cences or forming a thin layer on limited ally colliculose, with filiform periphyses areas of culms or leaves or within florets lining the ostiolar region; asci arising of the host. Ephelidial (conidial) fruc- from the base of the perithecium, cylin- tifications arising from the hypothallus. drical, with slime-cap; ascospores hy- discoid-cupulate to hysteriform; exciple aline, filiform, and pluri-septate at ma- or peridium variable to thick, glabrous turit}^; paraphyses delii^uescent with de- to hirsute; hymenium of simple but pseu- velopment of the asci. SYNONYMY Ophiodothis of Saccardo (1883) was issue with Atkinson (1905) respecting based primarily upon Dothidea vorax, a the inadequacy of the Indian specimen specific name given by Berkeley and of D. vorax as a type or the confusion Curtis (1854) with an inadequate de- resulting from the fact that the original scription, i.e., from the modern view- description included at least one other point, of an imperfect specimen found species, a condition that renders D. vorax upon the spike of an unknown species a nomen confusum and in its train in- of Carex in India. Nothing is yet known validates the genus Ophiodothis. of the life history of this species and of Hyalodothis of Patouillard and Hariot its accessory fructifications, which are (1893), based on H. clavus Pat. & here regarded as requisite for generic Hariot from an African specimen definition. It has been the writer's privi- gathered by Dybowski in the Congo, is lege to examine the type specimen still another genus without precise definition. preserved at Kew, and it is recognized Von Hohnel (1910) and Theiszen and that the specimen may be referred Sydow (1915) concluded that the genus equally well to Balansia or to one of was invalid, since the type specimen several other genera. represented material referable to Ophio- Despite Atkinson's (1905) erroneous dothis and also contained perithecia of statemient to the contrary, the name a species of Hyponectria. The writer has Ophiodothis antedates Balansia and an examined type material in the Patouil- understanding of the life history of lard Herbarium at the Farlov/ Herbarium Dothidea vorax Berk. & Curt, could ulti- and considers it a species of Balansia mately disturb the nomenclatorial status identical with the African Epichlo'e of one or both of these genera on a basis volkensii P. Henn.; this same conclusion of strict priority unless recourse be had has been drawn also by Castellani and to the safety of genera conservanda. No Ciferri (1937). The species is of no one, however, has been inclined to take further concern in this account, as it is BALANSIA AND THE BALANSIAE IN AMERICA 15 not known to occur in America. kinson (1905) later attempted to vali- Dothichloe of Atkinson (1894) pre- date his genus by showing that the first sents unusual nomenclatorial difficulties, of the two species of 1894, which he and a special explanation is in order to apparently considered the type, was not justify the present choice of D. aristidae the same as E. hypoxylon Pk., but was Atk. as the type species. His concept of the same as Hypocrea atramentosa Berk. the genus embraces those species of & Curt., which he erroneously considered Balansia (as treated in the present dis- to be the same as Ophiodothis vorax var, cussion) having effuse stromata. His atramentaria Berk. & Curt. (Dothidea discussion included descriptions of two atramentaria Berk. & Curt. ) . The first species: D. hypoxylon (Pk.) Atk. as a confusion was compounded and made new combination and D. aristidae Atk. still more complex by the attempted cor- as a new species. The first name was rection. As will be noted later, however, based upon Epichioe hypoxylon Pk., a H. atramentosa Berk. & Curt., as applied different species to which Atkinson's strictly, is still another species in an en- description does not apply, he having in tirely different genus! Atkinson (1905) mind the species afterward described by did not realize that his second species,

Moller (1901) 2.S Ophiodothis henning- D. aristidae, serves much more satisfac- siana. Moller evidently appreciated torily as a generic type, because it lacks Dothichloe as a nomen confusum, since the nomenclatorial confusion inescapable he dismissed it briefly with the statement in the case of his first species. "Dothichloe Atk. ist 2u streichen." At- THE CONIDIAL (EPHELIDIAL) STAGE - THE FORM-GENUS EPHELIS The name Ephelis was proposed by the several American species here re- Fries (1849) w^ith the italicized ''¥r. corded as well as in at least two exotic Fung. Mexic." followed by a two-line species of the genus. technical diagnosis but with no cited Sphaceliopsis cypericola, the type of species. Twenty years later Berkeley Spegazzini's (1910) form-genus 5"^ /^^r^- (1869) supplied the description of E. liopsis, appears from his illustration and mexicana Fr., based upon a Mexican meager description to be the same as the specimen furnished by Fries, thus typify- ephelidial stage of Balansia cyperi later ing the species and citing for it a speci- described by Edgerton (1919). Spegaz- men now in the Berkeley Herbarium at zini's genus and species thus become Kew. The first recognition of Ephelis as synonyms. Spegazzini did not assign a conidial stage for Balansia was noted his form-genus to any particular position by Cooke and Massee (1889) in their among the Fungi-imperfecti, but Hohnel account of B. trinitensis, here regarded placed it in the Tuberculariaceae and as synonymous with the type species. referred Ephelis to his "patelloid Excipu- Since that time the association of the latae" of his family "Pycnidiaceae." ephelidial stage has been observed in THE SUBGENERA - EUBALANSIA AND DOTHICHLOE The subgenus Euhalansia'^ is here pro- moreover, the ascostromata are stipitate, posed for those species having patellate but in B. cyperi, which qualifies as to the to cupulate ephelidial fructifications with ephelidial fructification, the ascostromata well-defined peridia and only incidental- are sessile and pulvinate. ly for those with ascostromata that are The subgenus Dothichloe^ with the discretely capitate or pulvinate as in the concept of Atkinson's (1894) genus of type of the genus. In Balansia claviceps, that name, which is invalid as a genus

* '' Eubalansia subgen. nov.: Typifacto Balansiae Dotlotctolo'e subgen, nov. ; Typifacto Dothichloe clavicipitidis Speg. ; ascostromatibus pulvinatis capi- aristidae Atk. ; ascostromatibus ettusis vel pulvinatis tatis, sessilibus vel stipiratis ; fructibus ephelidibus praecipue sessilibus; fructibus ephelidibus effusis vel papillaris vel cupulatis. elongato-hysreriformibus connatis. 16 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE nomenclatorially, typified by D. aristidae, nomen confusum, but it is obvious that is proposed to comprise those species Atkinson had in mind the same species with effuse to hysteriform ephelidial described by Moller (1901) as Gphio- fructifications and with ascostromata dotis henningsiana. Atkinson's second typically effuse but varying from effuse species, D. aristidae, closely related to to capitate-stipitate. There may be some that species and without its nomencla- question as to the choice of a type for torial disadvantages, is adequate as a Dothichloe. Atkinson (1894) proposed type. Balansia amhiens Moell., although the genus with descriptions of tv^'o spe- with distinctly capitate ascostromata, cies: "Dothichloe hypoxylon" and "D. possesses ephelidial fructifications typical aristidae." Dothichloe hypoxylon, as ad- of Dothichloe. B. discoidea P. Henn., by mitted by Atkinson in his 1905 treat- reason of its ephelidial fructifications ment, was misapplied to the species he easily referred to Dothichloe and normal- had later identified with D. atramentosa ly characterized by sessile ascostrom^ata, (Berk, & Curt.) Atk., the specific epithet appears rarely to develop even stipitate of which is also inapplicable on nomen- ascostromata. Primarily because of their clatorial grounds because it applied ephelidial structures these two are re- originally to still another species. This ferred to the subgenus Dothichloe, name (D. atramentosa) is clearly a

THE SPECIES OF BALANSIA

Key to American Species of Balansia a. Hypothallus well developed, thickened, enclosing tissues of an undeveloped infloiescence; ephelidial fructifications infundibuliform to patellate-effuse, ephemeral to persistent, ascostromata capitate, sessile to stipitate Eubalansia. b. Hypothallus becoming black, ephelidial fructification infundibuliform to patellate, ascostromata substipitate to long-stipitate. c. Ephelidial fructifications and ascostromata relatively tough and persistent. 1. B. claviceps. cc. Ephelidial fructifications and ascostromata delicate and ephemeral, developing only after the hypothallus has undergone a short rest period 2.B. obtecta. bb. Hypothallus remaining white, ascostromata sessile, black 3. 5. cyperi.

bbb. Hypothallus yellow, ephelidial fructifications and ascostromata yellow. . A. B. pallida. aa. Hypothallus as a thin layer on surface of host tissue; ephelidial fructifications irregularly hysteriform, ephemeral; ascostromata effuse, sometimes developing as separate pulvinuli or capitulae over the hypothallus, chiefly subsessile, rarely stipitate Dothichloe. b. Hypothallus encircling culm or culm and leaf bases of host.

c. Ascostromata continuous over the hypothallus, forming collars about the host. d. Ascostromata roughened-papillate by prominently extruding perithecia and ostioles. 11. B. aristidae. dd. Ascostromata smooth to granular or rugose, never prominently papillate-roughened. (cf. also B. cyperacearum) 5. B. strangulans. cc. Ascostromata as subsessile (rarely stipitate) heads or pulvinuli over the surface of the hypothallus. d. Ascostromata few, arranged as a single circlet about the culm below the node. 7. B. subnodosa. dd. Ascostromata numerous, covering large areas oi the hypothallus on culm or leaf

sheaths 6. B. strangulans f . discoidea. bb. Hypothallus and fructifications on specific areas of the leaf surfaces or in florets or surrounding inflorescences. c. Hypothallus and fructifications on leaf surfaces rarely encircling both leaf and culm. d. On adaxial leaf surfaces 10. 5. epichloe. dd. On abaxial leaf surfaces. e. On Cyperaceae 9. 5. cyperacearum. ee. On Gramineae;

/. On broad areas of leaf chiefly at base, ascostromata sometimes rugose. 8. B. henningsiana. 14. linearis. If. Restricted to single or double lines on leaf surfaces B. ..

BALANSIA AND THE BALANSIAE IN AMERICA 17

cc. Hypothallus and fructifications within florets or surrounding inflorescences. d. Hypothallus and fructifications developing within and ascostromata extruding from florets 12. B. hemicrypta. dd. Hypothallus surrounding undeveloped inflorescences within enclosing leaf, fruai- fications extruded in a single line from leaf margins 13. B. ambiem.

1 . Balansia claviceps form, crowded but not compressed, 200;Lt to 260^ by 120/z to l40/ji; perithecial Balansia claviceps Speg. Soc. Cient. walls 10^ to 15/x thick; asci subcylindri- Argentina An. 19: 45-46. 1885. cal, short-stipitate 106/^ to 195/x by 5ja "^'Ephelis mexicana Fr. ap. Berk., to 6IX with 8 spores; ascospores 100|U to p.p. Linn. Soc. London, Bot. 180;xby I IX to 1.5^. Jour. 10: 353. 1869. SPECIMENS CITED Balansia trinitensis Cke. & Mass., Ann. Bot. [London] 3: 35, pi. 4. On Panicum ? sp.—Pirayu, Paraguay, 1879, 1889. Balansa, PI. du Parag. No. 2755 ex type of Balansia claviceps Speg., ex Herb. ^Ephelis trinitensis Cke. Mass., & Spegazzini (A, W).9—Although the host Ann. Bot. [London] 3: 35, pi. 4. name in this specimen was labeled "Pani- 1889. cum" by Spegazzini, the published description reads "Setaria vel Pennisteum," Claviceps philippii Rehm, Hed- and his later record (1887) is Setaria. wigia 28: 302. 1889. On Setaria palmijolia Willd. — Trinidad, Balansia claviceps var. fasciculata B. W. L, 1888, J. H. Hart No. 1332, in in litt. Speg. herb. & type of B. trinitensis Cke. & Mass. (K);

J. H. Hart, type of Ephelis trinitensis variable in Hypothallus dimensions or Cke. & Mass. (K) ; 1889, J. H Hart ( A) robustness, rarely up to 400 mm. long, On Gramineae (indet.)—Cuba, C. Wright No. usually moderately robust, enclosing 729, paratype of Ephelis mexicana Fr. in culm and undeveloped inflorescence of Herb. Berkeley (K); in Herb. Curtis the host; context white to gray, waxy when moist compactly ** Explanation of abbreviations and of type termi- and prosenchy- nology used in citation of specimens: matous; exterior at first white, becoming (A) = Herbarium of G. F. Atkinson, Depart- ment of Plant Pathology, Cornell University, purple to black usually laccate a and as Ithaca, N. Y. rind. Ephelidial fructifications solitary (F) = Farlow Herbarium of Harvard University, or gregarious, erumpent from beneath Cambridge, Mass. (Fl) = Herbarium of the Florida Agricultural the rind of the hypothallus, at first Experiment Station, University of Florida, Gaines- ville, Fla. spherical, and exciple covered with short (I) =r Herbarium of the University of Illinois, whitish to ephemeral hairs, becoming Urbana, 111. subpatellate when moist, cupulate when (K) = Herbarium of the Royal Botanic Garden, Kew, England. dry, sessile; context fleshy, pseudoparen- (Mo) = Herbarium of the Missouri Botanical chymatous, pallid; hymenium pallid- Garden, St. Louis, Mo. (N) = Herbarium of the New York Botanical carneus to gray; conidiophores when Garden, New York, N. Y. simple I IX to 2^ in diameter, becoming (Pk) = Herbarium of the New York State Museum (containing the Herbarium of C. H. much elongate and swollen, irregularly Peck), Albany. N. Y. to subverticillately branched; conidia 24^ (W) = Mycological Collections of the U. S. Department of Agriculture and of the U. S. Na- to 40^ by 1/x to 1.5 fx. Ascostromata tional Museum, formerly housed at Washington, erumpent from the hypothallus, short to D. C, now at Plant Industry Station, Beltsville, Md. long stipitate, rarely subsessile (the stipe Type = The specimen actually examined and usually 5 to 20 by 2 to 4 mm.), sub- studied by the author of the species and utilized as the basis of the specific description (holotype). spherical, rarely flattened-discoid, surface Ex type = Fragment from the type (clastotype) first light brown then black, evenly Paratype = A specimen not the type in sensu stricto but cited by the author in an original de- granular-punctate to coilicular from the scription. ostioles; context pallid, subperithecial Type coll. = A specimen of the same original gathering (or collector's number) as the type (iso- stroma well developed; perithecia lageni- type). ;

18 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

(F); and in Herb. Farlow (F)10 [prob- TYPE LOCALITY ably Setaria sp.— D.]. W. W. Pirayu, Paraguay, on spikes (inflorescences) On Panicujn ghiesbreghtii Fourn.—Cuba, of Gramineae (? Panicum sp.). 1924, R. Weir and A. Faris (W); J. J. DISTRIBUTION Costa Rica, 1948, E. P. Imle (F, W). culms On P. maxbnum Jacq.—Venezuela, 1949, J. On and inflorescences of various StandenNo. 16 (W). grasses in the Paniceae from the West Indies to Argentina and Chile (fig. On Setaria geniculata (Lam.) Beauv.—Cuba. 6).

1920, C. V. Piper (W); 1924, J. A. Stevenson (W); 1927, W. A. Orton (W); 1933, A. G. Kevorkian (W); no date, S. C. Bruner and Arias No. 1452 (Mo, W); Venezuela, 1940. A. S. MuUer No. 3924 (W); Colombia, 1929, C. E.

Chardon and J. A. B. Nolla Nos. 260, 348 (W).

On S. lutescens (Weigel) F. T. Hubb.—Cuba,

1924, J. R. Weir and J. A. Faris (W) 1924, D. H. Linder (F).

On S. macrostachya H. B. K.—Rio Acre, Peru, 1916, E. UleNo. 3519 (F).

On S. palmifolia (Willd.) Stapf—Trinidad,

1889, J. H. Hart (N, F); 1911, J. B. Rohrer (F).

On S. poiretiana (Schult.) Kunth—Guatemala, 1907, W. A. Kellerman No. 6079 (W); 1908, No. 7543 (F, W); Trinidad, 1922, F. L. Stevens No. 969 (N, F); 1944,

J. M. Todd No. 184 (W); Venezuela, F. L. Stevens No. 209 (W); 1928, H. SydowNo. 209 (F). On Setaria sp.-Cuba, C. F. Baker No. 292 (in herb. R.P. White); 1924, F. S. Earle (W); 1907, 1908, H. Hasselbring (W); Figure 6.— Distribution of Balansia claviceps. Each Honduras, P. C. Standley, 1928, Nos. point indicates the approximate area for one 23475, 52853 (W). or more findings on each host-species. On Sorghum vulgare Pers. (cult.) —Cuba, 1921, S. C. Bruner (W). TAXONOMY AND NOMENCLATURE On Gramineae (indet.).—Costa Rica, 1921, here treated this species is pos- P. V. Siggers No. 5 (W); Venezuela, As 1936, E. P. Arberlaez No. 146 (W). sibly a composite of different races, forms, and varieties, some of which, it

ILLUSTRATIONS is to be expected, may be recognized

Plates 1 and 2; Cke. & Mass., Ann. Bot. 3: ultimately as distinct when more ade- plate 4; Atk., Jour. Mycol 11: plate 87, quate field observations, as well as evi- figure 21; Diehl, Jour. Agr. Res. 41 (11): dences from experimental culture and plate 1, figure 2; ISIarchionatto, Rev. Argent. tests, will have ac- Agron. 14 (2): figure 1; Baker, Trop. Agr. cross-inoculation 11: figures 1, 2; Rehm, Hedwigia 28: figure cumulated. If, however, segregates from 14, a, b, c, d. this composite species are finally distinguished, the identification of the type ^^ It mighc be contended, since Berkeley's description of Ephelis mexicana Fr. was published in an specimen of Balansia daiiceps has spe- account concerning Cuban fungi, that his cited Cuban cial significance. specimen, Charles Wright's No. 729, is the type. \z should nevertheless be obvious that the Mexican Spegazzini (1885) specifically cited specimen cited above is the r^-pe, since Berkeley cited it as "Fr. Fung. Mex." therefore accepting with as the type Balansa's Plantes du Paraguay Fries' manuscript name the type specimen also supplied by him. By modern praaice and rules Berkeley's No. 2755 from Paraguay. It probably (1869) publication rypified and validated Fries' contained material from more than one (1849) genus. Ephelis was described without naming any species, merely citing "Fr. Fung. Mex." in host, as is evidenced by the fact that a reference to the manuscript still unpublished a cen- (pi. tury later. fragment I, A) presented to the BALANSIA AND THE BALANSIAE IN AMERICA 19

writer by the late Prof. C. Spegazzini as recognition as the same species is the part of the type resembles somewhat lack of record of an Ephelis stage in closely the fungus from Cuba on Setaria, Spegazzini's type. That specimen (pi. 1, whereas a part of the supposedly same B) possesses scars on the hypothallus original collection, in fact the very speci- readily interpretable as the kind of scars men illustrated by Atkinson (pi. 87, fig. following abscission of ephelidial fruc- 21, 1905) appears to be different. The tifications in Cuban specimens of it on specimen sent to the writer is more ac- Setaria and in the type of B. trinitensis.

curately representative of Spegazzini's In this connection it is noteworthy that

description, and it bears a label in his Bresadola, Hennings, and Magnus handwriting including the term "typus." (1893), in reporting Ephelis mexicana That he recognized the likelihood of from Puerto Rico on an unnamed grass,

more than one host species in Balansa's considered it to be a stage of B. claviceps No. 2755 is indicated by his having pub- Speg. and also reduced Rehm's Claviceps lished (1885) as the host "Setariae vel philippii to synonymy with B. claviceps. Penniseti" and later (1887) "Setaria." A few years later Rehm (1898) likewise His type specimen, from which this frag- acknowledged his B. philippii as a ment is presumed to have come, is neces- synonym of B. claviceps. Other observers sarily restricted to but one host species. have tended to accept the relation of the Since the fragment bears his notation ephelidial fructification in this species "Panicum," that identification, presum- recorded either as B. claviceps or B. trini- ably representing his matured opinion, tensis.

is accepted here. This species is probably not repre- A twinned specimen (pi. 1, C), also sented among the specimens in Sac- made available years ago through Pro- cardo's Herbarium, although Gola fessor Spegazzini's generosity, bore his (1930) has recorded a single specimen, holographic label "B. clavkeps var. fas- not noted by number, from there by this ciculata," a variety name that appears name as in Ule's Mycotheca Brasiliensis. never to have published. for been Except In that set of 100 specimens there is the twin effect, most likely a character only one specimen referable to Balansia, originally of the host plant, it does not No. 78; but No. 78 happens to be a appear to differ from the type. duplicate of the type of B. asclerotiaca Cldv/ceps pb/lipp/f Rehm (1889) was P. Henn. based upon a Philippi specimen from Merguri, Chile, host not named. Rehm's QUESTIONABLE RECORDS AND ERRORS OF illustration is a perfect reproduction of IDENTIFICATION the specimen in Atkinson's herbarium but lacking data that might prove it to Spegazzini's (1891) second account be a part of the type. of Balansia claviceps with reference to Cooke and Massee (1889) described Balansa's No. 4320 in culms of Chloris what is here regarded as a synonym, sp. from Guarapi in Paraguay was recog- Balansia trlmtensts, from Trinidad as a nized as different from the type in that parasite on the ubiquitous Setaria palmi- it occurred on internodes of the culm jol'ui. Although this latter host is of rather than on the inflorescences. But he

Asiatic origin, the writer has found no did not include it with his Epichloe nigri- record of the fungus on this host except cans because of the capitate, even oc- in America. B. trinitensh (pi. 2, B) was casionally stipitate, ascostromata. He sug- described with the imperfect Ephelis gested the possibility that it belonged to trinitensis as an auxiliary fructification, B. pilulaeformis as follows: "an hue a feature not recorded by Spegazzini. Ophiodotis vorax (B. & Br.) var. pilu- The similarities of the two species are laeformis (B. & Br.) etiam?" His de- so evident that the only objection to their scription suggests Hennings' B. discoidea, 20 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE an identification verified by examination claviceps, insofar as it referred to the of Balansa's No. 4320 in the Farlow species on Cenchrus, applied to what is Herbarium. now regarded as a distinct species. The report by Bommer and Rousseau ADDITIONAL HOST AND GEOGRAPHIC (1896) of B. claviceps from Costa Rica RECORDS as on the surface of bamboo culms is Berkeley's publication of most certainly suspect. The identity of (1869) Ephelis mexicana was based upon two both fungus and host awaits examination specimens of somewhat different appear- of the specimen, a gathering of Tonduz. ance, most likely on different host spe- It is difficult indeed to evaluate Lin- cies; one from Mexico, locality unknown, dau's (1897) "original" illustration {his and the other from Cuba, locality also fig. 247-a on p. 371) reproduced by unknown. The Mexican specimen, pre- Gaumann (1926) and by Gaumann and sumably sent to Fries by F. M. Liebmann Dodge (1928). Although labeled ''B. and doubtless the material on which claviceps;'' it resembles more closely Fries (1849) based his then untypified specimens of certain Asiatic species; it genus Ephelis, was sent to Berkeley by is most certainly not an illustration of Fries. This specimen in the Herbarium Spegazzini's type. of the Royal Botanical Garden at Kew, Atkinson (1905) was clearly in error England, properly recognizable as the in ascribing 1880 in place of 1885 as type, is in poor condition but could well the date of publication of this genus and be the ephelidial stage of the related species. He was also confused in his as- species so common on Cenchrus echinatus sumption that Epbelis mexicana wis a in Florida. The paratype from Cuba at stage of Balansia hypoxylon. He was Kew (pi. 2, A), Charles Wright's No. unaware of or ignored the opinions on 729, of which replicates are in the Curtis this score offered by Bresadola and co- and Farlow Herbaria at Harvard Uni- authors (1893) and of Rehm (1898), versity, is obviously on a different host and he ignored entirely the report of species from the type, and it is different Cooke and Massee on the pertinent B. from any recognizable host otherwise of rrinitensis and E. trinitensis. record for this species. Seaver and Chardon appar- (1926), Rorer (1911) and Thorold (1931), ently accepting Atkinson's in- (1905) in reporting Balansia trinitensis from terpretation of the relation of Ephelis Trinidad, merely cited Hart's original mexicana, have reported the ascostromata findings on which the species was based. of this species as referable to the un- The Sydows (1916) reported as B. related Balansia hypoxylon. trinitensis a specimen on Setaria macro- There are various reports of Balansia stachya "Tr." (?) found by Ule No. claviceps on grasses in the old world; 3519 in Peru, a record quoted by Garcia but the Eurasian specimens so recorded, Rada and Stevenson (1942) ; a duplicate writer which the has been privileged thus specimen is cited in the above list. far to examine, have not proved to be Spegazzini's (1919) report of Balan- of that species. sia claviceps from Brazil on Setaria sp. Barger (1931) has included Claviceps referred to the twinned specimen after- philippii among the ergot fungi, prob- ward labeled "var. fasciculata" in his ably because of the name only, since he herbarium. lacked evidence or record of its having Sydow (1930) noted B. trinitensis any alkaloid content; he was doubtless from Venezuela on Chaetochloa sulcata not aware that this name had previously (i.e., Setaria poiretiana) in reference to been recognized as a synonym of B. LJle's specimen No. 3519 cited above. claviceps. Baker (1934), in reporting his ob- Diehl's (1930b) account of Balansia servations on Balansia trinitensis in . ,

BALANSIA AND THE BALANSIAE IN AMERICA 21

Trinidad on both Setaria pahnifolia and ervation, it has always been obvious that S.. poiretiana (as Chaetochloa sulcata)^ this foliage was of the same green color suggested the hkehhood of systemic in- as any healthy plant. Although the infection by a perennial mycelium. florescences were aborted and mummi- Although Spegazzini seems not to fied, the elongation of the rachis in most

have recorded his species from Argen- cases is only slightly inhibited. Also, the tina, Marchionatto (1947) has reported black pseudomorph is prominently evi-

it as frequent in Missiones (northern dent at the tip of the stem, particularly Argentina) on an unidentified species in species of Setaria and Sorghum, al- of Setaria. though in some species of Panictim the

As Ephelis mexicana the species has pseudomorph is fairly hidden within the been reported from Brazil as on Pamcum sheath of the terminal leaf. It would sp. by Hennings (1908), from Colombia seem that affected plants of Setaria and as on Chaetochloa geniculata (i.e., Sorghum, bearing the black pseudo- Setaria) by Chardon and Toro (1930), morphs well above the green foliage, and from Cuba on Holcus sorghum would be conspicuous in the field, (i.e., Sorghum vulgare) by Bruner whereas with the partially hidden (1931). Bruner's account refers to his pseudomorphs diseased plants of Pani- specimen cited above (pi. 2, D) cum would be very inconspicuous.

PATHOLOGICAL CONSIDERATIONS 2. Balansia obtecta obtecta spec. nov. Little of published record is available Balansia to indicate the incidence of plants in- "^Ephelis mexicana Fr. ap. Berk, fected by this fungus, although Baker p.p. typ., Linn. Soc. London, Bot. (1934) implies its local abundance in Jour. 10: 353. 1869. Trinidad on two species of Setaria. Nota- Balansia mexicana Atk. (hyponym) tions accompanying specimens and per- Jour. Mycol. 11: 253. 1905. sonal communications in 1925 by the Hypothallo sclerotioideo, corniformi, late H. Hasselbring, as well as recent cylindrico-clavato, inflorescentiam hos- statements in conversation by A. G. pitis amplexanti et toto in vagina folii Kevorkian, affirm that in Cuba affected cryptico, externe nigro-laccato, intus plants of Setaria, presumably S. lutescens albo; fructus et sporae Balansiam clavi- and 5". geniculata, are prevalent and very cipitem simulant sed peridia ascomatis numerous in some places. Bruner's et fructificationis ephelidialis textura (1931) account of infected Sorghum in delicatissima et coloribus fuscioribus dis- Cuba appears to be restricted to a single t!n.f?uintur.o instance. Hypothallus corniform, narrowly cyl- All specimens available from different indrical clavate sclerotioid, variable in sources reveal a complete mummification size but rarely exceeding 120 by 3 mm.; of the inflorescences that are enveloped surface at first white but becoming black by the encompassing hypothallus; and and laccate, encompassing aborted in- notations on specimens as well as pub- florescence and rachis of the host; con- lished accounts bear out the thesis of text waxy when moist, corneous when complete, never merely partial, mummi- dry; entirely enclosed within the sheath fication of the immature inflorescence. of the terminal leaf and with the enclos- Specimens of this fungus in herbaria ing leaf readily becoming detached when can afford only partial evidence of the mature by abscission from the culm. pathological effect on the host, but that Ephelidial fructifications solitary or evidence is fairly specific. Where the crowded, erumpent through the rind of specimen was accompanied by foliage of the hypothallus, sessile, at first subspheri-

the affected plant and it is clear that the cal, becoming cupulate to patellate with material was properly prepared for pres- a hairy exciple; excipular hairs white to 22 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE yellowish brown, persistent, simple to DISTRIBUTION septate, 40^. to 100^ by 2.5^ to 5/a (rare- On inflorescences of species of Cenchrus, Setaria, ly to 10/x in diam.) ; context prosenchyma- Cynodon, and known only from Florida, Cuba, tous to pseudoparenchymatous, delicate, Dominican Republic, and Mex- ico (fig. 7). carneus pallid to grayish brown; conidia borne in pinkish masses that become cirri; conidiophores simple to complex, 1^ to

2fJi in diameter, becoming swollen and irregularly branched; conidia 24^ to 40/ji by Ifx to 1.5/A. Ascostromata subspherical, 2 to 2.5 mm. in diameter, erumpent from the hypothallus, frequently through an ephelidial fructification, capitate and stipitate (stipes 5.0 by 0.3 to 0.8 mm.), yellowish pallid becoming dark brown, granular punctate to coUicuIar from the figure 7.— Distribution of Balans'ia obtecfo. ostioles; context pallid; subperithecial stroma well developed; perithecia elliptic- TAXONOMY AND PREVIOUS RECORDS lageniform, crowded but not compressed, The Species has much in common with Balansia clavkeps. The hypothallus is 2u0;a to 260fx by 120/^ to l40/x with walls 10/x to 15/A thick; asci subcylindrical, distinctly more laccate than that species. In all the specimens the hypothallus sur- short stipitate 101/x to 195/a by 3/a to not only the inflorescence but all 6jx, with eight filiform septate spores rounds of the rachis; and the hypothallus with 100/x to 180/x by Ifx to 1.5/x. contained host tissues is easily disarticu- SPECIMENS CITED lated along with the terminal leaf from On Cenchrus ech'inatus L.—Florida, 1892, W the node where it is attached. The fruc- T. Swingle No. 2995 (W); 1894, P. H tifications here are thus most likely to Rolfs (W); 1922 (W), 1923 (N, W) 1930, 1932, G. F. Weber (N); Gaines be developed from the disarticulated ville, 1939, G. F. Weber Type (W) hypothallus rather than from that struc- A. C. Foster A. 1925, (W); 1927, J ture while still attached to the apex of Bruman (W); 1928, H. R. Fulton (W) a grass shoot, as in B. clavkeps. Further- 1930, R. T. Gibbens (W); 1929, 1930 G. B. Sartoris (F, N, W); 1931, W. W more, in the specimens of B. obtecta thus Diehl (W); 1935, E. West (N); 1943 far seen the hypothallus is consistently V. K. Charles (W); Cuba, 1947, A. G matured while still within the sheath of Kevorkian (W). the terminal leaf of the host and is thus On C. gracillimus Nash-Florida, 1892, W. T. Swingle No. 3063 (W). practically hidden from view. On Cenchrus sp .-Florida, 1892, W. T. The few specimens of ascostromata Swingle No. 3104 (W); 1901-03, P. H. thus far available are lighter colored and Rolfs (W). stipes are longer and more slender On Cynodon dactylon (L.) Pers.— 1906, P. H. the Rolfs (Fl); 1924, O. F. Burger (Fl, W); than in B. clavkeps. In all the specimens, 1924, G. F. Weber (N). the peridia (exciples) of the ephelidial On Setaria setosa (Swartz) Beauv.—Domini- stage are more delicate also, while the can Republic, R. Ciferri No. 4434 (W). On Gramineae (indet.).—Mexico (probably excipular hairs are more prominent. The coll. F. M. Liebmann), type of Ephelis delayed production and sequence of

mexicana Fr. in Herb. Berkeley (K). fructifications is also a feature that ap-

ILLUSTRATIONS pears to distinguish it from B. clavkeps. Balansia obtecta have Plates 3 and 4, A to D; Diehl, Jour. Agr. Specimens of Res. 41 (11): plate 1, F to K; text figures been distributed in herbaria chiefly under 1 and 2 (sub B. claviceps). the names B. clavkeps, B. hypoxylon, TYPE LOCALITY and Ephelis mexicana, and in one of the

Gainesville, Fla. great herbaria of the United States some BALANSIA AND THE BALANSIAE IN AMERICA 23

specimens have even been referred to only. These areas were easily detected ErJospora. It has been reported by Weber because they were heavily grazed by (1924) as B. hypoxylon and by Diehl cattle, whereas adjacent areas bearing the (1930) as B. clavkeps. The earliest unpalatable burs were not grazed. Never- record of it is the original report by theless, at other seasons new shoots of Berkeley (1869) of the type of Epbelis normal plants developing before bur mexicana Fr. based on a Mexican speci- formation are said to be readily grazed men sent to him by Fries, which is by the same cattle that avoid them after doubtless the basis of Fries' (1849) pre- burs have formed. Quantitative and more viously untypiiied genus Epbelis. Atkin- specific observations are needed to ap- son (1905) suggested the likelihood that praise adequately the incidence of in- this name E. mexicana referred to the fection as well as the economic implica- ephelidial stage of the Balansia from tions. Texas, which he then referred to B. CULTURE OF THE FUNGUS hypoxylon, thus considering E. horealis attempts to cultivate Balan- Ell. & Ev. to be a synonym of E. mexi- Numerous culture cana, an assumption not justified by the sia ohtecta in pure on standard thus far been consistent facts. It is unfortunate that the type agar media have failures. Similarly, attempts to grow it specimen of E. mexicana is so fragmen- tary and that no additional specimens are in decoctions of sandbur plants and on agar incorporating a sandbur plant available from Mexico. It is to be ex- media pected, however, that search will reveal decoction were without success. The its presence in Mexico where Cenchrus media were inoculated by placing within

echinatus is to be found. and upon the surface viable ephelidial conidia or fragments from within the FIELD OBSERVATIONS hypothallus or the entire hypothallus, The writer's observations of Balansia using standard aseptic precautions. Al- ohtecta in the field have been restricted though conidia germinated here as readi- to two short periods in central Florida ly as in water, there was no mycelial (one in September 1931 and the other development after the reserve nutrients in January 1933) concerned with but in the conidia were spent. From the one host plant, the locally abundant hypothallus or fragments of it no hyphae Cenchrus echinatus. Of the thousands of grew onto or into the media, but after infected clumps of this grass there surface-sterilized pseudomorphs had been readily available, at least 99 percent were plunged partially into the media in tubes entirely sterilized by the fungus. Only a or placed upon agar plates ephelidial few were found to bear both normal fructifications developed on the surface (bur-bearing) and infected (fungus- of the hypothallus after 48 hours. When bearing) shoots. These few clumps, the surface of a sound pseudomorph had showing partial infection only, could been first sterilized by immersion in have been accounted for as having orig- mercuric chloride solution followed by inated from more than one seed, one rinsing in sterile water or flamed after healthy and one diseased. In one speci- momentary immersion in alcohol and men, however, found at Gainesville, then manipulated aseptically, the likeli- Fla., September 23, 1931, both healthy hood of contamination was reduced to and diseased branches arose unquestion- a minimum so that conidia were thus ably from one culm. Some areas ap- produced in pure culture. peared to be practically a pure stand of Obviously the fungus did not lend infected plants. Between Fort Myers and itself to the culture methods in common Tampa, Fla., in January 1933, certain use with saprophytic or hemiparasitic roadside areas of from 10 to 50 feet in fungi. There was, however, no material diameter were covered by infected plants need to produce conidia aseptically, and 24 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE the difficulties easily outweighed any morphs in quantity but reached more possible advantage. Some difficulty was than twice the normal height and showed always encountered regardless of the exceptional vigor compared with its be- techniques employed because of con- havior as noted under natural conditions taminating nematodes within the pseudo- in Florida, or under culture at Arlington, morphs. Under moist-chamber condi- Va., or even in the greenhouse at Cam- tions, aseptic or not, these nematodes bridge. usually infested most of the material, One striking development of this out- often largely consuming it after 15 to 30 door culture at Cambridge was the ap- days. In instances, however, where pearance of a normal shoot bearing burs. pseudomorphs over wet filter paper were In other words a healthy shoot arose in allowed to dry intermittently every few culture from an infected plant, duplicat- days, the nematodes were less numerous ing a condition observed in one instance and the hypothallus continued to produce in 1931 under natural conditions in ephelidial fructifications for as long as Florida. 3 months. In one instance at Cambridge, SNOCULATION EXPERIMENTS Mass., in February, 1931, after 2 months intermittent drying and wetting of the Attempts to secure infection of green- hypothallus in moist chamber there was house-grown, healthy plants of Cenchrus produced a single stipitate ascostroma echinatus by placing germinating epheli- (pi. 4, B) identical with ascostromata dial conidia of Balansia ohtecta on leaves found in 1923 by G. F. Weber, which (more than 200) and culms were made he reported (1924) as Balansia hypoxy- at various times at Cambridge in 1930 lon. In 1939 Dr. Weber very kindly and 1931 but without success. During presented the writer with abundant ma- the same period inoculations were at- terial of stipitate ascostromata produced tempted by injecting with a hypodermic by him in April of that year from pseudo- syringe germinating ephelidial conidia morphs in moist-chamber, which are here into apices of shoots of the host (more utilized in typifying the species (pi. 4, than 45 injections) with no evidences of D). It would appear that the ascostro- infection. Also an attempt to obtain in- mata develop normally after the pseudo- fection was made in December 1930 by morphs have fallen to the ground or on germinating more than 100 healthy seeds decumbent culms. of Cenchrus echinatus in tap water containing ephelidial conidia in process of As attempts to obtain growing myce- germination; then when the seedlings lium in pure culture were not successful, were approximately 1 cm. in height they the fungus was grown, perforce, on the were transplanted to flowerpots until living host. Since mycelium could be flowering in April 1932, by which time demonstrated in infected plants in culms there was no evidence of infection. and leaves the infection was naturally Attempts to obtain infection, further- assumed to be systemic. Infected plants more, were made at the same time by from Florida were grown under green- atomizing 60 inflorescences having re- house culture and also outdoors at ceptive stigmas with germinating conidia Arlington, Va., from 1928 to 1935 and and keeping them in a moist chamber at Cambridge, Mass., in 1931 and 1932. for 24 hours. Seeds that formed upon Vegetative propagation of the infected these inflorescences when planted re- host was a simple matter in that it roots sulted in a total of 112 plants, all normal. readily at the nodes and frequently At the same time conidia were atomized branches at the nodes, permitting a rapid on flowers of species of other grass increase of living material. The infected genera without result — Anthoxanthum, host plant outdoors during the summer Avena, Danthonia, Echinochloa, Pani- of 1931 at Cambridge produced pseudo- cum, and Setaria. BALANSIA AND THE BALANSIAE IN AMERICA 25

Another attempt at floret inoculation fected plants discloses slender hyphae

was made on March 9, 1931, by touching ramifying between cells in the vascular the stigmas of flowers on 10 inflores- region and especially in the apical meri- cences with a camel's hair brush that had stem where they are most abundant. been previously brushed against epheli- These intramatrical hyphae are prone to dial fructifications, then placing the take up readily and retain the dye from plants in a moist chamber for 24 hours. an acidulated glycerine solution of From the seeds that developed on these phloxine; and they are still more readily inflorescences there were germinated 20 visible when thin sections or mascera- plants, of which at the time of inflores- tions of affected tissue stained as above cence, January 3, 1932, 17 were normal, are mounted in clearcol (Zuck, 1947). healthy plants, but 3 bore the fungus in Hyphae in stems and leaves rarely exceed place of inflorescences. Similar treatment 1.5|M in diameter, but in the pseudo- without any positive result was also given morphs they commonly reach 5//,. to Avena sativa, Cenchrus gracillhnus, The endophytic mycelium is thus the C. incertus, C. paucifiorus, Danthonia active vegetative stage, and the only long- spicata, DigitarU sangtiinalls, and Echi- time dormancy permissible by the facts nochlod crusgalli. of its life history would be while quies- The results of these attempts to secure cent in the seed. infection, though meager, indicate that PHENOLOGY AND L!FE HISTORY infection of the ovule occurs by way of On Cenchrus echinatus, and most like- the stigma and that it is transmitted ly on other host plants, the pseudomorphs through the seed. This contention was of Balansia obtecta develop before and supported by the evidence afforded by during the time of flowering. The epheli- microscopic examination of three of the dial stage develops quickly under pro- normal-appearing seeds of Cenchrus pitious conditions of moisture, and it is echmatus that developed after this last probable that ephelidial conidia are inoculation attempt; it disclosed the chiefly responsible for infection. Since presence of abundant mycelium char- Cenchrus has a fairly long period of acteristic of Balansia in the embryo and flowering, even with the later develop- especially noticeable between the embryo ment of ascostromata, ascospores might and endosperm. The systemic nature of be expected also to function but in the endophyte in the living host is thus lessened volume of infection. accounted for. Whether natural dissemi- The life history of Balansia obtecta nation is the result of splashing by rain may be expressed by the following dia- or of insect transmission of conidia or gram (fig. 8). ascospores is still problematical, although the glutinous walls of these spores would TERATOLOGY seem ideally fitted for adhesion to insects. Unlike many plant pathogens Balansia obtecta, in common with most other ENDOPHYTISM Balansiae, does not produce any notice-

That the fungus {Balansia obtecta) is able chlorotic effect, as affected plants an endophyte is realized not only from are equally green, or possibly even a the fact that it is borne within the m.ore vivid green than normal plants. In seed coat and appears superficially only wintertime greenhouse culture in Vir- at the time of flowering of the host; it ginia and in Massachusetts, as well as is readily proved by examining stem under natural conditions in Florida, in- and even leaf tissues of infected plants. fected plants of Cenchrus appear some- It is apparent at the base and middle v/hat shorter than normal plants because of leaves but rarely at the tip and has of the lack of an inflorescence. But the not been demonstrated from roots. Care- affected plants are actually dwarfed ful examination of stem tissue of af- slightly, due to shorter internodes. Af- 26 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

HEALTHY SHOOTS FROM INFECTED PLANTS The production of normal shoots from infected plants of Cenchrus echinatus poses questions both as to a cause and to

a mechanism by which it can have resulted. There has appeared thus far no consistent evidence of a causal relation to recover)^ or escape from this type of systemic infection, but an explanation

of the mechanism is possible. As this type of infection in the grass host prog- resses only with the meristem, the fungus

is compelled to grow as fast as the apical

meristem is forced upward. If the grow-

ing tip produces behind it host cells at a more rapid rate than the upward growth of the neighboring hyphae, the apical meristem will have escaped the infecting hyphae, which are left behind and can- Figure 8.— Diagram to illustrate the life-cycle of not invade the new shoot that develops BaJansia as exemplified by B. daviceps and beyond that point. Hence, the plant has B. obtecfa. not recovered from the infection but has merely produced a vegetative shoot that fected plants tend to branch more ex- has escaped it. tensively, giving a bushy, almost a fasciated, appearance. In Florida, owing 3. Balansia cyperi of affected to the more intense grazing Balansia cyperi Edg.," Mycologia 11: plants, they appear decumbent but are 260-261. 1919. outdoors erect in greenhouse culture or '^Ephelis texensis Ell. & Ev., ined. in Virginia and Massachusetts. One plant in Herb. EUis. grown during the summer of 1931 in ^Sphaceliopsis cypericola Speg., the botanical garden at Cambridge, Mass., Buenos Aires Mus. Nac. de Hist. was actually taller and more vigorous Nat. An. 20: 457-458. 1910. than normal plants, a condition in strik- Hypothallus spherical to subspherical, ing contrast with those in the greenhouse usually 5 to 30 mm. in diameter, enclos- native habitat. No explanation and in the ing apex of culm with bases of involucral for this giant effect is offered, unless it leaves and undeveloped inflorescence; to photoperiodism, since similar be due surface white, becoming gray to black cultures at Arlington, Va., more outdoor with age; context white to gray, com- nearly resembled the plants in Florida. pactly prosenchymatous. Ephelidial fruc- The striking effect of infection in tifications gregarious, crowded, few to Cenchn^s is the complete suppression of many, aggregate to contiguous or solitary, any elongation and expansion of the erumpent beneath surface of the hypo- rachis and inflorescence at the time the thallus, sessile, cupulate to patellate and inflorescence has reached the length of convex; exciple thin, without hairs, the terminal leaf sheath or just as it has pseudoparenchymatous, gray, evanescent; exceeded it slightly (pi. 3, A, B, C) ; at hymenium pink to orange-colored; coni- that time it is fully covered by the diophores simple to irregularly branched, strangling hyphal structure that is the 1^ to 1.5;u, in diameter; conidia 15/z to hvpothallus. This pseudomorph is thus 30/i by 1.5^ to 2^. Ascostromata few, wholly or almost wholly enveloped by aggregate to contiguous or solitary; sur- the leaf sheath. face black, evenly granular-punctate from :

BALANSIA AND THE BALANSIAE IN AMERICA 27 the ostioles; context white; subperithecial stroma moderately robust; perithecia crowded but rarely compressed, ovate to lageniform, 360^ to 600 fx by 120^ to 200/x; perithecial walls l6/x to 25/x thick; ostioles elongate but very slightly exsurgent; asci cylindrical, 160^ to 225 fi by 6ju to 9fji', perithecial walls l6/x to 26jx thick; ostioles elongate but very slightly exsurgent; asci cylindrical, 160^ to 225ja by 6/i to 9ju,; ascospores 100^ to 200fi by 1.5/x to 3/x.

SPECIMENS CITED

On Cyperus virens Michx.—Louisiana, 1917, A. T. Bell (W); 1917, A. T. Bell and C. W. Edgerton (W, A); 1917, A. T. Bell and C. W. Edgerton, type (W, N); Texas, 1944, E. C. Tullis (W); vie. Belem, Para, Brazil, 1942, W. A. Archer No. H 446 (W). Figure 9.— Distribution of Bafansia cyperi (indi- On Cyperus sp.—Arkansas, 1884, F. L. Har- cated by black dots), B. pallida (circle), B, vey (F); Texas, 1890, H. S. Jennings, cyperacearum (black squares), and B, bsmi- type of EphelJs texensis in Herb. Ellis crypia (crosses within circles). (N, Mo); 1913, D. A. Saunders Nos. 1315, 1656 (W). little question that Edgerton's specimens ILLUSTRATIONS are-C. virens Michx. and scarcely more Plate 4, F and G; Edgerton, Mycologia 1 1 that Spegazzini's (1910) recorded host plate Spegazzini, Anal. Nac. B. A. 12; Mus. might have been C. vegetns Liebm. The 20: figure 70. latter name is a synonym of C. virens TYPE LOCALITY Michx., and both records refer to the St. Gabriel, La., on inflorescence of Cyperus same host and the same endophyte. In virens Michx, the above-cited specimens, as on Cyperus DISTRIBUTION sp., the host materials are indistinguish- On inflorescences of species of Cyperus, able from Edgerton's specimens. It is Louisiana, Arkansas, Texas, Brazil, and Ar- possible that B. cyperi Edg. may be re- gentina (fig. 9). stricted to this one ubiquitous species of DISCUSSION Cyperus. No Argentine specimen or type of Seymour's (1929) Host Index has Spegazzini's (1910) Sphaceliopsis cype- noted B. cyperi as on Scirpus sp., an ricola has been available for examination; entry (as revealed by the late Mr. Sey- nevertheless Spegazzini's illustration {his mour's card catalog) that is based on an fig. 70) leaves little doubt as to its iden- erroneous record, since corrected, of the tity with the ephelidial stage of Balansia United States Plant Disease Survey; this cyperi Edg. Spegazzini's type was found record was of the above-cited specimen on ''Cyperus vegetus" in the vicinity of of D. A. Saunders, No. 1656 (W), La Sabana, Chaco, Argentina. This host from Texas for which the host is a record of "C. vegetus" like Edgerton's species of Cyperus, probably C. virens, (1919) record of "C. virens" might be that had been labeled "Scirpus" by the a source of considerable confusion were collector. there much cause for doubt as to which The writer's knowledge of the peculiar of several authors these specific names features of this disease of Cyperus and should be credited. Fortunately there is of its incidence is restricted to the meager 28 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

technical description by Spegazzini phiaiides, and it is possible that study (1910) and the rather short account by of this fungus in living condition might Edgerton (1919). Spegazzini's record, demonstrate a primary typhodial layer. with a good illustration of his material If a typhodial layer is present in young from the vicinity of La Sabana in the specimens, the species would then be Argentine Chaco, calls special attention referable to Atkinsonella. to the pathological effect of thickened

coalescent bracts that surround and are 4. Balansia pallida incorporated as a part of the pseudo- Balmisia pallida Wint. in Rab., Fungi morph. This is a feature not especially given consideration by Edgerton, who Europaei [Exsiccati] Cent. 36: noted: "A bud from a diseased stool No. 3549. 1886, and Wint., has the appearance of being perfectly Hedwigia 26: 32, 1887. healthy, yet a cross section shows the Claviceps pallida (Wint.) P. mycelium to be abundant betv/een the Henn., Hedwigia (Beibl.) 38: different parts. The host cells, however, (64). 1889; nee C. pallida Pat, not appear to be injured in these do Soc. Mycol. de France, Bui. 15: very small buds." The pathological effects 206-207. 1899. of this species apparently parallel that of Hypothallus subspherical, chiefly Bdcinsia claviceps in that the fungus 0.5 to 2 in diameter, within florets of halts completely at a given point, the mm. the host, enclosing the ovary and par- expansion of the inflorescence, which is tially enclosing palea and lemma, sur- mummified by the encompassing sclero- face yellow, tizing hyphae of the hypothallus. Edger- whitish within, largely covered by a darker yellow layer composed of ton (1919) noted that "a large propor- convergent ephelidial fructifications and tion of the sedge plants" at St. Gabriel, bearing 1 stipitate ascostro- La., "were attacked by the fungus" and from to 6 ephelidial fructifications especially called attention to the fact mata; connivant excipular that a stool of the sedge that was healthy with ephemeral dissepiments, bore normal flowers, whereas on an conidia 44^ to 62yz by Iju to 1.5/a (Win- affected plant the flowering shoots bore ter notes 2^) ; acostromata stipitate, sub- only the pseudomorphs. This indicates globose, 0.3 to 1.5 mm. in diameter, surface yellow punctulate the clearly that the mycelium of this species, by brownish papillate ostioles; context white to yel- like that of B. claviceps, is systemic lowish; stipes curved-flexuose, angular- in the tissues of the host and suggests flattened to grooved, light yellow, to the likelihood that infection may be seed- 0.3 borne. As conidia and ascospores both 0.5 by 1 to 3 mm; perithecia ovate-lageniform, moderately crowded 290/x to mature at the time of flowering, it by 130/x to l60/x, asci cylindrical seems likely also that infection may well 320^ to to with distinct be through the stigma. 175/x 220;a by 3/a 5/x, slime cap not exceeding 5/x in diameter; Even in dried specimens the white ascospores 160;* to 200/x by X^k (described surface of the hypothallus resembles by Winter as breaking into fragments somewhat the superficial appearance of 44^ to 62ja by I^k) . the typhodial stage of Epichloe and At- SPECIMENS CITED kinsonella, although examinations of the dried specimens cited above have failed On Luziola peruviana GmeL, near Sao Fran- to reveal a typical typhodial layer. In one cisco, Santa Catharina, Brazil, May 1885, E. Ule, in Rab.—Fungi Europaei No. instance, on an unnumbered specimen 3549, ex type (F). gathered in Texas, May 1913, by D. A. Saunders, there are present extruding ILLUSTRATIONS hyphae that are suggestive of collapsed Plate 4, £. BALANSIA AND THE BALANSIAE IN AMERICA

DISTRIBUTION If Winter's species were not actually a Balansia some confusion in nomencla- Known only from the type locality, jn florets of Luziola peruviana (fig. 9)- ture might develop from the fact that Hennings (1899a) had referred it to MORPHOLOGY AND TAXONOMY Claviceps in the same year that Patouil- lard (1899) described a C pallida from Superficially Balansja pallida re- Guadeloupe, later given a new name, sembles B. dtadema Moell., but is dis- C. patouillardiana, by Hennings tinguished from it by the ephelidial (1899b). Patouillard's fungus is some- fructifications lacking in that species, what similar in color and general ap- which is characterized by its sphacelial pearance to Winter's species, but differs stage. In respect to its limitation to single in the decidedly more robust ascostro- florets it suggests B. hemicrypta (cf ) . B. mata and stipes and in having developed pallida is remarkable among American from larger blackened sclerotized caryop- species of the genus because of the yel- ses on the ground. This is well borne low color of the hypothallus and fructi- out by the type of Patouillard's species fications, a feature that is lacking in most at the Farlow Herbarium. Incidentally, species of the genus, but which alines Patouillard's species was correctly con- it with the time-honored concept of the sidered by Petch (1933) along with Hypocreales. It is unfortunate that this Moller's (1901) C. balanswides from distinctive species is known from only Brazil to be synonymous with the Cuban one gathering, thus precluding adequate C. fiavella (Berk. & Curt.) Petch. It is, record of it until additional material is however, curious that Duss (1903), in at hand. listing the fungi of Guadeloupe and NOMENCLATURE AND DISTRIBUTION Martinique, based on his specimens identified by Patouillard, did not note this The type gathering by Ule in 1885 synonymy of C. pallida with C. fiavella, was issued by Winter (see Rabenhorst, since he included both names. Never- 1886) in his exsiccati as No. 3549, ac- theless, the type of C. pallida Pat. (Duss companied by the original description. No. 1128) bears the notation in Patouil- This was in good condition and in suffi- "= lard's own handwriting Claviceps fia- cient quantity to be so distributed, vella Bk.," and it is on the same otherwise the species would be more herbarium sheet with the three similar readily confused with others. It appears specimens (Nos. 178, 963, and 1704) that since 1886 no other specimen of recorded by Duss (1903) as Cordyceps this species has been brought to notice, fiavella. Patouillard had thus anticipated the reports of Winter Pazschke (1887), the conclusion published by Petch that Saccardo and of Rehm (1892), (1895), C. pallida Pat. was a later synonym of referring to this original ma- (1898) C. fiavella. terial. Hennings' (1899a) transfer of the species to Claviceps was not accepted 5. Balansia strongulons by Rehm (1900), but was so recognized by Saccardo (1902) and by Bar- Balansia strangulans (Mont.) comb, ger (1931), each without offering evi- nov. dence in support of either contention; Cordyceps strangulans Mont., Ann. there is, however, no indication of des Sci. Nat. (ser. 4) 3: 98-99. additional gatherings. The species should 1855. still exist at least in the type locality and Epichloe strangulans (Mont.) Sacc, may well occur over much of the range Michelia 1: 323. 1878. of the host which, as noted by Hitchcock

(1936) , extends through and beyond the E. nigricans Speg., Soc. Cient. Ar- tropics in America. gentina An. 19: 47. 1885. .

30 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Dothichloe nigricans (Speg.) Sea- Florida, 1935, E. West No. 10744 (Fl, ver ap. Stevenson, Porto Rico N, W). On Hymenachne a?nplexicaulis (Rudge) Nees Dept. Agr. Jour. 2 (3) : 151. —Puerto Rico, 1912, J. R. Johnston No. 1918 (name only) 4206 (N, W); Trinidad, 1947, R. E. D. Baker No. 1733 (W). D. strangulans Diehl, in litt. On Panicum aciculare Desv..—Georgia, 1933, Hypothallus as an ephemeral, effuse, Geo. Crisfield (W). P. boliviense thin, white, prosenchymatous layer or On Hack.—Cuba, 1922, G. M. Fortun (W). film of variable size, usually covering a On P. clandestinum L.—New York, 1920, considerable area, 3 to 30 mm. long on J. Rubinger (F, W); 1920, W. W. Diehl the internodes just below the nodes or on (F, W); 1938, W. C. Muenscher No. the nodes, variable in thickness, rarely 7873 (W); Virginia, 1931, 1932, W. W. Diehl (F, W); Pennsylvania, 1939, up to thick. Ephelidial fructification 50/^ C B. Stifler No. 43 (N). ephemeral, elongate subhysteriform, On P. comtnonsianum Ashe—Delaware, 1939, white to gray, erumpent through the C. L. Lefebvre (W). On P. commutatum Schult.—Florida, 1898, R. hypothallus; hymenium of a palisade of Combs (W). conidia upon short, simple conidiophores On P. cyanescens Nees—Trinidad, 1947, R. E. rarely swollen and branched; conidia 20/x D.Baker No. 1522 (W). to AQ)(i by 0.5iM to 2/u. Typhodiumlike On P. depauperatum Muhl.—Massachusetts, 1892, W. A. Setchell (Calif. Univ. Herb. conidia developing under conditions of No. 48779); 1894, W. A. Setchell extreme moisture upon the ascostroma (Calif. Univ. Herb. No. 48778). as a layer of irregularly swollen to spatu- On P. hemitomon Schult.—Florida, 1940, late conidiophores, each with an apical C. L. Lefebvre (W). On P. huachucae Ash^-Virginia, 1932, A. sterigma bearing a single conidium, J. Stevenson and W. W. Diehl (W); conidia obovate up to 4^ by 3ja. Ascos- W. W. Diehl (W). tromata variable, as uniform effuse layer On P. implicatum Scribn.—New York, 1920, over the hypothallus or broken up into C. Drechsler (F, N, W). On P. laxuni Swartz—Venezuela, 1940, A. pulvinuli separated by sterile areas; sur- Chase Nos. 12546, 3916 (W); Trinidad, face uniform to rugulose, black punctate 1945, R. E. D. Baker No. 755 (W); from the slightly to prominently emer- 1947, R. E. D. Baker No. 1524 (W). P. lindheimeri Maryland, gent ostioles; context white; subperithe- On Nash— 1915, A. Chase (I, N); Virginia, 1915, C. V. cial stroma prominent; perithecia crowd- Piper (W, N); 1928, 1929, 1930, 1931, ed compressed, elongate elliptical to 1933 W. W. Diehl (W); 1928, R. W. elongate lageniform 210/^ to 500/x by Davidson and W. W. Diehl (W); 1932, A. Stevenson and W. Diehl (W); 100/x to 200/A with elongated ostiole; J. W. West Virginia, 1946, W. W. Diehl; perithecial walls 10/^ to thick; asci 18^ Tennessee, 1935, L. R. Hesler No. 9401 elongate clavate to cylindrical with elon- (W). gate stipelike base, 105/M to 210/u by On P. linearijolium Scribn.—Massachusetts, 1930, 1932, W. Diehl (F, W). 5/M to Gil with 8 spores; ascospores 95/w W. On P. lucidum Ashe-Florida, 1946, E. West to 205/Aby l/x to 1.5^. (W). On P. meridionale Ashe—Indiana, 1926, SPECIMENS CITED C. C. Deam (N, W); New York, 1927, W. C. Ferguson (P, F). On Gramineae {Paspalum ? sp.)—French Guiana, F. R. Lepr-ieur ex Herb. Mon- " The following corroborative notation has been tagne No. 971, Herb. Paris, according to kindly furnished by C. L. Shear (Feb. 1932), "Mon- photograph and notation by C. G. Lloyd tagne's type of this species in Crypt. Guayan. no. 468 of specimen of type of Cordyceps strangu- is represented by Leprieur's specimen in Montagne's Herbarium in Paris. This specimen bears Montagne's lans Mont. 11 autograph label 'Cordiceps strangularis' , and con- On Gramineae (indet.).—Guarapi, Paraguay, sists of two small stromata on the culm of a grass 1883, type of Epichloe nigricans Speg. which looks like a species of Panicum. The stromata are black, about Vs of an inch or less long, minutely in Bal- comm. C. Spegazzini (W); dupL roughened by the ostioles. There are no duplicates ansa Plantes du Paraguay No. 3741 (F, in the Herbarium according to Patouillard. N); 1887, C. Spegazzini, F. guarantici "Specimens of Epichloe nigricans Speg., PI. Parag. of Balansa 4337 on Panicum and 4321 and 3741 look (F, No. 31 N). like Montagne's specimen but the stromata are On Axonopus compress us (Swartz) Beauv.— longer. -C.L.S. Paris 1923." BALANSIA AND THE BALANSIAE IN AMERICA 31

On P. tnicrocarpon Mijhl.—Maryland, 1922, 1926, W. W. Diehl (F, W). On P. nitidum Lam. (?).—Cuba, 1903, N. L. Britton and P. Wilson No. 341 (N). On P. pilostim Swartz—Minas Geraes, Bra- zil, 1933, A. S. Muller (W). On P. polygonatum Schrad. (?).—British Guiana, 1919, A. S. Hitchcock No. 16,691 (F, N, W); Nicaragua, 1945, A. G. Kevorkian No. 50 (W). On P. scabriusculum Ell.—Mississippi, 1889, S. M. Tracy and F. S. Earle No. 220 (N); S. M. Tracy No. 200 (Mo). On P. schiffneri Hack.—Minas Geraes, Bra- zil, 1923, A. S. Muller No. 490 (W). On P. scoparium Lam.—Georgia, 1933, Geo. Crisfield (F, W); Virginia, 1937, 1941, C. L. Lefebvre and W. W. Diehl (W); Kansas, 1888, W. T. Swingle No. 1165 (W). On P. scribnertanum Nash—Nebraska, 1938,

J. L. Fults (W). On P. tsugetorum Nash—New York, 1892, S. E. Jelliffe (N). On P. zizanioides H. B. K.—Venezuela, 1940, A. S. Muller No. 3923 (W); Trinidad, 1947, W. T. Dale No. 1710 (W). On Panicum sp.—Massachusetts, 1893, W. A. Figure 70.— Distribution Balansia strangulans Setchell (Calif. Univ. Herb. No. 48780); of New York, 1892, S. E. Jelliffe (N); (black dots) and 6. strangulans f. discoidea

New Jersey, 1879, J. B. Ellis (N); (black squares). 1903, C. L. Shear (W); Delaware, 1908, H. S. Jackson (W); Mississippi, 1889, zini's (1885) Epichloe nigricans was S. M. Tracy (Mo, W) ; Cuba, C. Wright based upon longer stromata otherwise of No. 916 in Herb. Curtis (F); Brazil, 1898, F. Noack in Herb. Theiszen (F). similar character. Many specimens cited On Sacciolepis striata (L.) Nash—Florida, above are intermediate between the two 1931, W. W. Diehl (F, W). extremes; some are even considerably On Gramineae.—Florida, 1891, R. Thaxter longer and more sleevelike than Spegaz- (F); New York, 1890, B. D. Halsted in Seymour and Earle Econ. Fungi No. zini's type. Some with the shorter stro- 548 (F, N, Mo, W). mata approach the succeeding species of In ILLUSTRATIONS the present account. many instances stromata are not continuous but tend to Plate 5, A to F; Viegas, Bragantia 2: be small, separate, subdiscoid, or irregu- plates 120 and 121. lar units upon the collarlike hypothallus, TYPE LOCALITY apparently as a result of suppression of French Guiana on unknown grass. what would ordinarily be a continuous DISTRIBUTION stroma over certain areas of the hypo-

On culms and leaf sheaths of various thallus. This condition has naturally grasses in the Atlantic drainage of North given rise to confusion in distinguishing and South America from Massachusetts to this from other species. As a result some Paraguay and in the West Indies (fig. 10). published records unsupported by speci- QUESTIONABLE RECORDS AND ERRORS IN mens will remain indefinite. IDENTIFICATION Ascostromata of Balansia strangulans Montagne's Cordyceps strangulans frequently fail to develop uniformly over was described in 1855 and 1856 from the hypothallus, often developing as short effuse stromata, collarlike below separate small stromata with sterile areas the nodes of the host, whereas Spegaz- between. These small stromata in some 32 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE cases are uniformly discoid with rem- strangulans, is practically indistiguish- nants of the hypothallus as gray to black able from it. layers as a "collar" the sterile below Viegas, (1944a, p. 91) has reported nodes on grass culms. This condition is this species from Sao Paulo in Brazil as occasional on some grasses, as on Pani- Dothichloe nigricans as on Paspalum cum clandestinum, for instance; on other s chaffneri Griseb. and as D, suhnodosa grasses, as suggested by herbarium speci- on Panicum pilosum and Lasiacis ligu- mens, it is possibly a consistent feature. lata. His illustrations are convincing, and It is this discoid condition that has been the writer has examined a replicate, cited considered the diagnostic feature for above, of his specimen on P. pilosum B. discoidea. Where stromata are less (1933, A. S. MuUer). His host identi- regularly discoid the condition approxi- fication of Paspalum schaffneri Griseb. mates that described by Chardon (cf. is in disagreement with that of Mrs. Chardon and Toro 1934) for his Doth't- Agnes Chase, who has determined the chloe granulosa, based upon a Venezue- Department's (W) duplicate specimen lan specimen on a grass host as yet as Panicum schiffneri Hack.; this de- undetermined. Chardon's species was re- termination is in agreement with the ported by Baker and Dale (1948) from report on the same specimen by Chardon, Barbados and St. Vincent. These speci- Miller, and Muller (1940). Incidentally mens and those of Chardon's type are their report of a "Panicum ajfolyroides" intermediate between the forms discoidea as a host for this fungus is evidently a and typica. printer's error for "aff [inis] olyroides" Some confusion in citing the name (H.B.K.); Mrs. Agnes Chase suggests Dothichloe nigricans (Speg.) Seaver has that the plant could well be P. cervicatum arisen because the combination was pub- Chase. In original lished by Stevenson (1918, p. 151) with Montague's description of reference to the related species D. subno- the species and in the recapitulation in dosa (q.v.), but the name of the former his Sylloge (1856) the host was reported is frequently credited to Chardon (1927, as "Paspalum," a determination somewhat unlikely in the light of Lloyd's p. 296) , who first applied the combination to Spegazzini's species. photograph (pi. 7) which suggests a species of Panicum or possibly Lasiacis Like other species of Balansia this has or Sacciolepis. at different times, especially on herb- Spegazzini's type of Epichloe nigri- arium labels, been referred to Dothidea cans was described (1885, 1887) from vorax Berk. & Curt., as was done by a specimen on an unnamed grass that Berkeley (1876, p. 105) with reference could also be considered a Panicum. He to what must have been B. strangulans (1891) later described another specimen on Panicu?n. {Balansa No. 4321), also from an un- Although the species is fairly common named grass in the same area, which he and widely distributed in eastern North referred doubtfully to the same species America, it is surprising that Atkinson because of the larger asci, 150/x to l60fi had not given it any consideration. There by 6/x, which are actually moderate-sized is, however, one specimen of it in his for mature asci of this species. His type herbarium lacking any information as to described with asci 90/x to 100 fi by 4fi origin, which he labeled "Dothichloe to 5 /A is obviously immature; in fact the aristidae forma panici." Diehl (1930b, specimen bears remnants of the epheli- 765) in a discussion of its conidial p. dial stage. fructification likewise referred to it as D. aristidae. Actually the ephelidial stage PATHOLOGICAL CONSIDERATIONS of D. aristidae, although less conspicuous In order to keep the species in living and more transitory than that of B. condition under observation for long BALANSIA AND THE BALANSIAE IN AMERICA 33

periods of time, the writer has at various Various inoculation attempts have times transplanted grasses infected with been unsuccessful, but infected plants Balansia strangulans from the field to a showed every indication of bearing per- garden in Arlington County, Va., in ennial infection in culms. It has been certain instances in 1931 to 4-, 6-, and of considerable interest that since the 10-inch pots to a greenhouse in Arling- stromata appear as "collars" on the inter-

ton, and in 1931 and 1932 into a green- nodes the basal stroma is the oldest and

house at Cambridge, Mass. The garden the terminal one the youngest, hence it observations were as follows: 1929-35, is often possible to find both ephelidial Pankmn lindhelmeri; 1928-35, P. clan- and ascogenous fructifications at the desUnum\ 1931-48, Sacciolepis striata. same time. The greenhouse cultures at Cambridge If, as seems likely, the infection takes were 1930, 1932, Vanicum linearijolium, place by way of the stigma and ovule, and 1931-32 on Sacciolepis striata; and the infected seed would give rise to at Arlington were 1931-32, Panicum plants doomed to systemic infection and elandestinum, P. lindheimeri, and Saccio- consequent sterility. Nevertheless, the lepis striata. fact that in two instances infected grasses have lost superficial evidence of infection In all cultures the fungus fruited each suggests that they may outgrow the para- season even when the fungus-laden culms site or that the intramatrical parasite can were removed the previous season. die or be enfeebled. Marked clumps of Panicum clandes- Of the above-cited specimens one tinum after the fifth year, i.e., in 1936, (A. Chase No. 12546) on Panicum and thereafter failed to produce the laxum from Venezuela includes a normal fungus and were apparently no longer inflorescence borne upon a culm bearing infected. The transplants of Sacciolepis ascostromata of Balansia strangulans, to the garden likewise ceased to produce demonstrating for this species an escape the fungus after 1935, apparently being from infection possibly comparable to normal for the next 13 years. As in that here recorded also for B. obtecta on related species the infected plants were Cenchrus. West's (1945) report of this of a normal green color, superficially fungus on Axonopus from Florida has indistinguishable from healthy plants ex- been noted by Weiss (1943) to cause cept for a peculiar type of dwarfing. the condition termed "black ring." All the writer's cultures of plants infected by this fungus were characterized 6. Balansia strangulans forma discoidec by shortened internodes with consequent Balansia strangulans forma discoidea dwarfing of the culms and by narrowed (P. Henn.) comb. nov. and generally smaller leaves, giving clumps a "broomed" apearance. This has B. discoidea P. Henn., Hedwigia been especially noticeable in P. clandes- (Beibl.) 39: (77). 1900. tinum. In this condition it is of interest B. trichloridis Speg., Mus. Nac. de that Viegas (1944) has reported and Hist. Nat. An. Buenos Aires 23: illustrated a witches' -broom condition, 96. 1912. seemingly arising from a normal shoot Dothichloe gramdosa Chardon ap. of Lasiascis clearly in reference to this Chardon & Toro, Puerto Rico species, but there referred to as Dothi- Univ. Monog. Phys. and Biol. chloe subnodosa. In the writer's experi- Sci., ser. B., 2: 140. 1934. ence the "brooming" seems alv/ays to D. discoidea (P. Henn.) Diehl ap. affect an entire plant, or rarely a normal Chardon, Miller and Muller, My- shoot may arise from a diseased plant; cologia 32: 174. 1940. but both were of the same healthy In many instances ascostromata of Bal- coloration. ansia strangulans fail to develop uni- 34 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

formly over the hypothallus, often devel- H. von Tuerckheim ex U. S. Natl. Herb. No. oping as separate stromata with sterile 576813 (W). On Panicum sp.—Minas Geraes, Brazil, 1929, areas between. These small stromata in A. Chase No. 102881/, (W); 1934, some cases are uniformly discoid, with A. S. Muller No. 1136 (W). remnants of the hypothallus as gray to On Gramineae.—Miranda, Venezuela, 1932, C. E. Chardon, R. A. Toro, R. black sterile areas, each hypothallus as and Alamo No. 894 ex type of Dothichloe in B. strangulans initially forming a granulosa Chardon ex Herb. Dept. PI. "collar" below the node on grass culms. Path., Cornell Univ. (W). This condition is occasional on som.e ILLUSTRATIONS grasses, as on Panicum clandestinum, for Plate 5, G and H; Atkinson, Jour. Mycol. instance; on other infected grasses it is 11: plate 88, figures 22, 23, 24; Viegas, Rev. possibly a consistent feature as suggested Agron. 7 (4) : plate 4. by herbarium specimens. It is this discoid TYPE LOCALITY condition that has been considered the Blumenau, Santa Catharina, Brazil, on culm diagnostic feature for B. strangulans f. and leaf sheaths of Chloris distichophylla. dhcoidea. Where stromata are less regu- DISTRIBUTION larly discoid the condition approaches that described by Chardon for Dothkhloe On culms and leaf sheaths of panicoid grasses, Kansas and Central America to granulosa. Brazil and Paraguay (fig. 10).

SPECIMENS CITED DISCUSSION Ule's specimen as No. 3309 in the Andropogon On (?) sp.—Kansas, 1887, Farlow Herbarium from Huallaga, Bra- W. A. Kellerman and W. T. Swingle (N). zil, would seem to be from a different On Bouteloua curtipendula (Michx.) Ton.— place than his No. 3309 reported by Kansas, 1921, R. P. White No. 2,000 Hennings (1904b) from Sahnas de (W). Pilluana, Peru, as Balansia discoidea on On Chloris disiichophylla Lag.—Blumenau, Chloris sp. It is this latter specimen (not Santa Catharina, Brazil, 189— (?), E. Ule No. 1334 ex type of Balansia dis- seen by the writer) that has been re- coidea P. Henn. (A); Huallaga, Brazil, ported from Peru by Garcia Rada and 1903, E. Ule No. 3309 (F). Stevenson (1942) and incorrectly re- On C. polydactyla (L.) Swartz—Venezuela, to as the "type" of B. discoidea. 1937, A. S. Muller No. 2038 (W). ferred On Chloris sp.—Guarapi, Paraguay, 1884, Balansa's Plantes du Paraguay No. 4320 Balansa PI. du Parag. No. 4320 (F). was reported by Spegazzini (1891) as On Gymnopogon sp.—Goyaz, Brazil, 1930, his B. claviceps; it is actually very similar A. Chase No: 11329 (W). On Ichnanthus candicans (Nees) DoelL— to the above-cited Kansas specimen on Minas ,Geraes, Brazil, 1933, A. S. Andropogon (?), which was illustrated Muller No. 475 (W). by Atkinson (1905, fig. 24). Spegazzini On Lasiacis ligulata Hitchc. & Chase—British (1912) appears to have reported the Guiana, 1922, F. L. Stevens No. 971 as (F, N, W); Guatemala, 1932, More- same fungus on Trichloris mendocina land and Pettit (W). his B. trichloridis. On Oplismenus hirtellus (L.) Beauv.—Minas As noted previously, the specimens of Geraes, Brazil, 1933, A. S. Muller No. B. strangulans so often show a tendency 641 (W); Barbados, B. W. I., 1945, separate stromata, notably on R. E. D. Baker, I. C. T. A. Nos. 610, to bear 611 (W). Panicum clandestinum, that the separate On O. setarius (Lam.) Roem. & Schult.— stromata are rarely truly discoid; it is, St. Vincent, B. W. L, 1946, R. E. D. therefore, not included here. Although it Baker, I. C. T. A. No. 926 (W).

f . discoidea On Panicum boliviense Hack.—Panama, 1935, is likely that B. strangulans G. W. Martin No. 3038 (W). is merely a depauperate form of the Presl Rica, On P. haenkeanum —Costa 1923, species, it is noteworthy that B. strangu- F. L. Stevens 19-964 (W); No. 694 lans typica, with one exception, has not (F, N). far found upon any of the On P. millegrana Poir.—Guatemala, 1904, thus been BALANSIA AND THE BAlANSIAE IN AMERICA 35 host species known to bear the discoid perithecia obovate to lageniform, often forn-i. As Dothkhloe disco'idea it has crowded, 150/x to 300;x by 125/x, to 150/x, been reported and illustrated by Viegas ostioles short; perithecial walls 8^ to 16^ (1944b) from Rio Grande do Sul, in thick; asci 150^ to 200/x by 3/x to 4.5/x, Brazil, based on a specimen by Costa ascospores 120^ to 180^. Neto (No. 4791). Viegas here correctly SPECIMENS CITED pointed out that Hennings' type was On Ichnanthus leiocarpiis (Spreng.) Kunth from Blumenau in Santa Catharina. —Trinidad, 1947, R. E. D. Baker No. Chardon, Miller, and Muller (1940), 1731 (W). in noting this fungus from Minas On /. neynorosus (Swartz) Doell—Trinidad,

Geraes, Brazil, reported it as on Ichnan- 1947, R. E. D. Baker No. 1459 (W). On /. pall ens Munro—Puerto Rico, 1916 ihiis candidus, apparently in error for H. H. Whetzel and E. W. OIi%- Ni^

/, candka^'"^ ~ Mu.ll^«-'« Rr^-^Ui"'^ s>^<^ciinen r.9Q -'tx- --IJ 'N Y~l [U A .^-i^ ciuea aDovc sriows sume stromaca witK nie\^ .6 liiDcleci as on /. axillaris Hitchc. some short stipes (up to 1 mm. long), & Chase), 691 (N); 1913. F. L. Stevens No. 778 (A, W); 1916, F. L. a condition possibly more frequent under Stevens No. 691 (ex Herb. J. A. Steven- certain conditions. Stipitate stromata are son) (W); 1924, Whetzel, Kern, and easily broken off and are not so likely Goss Nos. 2503, 2532 (W); Mayera to be represented in herbarium specimens Island, Windward Islands, B. W. 1., 1932, P. H. Dorsett (F, W). as the sessile ones. Onl. tennuis (Presl) Hitchc—Trinidad, 1947, Baker and Dale (1948) have reported R. E. D. Baker No. 1458 (W). Dothkhloe granulosa from Barbados on On Panicum trichanthum Nees—Puerto Rico, Oplhmenus hirtellus and from St. Vin- 1912, J. R. Johnston No. 4212 (W); 1916, A. Stevenson No. 5798 (W, cent on O. setarius\ duplicates of these J. specimens made available through Dr. On Panicum sp.—Santa Catharina, Brazil, Baker's kindness are cited above. 1885, E. Ule in Rehm, Ascomyceten No. 924 (A, F, N), and in Rabenhorst, 7. Balansia subnodosa Fungi Europaei No. 3563 (F, Mo, N, K); Trinidad, B. W. I., F. J. Seaver Balansia subnodosa Atk. ap. Chardon, (N). Mycologial3: 287-288. 1921. ILLUSTRATIONS Dothkhloe subnodosa Chardon, Plate 5, / and /; Chardon, Mycologia 13: Mycologial3: 287. 1921. figure 1, plate 14, figure 14.

Hypothallus restricted to a limited TYPE LOCALITY area, rarely more than 3 mm. long, sur- Mayaguez, Puerto Rico, on Ichnanthus rounding the culm of the host plant at pallens Munro. the node or on the internode immediately DISTRIBUTION below the node, as an effuse prosen- On species of Ichnanthus and Panicum, chymatous film at first white, black with West Indies and Brazil (fig. 11). age, sometimes ephemeral. Ephelis fructifications ephemeral, covering the sur- DISCUSSION face of the hypothallus separated by exci- Although superficially distinct by rea- pular strands, conidia as a palisade upon son of its single ring of subglobose to simple conidiophores, each Iju, in diam- discoid stromata over a narrow hypo- eter (branched or swollen conidiophores thallus, it is obviously very close to Bal- not seen in dried specimens) ; conidia ansia strangulans, of which some speci- 18/x to 30^ by 1/x to 2^. Ascostromata mens show a very short "collar," often subglobose, slightly flattened, sessile, with irregular subglobose stromata cover- contiguous or distinctly separate, usually ing the hypothallus. In the type of B. as a circlet of 3 to 6 upon the hypothallus strangulans the collar is very short, al- in a single row about the culm of the though the ascostroma is continuous. host plant, 1 to 3 mm. in diameter; But until more can be learned about B. 36 AGRICULTURE MONOGRAPH 4. U. S. DEPT. OF AGRICULTLTIE subnodosa it is convenient to regard it as Jour. 10: 377. 1869. -nee in a species rather than a variety. sensu typis. Ule's 1885 specimen on an unknown PHypocrea subviridis Berk. & Curt. grass from Brazil, issued in both Rehm's ap. Berk., Grevillea 4: 15. Ascomyceten and in Rabenhorst- 1875. Pazschke Fungi Europaei reported by PHypocrella subviridis Berk. & Rehm (1888) as Ophiodothis rorax and Curt., ap. Cke., Grevillea 12: again (1897) as O. vorax var. p'llulae- 105. 1884. forfuis, is t)'pical of B. subnodosa. Two Dotbichloe hypoxylon Atk., Torrey gatherings in Puerto Rico, one on Ich- Bot. Club Bui. 21: 223. 1894. nanthus pallens reported by F. L. Stevens nee Pk., error ident., name ap- (1917) as Dotbichloe aristidae and one plies to Epichloe hypoxylon Pk. on Vanicum trkhanthtim reported by Ste- Ophiodothis henningsiay^a Moell., venson (1918) as Balansid hypoxylon, Phyc. u. Ascomyceten Bras., pp. are both B. subnodosa. Viegas' report 188, 302. 1901. (1944a) of B. subnodosa in Brazil on Hypothallus eifuse as a flat, thin, Lasiacis Ijgulata and on Vanicum pilosum white, prosenchymatous layer or film, are here considered to refer to B. becoming black vvith age, on abaxial strangulans; the specimen on P. pilosum surfaces of unrolling leaves, rarely ex- is possibly a borderline case. ceeding 15ju, in thickness (vertical diameter) and 80 mm. in horizontal (linear) diameter, rarely as separate units. Epheli- dial fructifications ephemeral, linear to hysteriform, excipies thin evanescent, covering the surface of the hypothallus, swollen and irregularly branched; coni-

dia 11/x to 30^ by l/>t to 2/x. Ascostro- mata variable, effuse maculiform, usually 10 to 20 mm. but often 80 mm. along the length of a leaf sheath, continuous or often as separate contiguous pulvinuli separated by sterile areas; surface uniform to rugulose, even collicular, black, sm.ooth to punctate, roughened by the slightly emergent ostioles; context white; subperithecial stroma scant}'; perithecia lageniform, often but not always crowded, compressed, 150^ to 300/a by 100^ to 150/x, ostiole short, sometimes exsurgent; perithecial walls 6/x to 15/x thick; asci 80/a to 180^ by 4/x to G^\ ascopores nearly the length of the ascus

and up to 15yLi in diameter.

Figure 7 7.— Distribution of Balansia subnodosa CITED (black squares) and B. henn/ngs/ano (black SPECIMENS dots). On Andropogon bicornis L.—Brazil: Minas Geraes, 1929, A. S. Muiler No. 19 (W); 8. Balansia henningsiana Matto Grosso, 1930, Agnes Chase No. Balansia hemiingsiana (Moell.) comb, 10954 (F, W). nov. On A. leucostachyus H. B. K.—Puerto Rico, 1915, F. L. Stevens No. 8211 (A, N); Hypocrea atramentosa Berk. & Curt, Minas Geraes, Brazil, 1933, A. S. Mui- ap. Berk., Linn. Soc. London Bot. ler No. 473 (W). BALANSIA AND THE BALANSIAE IN AMERICA 37

On A. scoparius Michx.—Kansas, 1935, C. L. Berk. & Curt.; Alabama, 1890, S. M. Lefebvre (W); Missouri, 1941, G. D. Tracy (A). Darker No. 6886 (Mo, W); Maryland, ILLUSTRATIONS 1946, W. W. Diehl (W); Virginia, 1939, 1940, 1941, W. W. Diehl (W); Plate 6; Moller, Phyc. u. Ascomyceten Bras. Florida, 1931, W. W. Diehl (F, W); figures 66, a, b, 70; Atkinson, Jour. Mycol. Louisiana, 1930, R. W. Davidson (W). 11: plate 85, 88; figures 25 to 27. A. virginkus L.— York, H. D. On New 1919, TYPE LOCALITY House (Pk.); 1940, R. Latham No. 2285 (W); Maryland, 1946, W. W. Blumenau, Santa Catharina, Brazil, on Diehl (W); North Carolina, 1949, leaves of Andropogon sp. W. W. Diehl (W); Tennessee, 1939, DISTRIBUTION R. W. Davidson (W); Alabama, 1891, G. F. Atkinson No. 2094 (A), No. On abaxial surfaces of leaves of various 1164 (A) and C. L. Newman No. 1965 panicoid grasses from Kansas, New York, (A); Georgia, 1933, George Crisfield Mexico, and the West Indies to Santa Cath- (W); Florida, 1911, F. L. Lewton (W); arina in Brazil (fig. 11). Dominican Republic, 1930, E. L. Ekman (W). NOMENCLATURE On Andropogon sp.—Alabama, (H. Beau- In Atkinson's (1894) initial treat- mont) paratype coll. of Hypocrea at- ment of his genus Dothichloe and of the ramentosa pro parte in Herb. Curtis and species under consideration, which was in Herb. Farlow (F); 1890, G. F. Atkin- son No. 1169 (A); S. C, Ravenel, Herb. the first species referred to that genus, Ellis (N); Georgia, 1940, C. L. Le- he appUed to it the newly combined febvre (W); Florida, 1892, T. W. name Dothichloe hypoxylon (Pk.) Atk., Swingle No. 3064 (W); 1896, P. H. on the mistaken assumption that Peck's Rolfs in Herb. Earle 22 (N, Fl); 1915, H. E. Stevens No. 5215 (Fl) Missis- (1875) name Epichloe hypoxylon was sippi, 188—, S. M. Tracy 28 (Mo, N); applicable. To correct this mistake Atkin- 1891, S. M. Tracy (W); Blumenau, son (1905) supphed for his species the Santa Catharina, Brazil, 1890-93, ex type of Ophiodothls henningsiana Moell., name D. atramentosa as a new combina- (A, F). tion based on Hypocrea atramentosa On Chloris sp.—Florida, 1924, G. F. Weber Berk. & Curt., but erroneously citing as (W, Fl); Tampico, Tamalipas, Mexico, the type a specimen on Andropogon 1910, A. S. Hitchcock No. 5798 (N). found in Alabama by H. Beaumont, No. On Gymnopogon ambiguus (Michx.) B. S. in the Berkeley Herb- P.-Mississippi, 1888, S. M. Tracy Nos. 4018, which was 31, 82 (N). arium at Kew. On Panicum agrostoides Spreng.—Mississippi, Hypocrea atramentosa was described 1889, S. M. Tracy in Ell. and Ev. North in Berkeley's (1869) article on Charles Amer. fungi No. 2373 as Hypocrea hypoxylon (A, F, N, W); 1890, S. M. Wright's collection of Cuban fungi, the Tracy No. 1550 (N). Latin diagnosis being clearly in reference On P. anceps Michx.—Mississippi, 1886, to Wright's specimen No. 419 as the S. M. Tracy No. 34 (N); 1888, S. M. type and with another specimen as para- Tracy No. 78 (Mo, N); 1892, M. J. 4018." White in Herb. Tracy No. 1847 (N); type, cited as "Alabama No. Missouri, 1924, C. Drechsler (F, W). Duplicates of both these specimens are On P. proliferum Lam.—Alabama, 1901, to be found in the Curtis Herbarium. G. W. Carver (F). Berkeley and Curtis undoubtedly con- On P. schiffneri Hack.—Minas Geraes, Brazil, 1933, A. S. Muller (W). sidered the two specimens conspecific, On P. stipitatum Nash—Virginia, 1932, although they are clearly different W. W. Diehl (F, W). species. The Cuban specimen No. 419 On P. vjrgatum L.—Kentucky, 1934, F. T. (type) is referable to Myriogenospora McFarland (W); Virginia, 1924, C. Drechsler (F, W). pasp all, described by Atkinson (1894) On Panicum sp.—Virginia, 1936, R. F. along with his first account of Dothi- Smart (W). chloe. It is pertinent to review the On Gramineae.—South Carolina, M. A. Cur- peculiar wording of the original diag- tis in Herb. Curtis (F) and in Michener Herb. (W), type of Hypocrea subviridis nosis and description of Hypocrea atra- 38 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTLUE mentosd. The diagnosis reads: "Effusa, incorrectly identified. Diehl (1930), for tenuis, elongata, atra, demum ragosa; instance, in reporting Sporoholus herte- peritheciis globosis ostiolisque inimersis Yoanus as a host for Dotbicbloe atra- (419)/' applying distinctly to No. 419, mentosa, in the light of present opinion, Myr20ge72ospora with its elongate stroma was concerned with 5". poiretii as a host and immersed ostioles. This is followed for D. epicbloe (cf.). In the account of by "On leaves of grasses Hah. Alabama. Chardon, Miller, and Muller (1940) On Andropogon, No. 4018, forming a S. poiretii is noted as a host for D. atra- thin stratum on the underside of the mentosa, but in that case also the fungus leaves," this latter applying to Atkin- is Balansia epicbloe. In the sam.e account son's concept of Dothicbloe. Unwitting- and in that of Viegas (1944a) as hosts ly, Berkeley and Curtis thus described for D. atramentosa there are reported separately and, in the light of present severally "Andropogon leptostacbyns'' information, distinguished both speci- and ''A. leptostachys Benth.," both ap- mens in their several species. Duplicaces parently in reference to the same speci- of both these specimens are in the Curtis men; in this case the host is most prob- Herbarium, and in the Curtis specim^en ably A. leiicostacbyus H. B. K., an of Wright's No. 419 is a notation by ubiquitous grass in Brazil, these errone- the late Roland Thaxter recogizing it as ous host-name spellings probably being Myriogenospora. D. atramentosa (Berk. merely errors of transcription.

& Curt.) Atk. is not applicable on It is possible, even likely, that Ule's nomenclatorial grounds to the species as Brazilian specimen No. 60 (not seen by Atkinson conceived it. the writer) reported by Pazschke (1892) Hypocrea suhviridis of Berkeley and as Epicbloe nigricans "ad caul, et fol. Curtis, referred to Hypocrella by Cooke Andropogonis macruri'' is referable to (1884), appears on the evidence of part Balansia benfiingsiana and that the host of the type in the Michener Herbarium is a species other than A. macrotiri and of another in the Curtis Herbarium Michx., synonymous with A. glomeratits to be the remnants of the ephelidial (Walt.) B. S. P., a species unknown in stage of BalansJa henmngsiana. But these Brazil. specimens are in too poor condition to be selected as types; and as bearing only PATHOLOGICAL CONSIDERATIONS conidial fructifications they are inconse- Where the v/riter has sought this quential in the nomenclature of an asco- fungus in likely places at times when mycete. a potential host was in flower, Balansia Balansia hennings'tana has been re- benningsiana has been found most readi- ported under the name of Dothicbloe ly by first noting individual clumps of atramentosa and its many synonyms in the grass that failed to bear inflorescences. various accounts, sometimes including Here the fungus would be found on the other related species, notably by Sac- under side of leaves. Earlier in the cardo (1879), Cooke (1884), Earle season before the time of flowering, (1900), Chardon (1921), Diehl grasses infected by the fungus were in-

(1930b), Miles ( 1935) , Maneval distinguishable in habit and coloring, (1937), Chardon, Miller, and Muller and the fungus could be defected only (1940),Miller (1941), Viegas (1944a), by critical examination at close range. and Presley (1947). As Hypocrella by- When infected grasses were transplanted poxylon it was reported by Tracy and to Arlington County, in Virginia, they Earle (1895) and Presley (1947). remained sterile, continuously failing to It is naturally to be expected that of produce any evidence of an inflores- the various hosts recorded in the litera- cence : Andropogon scoparius from Louisi- ture and especially on many of the labels ana, cultivated during the seasons 1930- of specimens in herbaria some have been 33; ^. scoparius from Virginia, seasons BALANSIA AND THE BALANSIAE IN AMHRICA 39

1939.41; Pau/cu/// stipilatum from Vir- usually broken up into separate contigu- ginia, seasons of 1932-34. As in inous flattened pulvinuli, separated by nar- stances of plants infected by related row sterile areas, 30^ to 90/a thick or species of Balansiae, in no case was effuse, surface reddish black, smooth to there any material chlorosis or other rugulose, ostioles inconspicuous; context evidence of disease than sterility; and no white to reddish; subperithecial stroma brooming or fasciation was observable. slight; perithecia not crov/ded, obovate- Nevertheless the culms of affected plants lageniform, 200^ to 380^ by 120;ji to being slightly dwarfed and stiffer than 200;x; perithecial walls 12;^ to 20/x, normal ones presented a crowded appear- thick; asci often long stipitate, clavate

ance simulating brooming. All the cul- cyhndrical, 90/x to 125/x, by 4^ to 6;*, tures bore every evidence of systemic sporiferous part 36/jt to 90^ by 4/x to 6/x; infection. For instance, for those infected ascospores 36^ to 100/x by l^u to \.'b^. plants wintered outdoors, the dead culms SPECIMENS CITED and leaves bearing fructifications of the On Cyperaceae.—Surinam, type in Herb. parasite were removed each /all, only \o Michenet (W), in Herb. Schv/einitz be followed the following season by (Philadelphia, Pa.), and in Herb. Curtis shoots bearing the fungus, the mycelium (F). of the endophyte being detectable in the On Cyperus diffusus VaU— British Guiana, 1922, F. L. Stevens No. 325 (W). tissues of new shoots well before the ad- On C. ovularts (Michx.) Torr.—Sneed's vent of its fructifications. No instance of Island, Manatee County, Fla., 1894, healthy shoots from infected plants was S. M. Tracy No. 6496, type of Ophio- observed. The abundance of ascospores dothis atramentosa Earle var. cyperi Earle (N), S. M. Tracy No. 6436 (F, produced coincident with the normal Mo, N). time of the flowering of the host pre- On C. retrorsus Chapm.—Florida, 1944, V. K. supposes stigmia infection and seed trans- Charles (W). mission. On C. strigosus L.—Florida, 1939, J. B. Mc- Farlin (W). 9. Balansia cyperacearum On Cyperus sp.—Alabama, 1899, F. S. Earle Balansia cyperacearum (Berk. & Curt.) (N). comb. nov. On Scleria bracteata Cav.—Venezuela, 1939, C. E. Chardon No. 2785 (W); Trini- Hypocrea cyperacearum Berk. & dad, 1948. E. Chenery, I. C. T. A. No. Curt., Acad. Nat. Sci. Phila. Jour. 1920 (W). (ser. 2) 2: 285. 1854. ILLUSTRATIONS Spbaeria cyperacearum Schw., mss. Plate 7, A and B: Berkeley and Curtis, ap. Berk. & Curt. 1854. Acad. Nat. Sci. Phila. Jour. 2 (ser. 2) pt. 4: Hypocrella cyperacearum (Berk. & figures 8, a, b. Curt.) Sacc. Sylloge Fungorum 2: TYPE LOCALITY

580-581. 1883. Surinam, on abaxial leaf surface of Cypera- Ophiodoth'is atramentosa (Berk. & ceae, indet. Curt.) Earle var. cyper'i Earle, DISTRIBUTION Torrey Bot. Club Bui. 27: 122. On abaxial leaf surfaces and culms of 1900. Cyperaceae, Florida and Venezuela to Suri- Dothichloe cyperacearum (Berk. & nam (fig. 9). Curt.) Diehl, in litt. DISCUSSION

Hypothallus effuse, gray through red- Balansia cyperacearum superficially re-

dish to black, thin (about 30/x to 40/x, sembles so closely B. hennings'iana and thick) upon abaxial leaf surfaces or B. strangulans as to be confused with partially surrounding the culm at nodes them. Of the several specimens cited with a longitudinal diameter of 5 to 23 above all are so matured or so parasitized mm. Conidial fructifications not ob- as to preclude determination of the na- served. Ascostroma rarely continuous, ture of the conidial fructifications. On i 40 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

two specimens, F. L. Stevens No. 525 Ophiodothis torax (Berk. & Curt.) and Chardon No. 2785, there are frag- Sacc. var. atramentaria Berk. & ments here considered excipular tissue Curt., in Sacc, Sylloge Fungorum of an ephelidial stage, but no ephelidial 2: 652. 1883. conidia have yet been observed. If these Dothichloe atramentosa Atk., p.p. fragments are not of an ephelidial stage Jour. Mycol. 11: 260-261. or if such stage is not present in the life 1905. nee Hypocrea atramentosa cycle of the fungus, the species would Berk. & Curt. 1869. be relegated to the genus Balansiopsis. U. epichloe Weese, Akad. Wiss. Saccardo (1879) cited Sphaeria cyper- Wien., Math.-Nat. Kl. Sitzber. acearu7n Schw^ as a synonym of Hypo- (Abt. 1) 128: 118. 1919. crella semiamplexa (Berk.) Sacc. This D. atramentaria (Berk. & Curt.) was doubtless a conclusion in- Atk., ap. Davis, derived J. J., Wis. Acad. correctly from Berkeley's (1869) discus- Sci. Arts, and Letters Trans. 24: sion accompanying the description of 299. 1929. Hypo erea atramentosa noting the rela- D. limitata Diehl, Jour. Agr. Res. tionship to H. semiamplexa and presum- 58: 948. 1939. ably citing S. cyperacearum as a synonym Hypothallus effuse, as white prosen- of the latter. H. semiamplexa on bamboo chyma becoming black with age, rarely from. India is, however, clearly distinct, exceeding 15^ in thickness (vertical and Saccardo in the second Sylloge diameter) and 200 mm. in horizontal (1883) recorded it as a separate species. diameter, very rarely as smaller disjunct A specimen on Cyperaceae from Rio units. Ephelidial fructifications ephe- de Janeiro (Ule No. 2450) referred by meral, but at first covering most of the Rehm (1900) to Epkhloe nigricans is surface of the hypothallus as a whitish most likely Balansia cyperacearu?n, al- film that splits irregularly in elongate though the writer has not been able to subhysteriform and coalescing pycnidia verify this opinion by examining the partly separated by the dissepiments of specimen. the surface layer as peridia; hymenium

It is possible that as here recognized as a palisade of conidia (scolecospores) this is a composite species, of which the 18^ to 30^ by 1/A to 2/x, on short simple type is on leaves, w^hereas all the other conidiophores that are rarely swollen or specimens cited are on culms and leaf branched. Ascostromata uniformly ef- bases and are in complete agreement fuse, maculiform, usually 10 to 15 mm. with Ophiodothis atramentosa var. cyperi in horizontal diameter, usually covering Earle (1900). most of the hypothallus, rarely exceeding 200^ in thickness, som.etimes, but rarely, 10. Balansia epichloe separated into pulvini by sterile areas: Balansia epichloe (Weese) comb. nov. surface uniform to rugulose. black, punc- Spbaeria epichloe G. Kunze, nom. tate from the slightly em.ergent ostioles: nud. in sched. in Weigelt's exsic- context white to black, perithecia close cati, 1837; nee Botryosphaeria but not crowded, ovate to lageniform. epichloe (G. Kunze ex Sacc.) 180/x to 315/x by SO^a to 130^, ostioles Sacc, Roy. Soc. Bot. de Belg. papillate, short, perithecial walls 6jM to Bui. 31 (pt. 2): 227. 1892. 12/x thick, asci 105/x to l60/x by 4ju to 6/x Dothidea atramentaria Berk. & with 4 or 8 spores, ascospores 90nx to Curt. nom. nud. ap. Berk., Gre- 120/a by 1/z to 1.5^ villea 4: 105. 1876. SPECIMENS CITED

D. atramentaria Berk. & Curt. f. On Andropogon scop.uius Michx.—North Carolina, 1940, C. L. Lefebvre (W). eragr OS t id is-capillar is Thnem. On Andropogon (?) sp.—South Carolina, nom. nud., Thuem., Myc. Univ. 1868, H. W. Ravenel (A); Alabama, No. 1068, 1879. 1890, G. F. Atkinson Nos. 1168, ll68-a, BALANSIA AND THE BALANSIAE IN AMERICA 41

1546 (A); Missouri, 1929, W. E. Mane- On Panicum agrostoides Spreng.— Missouri, val (W). 1906, H. L. Shantz (W); 1943, G. D. On Calamagrostis canadensis (Michx.) Beauv. Darker No. 7354 (W, Mo); 1943, D. M. Gaebler and G. D. Darker No. -Wisconsin, 1910, 1912, J. J. Davis (N); 1919, C. Drechsler; 1928, E. E. 7364 (W). Honey (W). (This specimen reported On Panicum sp.—Mississippi, 1920, L. E. by Davis, 1929, p. 299, as on "culms" Miles No. 323 (W), No. 719 (W); is actually on adaxial leaf surfaces.) 1930, R. J. Haskell (F, W). On C. inexpansa A. Gray (C. hyperborea On Sporobolus indicus (L.) R. Br.—Ala- Lange)—North Dakota, 1922, J. F. bama, 1904, A. S. Hitchcock (W); Brenckle F. Dak. No. 532 and N. Dak. F. Mississippi, 1922, L. E. Miles No. 719 No. 1492 (W). (W); Minas Geraes, Brazil, 1929, A. S. On Calamagrostis sp.—Wisconsin, 1910, Ex. Muller No. 37 (W); 1934, A. S. Muller Herb. Wis. Univ. (A). (This possibly No. 768 (W). 5". duplicate of J. J. Davis, 1910, specimen On poiretii (Roem. & Schult.) Hitchc— above). Arkansas, 1935, E. C. Tullis (W); On Chloris petraea Swartz—Cuba, 1911, N. Louisiana, 1931, R. W. Davidson (F, L. Overholts No. L. Britton and J. F. Cowell No. 9899 W); 1931, O. 15403 (N, W); Puerto Rico, 1916, H. H. (W); 1940, T. T. Ayers (W); 1948, Whetzel and E. W. Olive No. 694 L. S. Olive (W); Mississippi, 1920, (N, W); 1944, C. E. Chardon 134 1922, L. E. Miles No. 324 (W); 1930, (W); Florida, 1940, C. L. Lefebvre R. J. Haskell (F, W); 1942, J. A. (W); 1924, G. F. Weber (N); 1935, Pinckard (W); Alabama, 1898 (on G. F. Weber No. 10469 (N); 1936, Sporobolus radicans of label), G. W. E. West and R. K. Voorhees (N). Carver (Fl); 1929, J. L. Seal (type of On Chloris sp.—Mexico, 1910, A. S. Hitch- Dothichloe limitata) (F, W); 1931 cock (W). J. L. Seal and W. W. Diehl (F, W) On Ctenium aromaticum (Walt.) Wood- Georgia, 1908, H. A. Allard (W, N) B. Demaree (W); 1930 North Carolina, 1930, 1931, W. W. 1930, 1931, J. 1931, W. W. Diehl (F, W); 1932 Diehl (F, W). E. Benedict, Jr. (F, W); 1933, Geo On Eragroslis capillaris (L.) Nees—Ala- J. Crisfield (F, W); North Carolina, 1933. bama, 1891, G. F. Atkinson (A); South W. W. Diehl (W); Florida, 1928, G. F Carolina, 1876, H. W. Ravenel in De Weber and .. .. Nolen (Fl); 1936 Thum., Myc. Univ. No. 1068 as Dothi- E. West (N); 1931, W. W. Diehl dea atramentaria Berk. & Curt. f. era- (W); 1932, R. W. Davidson (W) grostidis capillaris (F, W, N). [This 1940, C. L. Lefebvre (W); 1943, V. K latter host could well be E. rejracta.— Charles (W); 1943, C. L. Shear (W) W. W. D.} Costa Rica, 1923, F. L. Stevens Nos. On E. hirsuta (Michx.) Nees—Georgia, 10, 111, 151, 157 (I, N, W); Minas 1933, H. Miller (W). J. Geraes, Brazil, 1929, A. Chase No. On E. rejracta (Muhl.) Scribn.—Alabama, 101991/2 (W). 1891, B. M. Duggar (A); 1899, G. F. On Sporobolus sp.—Alabama, 1904, A. S. Atkinson No. 2144 (A); Georgia, 1931, Hitchcock (W); Mississippi, 1920, L. E. W. W. Diehl; Florida, 1931, G. F. Miles (W); Florida, 1941, 1943, C. L. Weber, E. West, and W. W. Diehl Shear (W); Minas Geraes, Brazil, 1934, (F, W); Virginia, 1932, W. W. Diehl A. S. Muller (W). (F. W). On Thrasya petrosa (Trin.) Chase.—Goyaz, On E. triehoc olea Hack. & Arech.—Florida, Brazil, 1930, A. Chase No. 11287 (W). 1947, E. Benedict, Jr. No. 5398 (W). J. On Triodia jlava (L.) Smyth.—Alabama, On Eragrostis sp.—South Carolina ("Grami- 1899, F. S. Earle No. 2094 (N); Kan- neae" of label), H. W. Ravenel No. 259, sas, 1948, C. O. Johnston (W). in M. A. Curtis Herb. (F); {'Poa of On Gramineae (indet.).—Surinam, 1827, label), H. W. Ravenel, Fungi Amer. Weigelt's (1837) exsiccati, type coll. Exs. No. 100 as Dothidea atramen- of Sphaeria epichloe G. Kunze, (W); taria Berk. & Curt. (F, W, N); Ellis, Florida, 1907, R. Thaxter (A). North Amer. Fungi No. 683 (A, F, W, N); Florida, 1897, R. Thaxter in ILLUSTRATIONS Herb. Thaxter (F).

On Gymnopogon ambiguus (Michx.) B. S. P. Plate 7, C and D; Chardon, Mycologia 13: —Virginia, 1936, R. F. Smart (W); plate 14, figure 12; Diehl, Jour. Agr. Res. 1937, A. B. Massey (W); North Caro- 41: plate 2, D, E, and figure 3, D, as Dothi- lina, 1933, F. A. Wolf (W); Georgia, chloe atramentosa; Diehl, Jour. Agr. Res. 58: 1939, Mrs. Sarah M. Tiffney (W); plate 1; Viegas, Bragantia 2: plate 119 as locality (?), F. L. Stevens (A). D. limitata; Weese, Akad. Wiss. Wien, Math.- ,

42 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OE AGRICULTURE

Nat. Kl. Sitzber. 128 (Abt. 1): figures 5 Exsiccati No. 259 (1882) and in Ellis' to 7, as Dothichloe epichloe. North American Fungi No. 683 (1881) TYPE LOCALITY may be regarded as the type.

Surinam on unknown grass. The unnumbered specimen distributed

DISTRIBUTION inWeigelt's (1837) exs'iccsiti a.s Spbaeria epichloe Kunze, lacked an adequate diag- On adaxial leaf surfaces of various grasses nosis remained a practical from North Dakota and Wisconsin, Virginia, and nomen Louisiana, Florida, and southward in the nudum, except for Saccardo's (1892b) West Indies to Surinam and Brazil (fig. 12). Botryosphaeria epichloe, until validated under the name Dothichloe epichloe by Weese (1919), who supplied a suitable diagnosis. Botryosphaeria epichloe Sacc, based upon a Weigelt specimen in the

Brussels Herbarium, is here regarded as a secondary fungus not characteristic of the individual Weigelt collection of which the Balansia {Dothichloe epi-

chloe) is the chief feature. D. limitata was proposed by Diehl (1939) under the assumption that Saccardo's B. epichloe was acceptable for the Weigelt material, but examination of an additional Weigelt specimen since made available bears out Weese's description.

TAXONOMY AND HOST SELECTIVITY

The superficial similarity of Balansia epichloe to B. henningsiana readily ex- plains the general tendency in the past Figure 72.— Distribution of Balansia epichloe. to regard them as one species. In fact, NOMENCLATURE if the two were consistently present on It seems most likely that the epithet different host species their slight measur- "atramentaria" was first applied to this able differences might readily be ex- species because of a lapsus plumae or of plained as effects due to differing specific a printer's error in place of "atramen- hosts as habitats. The fact that both tosa." It was nevertheless applied as a species are present and distinguishable nomen nudum in reference to specimens on three different host species, Andro- of this species, Balansia epichloe. Be- pogon scoparius, Gymnopogon amht- cause of its superficial similarity to B. gtius, and Panicuni agrostoides, is con- henningsiana and because of the simi- trary to any such assumption. Further-

larity of names it was confused with more, in Virginia where the writer has that species. Regardless of the generic found one of these hosts, A. scoparius, allocation, these two species have been liberally infected with B. henningsiana, treated as synonyms by Saccardo (1883) none of the B. epichloe could be found, Atkinson (1894, 1897, 1905), Diehl although this as well as other potential (1930b), Miles (1935), and others. hosts were present. In some localities Fortunately, in the light of the confu- where the writer has noted B. epichloe sion with "atramentosa," "atramentaria" to be abundant in close proximity to seems never to have been validated by a hosts recorded for B. henningsiana, the description, although the specimen dis- latter was often not to be found after tributed in Ravenel's Fungi Americani thorough searching. BALANSIA AND THE BALANSIAE IN AMERICA 43

PATHOLOGICAL OBSERVATIONS hood of infection through the stigma and consequent seed transmission as reported Living infected specimens have been above for Bcilansia claviceps. kept under observation in greenhouse and outdoor culture on but three hosts: Attempts at securing the fungus in Cten'ium aromaticum from North Caro- culture in vitro were made without suc- five placing Hna in the greenhouse at Cambridge, cess on different occasions by Mass., in 1930 and 1931; Eragrostis re- aseptically bits of tissue from within fvdcta from Georgia in a greenhouse at living leaves on Ctenium containing my- Arhngton, Va., in 1931, and from south- celium of the fungus onto certain stan- ern Virginia outdoors in Arlington, in dard agar media: Thaxter's potato agar, 1932-34; Sporobolus poketn from North corn meal agar (Shear formula), prune Carolina in 1933 and 1934 in a green- agar, and a 2 percent agar medium living house at Arlington. These are all peren- using a decoction of Ctenium nial grasses and when kept in a vigorous leaves as the nutrient. On six occasions growing condition produced consistently similar unsuccessful attempts were made from one to three stromatic areas on to secure the fungus in culture on corn live tissue each leaf. meal agar, using infected from emerging leaves of Sporobolus When infected plants of Ctenium poiretii. were kept for 6 weeks in 1931 in a greenhouse where night temperatures Infected plants when cultivated were consistently near 7° C, these in- showed no less vigor or coloration than fected plants produced leaves free of healthy ones. On the spots where fructi- stromata, although certain areas on these fications were forming, however, there leaves were of a paler hue than the was frequently a slight chlorosis. Al- rest of the leaf surfaces. Critical exami- though mycelium of the Balansia could be nation disclosed the presence of my- detected elsewhere in the tissues of the celium typical of Balans/a v/ithin the same leaves, these other tissues showed tissues of these leaves. After these plants no discoloration whatever. Obviously the were removed for a month to a warmer parasite did not seem to impair normal greenhouse they again produced fructifi- vegetative vigor, the effect appeared cations typical of B. epichloe. No in- merely that of precluding development fected plants of Ctenium or of Sporobo- of any flowering shoots in Ctenium and lus have been observed to bear any in- in Sporobolus. The same effect was ob- florescence, although infected Eragros- served in the field on these hosts and tis refracta m Virginia and in Georgia less so on Eragrostis. The fact that in- have been found that bore some flower- fected plants are not observably different ing stems. in color from healthy ones, except for a Numerous unsuccessful attempts in very short tim.e in the areas of fungus 1930-31 were made to obtain infection fructifications, causes them to be over- of Ctenium by placing scolecoconidia looked except as a result of careful and ascospores upon uninfected potted search. Affected plants with no other plants kept for 3 days in a damp cham- symptoms than sterility easily escape de- ber. Furthermore, on three occasions tection. Except for the name "leaf dis- viable scolecoconidia were injected by ease," applied by Sprague (1946) to means of a hypodermic needle into the the effect on Calamagrostis inexpansa, no apices of unfolding leaves of four name appears to have been applied to healthy potted plants of Ctenium without this peculiar sterile condition of other any later evidence of infection. These grasses caused by the fungus. unsuccessful attempts at inoculations, 11. Balansia aristidae along with the observed evidences of systemic infection, suggest the likeli- Balansia aristidae (Atk.) comb. nov. 44 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Dothichloe aristidae Atk., Torrey On A. oligantha Michx.—Mississippi, 1890, S. M. Tracy (N, Mo). Bot. Club Bui. 21 : 224. 1894. On A. portoricensis Pilger—Puerto Rico, Ophiodothis aristidae Atk., ap. 1913, F. L. Stevens 6011 (N, F, W); Sacc. Hedwigia 35 (Kept. 7): 1916, H. H. Whetzel and E. W. Olive XXXIV. 1896. No. 695 (N). On A. purpurascens Poir.—Alabama, 1891, O. atramentosa Earle var. aristidae j B. M. Duggar (G. F. Atkinson No. j Earle, Torrey Bot. Club Bui. 27: 2145), type (A, F); 1897, F. S. Earle ^ 122. 1900. and C. F. Baker (N); Virginia, 1939, 1941, W. W. Diehl (W). Hypothallus effuse as a continuous On A. rejracta Griseb.—Dominican Repub- flat, thin, white to gray prosenchymatous lic, 1925, E. L. Ekman and R. Ciferri film early becoming black, on culms and No. 77 (W). On A. spiciformis Ell.-Florida, 1898, C. F. infrequently leaf sheaths, rarely over on Baker (W). 30/>i in thickness, usually encircling the On A. stricta Michx.—Georgia, 1933, George culm and covering a part of the adjacent Crisfield (W); Florida, 1936, E. West No. 11820 (N). adaxial leaf sheaths, rarely over 30/* in On A. tenuispica Hitchc—Florida, 1946, L. thickness, usually encircling the culm as J. Brass No. 15885 (W). a collar of variable length, rarely exceed- On A. vilfijoUa Henr.—Cuba, 1940, Bro. ing 20 mm. in length. Ephelidial fructi- Leon No. 17480 (W). On A. virgata Trin.—Georgia, George fications ephemeral, hysteriform to ef- 1933, Crisfield (F, W); Alabama, 1898, S. M. fuse, very small part covering a or most Tracy No. 5192 (N). of the surface of the hypothallus sepa- On Aristida sp.—Alabama, 1897, F. S. Earle rated by thin excipular fragments, hy- and C. F. Baker (I, Mo, W); 1892, S. M. Tracy (Mo); 1931, L. Seal menium as a palisade of conidia upon J. (F. W); Florida, 1891, R. Thaxter (F), short, simple conidiophores, usually not 1893, 1895, G. V. Nash No. 2225 (F, N, exceeding l/x in diameter, conidia short, Mo, W) . F. L. Scribner (N) ; 1895, P. H. 15/A to 20/i by 1.5/A to 2^, rarely con- Rolfs No. 44 (N),No. 16 (Fl); 1897, R. Thaxter (F); 1898, R. Thaxter (A); stricted. Ascostromata variable, effuse to 1932, G. B. Sartoris; 1935, E. West pulvinate over part or over the entire No. 10,304 (W); Mississippi, 1889, hypothallus (on culms, never noted on S. M. Tracy No. 197 (Mo) ; Cuba, 1907, leaves), usually broken up into pulvinuli C. F. Baker and H. Hasselbring No. 7224 (W); Venezuela, 1939, A. S. separated by slight or considerable areas Muller and H. H. Whetzel (W). of thinner sterile tissue; surface black, ILLUSTRATIONS prominently roughened to coUiculate by the prominent beaklike upper parts of Plate 7, E and F; Atkinson, Jour. Mycol. 11; plate 88, figures 28 and 29; Chardon, the perithecia; context gray to brown or Mycologia 13: plate 14, figure 13. black; subperithecial stroma very slightly TYPE LOCALITY developed; perithecia lageniform, 350/x to 400/x by 130/x to 200;*; perithecial Auburn, Ala., on Aristida purpurascens walls 15/x to 20/* thick; asci subcylindri- Poir. cal, l40ju, to 255/* by 5/* to 6/*; ascospores DISTRIBUTION one-half to three-fourths the length of On culms and leaves of Aristida spp., Vir- the ascus, 1/* to 2/* in diameter. ginia to Florida and Mississippi and southward through the West Indies to Venezuela SPECIMENS CITED and Brazil (fig. 13). On Aristida adscencionis L.—Venezuela, DISCUSSION 1939, C. E. Chardon No. 3896 (W). On A. dtchotoma Michx.—Mississippi, 1892, Balansia aristidae resembles B. S. M. Tracy, Ellis Herb. No. 1849 (N); strangulans in forming the collarlike Alabama, 1891, C. L. Newman (A). fructifiications on culms of the host, but On A. intermedia Scrib. & Ball (?) .—Florida, is readily distinguished by the prominent 1903, A. A. Eaton (W). ostiolar beaks, by the greater separation On A. longespica Poir.—Alabama, 1897, G. W. Carver No. 353 (N). and prominence of the individual peri- BALANSIA AND THE BALANSIAE IN AMERICA 45

as Euryachora, is sometimes confused with Balansia aristidae because of a superficial similarity, forming thin, black, sterile films on leaves and culms of species of Aristida in southern United States. It appears to lack the spore-forms characteristic of Balansia.

12. Balansia hemicrypta

Balansia hemicrypta sp. nov.

Dothichloe hemicrypta Diehl, in

litt.

Hypothallo strato tenui, albido, in floris praesertim in lemma hospitis, mox fugiente; fructu conidifero scolecosporo in hypothallo centro in lemma convoluto innato; conidiophoris ex hypothallo Cur- tis, simplicibus, usque 3^ in diam.; coni-

Figure 73.— Distribution of Balansia arisiidae. diis 18/A ad 40/a longis, l/>t ad 2/x latis; ascostromatibus paululo patefactis, pul- thecia, and by the more ephemeral ephel- vinatis, usque 0.5 mm. in diam., super- idial stage. These similarities have natur- ficie brunnea, granulata, contextu albido, ally involved confusion with other laxe prosenchymatico, peritheciis non species. Tracy and Earle (1895) re- maturis. ferred to this species on Aristida stricta Hypothallus as thin white hyphal in Mississippi as Hypocrella hypoxylon. a film in and upon the inner floral parts, On the other hand Stevens (1917), in chiefly within the lemma of the host reference to Chardon's as yet unpub- plant, soon evanescent. Ephelidial stage lished Dothkhloe suhnodosa on Ichnan- within the lemma, lining it and displac- thus in Puerto Rico, used the name D. ing the inner floral parts by a mass of aristidae, and Diehl (1930), in report- conidia; conidiophores from the hypo- ing the conidial fructifications of what is thallus layer, short, simple, not exceed- here recognized as B. strangulans, also ing 3;Lt in diameter; conidia 18/x to 40/x erroneously employed the name D. aristidae. by 1/x to 2/x. Ascostroma slightly developed as a blackish-brown stromatic papil- Originally described by Atkinson late protrusion at the base of the lemma (1894) from material on Aristida pur- not exceeding 0.5 mm. in diameter; sur- purascens from Alabama, it has been re- face granular, brown; context gray to ported from Puerto Rico by Stevens brown; perithecia not matured. (1917), Stevenson (1918), and Char- don (1920) on A. portoricensis and SPECIMENS CITED from San Marcos, Rio Bianco, Brazil, On Aristida cognata Trin. & Rupr.—Maturin, on A. folia seti by H. and P. Sydow Venezuela, 1940, A. Chase (W). (1916). The species has been reported On A. ]endleriana Steud.—New Mexico, 1937, also from Alabama by Earle (1901), Comm. M. W. Pladek (W). from Mississipi by Miles (1935), and On A. glauca (Nees) Walp.—Lampasas County, Tex., 1924, B. C. Tharp; from Florida by Earle (1900). 1930, B. C. Tharp and W. W. Diehl, The little-known but ubiquitous type (F, W). On Aristida sp.—Vic. Santa Rita do Paran- Dothidella aristidae (Schw.) Ell. & Ev., hyba, Goyaz, Brazil, 1930, Agnes Chase treated by Theiszen and Sydow (1915)' No. 12100, paratype (F, W). 46 AGRICULTURE :M0N0GRAPH 4, U. S. DEPT. OF AGRICULTUTIE

ILLUSTRATIONS lemma. In that short-grass prairie where this Plate 8, A to £; Diehl, Phytopathology fungus was found there were liter- 20: figures 1. 2. ally hundreds of acres containing a high

TYPE LOCALITY percentage of infected plants, and where diseased plants were counted it was evi- Lampasas County, Tex. dent that from 10 to 50 percent of the DISTRIBUTION stand was infected. In florets of Aristida spp., Texas, New The specific name Dothichloe hemi- and Goyaz in Brazil Mexico, Venezuela, crypta Diehl without a description was (fig. 9). reported by Weiss (1943) for the above- DISCUSSION cited Texas specimens, recording the Balansia hemkrypta might be re- pathological effect as "sterility disease." garded as a very depauperate form or 13. Balansia ambiens variety of B. arhtidae were its characters Balansia amhiens Moell., Phycom. u. not so consistently uniform in all the Ascomyceten,p. 204. 1901, specimens examined. It is noteworthy, emend. however, that none of its recorded hosts

is as yet known as a host for B. aristidae. Hypothallus as a linear gray to black

It is unfortunate that in no material prosenchymatous layer on the culm and have specimens of the ascogenous state inflorescence up to 1.5 mm. in width come to hand, all the specimens cited and nearly the length of the leaf sheath showing the well-developed ephelidial beneath which it develops. Ephelidial stage only. Sterile ascostromata were fructifications ephemeral, hysteriform, formed on infected Aristida glauca from exciple thin, conidia 18^. to 35/x by 1/a to Lampasas County, Tex., transplanted 1.5/x. Ascostromata pulvinate-capitate, in 1930 to an Arlington, Va., garden. subsessile in a single row upon the The host plants did not thrive in either hypothallus and emerging from between this garden or in greenhouse culture for the margins of the leaf sheath, becoming a long enough time even to permit black, up to 2 mm. in diameter; peri- maturity of the ascogenous fructifica- thecia sunken in the stroma, lageniform, tions. The immature ascostromata re- with thin walls; asci up to 225^ by 6/x sembled superficially those of B. gaduae. with rounded slime-cap and 4 or 8 spores The specimen on Aristida cognata, reaching 200^ by 1^. cited above, was found in a sample of SPECIMENS CITED seeds produced in a grass nursery at On Olyra sp.—Blumenau, Santa Catharina, Vaughan, N. Mex. Brazil, 1890-94 (?), A. M511er No. 972. In Lampasas County, Tex., where the ex type of Balansia ambiens (A, F). writer was enabled to see the effect of On Leersia grandiflora Prodoehl—Minas Geraes. Brazil, 1933, A. S. Muller No. this fungus in the field, the afl?ected 415 (W). host plants were easily distinguished by ILLUSTRATIONS their dwarfed condition in contrast to that of normal plants. As previously Plate 8, E and G: Moller Phycom. u. Ascom. plate 5, figure 66 a, b. noted (Diehl, 1930a) the eifect here, unlike that in most jB^/^;;,r/^- infected TYPE LOCALITY grasses, was dwarfing, including the pro- Blumenau, Santa Catharina, Brazil, on dixtion of dwarfed inflorescences into 0!)rJ sp. rlorets dwarfed and sterilized, but not DISTRIBUTION ^ oppression or abortion of inflorescences. Known only from the above-cited speci- The ephelidial stage was entirely hidden mens on Olyra sp. and Leersia grandijlora by the lemma and glumes, but in the from southern Brazil and as reported by Mollino (1930) on Olyra ciliafifolia Raddi cultivated specimens the immature ascos- in 1926 fiam Missiones in Argentina (fig. tromata were produced at the base of the 14). BALANSIA AND THE BALANSIAE IN AMERICA 47

Hypothallus inconspicuous, effuse, linear, limited to a yellow prosenchyma- t^ \l tous film becoming black with age. Ephelidial fructifications on definite

•> f points upon the hypothallus as sporo- \ 40 dochiumlike pulvinuli up to lOO/w in /tl/—v. 1 *^, ' ^ ^7^u diameter, made up of a mass of simple , A' to irregularly branched subgeniculate, / but not swollen, conidiophores not ex- y^ ^— 7 If ceeding 1/x in diameter and of variable '-._ \T / lengths, with conidia distinctly septate, "f ) / '"', linear to scythe-shaped, 60^ to 80/* by ( I ^ V " to early breaking at the septa. - 1.5/x — vS // 2^, ' i J linear-pulvinate, usually 1 Ascostromata 4 ri. y^ P long, to 10 mm. not known to exceed 30 mm. by 1 to 2 mm. in width and up

^ to in thickness, sessile, surface ^ i 500^ 1 black, smooth, very inconspicuously % punctulate from the minute ostioles, subperithecial stroma slightly developed; E•y 3^ 1 V r 1 perithecia rarely in one row, usually in two parallel rows, broadly ovate, 240ju Figure 14.— Distribution of Balansia ambiens to 390/x by 260/x to 335/*, ostioles short, {black squares) and 6. linearis (black dots). perithecial walls l4/x to 12^ thick; asci DISCUSSION cylindrical, 200/x to 300/x by 5/a to d^K with prominent slime cap diam.) Balansia ambiens simulates a sessile (7/x and with 4 to 8 spores; ascospores nearly B. redundans, but differs in the presence as long as the ascus, up to thick, of the ephelidial fructifications as well 2^ readily breaking into segments to as in the lack of a greenish hypothallus. 20/>i in length. MoUer (1901) implies that the infection 27/x is systemic. Since he points out that asco- SPECIMENS CITED spores in fresh condition are immediately On Olyra micrantha H. B. K.—Serra de germinable, i.e., at the normal time of Paraceima, Rio Branco, Amazonas, Bra- zil, In floret production, it is reasonable to as- 1909, E. Ule No. 3339 (W): Herb. Theiszen and in Herb. Hohnel sume that infection takes place at the (F). time of flowering through the stigma; On Chusquea sp.—Brazil, E. Ule ex type (?) therefore, resulting in a seed-borne infec- of Ophiodothis linearis in Theiszen tion. Herb. (F); vicinity of Rio de Janeiro, 1925, A. Chase No. 8496 (W). 14. Balansia linearis On Pariana sp.—Para, Brazil, 1934, J. R. Balansia linearis (Rehm) comb. nov. Swallen No. 3229 (W). On Gramineae.—Brazil, in Theiszen Herb. Ophiodothis linearis Rehm ap. as Ophiodothis rhaphidospora Rehm Pazschke, Hedwigia 35: 54. (F), possibly part of type. 1896. ILLUSTRATIONS

O. rhaphidospora Rehm, Hedwigia Plate 9; Moller, Phycom. u. Ascomyceten, 36: 380. 1897. plate 5, figure 69, a to d, Linearistroma lineare (Rehm) TYPE LOCALITY Hohn., Akad. der Wiss. Wien, Campo Bello, Rio de Janeiro, Brazil on Math.-Nat. Kl. Sitzber. (Abt. 1) Chusquea sp. 119: 938-939. 1910. DISTRIBUTION Dothichloe linearis (Rehm) Diehl, Brazil, on culms and abaxial surfaces of in litt. leaves of Gramineae (fig. 14). 48 AGRICLLTLRE MO:nOGR.\PH 4. U. S. DEFf. OF AGRICULTURE

DISCUSSION consistent, and the additional features Since the ephelidial conidia and the listed by him are found in other species conidiophores as well as the prominent of Balansia. ascus slime-cap of Bdansia linearis are Moller's (1901) account took special somewhat unlike those of the other cognizance of the peculiar spiral inroU- species discussed here, it is possible that ing of the leaves of the host and of the their differences, chiefly unknown to Von "hyphengeflecht" or hypothallus within Hohnel, may justify the recognition of the inroHed leaf. He recorded the lack his genus Linearistroma of which this of destruction of the leaf tissue and of species is the type. In other respects the the production of a pseudomorph com- species is referable to Balansia. parable to those of Balansia and Epi- The original description of the species cbloe. The peculiar inroUing of the as Ophiodothis linearis was credited to terminal leaf without its being otherwise Rehm in Pazschke's (1896, p. 34) ac- reduced in length is particularly striking count of one of Ule's collections of in the above-cited specimen, as Pariana Brazilian fungi with the host noted sp., Swallen No. 3229 (pi. 9, A). In simply as "Chusquea." It is likely that Mrs. Chase's specimen, No. 8496-^", on a part of this type specimen is the one Cbusquea, on the other hand, there is a on Chusquea cited above from Theiszen's m.arked brooming of the terminus of Herbarium at the Farlow Herbarium, the shoot with a decided dwarfing of where it lacks special designation as a the leaves (pi. 9, C) . In both these type. A year after Pazschke's account, instances it is noteworthy that this tera- Rehm (189", p. 380) again described tological effect is to be noted only on the species under the nevs' name 0. the tips of the shoots. rhapbidospora, based on material listed Ule's Brazilian specimen No. 3339 on as "Gramineae." He noted the presence 0'.)ra micrantba has been reported by of filiform paraphyses, feat^jres not since H, and P. Sydow (1916) as Lineari- obser^-ed. stroma.

Moller's Brazilian specimens listed as Doubtful and Excluded Species of on Olyra (presumably 0. micrantha) Balansia and on JMicrostachys speciosa Spr. (error for Merostachys) were reported bv both l.B. bypoxylon (Pk.) Atk. .cf. p. 49- Moller (1901) and Hennings (1902) 2. B. gaduae Rehxm. cf. p. 55. as 0. rhapbidospora. It remained for 3. B. clavula Lloyd cf. p. 56.

Von Hohnel (1910), however, to recog- 4. B. asclerotiaca P. Henn. . .cf. p. 58. nize that the t^'o species were identical. 5. B. diademaModl cf. p. 60. Using this species as the type he pro- 6. B. pallida var. orthocladae posed the genus Linearistroma because of P. Henn., type of Bal- the linear stromata, white hypothallus, ansielld cf. p. 60. and tv.-0-rowed arrangement of the peri- 7. B. cbusqueicola P. Henn. .cf. p. 61. thecia. This latter feature is actually not 8. Epbelis japonica P. Henn. cf. p. 61. Tho Genus A^kinsonella Atkhisonella gen. nov. Type species— Hypothallus prosenchymatous-sclero- Epicbloe hypoxylon Pk. tic to pseudoparenchymatous, context HypothaUis et ascostromatibus ut in VN-hite or yellow to black, surrounding genere Balansia, a Balansia propter fruc- or covering tissues of host plant. Typho- tificationes typhodios et ephelidios dis- dial fructification covering the surface tinguitur: fructificationibus t}-phodii ut of the hypothallus as a palisade layer of in genere Epicbloe, fructificationibus a inflated, apically aristate phialides bear- Balansia propter excipula phialidibus ty- ing dry amerospores as in Epicbloe. phodiis vestita differentibus. Ephelidial fructifications erumpent from BALANSIA AND THE BALANSIAE IN AMERICA 49 the hypothallus, discoid-cupulate to hy- (Ephelis borealis Ell. & Ev.) solitary, steriform, exciple (or peridium) hirsute few, erumpent from beneath the primary from the remnants of the typhodial layer. conidial fructification, sessile, at first Ascostromata as in Balansia. subspherical hysteriform, becoming cupulate or patellate, subimmersed; exciple 1 . Atkinsonella hypoxylon covered with amerosporous conidio- Atkinsonella hypoxylon (Pk.) comb. phores often elongate and hairlike but nov. nonseptate; context fleshy, pseudoparen- Epkhloe hypoxylon Pk., N. Y. chymatous, pallid; hymenium pallid- State Mus. Rpt. 27: 108. carnose to gray; conidiophores when 1875. simple becoming much elongate and Hypocrella hypoxylon (Pk.) Sacc, swollen at the base with tip l^w, to 1.5ju

Sylloge Fungorum 2 : 581. in diameter, sometimes verticillately to 1883. irregularly branched; conidia 25/jt to 38ju "^Ephelh borealis Ell. & Ev., Jour. by I IX to 1.5/A and when germinating Mycol. 1: 86. 1885. swollen to 2fx. Ascostromata few, sepa- Hypocrea hypoxylon (Pk.) Eli. & rate, erumpent from beneath the typho- Ev., Jour. Mycol. 2: 67. 1886, dial fructifications, sessile to subsessile, and North Amer. Fungi No. often separated from the hypothallus by 2373. 1890. Name only, error a constriction, hemispherical to flattened, in ident., specimen is Balansia pulvinate, surface black, 1 by 4 mm. henningsiana. broad by 1 to 2 mm. high, surface evenly Dothkhloe hypoxylon (Pk.) Atk., granular-punctate to collicular from the Torrey Bot Club Bui. 21: 223. ostioles; context pallid; subperithecial 1894. Name only, error in ident., stroma robust; perithecia lageniform, description is of Balansia hen- crowded but not compressed, 180/* to ningsiana. 420^ by 100/x to 200;*; asci subcylindri- Ophiodothis hypoxylon (Pk.) Sacc. cal, 120/x to 305;t4 by 4^ to 7/*, with 8 ap. Atk,, Cornell Univ. (Science) spores; ascospores 100/x to 210/x by l/* to Bui. 3: 10. 1897. Name only, 1.5/x. error in ident., description is of Balansia henningsiana. SPECIMENS CITED12 Balansia hypoxylon (Pk.) Atk., Quebec.-1938, I. H. Crowell (W).

Jour. Mycol. 11: 254-255. Nova Scotia.— 1909, J. Dearness in Bartho- 1905. lomew's F. Col. No. 3027 (A, F, W); 1909, J. Dearness (N, Pk). Hypothallus variable in dimensions Maine.~1885, F. L. Scribner (N, Pk, W); dependent chiefly upon character of host, 1897, R. Thaxter (F), Herb. Lloyd No. 6139 (W), Reliquiae Farlowianae No. 611 (F, usually 4 to 15 by 1.5 to 2 mm., con- W); 1907, R. Thaxter (A); 1931, R. Thaxter sistently enclosing culm and leaf base (W); 1915, J. G. Parlin (A). with undeveloped inflorescence of the New Hampshire.— 1904, 1907, P. Wilson host; surface and context at first gray (F, W); herb. C. G. Lloyd No. 6l40 (W), as on Pestuca ovina L. 1904 (F); 1922, then white to purple gray, waxy when D. H. Linder (F); 1917, P. Wilson (Herb. moist, compactly prosenchymatous; sur- Brooklyn Bot. Garden). face covered by typhodial fructification. Massachusetts.— 1877, B. D. Halsted (F, Primary (typhodial) conidial fructifica- Mo); 1883, A. B. Seymour (F); 1909, G. E. Morris (F, N, Pk); 1908, A. P. D. Piguet tion glistening white to gray, of one layer (palisade) of swollen or spatulate simple ^^ In these cases the evidence either of the labels or of the specimens themselves compels recognition conidiophores (phialides) , each with an of the hosts as species of Danthonia, most likely either D, spicata (L.) Beauv. or D. compressa Aust. apical projection (sterigma) bearing a It is suggested here that the host of the specimen single conidium; conidia simple, hyaline, from New Hampshire in the Farlow Herbarium noted as Festuca is very likely a species of Danthonia also. obovate, 3/x to 8/x by 3;u, to A^, early The condition of most host material in fungus col- on labels unless there evanescent. Ephelidial lections compels chief reliance fructifications are satisfactory indications to the contrary. 50 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

(F); \V. G. Farlow in Ellis Herb. No. 383 ILLUSTRATIONS (N); 1909, [N. L.] Britton (W); 1925, Plate 10, to G; Atkinson, jDur. 1941, D. H. Linder (F, W); 1930, 1931, A Mycol. 11: plate 81, figures 4 to plates W. W. Diehl (F, W); 1931, G. D. Darker 6; 82, 86, 87 (figs. 17, 18); Schwarze, N. J. Agr. Expt. (W); 1932, J. D. McLachlan (W); 1934, Sta. Bui. figures to Diehl, A. G. Kevorkian (W); 1938, L. E. and G. D. 313, 149 152; Jour. Darker No. 6543 (W). Agr. Res. 41: plate 2, A, B, figure 3, A. Connecticut.— 1889, R. Thaxter (A); TYPE LOCALITY 1900, J. L. Sheldon (N); 1901, J. L. Sheldon (Pk); 1930, R. P. Marshall (W). Sandlake, N. Y. New York—1873, C. H. Peck t}'pe (Pk); DISTRIBUTION 1885, [G. A.} Rex (N): 1891, \T. R. Dudlev (W). On aborted inflorescences of Danthonia compressa Austin and D. spicata (L). Beauv. Pennsylvania.— 1929, C. S. Parker No. Quebec and Nova Scotia to Iowa and Geor- 1283 (W); 1929, L. O. Overholts (Herb. gia; also on unknown host in Brazil (fig. L. O. O.); [no date], Wm. Herbst (Pk). 15). New Jersey.— 1896, J. B. Ellis in North Amer. Fungi No. 3467 and Fungi col. No. ERRORS IN IDENTIFICATION AND 1092 (F, N, W, Mo); 1906, P. L. Ricker QUESTIONABLE RECORDS No. 3507 (W); 1947, W. W. Diehl (W). Farlow's (1878) record of Atkinson- Delaware.— 1929, C. Drechsler (W). ella hypoxylon, possibly the first after Maryland.-1900, C. L. Shear (W); 1905, Peck's, is C. F. Wheeler (W); 1912, H. S. Barber listed as ''Dothidea torax B. & (W); 1929, 1930, 1932, 1939, 1944, 1945, C. on stems of grasses. Bussey "Woods. 1946, 1947, 1948, W. W. Diehl (F, W); Rare." In the succeeding remarks (p. 1933, E. Benedict, Jr. (W); 1944, L. J. 249-250), however, he identified the Shanor and W. B. Allington (W); 1935. A. G. Johnson (W). specimen as found by Halsted and calls Virginia.-1918, 1921, 1929, 1933, H. A. attention to Peck's name, which he con- Allard (W); 1929, 1930, 1931, 1935, 1947, sidered along with D. rorax Berk. & 1948, 1949, W. W. Diehl (F, W). Curt., D. p/lulaeformis, and D. atramen- Washington, D. C— 1886, H. Pergandy (W). taria to be referable to Sphaeria rorax (Berk. Curt. ) Farlow West Virginia.-1928, W. A. Archer No. & as a new combi- 2060 (W); 1941, H. A. Allard Nos. 9034, nation. The Halsted specimen now in the 9035, 9053, 9211, 9212, 9230 (W); 1947, Farlow Herbarium, as cited in the present H. A. Allard No. 12,283; 1940, 1946, 1949, W. W. Diehl (W).

North Carolina.— 1933, J. H. Miller and W. W. Diehl (W); 1933, L. O. Overholts (Herb. L. O. O. as examined by J. A. Stevenson); 1947, J. A. Stevenson (W), W. W. Diehl (W). Kentuck}'.-1949, W. W. Diehl (W). Tennessee.-1934, L. R. Hesler No. 3533 (W, N); 1939, R. W. Davidson (W); 1934,

J. H. Miller No. 3533 (W). Georgia.— 1940, G. E. Thompson and J. H. Miller (W); 1946, J. H. Miller (W); 1947, W. W. Diehl (W). Ohio.-1901, W. A. Kellerman (A, F, Brooklyn Bot. Garden); 1946. G. D. Darker (W); 1948, 1949, W. W. Diehl (W). Indiana.— 1949, C. C. Deam (W); 1949, W. W. Diehl (W). Illinois.— 1947, V. H. Chase No. 8942 (F, W). Iowa.-1903, R. E. Buchanan (N). Brazil (Serra Geral).-E. Ule 1329 (A) on unknown host. (The Saccardo Herbarium at Padua, ac- Figure 75.— Distribution of Atkinsonella hypoxy- cording to Gola (1930), contains but one lon (black dots) and A. hypoxylon var. iexensis specimen of this species: "t}'pe.") (black squares). .

BALANSIA AND THE BALANSIAE IN AMERICA 51 list, is typical of the species. Of late Atkinson had obtained this from Sac- years the fungus is far from rare near cardo in a personal communication. the old Bussey habitation in the Arnold Stevenson's (1918) report of Balan- Arboretum in Massachusetts. sia hypoxylon from Puerto Rico on Pani- Included with certain specimens re- cum tricanihuni referred to specimens ported by Ellis and Everhart (1886) as later distinguished by Chardon (1921) "Hypocrea hypoxylon Pk.," there was as his Dothichloe subnodosa, although it one on ^'Pankum (capHlare?) surround- was reiterated equivocally later as B. hy- ing the culm just below the internodes." poxylon by Seaver and Chardon (1926) This is doubtless the 1879 Ellis speci- At the same time they reported Ephelis men (W) of Balansia strangulans from wexicana, host not noted, as a conidial New Jersey cited in the present account stage for B. hypoxylon, most likely hav- as on Panicum sp. ing in mind, but not citing at that point, The Brazilian specimen, E. Ule No. the record of E. mexicana on "Grami- 1329, cited above, is represented in the neae" from Puerto Rico by Bresadola, Atkinson Herbarium as a fragment from Hennings, and Magnus (1893), who the Berlin Herbarium. It was recorded had, however, accepted Rehm's (1897) by Rehm (1897) as Ophiodothis vorax. previous identification of it as a stage of Although a single piece, hypothallus and B. claviceps. a few ascostromata, it appears to be Ellis and Gerard (1889) reported Atkinsonella hypoxylon. As Hypocrella Epichloe hypoxylon "on leaves of Pani- hypoxylon it has, however, been reported cum" from New Jersey. According to the from Brazil by Viegas (1944, p. 94), present interpretation, this cannot be with reference to a different fungus Peck's species but would seem to be either growing on an insect on oleander. This of the two species of Balansia that fruit is evidently a mistaken identification on leaf surfaces. The writer has not, how- since, like all other known species of ever, found any such specimen of pre- the genus, it is strictly phytophilous. vious date among the Ellis specimens Balansia hypoxylon has been reported anywhere and suspects that the record by Dearness and House (1925, p. 69) in question refers to an Ellis specimen on leaves of Spartina pectinata. This dated 1879, which is B. strangulans seems to be the sole report of any species (q. v.). of the Balansiae on this host and as it Ellis and Everhart's (1892, p. 91) is said to be on leaves, not on culms or treatment of the species included practi- inflorescences, it cannot be the species cally all the Balansiae in the United reported. States, no one of which was distinguish- Atkinson's (1894, 1897) early use able. Tracy and Earle (1895), in re- of the specific epithet "hypoxylon" ap- porting it from Mississippi as on species plied not to this species but to the one of Andropogon, Panicum, Aristida, and recognized in the present discussion as Uniola, were actually concerned with B. henningsiana, for which Atkinson other species (q. v.) on these hosts. (1905), in an attempted correction of Saccardo (1897, 1898) compiled these his earlier usage, erroneously employed early interpretations as a permanent the name Dothichloe atramentosa. His source of confusion. inclusion in synonymy of ''Ophiodothis The question of host identity in the hypoxylon (Pk.) Sacc. 1897" has pre- various records of this species would be sented a problem, inasmuch as it has not especially difficult if the evidence of been possible, to date at least, to find more recent field observations did not any publication by Saccardo in 1897, or indicate that the host plants are species in any other year for that matter, in of Danthonia, either D. spicata or D. which this combination appears. The as- compressa. This clarifies the host records sumption is, therefore, warranted that as "grass" noted by various observers: 52 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Trelease (1887), locality not stated, Herbarium, and Maneval (1937) has Harvey (1896) and Ricker (1902) for reported it on the same host from one Maine, Pammel and coworkers (1901) gathering in Missouri. Diehl's (1930b) for Iowa, and Schwarze (1917) for New discussion was concerned chiefly with Jersey. Jones (1898), in reporting the gatherings in Maryland and Virginia. fungus on "Poa" from Ohio, doubtless Miller (1941) has reported it from failed to recognize that the host was Georgia growing upon D. sericea Nutt., Danthoma growing along with unin- but from the writer's examination of the fected Voa. Weber's report of B. hypoxy- Thompson and Miller specimen cited lon on Cenchrus from Florida refers to above, he is inclined to regard the hoSc the species here recognized as B. ohtecta as D. spicata. Hardison (1945) has re- (q. v.). corded it on D. spicata from Kentucky.

It was reported from Wisconsin by The species, including the variety Davis (1914, p. 861) on Calamagrostis texensis, on the basis of printed records canadensis, but this is doubtless a mis- and of specimens examined is thus far identification for Balansia epicbloe in the known only from the United States east light of a 1912 specimen of that species of the Great Plains, from northeastern (cf.) in the New York Botanical Gar- Canada, and possibly from the one place Davis. this last instance, den gathered by J. J. in Brazil. Except for Atkinson's (1905, p. 255) question- the only host species thus far recognizing inclusion of Ephelis mexkana from able are Danthonia spicata, D. compres-

Mexico is in reference to the type speci- sa, Stipa leucotricha, S. riridula, and, m.en of that species, which is here re- questionably, Elyonurus barbiculmis, the ferred to Balansia ohtecta (cf.). last three as hosts for var. texensis. Criti- The Herbst specimen cited above from cal search can be expected to enlarge the the Peck Herbarium has special interest, list of host species and extend the present limited area where the fungus has been because it may be part of the material found. reported by Herbst (1899 p. 197) as on nodes of Eriophorum virginicum. PATHOLOGICAL CONSIDERATIONS From his notation as to nodes, it sug- The writer's observations of Atkinson- gests, however, another species, B. stran- ella hypoxylon in living condition have gulans. The writer has not been able to been limited to comparative studies of find any such specimen in the Herbst diseased and healthy plants of Danthonia Herbarium at Philadelphia. In any case spicata and D. compressa in the field and there is grave doubt as to the identity in greenhouse culture, including attempts of the host plant. at inoculation of healthy plants. Infected ADDITIONAL RECORDS WITH REFERENCE and healthy clumps of these grasses have TO RANGE always grown readily after being transplanted to suitable soil in garden and Seaver's monographs (1910 a, b, results 1911) noted the species as on Danthoma greenhouse. But most striking transplanting spicata "and other grasses" from Maine have been obtained by these grasses during the late to South Carolina and Iowa. This is the clumps of in the greenhouse. Such only report of it from South Carolina. winter to pots transplantings of D. spicata and D. com- Schwarze (1917) noted it "on the although most grasses" in New Jersey. Oilman and pressa behave identically, were Archer (1929) followed by Fitzgerald of the observations here recorded (1949) have recorded a specimen on concerned with D. spicata. D. spicata obtained in Iowa in 1909 by The phenology of the fruiting fungus,

R. E. Buchanan, which is of later date outdoors and in greenhouse transplants, than the 1903 Iowa specimen cited above is synchronous with that of the inflores- from the New York Botanical Garden cence on a healthy host plant, develop- BALANSIA AND THE BALANSIAE IN AMERICA 53 jng the typhodial stage slightly before been ineffective, except in two doubt- anthesis, the ephelidial stage at anthesis, fully successful instances that followed and the matured ascostromata well after placing germinable ephelis conidia on the time of seed maturity on healthy stigmas. Germinating conidia placed on plants. At Cambridge, Mass., in 1932 leaves, culms, and roots and injected transplants from field to greenhouse on within nodes and internodes failed to February 22 reached anthesis by April produce infection. It would seem likely 15, with coincident development of the that this species follows the pattern of fungus on infected plants, while those Bdansia obtecta with stigma infection by left on the field reached the same stage ephelidial conidia, or more rarely by by mid-June. overwintered ascospores, resulting in In the region of Washington, D. C, seeds bearing a perennial mycelium the purplish-gray hypothallus is usually transmissible on germination to the new apparent from May 15 to 20, then soon host plants that perennially bear the char- covered by the pearly- white typhodial acteristic fructifications of the parasite. layer or amerosporous fructification The diseased host shows none of the (Diehl 1930), and by May 30 bearing usual evidences of disease such as chloro- ephelidial fructifications and a few mi- sis or death of tissues. The culms and nute warts that mark the initials of ascos- leaves of diseased plants show no appre- tromata. Although these ascostromata are ciable difference in color from healthy mature by July 4, the asci and ascospores plants, merely being sterilized and often remain intact within the perithecia dwarfed by the fungus. As a result in- for a month or more, the spores being fected plants, being undersized and of ejected intermittently; by September the normal coloration, usually escape the at- perithecia are usually empty. Neverthe- tention of collectors, as well as of plant less, in Maryland in 1946 and 1948, the pathologists who may traverse an area writer has found some overwintered as- where as many as 50 percent of the costromata still full of asci with germin- Danthonia plants are infected without able ascospores as late as June 1 of the detecting them. year after their formation. In New Eng- Although inconspicuous, the effect on land healthy Danthonia spicata normally the host is striking. The parasite appears develops some inflorescences intermit- to have choked the developing culm and tently during the summ.er, and coinci- inflorescence, hence simulating slightly dentally plants infected by Atkinsonella the effect of Epichloe typhina that causes also produce some fungus-bearing shoots the "choke" disease of various other throughout the season. grasses. In this latter case, however, culm

The mycelium of this species is inter- elongation is not so drastically inhibited. cellular and perennial in the tissues of The name "choke" in reference to the the host, and similar mycelium has been disease caused by Epichloe is even more detected in seeds of seemingly healthy fitting for the e^cct produced by Atkin- plants growing adjacent to infected sonella. The name "purple choke" has plants. When diseased plants were trans- been applied to this disease in Kentucky planted uniformly to warm greenhouse by Hardison (1945), because of the culture, they sent up shoots bearing the early purple coloration of the hypothal- fungus, whereas, labeled, similar in- lus, and "black choke" by Weiss (1945), fected clumps outdoors often sent up doubtless because of the coloration of the some seemingly healthy shoots. This last ascostromata. When the hypothalli are was especially noticeable in plants in the first apparent the culms will have reached vicinity of Cambridge, Mass., although a height of from 4 to 6 cm., rarely 15 rarely in the region of Washington, cm., with culms of healthy plants only D. C. slightly if at all higher. From that time Inoculation experiments have thus far onward the healthy culms continue to 54 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE elongate, reaching a normal height of ILLUSTRATIONS up to 50 to 70 cm., while the infected Atkinson, Jour. Mycol. 11: plate 81, figures culms develop no further and remain 1 to 3, and plate 83; Heald and Wolf, Bur. dwarfed. The fruiting hypothallus en- Plant Indus. Bui. 226: plate 11, figure 1. circles and mummifies the apex of a culm TYPE LOCALITY and two or three emerging leaves and the health)'-appearing, nascent inflorescence; San Antonio, Tex. it has acLually caused a sudden stoppage DISTRIBUTION of growth of the host before the inflorescence has expanded. Normally with this Texas, on aborted inflorescences of species of Stipa and possibly other Gramineae (fig. dwarfing there is also a failure to pro- 15). duce any cleistogenes. DISCUSSION 2. Atkinsonella hypoxylon var. texensis Under the name Balans'ia hypoxylon Atk'msonella hypoxylon var. texensis this variety was reported by Heald and var. nov. Wolf (1912) as causing the disease A specie dififert in hypothallibus et as- there named "Balansia blight," which costromatibus et fructibus ephelidibus destroyed the inflorescence of Stipa leu- robustioribus. cotricha. It appears not to have been This differs from the species chiefly accorded other attention in the literature, in its greater size with hypothalli of from unless Weiss' (1945) Check List inclu- 100 to 200 by 3 to 6 mm. and ascostro- sion of B. hypoxylon on Stipa pulchra in mata reaching a diameter of 4 mm., with California is concerned with it. Unfor- larger ephelidial fructifications but with tunately, neither Dr. Weiss nor the all spore forms as in the species. writer has been able thus far to substan- SPECIMENS CITED tiate this by any written report, printed On Stipa leucotrtcha Trin. & Rupr.- San account, or specimen record. They are, Antonio, Tex., May 17 1941, H. therefore, forced to conclude that the Johnson, type (W, F). above was an error in transcription and On S. viridula Trin.— (Possibly error for in S. leucotrkha) Greenville, Tex., May that B. hypoxylon has not been found 19, 1913, D. A. Saunders (W); June 2, California thus far, nor on S. pulchra 1913, C. G. Marshall and D. A. Saunders anywhere. The specimen in the Farlow (W). Herbarium with the host labeled '^ Elyon- On Stipa sp.-Marble Falls, Tex., 1905, H. F. Fuchs (W). urus barbiculmis" also provides a very On Gramineae (? Elyonurus barbiculmh doubtful host record; but the material Hack.).—Bracken, Tex., 1903, B. H. contains so little of diagnostic host ma- Groth No. 238 (F). terial that the label cannot readily be On Gramineae.—Austin, Tex., 1901, W. H. Long, Jr., Nos. 756, 774 (A). contradicted.

The Genus Balansiopsis

Balansiopsis Hohn., Akad. der Wiss. Hypothallus scant, prosenchymatous, Wien. Math.-Nat. Kl. Sitzber. ephemeral, on tissue of undeveloped in- 119 (Abt. 1): 936. 1910 florescences or forming a thin layer on (emend.). Type species: Ophio- limited areas of the host. Conidial fruc- dothis gaduae Rehm. tifications lacking. Ascostromata as in Balansia. BALANSIA AND THE BALANSIAE IN AMERICA 55

THE SPECIES OF BALANSIOPSIS KEY TO THE SPECIES a. Ascostromata sessile, pulvinate-hemispherical, each arising from a minute hypothallus at a node of the host 1. B. gaduae. aa. Ascostromata substipitate to stipitate arising irregularly from various points on culm, foliage or aborted inflorescence of host. b. Ascostromata stipitate or substipitate, arising from either terminal leaf or aborted inflorescence within terminal leaf 2. B. pilulaejormis. bh. Ascostromata stipitate arising from culm and aborted inflorescence. .3. 5. asclerotiaca.

1. Balansiopsis gaduae On Guadua sp.—Brazil, E. Ule in Herb. F. Theiszen as Balansiopsis guaduae Bala?7swpsn gaduae (Rehm) Hohn., (Rehm) Hoehn. (F); Porto Bello, Bra- Akad. der Wiss. Wien., Math.- zil, 1894, E. Ule No'. 2106, type of Ophiodothis gaduae in Elerb. Hoehn. Nat. Kl. Sitzber. 119 (Abt. 1): (F); Amazonas, Brazil, 1901, E. Ule, 936. 1910. Mycotheca Bras. No. 79 (F, N, W).

Ophiodothis gaduae Rehm ap. ILLUSTRATIONS Pazschke Hedwigia 35: 53. Plate 10, li\ Moller, Phyc. u. Ascomyceten, 1896. figures 69, a to d; Viegas, Bragantia 4: plate Balansia regtdaris Moell., Phycom. 115. u. Ascomyceten, p. 303. 1901. TYPE LOCALITY Dothichloe gaduae (Rehm) Diehl, Campo Bello, Rio de Janeiro, Brazil, on in htt. Guadua sp.

Hypothallus as a thin prosenchymat- DISTRIBUTION ous film not exceeding 3 mm. in diam- Brazil, on culms of Bambusae (fig. 16). eter at the nodes of the host on the culm opposite the leaf bases, early covered by the ascostroma. Ascostromata sessile, pulvinate-hemispherical, not exceeding 2 mm. in horizontal diameter and 1 mm. in vertical diameter; context whitish, surface black, finely granular from the slightly emergent ostioles; subperithecial stroma robust, pseudoparenchymatous, perithecia not crowded—compressed, lageniform 350^ to 400/^ by 180/a to 255^; perithecial walls 10/x to 20^ thick; asci 105/x to 300^ by 4^ to 6/a with prominent subsphaerical slime-cap and from 3 to 8 spores; ascospores 96^ to

210/xby 1.5/Ato 2fx.

SPECIMENS CITED

On Bambusa arundinacea Willd.—Estado Rio de Janeiro, Brazil, 1930, A. Chase No. 12173 (F, W). On Chusquea unijlora Steud.—Chile, 1888, R. A. Philippi, ex U. S. Natl. Herb. No. 821529 (W); Bolivia, 1923, A. S. Hitch- cock, ex U. S. Natl., Herb. No. 1169677

(W). Figure 7 6.— Distribution of Balansiopsis gaduae On Guadua tagoara (Nees) Kunth—Santa (black squares), B. pilulaeformis (black dots), Catharina, Brazil (1890-93), A. Moller B. asclerotiaca (circle), and Ephelis japonica No. 830, type of B. regularh (A). (cross). 56 AGRICULTURE MONOGRAPH 4. U. S. DEPT. OF AGRICULTURE

DISCUSSION in Sacc, Sylloge Fungorum 2: 652. 1883. Balansiopsis gadiiae appears to have Hypocrella hypoxylof? (Pk.) Sacc. been gathered only rarely and to have re- var. pilulaeformis Berk. & Curt. ceived little notice except for the records in Ell. and Ev., North Amer. of Pazschke (1896), MoUer (1901), Pyrenomycetes, 91. 1892. Hennings (1904b), Von Hohnel p. Ophiodothis atramentosa Earle var. (1910), and Viegas (1944a), the latter pilulaeformis Earle, Torrev Bot. from Sao Paulo. Von Hohnel recognized Club Bui. 27: 121-122. 'l900. that Moller's Balansia regularis was the Balansia redundans Moell., Phycom. same species as Ophiodothis gaduae u. Ascomyceten, p. 204. 1901. Rehm. MoUer and Viegas pointed out (?) B. claiula (Berk. & Curt.) that the fungus produced a striking Lloyd, Mycol. Notes No. 71: witches'-broom effect on the host. As 1259. 1924. shown in plate H, this brooming is 10, B. pilulaeformis (Berk. & Curt.) characterized by an angular, twisted ef- Miller, Plant Diehl ap. J. H. Dis. fect. The regular orientation of the stro- Rptr. Sup. 131: 55. 1941. mata upon the nodes opposite each leaf is a distinctive feature. Moller noted but Hypothallus variable as an irregular, linear effuse cover- three or four spores to the ascus, but as to subfusiform, mass Viegas has pointed out the normal num- ing the apex of the culm or part of the inner surface of an infolded leaf sheath ber is eight. or blade of the host, 10 to 30 by 2 to 4 The original spelling of the epithet mm. in linear and horizontal diameter: as '' gaduae," possibly an editorial or surface and context greenish yellow to printer's error, was consistent at the greenish black. Ascostromata subpul- same time with the spelling of the host vinate to subspherical, capitate, often name as "Gadua" in place of the stan- somewhat flattened-pulvinate, subsessile dard "Guadua." Saccardo re- (1898) to stipitate, scattered to aggregate upon peated Rehm's spellings, but Hohnel the surface of the hypothallus; context

has transcribed '^ Gaduae' \o (1910) yellow to yellow-green; subperithecial ^^Guaduae" in line with his corrected stroma robust; surface black, punctulate- spelling of the name of the host. roughened from the ostioles; perithecia not usually crowded or compressed, la- 2. Balansiopsis pilulaeformis geniform, ostioles very slightly emergent; Balansiopsis pilulaeformis (Berk. & perithecia 225^ to 250^ by 60^ to 130/x; Curt.) comb. nov. perithecial walls 12^ to 15/x thick; asci

subcylindrical 105 p. to 300^ by Ajx to Dothidea pituJaeformis'^^ Berk. & 6^; ascospores approaching the ascus in Curt. ap. Berk., Nom. nud. length and 1^ to 2^ in diameter. Hooker's Jour. Bot. 6: 227. 1854. SPECIMENS CITED

(?) Xylaria clavulus Berk. & Curt. On Bambuseae—hid.zi\. 1897, F. Noack in ap. Berk., Grevillea 4: 47-48. Herb. Theiszen (F). 1876. On Paspalutn ciliatijolium Michx.—Florida, 1906, R. Y. Winters (Fl); 1934, G. F. Dothidea pilulaeformis Berk. & Weber and E. West No. 9716 (Fl, W); Curt., Grevillea 4: 105. 1876. 1935, G. F. Weber (N); 1935, E. West (N). Ophiodothis vorax Berk. & Curt. On P. pub escens Muhl.—Georgia, 1933, var. pilulaeformis Berk. & Curt. George Crisfield (W).

'^ The spelling "pitulaeformis," as pointed out by On Uniola laxa (L.) B. S. P.—Mississippi, Atkinson (1905), appears as a typographical error, 1889, S. M. Tracy No. 197 (N, W), since Berkeley and Cunis afterwards consistently as on U. gracilis F. S. Earle No. 197 v,-rote "pilulaeformis" of which the origin, from Latin pilula. little ball, appears clear. (N). s

BALANSIA AND THE BALANSIAE IN AMERICA 57

On Uniola sp. (probably U. laxa).—Sonih and illustrated by Moller (1901) with Carolina, H. W. Ravenel, type in Berke- long-stipitate ascostromata. It has been ley Herb. (K) and in Herb. Curtis (F), impossible to examine authentic speci- Herb. J. S. Billings (N) as Ophiodothh vorax, data lacking (A). mens of B. redundans except for a frag- On Gramineae.-Texas, C. Wright No. 3150, ment in the Atkinson Herbarium, since type of" Xylaria clavulus Berk. & Curt, the type in the Berlin Herbarium is said in Herb. Berkeley (K) and in Herb. to have been destroyed and no other Curtis (F); Blumenau, Santa Catharina, replicates have been available. length Brazil, 1892?, A. Moller No. 830, ex But type (?) of Balansia redundans Moller of stipe in this species is of little conse- (A). quence, as in the writer's experience in ILLUSTRATIONS growing infected host material (cf. below) there is great variability in length Plate II, A to F; Lloyd, Mycol. Notes 7: figure 2745; Moller, Phycom. u. Ascomyceten; of stipes in a single culture. figures 67, a, b. RANGE TYPE LOCALITY The species does not seem to have on culms of Uniola Berkeley County, S. C, been reported from South Carolina, ex- sp. (probably U. laxa (L.) B. S. P.) cept as based on the original gathering DISTRIBUTION by H. W. Ravenel along the old Santee South Carolina to Texas and Brazil on Canal in Berkeley County. In September leaves and culms of Uniola, Paspalum, and E. 1931, accompanied by Mr. J. Bene- other Gramineae (fig. 16). dict, Jr., the writer attempted to find SYNONYMY the species along the line of the old

Balansiopsis pilulaeformis is distinc- Canal in the vicinity of Ravenel' tive in the light-greenish to yellow color Pouchee Plantation, then largely aban- of the context of both hypothallus and doned. The water table and the local ascostroma and in the yellow-green to conditions were doubtless far different in black color of the surface of the hypo- 1931 from those in Ravenel' s day. Lately, thallus. The several pieces of the type much of this area has been covered by specimen of Dothidea pilulaeformis are the water of a reservoir. in fairly good condition, providing a The species is probably much more more definite basis for the specific name common and on more host species than than the type material of Xylaria clavula, the meager number of specimens cited which is very similar except for the lack above might indicate. West (1941) has of the greenish color. The latter is an reported it in Florida upon Paspalum old and weathered specimen from which debile Michx., as well as on P. ciliati- the color could well have faded. Curi- folium. ously, Ellis and Everhart (1887), in CULTURE FEATURES AND PATHOLOGICAL discussing it, failed to appreciate its rela- CONDITIONS tion to the Clavicipitaceae, and it re- As none of the herbarium materials recognition until mained without other showed evidences of conidial stages, it Lloyd's (1924) account. was thought that they would be apparent Farlow (1878) used the name D. pilu- during the development of the fungus laeformis in reference to Epichloe by- on host plants under culture. Hence, poxylon, but Ellis and Everhart (1892) when George Crisfield in 1933 supplied recognized it as a variety of that species living material of infected Paspalum and Earle (1900) as a variety of his pubescens from Savannah, Ga., the plants Ophiodothis atramentosa. The type speci- were propagated in the greenhouse in mens of both D. pilulaeformis and 8- and 10-inch pots at Arlington, Va., Xylaria clavula possess subsessile asco- fo. 3 years. Infected specimens of P. stroma, whereas Balansia redundans on cHiatifolium from Florida, sent by G. F. Panicum sp. from Brazil was described Weber and Erdman West in 1935, were 5 —

58 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE similarly grown for 3 years in a green- text and grayish-black surface. Asco- house at Arlington and in one at Wash- stromata erumpent from within the ington, D. C. Conidial fructifications spherical thickenings of the hypothallus were lacking in all the cultures, although with brownish-gray, longitudinally stri- there was a profuse crop on both culms ated prosenchymatous stipe varying up and leaves of ascostrom.ata, varying from to 5 mm. long by 0.5 to 1.5 mm. thick, subsessile to definitely stipitate. regularly to irregularly subspherical, The infected host plants compared rarely somewhat flattened at the base, wîth normal ones growing nearby 1 to 2.5 mm. in diameter; surface showed practically no pathological affect whitish to black and granular, punctate except suppression of inflorescences and from the ostioles; context tan, subperi- some increase in the number of culms, thecial stroma robust; perithecia broadly i.e., a tendency toward brooming. Under lageniform, rarely crov\'ded, 200^ to some conditions normal inflorescences 300/x by 100/x to 185;a; perithecial wâlls developed upon infected plants. It has 9fx to 18/x thick; asci variable, cylindri- never been possible to detect mycelium co-clavate to elongate cylindrical, 95/" to in these normal-appearing shoots, and 220^. by 4^. to 6jx, with 8 spores; asco- the means by w^hich these healthy shoots spores 95/.1 to 150 jx by 1/x to 1.5^.. arose infected plants has not been from SPECIMENS CITED determined. Since these normal shoots On Orthoclada laxa (L. Rich.) Beauv. developed chiefly in midwinter when the Jurua-Miiy, Amazonas, Brazil, 1901, E. greenhouse temperatures were sometimes Ule. Myc. Brasil. No. 78 (as on O. low-ered, it is possible that an occasional ratiflora), type coll. (?) of 5. asclerotiaca (F. N. W); Amazonas, Brazil, lowered temperature inhibited growth of 1923, J. R. Weir (N, F, W). the parasite but permitted normal culm- elongation, enaoling certain rapidly ILLUSTRATIONS growing shoots to outrun the pathogen. Plate 11, G; P. Herxnings, Hedwigia 43: 259, figures 1 to 3. UNVERIFIED RECORD TYPE LOCALITY Under the name of Ophiodothis vorax Jurua-Mir}', Amazonas, Brazil on Ortho- var. piliilaeformis, Sydow (1914) has clada laxa Beauv. reported this species from Madras in India on Panicum prostratum. If the DISTRIBUTION identification is correct, as is very likely, Known only on culms and leaf sheaths of Orthoclada laxa Beauv. in Brazil and Peru this is the outstanding example of an (fig. 16). American species of Balansia on an American host, both adventive in Asia. DISCUSSION asclerotiaca was described 3. Balansiopsis asclerotiaca Balansiopsis by Hennings (1904b) as on Orthoclada Balansiopsis asclerotiaca (P. Henn.) rariflora (i.e., O. laxa) from a 1901 comb. nov. specimen, Ule No. 3139,. in the Berlin Balansia asclerotiaca P. Henn., Hed- Herbarium. Ule also issued 1901 ma- wMgia43: 259- 1904. terial, possibly from the same gathering Hypothallus reduced to a thin layer, as the type, as No. 78 in his Mycotheca at first white then grayish brown to Brasiliensis (1905). It is very likely black, chiefly effuse, often somewhat that Ule's No. 3139 was destroyed in linear, rarely exceeding 3 cm. in length, 1945 along with the destruction of other but often aggregated over a length of 1 specimens of fungi during the recent cm. upon culms or leaf sheaths, often war. In that case, even if Ule's dis- nearly lacking except for spherical, often tributed specimens No. 78 should prove connate thickenings up to 1 mm. in to be a different gathering, it may well diameter wîth gray prosenchymatous con- be regarded as a substitute type. The )

BALANSIA AND THE BALANSIAE IN AMERICA 59 host, O. rariflora, c|uotecl by Saccardo from Peru by Sydow (191 6). (1905, p. 821) as "ramiflorci," is here The species resembles B. p/iulde- reported as O. laxa in the Hght of Hitch- jortnis, except for its relatively longer cock's (1936) treatment of it as a syno- stipes, its color differences, its generally nym. The fungus has also been reported swollen size, and its more delicate aspect.

Doubtful Species or Species not Included in the Balansiae

DOTHICHLOE ATRAMENTOSA as was well appreciated by Moller, who As previously noted Dothkhloe atra- noted that his species lacked the distinc- mentosa (Berk. & Curt.) Atk. has been tive ephelidial conidia included by Win- applied mistakenly, even if almost uni- ter 1887 (see Rabenhorst, 1886) in the versally, to the species here recognized description of that species. The coni- as Balctnsia henningsiana. The initial ap- dia noted by Moller for his B. diadema plication of Hypocrea atramenlosa by were not formed in any particular fructi- Berkeley and Curtis (1869) to the fungus fication as in B. pallida, but were devel- later called Myriogenospora paspali by oped on germ tubes from sprouted asco- Atkinson (1894) renders the latter ille- spores. Moller seems not to have ob- gitimate, requiring reallocation of the served any conidia or conidial layer on prior epithet: the surface of the hypothallus; and Myriogenospora atramentosa (Berk. & whether they were present or not in his Curt.), comb. nov. specimens can only be deduced from the evidence provided by other specimens Hypocrea atramentosa Berk. & Curt. thought to be of the same species, as ap. Berk., Linn. Soc. (London) none of his type materials are now Jour. Bot. 10: 377. 1869. known to be extant. His specimens at Hypocrella atramentosa (Berk. & Berlin are supposed to have been de- Curt.) Sacc, Michelia 1: 323. stroyed during the recent war, and no 1879. duplicates have thus far been available Epichioe atramentosa (Berk. & elsewhere. Curt.) Cke., Grevillea 12: 103. 1884. Fortunately, there have come to hand Myriogenospora paspali Atk., Tor- four specimens from various localities in rey Bot. Club Bui. 21: 225. South America, which by comparison 1894. with MoUer's description and illustra- Ophiodothis atramentosa (Berk. & tions would seem referable to his B. dia- Curt.) Earle, Torrey Bot. Club dema, except for the evidence upon them Bui. 27: 121. 1900. (Name of sphacelial fructifications bearing mi- only. nute conidia— the very type of fructifica- Dothichloe atramentosa (Berk. & tion most likely to have escaped Moller's Curt.) Atk., Jour. Mycol. 11: notice. The hosts for these four speci- 260. 1905. (Name only.) mens are identified as species of Ichnan- thus. Actually Moller's "rispigen Pani- BALANSIA DIADEMA cum" could well have been an Ichnan-

Balansia diadema (Moeil.) , hitherto of thus. Of these four specimens, one, record only from Santa Catharina in W. A. Archer No. H-253, bears abun- Brazil, was described and illustrated by dantly minute sphacelial conidia or phia-

Moller (1901) as especially distin- lospores 1/x to 1.5/x by 2/a to 3/a, on fili- guished by its bright yellow ascostromata form phialides of \2ii to 15/" in length, arising from mycelial wefts enveloping or Vs^hen branching up to 45 /x and not individual spikelets of a grass identified exceeding 1.5/x in diameter. These phial- as a "rispigen Panicum." It bears a ides might possibly be confused with the marked resemblance to B. pallida Wint., ephelidial conidia or scolecospores of 60 AGRICULTURE :vIOXOGRAPH 4. U. S. DEFT. OF AGRICULTURE

Bala-/2sia pallida Wine., which, however, sioides Moell.), is thus contrary to At- are acute at both ends, having the usual kinson's concept that Claiiceps has the conformation of conidia of the form- sclerotium (h\'pothallus) restricted by genus Epheln. The presence of this spha- the ovary wall. celial layer identifies the species as BALANSIA PALLIDA VAR. referable to Claiiceps rather than to Balansia. ORTHOCLADAE AND THE GENUS Although Hennings (1904a) had con- BALANSIELLA sidered B. diadema Moell. to be synony- Hennings (1904a) described the mous with his Balausiella orthocl.tdae. genus Balansiella with the type, B. ortho- rhat disposition does not seem tenable. cladae as a new combination for his (1900 and 1901) own previously de- Claiiceps diadema (Moell.) comb. nov. scribed variet}^ orthocladae of Claiiceps BaUnsia diadema Moell., Phycom. pallida Wint. That variety was t^'pified u. Ascom., pp., 197-199, 304. by a specimen reported as on Orthoclada 1901. sp. in Brazil, from Para found by J. Huber (Xo. 18). Hennings distin- SPECIMENS CITED guished it from B. pallida because the On Ichnantkus riedelii (Trin.) DoelL— stipitace ascostromata arose from florets British Guiana, 1888. G. S. lenman No. 5975 (W); 1934. \r. A. '.\rcher No. of the living plant, a character that H-253 (W). by itself can hardly justify specific, let On 7. verticillattis Mez—Venezulela. 1941. F. alone generic, recognition. It is extreme- TomayoNo. 1999 (W). ly doubtful if Hennings' type at Berlin On Ichnantkus sp.—Minas Geraes. Brazil. 1925, A. Chase No. 96301/9 (W). has suryived the eflfects of the recent armed conflict; and it is equally unlikely ILLUSTRATIONS that any replicate in the Goeldi Herb- Moller, Phvcom. u. Ascom.. plate 5: figure -4. arium at Belem in Brazil is extant, since information is available to the ettea that TYPE LOCALITY most of the fungus specimens there have Blumenau, Santa Catharina. Brazil, in flo- not endured the vicissimdes of local con- rets of Panicum (?) sp. (Tj-pe specimens ditions. Another gathering (Ule. Myc. supposedly not extant.) Brasil. No. 80) from Rio de Janeiro. DISTRIBUTION doubtless authenticated by Hennings, is In florets of Pariictim and Ichnantkus. well represented in various herbaria un- Venezuela and British Guiana to Santa Cath- der the name B. orthocladae P. Henn; arina in Brazil. it thus has an inherent advantage as a IMPLICATIONS lectot}-pe.

This species is of special significance The specimen resembles superficially in that, although referable to the genus the t}'pe of Balansia pallida Wint. in its

Claiiceps because of its sphacelial fructi- production of yellow ascostromata from fication, it possesses a hypothallus that mycelium-infected ovaries, but it differs encompasses the ovary wall and adjacent from that species in possessing a spha- tissues of the spikelet, a pseudomorph celial fructification and in the lack of proposed by Atkinson (1905) as distin- any ephelidial structure. The genus Bal- >cuishing the genus Balansia. On the other a-risiella is, therefore, a synonym of Claii- hand it has generally been an accepted ceps, and the t}'pe species is referable to notion that in Claiiceps the h\^othallus that genus as Claiiceps orthocladae (P. did not affect the tissues of the spikelet, Henn.), comb. nov. (syn. — C. pallida but was limited by the confines of the Wint. var. orthocladae P. Henn., Hed- ovar)' wall except for apical expansion. wigia 39: 77. 1900; Balansiella ortho- The sclerotium in this species, as in cladae P. Henn., Hedwigia 43: 85. C. patella Berk. & Curt. (syn. C. balan- 1904). BALANSIA AND THE BALANSIAE IN AMERICA 61

SPECIMENS CITED species from Ecuador. All these fungi

On Panicum (?) sp.—Rio de Janeiro, Brazil, grow on species of Chusquea. Their 1900, E. Ule Mycotheca Brasil. No. 80, nomenclature and morphology present lectotype (F, W). complications in addition to those antici- On Orthoclada sp.—Para, Brazil, Huber J. pated in determining their life cycles. No. 18, type (not seen). It is, nevertheless, to be expected that BALANSIA CHUSQUEICOLA these species characterized by pleuro- Bdansia chusqueicola P. Henn. was genous, sphaeroid conidia on specialized described by Hennings (1904c) from a conidiophores that arise from the base Costa Rican specimen found on an un- of pycnidiumlike pits in the periphery identified species of Chusquea in 1854 of a sclerotic hypothallus, might well

by C. Hoffmann. It is recorded as form- have ascostromata homologous with that ing black-encrusted hemispherical asco- on the Panamanian specimen. The type stromata that may reach 1 cm. in diam- of conidial formation in these species eter. Occurring singly on nodes of the indicates a definite relationship to that in deformed (witches'-broomed) host, it Vstilaginoidea, of which but one species,

seemed to Hennings to be a gigantic U . setariae, as recorded by Brefeld equivalent of Balansia gaduae\ but there (1891), has previously been of record is no indication that like the latter it with a similar Balansia-like ascostroma. develops an ascostroma at each node. The ustilaginoid conidial structure ex- Without other evidence than Hennings' cludes these species from the Balansiae. description one would assume that, like EPHELIS JAPONICA IN PUERTO RICO B. gaduae, it lacked accessory conidial fructifications and would be referable to Ephelis japonica P. Henn. has been re- Balansiopsis. A specimen is, however, ported by Weiss (1945) from Puerto available, bearing an accessory fructifica- Rico on Eriochloa polystachya H.B.K., tion that prompts a different allocation. causing a condition termed "blackhead," This specimen, found in 1911 in and constitutes the sole record of this Chiriqui, Panama, by W. R. Maxon, species from the Western Hemisphere (No. 5664) is unfortunately meager; it and the only record of it so far as known is a single, maturing ascostroma sur- upon any species of Eriochloa. As the mounting a stroma of Shropshiria chus- writer had been personally responsible queae F. L. Stevens. Although admit- for the original determination of the tedly inadequate for satisfactory conclu- pathogen in this case, and since there sions, the propinquity of the two struc- are involved certain implications of pos- tures suggests the likelihood of a genetic sible practical significance to agriculture

relation. The less likely possibility is as well as to the taxonomy and mycogeo-

that superimposed ascostromata might graphy of the Balansiae, it seems desir- represent a parasite of the Shropshiria. able here to consider these involvements More adequate materials and observa- in detail. tions are necessary to prove whether the The host:—The specimen (W), con- ascogenous structure is the perfect stage sisting of three depauperate grass spikes

of this better known conidial fructifica- (pi. 8, H), was found on February 2, tion. 1929, in the vicinity of Rio Piedras E. Marchionatto (1940) has concluded by C. Chardon and J. A. B. Nolla, that Shropshiria chusqueae from Central who were under the impression that the America, along with JJstilaginoidea stru- host was E. subglabra (Nash) Hitchc. mosa (Cke.) Clint, from Jamaica, are (syn. E. polystachya H.B.K.). But since referable to the form-genus Munkia rep- the evidence of the specimen itself sug- resented by M. martyris Speg. from Para- gested that the host was much less robust guay, and Petrak (1947) has described a than was to be expected for E. poly- similar genus Neomunkia with one stachya, the aid of Mrs. Agnes Chase 62 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE was enlisted toward clarifying its iden- course, possible that this specimen is not tity. By comparing it with specimens in properly referable to E. japonica but is her charge she concluded that it was merely an aberrant example of Balansia most likely E. ptmctata (L.) Desv., and claviceps or B. obtecta, in which case the that it could not be E. polystacbya. appearance is different on this host from The fungus:—Epbeln japOJiica was that on those hitherto recorded. Al- described by Hennings (1904d) from though on the various recognizable Japanese specimens on inflorescences of hosts the hypothallus of these two species Miscanthus tmctor'm Hack, and of Vas- may exhibit a little variation, it is never pdum thunhergii H.B.K., neither of so definitely suppressed as in the speci- which is recorded from America. The men under consideration. somewhat superficially similar E. mexi- IMPLICATIONS cana Fr. {Balansia ohtectci) , growing on Since this fungus is most likely seed- grasses of closely related genera, is not borne, it is possibly adventive in Puerto known definitely to occur in Asia. Both Rico from Asia and in seed of some species of Epbelis have similar fructifi- other host if not of Eriochloa punctata, cations with scolecoconidia that are which is a ubiquitous tropical grass. Al- practically indistiguishable. In E. niexi- though other hosts of definite Asiatic cana the hypothallus is well developed origin bearing infection of Epbelis ja- with a black laccate rind, whereas in ponica or of E. oryzae have not yet E. japonica the hypothallus is merely a been reported from the Western Hemi- thin grayish to gray-black film over the sphere, it is realized that as yet they surface of the stunted inflorescence. E. may merely have escaped detection. When japonica is thus similar to, and may be found, their early effective eradication the same species as E. oryzae Syd. found would be expedient. Furthermore, inocu- on rice in the Orient and also unknown lation experiments to determine the pos- in America. The Puerto Rican specimen sible susceptibility of Eriocbloa to def- on Eriochloa, having a very thin gray to inite endemic Puerto Rican strains of black film over the inflorescence, is thus Balansia obtecta should have pertinent considered to be E. japonica. It is, of significance.

HOST RELATIONS IN THE BALANSIAE

Host Specificity

In the usual experience of a collector In any locality a particular species of any species of the Balansiae after with several known host-species, as, for the first specimen has been noted, it is instance, Balansia strangulans with 26 soon obvious that in the immediate here recorded, is found almost invariably vicinity there are many other specimens only upon one host, although other of the same fungus in about the same grasses known as hosts elsewhere may stage of development on plants of the be growing in the immediate vicinity same host species. Rarely, indeed, is it or even intermixed in the plant associa- possible to find more than one host tion. The most ready explanation is that species bearing the same species in the this condition is the result of distinct immediate environment. There are nearly strains of the endophyte to w^hich only always islands, or colonies; of infected a particular host species, or perhaps a host plants, sometimes in almost pure strain of that host species, is susceptible. stands, but usually as colonies with nor- Such susceptibility or its lack may, of mal plants intermixed. course, be due to protoplasmic compati- BALANSIA AND THE BALANSIAE IN AMERICA 63 bility or antagonism, or may be merely a yet of record. Nevertheless, the present phenological phenomenon dependent up- list reveals a general limitation of a on the coincidence of spore production particular species of the Balansiae to and dissemination of the parasite with special groups of hosts. For instance, the momentary exposure of critically sus- Balansia cyperi and B. cyperacearum are ceptible host tissue, as of stigmas in restricted to Cyperaceae. Furthermore, Cenchrus, for example. Regardless of among the grasses certain taxonomic the nature of resistance or escape of a units, recognized as "tribes," are groups particular host, there is clearly some of hosts to which individual species of the specific relation between host and para- Balansiae appear to be restricted, al- site that results ordinarily in infected though this restriction is obviously not plants of but one host-species in a par- entirely consistent. The following out- ticular locality where several other species line summarizes the present information recorded as hosts are also present but listing the Balansiae recorded for each free from infection by the endophyte. tribe of the Gramineae but omitting cer- It is to be expected that further search tain records here considered obviously will disclose relations to host plants not questionable or erroneous.

^^^^nsia linearis. 1. Bambuseae \ \ Balansiopsis gaduae. _ _ i Balansia epichloe. 2. Festuceae J 5_ pilulaejormis. \Balansiopsis asclerotiaca. 3. Hordeae Balansia linearis. 4. Aveneae Atkinsonella hypoxylon. (Balansia aristidae.

5. Agrostideae )b. hemicrypta. \ B. epichloe. \Atkinsonella hypoxylon var. texensis. 6. Zoysieae None.

f. discoidea. 7. Chlorideae \ Balansia strangulans

( B. epichloe. 8. Phalarideae None. 9. Oryzeae Balansia ambiens. 10. Zizanieae b. pallida. 11. Melinideae None. (Balansia claviceps. B. obtecta. I B. strangulans. I B. strangulans f. discoidea. j 12. Paniceae ) B. suhnodosa.

"j B. henningsiana. B. epichloe. I IB. ambiens. . Balansiopsis pilulaejormis. yEphelis japonica. (Balansia claviceps. 13. Andropogoneae Jb. henningsiana. is. epichloe. 14. Tripsaceae None.

Teratology

The aberrant effects on hosts by the may conveniently be summarized in American species of Balansiae have in table 1. most cases been discussed severally; they .—..

64 AGRICULTURE MONOGRAPH 4, U. S. DEFf. OF AGRICULTURE

Table 1 . Teratology induced by species of the Balansiae

Name Cul- Sterility Dwarf- Gigan- Broom- Foliage ture ness tism ing deformation

Balansia claviceps — + p p p Terminal leaf +. Balansia obtecta + + + ± + Do. Balansia cyperi + + 7 ? Terminal bracts swollen. — ? Balansia pallida + ? p p — Balansia strangulans. . + + + + Terminal leaf reduced. Balansia strangulans f. discoidea _ + + — + Do. Balansia subnodosa — + + — + Do.

. Balansia henningsiana + + + i ± Do. — > Balansia cyperacearum + ? 1 ? p Balansia epichlo'e + + + — ± Slight. Balansia aristidae + + — ± — — — — Balansia hemicrypta . . + + + Balansia ambiens ? p — p p Balansia linearis + + + Terminal leaves reduced and rolled. - — Atkinsonella hypoxylon. . + + + + A. hypoxylon var. texensis — + + ? p ? > — — Balansiopsis gaduae . . + + + Reduction and rolling of leaf. Balansiopis pilulaeformis + + + + Reduaion of ligule on terminal leaf. Balansiopis asclerotiaca _ + + P p > *? Ephelis japonica — + + P p Epichlo'e typhina + + + + Reduaion in leaf area

Mode of Infection and Dissemination

Although only 7 of the 19 recognized eased plants compared with healthy ones species and segregates of the Balansiae kept under observation in parallel cul- here treated have been under observa- ture or by records of coincidence of tion in living condition on hosts under phenology of the endophyte and of experimental culture, these 7 may be con- healthy host to suggest the likelihood sidered as representative of the 19. All there also of stigma infection. The above of them are systemic endophytes. For conclusions are summarized in table 2. 9 others the evidence of herbarium ma- In all 19 species and their segregates, terials or of published accounts suggests the spores, whether ephelidial conidia or systemic infection also. In only one of ascospores, are gelatinous-glutinous, sug- the species, Balansia obtecta, has the gesting the possibility of transmission to nature of the infection been proved by stigmas through the medium of insects. experiment to be by penetration of the There is nothing, however, to preclude host through the stigma and style to the transmission by the drift of spores ex- ovule, resulting in an infected seed pelled into the air, as has been so well that develops into a host plant bearing shown by Ingold (1948) in the case of the endophyte. In at least 15 other Epichlo'e typhina, or by splashing rains. species, however, there is some corollary Spores have been observed to "squirt" evidence provided by the habits of dis- or shoot in the manner noted by Ingold — .

BALANSIA AND THE BALANSIAE IN AMERICA 65

Table 2. Evidences oj systemic injection in American Balansiae

Stigma Ob- Ob- Evi- Deduc- Stigma infec- served served dence tion infection Name in in of from tion suggest- nature culture speci- printed proved ed by mens accounts evidence

Balansia obtecta + + + + + + B, claviceps — — + + B. cyperi + + B. pallida _ _ + ? -? _ p B. strangulans + + + — + — — — — B. strangulans f, discoidea — _ + + B. suhnodosa + ? — + B. henningsiana + + + + — + B, cyperacearum + + B. epichloe + + _ — + + + B. aristidae + + + ? — + B. hemtcrypta ...,., + + + + — + amhiens 7 B, _ B, linearis _ + ) _ + ? — Atkinsonella hypoxylon . . . T" + + + + A. hypoxylon var. texensis. — — ? + — + Balansiopsis gaduae — — + ? > — + ? — — J5. pilulaeformis + + ? 4- B. asclerotiaca — — + ? 7 — p Epichloe typhina + + + + +

+ Present. Absent.

in 5 of those species: Balansia strangu- germinating conidia of these two species lans, B. henningsiana, B. epichloe, At- in water onto stigmas and the only suc- kinsonella hypoxylon, and Balansiopsis cessful inoculations by B. obtecta wert pilulaeformis. accomplished by brushing stigmas with

When drops of water were allowed to conidia, it was assumed that splashed fall upon wetted spore-bearing surfaces raindrops containing viable conidia were of ephelidial fructifications of Balansia not the natural vehicle for successful in- obtecta and of Atkinsonella hypoxylon, oculation. After conidia had been sprayed

it was found that conidia were readily in water droplets onto stigmas the ovules dislodged and splashed into the air, a failed to develop into seeds, the re- condition comparable to the effect of sultant sterility being credited to the splashing by rain. Such splashed droplets failure of pollination because of the caught on glass slides contained conidia bursting of pollen when immersed in that germinated readily. But since no suc- water rather than to any effect of the cess attended inoculation by atomizing fungus.

Incidence, Epidemiology, and Control

As previously noted, Balansia obtecta rarely on this host elsewhere, although is abundant on Cenchrus echinatus in it has been observed in some abundance some places in Florida, and B. claviceps at certain places in Virginia and Mary- likewise appears to be common on Setaria land on Panicum liyidheimeri and P. at certain places in Cuba. B. strangulans huachucae. Thousands of acres of Aris- has been observed to affect thousands of tida glauca in Texas were observed in plants of Panicum clandestinum at one 1930 to have more than 10 percent of place in New York but has been found the plants infected by B, hemicrypta. 66 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

The actual incidence generally of grasses seem that where a susceptible host species affected by Balansiae is practically un- is to be planted only seed should be known, because infected plants are diffi- chosen that had been produced far re- cult of detection and so few persons have moved from plants bearing the infection. been interested that the records have Where it may be desired to produce in- largely been as a result of mere chance fected plants, as of bur-free Cenchrus observations; no systematic, large-scale for forage, infected seed or, still better, surv^eys having been undertaken with cuttings of infected plants would be the Balansiae as the object. desideratum. Some naturally infected seed No experience is available on which could doubtless be obtained from plants to base attempts at prevention or control growing near infection, but a large pro- of these fungi as undesirables. As they portion of such seed might chance to be are probably all seed-borne, it would healthy.

Susceptibility and Resistance

Practically nothing is of record rela- species. Atk'insonella hypoxylon is tive to specific host-susceptibilities or represent chiefly in old stands of either sistances to infection by Balansiae. Al- Danthonia compressa or D. spicata, be- though the appended host-index listing ing rare in obviously young associa- all known suscepts is suggestive, it is tions where populations of the parasite realized that this list will no doubt be are small because they have not yet had augmented by further observations. time to build up in numbers. Presumably,

There is, furthermore, the likelihood of here, as in Balansia obtecta, with enough a complexity of genetic susceptibilities inoculum present many of the seeds and resistances within any species, as was that are set are likely to bear the endo- suggested heretofore, to explain the in- phyte. It is probable that a similar rela- stances of plants of susceptible species tionship between host and parasite exists being free from infection even contigu- for all the Balansiae. ous to infected plants of other host

Recovery From Infection

The occasional development of nor- outdoors in Cambridge, Mass., in 1931, mal, healthy shoots from occasional Bd- it developed one shoot bearing a normal ^?2j-/rf-infected plants of Cenchrus, An- inflorescence entirely free from infection. dropogon, and Paspalum has been ob- When clumps of a grass of unknown served in plants under cultivation. The heritage outdoors are found to bear both same phenomenon has been observed in healthy and diseased shoots, it is reason- the native plants of Cenchrus in Florida, able to account for the condition as the and in New England it is commonplace product of at least t^'o seeds—one healthy, in midsummer to see individual clumps one infected. But the observ^ed growth of Danthonia spicata producing normal of a healthy shoot of Cenchrus from a inflorescences and also aborted ones bear- diseased plant requires a difl^erent ex-

ing Atkinsonella. When infected clumps planation. Clearly this healthy shoot is of Danthonia were transplanted to a an escape of new healthy tissue from the warm greenhouse in Cambridge, Mass., endophyte that was resident in the parent in 1932, however, only infected shoots clone. At present no explanation for this developed, resembling the condition escape from infection is forthcoming, commonly noted outdoors in Virginia although there is strong suspicion that and Maq'land. at critically lowered temperatures the When a plant of Cenchrus echinatus host grows faster than the endophyte, infected by Balansia obtecta was grown thus escaping the infection, because the BALANSIA AND THE BALANSIAE IN AMERICA 67

Graminales have no meristem in the the young, parent cells were infected then internodes through which the parasite this noninfected internode would act as might make its progress upwards in a barrier of matured but healthy tissue the stem. As the mycelium appears to through which the endophyte could not

grow only in meristem and with it, if grow. an internode has once elongated before

CLIMATE AND GEOGRAPHIC DISTRIBUTION Temperature Relations

The cited records for the species of period, although endophytic mycelia were Balansia and Balansiopsis show a range detectable. After removal to a tempera- primarily within the warmer parts of ture of 20° to 32° C. the fungus fruited the continent, with few outliers in re- normally. Evidently there was a tempera- gions having minimum temperatures ture differential between vegetative much below —20° C. The host plants, growth and the development of fruiting however, although they are species found structures, both conidial and ascostro- over a greater range into the higher lati- matic. tudes with lower temperatures, tend to Atkinsonella hypoxylon, in the light follow the same general pattern. This of its geographic distribution, would ap- distribution points to limitations due to pear to have a somewhat lower tempera- temperature. There are nevertheless some ture tolerance or preference than other notable exceptions that lack agreement species of Balansiae, its present known with this hypothesis. Balansia epkhloe range embracing most of the eastern from Virginia in the United States to half of that of the host species; and it Minas Geraes in Brazil on hosts that has been the writer's experience to find are predominantly subtropical in their it most plentiful in the cooler parts of distribution is also recorded from North its range. When infected plants of Dan- Dakota and Wisconsin on species of Cal- thonia spicata were "forced" in a green- amagrostis, which have a much more house at 13° C, only the typhodial northerly range. There is, however, a gap stage was produced. But when some of between this recorded northern area and these same forced plants were exposed the known more southern distribution in to day temperatures of 22° to 34° C, which the fungus apparently has never ephelidial fructifications were soon in been found. Balansia strangulans, sensu evidence and ascostromata were devel- lato, probably most common in the oped within 2 weeks. When comparable tropics, is found on various species of plants were forced at these higher tem- grasses all the way from Massachusetts peratures the typhodial stage was defi- 42** (lat. N.) to Paraguay (lat. lA" S.) . It nitely inhibited, at least it was present is possible that this species is made up of for only a short time and the initials strains having different temperature of ascostromata were soon apparent, limitations, as was shown by its failure to early suppressing the conidial fructifica- fruit on infected plants of Florida origin tions altogether. Evidently higher tem- after several years' growth in Virginia, peratures than those suitable for coni- whereas on native infected plants in both dial formation may be required for the

Virginia and Massachusetts it continues development of ascostromata. In five of to fruit year after year. the species of Balansia the ascostromata When Sporobolus and Ctenium, in- developed rapidly when grown at the fected by B. epichloe^ and Cenchrus, in- higher temperatures (26° to 35° C) fected by B. ohtecta, were kept at 13° quickly displacing ephelidial fructifica- to 15° C. for 30 days, they grew slowly; tions. but the fungi did not fruit during that 68 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Moisture Relations

In the United States and Canada all susceptible grasses, since generally to the the findings of these species, except for westward the grasses blossom during the one specimen of Bala??sia hemkrypta periods that are relatively dry. The seem- from New Mexico, have been made ing striking discontinuity in the range east of 100° longitude W., and in South of B. hemicrypta, known only from America except for t^'o records from Texas and Brazil, may be merely lack of Chile all findings have been east of the records; but if the record is adequate,

Andes. This is somewhat different from it may be the result of the accident of the distribution of Clavkeps, which oc- geological history separating by climatic

curs also on the Pacific coast in both changes a former continuous range. It is Americas. This eastern distribution sug- possible, of course, that the distribution gests a restriction to relatively humid of other species has been equally for- climates at the time of flowering of the tuitous.

Phenology

The time of fruiting of these species of the host there continues late into the coincides in general with the normal season. If all these species infect the

times of flowering of their particular o^oile through initial stigma infection, it

hosts. For some, notably Bala7isia stran- is to be expected that infection could gulans, B. hennmgstana, B. epichloe, not take place at all unless temperature and B. aristidae, the fructifications, in- and humidit}" relations are favorable at cluding the ascostromata, tend to be the time of flowering of the host. Where ripened before and continue to be de- these favorable conditions plus the pres- veloped until long after anthesis. As ence of adequate inoculum are not avail- noted previously, Atkinsonella hypoxylon able at anthesis, as m,ay well be the case in the vicinit}' of Washington, D. C, on the Pacific slope of North America produced its t}-phodial stage well before, where these fungi have not yet been the ephelidial stage during, and the as- found, infection does not take place. The costromata well after, anthesis, thus more lack of favorable conditions and of coin- than synchronizing with the time of cident phenolog)' of host and parasite at flowering and the temperature changes in some moment in past geological history the march of the season. On the other could easily explain present discontinui- hand, farther north in New England no ties in the range of any species having such consistently progressive develop- momentarily so critical a requirement. ment was obser^-able, as some flowering

Endemism

The 17 species of Balansiae considered tion for this apparent anomaly: Of the here are known with certainty only from total 124 authentic host species for these the Americas. This seems surprising, if American Balansiae, 106 are endemic in the assumption that these species are America, and comparatively few of these seed-borne be accepted, especially since endemics are known to have established the comparable seed-borne Epichloe ty- themselves elsewhere, possibly because

phina has been found in both hemi- none of them is of sufficient value eco- spheres. It is to be expected that seed- nomically to have commended its pur- borne parasites would be widely dis- poseful introduction abroad and because tributed over the earth because infected the few of these host species that have seeds are as easily transported as healthy become adventives in the Eastern Hemi- ones. sphere may well have been derived ini- There may, however, be an explana- tially from healthy seeds or else infec- BALANSIA AND THE BALANSIAE IN AMERICA 69

tions there have not yet been found. by accident. In their native habitats these Of the 18 host species of foreign origin, foreign species of susceptible grasses 5 were doubtless introduced purposefully have apparently lacked the opportunity as grasses of some economic worth: for infection possible only through inoc-

Axonopus compressus, Bambusa arundi- ulum available in America, and it is like- nacea, Cynodon dactylon, Setaria palmi- wise reasonable to assume that they folia, and Sorghum vulgare. Another have become infected only after exposure cosmopolitan and tropical host, Sporo- to inoculum in the Western Hemisphere. bolus poiretii, may have been adventive

ECONOMIC CONSIDERATIONS

The effects of species of Bdansia and highland and in New England Dan- closely related genera on their hosts have thonia spicata and D. compressa are com- generally escaped attention because these mon native pasture grasses of more worth fungi do not cause necrotic lesions nor than is usually appreciated. do they kill the host plants outright If the failure of infected seeds to but merely prevent the setting of seed. germinate, found to be so common in No comprehensive attempt in the way of experimental plantings, is not merely a survey has ever been made to deter- fortuitous, that defect alone would be of mine the distribution of these very in- economic significance to both vendor conspicuous fungi except for the present and purchaser of such seeds. It is obvious recording of specimens in herbaria, and also that where seed production is the it is most certain that their actual host intent of the grower it would be to his and geographic ranges are considerably decided disadvantage to plant Balansia- in excess of those indicated by present infected seed because the plants that meager records. In the total absence of develop from them can yield but little any such survey no adequate estimate return. It is needful to plant uninfected of the economic effect of these parasites seed when it is desired to establish a can be offered and the present remarks stand of perennial grasses, because Bal- attempt merely to call attention to some ^/;j7'^-infected plants could have little rather evident possibilities. progeny, whereas healthy plants would in From a perusal of the grasses known turn produce seed tending to make the to be hosts for the 13 American species stand denser. A few years ago a practical of Bdansia and the 4 species in closely problem concerned with this requirement related genera, it can be seen that these was encountered vv^hen efforts were made parasites cannot be entirely without to reestablish stands of native grasses in economic significance. If we except Se- the North American "dust bowl." Among taria and Sorghum it is true that none the native short-grass species used was of these is normally used as human food. Aristida fendleriana. Inspection of some Some are probably of no concern to of the seed of that grass harvested in civilized man, but most of the species New Mexico for this purpose revealed are pasture plants of a lesser sort and a high percentage of sterile florets in- certain ones are major forage plants, fected by Balansia hemicrypta. especially in the more torrid climates. With modern ease of transportation The more important of these hosts there is properly an increasing tendency are species of Andropogon, Cenchrus, for plants to be introduced accidentally Cynodon, Panicum, Setaria, and Sor- into regions where they have never be- ghum. In the North Calamagrostis and fore been present. Seed-borne parasites, Bouteloua are the more important eco- so difficult to guard against, are an ever nomic plants known to be affected by possible menace, although it must be ad- these fungi, although in the Appalachian mitted that there is thus far little ade- 70 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE quate evidence of international carriage ly with no deleterious effect. In many of species of Balansiae in commercial areas of the Appalachian highland, where seed, although Wernham (1942) has Dantho7iia compressa is one of the chief noted the presence of Epichloe typhina forage plants and where some stands of in fescue seed from Hungary. A possible that species may show over 50 percent menace abroad—not to America—is At- infection by Atkinsonella, cattle appear kinsonella hypoxylon. Its known hosts, to consume the pseudomorphs with im- two species of Danthonia and at least punity. one species of Stipa, are less important These pseudomorphs when first in America than are species of the same formed have a sweetish taste and when a genera in New Zealand and Australia small handful was chewed by one ex- (Bews, 1929), which are esteemed perimenter he suffered no ill effects. To among the more valuable of their forage determine more adequately if animals grasses. It is conceivable that if seeds of could suffer any deleterious effects from American species of Danthonia bearing eating these pseudomorphs a simple ex- Atkinsonella hypoxylon were introduced periment was resorted to in 1932. Three into those regions, some of the native guinea pigs, one rabbit, and three white grasses there might prove susceptible. rats were subjected for 2 days to a diet Likewise the similar African and Asiatic limited to water and pseudomorphs only, Balansiae, so little known, but usually half of Balansia obtecta harvested from referred to Epichloe, Balansia, or Ophio- Cenchrus echinatus in Florida and half dothis, constitute, until more properly of Atkinsonella hypoxylon from Dan- understood, an equally serious threat to thonia spicata in Virginia. The animals many grasses of present or potential ate pseudomorphs but showed no dele- human concern in America. The sedges terious effects and remained in good thus far recorded as hosts are not of health for a week following their return significant economic moment. to a normal diet. It is possible that the sterility induced To determine whether the ergotlike by these species might well tend, especial- pseudomorphs of Balansia obtecta pos- ly in annuals, toward their ultimate sup- sessed any ergotlike quality they were pression or even extinction as a com- utilized in treating domestic cocks ex- ponent of a local flora. The Florida actly according to the requirements at sandbur {Cenchrus echinatus) , however, that time of the American Pharmaco- thrives as a pest because of its annoying paeia (Barger, 1931). The decoction burs, despite an abundance of inoculum. was very kindly prepared by the late Susceptible perennial grasses, Danthonia O. F. Black, then of the Division of compressa, D. spicata, and Aristida glau- Drug Plants, United States Bureau of ca, appear to hold their ground with no Plant Industry, and three of the domestic indication of extinction. Species of Pani- cocks used, Rhode Island Reds, were pro- cum and related genera, likewise com- vided by the United States Bureau of Ani- monly infected by Balansiae, show little mal Industry. Within 2 weeks after two sign of suppression because enough nor- subjects were injected they showed normal plants are present to provide an mal health; the third rooster, untreated available seed supply. and kept as a control, died on the third The evident relationship of the Balan- day from causes unknown. This opera- siae to Claviceps, especially those species tion was later repeated, using three producing pseudomorphs simulating er- Leghorn roosters with, however, no dele- gots, has suggested the possibility that terious effect on either the treated sub- these pseudomorphs might cause poison- jects or the control. One is forced to ing if they were eaten by animals or by conclude that as a component of forage man. In Florida where acres of infected these pseudomorphs are innocuous. It is Cenchrus echinatus produce quantities of even possible that they may help the diet the ergotlike pseudomorphs of Balansia through the addition of vitamins and ohtecta, cattle eat this material seeming- soluble carbohydrates. BALANSIA AND THE BALANSIAE IN AMERICA 71

On at least one host the effect of a bur can spread by runners rooting at the species of Balansia is an economic ad- nodes, individual plants, especially those vantage. The Florida sandbur, Cenchrus sterilized by this fungus, tend to make ecbinatus, is good forage for cattle up pads covering areas from a square foot to the time of flowering when the "burs" to a square rod or more in size. In Flori- make it offensive to them, but they ap- da during late summer and autumn such pear to find no objection at any time sterile pads are eaten by cattle, whereas to plants sterilized by Balansia obtecta normal plants of the sandbur are un- and bearing pseudomorphs. Since sand- touched.

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Hennings, p. Ellis. J. B., and Gerard, W. R. 1889 (1890). CLASS 3—FUNGL (Pp. 468- 1900. FUNGI PARAENSES I. Hedwigia 39 602.) In Britton, N. L., Cata- (Beibh): (76)-(80). logue of Plants Found in New Jer- Rpt. 2: sey. N. J. Geoi. Survey 1901. FUNGI PARAENSES (l). Bol. de 27-642. Mus. Paraense Hist. Nat. e Eth- Farlow, W. G., nog. 3: 231-237. 1878. LIST OF FUNGI FOUND IN THE VICINI- TY OF BOSTON, PART II. Harvard 1902. FUNGI BLUMENAVIENSIS II A CL. Univ. Bussey Ins. Bui. 2 (pt. 3): ALFR. MOLLLER LECTi. Hedwigia 224-252. 41: 1-33. Fitzgerald, L. 1949. hypocreales of iowa. Iowa Univ. 1904a. FUNGI FLUMINENSES A. CL. E. ULE Studies in Nat. Hist. 19(2) : 1-32, illus. COLLECTI. Hedwigia 43: 78-95. Fries, E. [M.] 1849. SUMMA vegetabilium scandinaviae 1904b. FUNGI AMAZONICI II. A CL. ERNESTO (sect. post.), pp. 259-564. ULE COLLECTI. Hedwigia 43: 242-273, illus. Garcia Rada, G., and Stevenson, J. A. 1942. LA flora fungosa peruana. 122 pp. Lima, Peru. 1904c. EINIGE NEUE PILZE AUS COSTARICA Gaumann, E. [A.] UND PARAGUAY. Hedwigia 43: 1926. vergleichende morphologie der 147-149. piLZE. 626 pp., illus. Jena.

1904d. EINIGE NEUE PILZE AUS JAPAN II. 1949. DIE PILZE, GRUNDZiJGE IHRER ENT- Hedwigia 43: 150-153. WICKLUNGSGESCHICHTE UND MOR- PHOLOGIE. 382 pp., illus. Basel, 1908. FUNGI BAHIENSIS A CL, E, ULE COL- and Dodge, C. W. LECTI. Hedwigia 47: 266-270. 1928. COMPARATIVE MORPHOLOGY OF FUN- Herbst, GI. 701 pp., illus. New York W. and London. 1899. FUNGAL FLORA OF THE LEHIGH VAL- LEY, PA, 229 pp., illus. Allen- Gilman, J. C, and Archer, W. A. 1929. THE FUNGI OF IOWA PARASITIC ON town, Pa. PLANTS. Iowa State Col. Jour. Hitchcock, A. S. Sci. 3: 299-507, illus. 1935. MANUAL OF THE GRASSES OF THE Gola, G. UNITED STATES. U. S. Dept. Agr. 1930. l'erbario micologico di p. a. Misc. Pub. 200, 1040 pp., illus. SACCARDO. CATOLOGO. 328 pp., Washington. illus. Padova.

Hardison, J. R. 1936. MANUAL OF THE GRASSES OF THE 1945. observations ON GRASS DISEASES IN WEST INDIES. U. S. Dept. Agr. KENTUCKY, SEPTEMBER, 1942 TO Misc. Pub. 243, 439 pp., illus. SEPTEMBER, 1944 AND A PRELIMI- HOHNEL, F. VON. NARY CHECK LIST. U. S. Bur. 1909. FRAGMENTE ZUR MYKOLOGIE: 21 Plant Indus., Soils, and Agr. En- 6, 217. Akad. der Wiss. Wien, Math.- gin. Plant Dis. Rptr. 29: 76-85. Nat. Kl. Sitzber. (Abt. 1) 118: Harvey, F. L. 312-315, illus. 1896. contributions to the pyrenomy- CETES OF MAINE—I. Torrey Bot. Club Bui. 23: 50-58. 1910. FRAGMENTE ZUR MYKOLOGIE: 630. Akad. der Wiss. Math.- Heald, F. D., and Wolf, F. A. Wien, 1912. A PLANT-DISEASE SURVEY IN THE Nar. Kl. Sitzber. (Abt. 1) 119: VICINITY OF SAN ANTONIO, TEXAS. 933-941. U. S. Dept. Agr. Bur. Plant Indus. Bui. 226, 129 pp. illus. 1923. SYSTEM DER FUNGI IMPERFECTI Hennings, p. FUCKEL. In Falck, R., Mykolo- 1899a. xylariodiscus nov. gen, und gische Untersuchungen und Be- einige neue brasilianische as- richte 1 (heft 3) : 301-369. COMYCETEN DES E. ULE'SCHEN (J. Weese, edit.) HERBARS. Hedwigia 38 (Beibl.): (63)-(65), illus. 1925.UBER HETEROSPHAERIA POAE FUCKEL. Mitt, aus dem Bot. Inst, 1899b. [Editorial footnote.} Hedwigia 38 der Techn. Hochsch. Wien 2 (heft (BeibL): (219). 3): 93-95. 74 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Ingold, C. T. MOLLER, A. 1948. the water-relations of spore 1901. PHYCOMYCETEN UND ASCOxMYCETEN, DISCHARGE IN EPICHLOE. Brit. UNTERSUCHUNGEN AUS BRASILIEN. MycoL Soc. Trans. 31: 277-280, In Schimper, A. F. W., Botanische illus. Mitteil. aus den Tropen. 9, 319 pp.. Jones, H. L. illus. Jena. 1898. list of ohio plants not recorded MOLFINO, F. in the latest state catalogue. J. 1930. NOVED.\DES MICOLOGICAS ARGEN- Oberlin CoL Lab. BuL 9: 4-8. TINAS (m). Soc. Cient. Argen- KiLLLAN, C. H. tina An. 109: 127-131. 1919. SUR LA SEXUALITE DE L' ERGOT DE SEIGLE, LE CLAVICEPS PURPUREA MONTAGNE, [J. F.] C. (tulasne). Soc. MycoL de 1855. CRYPTOGAMIA GUYANENSIS, Ann. France, BuL Trimest. 35: 182-197, des Sci. Nat., ser. 4, 3: 91-173, illus. illus. King, A. M. 1919. NOTES on the GENUS BALANSIA. 1856. SYLLOGE GENERUM SPECIERUMQUE So. African Jour. Sci. 15: 670- CRYPTOGAMARUM. 498 pp. 673, illus. [Paris.] Lindau, G. Nannfeldt, A. 1892-1897. dothideales. J. hypocreales; 1932. STUDIEN iJBER DIE MORPHOLOGIF In Eng- (Pp. 343-372, 373-383.) UND SYSTEMATIK DER NICHTLI- ler, A., and Prantl, K., Die Natiir- CHENISIERTEN INOPERCULATEN lichen Pflanzenfamilien ihren Gat- DISCOMYCETEN. Nova Acta Reg. rungen. pt. 1 (Abt. 1): 513 pp., Soc. Sci. Uppsal., ser. 4, 8 (2), illus. 368 pp., illus. Link, H. F. Pammel, L. H., Weems, B., and Lamson- 1826. ENTWURF FINES phytologischen J. SCRIBNER, F. PFLANZENSYSTEMS NEBST FINER 1901a. FUNGUS DISEASES OF GRASSES. ANORDNUNG DER KRYPTOPHYTEN. (Pp. 253-344). In Grasses of Akad. der Wiss. Berlin (Phys. Iowa, Part I. Iowa Agr. Expt. Sta. Kl.) 1824: 145-194. Bui. 54: 71-344, illus. Lloyd, C. G. Weems, J. B., and Lamson-Scrib- 1924. mycological notes. No. 71 (v. 7, NER, F. No. 6): 1237-1268, illus. 1901b. FUNGUS DISEASES OF GRASSES. Maneval, W. E. (Pp. 185-292.) In Grasses of 1937. a list of MISSOURI FUNGL Mo. Iowa, Part I. Iowa Geol. Survey Univ. Studies 12 (3): 3-150. Bui. 1, 525 pp., illus. Des Moines.

M.\RCHIONATTO, J. B. Patouillard, N. 1940. NOTA TAXONOMICA SOBRE MUNKIA 1899. champignons de Guadeloupe. MARTYRIS, USTILAGINOIDEA STRU- Soc. MycoL de France, Bui. 15: MOSA Y SHROPSHIRIA CHUSQUEAE. 191-209, illus. Rev. Argentina de Agron. 7: 172- and Hariot, P. 175, illus. 1893. fungos aliquot novos in regione CONGOANA COLLECTOS. SoC. My- 1947. NOTA SOBRE BALANSIA CLAVICEPS, col. de France, Bui. 9: 206-211. HONGO PARASITO DE LAS GRAM- INEAS. Rev. Argentina de Pazschke, O. Agron. 14: 175-176, illus. 1892. erstes verzeichniss der von e. Miles, L. E. ULE IN DEN JAHREN 1883-87 IN 1935. THE ASCOMYCETES of MISSISSIPPI. BRASILIEN GESAMMELTEN PILZE. U. S. Bur. Plant Indus., Soils, and Hedwigia 31: 93-114. Agr. Engin. Plant Dis. Rptr. 19: 39-66. [Processed.] 1896. U. VERZEICHNISS BRASILLAJsJISCHER

Miller, J. H. VON E. ULE GESAMMELTER PILZE. 1941. THE ASCOMYCETES OF GEORGIA. Hedwigia 35: 50-55. U, S. Bur. Plant Indus., Soils, and Peck, C H. Agr. Engin. Plant Dis. Rptr. Sup. 1875. report of the botanist. n. y. 131: 31-93. [Processed.] State Mus. Rpt. (1873) 27: 73- 116, illus. 1949. A REVISION OF THE CLASSIFICATION OF THE ASCOMYCETES WITH SPE- QAL EMPHASIS ON THE PYRENO- 1887. NAMES OF NEW YORK STATE PYREN- MYCETES. Mycologia 41: 99-127, OMYCETOUS FUNGI. N. Y. State illus. Mus. Bui. I (2): 49-56. BALANSIA AND THE BALANSIAE IN AMERICA 75

Peck, C. H. Saccardo, p. a. 1909. list of speaes and varieties of 1879. MICHELIA [1877-79]. v. 1, 619 fungi described by c. h, peck, pp. Patavii. STATE BOTANIST, N. Y. State

Mus. Bui. 131: 59-190. 1882. MICHELIA [1880-82]. v. 2, 682 Petch, T. pp. Patavii. 1920. HYPOCREACEAE ZEYLANICAE. Roy. Bot. Gard, Peradeniya Ann. 7: 85- 1883. SYLLOGE FUNGORUM. V. 2, 815 pp., 138. illus. Patavii.

1933. NOTES ON ENTOMOGENOUS FUNGI. 49. CORDYCBPS FLAVELLA B. & C. 1884. SYLLOGE FUNGORUM. V. 3, 860 pp. Brit. Mycol. Soc. Trans. 18: 48- Patavii. 49. Petrak, F. 1886. SYLLOGE FUNGORUM. V. 4, 808 pp. 1947. ijber die gattungfn shropshiria Patavii. stevens, munkia spec. und neo- MUNKIA N. GEN. Sydowia 1: 1886. SYLLOGE FUNGORUM. ADDIMENTA. 328-331. ad. V. 1-4, 484 pp. Patavia. Presley, J. T. 1947. A HOST INDEX OF MISSISSIPPI PLANT 1891. SYLLOGE FUNGORUM. V. 9, ll4l pp. DISEASES. U. S. Bur. Plant Indus., Patavii. Soils, and Agr. Engin. Plant Dis. Rptr. Sup. 169: 55-168. [Proc- essed.] 1892a. SYLLOGE FUNGORUM. V. 10, 964 Rabenhorst, L. (G. Winter, edit.) pp. Patavii. 1886. FUNGI EUROPAEI EXSICCATI KLOTZ- SCHII HERBARII VIVI MYCOLOGICI 1892b. FUNGILLI ALIQUOT HERBARII REGII CONTiNUATio. New ed., (ser. 2) BRUXELLENSIS. Roy. Soc. Bot. dc Century 36: Nos. 3501-3600. Belg. Bui. (pt. 2) 31: 224-239. Raqborski, M. PARASITISCHE ALGEN UND PILZE 1900. 1895. SYLLOGE FUNGORUM. V. 11, 753 pp. JAVA'S. Pt. 2, 46 pp. Buiten- Patavii. zorg Inst. Bot. Batavia. Ravenel, H. W. (M. C. Cooke, edit.) 1896. ELENCHUS FUNGORUM NOVORUM. 1882. fungi AMERICANI EXSICCATI. Fas- Hedwigia 35 (Repert. f. krypto- cicle 1: Nos. 1-100. gamische lit. Beibl. No. 7) : I-L. Rehm, H. 1888. ASCOMYCETEN. FASC. XIX. Hed- 1897. SYLLOGE FUNGORUM. V. 12, 1053 pp. wigia 27: 163-175.

1898. SYLLOGE FUNGORUM. V. 13, 1340 1889. EXOTISCHE ASCOMYCETEN. Hed- pp. Berolini. wigia 28: 295-303, illus. —— and Saccardo, D. 1897. beitrage zur pilzflora von sud- 1905. SYLLOGE FUNGORUM. V, 17, 991 pp. AMERiKA. m. Hedwigia 36: 366- Patavii. 380, illus. and Sydow, P. 1902. SYLLOGE FUNGORUM. V. l6, 1291 1898. beitrage ZUR pilzflora von SiJD- pp. Berolini. amerika. IV. Hedwigia 37: 189- 201, illus. 1926. SYLLOGE FUNGORUM. V. 24 (seC. 1), 703 pp. Pergola. 1900. beitrage zur pilzflora von sud- Sampson, K. AMERIKA. IX. Hedwigia 39: 221- 1933. the systemic infection of grasses 226. by epichloe typhina (pers.) Ricker, P. L. TUL. Brit. Mycol. Soc. Trans. 18: 1902. A preliminary list of MAINE FUN- 30-47, illus. GI. Maine Univ. Studies 3: 86 pp. and Western, J. H. 1941. DISEASES OF BRITISH GRASSES AND Rorer, J. B. 1911. A PRELIMINARY LIST OF TRINIDAD HERBAGE LEGUMES. 85 pp., illus. FUNGI. (Pp. 37-44). In Report Cambridge. of Mycologist for year ending SCHWARZE, C. A. March 31, 1911 (pt. 2). Trinidad 1917. THE PARASITIC FUNGI OF NEW JER- and Tobago Dept. Agr. Cir. 4, SEY. N. J. Agr. Expt. Sta. Bul. 62 pp., illus. 313, 226 pp., illus. 76 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Seaver, F. J. STATES. U. S. Bur. Plant Indus., 1910a. THE HYPOCREALES OF NORTH Soils, and Agr. Engin. Plant Dis. AMERICA—m. Mycologia 2: 48- Rptr. Sup. 163: 101-268. [Pro- 92, illus. cessed.] Stevens, F. L. 1910b. HYPOCREACEAE. North Amer. 1917. porto rican fungi, old and new. 31-56. Flora 3 (pt. 1): 111. Acad. Sci. Trans. 10: 162-218, illus. 1911. THE HYPOCREALES OF NORTH AMERI- CA—IV. Mycologia 3: 207-230, 1927. NEW TROPICAL FUNGI, Mycologia illus. 19: 231-238, illus. and Chardon, C. E. Stevenson, J. A. 1926. SCIENTIFIC survey OF PORTO RICO 1918. A CHECK LIST OF PORTO RICAN FUN- AND THE VIRGIN ISLANDS. N. Y. GI AND A HOST INDEX, PortO Rico Acad. Sci. 8 (pt. 1), 208 pp. Dept. Agr. Jour. 2: 125-264. Seymour, A. B. SUNDARARAMAN, S. 1929. HOST INDEX OF THE FUNGI OF 1921. ON THE PRESENCE OF AN EPHELIS NORTH AMERICA, 732 pp. Cam- AND A BALANSIA ON A COMMON bridge, Mass. FODDER GRASS AT CENTRAL FARM.

• and Earle, F. S. coiMBATORE. Madras Agr. Dept. 1899. ECONOMIC FUNGI [Exsiccati.] Fas- Yearbook 1920-21: 46-48. cicle 11: Nos. 251-400. Cam- Sydow, H. bridge, Mass. 1914. beitrage zur kenntnis der pilz- Spegazzini, C. flora des sudlichen ostindiens II, Ann. Mycol. 12: 484-490. 1885. fungi guaranitici, pugillus i. — Soc. Cient. Argentina An. 19: 34- 48, 91-96, 241-265. 1930. FUNGI VENEZUELANi. Ann. Mycol. 28: 29-224. and Sydow, P. 1886. FUNGI GUARANITICI. PUG. I. SoC. Cient. Argentina An. 22: 186- 1916. FUNGI AMAZONICI A CL. E. ULE 224. LECTI. Ann. Mycol. 14: 65-97, illus. Theiszen, F., and Sydow, H. 1887. FUNGI GUARANITICI. PUGILL. I. Rev. Mycol. 9: 89-98. 1914. dothideazeen studien. Ann. My- col. 12: 176-194. 1891. FUNGI GUARANITICI NONNULLI NOVI and Sydow, H. V. CRiTici. Rev. Argent. Hist. 1915. die dothideales. Ann. Mycol. 13: Nat. 1: 101-111, 168-177, 398- 149-746, illus. 432. Thorold, C. a. 1931. a further preliminary list of in- 1910. MYCETES ARGENTINENSES SER. V. TRINIDAD FUNGI. 30 pp., plus Buenos Aires Mus. Nac. de Hist. dex. Port-of-Spain. Nat. An. 20: 329-467, illus. ThUMEN, F. DE. 1879. MYCOTHECA UNIVERSALIS [Exsicca- ti.} Century 11: Nos. 1001- 1912. MYCETES ARGENTINENSES SER. VI. 1100. Buenos Aires Mus. Nac. de Hist. Tracy, S. M., and Earle, F. S. Nat. An. 23: 1-130, illus. 1895. MISSISSIPPI FUNGI. Miss. Agr. Expt. Sta. Bui. 34: 80-122. 1919. RELIQUIAE MYCOLOGICAE TROPICAE Trelease, W. FT FUNGI COSTARICENSES NON- 1887. the fungi of forage plants. NULLI. Acad. Nac. Cient. Cordo- (Pp. 413-431). In Beal, W. J.. ba Bol. 23: 365-609, illus. Grasses of North America for Farmers and Students. 457 pp. 1921. DESCRIPCION DE HONGOS MIRMECO- Lansing. (Reprinted in 1896, New FiLOS. Rev. Mus. La Plata 26: York.) 166-174, illus. TULASNE, L. R. 1851. NOTE SUR L'ERGOT DU SEIGLE, SCLER- 1922. FUNGI PARAGUAYENSES. BuenOS OTiUM CLAvus DC, Compt, Rend. Aires Mus. Nac. de Hist. Nat. An. Acad. Sci. [Paris] 33: 645-647, 31: 355-450, illus. illus. Sprague, R. 1946. ROOTROTS AND LEAFSPOTS OF GRAINS 1853. MEMOIRE SUR L' ERGOT DES GLUMA- AND GRASSES IN THE NORTHERN CEES. Ann. des Sci. Nat., Bot. GREAT PLAINS AND WESTERN (Ser. 3) 20: 5-56, illus. BALANSIA AND THE BALANSIAE IN AMERICA 77

Ule, E. Weiss, F. A. 1905. MYCOTHECA 3RASILIENSIS. [Exsicca- 1943. CHECK list revision. U. S. Bur. ti.} Century 1: Nos, 1-100. Plant Indus., Soils, and Agr. En- Berlin. gin. Plant Dis. Rptr. 27: 175-184, Underwood, L. M., and Earle, F. S. 188-193. [Processed.] 1897. A PRELIMINARY LIST OF ALABAMA Sta. FUNGI. Ala. Agr. Expt. Bui. 1945. CHECK list revision. U. S. Bur. 80: 113-283. Plant Indus., Soils, and Agr. En- ViEGAS, A. P. gin. Plant Dis. Rptr. 29: 68-76, 1944a. suMMARio alguns fungos do bra- 132-139, 576-583. [Processed.] SiL II. Bragantia 4, 392 pp., illus. Wernham, C. C. 1944b. alguns fungos esparsos brasil- 1942. epichloe typhina on imported EIROS. Revista Agron. 7: 367- FESCUE seed. Phytopathology 32: 383, illus. 1093. Vincens, F. West, E. 1917. rf.cherches organogeniques sur 1941. NOTES on FLORIDA FUNGI. U. My- quelques hypocreales. Faculte cologia 33: 38-49, illus. des Sci., Paris, Ser. A, No. 800, 166 pp., illus. (Thesis, D. Nat. 1945. NOTES ON FLORIDA FUNGI. III. My- Sci. ) Lons-le-Saunier. cologia 37: 65-79, illus. Weber, G. F. 1924. EPHELIS MEXICANA FR., BALANSIA Winter, G. (pk.) hypoxylon atk. on sand- 1887. REPERTORIUM RABENHORSTII FUNGI bur, echinatus l.) (cenchrus EUROPAEI ET EXTRA EUROPAEI EX- Phytopathology 14: 66. SICCATI, CURA G. WINTER, CENTURIA Weese, J. XXXV ET XXXVI. Hedwigia 26: 24- 1919. iJBER sphaeria epichloe kunze. 35. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. (Abt. 1) 128: 715- ZuCK, R. K. 719, illus. 1947. SIMPLIFIED PERMANENT ACETO-CAR- Weigelt, C. E. von. MINE SMEARS WITH A WATER- 1837. [fungi exsiccati.} Not numbered, MISCIBLE MOUNTANT. Stain distributed by Gustav Kunze. Technol. 22: 109-110.

HOST INDEX'*

Andropogon bkornis L Balansia henningsiana. A. leucostachyus. H.B.K B. henningsiana. A. scoparius Michx B. epichloe, B. henningsiana. A. virginicus L B. henningsiana. Andropogon sp B. claviceps, B. epichloe, B. henningsiana, B. strangulans, B. strangulans forma discoidea. Aristida adscensionis L B. aristidae. A. cognata Trin. & Rupr B. hemicrypta. A. dichotoma Michx B. aristidae. A. jendleriana Steud B. hemicrypta, A. glauca (Nees) Walp B. hemicrypta. A. intermedia Scribn. & Ball B. aristidae. A. longespica Poir B. aristidae. A.. oUgantha Michx B. aristidae. A. portoricensis Pilger B. aristidae. A. purpurascens Poir B. aristidae.

A. refracta Griseb . B. aristidae. A. setifolia H.B.K B. aristidae. A. spiciformis Ell B. aristidae. A. stricta Michx B. aristidae. A. tenuispica Hitchc B. aristidae. A. vilfifolia Henr B. aristidae. A. virgata Trin B. aristidae.

" In attempting to catalog the hosts for each species under consideration it was necessary to evaluate all available records, whether in published accounts or on herbarium labels. Since it was inevitable that the recorded names of host plants were sometimes suspect, every effort has been made to correct anj? seeming mis- identification. This has been possible where herbarium specimens contained suitable diagnostic features. In cases where specimens were not available or where host materials were inadequate, the professional standing of the taxonomist was necessarily a standard. Sometimes the evidence of comparable specimens provided helpful indications of correct host identities. In case of doubt the host name is followed by a question mark '"?". The nomenclature of the Gramineae is in general agreement with that of Hitchcock (1935. 1936). 78 AGRICULTURE MONOGRAPH 4, U. S. DEFT. OF AGRICULTURE

Aristida sp B. aristtdae, B. hemicrypta. Axonopus affinis Chase B. sirangulans. A. compressus (Swart2) Beauv B. sirangulans. Bambusa arundinacea Retz Balansiopsis gaduae. Bambuseae (indet.) B. pilulaejormis. Bouteloua curtipendula (Michx.) Torr Balansia sirangulans forma discoidea. Calamagrostts canadensis (Michx.) Beauv B. epichloe. C. inexpansa A. Gray B. epichloe.

Calamagrosiis sp. . -S. epichloe. Cenchrus echinatus L B. ohiecia. C. gracillimus Nash B. obiecta. Cenchrus sp B. obiecta. Chloris disiichophylla Lag B. sirangulans forma discoidea. C. petraea Swart2 B. epichloe. C. polydaciyla (L.) Swart2 B. sirangulans forma discoidea. Chloris sp B. epichloe, B. henningsiana, B. sirangulans forma discoidea. Chusquea uniflora Steud Balansiopsis gaduae. Chusquea sp Balansia chusqueicola, B. linearis. Cienium aromaiicujn (Walt.) Wood. B. epichloe. Cynodon daciylon (L.) Pers B. obtecia. Cyperaceae (indet.) B. cyperacearum. Cyperus diffusus Vahl B. cyperacearum. C. ovluaris (Michx.) Torr B. cyperacearum. C. reirorsus Chapm B. cyperacearum. C. strigosus L B. cyperacearum. C. vegeius ? B. cyperi. C. virens Michx B. cyperi. Cyperus sp B. cyperi, B. cyperacearum. Danthonia compressa Austin Atkinsonella hypoxylon. D. sericea Nutt (?) A. hypoxylon. D. spicata (L.) Beauv A. hypoxylon.

Elyonurus barbiculmis Hack. (?) . . A. hypoxylon. Eragrosiis capillaris (L.) Nees .... Balansia epichloe. E. hirsuia (Michx.) Nees B. epichloe. E. refracia (MuhL) Scribn B. epichloe. E. irichocolea Hack. & Arech B. epichloe. Eragrosiis sp B. epichloe. Eriochloa punciaia (L.) Desv Ephelis japonica. Eriophorum virginicum L. (?) Atkinsonella hypoxlon. ' Eusiachys (sp.), cf. Chloris spp. Festuca ovina L. (?) A. hypoxylon. Gramineae (indet.) cf. end of list. Guadua tagoara (Nees) Kunth .... Balansiopsis gaduae. Guadua sp B. gaduae. Gymnopogon arnbiguus (Michx.) B.S.P Balansia henningsiana, B. epichloe. Gymnopogon sp B. straugulans forma discoidea. Hymenachne amplexicaulis (Rudge) Nees n. sirangulans. J Jchnanthus candicans (Nees) Doell. B. sirangulans forma discoidea. J I. leiocarpus (Spreng.) Kunth B.subnodosa. I. nemorosus (Swartz) Doell B. subnodosa. I. pallens (Swartz) Munro B. subnodosa. I. riedelii (Trin.) Doell B. diadema = Claviceps diadema. I. tenuis (Presl) Hitchc. & Chase . . B.subnodosa. I. veriicillatus Mez B. diadema = Claviceps diadema.

Jchnanthus sp -. B. diadema = Claviceps diadema. Lasiacis ligulata Hitch. & Chase .... B. sirangulans forma discoidea. Leersia grandiflora (Doell) Pro- doehl B. ambiens. Luziola peruviana Gmel B. pallida. ..

BALANSIA AND THE BALANSIAE IN AMERICA 79

Merostachys speciosa Spreng B. linearis. Microstachys speciosa = error for Merostachys cf B. linearis. Olyra ciliatifolia Raddi B. ambiens. O. micrantha H.B.K B. linearis. Olyra sp B. ambiens, B. linearis. Oplismenus hirtellus (L.) Beauv. ., B. strangidans forma discoidea. O. setarius (Lam.) Roem. & Schult. B. strangidans forma discoidea.

Orthoclada laxa (L. Rich.) Beauv. . . Balansiopsis asclerotiaca. Orthoclada sp Balansia pallida var. orthocladae = Claviceps orthocladae. Panicum aciculare Desv B. strangulans. P. affolyroides (?) B. strangulans. P. agrostoides Spreng B. henningsiana, B. epichloe. P. anceps Michx B. henningsiana. P. boliviense Hack B. strangulans, B. strangulans forma, discoidea. P. cervicatum Chase B. strangulans. P. clandestinum L B. strangulans. P. commonsianum Ashe B. strangulans. P. commutatum Schult B. strangulans. P. cyanescens Nees B. strangulans. P. depauperatum Muhl B. strangulans. P. dichotomiflorum Michx B. henningsiana. P. ghiesbreghtii Fourn B. claviceps. P. haenkeanum PresI B. strangulans forma discoidea. P. hemitomon Schult B. strangulans. P. huachucae Ashe B. strangulans. P. implicatum Scribn B. strangulans. P. laxum Swartz B. strangulans. P. lindheimeri Nash B. strangulans. P. linearijolium Scribn B. strangulans. P. lucidum Ashe B. strangulans. P. maximum Jacq B. claviceps. P. meridionale Ashe B. strangulans. P. microcarpon Muhl B. strangulans. P. millegrana Poir B. strangulans forma discoidea. P. nitidum Lam B. strangulans. P. pilosum Swartz B. strangulans, B. subnodnsa. P. polygonatum Schrad B. strangulans.

P. proliferum Lam ; B. henningsiana.

P. scabriusculum Ell. . . B. strangulans. P. schiffneri Hack B. henningsiana, B. strangulans. P. scoparium Lam B. strangtdans. P. scribnerianum Nash B. strangulans. P. stipitatum Nash B. henningsiana. P. trichanthum Nees B. subnodosa. P. tsugetorum Nash B. strangidans. P. virgatum L B. henningsiana. P. zizanioides H.B.K B. strangulans. Panicum sp B. claviceps, B. epichloe, B. henningsiana, B. strangulans, B. strangulans forma discoidea, B. subnodosa, B. diadema = Claviceps diadema, B. pallida var. orthocladae = Claviceps orthocladae. Pariana sp B. linearis. Paspalwm ciliatijolium Michx Balansiopsis pilulaeformis, Balansia henningsiana. P. debile Michx Balansia pilulaeformis. P. pubescens Muhl B. pilulaeformis. P. schaffneri Griseb., erroneous report (?) for Panicum schiffneri Hack, (cf.) Paspalum sp Balansia epichloe, B. diadema = Claviceps, B. strangulans, Balansiopsis pilulaeformis. Poa (?) sp. (probably Eragrostis

sp. ) Balansia epichloe. Sacciolepis striata (L.) Nash B. strangulans. 80 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Scler'm bracteata Cav B. cyperacearum.

Setaria geniculata (Lam.) Beauv. . . B. claviceps.

S. lutescens (Weigel) F. T. Hubb. . . B. claviceps. S. macrostachya H.B.K B. claviceps. S. palmifolia (Willd.) Stapf B. claviceps. S. poiretiana (Schult.) Kunth B. claviceps. S. setosa (Swartz) Beauv B. obtecta. Setaria sp B. claviceps. Sorghum vulgare Pers B. claviceps. Spartina pectinata Lk., doubtful report of Atkinsonella hypoxylon. hypoxylon. Sporobolus indicus (L.) R. Br Balansia epichlo'e. S. poiretii (Roem. & Schult.) Hitchc B. epichlo'e. Stipa leucotricha Trin. & Rupr Atkinsonella hypoxylon var. texensis. S. viridiila Trin Atkinsonella hypoxylon var. texensis. Stipa sp Atkinsonella hypoxylon var. texensis. Thrasya petrosa (Trin.) Chase .... Balansia epichlo'e. Trichloris mendocina (Phil.) Kurtz. B. strangulans forma discoidea. Triodia jiava (L.) Smyth B. ep'tchloe. Uniola laxa (L.) B.S.P Balansiopsis pilulaejormis. Uniola sp B. pilulaejormis. Gramineae (indet.) Atkinsonella hypoxylon, Balansia claviceps. B. epichlo'e, B. henningsiana, B. linearis, B. strangulans, B. strangulans forma discoidea, Balansiopsis pilulaejormis. BALANSIA AND THE BALANSIAE IN AMERICA 81

INDEX OF FUNGUS NAMES [Names of Balansiae accepted as valid are printed in boldface.]

Page Page Atkinsonella 48 C. flavella 29 A. hypoxylon 49 C. strangulans = Balansia strangulans 29 A. hypoxylon var. texensis 54 Dothichloe 13, 15 Balansia 13 D. aristidae = Balansia aristidae .... 44 B. ambiens 46 D. atramentaria = Balansia epichloe 40 B. aritidae 43 D. atramentosa = Myriogenospora at- B. asclerotiaca = Balansiopsis asclero- ramentosa 59 tiaca 58 D. cyperacearum = Balansia cypera- B. ckusque'tcola 61 cearum 39 B. claviceps 17 D. discoidea = B. strangulans forma B. clarJceps var. jasciculata = B. discoidea 33 claviceps 17 D, epichloe = Balansia epichloe .... 40 B. clarnla = Balansiopsis pilulae- D. gaduae = Balansiopsis gaduae ... 55 formis 56 D. granulosa = B. strangulans forma B. typeri 26 discoidea 33 B. c\'peracearum 39 D. hemicrypta = B. hemicrypta 45 B. diadem a = Claviceps diadema. ... 59, 60 D. hypoxylon = Atkinsonella hy- B. discoidea = B. strangulans forma poxylon 36, 49 discoidea 33 D. limitata = Balansia epichloe .... 40 B. epichloe 40 D. linearis = Balansia linearis 47 B. gaduae = Balansiopsis gaduae. ... 55 D. nigricans = Balansia strangulans. 30 B. hemicrypta 45 D. strangulans = Balansia strangu- B. htnningsiana 36 lans 30 B. hypoxylon = Atkinsonella hypoxy- D. subnodosa = Balansia subnodosa. 35 lon 49 Dothidea atramentaria = Balansia B. linearis 47 epichloe 40 B. mexicana = B. obtecta 21 D. atramentaria forma Eragrosiidis-ca- B. obtecta 21 pillaris = Balansia epichloe .... 40 B. pallida 28 D. pilulaeformis = Balansiopsis pilu- B. pallida var. orthocladae = Clavi- laeformis 56 ceps orthocladae 60 D. pitulaeformis = Balansiopsis pilu- B. pilulaejormis = Balansiopsis pilulaeformis 56 laeformis 56 D. vorax 3, 14, 50 B. redundans = Balansiopsis pilu- Ephelis 13, 15 laeformis 56 E. borealis = Atkinsonella hypoxylon 49

B. regularis = Balansiopsis gaduae. . 55 E. japonica 61 B. strangulans 29 E. mexicam = Balansia obtecta .... 21 B. strangulans forma discoidea 33 E. texensis = Balansia cyperi 26 B. subnodosa 35 E. trinitensis = Balansia claviceps ... 17 B. irichloridis = B. strangulans forma Epichloe 1, 12 discoidea 33 E. atramentosa = Myriogenospora at- B. trinitensis =: B. claviceps 17 ramentosa 59 Balansiella 60 E. bertonii 12, 13 B. orthocladae = Claviceps orthocla- E. hypoxylon = Atkinsonella hypoxy- dae 60 lon 49 '= Balansiopsis 54 E. nigricans Balansia strangulans . . 29 Balansiopsis asclerotiaca 58 E. Strangulans = Balansia atrangulans 29 B. gaduae 55 E. typhina 13 B. pilulaeformis 56 E. volkensii 14 Botryosphaeria epichloe 40, 42 Hyalodothis 13, 14 Claviceps 10, 11, 12 H. clavus = Epichloe volkensii 14 C. balansiaides = C. flavella 29 Hypocrea atramentosa = Myriogeno- C. diadema 60 spora atramentosa 36, 59 C. flavella 29 H. atramentaria = Balansia epichloe. 40 C. pallida = C. flavella 29 H. cyperacearum = Balansia cypera- C. pallida = Balansia pallida 28 cearum 39 C. pallida var. orthocladae = C. ortho- H. hypoxylon = Atkinsonella hy- cladae 60 poxylon 49 C. philipii = Balansia claviceps .... 17 H. subviridis = Balansia henningsi- Cordyceps 12 ana 36 82 AGRICULTURE MONOGRAPH 4, U. S. DEPT. OF AGRICULTURE

Page Page Hypocrella atramentosa = Myriogeno- O. gaduae = Balansiopsis gaduae ... 55 spora atramentosa 59 O. henningsiana = Balansia henning- H. cyperacearum = Balansia cypera- siana 36 cearum 39 O. hypoxylon = Atkinsonella hy- H. hypoxylon = Atkinsonella hypoxy- poxylon 49 lon ' 49 O. linearis = Balansia linearis 47 H. hypoxylon var. pilulaejormis =^ O. rhaphidospora = Balansia linearis 47 Balansiopsis pilulaeformis .... 56 O. vorax 14 H. sem/amplexa 40 O. vorax var. atramentaria = Balansia H. stibviridJs = Balansia henningsi- epichloe 40 ana 36 O. vorax var. pilulaejormis = Balan- Linearistroma 14 siopsis pilulaeformis 19, 56, 58 Linearistroma lineare = Balansia line- Poroniopsis 11 aris 47 P. bruchi 12 Munkia 12, 61 Shropshira ^= Alunkia 12, 61 M. martyris 61 S. chusqueae 61 Myriogenospora atramentosa 59 M. paspali = M. atramentosa 59 Sphaceliopsis 14 Neomunkia = Shropshiria 12, 61 5. cyperi cola ^= Balansia cyperi 26 Ophiodothis 13, 14 Sphaeria cyperacearum = Balansia cy- O. aristidae = Balansia aristidae .... 44 peracearum 39 O. atramentosa = Myriogenospora at- S. epichloe = Balansia epichloe 40, 42 ramentosa 59 S. vorax 50 O. atramentosa var. aristidae ^ Bal- Ustilaginoidea 12, 61 ansia aristidae 44 U. setariae 61 O. atramentosa var. cyperi = Balansia cyperacearum 39 U. strumosa 61 O. atramentosa var. pilulaeformis = Xylaria clavulus = Balansiopsis pilu- Balansiopsis pilulaeformis 56 laeformis 56

•i^U. S. GOVERNMENT PRINTING OFFICE: 1950

Agriculture Monograph 4, U. S. Department of Agriculture Plate 1

Balans.a claviceps. A, Part of type ex Herb. Spegazzini (W). X h B, Detail of A, showing stipitate ascostromata and remnants of ephelidial fructifications on black hypothallus (W). X 12. C, Twinned specimen (B. claviceps war. fasciculata Speg. ined.) ex Herb. Spegazzini (W). X 1. Agriculture Monograph 4, U. S. Department of Agriculture

ialonsia daviceps. A, Paratype of Ephelis mexicana in Herb. Berkeley (K), showing black hypothalli

bearing sessile ephelidial fructifications. X 1 . 6, At left, part of type of E. trinitensis (K), showing blackish hypothailus bearing sessile ephelidial fructifications; at right, part of type of 6. trinitensis (K), showing blackish hypothailus bearing stipitote oscostromata and some sessile ephelidial fructifications. X 1. C, Detail of 6. X 6.5. D, Apex of culm of Sorghum vulgare (1921, S. C, Bruner) covered by block hypothailus bearing ephelidial fructifications (W). X 1. Agriculture Monograph 4, U. S. Department of Agriculture

Balansia obtecta. A, Infected plant of Cenchrus echinatus In greenhouse at Cambridge, Mass., grown from an infected seed that had matured after stigma inoculation with ephelidial conidia; the sheath of each apical leaf enfolds a pseudomorph consisting of an aborted inflorescence surrounded by sclerotic tissue (hypothallus) of the fungus. X 0.7. 6, Pseudomorph from an infected plant of Cenchrus echinafus with enfolding leaf sheath. X 1. C, Pseudomorph with part of hypothallus removed, revealing the aborted (mummified) inflorescence. X 1. D, Type of Ephelis mexicana in Herb. Berkeley (K). X 1. E, Detail of D, showing characteristic blackish hypothallus bearing sessile ephelidial fructifications. X 12. Agriculture Monograph 4, U. S. Department ot Agriculture , Balansia obtecta: EphelididI fructification with cirrhi developed from hypothallus (Florida material) after 48 hours in moist chamber, at Washington, D. C, 1931 (W). X 10. B, B. obtecto: Hypothallus bearing a stipitate ascostroma after 8 weeks in moist chamber (Florida material) at Cambridge, Mass., 1931 (W). X 1. C, B. obtecta: Detail of B, showing ascostroma with stipe emergent from site of an ephelidial fructification. X 6.5. D, B. obtecta.- Sessile hirsute ephelidiol fructifications and stipitate ascostromata produced in moist chamber at Gainesville, Flo., 1939, by G. F. Weber, ex type of B. obtecta (W), X 6.5. E, 6. pallida: Yellow ascostromata arising from yellow hypothallus on florets of Luziola peruviana (F). X 13. F, Balansia cyperi: White hypothallus bearing light, orange-colored ephelidial fructifications on inflorescence of Cyperus sp., Texas, 1913, D. H. Sanders (W). X 1. G, B. cyperi: Black ascostromata over hypothallus on Cyperus virens ex type (W). X 1- Agriculture Monograph 4, U. S. Department of Agriculture

I A, Halans'ia strangulans: Type of Cordyceps sfrangulans in Herb. AAontagne, Paris, showing black ascostromota one (at right) with whitish border. X ^. B, B. strangulans: Type collection (syntype) of Epichloe nigricans ex Balansa, Plantes du Paraguay No. 3741 in Herb. Paris, showing black oscostroma with whitish border (hypothallus). X 1. C, B. sfrangulans: Infected plant of Sacciolepis striata from Florida in culture at Cambridge, Mass., 1931, showing black ascostromota bordered by white remnants of hypothalli and ephelidial fructifications surrounding internodes just beiov/ nodes. X 1. D, B. strangulans: Detail of C. E, B. strangu/ans. Black ascostromota on internodes of Panicum clandestinum, Virginia, 1931, showing suppression of inflorescence (center) and (at right) aberrant discoid stroma on remnant of hypothallus and ephelidial fructifications. X 1. F, B. strangulans: Photomicrograph of fragment of leaf tissue of infected Sacciolepis striata, showing phloxine-stained hyphae. X 3,000. G, B. strangulans forma discoidea: Ascostromota on internodes of Chloris polydactyla, Venezuela, 1937, Muller No. 2,038 (V/). X 1. H, B. strangulans vormo discoidea: Discoid oscostromata on internodes of Chloris sp. ex Balansa, Plantes du Paraguay in

Herb. Paris. X 1. ', Balansia subnodosa: Circlet of convergent buttonlike ascostromota just below nodes of Panicum sp. in Rabh., Fungi Europaei No. 3563 (V^). X 1. J, B. subnodosa.- Detail of I. X 4. Agriculture Monograph 4, U. S. Department of Agriculture ^™F f

/J/cAo, B (/*V

\ \ \

Balansia benningsiana. A, Mature ascostromata on leaves of Andropogon sp., Alabama, paratype of Hypocrea atramentosa :n Herb. Curtis (F). X 1. 6, Copy of Moller's illustration (type) of Ophiodoihis henningsiana, showing section through oscostroma on leaf on Andropogon sp. X 15. C, Ephelidial fructification on unknown grass. South Carolina, type of Hypocrea subviridis in Herb. Curtis (F). X 1. D, Mature ascostromata on leaves of Andropogon scoparius, Louisiana, 1931, R. W. Davidson (W). X 1. C, Mature ascostromata on leaves of Andropogon scoparius, Va., 1941 (F), showing black ascostromata bordered by whitish remnants of ephelidial fructifications. X 4. Agriculture Monograph 4, U. S. Department of Agriculture Plate 7

5iiro»o"»-

B D

A, Balansia cyperacearum: Black ascostromata on leaves of unknown sedge, Surinam, type in Herb. Curtis (F). X 1. B, 6. cyperacearum: Black ascostromata on culms of Scleria bracfeata, Venezuela, 1939, Chardon and Alston No. 2785 (W). X 1. C, Balansia epichloe-. Black ascostromata on adaxial leaf surfaces of unknown grass, Surinam, 1827, ex type (syntype) of Dothichloe epichloe in Weigelt's exs. (W). X 1. D, B. epichloe: White ephelidial fructifications and black ascostromata on adaxial leaf surfaces of Sporobolus poiretii, Alabama, 1929, J. L. Seal (type of Doihichloe limitafa (W). X 1. f/ Balansia aristidae: Black ascostromata with characteristic ostioiar pro- tuberances on Aristida purpurascens, Virginia, 1941, W. W. Diehl (W). X 3.5. F, B. aristidae: Ex type, Alabama, G. F. Atkinson No. 2145 (F). X 1. Agriculture Monograph 4, U. S. Department of Agriculture Plate 8 A, Balansia hemicrypfa: On Arist.da glauca, causing dwarfed and sterile spikelets (at right) and comparable normal spikelets at left, Texas, 1924, B. C. Tharp, type (W). X 1. B, B. hemicrypfa: Detail from above (right), showing dwarfed and sterile florets with extruding ephelidial conidia as the whitish exudate. X 6. C, B. hemicrypfa: Photomicrograph of free-hand section of part of floret of infected A. glauca, showing conidiiferous hyphal mat within. X 120. D, B. hemicrypfa: Photomicrograph of free-hand section of part of floret, showing needle^like ephelidial conidia lining lemma. X 192. E, 6. hemicrypfa: Sessile black ascostromata and ephelidial fructifications on Leersia grandiflora, Brazil, 1933, A. S. Muller No. 415 (W). X 1. F, Balansia ambiens: On Olyra sp., Brazil, copy of Moller's illustration of type. X 0.75. G, B. ambiens: Ephelidial fructifications, detail of E. X 10. H, Ephelis japor^ica (?) on Eriochloa punctata, Puerto Rico, 1929, C E. Chardon and J. A. B. Nolla (W). X 1. Agriculture Monograph 4, U. S. Department of Agriculture Plate 9

A, Balansia linearis-. Apex of shoot of Pariana sp., showing normal large leaves (below) and (above) inrolled, dwarfed leaves bearing black ascostromota, Brazil, 1934, J. R. Swollen No. 3229 (W). X 1- Bf Detail of A, showing linear black ascostromota emerging from leaves. X 1. C, Apex of shoot of Chusquea sp., showing much-dwarfed apical leaves bearing very small ascostromota, Brazil, 1925, A. Chase No. 8496 (W). X.l- D, Linear ascostromota on inrolled leaves of Chusquea sp., Brazil,

ex type (?) in Herb. Theiszen (F). X 1. Agriculture Monograph 4, U. S. Department of Agriculture Plate lO

A, Atkinsonella hypoxylon-. Infected culm of Danthonia spicata with white typhodial layer and one ephelidial fructification over hypothallus at apex of culm surrounding aborted inflorescence, Virginia, 1929. X 1. 6, A. hypoxylon: Detail of A. X 8. C, A. hypoxylon: Photomicrograph

of longitudinal section of culm of infected D. spicafa at a point of 1 cm. below hypothallus, showing phloxine-stained hyphae of the endophyte, Maryland, 1946. X 2,000. D, A. hypoxylon: Subsessile oscostromato on hypothallus covering aborted inflorescence of D. spicafa culture in greenhouse, Cambridge, Mass., 1931. X 1. E, A. hypoxylon: Photomicrograph of immature perithecia as squeezed out of oscostromato. X 100. F, A. hypoxylon: Normal healthy plant of D. spicafa transplanted when dormant and forced in greenhouse, Cambridge, Mass., 1932. X 0.16. G, A. hypoxylon: Comparable healthy-appearing plant of D. spicata, bearing endophytic mycelium of A. hypoxylon, transplanted when dormant and forced under like conditions and for same length of time as healthy plant (F), Cambridge, Mass., 1932. X 0.16 (Note dwarfed condition and absence of inflorescences.) H, Balansiopsis gaduae: Small black oscostromola at nodes of fasciated and dwarfed Bambusa arundinacea, Brazil, 1930. X 1. 1

Agriculture Monograph 4, S. Department of Agriculture U. PLATE 1

, Balanciopsis pilulaeformis: Part of type of Xylaria clavulus (sub Hypoxylon) in Herb. Curtis (F) showing blacic subsessile ascostromata. X 1.3. B, B. pHulaeformis: Detail of part of type of Xylaria clavulus (K), duplicate of above, showing a substipitate ascostroma. X 13. C, B. pilulaeformis: Part of type of Dothidea pilulaeformis (K), showing hypothallus and ascostromata on tip of culm and subtending apical leaf. X 1. D, B. pilulaeformis-. Detail of C. X 5. E, 6. pilulaeformis: On Uniola laxa, S. M. Tracy No. 197 (N), showing yellow hypothallus and black ascostromata. X 5. F, 6. pilulaeformis: On Paspalum ciliafifolium from Florida culture in g.'eenhous3 a? Arilngfon, Va., 1938. (At left, ascostromata emerging from sheath of terminal leaf, at right (below) ncrmol inflorescence emerging from enclosing leaf sheath of infected plant. Both left and right figures show deformed terminal leaves locking ligules.) X 1. G, B. asclerotiaca: Small stipitate ascostroma on sheath of epical leaf of Orthcclcda laxa, Brazil, 1901, E. Ule Mycotheca Bras. No. 78, type

coll. (?) (W). XI.