Illinois Natural History Survey bulletin 5 NATUSAl KI5TC3Y SUT/Vf ?)Vf.C, ILLINOIS NATURAL .^ HISTORY MAY 10 1994 SURVEY LIBRARY Systematics of Leptosphaeria Species Found on the Rosaceae Sabine M, Huhndorf Illinois Natural History Survey Bulletin Volume 34, Article 5 May 1992 OEPOSITORY AT URBANa-CHAMPAIGN ILLINOIS NATURAL HISTORY SURVEY Systematics of Leptosphaeria Species Found on the Rosaceae Sabine M. Huhndorf Illinois Natural History Survey Illinois Natural History Survey Bulletin Volume 34, Article 5 May 1992 Illinois Natural History Survey. Lorin I. Nevling. Chief A Division of the Illinois Department of Energy and Natural Resources A catalog of the publications of the Illinois Natural History Surrey is available without charge from the address below. A price list and an order blank are included with the catalog. Illinois Natural History Survey Distribution Center Natural Resources Building 607 East Peabody Drive Champaign, Illinois 61820 Citation: Huhndorf, S.M. 1992. Systematics of Leptosphaeria species found on the Rosaceae. Illinois Natural History Survey Bulletin 34(5):479-534. Editor: John P. Ballenot Author's current address: The New York Botanical Garden. Bronx, NY 10458. US ISSN 0073-4918 Printed by Authority of the State of Illinois (X04 1 5- 1,200-5-92) Contents Acknowledgments iv Introduction 479 Materials and Methods 480 Discussion of Characters 480 Leptosphaeria Species Referable to the Pleosporales 482 Leptosphaeriaceae 482 Phaeosphaeriaceae 492 Lophiostomataceae 503 Leptosphaeria Species Referable to the Melanommatales 505 Leptosphaeria Species Referable to the Dothideales 508 Dothioraceae 508 Pseudosphaeriaceae 5 1 Leptosphaeria Species Referable to the Hymenoascomycetes 5 1 8 Clypeosphaeriaceae 5 1 8 Diaporthaceae 520 Amphisphaeriaceae 520 Species incertae sedis 525 \pYtend\\: Leptosphaeria Species Described from the Rosaceae 526 Literature Cited 532 Index to Taxa 534 Acknowledgments This research was supported by National Sci- ence Foundation grant BSR 87-00065 to C.A. Shearer, J.L. Crane, and D.L. Swofford and by the Floyd Ingersoll Fellowship, Department of Plant Pathology, University of Illinois, Fall 1989. The Friends of the Farlow Fellowship supported research on specimens at the Farlow Herbarium, and the H.H. Ross Memorial Grant from the Illinois Natural History Survey sup- ported research on specimens at the New York Botanical Garden. I would also like to acknowl- edge and thank the Morton Arboretum for the Baker Fellowship, which helped in the completion of the manuscript. I thank the curators of the herbaria for the material made available for study. I thank J.L. Crane, D.A. Glawe, and M.E. Barr Bigelow for reviewing the manuscript. This work was originally part of a dis- sertation submitted to the Graduate College of the University of Illinois at Urbana- Champaign in partial fulfillment of the re- quirements for the degree of Doctor of Phi- losophy in plant pathology. Introduction The genus Leptosphacria was established by Cesati and de Notaris ( 1 863) to include 26 species. The original description was superfi- cial by modem standards and relied primarily on ascospore characteristics to delimit the genus. Cesati and de Notaris described ascospores as oblong or fusoid, two- to many- celled, and hyaline becoming yellow to dark brown. Because other structural features were poorly defined, a wide range of ascomycetes has been included in this genus. The 1,689 taxa described in Leptosphacria (Crane and Shearer 1991 ) represent, according to current concepts of ascomycete classification, a mixture of Hymenoascomycetes and Loculoascomycetes. Crane and Shearer (1991) and Muller ( 1950) provide good reviews of the historical back- ground of the genus Leptosphacria. Holm (1957) and Shoemaker ( 1984a) consider a limited number of species. A large number of intergeneric transfers of Leptosphacria species have been made in the past 50 years. Five genera — Phaeosphaeria Miyake, Paraphacosphacria Eriksson, Nodulosphaeria Riess, Entodesmium Riess, and Ophioholus Riess—have become accepted repositories for many Leptosphacria species (Holm 1957: Leuchtmann 1984; Shoemaker 1976, 1984a,b). Paraphacosphacria. Nodulosphaeria, and Entodesmium are well defined by several morphological features, including ascocarp wall structure and ascospore characteristics (Shoemaker 1984b, Shoemaker and Babcock 1985). Phaeosphaeria \s scpa- rated from Leptosphacria by several morpho- logical characters and host specialization but has itself become a very large and diverse genus (Shoemaker and Babcock 1989b). Ophioholus species intergrade with long-spored species of Leptosphacria (Shoemaker 1976). For a key to genera variously allied with Leptosphacria, see Shoemaker ( 1984a) or Leuchtmann (1984). Holm (1957) considered 62 species of Leptosphacria and restricted the genus to those species most similar to the type of the genus, Leptosphacria doliolum (Pers.iFr.) Ces. & de Not. He emphasized the anatomy of the ascocarp wall and found that in most Leptosphacria species the wall consisted of thick-walled cells tenned scleroplectenchyma. He also emphasized the identity of the substrate and the structure of the ascocarp relative to the substrate in relegating a number of Leptosphacria species to other genera. Ex- cluded species were distributed in Nodulosphaeria. Phaeosphaeria. and Entodesmium. He gave considerable weight to the family of host plants in distinguishing these genera. Holm's ( 1957) scheme was accepted by Shoemaker ( 1984a) in his treatment of Canadian species of Leptosphacria. In recent years, Leptosphacria has been included in various orders in the Bitunicatae or the Loculoascomycetes. Luttrell ( 1973) placed Leptosphacria in the family Pleosporaceae of the Pleosporales. The single order Dothideales was used by von Arx and Miiller ( 1975) for all fungi classified as Loculoascomycetes; they retained Leptosphacria in the Pleosporaceae. Eriksson and Hawksworth ( 1986) classified the genus in the family Phaeosphaeriaceae of the Dothideales. Barr ( 1987a) placed Leptosphacria in the family Leptosphaeriaceae of the Pleosporales. Most recently, Eriksson and Hawksworth (1990) accepted the family Leptosphaeriaceae in the order Dothideales. Barr's classification of the Ascomycota ( 1983, 1987a) is employed in this dissertation because of its relationship to previous classification schemes (Luttrell 1973) and its detailed explanation of the relationship of taxa, Barr's orders and families are based upon coinbina- tions of developmental and morphological characteristics that reflect the widespread diversity within the group. Barr's classification provides a key to higher taxa and is practical iM) Illinois Natural History Survey Bulletin Vol. 34 An. 5 lor identification and for determining the placement of organisms not yet included in the scheme. The present research reassesses the taxonomic position of species of Leptosphaeria described from plants in the family Rosaceae. In doing so, this wori< emphasizes a concept of Leptosphaeria based on its type species. Because discrete groups of morphologically similar species were found on related hosts (Nixlulosphaeria on Compositae, Phaeo- sphaeria on Gramineae and Entodesmiitm on Leguminosae), a goal of the study was to determine whether a distinct group of related species could be found on Rosaceous hosts. Also, approaching the taxonomy of this large, complex genus on the basis of host family is a convenient way of dividing the large numbers of Leptosphaeria species into smaller, work- able groups. Fifty-five species described from Rosaceous hosts have been included in Leptosphaeria (see appendix). Efforts were made to locate and obtain type material for all of the described species. The names of herbaria providing material are abbreviated in this dissertation according to Index Herbariorum (Holmgren et al. 1990). When possible, type specimens were studied, supple- mented by other collections, and the species were redescribed and illustrated from the type material. Unfortunately, type material could not be located for all of the species; in such cases, descriptions were based on other material only when the resulting concept of the species was obviously in accordance with the original description. From this work, it became apparent that no discrete group of related species is specialized on the Rosaceae. The species studied were determined to represent a mixture of Loculoascomycetes and Hymenoascomycetes, and non-Leptosphaeria species were reassigned to more appropriate genera. Materials and Methods Several techniques were employed to assess character states of specimens. Fungi were observed microscopically in water mounts for details of asci, ascospores, hamathecium. centrum, and ascocarp wall surface, using bright-field and Nomarski differential interfer- ence contrast. India ink was used as a nesiative stain to reveal ascospore sheaths. Melzer's reagent (fJ.S g iodine, 1.5 g KI, 2(J.O g chloral hydrate. 20.0 ml distilled water) was used to observe amyloid reactions in unitunicate asci. Semipermanent mounts were made in lacto- phenol, and many of the photomicrographs were made from these mounts. Micrographs were made using Kodak T Max KXJ and Kodak Technical Pan film 2415 (Eastman Kodak Company, Rochester, New York). For scan- ning electron microscopy, dry ascocarps and substrate were used direct from the herbarium specimen, without any preparation. Specimens were viewed with an Amray 1830 scanning electron microscope. The order of septation in ascospores is given following Shoemaker { 19S4a). The sequence of septation is recorded chronologi- cally, with 1 being the first septum formed. A series of numbers separated by colons indicates the sequence in which the septa form. TTie first number represents the septum nearest the ascospore apex, and the last numljer is the septum nearest the base. Unusual or infrequent occurrences are recorded w ithin parentheses (Shoemaker 1984a). To observe details of ascocarp anatomy, semi-thin sections of the ascocarp were needed for light microscopy. Fixation and embedding techniques for plastic were developed to obtain good, uniform sections from dried fungal material in a relatively short period of time. The techniques for fixation and embedding are given in Huhndorf ( 1991 ). Discussion of Characters Semi-thin sections of ascocarps were used to assess characteristics of the ascocarp w alls. Wall characteristics have proved useful in taxonomic placement of species but may have limitations in the routine identification of specimens. It may be possible to observe this feature with hand sections or frozen microtome sections, as done by Shoemaker ( l^S4a). Ascocarp walls of cells radiating in surface view (textura prismatica) and thick-walled cells (scleroplectench\ ma) are good indicators within Leptosphaeria of alliance with L. doliolum. Wall cells in surface \ icw are seen easily from crush mounts. Semi-thin sections and scanning electron microscopy give a good indication of the relationship of the a.scocarp to the substrate. This character has ai.so proved May 1992 LfpiDsphaeiia Species on ilie Rosaceae 481 important in delimiting Leptosphaeria; in- cluded species have ascocarps that are superfi- cial or thai become superficial. Semi-thin sections also show ostiole structure, a charac- teristic that is important in related genera such as Nodulosphaeria. Ascospore characters, in combination with other characters, are useful for placement of species. Lepiosphaeria species have more or less fusiform, three- to multi-septate ascospores with some pigmentation. These character states are continuous rather than discrete, and in practice, divisions are made in a somewhat arbitrary manner. Also, ascospore pigmenta- tion varies within a species and may reflect age, environmental influences, and/or substrate. Ascospore wall ornamentation often is difficult to assess, especially when using type material, which frequently is old and in poor condition. In addition, type material is usually of uniform developmental stage and frequently is sparse and depleted by the activities of previous researchers. Ascospore wall ornamentation is difficult to assess with the light microscope because of the limits of resolution, except in cases for which there may be unusually distinctive surface ornamentation. Using the scanning electron microscope to resolve surface features requires sufficient material of an appropriate developmental stage, with the ascospores released from the asci and with surface detail not obscured by sheaths or mucilaginous material. Unfortunately, this situation does not usually exist in type material. In recent collections with different develop- mental stages it is easier to view ascospore wall ornamentation. Although the taxonomic significance of this character is unclear at this time, largely because of the difficulties in- volved in studying it, further studies would be useful. The importance of host specificity or substrate preference in delimiting species or genera needs further clarification. Whether substrate preference does occur and can be used as a taxonomic character can be demonstrated only by extensive collection and comparison of species from a variety of substrates, by com- parative study of the morphology of pure cultures of species obtained from a variety of substrates and grown under identical environ- mental conditions, and by determining experi- mentally the range of substrates on which isolates of species will grow and reproduce. The structure of the interascal filaments, part of what Eriksson ( 1 98 1 ) termed the hamathecium. plays an important role in distinguishing members of the Melanom- matales from those in the Pleosporales accord- ing to Barr's (1987a) classification. Only one member of the Melanommatales was found in this group of species, and the diagnostically important trabeculate pseudoparaphyses were difficult to recognize. It was placed in that group because of a combination of other characters. The distinction between cellular pseudoparaphyses (filaments that appear .septate at 450x magnification and branch and anastomose) and trabeculate pseudoparaphyses (thin filaments that do not appear septate at 450x magnification and branch and anasto- mose) seems to be clear only in certain cases; the pseudoparaphyses often appear as continu- ous, rather than discrete, character states. Trabeculate pseudoparaphyses can be mistaken for cellular ones if septa appear regularly and anastomoses are sparse. Ascus morphology, emphasized by Eriksson (1981), was not employed in this study because of the difficulty in observing modes of ascus dehiscence and details of ascus wall layers in type and other herbarium specimens. Also, in most cases, the use of stains to enhance ascus wall layers gave virtually no results. Leptosphaeria Species Referable to the Pleosporales All of the species in this chapter belong in the Pleosporales (sensii Barr 1987a) and share a combination of character states, any one of which may deviate somewhat for a particular taxon. These characters include cellular pseudoparaphyses, asci in a basal layer, a peridium that is usually pseudoparenchyma- tous. and bipolarly asymmetrical ascospores. The families included within the order are the Leptosphaeriaceae, Phaeosphaeriaceae, and Lophiostomataceae. Leptosphaeriaceae The Leptosphaeriaceae as defined by Barr (1987a) include five genera united by the characters of coelomycetous anamorphs, asci that are narrower and thinner-walled than in the Pleosporaceae, and ascocarp walls that consist of relatively large, thick-walled or sclero- plectenchymatous cells. The only genus in the family treated here is Leptosphaeria. Leptosphaeria V. Cesati & G. de Notaris, Commentario della Societa Crittogamologica Italiana. Milan 1:234. 1863. Nomen conser- vandum. Lectotype: Sphaeria doliolum C.H. Persoon: E.M. Fries. Ascocarps scattered or clustered, superficial or immersed beneath epidermis, becoming superficial as epidemiis is shed, conic globose, suhglobose or depressed, glabrous, papillate. Ascocarp wall often of radiating textura prismatica in surface view; in longitudinal section usually more than three cells thick, composed of sclcroplectenchyma. often with an external crust. Papilla central, short, some- times longer and hcaklikc. without hvalinc periphyscs. Pseudoparaphyses broad or narrow, septate, anastomosing, with or without guttules, .septal thickenings, and gelatinous coating. Asci bitunicate. numerous in a broad basal hymenium. cylindrical to clavate. short- stalked. 8-spored in most, with spores often biseriate or overlapping uniseriate. Ascospores fusiform, clavate, 3 or more septate, primary septum often median and often constricted, some shade of yellow or brown, appendages or sheath may or may not be present. Anamorphs coelomycetous where known (see Crane and Shearer 1991). The concept of Leptosphaeria accepted here is essentially that of Holm ( 1957). later adopted by Shoemaker (1984a) with some modification. The lectotypification of Z,. doliohtm (Shearer et al. 1990) represents a basis for circumscribing the genus. More emphasis is placed on sclcroplectenchyma found in the ascocarp walls than is considered important by Shoemaker (1984a). As Shoe- maker ( 1984a) noted, thin sections are essential to show this thick-walled cell structure in which the lumen of the cells is very small. Emphasis is also placed on the wall tissue arrangements in surface view . described using the textura types of Kort' ( 19.*i8). Species having a tissue arrangement similar to that of Z.. doliolum. of cells radiating in surt'ace view (textura prismatica), are regarded as most typical, but species w ithout this character state are not necessarily excluded. The position of the ascocarp relati\e to the substrate is \ er\ important, w ith included species having either superficial ascocarps or a.scocarps immersed beneath the epidermis and becoming superficial as the epidermis is shed. Because onh tyjje or herbarium specimens were studied, anamorphs were not considered because cultures could not be made from this dead material. Examination of 28 purported species of Leptosphaeria from the Rosaceae re\ealed five species that fit the adopted generic concept. The excluded species are treated follow ing these five species. May 1992 Lepiiisphat'iia Species on the Rosaceae 483 Key to species of Leptosphaeria on the Rosaceae la. Ascocarp wall cells radiate 2 lb. Ascocarp wall cells randomly oriented 4 2a. Ascocarp superficial or immersed becoming superficial, ascospores mature inside centrum 3 2b. Ascocarp immersed beneath host cuticle, ascospores mostly maturing on substrate surface L. itmhrosa 3a. Neck papillate, short conic, ascospores narrowly fusiform, 22-28 x 4—5 |im L. dnliohim 3b. Neck cylindrical, beaklike, ascospores broadly fusiform, 17-20 x 5-8 |im L. praetermissa 4a. Ascocarp immersed-subepidermal, wall pseudoparenchyma, or partly scleroplectenchyma, on leaves of Cercocarpus (provisionally retained within Leptosphaeria) L. cercocarpi 4b. Ascocarp erumpent to superficial, wall scleroplectenchyma, on Dryas L. dryadophila •Leptosphaeria doliolimi (C.H. Persoon:E.M. Fries) V. Cesati & G. de Notaris, Commentario della Societa Crittogamologica Italiana, Milan 1:234-235. 1863. See Figure I. = Sphaeria doliolum C.H. Persoon:E.M. Fries, Icones et Descriptiones Fungorum Minus Cognitorum, p. 39. 1800; E.M. Fries, Systema Mycologicum Sistens Fungorum 2:509. 1823. = Leptosphaeria rustica P.A. Karsten, Fungi Fenniae Exsiccati, Century 10, No. 964. Anno 1870. = Metasphaeria rustica (P.A. Karsten) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:157. 1883. Ascocarps scattered, sparse, immersed- subcuticular to erumpent, papillate, glabrous to tomentose at base, globose, flattened at base, 300-325 nm diameter, 250-300 nm high. Ascocarp wall of textura prismatica in surface view, composed of cells radiating from apex outward; in longitudinal section 45-55 |J.m thick at sides, 20-27 |im thick at base, com- posed of 12-15 layers of hyaline, polygonal, isodiametric, scleroplectenchymatic cells (3.6- 6.5 |im diameter), outer 2-3 layers somewhat brown-melanized, with an external brown- melanized crust. Papilla broadly rounded, not fully fomied. 130-135 |im high, 70-75 |im wide; wall 40-50 |am thick, composed of 15-20 layers of small, brown-melani/ed, thick-walled, polygonal, isodiametric cells (2.5-6.5 \im diameter), outer layers heavily brown- melanized, inner layers hyaline, surrounding a 20-25 |im wide immature circular ostiole formed from small hyaline cells, without periphyses. Pseudoparaphyses 1.0-2.0 |im wide, 150-175 \xm long (height of the ascocarp cavity), numerous, narrowly cellular, with guttule-like thickenings at septa, without gelatinous coating. Asci 90-125 x 5-9 |im, numerous, basal, cylindrical-clavate, with 8 overlapping uniseriate ascospores. Ascospores 22-28 X 4—5 jim, narrowly fusiform, with acute end cell shape, second cell slightly enlarged, straight to slightly curved, 3-septate, septa evenly distributed, order of septation 2: 1 :2, primary septum median and slightly con- stricted; light brown, guttulate, without appendages, surrounded by gelatinous material. Lectotype: As Sphaeria doliolum Pers., 910.270-650 (L). Exsiccatae: Finland: Tammela, on Spireae ulmaria L. {=Filipendida ulmaria Maxim.), Sept., Fungi Fenniae Exsiccati, Century 10, No. 964 (isotype of L. rustica, H, FH). Comments: The collections of /.. rustica, on stems of Spiraea ulmaria L. in both of the exsiccatae sets, were immature. No ascospores were found, and asci were only beginning to form. Karsten ( 1873) describes the ascospores as elongate-fusoid. subhyaline, uniseptate with several guttules in each cell, and 30-36 x 5-6 \im in size. He probably described immature spores. This information—along with the Illinois Natural History Survey Bulletin Vol. 34 An. 5 (. f Figure 1. Leptosphaciui dolutlum. a. SEM of erumpent ascocarp. xl25. b. longitudinal median section through ascocarp, xl50. c. longitudinal, median section through ascocarp wall. x950. d. ascus. x525. e. ascocarp wall surface with radiating cells. x5.S0. All from isotype of i.. rustica. radiating wall cells in suiface view, the sclcroplectenchytnatic wail cells in section, and the location of the ascocarp relative to the substrate—is consistent with the character of Z.. doliolunu the type of the genus. Leptosphaeria doUolum is found on many dead herbaceous stems, although not previously described from Spiraea. •Leptosphaeria dryadophila S.M. Huhndoif nom. nov. See Figure 2. Etymology: Gr. philos = having affinity for. = Mchinonmhi dryadis C.J. Johanson in Rabcnhorst, 1-ungi europaei no. 3659 (1890). (Basionym). = Leptosphaeria dryadis E. Rostrup, Botanisk Tidsskrift 25:305. 1903. Ascocarps clustered, erumpent to superficial, papillate, glabrous to slighll\ tomentose toward base, tomenlum of dark brow n hyphae. conic- globose, 150-250 |im diameter. 180-275 |im high. Ascocarp wall of textura angulaiis in sutface \ iew : in longitudinal section uniformly 12-20 |im thick (up to 27 )itn thick near apex), composed of 4-5 layers of polygonal, isodia- metric to slightly elongate, scleroplecten- chymatic cells, outer 2-3 layers composed of hrown-melani/ed cells (3.5-4.3 x 5.5-6.8 |im). inner 2-3 layers composed of hyaline, com- pressed cells (2.1-3.5 X 10.0-12.3 ^m). Papilla conical, 60-65 |im high, 25-50 \xm wide at the apex. 90-1 10 )itii wide at the base, composed of 7-9 layers of isodiametric cells (2.8-3.6 |am diameter), wall 10-12 |im thick surrounding a 35—45 |im w ide ostiole without May 1992 Leplosphufiia Species on ihc Rosaceae 485 Figure 2. Lcptosphai'iia dryadnpluUi. a. SEM of erumpcnl ascocarps. xSO. b. ascocarp wall surface. xSSO. c. longitudinal median section through ascocarp, x25(). d. cellular pseudoparaphyses. x2.()()0. e. longitudinal median section through ascocarp wall. x95(). f. ascus, x5()(). g. ascospore, xl.l.'iO. a. c. c. and f from holotype of Z.. dryaJis: b. d. and g from isotype of/,. dryudophiUi. linols Natural Hislor> Survey Bulletin Vol. 34 An. 5 periphyses. Pseudoparaphyses 0.7-3.0 ^m wide, 1 10-175 )im long (height of the ascocarp cavity), numerous, narrowly cellular, with gelatinous coating. Asci (72-)90-l 18 x 10.8- 13.6 ^xm, numerous, basal to slightly lateral, cylindrical, thin-walled but with a thickened rounded apex, short-stalked, with 8 biseriate ascospores. Ascospores 20.1-24.5 x 5.7-7.2 |im. fusiform to slightly clavate. with acute end-cell shape, slightly curved; 3(-4-5)-septate. septa evenly distributed, order of septation 2: 1 :2(:3:4). without constrictions: pale brown- ish yellow, with ornamented wall, without sheath or appendages. Isotype: Swkijkn: Jamtland, Renfjallet, c. 900m. in fructibus et calycibus emortuis Dixadis octopetalae L. 13, Juli 1884, leg. C.J. Johanson. Rabenhorst. Fungi europaei no. 3659 (1890) (FH. NY). Exsiccatae: Sweden: Jamtland, Renfjallet, 13, Juli 1884, leg. C.J. Johanson, Vgr., Micr. rar. sel. 105 (FH). Other material examined: IrEi and: Sponsgerdi, on Diyas octopetalci. Juli 12, 1901, O. Davidson (type of Leptosphaeria dryadis. C). Comments: This fungus was originally de- scribed in Mclanomma and was retained with some reservation in that genus by Holm ( 1957, 1979). who stated. "The species is hardly a Mclaiunnma but its true affinities are doubtful to me." Mc'lciiiimumi dryadis differs from the current concept of Mclanomma (Barr 1987a) in not having asci that arise peripherally within the centrum and not having trabeculate pseudoparaphyses. Barr ( 1982) erected the genus Bricodkca for a biologically similar species. Bricookea scpalnnim (Vleugal) Barr. which is found on the inflorescences o( Jiintiis species. MeUmomma dryadis differs from this fungus in several respects: cells of the ascocarp wall are sclcroplectenchymatous. not pseudo- parenchymatous; the ascocarp apex opens by a pore, not by a slit; and the ascospores are not hyaline. Mclanomma dryadis. occurring in and on the dead llowers and fruits of D/\v;.v octopclahi. is a true Lcpiospliacria. ha\ ing the characteristics of an erumpeni to superficial ascocarp with a wall of sclcroplectenchymatous cells and 3-septate yellow-brown a.scospores. It does not, however, show the radiating arrange- ment of cells of the ascocarp surface character- istic of L. doliolum. Because the epithet dryadis is pre-empted in Leptosphaeria by L. dryadis Rostrup, a new name, Leptosphaeria dryadophila is proposed for Mclanomma dryadis Johanson. 'Leptosphaeria praetermissa (P.A. Karsten) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:26. 1883. See Figure 3. = Sphaeria praetermissa P.A. Karsten, Fungi Fenniae Exsiccati. Century 9, No. 852.^Anno 1869. Ascocarps clustered, sparse, immersed- subepidermal becoming erumpent. papillate. glabrous to tomentose at base, globose, flattened at base, 350-625 |im diameter. 300- 500 )im high. Ascocarp wall of textura prismatica in surface view, composed of cells radiating from apex outward: in longitudinal section uniformly 32-58 |im thick at sides and base, composed of 7-8 layers of hyaline, polygonal, isodiametric, scleroplectenchymatic cells (5.7-13.6 x 3.6-7.2 |im). inner 2-3 layers somewhat elongate and compressed, with an external brown-melanized crust. Papilla beaklike, cylindrical, erumpent. 120-180 |im high. 75-135 |im wide: wall 25-36 |im thick, composed of 5-6 la\ ers of small, brow n- melanized. thick-walled, polygonal, isodiamet- ric cells (4.3-7.2 (im diameter), surrounding a 35-80 |im v\ ide circular ostiole formed from small hyaline cells, w ithout periph\ses. Pseudoparaphyses 1 .0-2.0 |im wide. 180-260 |am long (height of the a.scocarp ca\ ity). numerous, narrowly cellular, without gelati- nous coating. Asci 100-130(-150) X 7-13 |im. numerous, basal, cylindrical, thin-walled, shon- stalked. rounded apex with apical chamber present, with 8 biseriate ascospores. .Ascos- pores 17-20(-25) X 5.5-8.0 |im, broadly fusifonn. end cells acute to rounded, straight to slightl> cur\ed: 3-septate, septa e\enl\ distrib- uted, order of septation 2:1:3 or 2:1:2, primary septum median, w ith constrictions at all septa, second cell occasionally enlarged (wider): brownish-Ncllow. smooth, without sheath or appendages, Isotype: Finland: In caulibus emortuis Riihi odoraii et R. idaci in par. Tammela sat fre- May 1992 Lfplosi'hufiid Species i)n the Rosaceae 487 488 Illinois Natural History Survey Bulletin Vol. 34 An. 5 quenter per annum obvia. Fungi Fenniae Exsiccati, Century 9. No. 852 (H, FH). Other material examined: Canada: Quebec: Gatineau Provincial Park, Chemin Ridge Road, on Rithus odoratus. 20 Jun. 1987, C.T. Rogerson (NY): Sweden: Jemtland: Five collections, all on Riihus ideaus. Leg. A.G. Eliasson, 20 May \9?,i), 27 Jun. 1930. 17 Jun. 1931, 19 Jun. 1931, 28 Jul. 1931 (S); Umea, Apr. 1908,J. Vleugel(S). Comments: Leptosphaeria praetermissa warrants inclusion in the genus because of its large erumpent ascocarp with a wall composed of scleroplectenchymatous cells. As in L. doliohim, the wall has an external crust, and the cells form a radiating pattern in surface view. It differs from L. doUolum in having a rather long, cylindrical neck that grows through the host epidermis before the ascocarp becomes erumpent. This species seems to be found predominantly in Europe but is probably not common there. It was not encountered in any collections of Ruhiis sp. in the United States but was found in one collection from Canada. •Leptosphaeria umhrosa G, NiessI in G.L. Rabenhorst, Fungi Europaei Exsiccati, Klotschii Herbarii vivi Mycologici Continuatio, Edition 3 (Edita Nova), Series 2, Century 20, No. 1934. Anno 1875; Just's Botanisch Jahresberichte 3:262. 1887. See Figure 4. = Massciha umhrosa (G. NiessI) H. Rehm in P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 9:761. 1891. Ascocarps scattered, immersed-subcuticular, papillate, glabrous, globose, flattened at base, 1 50-300 |am diameter, 90-200 nm high. Ascocarp wall of textura prismatica in surface view, composed of cells radiating from apex outward; in longitudinal section 4-6 |am thick at the base, composed of thin, hyaline, com- pressed cells, 10.8-22 |im thick at the sides, composed of 5-6 layers of polygonal, scleroplcctenchymatic cells, outer 2-3 layers of isodiamelric to slightly elongate, brown- melanized cells (3.5 x 7-10 |am) surrounded by a brown-melanized crust, inner 2-3 layers of hyaline, elongate-compressed cells (2-3 x 10- 18 \im). Papilla bluntly conical, (18-)25-54 MMi high, 36-56 nm wide at the apex, 60-70 |im w ide at the base, composed of 5-6 layers of small, isodiametric, heavily brown-melanized, thick-walled cells (2-3.6 |im diameter), wall 14.4—22 nm thick surrounding a 25—45 ^m wide ostiole without periphyses. Pseudo- paraphyses 0.7-2.0 \im wide. 80-1 10 ^m long (height of the ascocarp cavity), numerous, narrowly cellular, with guttule-like thickenings at septa, without gelatinous coating. A.sci (72-) 80-1 ()()(-] 17) X 10.8-15.0 lam. numerous, basal, cylindrical-clavate. thick-walled, rounded apex with apical chamber, short- stalked, with 8 tri- to biseriate ascospores. most commonly immature. Ascospores 27.3-33.0 x 4.3-7.2 |im. hyaline to subhyaline when immature, 28-37.4 x 5.7-9.3 [im. pale brown when mature, narrowly fusiform, with acute end-cell shape, slightly curved: 4-septate, septa unevenly distributed, order of septation 2: 1 :2:3, primary septum supramedian and slighth constricted, second cell slightly enlarged at maturity, slightly roughened wall, continuous sheath (0.7-1.5 (im thick) surrounding imma- ture spore, without appendages. Holotype: Austria: An Spiraea Anincus [=Anincus dioicus (Walt.) Fern.] bei Voitsberg in Steiermark. Septbr. 1874 (M). Exsiccatae: Austria: Voitsberg, Steiermark, Septbr., G. v. NiessI, Rabenhorst, F. europaei exs. 1934 (FH, NY isotypes); Voitsberg. Steiermark. Aug. 1882, G. v. NiessI, Rehm, Ascomyceten 690 (FH. NY authentic material); Voitsberg, Steiermark, G. v. NiessI. Weese, Eumycetes sel. exs, 638 (FH). Other material examined: Germany: Frauenfeld, on Solidago, October (NY). Comments: Leptosphaeria umhrosa is interest- ing because initially it seems to resemble members of the Phaeosphaeriaceae having small, pseudoparenchNniaiic-w ailed, subcuticular ascocarps that lift the host cuticle on maturity. Upon closer examination, it reveals the characteristic features of Leptosphaeria such as ascocarp walls com- posed of scleroplectenchyma surrounded by an external brown crust and wall cells that radiate in surface view. Other distinctive characteris- tics that ma\ aid in the identification of this species are the wide pseudoparaphy ses with proniinant thickenings at the septa and mature May 1992 LeptiKsphcicria Species on the Rosaceae 489 Figure 4. Leptosphaeria umhrosa. a. SEM of immersed ascocarps, xl50. b. ascocarp wall Mirfacc vviili radiating cells. x550. c. longitudinal median section through ascocarp. x2()(). d. cellular pseudoparaphyses with thickenings at septa, x2.()()(). e. longitudinal median section through ascocarp wall, x'->%). f. ascus w ith immature ascospores. x^lS. g. ascus with mature ascospores. x.'i25. h. immature ascospore. x 1.1.SO. i. mature ascospore from substrate surface, x 1.1 30. All Irom holotype of/,, iiiiihidsa. Ilinois Natural History Survey Bulletin Vol. 34 Art. 5 brown ascospores that are often seen only on the suiface of the substrate suirounding the ascocarp. Also diagnostic are the thin a.scocarp bases often left behind on the substrate when ascocarps are removed from the substrate suiface. •Leptosphaeria cercocarpi H. Sydow & P. Sydow, Annales Mycologici 5:339. 1907. See Figure 5. Ascocarps clustered, abundant, immersed- subepidermal, with surrounding stroma extending into substrate, papillate, glabrous, globose, 175-240 |im diameter, 1 75-240 ^m high. Ascocarp wall of textura angularis- globulosa in surface view; in longitudinal section 22-31 |J.m thick at sides and base, up to 50 |im thick near apex, composed of 6-8 layers of polygonal, isodiametric-elongate, pseudo- parenchymatic cells (3.6-7.2 x 5.4-l2.6(-16) |im|, outer 2-A layers of brown-melanized cells at sides and base, inner 4-5 layers of hyaline- compressed cells at base. Papilla very short, erumpent, rounded-conical, 31—45 |im high, 62-93 |im wide, wall 25-30 |im thick, com- posed of 8-10 layers of small, isodiametric, scleroplectenchymatic cells (2.0-7.2 |am diameter), outer 5-6 layers brown-melanized, inner 3^ layers hyaline, surrounding a 30-40 |im wide circular ostiole without periphyses. Pseudoparaphyses 1.0-2.0 |im wide, 125-200 Jim long (height of the ascocarp cavity), numerous, narrowly cellular, with gelatinous coating. Asci 95-120 x 17-21 |am, numerous, basal, cylindrical-oblong, thick-walled, short- stalked, rounded apex, with 8 biseriate asco- spores. Ascospores (25-)27-3l(-33) x 8.5- 1 1.5 |am, broadly fusifomi to slightly clavate, end cells rounded, straight to slightly curved, 3-septate, .septa evenly distributed, order of septation 2:1:2, primary septum median, slight constrictions at all septa, brown, ornamented walls, without sheath or appendages. Hololype: United States: In foliis emortuis Ccnocaipi Icclifalii. Mill Creek Canyon, Salt Lake Co., Utah Amcricae bor. leg. A.O. Ganett no. 677 (S). Other material examined: United States: Utah: Box Elder Co., One Mile Creek, north side of Rafl River Mts., Aug. 25, 1986; Juab Co., 5.2 mi up Granite Creek Canyon, east side of Deep Creek Mts, Aug. 26, 1986: Rich Co., Sunrise Campground. Cache National Forest, Route 89, west of Garden City, July 13, 1985; Weber Co., Malans Peak, Aug. 1 1 .'l972; Weber Co.. south slope of Malans Peak. Wa.satch Mts., east of Ogden, July 1, 1976; Weber Co., mouth of Taylor's Canyon, Wasatch Mts., east of Ogden, May 18. 1972, Apr. 29. 1981. May 6. 1982. all on Cercocarpus ledifoliii.s Torre> in Torrey & Gray var. internumtaniis (Brittonia 39:424. 1987). all C.T. Rogerson (all NY). Comments: Leptosphaeria cercocarpi is an easily recognized species on Cercocarpus ledifolius. The ascocarps usually entirely cover both surfaces of dead leaves. Retaining this species in Leptosphaeria is not totally satisfac- tory, but placing it elsewhere is also problem- atic. Certain characteristics resemble those found in the Phaeosphaeriaceae isensii Barr 1987a). The ascocarps are small-medium sized and immersed in the substrate, and there appears to be some kind of hyphal grou th or stroma surrounding the ascocarps and extend- ing into the substrate. Other characteristics do not coincide with inclusion in that group. Although the lateral walls of the ascocarp of Z,. cercocarpi consist of pseudoparenchy matous cells, they are not thin and soft as is character- istic of the Phaeosphaeriaceae. In fact, the walls in the upper regions of the ascocarp consist of thick-walled scleroplectench\malous cells. In Barr's (1987a) key to this family , the only two genera in which this fungus could be placed are Phacosphaeria and Kulmusia. The ascocarps are not small and delicate enough for inclusion within Phacosphaeria. and the current concept includes only species on monocots (Shoemaker and Babcock 1989b) (but see also the discus- sions for P. pomoiui and P. Iiicilla herein). The current concept o\' Kalmiisia. which includes K. clivensis (see discussion herein), does not adequately accommodate L. cercocarpi. Therefore, unlike other leaf-inhabiting species of Leptosphaeria belonging elsewhere (i.e.. Phaeosphaeriaceae). this species is retained within Leptosphaeria because of the larger, robust nature of the ascocarp and w all. May 1992 LcfUospluit'iiu Species on the Rosaceae 491 Figure 5. Lepiosphaeria cercocarpi. a. SEM of immersed ascotarps. x5(). b. ascocarp wall surface. x550. c. longitudinal median section through ascocarp. x225. d. section through ascocarp wall neck region. x9.')(). e. longitudinal median section through ascocarp wall. x950. f. ascus, xSl.S. g. h. cellular pscudoparaphyses, x2,()()0. i. ascospore, x 1.1 50. All troin holotype of /,. cercocarpi. 492 Illinois Natural Hislory Survey Bulletin Vol. 34 Art. 5 Phaeosphaeriaceae I he species considered in this section all possess characteristics of the Phaeo- sphaeriaceae (Barr 1987a). Species in the Phaeosphaeriaceae are distinguished from those in the Leptosphaeriaceae by the smaller ascomatal size, the thin, soft ascocarp wall composed of pseudoparenchymatous cells, and the rather sparse pseudoparaphyses (Barr 1987a). The genera represented within the family are Paraphaeosphaeria Eriksson, Phaeosphaeria Miyake, and Kalmusia Niessl. 'Paraphaeosphaeria conccntrica (J.B. Ellis & B.M. Everhart) S.M. Huhndorf comb. nov. See Figure. 6. = Leptosphaeria concentrica J.B. Ellis & B.M. Everhart, The North American Pyrenomycetes. A Contribution to Mycologic Botany, p. 354. 1892. (Basionym). Ascocarps scattered, sparse, immersed- subepidermal, globose, papillate, glabrous. 90- 100 |im diameter, 100-125 |xm high. Ascocarp wall of textura angularis in surface view; in longitudinal section 7.2-10.8 |im thick at sides and base, composed of 2-.^ layers of elongate- compressed, hyaline, pseudoparenchyniatic cells (2.8-3.6 x 12-14.5 \im), wall up to 13 ^im thick near apex, composed of 3^ layers of isodiametric-elongate. slightly brown- melanized cells (3.6-5.6 x 7.2-12.2 \xm). Papilla very short, erumpent. bluntly conical. 15-20 |im high, 25-30 |im wide, 10-13 |im thick near base of papilla, papilla apex wall cells hyaline, thin-walled surrounding a circular ostiole without periphyses. Pseudoparaphyses 1 .4-2.0 (im wide. 70-90 |im long (height of the ascocarp cavity), numerous, narrowly cellular, without gelatinous coating. Asci (43-)54-66 x 10.8-12.2 |jm. numerous, basal, cylindrical- clavate, thick-walled, shoin-stalked. rounded apex, with apical chamber, with 8 biseriate ascospores. Ascospores (14.5-)16.5-18.7 x (3.6-) 4.3 ;i.7(-6.5) |am. cylindrical, with rounded end cells, end cells longer than central cell, straight; 2-septate. septa unevenly distrib- uted, order of septation 2; I . primary septum submedian and constricted; brown, wall roughened, thin cellular shealh entirely surrounding spore, without appendages. Holotype: Umted States: On apple leaves (Mains sp.). Columbia. Missouri (H. Dorsett), and Louisiana (Langlois) (NY). Comments: Paraphaeosphaeria sjjecies are distinguished by ascocarps that form below or within the host epidermis, an ascocarp wall consisting of a few layers of pseudoparen- chyma, and cylindrical, usually echinulate ascospores with the primary septum forming in the lower half of the spore. Although Paraphaeosphaeria species are commonly found on monocotyledonous plants, species on dicot plants have also been placed in the genus (Hedjaroude 1969, Shoemaker and Babcock 1985). Such is the case here with Paraphaeosphaeria concentrica described from apple leaves. The description of P. concentrica closely resembles the description of the type of the genus. P. michotii (Westendorp) Eriksson, with the only difference being the dicotyledon- ous host plant. Paraphaeosphaeria michotii and its synonyms have so far been found exclusively on monocot hosts (Shoemaker and Eriksson 1967. Shoemaker and Babcock 1985). Overlooking substratum preference. P. concentrica would be synonymous \\ ith P. michotii. Because of the historical value placed on this character and the need for experimental evidence to demonstrate the range of substrata on which species will grow and the total lack of collections of P. michotii on dicot hosts. P. concentrica is retained as a separate species. 'Phaeosphaeria pomona (P.A. Saccardo) S.M. Huhndorf comb. nov. See Figure 7. = Leptosphaeria iLeptosphaercUa) pomona P.A. Saccardo, Nuovo Giomale Botanico Italiano e Bolletino della Societa Botanica Italiana 8:176. 1876. (Basionym). Ascocarps scattered, sparse, immersed- subcuticular. globose, papillate, glabrous to slightly tomentose near base, with loose, dark hyphal growth on host cuticle surrounding ascocarps. (55-)90-125 |.im diameter. 75-1 15 |.im high, .\scocarp wall of textura angularis- globulosa in surface view; in longitudinal section unifomily 6.1-7.7 (im thick, composed of 2-3 layers of brown-melanized. compressed, isodiametric-to-elongate pseudoparench\nia- tous cells ( 1.5-2.0 X 8-12 \im). Papilla \er\ short, crumpeni, bluntl\ conical. 15-21 pm Mav 1992 Lfpliispluienu Species on the Rosaceae 493 Figure 6. Paiaphaeosphaeria concenlrica. a. longitudinal median section through ascocarp, x375. b. cellular pseudoparaphyses, x2,()()0. c. longitudinal median section through ascocarp wall, x950. d, e. asci, x 1,050. r, g. ascospores, x 1,1 50. All from holotype oi P cnncciilrica. high, 31-37 )im wide, composed of 3^ layers of small, brown -melanized, isodiametric cells (2.6-4.6 \im diameter), (9-)12-17 urn thick near base of papilla, apex of papilla wall cells hyaline, thin-walled surrounding a circular o.stiole without periphyses. Pseudoparaphyses 1.0-1.3 jam wide. (46-)77-93 )im long (height of the ascocarp cavity), numerou.s, narrowly cellular, with guttule-like thickenings at septa, without gelatinous coating. Asci 55.5-65 x 494 Illinois Natural Histor> Survey Bulletin Vol. 34 Art. 5 Figure 7. Phacosphaeiia pomona. a. longitudinal median section through ascocarp. x375. b. ascocarp wall surface, x.'i.SO. c. longitudinal median section through ascocarp wall. x950. d. ascus. x 1.050. e. ascospore. X 1 , 1 ."iO. All from holotype of P. pomona. 8.3-1 1.5 |im, numerous, basal, cylindrical, thick-walled, shon-stalkcd. rounded apex, with apiial chamber, with 8 biseriate ascospores. Astospores 20.1-26.5 x (2.8-)3.6-5.0 |im, naiTowly t' Ifoirn, with acute end cells, second cell from the top occasionally enlarged, straight; 5-scptate, septa unevenly distributed, w ithout constrictions, order of septation unknown, pale brownish yellow, guttules present in ascospore cells, smooth, thin cellular sheath entirel) surrounding spore (0.7-2.0 |.tm thick). Holotype: Italy: In pag. super, folior. Pyri Mali, socia \ crniiciiUirlii Pomona, a selva (Treviso). Sept. 1875. raro (P.AD). Comments: See under Pluwosphacha liicilla. 'Plhicosphacria hicilhi (P..A. Saccardo) S.M. Huhndorf comb. nov. See Figure 8. = Lcptosphaeria liicilla P.A. Saccardo. Nuovo Giomale Bolanico Italiano e Bolletino dclla Socicla Botanica Italiana 7:310-311. 1875. (Basionym). May 1992 Leplosphaciiii Species on the Rosaccae 495 m^\ .\:^*> } ~,^--' Figure 8. Phaeosphaeria tucitla. a. longitudinal median section through ascocarp, x375. b. cellular pseudoparaphyses. x2,000. c. longitudinal median section through ascocarp wall. x93n. d. asci. xI.OSO. e. ascocarp wall surface, x550. f. g. ascospores. xl.lSO. All from holotype of P. lucilUi. Ascocarps scattered, sparse, immersed- subcuticular, papillate, glabrous, depressed- globose, 1 If)- 130 |im diameter, 90-1 15 urn high. Ascocarp wall of textura angularis in surface view; in longitudinal section uniformly 6.0-7.7 |im thick, composed of .3^ layers of brown-melani/ed, compressed, isodiametric-to- elongate, pseudoparenchymatous ceils (2.3-3.8 X 6.9-10.0 |im). Papilla very short, erumpent, conical, 20-25 |am high, M)-^() |im wide, composed of 4-5 layers of browii-mclani/ed, isodiametric cells (3.0-4.6 |am diameter), 10.7- .')6 linois Natural History Survey Bulletin Vol. 34 An. 5 13.0 )im thick near base of papilla, apex of papilla wall cells hyaline, thin-walled surround- ing a circular ostiole without periphyses. Pseudoparaphyses 1.5-2.0 ^trn wide, 62-71 fiiTi long (height of the ascocarp cavity), numerous, narrowly cellular, without gelati- nous coating. Asci 40-5 1(-56) x 8.4-10(-l 1.5) )im, numerous, basal, in a broad hymenium, cylindrical, thin-walled, sessile, rounded apex, with 8 hi- to triseriate ascospores. Ascospores 15.3_18.4(-20) X 3.0-4.6 ^m, fusiform, with acute end cells, second cell from apex slightly enlarged, straight to slightly curved; 3-septate, septa evenly distributed, order of septation 2: 1 :2, primary septum median and slightly constricted; pale yellowish brown, wall smooth, without sheath or appendages. Holotype; Italy: In foliis languidis Pyri communis in agro Tarvisino et Patavino (PAD). Comments: Fungi in the genus Phaeosphaeria are characterized by immersed ascocarps with thin walls of pseudoparenchyma, fusiform 3- to multiseptate ascospores and monocotyledonous hosts. Nonetheless, Leuchtmann's ( 1984) treatment oi Phaeosphaeria includes a group of species found on the leaves and stems of plants in the dicot family Caryophyllaceae. The placement of dicotyledonous Leptosphaeria species, morphologically similar to Phaeosphaeria. into the genus Phaeosphaeria is contrary to Shoemaker and Babcock's (1989b) concept of the genus. These authors do not, however, suggest an alternative genus for these species. They do place certain species on dicots in Leptosphaeria. citing the presence of thick-walled cells in the ascocarp wall. Phaeosphaeria pomona and P. Iiiciila are morphologically similar to species of Phaeosphaeria found on monocot hosts. They do not have the ascocarp and wall characteris- tics of Leptosphaeria. When P. pomona and P. Iiuilla were described. Saccardo (1875, 1876) placed them in Leptosphaeria subgenus Lepiosphaerella Sacc, which included fungi from leaves of dicotyledonous plants. Leptosphaeria subgenus Leptosphaerella was subsequently placed in synonymy with Phaeosphaeria (Leuchtmann 1984). At this time, there is no other genus suitable for the placement of these "dicot Phaeosphaeria" species. Erecting a new genus for these species based solely on substrate preference, with no morphological differences from Phaeosphaeria seems unw ise. With proof of strict substrate preference requiring experi- mental evidence, there seems to be some justification for their placement within Phaeosphaeria until such work is done. 'Phaeosphaeria thomasiana (P.A. Saccardo & C. Roumeguere) S.M, Huhndorf comb. nov. See Figure 9. = Leptosphaeria thomasiana P.A. Saccardo & C. Roumeguere. Revue Mycologique 5:236. 1883. (Basionym). Ascocarps scattered, numerous, immersed- subcuticular. globose, flattened at base, papillate, glabrous. 150-175 \xm diameter. 1 10-125 |im high. Ascocarp wall of textura angularis-globulosa in surface view; in longitu- dinal section uniformly 9.3-12.2 |im thick, composed of 4—5 layers of polygonal, pseudo- parenchymatous cells, outer 2-3 layers of polygonal-to-elongate, brown-melanized cells (2.2-5.0 X 6.5-8.6 ^m). inner 1-2 layers of elongate-compressed, hyaline cells (0.7-1.5 x 8.0-10.8 |im). Papilla very short, erumpent. bluntly conical. 12-30 |im high. 10-36 |im wide, composed of 7-8 layers of small, brown- melanized. isodiametric cells (2.6—4.6 \im diameter). 14—18 (im thick near base of papilla, apex of papilla wall cells hyaline, thin-walled surrounding a 12-18 |im wide circular ostiole without periphyses. Pseudoparaphyses 1.0- 1.5 |im wide. 80-85 |im long (height of the ascocarp cavity), numerous, narrowly cellular, without gelatinous coating. .\sci 55.5-65.5 x 8.0-10.8 \xm. numerous, basal, cylindrical- clavate. thick-walled, short-stalked, rounded apex, with apical chamber, w ith 8 biseriate ascospores. Ascospores 15.0-18 x 3.6-4.5 \im. fusiform, with acute end cells, straight or slightly cur\'ed; 3-septate. septa evenly distrib- uted, order of septation 2:1:2. primary septum median, without constrictions; subh>aline to pale brow nish yellow . guttules absent in ascospore cells, smooth, w ithout sheath or appendages. Holotype: Fr.ance: In samientis Riihi emortuis (305-Reliquiae Libertianae) n.v. May 1992 Leptosphaeria Species on the Rosaceae 497 'w 'l^:-^ "^>^. 1 A (^ x Figure 9. Phaeosphaeria thomasiana. a. longitudinal median section through ascocarp, x375. b. cellular pseudoparaphyses, x2,000. c. longitudinal median section through ascocarp wall. x950. d. asci, xl,050. e. ascocarp wall surface, x550. f. g. ascospores. x 1,1 30. All from Roum. F. sel exs. 6039. 498 Illinois Natural History Survey Bulleiin Vol. 34 Art. 5 Exsiccatae: Fra.nce: Bois des Roches (Noidan), May 1891, F. Fautrey, Roumeguere, F. sei exs. 6039 (NY). Other material examined: United States: Oregon: Corvallis, on loganberry (Riihiis logannhaccus Bailey). Mar. 12, 1930, S.M. Zeller; on loganberry. Mar. 15, 1916, A. Frank (all NY). Comments: Leptosphaeria thomasiana is placed in Phaeosphaeria because of its small ascocarps immersed beneath the host cuticle and its ascocarp wall of pseudoparenchymatous cells. 1 was unable to see the type specimen, which is number 305 in the exsiccatae set Reliquiae Libertianae. This is not a regular exsiccatae set (Pfister 1985), and the set at FH did not contain this specimen. The description and plate herein were prepared from Roumeguere, F. sel exs. 6039 specimen from NY. which included few ascocarps. I did not find any ascocarps of P. thomasiana in the collections from Oregon, but the canes did have large gray patches on them as Zeller (1927) described. He also mentions that a cane blight or "Loganberry gray bark disease" has been ascribed to this organism in western Washing- ton. 1 have seen no reports from other regions of this organism as a pathogen of Riibiis. 'Kalmiisia clivensis (M.J. Berkeley & C.E. Broome I M.E. Barr, Mycotaxon. An Interna- tional Journal Designed to Expedite Publication of Research on Taxonomy & Nomenclature of Fungi & Lichens 29:504. 1987. See Figure 10. = Sphaeria (Caitlicolae) clivensis M.J. Berkeley & C.E. Broome, Annals and Magazine of Natural History, Series 2, 9:379. 1852. = Leptosphaeria clivensis (M.J. Berkeley & C.E. Broome) P. A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:16. 1883. = Diapleella clivensis (M.J. Berkeley & C.E. Broome) A. Munk, Dansk Botanisk Arkiv 15(2):75. 1953. = Leptosphaeria iialionim P.A. Saccardo var. lapsanac P.A. Saccardo & P.A. Briard. Revue Mycologique 7:209. 1885. = Leptosphaeria steironematis J.B. Ellis & B.M. Evcrhart. Proceedings of the Academy of Natural .Sciences of Philadelphia 1890: 2.^7.(1890) 1891. = Leptosphaeria arunci S.M. Zeller. Mycologia 19:134-135. 1927. = Leptosphaeria longipedicellata J.H. Miller & G. Burton. Mycologia 34:2-3. 1942. Ascocarps scattered, sparse, immersed- subepidermal. papilla erumpent. at times with a surrounding clypeus. glabrous, depressed- globose, 275—400 \xm diameter, 175-375 |im high. Ascocarp wall of textura prismatica in surface view: in longitudinal section uniformly 15-20 |im thick, composed of 5-6 layers of parallel, elongate, prismatic, scleroplecten- chymatic cells (0.7-1.5 x 15-20 nm), outer 3^ cell layers slightly brown-melanized, inner 2-3 layers hyaline: at the base cells are compressed, flattened, hyaline. Papilla conical, short- intermediate. (50-)90-100(-130) urn high. 40- 80 |im wide at the apex, 50-100 )im wide at the base, composed of 6-8 layers of small, light- brown pigmented, isodiametric cells with no external, melanized crust (2-5 fim diameter), 10-13 |im thick, surrounding a circular ostiole 18-20 fim wide, composed of thin-walled, hyaline, compressed cells, without periphyses. Pseudoparaphyses 0.5-1.5 |im wide, 2CK3-225 |am long (height of the ascocarp cavity), numerous, narrow ly cellular. \\ ithout gelati- nous coating. Asci 95-1 10(-150) X n-15(-21) |im, numerous, basal, clavate, thin-u ailed, long-stalked (27-39 )im long), rounded apex, with 8 biseriate ascospores. Ascospores 19- 25(-32) X 5-8.5(-l 1 ) ^m, fusiform, with rounded to acute end cells slighth longer than central cells, straight to slightly curved: 3- septate, septa slightly unevenly distributed, order of septation 2: 1 :2, w ith slight constric- tions at all septa: dark brown, smooth, without sheath or appendages. Holotype: Great Britain: King's Cliffe, on dead stems of Pastinaca sativa. Jul 1 850. Herb. Berk. 1879 (K). Exsiccatae: Canada: London, on Steironema ciliatum. as Leptosphaeria steironematis, Ma\' 1890, Ell. & Ev.. North American Fungi 2615 (NY):Sphaer. Brit. 11160. (FH). Other material examined: Canada: London, on Steironema ciliatum. May 1890, with 1640. Deamess (Holot>pe of Leptosphaeria stcironetnatis. N^); 1640. London, as Leptosphaeria steironematis. 1 9 Apr., 1 890 May 1992 Lepiospliaeria Species on the Rosaceae 499 Figure 10. Kalmusia clivensis. a. SEM of immersed ascocarps with crumpent papilla, xlOO. b. ascocarp wall surface. x550. c. longitudinal median section through ascocarp. xl25. d. cellular pseudoparaphyses. x2,()0(). e. longitudinal median section through ascocarp wall. x95(). f. g. asci, x.'>2.'i. h-j. ascospores. xl.L'iO. a. c. d, e, f. and h from holotype o( l.epuisphacria arum i; b, g. and i from hololypc of /C clivensis: j from lectotype of/,, liingipedicelhila. .,00 Illinois Natural History Survey Bulletin Vol. 34 An. 5 (NY). Great Britain: England, Chute Meadow, Lambriggan, W. Cornwall, on Centaiirea nigra. May 9, 1942, F. Rilstone (NY-MEBB coll); Scotland, Wester Ross, Rassal N.N. Resei^'e, on ? Seneciojacohoea, June 2, 1982. P.P. Cannon (NY). Italy: Briard no. 8, on Lapsana communis. June 12, 1885 (holotype of Leptosphaeria galiorum var. lapsanae. PAD). United Si atf.s: Colorado: Larimer Co., 3 mi W. Redfeather Lakes, 8100' Roosevelt National Forest, on Rosa ,sp., Aug. 4, 1984, MEBB # 7008 (NY); Georgia: 7684, Clarke Co., Athens, South Campus, University of Georgia, on Daiwus carota. Sept. II, 1939, J.H. Miller (Lectotype q{ Leptosphaeria longipedicellala. GAM); Michigan: Emmet Co., Gill and Elder Roads, Carp Lake, on ? Acer, Sept. 1 1 , 1969, MEBB #5527 (NY); Oregon: Multnomah Falls, April, on dead stems oi Anmciis Silvester (holotype of Leptosphaeria ariinci in Zeller Herb. 6811, NY). Comments: Four of the putative Leptosphaeria species that were examined are synonymous with Kalmusia cHvensis; one was found on a Rosaceous host {L. arunci on Aruncus). The most distinctive feature of this species is the presence of long-stipitate asci with prominent dark brown, 3-septate ascospores. The long stipe, as well as the thin ascus wall, has led previous workers to treat it as a unitunicate fungus in the monotypic genus Diapleella (Munk 1957, Dennis 1978). Shoemaker ( 1984a) retained the genus Diapleella but treated it as bitunicate. Barr (1987b) trans- ferred D. clivensis to the genus Kalmusia Niessl without much explanation. The original description of Kalmusia (Niessl 1871) and the illustration of the genus in Berlese (1890) show a fungus with long-stipitate asci and dark brown, 3-septate ascospores. Kalmusia clivensis appears to be well-placed within this genus. Shoemaker (1984a) looked at one collection of Leptosphaeria longipcdicellata on Solidago caesia L., which is cited in the original description of the species. He men- tions that "the original description of Z.. longipedicellata is strongly suggestive of Diapleella clivensis." This collection was in fact L. macrospora (Fuckel) Thiimen, and Shoemaker did not resolve the placement of L. longipcdicellata. The collection on Daucus carota L., cited in the description and marked as type on the herbarium packet, matches the original description exactly and is the same as Kalmusia clivensis except that the ascospores and asci are slightly larger than those in the type collection of K. clivensis. The ascocarp and wall in section appear exactly the same. I believe the size differences are probably due to environmental differences or other individual variation and that L. longipcdicellata is synonymous with K. clivensis. Collection 7684, Clarke Co., Athens, South Campus. University of Georgia, on Daucus carota. Sept. 11, 1939, J.H. Miller (GAM) is chosen as the lectotype of Leptosphaeria longipedicellala. 'Kalmusia coniothyrium (L. Fuckel) S.M. Huhndorf comb. nov. See Figure 11. = Sphaeria coniothyrium L. Fuckel, Symbolae Mycologicae, p. 115. 1870. (Basionym). s Leptosphaeria coniothyrium (L. Fuckel) P.A. Saccardo, Nuovo Giomale Botanico Italiano e Bolletino della Societa Botanica Italiana 7:317. 1875. = Melanomma coniothyrium (L. Fuckel) L. Holm, Symbolae Botanicae Upsalienses 14(3):56-57, 1957. = Diapleella coniothyrium (L. Fuckel) M.E. Barr in M.E. Barr. C.T. Rogerson. S.J. Smith, and J.H. Haines. Bulletin of the New York State Museum 459:30. 1986. = Sphaeria (Ohtectae) hendersonia J.B. Ellis ;;; M.C. Cooke and J.B. Ellis. Grevillea 6:14-15. 1877. = Clypeosphaeria hendersoniae (J.B. Ellis) P. A. Saccardo, S> lloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:91. 1883. = Leptosphaeria {Clypeosphaeria) hendersoniae (J.B. Ellis) M.C. Cooke, Grevillea 17:91. 1889. Ascocarps clustered, numerous, immersed- subepidemial. depressed, globose, flattened at top and base, sometimes beneath blackened clypei. papillate, glabrous. 175-300 |im diameter, 175-200 |.ini high. .Ascocarp wall of textura angularis-globulosa in surface view ; in longitudinal section 15-25 |im thick at the sides, 13-18 |.im at the base, composed of S-12 layers of pol\gonal. pseudoparenchymalous cells, outer 3-5 layers of isodiametric-to- slighlly-elongate, light brown cells (5.0-9.5 x May 1992 Leplosphaeria Species on the Rosaceae 501 W^' ' . ^^^^^ , •4r^^J^ Figure 1 1. Kalmusia conioihyhum. a. longitudinal median section through ascocarp. x250. b. ascocarp wall surface, x550. c. longitudinal median section through ascocarp wall. x950. d, c. asci, xl,()5(). f, g. cellular pseudoparaphyses, x2,(K)(). h-j. ascospores, xl,150. a-d, f-i from holotype of /C. conioihyhum; e, j from neotype of Sphaeria hendersonia. .02 Illinois Natural History Survey Bulletin Vol. 34 Art. 5 <.ty-5.() |im), inner 5-7 layers of .small, isodiametric-eiongate. compressed, hyaline ceils (3.6-4.3 x 2.0-3.6 |im). Papilla short, erumpent. bluntly conical, 45-55 |im high. 25- 35 (im wide, 16-18 ^m thick, composed of 5- 10 layers of small, hyaline, isodiametric cells (1.4-2.8 nm diameter), surrounding a 10-20 |im wide, circular ostiole without periphyses. Pseudoparaphyses 1.0-1.5 |im wide, 75-100 |.un long (height of the ascocarp cavity). numerous, narrowly cellular, with guttule-like thickenings at septa, with gelatinous coating. Asci 60-75 X 5.5-7.5 ^m. numerous, basal in a broad hymenium, cylindrical, thin-walled, short-stalked, rounded apex, with apical chamber, with 8 overlapping, uniseriate ascospores. Ascospores 1 1.5-14.4(-15. 8) x 3.6-4.5 |im, fusiform to ellipsoidal, with acute end cells, second cell somewhat enlarged, straight or slightly curved; 3-septate, septa slightly unevenly distributed, order of septation 2:1:2, primary septum median and constricted; brownish yellow, guttules lacking, smooth, without sheath or appendages. Holotype: Austria: Auf durren Ranken von Riihiis fniticosus. selten, im Fruhling. An der Heimbach bei Oestrich (G). Exsiccatae: Rehm, Asc. 388, on Riihiis fniticosus (NY); Krieger, Fungi saxon, 18. 1120, 1121 (NY); Petrak Kryptogamaeexsic. 2318 (NY); Sacc. Mycotheca Veneta 72. as Spluieria fiiscella f. Ampelopsidis hederaceae. (FH); Petrak, Fl. Boh. et Mor. exsic. U no. 4, as Clypeosphaeria notarisii (FH); Ellis N. American Fungi 581 as Sphaeria Heiulcrsonia. Mar, 1878(ILL, FH, NY). Other material examined: Canad.a: Ottawa, on Sambucus racemosa. Mar. 10, 1897 (NY); London, as Clxpeosphaeria Hendersonia. 1883, 3 May 1892 (NY). Italy: 198 as Sphaeria clypcala. de Notaris (RO). United States; Delaware: Faulkland. as Sphaeria Hendersonia. Mar. 20. 1887 (NY); New Jersey: Newfield. on Riihiis srrii;(>si. d, e. trabeculaie pseudoparaphyses. x2.(X)0. f. longitudinal median section through ascocarp wall. x930. g. ascus. x525. h. i ascospores. x 1.1 50, holotype of Cladoxphaeiia rimkola. I . x5.'iO. All from May 1 992 Lepiosphaeria Species on ihe Rosaceae 507 Other material examined: Switzerland: Bremengartenwald, on Prunus avium L. (holotype of CluJospluieriu rimicohi. BERN). United States: California: Spruce Cove Heads, Trinidad, Humbolt Co., on Ruhiis parrifloriis. 30 Jan. 1941, H.E. Parks 6313 (FH). Comments: Leptosphaeria rimicola. described from the fallen branches of Prunus avium L., is synonymous with MeUinomma pulvis-pyrius. Leptosphaeria rimicola has the overall appear- ance of M. pulvis-pyrius. with large, gregarious, superficial ascocarps and ascospores with the characteristic Melanomma shape: 3-septate w ith the primary median septum dividing the spore into a wider anterior and a narrower posterior part. The measurements of the ascospores of £,. rimicola are somewhat larger than were found by Chesters ( 1938) for M. pulvis-pyrius. but they are not beyond the range for M. pulvis-pyrius given by Saccardo ( 1878). The asci peripherally lining the centrum about halfway up the wall and the ascocarp wall composed of small thickened cells correspond to Barr's (1987a) concept of the genus Melanomma. The pseudoparaphyses are thin and flexuous. but the branchings and anastomo- ses are infrequent and the septa often show thickenings, making them difficult to accurately identify as trabeculate. Leptosphaeria Species Referable to the Dothideales The species in this chapter all belong in the Dothideales (sensii Barr 1987a). Barr"s concept of this order differs considerably from the all-inclusive concept of von Arx and Miiller (1975) or the broad concept of Eriksson and Hawksworth{1985, 1986). The Dothideales (Barr 1987a) are characterized by ascocarps without a hamathecium (although interthecial cells are often present) and by asci that tend to be ovoid to saccate, arranged in a basal fascicle or a basal layer. The families represented are the Dothioraceae and the Pseudosphaeriaceae. Dothioraceae •SaccDtheciuni sepincola (E.M. Fries:E.M. Fries) E.M. Fries, Summa Vegetabilium Scandinaviae, p. 398. 1849. See Figure 14. = Sphaeria sepincola E.M. Fries, Observationes Mycologicae. 1:181. 1815; Systema Mycologicum Sistens Fungorum 2:498.1823. [h^ saepincola.] = Metasphaeha sepincola (E.M. Fries: E.M. Fries) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitoruin Digessit P.A. Saccardo 2:164. 1883. [As Fr? Fuckel.) = Leptosphaeria sepincola (E.M. Fries:E.M. Fries) H.G. Winter. Dr. L. Rabenhorst's Kryptogaman-Flora von Deutschland, Oesterreich und der Schweiz, Second edition, 1(2):473. 1885. = Sphaerulina sepincola (E.M. Fries: E.M. Fries) K. Starbiick. Botaniska Notiser 1890:1 17. 1890: Botanisches Zentralblatt, 46:261. 1891. = Piingslieimia sepincola (E.M. Fries: E.M. Fries) F. v. Hohnel. Annales Mycologici 18:97. 1920. = Pleospluwrulina sepincola (E.M. Fries: E.M. Fries) H. Rehm //; F. v. Hohnel. Annales Mycologici 18:96. 1920. = Sclerodolhis sepincola (E.M. Fries: E.M. Fries) F. Petrak, Annales Mycologici 19:41. 1921. For other synonyms see Barr (1972). Ascocarps thickly scattered, immersed- subepidermal, globose, glabrous. 180-200 ^m diameter, 170-180 jam high. Ascocarp wall of textura angularis in surface view: in longitudi- nal section 25-36 |im thick at sides and base, up to 45 |im thick at apex, composed of 7-8 layers of pseudoparenchymatic, polygonal cells (5-6 X 6-1 2 )im), outer 2-3 layers of isodiametric-to-elongate, brown-melanized cells, inner 4—5 layers of hyaline-subhyaline, thin-walled, elongate-compressed cells, at the base giving rise to a central column ( 1 8-23 |im high, 27-36 ^im wide) of hyaline, isodiametric cells (3^ |im diameter, but cell boundaries becoming obscured in mass) on which the asci are borne. Papilla broadly rounded, bluntly conical, 45-55 |im high, 55-70 |im wide. Asci 40-60 X (13-) 15-25 |im, numerous, fasciculate, borne on a basal column, oblong-ovate to clavate, thick-walled, short-stalked, apex rounded and thickened, w ith 8 bi- to triseriate ascospores. Ascospores ( 17-)18.5-20.5(-22.5) X 5.0-7.2 |im, obovate, with acute end cells, broad above and tapering to a narrower base, straight to slightly cuned; 4-5(-6)-septate, septa unevenly distributed, septaiion order 3:2: 1 :2:4 or 4:2: 1 :3:5, primary septum submedian, vertical septum occasionally present in second or third cell from the top: hyaline, wall smooth, without sheath or appendages. Lectotype: Sweden: Ad ramos emortuos Rosae alionimque fruticum (UPS). Exsiccatac: .Austria: Nassau, on Rosa canina. as Metasphaeria sepincola. 1894, Fuckel. Herbier Barbey-Boissier 385 (FH): Nassau, on Rubus fniticosiis. as Sphaerulina iniermixia (Berk & Br) Sacc, 1894, Fuckel, Herbier May 1992 Lepldsphuiriu Species on ihe Rosaceae 509 Figure 14. Saccothecium sepincola. a. SEM of immersed ascocarps. xl25. b. longitudinal median section through ascocarp wall. x950. c. longitudinal median section through ascocarp. x22.'i. d. asci. x 1.050. e. ascocarp wall surface, x550. f. ascosporcs. x 1.1 50. All from Icctotype of S. sepincola. 510 Illinois Natural History Survey Bulletin Vol. 34 An. 5 Barbey-Boissier 501 (FH); Czechoslovakia: Weisskirchen, Ohrensdorf, as Sphaerulina Intermixta, 25 Jan. 1912, Petrak, Fl. Boh. et Mor. exsic. Lfg. 4 Nr. 174. (FH); Germany: Leihterfeld bei Berlin, on Philadelphus sp., 1890, Sydow. Myc. March. 2934 (FH); Italy: Selva, as Sphaerulina intermixta, Sept. 1878, Saccardo, Myc. Ven. 1367 (FH). 'Saccothecium sepincnla var. ahhreviata (M.C. Cooke) S.M. Huhndort' comb. nov. See Figure 15. = Sphaeria abhreviata M.C. Cooke, Handbook of British Fungi, p. 893. 1 87 1 . (Basionym). = Leptosphaeria ahhreviata (M.C. Cooke) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:26. 1883. Ascocarps 1 17-144 |im diameter, 81-99 jam high; wall 14—21 \im thick at the sides and apex, up to 27 |im thick at the base, composed of 7-8 layers of pseudoparenchymatic, poly- gonal cells (5-6 X 6-12 |im). Papilla broadly rounded. Asci (32-)41-61 x (13-)16.5-24.5 |im. Ascospores (12-)13.7-16.6 x 3.6-5.7 |im, 4-5(-6)-septate. Holotype: Great Britain: On dead stems of bramble. Jan.-April (K). Comments: Saccothecium sepincnla is not uncommon, but it has been confused by mycologists over the years as exemplified by the number of name changes based on the fungus. Wehmeyer (1957) gave an account of the history and nomenclatural confusion surrounding this species, including its relation- ships with other organisms and its relationship to the later genus Pringsheimia Schulzer von Miiggenburg. He also included a lectotypifi- cation of Saccdthccinm Fr. Barr ( 1972) lists the taxonomic synonyms of Saccothecium sepincnla. In some treatments, Pringsheimia sepincnla is still used as the name for this species (I-roidevaux 1973. von Arx and Miiller 1975,Sivanesan 1984). Holm (1975) argued for the lectotypification of Saccnthecium Fr. 1835 by 5. sepincnla (Fr.) Fr. 1849. which, as mentioned above, apparently was already done by Wehmeyer (1957) (see Dennis 1978). Holm (1975) does not mention the lectotypification by Wehmeyer, so it is unknown if this lectotypification was not accepted or if Holm overlooked Wehmeyer's work. In any case, with lectotypification, Saccothecium is the correct name and Pringsheimia becomes a synonym. Saccothecium sepincnla is placed in the family Dothioraceae by Barr ( 1987a) and is characterized by sphaeroid ascomata with walls of pseudoparenchymatous cells and oblong to clavate asci w ith a thickened apex v. hich arise from a central basal column or mound of hypothecial cells. The septate, hyaline, obovate ascospores usually have a vertical septum present in one or more of the central cells. These characters suggest a similarity to some Dnthiora species with raised basal areas (Barr 1972). Leptosphaeria abhreviata is regarded as a variety of Saccothecium sepincnla because ascospore sizes differ. The ascospores of S. sepincnla are 17-22 x 5-7 |im. whereas the spores of S. sepincnla var. ahhreviata are 1 2- 16 X 3-6 nm. The ascospores appear to be mature in specimens of both species. The ascocarps also differ in size betvv.een the two species in the specimens seen. Some discrepancy exists between Cooke's ( 1 87 1 ) description of Sphaeria ahhreviata and the fungus that w as present on the type specimen. Cooke's ( 1871 ) description includes "perithecia minute, in short parallel lines" and "asci very short and broad, elliptical, pyriform or obovate." which matches the fungus in the type. But then he describes spores that are "triseptate. slightly torulose and pale brown," which does not match this fungus, although occasionally the spores when mature may appear slightly pale brown. It is unclear whether his description w as simply inaccurate or whether he was looking at two different fungi. Only one fungus is present on the type specimen and it w as similar to 5. sepincola. Pseudosphaeriaceae •Leptosphaerulina pulchra (H.G. Winter) M.E. Barr, Contributions de ITnslitut Botanique de L'Universite de Montreal 73:7. 1959. See Figure 16. = Sphaerella pulchra H.G. Winter, Hedwigia 11:145-146. 1872. = Leptosphaeria pulchra (H.G. Winter) P.A. Saccardo. S> lloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:53-54^ 1 883. May 1992 Li'plosphaeiia Species on the Rosaceae 511 I ,^ V*. Figure 15. Saccnihecium sepincola var. ahhreviala. a. longitudinal median section through ascocarp. x375. b. ascocarp wall surface, x550. c. longitudinal median section through ascocarp wall, xy5(). d. ascus, x 1,050. e. ascospores within an ascus, x 1,1 50. All from hololype of 5. sepincola var. cihhrcviaki. = Mycotodea pulchra (H.G. Winter) W. Kirschstein, Annales Mycologici .34:201. 1936: Kryptogamennora de Mark Brandenburg und Angrenzender Gebiete herausgegeben von den Botanischen Verein der Provinz Brandenburg 7(3):433. 1938. = Leplosphaeria oli^olheca F. Petrak & H. Sydow, Annales Mycologici 22:359. 1924. (Described from type material oi Laestadia potentillae E. Rostrup, Botanisk Tidsskrift 25:300. 1903.1 For other synonym.s see Barr ( 1959). Ascocarps thickly scattered, immersed- subcuticular, globose, papillate, glabrous, 50- 75 |jin diameter, 55-75 )im high. Ascocarp wall of textura globulosa in surface view: in longitudinal section unifonnly 7.2-10.8(-l2.7) \xm thick, composed of 2-3 layers of pseudoparenchymatic. polygonal cells, outer 1- 2 layers of brown-melanized cells (3.6-5.6 x 5.6-10.1 |im) covered by a brown-pigmcnted crust, inner 1-2 layers of hyaline, elongate- compressed cells (2.1-3.6 X 8.6-10.1 |im). Papilla short, erumpent, bluntly conical, 10-20 fim high, 10-21 |im wide at apex, 21-29 )im 512 Illinois Natural History Survey Bulletin Vol. 34 An. 5 if f h Figure 16. Leptospluu i iiltna pukhra. a. SEM of ascocaqi. x250. b. ascocarp wall surface. x350. c. d. longi- ludinal median section through ascocarps. x650. e. longitudinal median sections through ascocarp wall. x930. I. ascus. X 1.0.30. g, h. ascospores, x 1.1 50. a, b. d. and f from holotype oi Lcptosphacna oligotlwca: c. e.g. and h from neotype of Leptosphaerulina pulchra. May 1992 Leptosphaeria Species on the Rosaceae 513 wide at base, neck wall 3.6-8.6 nm thick, composed of 2-3 layers of isodiametric- elongate cells (3.6-4.3 x 3.6-6.5 |im) sur- rounding a circular ostiole (9-12 |im wide) without periphyses. Remnants of interthecial tissue remain between and above the asci. Asci 32-45 X 15-20 |im. few. basal, fasciculate, ovoid-saccate, thick-walled, sessile, rounded apex, without apical chamber, with 8 tri- to tetraseriate ascospores. Ascospores 17.2-20.8 X 5.7-7.2 |im, clavate to obovoid, bipolarly asymmetrical with broadly rounded apical end cell and acutely rounded basal end cell, straight; 4(-5)-septate, septa unevenly distrib- uted, order of septation 3: 1 :2:4, primary septum supramedian and constricted, a single vertical septum occasionally present in the central cells; subhyaline to pale brownish yellow, wall smooth, without sheath or appendages. Holotype: Austria: Ad caules foliaque aridos Potentillae caiilesceiuis. "am Martinstein bei Seis in Tyrol" 1870 von v. Hausmann gesammelt (B), (n.v.), presumed destroyed. Neotype: United States: Maine: Basin Pond, Baxter St. Park, July 5, 1962, MEBB # 3316 (NY). Other material examined: Canada: Labrador, Newfoundland, Blanc Sablon, July 19, 1957, R.T. Wilce#161 (NY). Iceland: on Potentilla maculata. type material of Laesladia potentillae Rostrup. July 14, 1884 (Holotype of Leptosphaeria olifiotheca. C). United States: Maine: Mt. Katahdin, Baxter St. Park, Table- lands from Saddle Trail, ca. 4200', Aug. 3, 1962. MEBB # 3546 (NY); New Hampshire: Mt. Washington, on Potentilla tridentata, June 9, 1894 (FH as Sphacrullna potentillae); Mt. Monadnock, near Dublin, June 27, 1961, MEBB # 2927 (NY). Comments: Leptosphaerulina pulchra is placed in the family Pseudosphaeriaceae because of the minute, sphaeroid-globose ascocarps with a very thin wall composed of pseudoparenchy- matous cells and the interthecial tissues. The species is recognized readily on the basis of its saccate asci, ascospore shape, and septation. Barr ( 1959) mentions the presence of a vertical septum in the central cells, but in most of the collections seen, vertical septa were lacking. Still, the obovate, asymmetrical ascospore shape is distinctive. The holotype of Leptosphaerulina pulchra at B. not located, is presumed to have been destroyed with the rest of the ascomycete collection during World War II. Also unsuc- cessful were attempts to locate authentic material of Winter or of the collector, von Hausmann. Because of this, a neotype for L. pulchra was chosen (Maine: Basin Pond, Baxter St. Park. July 5, 1962. MEBB # 3316) from the M.E. Barr collection now at NY. The neotype reflects both the original description and the current, well-established concept of this species. Leptosphaeria oligotheca is synonymous with Leptosphaerulina pulchra. When Petrak and Sydow ( 1924) originally described Leptosphaeria oligotheca from material on the type of Laestadia potentillae. on stems of Potentilla maculata. they did not segregate any of the material as the holotype for L. oligotheca and none could be found with their herbarium specimens at W or S. The type specimen of Laestadia potentillae at C contained abundant material matching the description of Leptosphaeria oligotheca. so a portion was segregated as the holotype of that fungus. Barr (1959) mentions several other species whose descriptions suggest they may be synonymous with Leptosphaerulina pulchra. One of these was Sphaeria minima Duby in C. Roumeguere, Fungi Selecti Gallici Exsiccati, Century 7, No. 694. Anno 1880. Examining specimens from the exsiccatae sets at G, FH, NY, and ILL showed that they included many ascocarps. but none contained asci or asco- spores. Therefore, it was impossible to resolve the placement of this species. •Diadema ohtusa R.A. Shoemaker & C.E. Babcock, Canadian Journal of Botany 67: 1353-1354. 1989. See Figure 17. Ascocarps thickly scattered, immersed- subcuticular becoming erumpent, globo.se, glabrous to slightly tomentose, with smooth brown hairs, 100-150 |im diameter, 120-150 |im high. Ascocarp wall of lextura angularis in surface view; in longitudinal section (7,5-) 10-18 |jm thick at sides and ba.se, composed of 2-3 layers of brown-melanized, polygonal, pseudoparenchymatic cells ( 1 .8-4.3 x 7-1 1 514 Illinois Natural History Survey Bulletin Vol. 34 An. 5 ^-^'v k^ -r \ ^ c e f Figure 17. Diadcma ohiusa. a. longitudinal median section through ascocarp. x2.'iO. b. ascocarp wall surface. x.'i.'SO. c. hyphae on substrate surface. x.'i.'iO. d. longitudinal median section through ascocarp wall, )^^)%), c. ascus. x515. f. ascospore. xl.O.SO. All from collection 12.^871. on Polciuilla agrophylta. (.1111). Apex broadly rounded, 18-27 (.mi thick. 4.'S~6.S |.ini diameter, composed of 2-3 outer layers of brown-nielani/ed. isodiamelric cells (.^.6-5.4 |im diameter) with 6-8 layers of hyaline pseudoparenchyma below center, opening area not seen, possibly discoid, caplike opening lacking, ostiole not seen. Interthecial tissues present. Asci (100-)120-145 x (30-) 5{)-70 (.im, few. basal, fasciculate, ovoid- saccate to cla\ate. thick-walled, short-stalked, rounded apex, w ith apical chamber, with 8 tri- to tetraseriate ascospores. Ascospores (41.5-) 4.';-50(-.'i5) X (15-1 7-) 19-23 \.im. broadly fusifomi. acutely rounded end cells, second cell May iy92 Leplosphaiita Species on Ihe Rosaceae 515 enlarged, straight to slightly curved, occasion- ally flattened in one plane: 3-septate. septa slightly unevenly distributed, order of septation 2:1:2, primary septum median and constricted, slightly constricted at other septa: brown, wall smooth, sharply delimited sheath entirely surrounding spore (5-7.5 |im thick), without appendages. Holotype: India: Kashmir: 123831(b). on Trisetiim spicatiim. Pensi La, Zaskar, 16,500 ft. W. Koelz (5906). 23 July 1933. TYPE, ex Herb. Wehmeyer, as Leptosphaeria hollosiana nom. nov. (DAOM). Other material examined: India: 123871, on Potentilla agrophxUa Wall.. Spiti Valley. Bara Lacha Pass, 16,500 ft, Bhagwan Singh (5), 8 July 1932, ex Herb. Wehmeyer, as Leptosphaeria hollosiana nom. nov. (DAOM). Comments: Shoemaker and Babcock ( 1989a) established the new genus Diadcma for a group of alpine fungi with relatively large, very dark brown ascospores. They did not place the genus within a family or order. Eriksson and Hawksworth ( 1990) placed it in the Dothideales but did not designate a family. In the present report I treat it under the Pseudosphaeriaceae because it resembles Wettsteinina Hohnel: for example, some species have a peculiar disclike opening of the ascocarp and have interthecial tissue in the centrum. Diadema is characterized by globose, subcuticular ascocarps with a thin wall of brown polygonal cells and broadly fusiform ascospores that are dark brown and have a prominent sheath. Wehmeyer ( 1963) published the name Leptosphaeria hollosiana for Leptosphaeria maritinia L. Hollos because the latter binomial was predated by L. maritima (Cke. & Plowr.) Sacc. He applied the new name to several collections from India and Pakistan, including two collections on Potentilla. Shoemaker and Babcock ( 1989a) described two new species of Diadema from these collections, including the type of the genus, D. acuta, on Trisctiim spicatiim (L.) Richt.. and D. ohtusa. also on that host. The collections on Potentilla were also determined to be D. ohtusa. The collection on Potentilla served as the basis for this descrip- tion and plate because of the Rosaceous host and because it had numerous ascocarps. The other collections of D. ohtusa were sparse. As Shoemaker and Babcock ( 1989a) noted, the ascocarp opening mechanism in this species is not clearly understood. It is not distinctly caplike but in vertical section becomes almost papillate (Figure 17a). The entire upper section of the wall is subtended by hyaline pseudoparenchyma. Interthecial pseudoparenchymatic threads occurred between the asci. Also, coarse, brown hyphae were seen on the host surrounding several of the asco- carps, and some of the ascocarps were not glabrous but had tapered hairs at the top. I did not see the ascospores flattened in one plane, as did Shoemaker and Babcock (1989a). Shoe- maker and Babcock (1989a) suggested that the generic placement of this species is not ideal because several characters were not identical to those of the type species. The overall aspect of this species, however, suggests an affinity with the type species, and that it is adequately placed in this genus. It appears that certain characters, such as the ascocarp cap and interthecial tissues, are variable among collections and may depend on the maturity of the specimen, as was also noted by Shoemaker and Babcock (1989a). Species of Wettsteinina are similarly variable, either showing a disclike cap or an ostiolar opening to the ascocarp: species with either character state are adequately retained within the genus because of overall resemblances (Shoemaker and Babcock 1987). Thus, such variation does not appear sufficient to exclude species from these genera. 'Diadema sieversiae (C.H. Peck) S.M. Huhndorf comb. nov. See Figure 18. = Lophiostoma sieversiae C.H. Peck, New York State Museum Bulletin 167:44. 1913. (Basionym). = Wettsteinina sieversiae (C.H. Peck) M.E. Barr. Canadian Journal of Botany 45:1042. 1967. = Massaria sieversiae F.E. Clements. Cryptogamae Formationum Coloradensium, Century 111. No. 234. Anno 1906. Nom. nud. Art. 36.1. = Leptosphaeria sieversiae (F.E. Clements) F. Petrak. Sydowia. Annales Mycologici 6(l-4):6. 1952. [Combina- tion is not \ alid because the basionym is an invalidly published species.] A.scocarps thickly scaitcretl. immcrsed- subcuticular becoming crumpent. globose, 5)6 linois Natural History Survey Bulletin Vol. 34 Art. 5 Figure 18. Diatlema sieveisiae. a. SEM of erumpent ascocarps. xl2,'i. b. ascocaip wall suiface. x550. c. longitudinal median section through ascocarp, x2.^0. d. hxphae on substrate surface. x200. e. elongated ascus. x2.'S(). f. longitudinal median section through ascocarp wall. x950. g. ascus. x525. h. ascospore. X I .O.'iO. a, b from lKilot> pe of D. sicvcr.siac: c-h IVoni isot> pe of Ma.ssaria sieveisiae. May 1992 LvplDsphavrui Species on the Rosaceae 517 glabrous to slightly tomentose, with loose dark- walled, longitudinal hyphae on the substrate, surrounding and connecting the ascocarps, 130-190 (im diameter, 120-175 ^m high. Ascocarp wall of textura angularis in surface view; in longitudinal section uniformly 12.6-20 (im thick, composed of 3—1 layers of brown- melanized, pseudoparenchymatic, polygonal, isodiametric-elongate cells (3.6-5.6 x 7.2-13 )im) with a brown-melanized crust, at base 3—4 inner layers of small, hyaline pseudoparen- chyma. Apex broadly rounded, with inner 3—4 layers of hyaline, pseudoparenchymatic cells (2-5 nm diameter), no evidence of papillate or caplike opening, ostiole not seen. Interascal pseudoparenchyma present. Asci ( 1 10-) 130- 170 X 40-70 (im, few, basal, fasciculate, ovoid- saccate, thick-walled, sessile, rounded apex, with apical chamber, with 8 tri- to tetraseriate ascospores. Ascospores (43.5-)50.6-56.0 (-57.9) X (16-)20-23.5(-25.5) Mm, broadly fusiform, with acutely rounded end cells, second cell enlarged, straight to slightly curved; 3-septate, septa slightly unevenly distributed, order of septation 2: 1 :2, primary septum slightly supramedian and constricted, occasion- ally constricted at other septa; brown, wall smooth, sharply delimited sheath entirely surrounding spore |2.0-5.5(-l0.5) (im thick, often constricted at midseptum, sheath occa- sionally gelatinizing and becoming amorphous (7-12 (im thick)], without appendages. or disclike opening mechanism in D. sicversiac. Interthecial tissues were seen between the asci. Diadenia sieversiae was described by Peck ( 1913) as a species of Lophiostomu, but it bears no compressed beak or ostiole that would warrant inclusion in that genus. Barr (1967) transferred it to Wettstelnina because of the medium- to small-sized ascocarps immersed in dead host tissues and the oblong to saccate asci interspersed with interthecial tissue. Shoe- maker and Babcock ( 1987) excluded it from Weltsteinina because of the dark brown ascospores with thick septa that are unlike the usual spores for Wettsteininci. It was left in Lophiosroma and later (Shoemaker and Babcock 1989a) was not included in Diadema. Clearly this fungus does not belong in Lophiostoma, and it bears a much greater resemblance to Diadema than to Wettsteinina. Even if it ultimately does not stay in Diadema, because of the lack of a disclike cap opening (as suggested for D. ohriisa by Shoemaker and Babcock 1989a), it seems useful to place this species in Diadema. in the hope that additional collections may be discovered. Massaria sieversiae, which was trans- ferred to Leptosphaeria. is synonymous with Diadema sieversiae. Holotype: United States: Utah; 193727, on Sieversia tiirhiiiata (Rydb.) Greene, Big Cottonwood Canyon, Salt Lake Co., A.O. Garrett 702, 3 July 1913 (NYS). Exsiccatae; United States: Colorado: Bottom- less Pit. July 13, 1906, Clements, Crypt. Form. Colorad. 234 (isotype of Massaria sieversiae, NY, FH). Comments: Diadema sieversiae. found on stems of Acomastylis liirhinata. is characterized by globose ascocarps with thin walls of brown polygonal cells, dark-walled hyphae on the substrate, and broadly fusifomi. dark brown ascospores with a prominant gelatinous sheath. It bears a strong resemblance to D. ohtiisa. but in that species the spores arc slightly shorter and the dark brown hyphae in the host arc not as prevalent. The ascocarp shapes differ somewhat, but 1 also saw no indication of a cap Leptosphaeria Species Referable to the Hymenoascomycetes The following Leptosphaeria species are all referable to the class Hymenoascomycetes (sensu Barr 1987a). The presence of perithecia and unitunicate asci separates these fungi from those in the Loculoascomycetes possessing pseudothecia and bitunicate asci. The families represented are Clypeosphaeriaceae, Diaporthaceae, and Amphisphaeriaceae. Clypeosphaeriaceae 'Clypeosphaeria mamiUana (E.M. Fries:E.M. Fries) J.B.E. Lambotte, Memoires de la Societe Royale des Sciences de Liege, ser 2. 14:128. 1887. See Figure 19. = Sphaeria mamiUana E.M. Fries:E.M. Fries, Systema Mycologicum 2:487. 1823. = Sphaeria clypeiformis G. de Notaris, Memorie della Accademie della Scienze di Torino Series 2, No. 7, p. 1 13. 1853; Micromycetes Italici Novi vel Minus Cogniti 7, p. 113. 1845. Non Sphaeria clypeiformis L.V. de Lacroix in G.L. Rabenhorst. = Clypeosphaeria notarisii L. Fuckel, Symbolae Mycologicae, p. 117. 1870. = Leptosphaeria (Clypeosphaeria) notarisii M.C. Cooke, Grevillea 17:91. 1889. See Barr (1989) for other synonyms. Ascocarp.s .scattered, separate or gregarious, inmiersed-subepidcmial beneath blackened clypci, papilla cniiiipont, glabrous, conic- globose, 300-400 |am diameter, 400-450 |jm high. Ascocarp wall of textura angularis in surface view: in longitudinal section unifomilv 1 8 25 pm thick, composed of 10-15 layers of hyaline, elongate-compressed, scleroplecten- chymatic cells (0.7-1 .5 x 7-15 |im), outer 2-3 layers of hyaline, rounded. pol\gonal cells (2.0-3.6 pm diameter). Papilla conic, 130-160 pm high, .50-75 pm wide at the apex, 150-225 pm wide at the base: wall 15-30 pm thick at apex, 35-65 pm thick at base, composed of 10- 15 layers of small, brown-melanized, thick- walled, rounded, polygonal cells at base (1.5- 3.6 pm diam), cells converging and appearing as elongate, setaelike structures at apex, surrounding a 25-50 pm w ide circular ostiole with periphyses. Paraphyses 0.5-1.5 pm wide, numerous. Asci 150-170 x 8-10 pm. unitunicate, numerous, basal to peripheral, cylindrical, short-stalked, rounded ai>ex, apical ring i+. with 8 overlapping uniseriate ascos- pores. Ascospores 21-26 x 5.7-8 pm. ovoid- oblong, straight to slightly cur\ed: at times 1- septate near base, large cell appearing 3-septate with .separation of cytoplasm: brow n. smooth, without sheath or appendages, germ slits not seen. Exsiccatae: Austria: On Riihiis fruticosus. Fuckel, F. rhen. 1823 (FH); Mappen. on Epilohiitm angiistifolii. as Sordaria clypeiformis f. Epilohii. Fuckel, F. rhen. 2036 (FH); Herb. Barbey-Boissier 146 = F. rhen. 1823, as C. notarisii (NY); Great Britain: Lynn, on Epilohium hirsutum. as Sphaeria clypeata Nees., Jan. 1877, Plowright, Sphaer, Brit. 57 (FH); Italy: on Ruhus fruticosus. as Sphaeria clypeiformis. .Aug. 1857, leg. Caldesi, Rabenhorst. Herb. myc. 645 (FH); as Sphaeria clypeiformis. 1857. leg. Caldesi. de Notaris. Erb. Critt. Ital. 47 (FH); La Spezia, as C. notarisii. Aug. 1880. Roumeguere, F. Gall. exs. 2095 (NY); Vittorio (Treviso"), on Ruhus fruticosus. as C. notarisii. Sept. 1897. Saccardo. Mycotheca ital. 101 (NY); Conegliano, as C. notarisii. Aug. 1877, Mycotheca \eneta 1 159 (NY): St. Romani-ad- Vigennam, as S. clypeiformis de Lcrx., 1859. leg. T. de Lacroix. Rabenhorst, F. europ. 331 (RO). Other materia] examined: Italy; Capraria, on Ruhus. as Sphaeria sepincola ruhorum. 1837. May 1992 Lepiosphaeria Species on the Rosaceae 519 Km»i' a b <^\ u^ *iiii % .e Figure 19. Clypensphaeria mamillana. a. longitudinal median section through ascocarp. xlOO. b, c. asco- spores. xl,150. d. longitudinal median section through ascocarp wall, x950. e. ascus. x523. All from lectotype of C. nolarisit. de Notaris (Lectotype of Sphaeiia clvpeiformis): two collections of Sphaeria ciypeaia. 1840; Spezia. Aug. 1857. Herb. N.A. Pedicino, 1 883; on Riihus. as S. chpeiformis. Mar. 17, 1842, de Notaris (all RO). Comments: Clypeosphaeria notarisii, a synonym of Clypeosphaeria mamillana, is characterized by cylindrical asci with an elongate, J+ apical ring and curved, brown ascospores appearing .3-septate with separation of the cytoplasm. Barr (1989) gives the synonymy and a short history of Clypeosphaeria and argues for the utilization of the family Clypeosphaeriaceae. Clypeosphaeria was established by Fuckel ( 1870). who included C. notarisii as a new name for Sphaeria chpeiformis de Not. A specimen of Sphaeria sepincola ruhorum was mentioned in the description of Sphaeria chpeiformis. and a collection of this at RO is here designated as the lectot) pe of .V. chpeiformis (Italy: Capraria. on Ruhus. 1837, de Notaris, RO). Ilinois Natural Hislory Survey Bulletin Vol. 34 An. 5 Diapurthaceae 'Diaporthe eres T.R.J. Nit.schke. Pyrenomy- cetes germanici, p. 245. 1867. See Figure 20. = Spliaeria controversa J. Desmazieres (pro parte), Annales des Sciences Naturelles, Paris, Botanique, Series 2, 17:102. 1842. = Leptosphaeria controversa (J. Desmazieres) V. Cesati & G. de Notaris, Commentario della Societa Crittogamologica Italiana 1:235. 1863. = Diaporthe controversa (J. Desmazieres) T.R.J. Nitschke in L. Fuckel, Symbolae Mycologicae Nachtrag 1:319. 1871. See Wehmeyer (1933) for other synonyms. Ascocarps scattered, separate or clustered, immersed-subepidermal often beneath black- ened stromatic tissue, papilla erumpent singly or in loose clusters, glabrous, globose, flattened at top and ba.se, 300-400 \im diameter, 250- 350 |im high. Ascocarp wall of textura angularis in surface view; in longitudinal section 25-35 \xm thick at the sides. 15-20 |im thick at the base, composed of 7-8 layers of hyaline, elongate-compressed, scleroplecten- chymatic cells (0.7-1.5 x 10-15 |Xm), wall near apex 45-50 |im thick, composed of 10-12 layers of brown-melanized. polygonal, isodia- metric cells (2-3 \xm diam). Papilla long cylindrical, central or eccentric, 125-180 |im high, 50-75 (im wide; wall 25-35 |im thick, composed of small, brown-melanized, thick- walled, rounded, polygonal cells ( 1.5-3.0 )im diameter), surrounding a 25^0 |im wide circular ostiole with periphyses. Paraphyses 0.5-1.5 )am wide, numerous, septate, un- bratiched. Asci 38-55 x 6-8.5 |im, unitunicate, numerous, basal to peripheral, dehiscent, clavate, short-stalked, rounded apex, with 8 biseriate ascospores. Ascospores 10.5-13.5 x 2.0-3.6 i^m, fusiform, .straight to slightly curved; 1 -septate, constricted; hyaline, with four distinct oil droplets, smooth, without sheath, occasionally with small, cellular bipolar appendages. Isoiype: pRANrn: Habitat in caulibus plantarum. in ramis exsiccatis F-'raxini, Rubi, Aceris Negundinis, etc. Desm. PI. Crypt. Fr. exs. XXVI 1255(PC, FH. BPI). Comments: Sphacria controversa (pro parte) is synoiwinous with Diaporthe eres (Wehmeyer 1933) and is characterized by ascocarps immersed beneath blackened stromata, with central or eccentric, long cylindrical necks with periphyses. As they mature, the numerous asci loosen and become free \v ithin the centrum. Ascospores are hyaline and 1 -septate and have four distinct oil droplets. The exsiccati collections all consisted of several different twigs of unknown identity, none of which appeared to be Ruhiis. most with varying amounts of the fungus on them. Amphisphaeriaceae 'Discostroma fuscella (M.J. Berkeley & C.E. Broome) S.M. Huhndorf comb. nov. See Figure 21. = Spliaeria (Ohtectae)fusceUa M.J. Berkeley & C.E. Broome. Annals and Magazine of Natural History . London. Series 2, 9:325. 1852. (Basionym). = Leptosphaeria fuscella (M.J. Berkeley & C.E. Broome) V. Cesati & G. de Notaris. Commentario della Societa Crittogamologica Italiana 1:236. 1863. = Sphaeria corticola L. Fuckel. Symbolae Mycologicae. p. 114, 1870. = Leptosphaeria corticola (L. Fuckel) P.A. Saccardo, Michelia Commentarium Mycologicum Fungos in Primis Italicos Illustrans 1:342. 1878; Fungi Italici autographice delineati (additis nonnuUis extra-italicis asterisco notatis). Patavii, table 288. 1878. = Metasphaeria corticola (L. Fuckel) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2: 166. 1883. = Griphosphacria corticola (L. Fuckel) F. V. Hohnel. Annales Mvcologici 16:87. 1918. = Chithridium corticola (L. Fuckel) R..-\. Shoemaker & E. Miiller. Canadian Journal of Botany 42:404. 1964. = Discostroma corticola (L. Fuckel) 1. Brockmann. Sydowia 28:313. 1975. = Sphaeria {Leptosphaeria) leioste\ia J.B. Ellis, Bulletin of the Torre\ Botanical Club (and Torreya) 8:91. 1881. ' = Leptosphaeria leiostega (J.B. Ellis) J.B. Ellis, Catalogue of Plants Found in New Jersey. Geological Sur\ e\ of New Jersey, Final Report of the Slate Geologist 2(1):525. 1889. May 1992 Leplosphaeria Species on ihe Rosaceae 521 Figure 20. Diaporthe eres. a. longitudinal median section through ascocarp. x 150. b. section through ascocarp neck showing periphyses, x40(). c. longitudinal median section through ascocarp wall, x950. d. ascocarp wall surface. x550. e. ascus. xl,()50. f. ascospore, xl,l5(). All from type of Leplosphaeria conlrover.ta. = Leplosphaeria lejostei^a ( J.B. Ellis) F. Hazslinszky, Matematikai es Termeszettudomanyi Kozlemenyek Vonatkozolag a Hazai Viszonyokra 25(2):148. 1892. = Metasphaeria lejostet^a (J.B. Ellis) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:164. 1883. For other synonyms see Brockmann (1975). Ascocarps scattered, separate or clustered, immersed-subepidermal beneath blackened clypei, glabrous, depressed globose. Ilattened top and bottom, with papilla protruding through surface, 3()0-5()() |im diameter, 150-250 ^m " high. Ascocarp wall of textura prismatica in surface view; in longitudinal section 8-18 |im thick at base, 18-27 |im thick at sides, com- posed of 6-8 layers of brown, elongate- compressed, scleroplectenchymatic cells (12- linois Natural History Survey Bulletin Vol. 34 Art. 5 N- V ^-.» '^**^^%^ lf\ ^ i fir L V «» ^- ^ '^> HP •^^ av* I Figure 21. Disiosinmui fiisd-lla. a. longituditial median section through ascocarp. xl.'iO. b. ascocarp wall surface, x.S.SO. c. section through ascocarp wall neck region, x.'i.^(). d. longitudinal median section through \aip. xl5(). e. longitudinal median section through ascocarp wall. x^50. f-h. asci. x.'>2.'>. i-k. ascospores. ' ;i-<-. e. t", and i from holoty pe of D.fii.sicllu: d. g. and j from lectotype of Spluicria leiostega: I'lypeof S. corticola. May 1992 Lepiosphaeria Species on the Rosaceae 523 25 X 2-3 |Jm); near apex, wall 21-36 \xm thick, composed of 6-7 outer layers of brown, polygonal-isodiametric cells (7-1 1.5 x 2-3; 2.1-3.6 |im diameter) and 6-7 inner layers of compressed-elongate, scleroplectenchymatic cells. Papilla conical, 50-75 ^m high, 20-30 |im wide at the apex. 35-50 |im wide at the base; wall 7-10 |im thick, composed of 7-10 layers of small, brown-melanized, thick-walled, rounded, polygonal cells, surrounding a 10-20 )im wide circular ostiole with periphyses. Paraphyses 0.5-1.5 |im wide, numerous, septate, unbranched. Asci (lOl-)l 15-132 (-137) X 7-12(-14) |im, unitunicate, numerous, basal in a broad hymenium, cylindrical, short- stalked, rounded apex, apical ring J-i-, with 8 overlapping uniseriate ascospores. Ascospores 15-21 X 5-9.5 urn. ellipsoidal, end cells rounded; 3-septate, septa often close together and end cells longer, seldom with 4 or more septa (up to 7), occasionally with vertical septa in central cells only or in every cell, order of septation variable, often 2; 1:2, primary septum median and occasionally constricted; hyaline, smooth, without sheath or appendages. Holotype: Great Britain: Easton, Norths., on dead twigs of rose (Rosa sp.), March 9, 1850 (K). Exsiccatae: Austria: Fuckel, Fungi Rhenani 91 1 (Isotype of Sphaeria cortkola, G, FH); Czechoslovakia: Welka, on Pnimis spinosa. as Metasphaeha cortkola f. Pruni. Oct. 3, 1912, Petrak, Fl. Boh et Mor. exsic. 61 (FH); Welka, on Rosa canina. as Metasphaeria coriicola f. Rosae, Apr. 17, 1912. Petrak, Fl. Boh et Mor. exsic. 62 (FH); Ohrensdorf. on Riihiis ihyrsoideus. as Metasphaeria corticola f. Ruhi, Sept. 25. 191 1. Petrak. Fl. Boh et Mor. exsic. 63 (FH); Sweden: Lockerud, Wenenborg, on Ruhusfrnticosus, as Lepiosphaeria lejostega (Ellis) Eliasson. Aug. 1892. Rehm. Asc. 1 189 (FH); United States: New Jersey: Newfield, as Sphaeria (Lept.l lelostega, Ellis, N. American Fungi 888 (NY). Other material examined: Austria: Nassau, on Pnimis domestica and P. spinosa. Fuckel (Isotype, as Metasphaeria corticola (Fuckel) Sacc, FH); United States: Kansas: Rockport, Rooks Co., on Cornus asperlfolia. as Metasphaeria lelosteiia. Dec. 24. 1894. E. Bartholomew (FH); Rooks Co., on Rlhes aureum, as Clypeosphaeria hendersonia 2555, Mar. 15, 1899. E. Bartholomew (NY); Michi- gan: 573, Ann Arbor, on Rlhes frondium, as Metasphaeria leiostega. Apr. 1. 1893, L.N. Johnson (NY); New Jersey: Newfield, on wild rose (Rosa haida ?), May 6, 1879, J.B. Ellis (lectotype of Sphaeria leiostega. NY); Newfield, on Vlhurnum lentago, as Sphaeria leiostega. Dec. 1878, J.B. Ellis (NY); Newfield, Gloucester Co.. on Hickory, as Sphaeria lelostega (NY); on Staphylea trifolia, as Metasphaeria lelostega, Dec. 23. 1891 (NY); on Viburnum, as Metasphaeria lelostega (NY); 1273, on wild currant, as Metasphaeria lelostega. Jan. 4, 1890 (NY); on Comptonia, as Metasphaeria lelostega Mar. 1890, J.B. Ellis (NY); Vineland, on Vacclniiim corxmhosum. as Sphaeria lelostega. Jan. 22, 1878; Jan. 10, 1880 (NY); New York: Washington Co., Vaughans, J. Vaughan's Copse, near Devines Woods, north of Hudson Falls, on Rlhes americanum (floridum), as Metasphaeria lelostega. Apr. 24, 1917, P.P. 25255, S.H. Burham (FH). Comments: Discostroma fiiscella is character- ized by subepidermal perithecia with short protruding papillae, cylindrical asci with a J-i- apical ring and hyaline. 3-septate, elliptical ascospores with broadly rounded ends. Certain collections have wider spores and vertical septa, most often in the central cells, but often also in the end cells. Most notably, vertical septa are found in the lectotype of Sphaeria lelostega. All other characters are identical to those of D. fiiscella. Discostroma corticola. the most common name of this fungus, is identical to the earlier named Sphaeria fiiscella. Previous interpreta- tions of Sphaeria fiiscella have caused some confusion in the literature. The type material contains a fungus resembling D. corticola, with hyaline ascospores. However, it has been considered to be a 3-septate, brown-spored fungus because of Berkeley and Broome's ( 1852) description of ascospores as "pale brown, oblong-elliptic, obtuse, triseptate, sometimes slightly curved." In the accompany- ing Latin description they do not mention the color of the ascospores. On the type material (K). labeled "Sphaeria fiiscella B & B. In Rosa, Easton, March 9, 1850," are three rose twigs, one of which is labeled (with a dotted line and a figure) as Sphaeria iSuhtectae) intermixta B & B. The other two twigs contain abundant 24 Illinois Natural History Survey Bulletin Vol. 34 An. 5 material of the hyaline-spored fungus resem- bling D. corlicola. Also on this material is a figure oi Spliaeria fiiscella showing an ascus and spores with "hyaline" written next to the spores. Wehmeyer (1957) suggested that a mixture of fungi in this and another collection of S.fusrella resulted in a confused description. He felt the name should be a nomen dubium and not used as a basis for later names. I believe, from the description and the fungus found on the type material, and from the figure on the type material, that Berkeley and Broome's confused description was not based on a mixture of fungi but was a miscommunica- tion of the actual information. I believe they were aware that S.fuscella had hyaline, not brown spores; thus, it is suitable for use as a basis for later names. Species incertae sedis The following species were examined but could not be placed in any taxa with certainty because the type specimens contained no ascocarps and no other collections were available on which to base the descriptions. •Leptosphaeria eriohotryae H. Sydow, P. Sydow & E. Butler, Annales Mycologici 9:409. 1911. Holotype: Hab. in foliis Eriohotrya japonicae, Saharanpur. 27. 6. 1906, leg. Inayat (E.J. Butler no. 1365). (S) Isotype: (W, HCIO) 'Leptosphaeria minima (J.E. Duby) ex P.A. Saccardo. Michelia Commentarium Mycologicum Fungos in Primis Italicos Illustrans 2:320. 1881. = Sphaeria minima J.E. Duby in C. Roumeguere, Fungi Selecti Gallici Exsiccati, Century 7, No. 694. Anno 1880. Norn. nud. = Mycotodea minima (J.E. Duby ex P.A. Saccardo) W. Kirschstein in O.C. Schmidt, Kryptogamenflora der Mark Brandenburg und Angrenzender Gebiete herausgegeben von den Botanishen Verein der Provinz Brandenburg 7(3):432. 1938. Isotyf)e: Sur les feuilles du Potentilla argenlea L., Mende, Prost. (G, FH, ILL. NY) 'Leptosphaeria sorhi A. Jaczewski, Annales Mycologici 1:30. 19(J3. Holotype: in foliis vivis Sorhi aucupariae. IX. 18 1914 , Gjatsk. Smolensk. Gouv. Jaczewski. (LE) Appendix: Leptosphaeria Species Described from the Rosaceae The following is a list of all Leptosphaeria species described from the Rosaceae and the species" nomenclatural synomyms. This information is taken from Crane and Shearer ( 1991 ). The taxonomic synonyms applied herein and the herbaria contacted have been added. The fungal specimens cited in this dissertation were studied at or on loan from various herbaria. Requests were made to various herbaria for type material for all of the described species. Unfortunately, type material could not be located for all of the species because either the material was destroyed, lost, or not deposited at a certain institution, or the institutions involved did not reply or do not lend specimens. Abbreviations follow Holmgren etal.{ 1981). L. ahhreviaia (M.C. Cooke) P. A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:26. 1883. ^Spluieria ahhreviaia M.C. Cooke, Handbook of British Fungi, p. 893. 1871. On dead stems of Rubus sp., Rosaceae. Great Britain. =Saccolhecium sepincola (E.M. Fries:E.M. Fries) E.M. Fries var. ahhreviaia S.M. Huhndorf, comb. nov. L. acideorum G. Passerini, Atti della R. Accademia dei Lincei Memoriae. Rome, Series 4, 6:458- 4.59. (1889) 1890. On stems oi Rosa canina L., Rosaceae. Italy, (nonvidi). Type material was not found at PARMA, PISA, PAD. L. ariinci S.M. Zeller, Mycologia, Lancaster. Pennsylvania 19:134-135. 1927. On dead stems of Aninciis silveslris Kostel. (=AniiHiyrophylla Wallich ex Lehm., Poten!illafrai;arioidcs L., Triglochin maiilima L., Triselum spicalum (L.) K. Richter, Geraniaceae, Gramineae, Juncaceae, Rosaceae. India, (n. v.). Type material was not found at BP, PAV, PRE. L. leioslef;a (J.B. Ellis) J.B. Ellis, Catalogue of Plants Found in New Jersey. Geological Survey of New Jersey, Final Report of the State Geologist, Trenton 2( 1 ):525. 1889. ^Sphaeiia (Leplosphaeria) leiostef>a J.B. Ellis, Bulletin of the Torrey Botanical Club (and Torreya). New York 8:91. 1881. On various dead twigs and limbs of Caiya sp., Rosa sp., Vaccinium sp., Juglandaceae, Rosaceae, Ericaceae. United States (New Jersey). =Discoslroma fuscella (M.J. Berkeley & C.E. Broome) S.M. Huhndorf, comb. nov. L. lejosle^a (J.B. Ellis) F. Hazslinszky, Matematikai es Termeszetludomanyi Kozlemenyek Vonatkozolag a Hazai Viszonyokra, Budapest 25(2): 148. 1892. ^phaeria (Leplosphaeria) leiosle^a J.B. Ellis, Bulletin of the Torrey Botanical Club (and Torreya), New York 8:91. 1881. ^Melasphaeria lejosle^a (LB. Ellis) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:164. 1883. On branches of Ci/nu sp., Rosa .sp., Vaccinium sp., Juglandaceae, Rosaceae, Ericaceae. United States (New Jersey). L. leiicoplaca (K.F. Wallroth) E.A. Vainio, Acta Societatis pro Fauna el Flora Fennica, Helsingforsiae 49( 2 ): 1 43- 1 44. 1 92 1 . =\'erriicuria leiicophica K.F. Wallroth, Flora Cryptogamica Gcmianiae 1:299. 1831. =Pyrenula leiicoplaca (E.G. Wallroth) G.W. Koerber, Syslema Lichenum Germaniae. p. 361. 1855. =VeiriH aria aiha H. A. Schrader var. leiicoplaca (F.G. Wallroth) I.E. Schaerer in Enumeralio Critica Lichenum Europaeorum. Quos E.\ Nova Methodo Digerit, p. 219. 1850. =Eopyrcnula leiico- placa (K.F. Wallroth) R.C. Harris, The Michigan Botanist, Ann Arbor 12:19. 1973. 5?8 Illinois Natural History Survey Bulletin Vol. 34 An. 5 On bark of Acer sp., AInus sp., Caiya sp., Corylus sp., Fraxinus sp., Julians sp., Populus sp., Quercus sp., 5a//.v sp., Sorhus sp., 7"/7/« sp., Utmus sp., Aceraceae, Juglandaccae, Betulaceae, Oliaceae, Salicaceae, Fagaceae, Rosaceae, Tiliaceae, Ulmaceae. Finland, (n. v.). No reply from ,STR. L. Iiuilhi P. A. Saccardo, Nuovo Giomale Botanico Italiano e Bolletino della Societa Botanica Italiana, Firenze 7:310-311. ISV.'i. Indiscrimi- nately on wilted leaves of Pyrus communis L.. Rosaceae. Italy. =Phaeosphaeriu liiciUa (P.A. Saccardo) S.M. Huhndorf, comb. nov. L. mandshiirica M. Miura, Flora of Manchuria and East Mongolia, Part III, Cryptogams, Fungi (Contributions of the Southern Manchuria Railway Company 27): 175. 1928. On leaves of PxiHs mains L. |as Mains domestica Borkh. I, Rosaceae. China, (n. v.). Type material was not found at TI, TKB. No reply from TFM, TOFO, TNS. L. minima (J.E. Duby) ex P.A. Saccardo, Michelia Commentarium Mycologicum Fungos in Primis Italicos Illustrans 2:320. I88I. =Spluieria minima J.E. Duby in C. Roumeguere, Fungi Selecli Gallici Exsiccati. Century 7, No. 694. Anno 1880. Nom. nud. =Mycol<)dea minima (J.E. Duby ex P.A. Saccardo) W. Kirschstein in O.C. Schmidt, Kryptogamenllora der Mark Brandenburg und Angrenzender Gebiete herausgegeben von den Bolanishen Vcrein der Provinz Brandenburg, Leipzig 7(3):432. 1938. On leaves of Powntilla argenlea L., Rosaceae. France. Species incerlae sedis. L. miyakeana P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 24:999. 1928. =Phaeosphaena eriohotiyae I. Miyake, Botanical Magazine, Tokyo 27:41-42. 1913. =TrematosphaereUa erioholiyae (I. Miyake) F.L. Tai, Sylloge Fungorum Sinicorum. p. 330. 1979. On leaves of Eiioholiya japonica (Thunb.) Lindlcy, Rosaceae. China, (n. v.). Type material was not found at TI, TKB. No reply from TFM. TOFO, TNS. It is not known whether type material is at PAD since this institution has stopped lending specimens until further notice. L. nuinic K. Hara, Byochu-gai Zasschi (Journal of Plant Protection), Tokyo .5:884. 1918. (As miinc in original publication.] On dead branch of Pniniis iniimc Sieb. & Zucc, Rosaceae. Japan, (n. v.). Type material was not found at TI, TKB. No reply from TFM, TOFO, TNS. L. nashi (K. Hara) P.A. Saccardo in A. Trotter, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 24:999. 1928. =Leplosphaerella nashi K. Hara, Kajyu-byogai-ron Nihon-kankitsu-Kai, Schiznoka (Fruit Tree Diseases), pp. 142-145. 1916. =Phaeospnra nashi (K. Hara) K. Hara, Jitsuyo-Sakumotsu-Byorigaku (Manual of Plant Pathology], p. 270. 1925. On leaves of Pyrus sinensis Hon.. Ko^acede. Japan, (n. v.). Type material was not found at TI. TKB. No reply from TFM, TOFO. TNS. L. (Clypeosphaeria) norarisii M.C. Cooke, Grevillea. London 17:91. 1889. ^phaeria clypeiformis G. de Notaris. Memorie Della Accademia delle Scienze di Torino, Series 2. No. 7, p. 1 13. 1853; Micromycetes Italici Novi vel Minus Cogniti 7. p. 113. 1845. Non Sphaeria clypeiformis L.V. de Lacroix in G.L. Rabenhorst. ^lypeosphaeria notarisii L. Fuckel. Symbolae Mycologicae, p. 117. 1870. On barky runners of Riihiis fruticosus L., Rosaceae. Italy. =Clypeosphaeria mamillana (E.M. Fries: E.M. Fries) J.B.E. Lambotte. L. ohesula P.A. Saccardo in V. Mouton, Bulletin. Societe R. de Botanique de Belgique, Bruxelles 26:176. 1887. On stems of 4ff/- pseiidoplatanus L., Angelica sp.. Spiraea ulmaria L. (=Filipendida idmaria Maxim.), Aceraceae. Umbelliferae, Rosaceae. Belgium. (n. v.). Type material was not found at DUKE. No reply from AUT, CGE. LD. It is not known whether type material is at PAD because this institution has stopped lending specimens until further notice. L. oligolhcca F. Petrak & H. Sydow, Annales Mycologici, Berlin 22:359. 1924. (Described from the type material of Laestadia polenlillac E. Rostrup, Botanisk Tidsskrift. Kjobenhavn 25:300. 1903.] On leaves of Poteniilla macidala Pour. i.=P(Hentilla aurea L). Rosaceae. Denmark. Iceland. =Lcpti>sphaeridiiia pidchra (H.G. Winter) M.E. Ban-. L. (Clypcdsphaeria) oscidanda (C.G.T. Preuss) M.C. Cooke. Grevillea, London 17:91. 1889. Sphaeria osculanda G.T. Preuss, Linnaea. Institut botanique de I'Universite de Geneve 26:713. 1853. ^lypeosphaeria osculanda (G.T. Preussl P..^. Saccardo. Sylloge Fungorum Omnium Hucusque Cognitorum Digessit PA. Saccardo 2:91. 188.^"^ On branches of Rubus sp.. Rosaceae. Germany. (n. v.). Type material was not found ai B. L. o.xyspora J. Feltgen. Vorstudien zu einer Pilz- Flora des Grossherzogthums. Luxemburg I(2):165-I66. 1901. On dry stems of /fii/jiw idaeus L., Rosaceae. Luxemburg, (n. v.). May 1992 Leptosphaeria Species on the Rosaceae 529 Type material Is probably at LUX but was not obtained because the collections are undergo- ing reorganization. L. pachythcca P.A. Saccardo. Nuovo Giomale Botanico Italiano e Bolletino della Societa Botanica Italiana, Firenze, New Series 22:39. 191.'). Norn, illeg. Art. 64.1. On dead branches of Amywdaliis communis L. [=Pnuuis chillis (Mill.) D.A. Webb), Rosaceae. Yugoslavia, (n. v.). It is not known whether type material is at PAD because this institution has stopped lending specimens until further notice. L. (Lepiospluieiella) pomonci P.A. Saccardo, Nuovo Giomale Botanico Italiano e Bolletino della Societa Botanica Italiana, Firenze 8; 176. 1876. On upper surface of leaves oi Pyius mains L. [-Malus domestica Borkh.), Rosaceae. Italy. =Phaeosphaeria pomona (P.A. Saccardo) S.M. Huhndorf. comb. nov. L. pomona P.A. Saccardo forma Iransilvanica A. Negru et I. Ditzu, Notulae Systeniaticae e Sectione Cryptogamica Instituti Botanici Nomine V.L. Komarov Academicae Scientiarum U.R.S.S., Petropolis 16:152-153. 1963. On fruits of Crataei;iis monogyna Jacq., Rosaceae. Rumania, (n. v.). Type material was not found at BUCM. No reply from BUG. BUCA, BUCF, CL. L. poierii G. Passerini, Atti dell' Accademia Nazionale dei Lincei. Memoire, Rome, Series 4,6:459.(1889) 1890. On wilted stems of Poleriiim san\>iiisorha L., Rosaceae. Italy, (n. v.). Type material was not found at PARMA, PISA, PAD. L. praeiermissa (P.A. Karsten) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:26. 1883. ^phaeria praeiermissa P.A. Karsten, Fungi Fenniae Exsiccati, Century 9, No. 852. Anno 1869. On dried canes of Riihiis idaeiis L., Kiihiis odoraltis L., Rosaceae. Finland. L. priini N.N. Woronichin, Vestnik Tiflisskogo Botanicheskogo Sada, Tlflis (Moniteur du Jardin Botanique de Tinis) 28:17-18. 1913. On living leaves of Primus domeslica L., Rosaceae. U.S.S.R. (Caucasus), (n. v.). Type material was not found at LE, LEP, TBI. No reply from TGM. L. piilchra (H.G. Winter) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:5.3-54. 1S83. ^phaerella piik lira H.G. Winter, Hedwigia, Dresden 11:145-146. 1872. =Mycodoleu piilchra (H.G. Winter) W. Kirschstein, Annalcs Mycologici, Berlin 34:201, 1936; Kryptogamentlora de Mark Brandenburg und Angrenzender Gebiete herausgegebcn von den Botanischen Verein der Provin/ Brandenburg, Leipzig 7(3):433, 1938. On dry stems and leaves of Polenlilla caulescens L., Rosaceae. Austria. Type material was not found at B. CORD, IBF, JE, LZ, WRSL, No reply from PI, IB, TO, WAG. =Leplosphaenilina piilchra (H.G. Winter) M.E. Barr, Contributions de I'lnstitut Botanique de L'Univcrsitc de Montreal 73:7. 1959. L. piillemansii A. Maublanc, Bulletin de la Societe Mycologique de France. Paris 21 :88-89. 1905. On \ea\es of Erioholrya japonica (Thunb.) Lindley, Rosaceae. Brazil, (n. v.). Type material was not found at IBI. PC, RSA- POM, SP. No reply from RBR. L. rimicola (G.H. Otth) P.A. Saccardo, Hedwigia, Dresden 35:XXIX. 1896: Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo ILXXIX. 1896. =Cladosphaeria rimicola G.H. Otth, Mitteilungen der Naturforschenden Gesellschaft in Bern 1870:106.1871. Nom inval. Art. 43.1. (The genus Cladosphaeria was validated in I894.| On fallen branches of Primus avium L., Rosaceae, Switzerland. =Melanomma pulvis-pyrius (C.H. Persoon:E.M. Fries) L. Fuckel. L. roslrupii P.A. Saccardo & D. Saccardo in P.A, Saccardo, Sylloge Fungorum Omnium Hucu.sque Cognitorum Digessit P,A. Saccardo 17:721. 1905. =Leplosphaeria dryadisE. Rostrup. Non Rostrup 1903. On upper surface of dried leaves of Dryas oclopelala L., Rosaceae. Norway, (n. v.). It is not known whether type material is at PAD since this institution has stopped lending specimens until further notice. L. nisiica P.A, Karsten, Fungi Fenniae Exsiccati, Century 10, No, 964. Anno 1870. =Metasphaeria ruslica (P.A. Karsten) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:157. 1883. On slems of Spiraea ulmariah. (=Filipendula ulmaria Maxim.), Rosaceae, Finland. =Leptosphaeria dolioliim (C.H. Persoon:E.M. Fries) V. Cesati & G. de Notaris. L. saccardiana J.H. Fabre. Annales dcs Sciences Naturellcs, Paris, Botanique, Series 6, 9:89. 1878. =Requienella saccardiana (J.H. Fabre) J.H. Fabre. Annales des Sciences Naturellcs, Paris. Botanique, Scries 6, 15:56. 1883. On dried stems of Crataegus oxyacantha L., Rosaceae. France, (n. v,). Type material was not found at PC. No reply from LHARMAS. L. sani^uisorhac P.A, Karsten in A,N. Berlese, Icones Fungorum Omnium Hucusque Cognitorum ad usum Sylloges Saccardianac 530 Illinois Natural Historj Suney Bulletin Vol. 34 An. 5 Adcommodaiae 1:78. 1892. On branches of Siiiigtdsorhii offii'inalis L., Rosaceae. Germany, (n. v.). Type material was not found at H. UPS. L. saimiiisorhac P.A. Karsten, Enumeratio Fungorum ct Myxomycetum in Lapponia Onentali .Aestaie 1861 Lectorum. p. 214. 1886; Noliser Sallskapets Pro Fauna et Flora Fordhandlmgar 8:214. 1866. [As Spluicria (Li'piospliaerial sangvisorhiic] On dead stems of Sangiiisorha pol\f;iimii F. Nyl.. Rosaceae. U.S.S.R. (n. v.). L. scpiiuolii (E.M. Fries:E.M. Fries) H.G. Winter, Dr. L. Rabenhorsl's Kr\ ptogaman-Flora von Deulschland. Oestcrreich und der Schweiz, Second edition. 1(2):473. 1885. jSphaeria sepinc(>la E.M. Fries. Obser\ationes Mycologieae. Seu Descripiiones tarn Novarum, quam Notabilium Fungorum Exhibitae a C.H. Persoon. Lipsiae 1:181. 1815; Systcma Mycologicum Sistens Fungorum 2:498. 1823. [As sacpincola.] sPringsheimia sepincola (E.M. Fries:E.M. Fries) F. v. Hoehnel. Annales Mycologici, Berlin 18:97. 1920. =Meiasphacrio sepincola (EM. Fries:E.M. Fries) PA. Saccardo. Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:164. 1883. (AsFr?Fuckel.l ^phaerulma sepincola (E.M. Fries:E.M. Fries) K.Siarback. Botaniska Notiser. Lund 1890:1 17. 1890; Botanisches Zentralblatt. Jena & Dresden 46:261. 1891. =Pleosphacnilina sepincola (EM. Fries:E.M. Fries) H. Rehm in F. v. Hoehnel. Annales Mycologici. Berlin 18:96. 1920. ^clerodolhis sepincola (E.M. Fries:E.M. Fries) F. Petrak. Annales Mycologici. Berlin 19:41. 1921. On dry stems of Rosa sp.. Riihus sp., Rosaceae. Sweden. =Saccoihecium sepincola (E.M. Fries:E.M. Fries) EM. Fries. Summa Vegeiabilium Scandinaviae. Seu Enumeratio. Systematica et Critica. Plantarum tum Cotyledonearum. Quam Nemearum Inter Mare Occidentale Et Album. Inter Eidoram et Nordkop. Haclenus Lectorum. una Cum Singulae Distributione Geographica. p. 398. 1849. L. sieversiae (F.E. Clements) F. Petrak. Sydowia. Annales Mycologici, Horn, N.O. 6(l-4):6. 1952. =Massaria sieversiae F.E. Clements, Cryptogamae Formationum Coloradensium, Century III, No. 234. Anno 1906. Nom. nud. Art. 36.1. [Combination is not valid because the basionym is an invalidly published species.) On standing, dead stems of Sieversia turhinata (Rydb.) Greene, Rosaceae. U.S.A. =Diadema sieversiae (C.H. Peck) S.M. Huhndorf, comb. nov. L. sorhi A. Jaczewski, Annales Mycologici, Berlin 1:30. 1903. On kiwet^ of Sorhiis aiicuparia L., Rosaceae. U.S.S.R. Species incertae .sedis. L. spiraeac P.A. Karsten, Hedwigia, Dresden 27:260. 1888. On dry fallen branches of Spiraea sorhifolia L. (=S(>rharia sorhifolia A. Br.). Rosaceae. Finland, (n. v.). Type material was not found al H, UPS. L. siihciihiiwa (M.C. Cooke & J.B. Ellis) J.B. Ellis in N.L. Britton. Catalogue of Plants Found in New Jersey. Geological Survey of New Jersey, Final Report of the State Geologist. Trenton 2(1 ):525. 1889. ^phaeria fsiihlcctac) suhciitanea M.C. Cooke & J.B. Ellis, Grevillea, London 7:41. 1878. =Mctasphacria suhciitanea (M.C. Cooke & J.B. Ellis) P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:167, 1883. On decorticated limbs of Pyrus communis L., Rosaceae. United States. (New Jersey). =Lophiost(>ma suhciilanca (M.C. Cooke & J.B. Ellis) S.M. Huhndorf. comb. nov. L. superficialis F.A. Hazslinszky, Malematikai es Teniieszettudomanyi Kozlemenyek Vonatkozolag a Hazai Viszonyokra, Budapest 25(2):146. 1892. On unknown substrate of Polcntilla argcntea L., Rosaceae. Hungary, (n. v.). Type material was not found at B, BP, F, FI, L, NBG, UPS, W, WAG. No reply from IBF. TNP. L. lernala F. Hazslinszky. Matematikai es Termeszetludomanyi Kozlemenyek Vonatkozolag a Hazai Viszonyokra, Budapest 25(2):150. 1892. On branches of Pvn/.v :?!«/(« L. {=Malus domeslica Borkh.). Rosaceae. Hungary, (n. v.). Type material was not found at B. BP, F, FI, L, NBG, UPS, W, WAG. No reply from IBF, TNP. L. ihomasiana P.A. Saccardo & C. Roumeguere, Revue Mycologique. Toulouse 5:236. 1883. On dead runners of Ruhus sp., Rosaceae. France. It is not known whether type material is at PAD because this institution has slopped lending specimens until further notice. =Phaeosphaeria ihomasiana (P.A. Saccardo & C. Roumeguere) S.M. Huhndorf, comb, nov. L. umbrosa G. Niessl in G.L. Rabenhorst, Fungi Europaei Exsiccati. Klotschii Herbarii vivi Mycologici Continuatio, Edition 3 (Edita Nova), Series 2, Century 20, No. 1934. Anno 1875; Just's Botanisch Jahresberichte, Berlin 3:262. 1887. =Massaria umbrosa {Q.H\fi%\) H. Rehm //( P.A. Saccardo, Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 9:761. 1891. On unknown substrate of Aciaea spicata L., Astrantia major L., May 1992 Lepmsphaeiia Species on the Rosaceae 531 Gentiana asclepiadea L.. Solanum dulcamara L., Spiraea aruiuits L. {=Aniiuiis dioicus Fern.), Ranunculaceae. Umbelliferae, Gentianaceae, Solanaceae. Rosaceae. Austria. L. vagahiinda P.A. Saccardo. Nuovo Giomale Botanico Italiano e Bolletino della Societa Botanica Italiana, Firenze 7:318. 1875. ^phaeria fuscella P.A. Saccardo, Atti dell' Accademia Scientifica Veneto-Trentino- Istriana. Padova2(l):145. 1873. [As Sphaeria fuscella M.J. Berkeley & C.E. Broome. Non Leplnsphaeria fuscella (M.J. Berkeley & C.E. Broome) V. Cesati & G. de Notaris.] On branches of Abies excelsa (Lam.) Poiret, Acer campeslre L., Alnus glutinosa (L.) Gaertner. Ampelopsis heterophylla (Thunb.) Sieb. & Zucc, Clematis vilalha L.. Cormis sanguinea L., Corylus avellaiia L.. Dulcamara sp.. Hypericum calycinum L., Kerria japonica (L.) DC.. Quercus pedunculata Ehrh.. Salix purpurea L., Pinaceae. Aceraceae. Betulaceae, Vitaceae. Ranunculaceae. Comaceae, Solanaceae, Guttiferae. Rosaceae, Fagaceae, Salicaceae. Italy, (n. v.). It is not known whether type material is at PAD because this institution has stopped lending specimens until further notice. L. vagahunda P.A. Saccardo var. divergens P.A. Saccardo, Fungi Italici autographice delineati (additis nonnullis extra-italicis asterisco notatis), Patavii, No. 518. 1879: Sylloge Fungorum Omnium Hucusque Cognitorum Digessit P.A. Saccardo 2:31. 1883. On branches of /ferna sp., Rosaceae. Italy, (n. v.). It is not known whether type material is at PAD because this institution has stopped lending specimens until further notice. L. vagahunda P.A. Saccardo var. sarmenti P.A. Saccardo, Fungi Italici autographice delineati (additis nonnullis extra-italicis asterisco notatis), Patavii. No. 428. 1879: Sylloge Fungorum Omnium Hucusque Congitorum Digessit P.A. Saccardo 2:31.1 883. On runners of /?«A>H.v sp., Rosaceae. Italy, (n. v.). It is not known whether type material is at PAD because this institution has stopped lending specimens until further notice. Literature Cited Arx, J. a. vf)N, and E. Muller. 1975. A re-evaluation of the bitunicate Ascomycetes with keys to families and genera. Studies in Mycology 9:1-159. Barr, M.E. 1959. Northern Pyrenomycetes I. Canadian eastern arctic. Contributions de I'institut botanique de I'universite de Montreal 73:1-101. Barr, M.E. 1967. Northern Pyrenomycetes III. Western species. Canadian Journal of Botany 45:1041-1046. Barr, M.E. 1972. Preliminary studies on the Dothideales in temperate North America. Contribu- tions from the University of Michigan herbarium 9:523-638. Barr, M.E. 1982. Leptosphaeria sepalorum. Mycotaxon 15:345-348. Barr. M.E. 1983. The Ascomycete connection. Mycologia 75:1-13. Barr, M.E. 1987a. Prodromus to class Loculoascomycetes. Hamilton I. Newell, Amherst. Massachusetts. 168 pp. Barr. M.E. 1987b. New taxa and combinations in the Loculoascomycetes. Mycotaxon 29:501-505. Barr, M.E. 1989. Clypeosphaeria and the Clypeo- sphaeriaceae. Systema A.scomycetum 8(1 ):l-8. Berkeley, J.M., and C.E. Broome. 1852. Notices of British fungi. Annals and Magazine of Natural History 119:317-329. Berlese. A.N. 1890. Icones Fungoruni. vol. 1, pp. 44-45. Abcllini. Brockman, I. 1975. Untersuchungcn iiberdie Gattung Discoslronui Clements (Ascomycetes). Sydowia 28:275-338. Cesati, v.. and G, de Notaris. 1863. Schema di Classificazione degli Sferiacei italici aschigeri. Commcntario della Socicla Crittoganiologia italiana. Milan 1:177-240. Chesters, C.G.C. 1938. Studies on British Pyrenomycetes II. A comparative study of Mekinomma puhis-pyrius (Pers.) Fuckel. Melanomma fuscidulum Sacc. and Thyiidaria nthronotala (B. & Br.) Sacc. Transactions of the British Mycological Society 22:1 16-150. Cooke, M.C. 1871. Handbook of British fungi. Macmillan, London and New York. 981 pp. Crane, J.L., and C.A. Shearer. 1991. A nomenclator of Leptosphaeria V. Cesati and G. de Notaris (Mycota-Ascomycotina-Loculoascomyceies). Illinois Natural History Survey Bulletin 34(3):195- 355. Dennis, R.W.G. 1978. British Ascomycetes. J. Cramer, Vaduz. 585 pp. Eriksson, O. 1 98 1 . The families of bitunicate ascomycetes. Opera Botanica 60:1-209. Eriksson, O., and D.L. Hawksworth. 1985. Outline of the Ascomycetes— 1985. Systema Ascomvcetum 4:1-79. Eriksson, O., and D.L. Hawksworth. 1986. Outline of the Ascomycetes— 1986. Svstema Ascomycetum 5(2): 1 85-324. Eriksson, O.. and D.L. Hawksworth. 1990. Outline of the Ascomycetes— 1 989. Systema Ascomycetum 8(2):l 19-318. Froidevaux. L. 1973( 1972). Contribution a I'etude des Dothioracees (Ascomycetes). No\a Hedwigia 23:679-734. FirKEL. L. 1870. Symbolae Mycologicae. Jahr- bucher des Nassauischen Vereins fur Naturkunde 23-24:1-159. Hedjaroude, G.A. 1968 (1969). Etudes taxonomiques sur Ics PluwosplMcrio Miyake et leurs formes voisines (Ascomycetes). Sydowia 22:57-107. HoiM, L. 1957. Etudes taxonomiques sur les Pleosporacees. S\ nibolac Boianicae Upsalienses 14(3):1-188. May 1992 Leplosphaciia Species on the Rosaceae 533 Holm, L. 1975. Taxonomic notes oti Pyrenomycetes. Taxon 24(4):475-488. Saccardo, P.A. 1876. Fungi Veneti novi vel critici. Series V. Niiovo Giomalc Botanico Italiano 8: 1 76. Holm. L. 1979. Microfungi on Dn'cis. Botaniska Notiser 132:77-92. Holm, L., and K. Holm. 1988. Studies in the Lophio- stomataceae with emphasis on the Swedish species. Symbolae Botanicae Upsalienses 28(2): 1-50. Holmgren. P.K.. W. Keuken, and E.K. Schofield. 1981. Index herbariorum. Part I. The herbaria of the world. 7th ed. Oosthoek. Scheltema & Holkema. Utrecht. HuHNDORF. S.M. 1991. A method tor sectioning ascomycete herbarium specimens for light micro- scopy. Mycologia 83:520-524. Karsten. P.A. 1873. Pyrenomycetes. Mycologia Fennica 2:105. Korf. R.P. 1958. Japanese discomycete notes I-Viii. Science Reports of the Yokohama National University, sect. 2. 7:7-35. Leuchtmann, a. 1984. \jber Pluieospluieria Miyake und andere bitunicate Ascomyceten mil mehrfach querseptierten Ascosporen. Sydowia 37:75-194. LuTTRELL, E.S. 1973. Loculoascomycetes. Pages 135-219 in G.C. Ainsworth, F.K. Sparrow, and A.S. Sussman (eds.) The fungi, an advanced treatise. Vol. IVA. A taxonomic review with keys: Ascomycetes and Fungi Imperfecti. Academic Press, New York and London. MuLLER, E. 1950. Die schweizerischen Arten der Gattung Leplosphaeria und ihrer Verwandten. Sydowia 4: 18.5-3 19. MuNK, A. 1957. Danish Pyrenomycetes. Dansk Botanisk Arkiv 17{l):l^91. NiESSL. G.V. ( 1 87 1 ) 1 872. Beitrage zur Kenntniss der Pilze. Verhandlungen des naturforschenden Vereines in Briinn 10:153-215. Peck, C.H. 1913. Report of the State Botanist 1912. New York State Museum Bulletin 167:1-137. Saccardo. P.A. 1878. Fungi Veneti novi vel critici. Series III. (ex Hedwigia. XIV (1875)) Michelia 1 :446-452. Shearer, C.A., J.L. Crane, and K.R. Chandra Reddy. 1990. Studies in Leplosphaeria. Lectotypification of Spluieria dotiolum. Mycologia 82:496-500. Shoemaker, R.A. 1976. Canadian and some extralimital Ophioholus species. Canadian Journal of Botany 54:2365-2404. Shoemaker, R.A. 1984a. Canadian and some extralimital Leplosphaeria species. Canadian Journal of Botany 62:2688-2729. Shoemaker. R.A. 1984b. Canadian and some extralimital Nodidosphaeria and Enlodesmhim species. Canadian Journal of Botany 62:2730-2753. Shoemaker, R.A., and C.E. Babcock. 1985. Canadian and some extralimital Paraphaeo- sphaeria species. Canadian Journal of Botany 63: 1284-1291. Shoemaker, R.A.. and C.E. Babcock. 1987. Weti- sleinina. Canadian Journal of Botany 65:373-405. Shoemaker. R.A.. and C.E. Babcock. 1989a. Diadema. Canadian Journal of Botany 67: 1 349- 1355. Shoemaker, R.A., and C.E. Babcock. 1989b. Phaeosphaeria. Canadian Journal of Botany 67:1500-1599. Shoemaker, R.A.. and O. Eriksson. 1967. Paraphaeosphaeria micholii. Canadian Journal of Botany 45:1605-1608. SivANESAN, A. 1984. The bitunicate Ascomycetes and their anamorphs. J. Cramer, Vaduz. 701 pp. Wehmeyer, L.E. 1933. The genus Diaporihe Nitschke and its segregates. University of Michigan studies. Scientific series 9:1-349. Petrak, F., and H. Sydow. 1924. Kritisch- systematische Originaluntersuchungen ijber Pyrenomyzeten, Sphaeropsideen und Melanconieen II. Annales Mycologici 22:359. Pflster, D.H. 1985. A bibliographic account of exsiccatae containing fungi. Mycotaxon 23:1-139. Saccardo, P.A. 1875. Fungi Veneti novi vel critici. Series II. Nuovo Giomalc Botanico Italiano 7:310- 311. Wehmeyer, L.E. 1957. The genera Saccolhecium. Prinfisheiniia. Pleosphaendina and Pseudoplea. Mycologia 49:8.V94. Wehmeyer, L.E. 1963. Himalayan Ascomycetes. Mycologia 55:319. Zeller, S.M. 1927. Contributions to our knowledge of Oregon fungi-II, Mycological notes for 1925. Mycologia 19:130-143. Index to Taxa ahhreviala 5 1 0, 326 aculeorum 526 arunci 498, 499, 326 cercocarpi 490.491.326 clivensis 498. 499 Ch'pvosphacria 3 1 8 concenlrica 492. 493, 526 coniotliyhum 500, 501, 526 conlroversa 520, 521, 326 iorucola 320.522,526 Diadema 513,514,515,516 Diaporthe 520 Discostroma 520 doUolum 483, 484 drwdis 484, 485. 527 dnadophila 484. 485. 527 e;-e,5 520, 521 erioholryae 525, 527 fiiscella 520,522,527 i'aliontm var. Uipsunae 498 hazslinszkii 527 hendcisonia 500, 501, 527 hollosiana 527 Kalmusia 498. 499. 500. 501 IcioslcKii 520. 522. 527 lejoslena 521.527 Lcplosphaerulina 510,512 k'ucnpUica 527 liiiigipedicellata 498, 499 Lophiosloma 503 ///(///<; 494, 495, 528 nhiniillana 518,519 niandshurica 528 McUinomma 505. 506 minima 525. 528 miyakeana 528 miimc 528 «/jm.?o 513,514 oliaoiheca 511,512,528 osvulanda 528 oxyspora 528 paihylheca 529 Paniphaeosphaeria 492, 493 Phacosphacria 492. 494. 495 "?;,•/!<; 492.494.529 pomona forma transihanica 529 polerii 529 praetermissa 486. 487, 529 prM«i 529 /j«/c/ira 510.512.529 pulvis-pyrius 505. 506 puttemansii 529 rimicola 505, 506, 529 rostrupii 529 rtt^r/ffl 483, 484, 329 saccardiana 329 Saccnthecium 508, 509, 510, 511 sanguisorbae 529, 530 sepincola 508, 509. 530 sepincola var. ahhreviala 510.511 i;eve«(ae 313.516.330 5orW 525. 530 spiraeae 530 sleironemalis 498 subciitanea 503. 504. 530 supeificialis 530 ternala 530 tlwmasiana 496. 497. 330 umhrosa 488. 489. 530 vagahiinda 53 1 vagahiinda var. divergens 53 1 vagahunda var. sarmenli 53 1 Illinois Natural History Survey Natural Resources Building 607 East Peabody Drive Champaign, Illinois 61820 217-333-6880 A Division of the Illinois Department of Energy and Natural Resources