Mjjcolog~a,85(2), 1993, pp. 294-3 10.

C 1993, by The New York Botanical Garden, Bronx, NY 10458-5126

A TAXONOMIC REVISION OF THE GENUS ENDOXYLA

Department of'Botany, I1niversity of Toronto, Toronto. Ontarlo M5S 3 8 2 Canada

ABSTRACT

The pyrenomycete genus Endoxyla (Ascomycotina, Clypeosphaeriaceae) is revised based on the examination of type and authentic material. Five species, E. macrostoma, E. munkii sp. nov., E. operculata, E. parallela and E. xanthostronla sp. no\$.,are included and a key to the species is given. Endoxjlla macrostoma is designated as holotype. Sixteen species and one variety are excluded from Endoxyla and three species are treated as doubtful. The synonymy of Ceratostomella with Endoxvla is rejected and a new combination Ceratostornella hyalostorna is proposed. The taxonomic position of Endoxyla is discussed. Key Words: Ascomycete, Ceratostomella, Clypeosphaeriaceae, Endo,x.vla, pyrenomycete, Scotiosphaeria, taxonomy

The genus Endoxyla was established by Fuckel (1871) for stromatic species possessing immersed ascomata with stout, apically thickened necks, clavate asci, dark, aseptate ascospores and abundant, filiform paraphyses. H e referred three species to the genus; Endo,~jllarnacrostorna Fuckel, Sphaeria parallela Fr. and S. operculatu Alb. & Schw. Saccardo (1882) formally combined the latter two species in Erzdo~xyla and described the ascospores of E . parallela as aseptate. He included the genus in the section Allantosporae of the Sphaeriaceae, an artificial assemblage of pyrenomycetes characterized by their allantoid ascospores. Winter (1886) treated Etzdo,~.~~lu a as subgenus of L'alsa Fr.. which he placed in the Valseae. Ellis and Everhart (1892, 1894) maintained Endo.xj>/aas a separate genus in this family and accepted E . tnucrostoma with aseptate ascospores, E. parallela with uniseptate ascospores, and four species with phragmosporous ascospores including E . fiaxitzi (Ell. & Ev.) Ell. & Ev. Berlese (1902, 1905) included only species with aseptate ascospores in Endo.s,.la and transferred E . parallela to Endo-~ylinaRomell. He considered Endos~tlato be related to Cryptosphaeria Grev. Von Hohnel (1915) removed the genus from the Valseae and treated E n d o q l a as a subgenus of Anthostoma Nitschke. H e restricted Anthostorna subg. Endosj~lato species with aseptate ascospores and referred E . parallcla to E u t ~ p o p sis P. Karsten. Von Hohnel (1918) later established the Allantosphaeriaceae (Allantosphaeriaceen) to include allantosporous pyrenomycetes

but he excluded ./lnthostorna subg. Endoxyla and taxa with septate ascospores from this family. Von Hohnel(1923) subsequently revised the circumscription of the Allantosphaeriaceae to include Endoxyla and genera such as Eutypopsis with septate ascospores. Wehmeyer (1926a) included Endoq9la in the Allantosphaeriaceae (Diatrypeen sensu Hohnel, 1918) and described the genus as possessing an effuse entostroma and septate ascospores. Wehmeyer later (1975) included Endoxyla in Cryptosphacria (Diatrypaceae). Dennis (1981) also included Endo.qtla in the Diatrypaceae but noted that its systematic position was unclear. Von Arx and Miiller (1954) placed E n d o - ~ j . 1 ~ the Diain porthaceae citing C'eratostornella Sacc. as a synonym. They delimited the genus to include diaporthaceous species with immersed perithecia. elongate necks, and aseptate or pseudoseptate, hyaline o r smoky-brown ascospores. They considered E. operculata and E. [rurullela to be conspecific and, following the lectotypification of Clements and Shear (1931). adopted the former as type species. In 1965 Munk emended Endo..ryla sensu von Arx & Miiller and expanded the genus to include six species characterized by their immersed perithecia, prominent. elongate necks. loosening asci. and unicellular, hyaline or brown ascospores. Munk (1957) had previously included E n d o q l a with Cerutosphaeria Niessl. Deharyella Hohnel. Lerztomita Niessl, Rhatnphoria Niessl and Zignoclla Sacc. in the Rhamphorioideae of the Diaporthaceae but he was later uncertain of the systematic position of the genus

(Munk, 1965). Barr (1978) excluded Endoxyla sensu von Arx & Muller from the Diaporthales and referred the genus, along with Ceratosphaeria and Lentomita, to the Lasiosphaeriaceae. She also noted the similarity of E. parallela to species of Cryptosphaeria. Recently, Barr (1990) placed Endoxvla in the Clypeosphaeriaceae and described the ascospores of its species as hyaline to light brown, septate or aseptate and as possessing a germ pore when brown. Twenty-two species and one variety have been assigned to Endoxyla. The genus presently contains an assemblage of unrelated fungi and it is not clearly delimited from other lignicolous pyrenomycetes. Recent treatments demonstrate that different concepts exist in the modern taxonomic literature for Endox~vla.Confusion concerning the taxonomy and systematic position of the genus is due primarily to the proposed synonymy of Endox.vla and Ceratostomella. This problem has been compounded by the questionable typification of the genus. Ceratostomella has been treated as a synonym of Endoxyla (von Arx and Miiller, 1954) but these taxa are not congeneric. Species of Ceratostomella possess immersed ascomata with elongate necks and pseudoparenchymatous peridia, small, loosening asci with a chitinoid, inamyloid apical ring and hyaline, aseptate or uniseptate ascospores. Ceratostornella has been included in the Diaporthaceae (Gilman et al., 1959; Muller and von Arx, 1973; Munk, 1953, 1957; von Arx and Miiller, 1954). Barr (1978) excluded the genus from the Diaporthales because of the carbonaceous peridia of the ascomata and occurrence on wood as secondary saprobes. Endo.xj~la differs from Ceratostomella in the brown. apically porate ascospores, the clavate, long-stipitate, non-loosening asci and the ascomata with a prosenchymatous peridium and stout, cylindrical necks which are confluent with or project only slightly above the surface of the wood. The affiliation of Endoxyla with members of the Diatrypaceae, which dates from the inclusion of the genus in the Allantosporae, is based primarily on ascospore shape and was strengthened by the inclusion of E. populi Rome11 and E. fraxini in the genus. The former species is comparable to Cryptosphaeria and the latter is synonymous with Cr~:ptosphaeriaeunomia (Fr. : Fr.) Fuckel. Endoxyla is not a member of the Diatrypaceae. The porate ascospores, asci with an

inamyloid apical ring, poorly developed stroma and occurrence on the wood of gymnosperms are features uncharacteristic of this family. Endoxyla shows little affinity to Endoxylina, a member of the Diatrypaceae treated by Rappaz (1987) as a synonym of Eutypa Tul. Porate ascospores and asci with an inamyloid apical ring are definitive characters of Endox.vla and are also found in the Boliniaceae, Sordariaceae and Clypeosphaeriaceae. The Boliniaceae are characterized by cylindric asci with an indistinct apical ring and small, laterally compressed, apically porate, brown ascospores. Species of Camarops P. Karsten, the largest genus in the family. possess tubular, monostichous or polystichous ascomata with membranous or elastic walls in a well developed, superficial, applanate to turbinate stroma (Nannfeldt, 1972). Camarops lutea (Alb. & Schw. : Fr.) Nannf. is characterized by a yellow stroma (Nannfeldt, 1972) and in this respect is similar to Endosj1la santhostrorna. Apiocamarops Samuels & Rogers, another member of this family, resembles species ofEndoxylain its terete ascospores which possess a small, terminal hyaline cell and cylindric-clavate asci (Samuels and Rogers 1987; Rogers and Samuels, 1988). Despite these similarities, Endo.xjda is not a member ofthe Boliniaceae; its species are distinguished from this family in the larger, terete ascospores, the morphology and arrangement of the ascomata and the absence of a superficial stroma. The Sordariaceae sensu lato encompass pyrenomycetes defined primarily by porate ascospores (Malloch and Cain, 1971) and asci possessing an inamyloid apical ring (Lundqvist, 1972). Genera with unequally bicellular ascospores consisting of a pigmented, porate cell and a hyaline, aporate cell are common in this family. The apical ring stains in congo red and often in cotton blue, and the asci are costate when empty (Lundqvist, 1972). The apical ring of species of Endoxyla does not react in these stains but costae are visible in discharged asci in cotton blue. The importance of cytoplasmic costae has been stressed at the family level (Carroll and Munk, 1964; Lundqvist. 1972) but other workers consider this feature to be nebulous (Cain and Krug, 1974). In surface view, the peridia of members of the Sordariaceae are pseudoparenchymatous (Jensen, 1985; Lundqvist, 1972). In some members of this family, however, the outer layer of the peridium appears areolate (Lundqvist, 1972)

and closely resembles the peridium of species of Endo-uyla. A further similarity between species of Endo.~ylaand members of the Sordariaceae is the occurrence of Munk pores in the walls of peridial cells. This feature was considered to be suggestive of the affinity of Lasiosphaeria Ces. & de Not. to the Nitschkiaceae (Carroll and Munk. 1964), but Munk pores have also been observed in species of the Diaporthaceae and Xylariaceae (Jensen, 1985). Despite its similarity to species included in the Sordariaceae, EndosjYu is not considered to be a member of this family. The absence of a fixed orientation of the ascospores within the asci. the staining properties of the apical ring and the diffuse entostroma are features of species of Endosyla which are uncharacteristic of this family. The Clypeosphaeriaceae was used by Barr (1989. 1990) for Cljpeospharria Fuckel and other genera related to the Amphisphaeriaceae. It includes pyrenomycetes with cylindric or clavate asci with an apical ring that is usually large and inamyloid and ascospores that vary in pigmentation, septation and in the presence of a germ pore and hyaline appendages. The stromatic tissues range from a thin. hyphal layer or a clypeus that overlies the ascomata, to a compact, prosenchymatous stroma in which the ascomata are immersed. The Clypeosphaeriaceae appears to be a heterogeneous family. Species of Endosyla differ from many of its members in features of the asci. ascospores and stroma. Within the family the closest affinities of the genus lie with Apiorhj~nchostomac1trrej3i (Rabenh.) Muller (Muller and von Arx. 1962). Dennis (1981) also compared E. operculata to .4piorhync~hostotna Petrak currej,i. .-fpiorh~~nchostor)~a and species of Endo.uj)la share a number of features, including their g y m n o s p e r m o u s substrates. ascomatal morphology and arrangement of the ascomata within a prosenchymatous stroma, asci with an apical ring not stained in Melzer's solution. congo red. and cotton blue. and porate. septate ascospores with a hyaline cell that lack a fixed orientation within the asci. Although the slstematic position of Endo.uj,la is problematic. Barr's (1990) placement of the genus in the Clypeosphaeriaceae is accepted.
MATERIALS AND METHODS

features of specimens were examined, measured and drawn in water mounts. Semipermanent mounts were made in lactophenol-cotton blue (Hawksworth et al.. 1983), a mounting medium useful for observing the germ pore of the ascospores. Freehand sections of the ascomata and substrate were examined to determine the extent of stromatal development. In order to obtain thin sections, the ascomata were fixed in 4% glutaraldehyde. dehydrated in a graded acetone series, and embedded in Spurr's embedding medium (O'Brien and McCulley. 1981). Sections were cut at a thickness of 3-5 Fm. Observations of peridium surface morphology were made from permanent mounts in glycerine jelly (Hawksworth et al.. 1983). The terminology used to describe peridial tissue morphology is taken from Korf (1973).The wood of specimens lacking substratum data was examined in toluidine blue 0 (O'Brien and McCulley, 1981) for the presence of vessels. Attempts to culture species of Endo,q.la from fresh and air-dried collections were made using modified Lconian's agar. Czapek's solution agar, V-8 agar (Malloch, 1981). Shear's corn meal agar (Rawlins. 1933). 1.5%water agar, Kauffman's synthetic agar, and woodchip infusion agar (Lohman. 1930). All plates were incubated at room temperaturc. Ascospore germination was not observed and cultures were not obtained from peridial cells or the centrum.
TAXONOMY

Endo.uyla Fuckel, Jb. Nassau. Ver. Naturk. 2526: 321. 1871.

= lTalsaFr. subg. Endo.xy/a (Fuckel) Winter, Rabenh. Krypt.-FI. l(2): 688. 1886. = .-Inthostoma Nits. subg. Endosyla (Fuckel) Hohnel, Sitzungsber. Kaiser]. Akad. Wiss., Math.-Natunviss. CI.. Abt. I, 124: 66. 19 15.'

Stromata pustulate or poorly developed, composed of yellow o r brown hyphae penetrating the substrate around the ascomata, often evident onll as a discoloration of the substrate. Ascomata perithecioid, immersed, monostichous, solitarto gregarious, black. Necks stout. cylindrical. separately erumpent. apices confluent with or projecting slightly above the surface of the wood. Peridium coriaceous, in surface view a tc.uturu irltricatu with cells arranged in radiate patches. Asci unitunicate, eight-spored, clavate-cylindrical. long-stipitate, with an indistinct, inamyloid apical ring. Ascospores elliptic-cylindrical or suballantoid, brown. with an apical germ pore. septate o r aseptate, with or without a hyaline cell. uniseriate o r biseriate becoming uniseriate below. with variable orientation within the asci.
I Von Hohnel(1915) treated Endoxyla as a subgenus of .-Inthostomabut did not formally transfer any of its species to Anthostoma.

This study was based primanly on the examination of herbarium specimens. The acronyms of herbaria are those used by Holmgren et al. (1990). The microscopic

Paraphyses filiform or filiform-ventricose. septate, sparse or absent in mature perithecia. On decorticated wood. Anamorphs not known. TYPE SPECIES: Endoxvla rnacrostotna Fuckel, Jb. Nassau. Ver. Naturk. 25-26: 322. 1871. As suggested by Holm (1975) the lectotypification of Endoxyla with E. operculata (elements and Shear, 1931) is debatable. When Fuckel (1871) established E n d o x ~ ~ lhe referred three a species to the genus. He described E. rnacrostoma at length and mentioned that Sphaeria parallela and S. operculata belonged to the genus. The combinations for these species in Etzdo-~yla were later published by Saccardo (1882). Endoxy/a parallela and E. operculata should not be considered for the typification of Endo,xj~Iabecause these names were not definitely included in the protologue of the genus. Endosyla macrostoma, the single element included in the genus by Fuckel, is therefore regarded as holotype.

1. 1.

3. 3.

or projecting slightly above the surface of the wood. Wall of venter brown, 15-25 pm thick; external layers composed of interwoven, tubular hyphae and globose cells 1.7-3.1 pm diam; internal layers composed of thinner-walled, narrowly rectangular cells. Wall of neck brown, external layers similar in composition to those of the venter; internal layers composed of compressed, rectangular-tubular cells. Wall of neck apex brown, 80-97 pm thick, composed of elongate, tubular cells 2.7-4.1 pm diam that grade inwardly into compressed, rectangular cells. Outer peridial layer of venter a te-xtura intricata in surface view, composed of dark brown, aseptate hyphae 2.9-6.6 pm diam with cell walls <0.5 pm thick, hyphae arranged in radiate patches: at lower magnifications appearing composed of highly interwoven hyphae. Munk pores not observed. Outer peridial layer of neck a textura porrecta to a t. intricuta in surface view, composed of cells 3.1-4.7 pm diam with cell walls 1 0 . 5 pm thick. Asci eight-spored, clavate-cylindrical, long-stipitate, total length 69-1 37 pm, pars Ascospores aseptate . . . . . . . . . . . . . .E. operculata
sporrfera 39-60 x 5.5-7.2 pm, with an indistinct, Ascospores septate. septum delimiting a hyaline inamyloid apical ring. Ascospores suballantoid, cell or dividing the pigmented portion of the x 2.2-3 4(-3.9) ascospore, or both . . . . . . . . . . . . . . . . . . . . . . . 2 curved, (8.9-)9.7-12.3(-13.3) 2. Ascospores uniseptate, without a hyaline pm, bicellular, composed of a hyaline cell 0.8cell . . . . . . . . . . . . . . . . . . . . . . . . . .E. parallela 1.7 x 1.5-2. I pm, and a brown cell with an apical 2. Ascospores uniseptate or biseptate, with a hyaline cell . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 germ pore. porate end acute, biseriate or apically biseriate, becoming uniseriate below. Apical and Pigmented portion of the ascospores with a septum, ascospores(l0.2-)I 1.3-14.8(-17.8) x 2.6basal ascospores with germ pores often oriented 3.6(-4) pm . . . . . . . . . . . . . . . . . . . . . . . . E. munkii toward one another, the remaining ascospores . Pigmented portion of the ascospores aseptate. ascospores smaller . . . . . . . . . . . . . . . . . . . . . . . . 4 randomly oriented. Paraphyses filiform. infrequently septate, 1.5-2 pm diam, absent in mature 4. Ascospores suballantoid, porate end acute, (8.9-)9.7-12.3(-13.3) x 2.2-3.4(-3.9) prn ascomata. Anamorph not known.
. . . . . . . . . . . . . . . . . . . . . . . . . .E. rnacrostorna

4. Ascospores ellipsoid-cylindrical, porate end broadly rounded, 8.3-10.9 x 2.3-3.5 um . . . . . . . . . . . . . . . . . . . .E. .uanthostroma

Endo,~ylarnacrostoma Fuckel, Jb. Nassau. Ver.

Naturk. 25-26: 322. 187 1.

FIGS. 1-7

Valsa rnacrostorna (Fuckel) Winter, Rabenh. Krypt.-Fl. l(2): 689. 1886.

HABITAT. decorticated wood. On HOLOTYPE. GERMANY. Fungi rhenani 232 1, Nassau. Winkler Woods at Bachweg, on wood of Quercus sp., L. F~cckel(G). SPECIMENS EXAMINED. GERMANY. ADDITIONAL Fungi rhenani 2321. L. Fuckel (SYNTYPES FH, G); L. Fuckel, as Sphaeria parallela Fr.? (G). SWEDEN. Scleromyceti Sueciz no. 268 Sphaeria operculata Fr. ( A. S.) (1st ed.), pro parte. E. 6 Fries (FH).

Stromata poorly developed, consisting of brown. septate hyphae 2.7-3.9 pm diam that darken the wood around the necks ofthe ascomata. Ascomata immersed, solitary to gregarious, often in short linear groups, ampulliform, 680990 x 495-550 pm, black, with stout, cylindrical, separately erumpent necks that widen towards their apices. 220-440 x 150-330 pm: apices of necks plane or planoconvex, confluent with

Endo.xj~la macrostotna is distinguished from other species of Endoxyla by its suballantoid. curved ascospores that possess a hyaline cell and an acute porate end. Berlese (1905) is the only worker to have previously described and illustrated the hyaline cell of the ascospores. His description of E. macrostoma, based on Fungi rhenani 232 1, differs little from the one given above. Berlese's illustration does not, however,

depict a stroma, suggesting that this feature may be variable in this species. Endoxj>/a wzacrostom a is known only from a single collection outside of Fuckel's herbarium and exsiccata.

Endoxyla munkii Untereiner, sp. nov. FIGS. -1 1 8

Stromata debiliter evoluta, superficiem ligni et lignum circa ascomata denigrantia, saepe entostroma compactum formantia turmas ascomatum circumdans: composita ex hyphis septatis brunneis 2.3-3 pm latis. Ascomata immersa, raro solitaria, vulgo seriebus 2-20 mm longis lineariter plus minusve regulariter aggregata, ampulliformia, 495-825 x 350-450 pm, nigra, collis crassis, cylindricis, discretim erumpentibus. apicem versus dilatatis, 120-210 x 120-250 pm; apices planoconvexi vel pulvinati, protrudentes. Paries ventris brunneus, 15-25 pm latus, stratum externum aspectu superficiali textura intricata. Asci octospori, clavati-cylindrici, longipedicellati, 1 19-1 68 pm longi, pars sporifera 40-79 x 6.4-9.7 pm, annulo indistinct0 inamyloideo. Ascos~orae suballantoideae, ( 10.24 11.314.8(-17.8) x 2.6-3.6(-4) pm. biseptatae, ex duabus cellulis brunneis aequalibus porogerminationis apical;. et cellula hyalina 0.8-1.7 x 1.7-2.3 pm compositae, biseriatae in parte superiore, infra uniseriatae. Ascosporae apicales et basales pons germinationis saepe aspicientes inter se. aliae irregulariter dispositae. Paraphyses filiformes vel modice ventricosae. septatae, 1.8-2.5 pm latae, sparsae. Habitat in ligno decorticato petrefacto Pini s,ylvestris. HOLOTYPUS. RUSSIA. Karelia, 7 July 1937, ,Wattl Laurila (H. e-x HPP 7 128).

Stromata poorly developed, evident as a darkening of the surface of the wood and of the wood around the ascomata, often as a compact entostroma that laterally encloses groups of ascomata; composed of brown, septate hyphae 2.3-3 pm diam. Ascomata immersed, rarely solitary, usually in regular or irregular linear groups 2-20 mm in length, ampulliform, 495-825 x 350-450 wm. black, with stout, cylindrical, separately erumpent necks that widen towards their apices, 1202 10 x 120-250 pm; apices of necks planoconvex or pulvinate, projecting above the surface of the wood. Wall of venter dark brown, 15-25 pm thick; external layers composed of polygonal cells 3.7-9.2 x 2.1-3.5 pm with cell walls >0.5 pm thick: internal layers composed of compressed, thinner-walled, rectangular cells 4.1-1 3.6 x 1.5-

2.5 pm. Wall of neck dark brown, with external layers composed of polygonal cells 5-7.5 x 2.94.1 pm that grade inwardly into compressed, rectangular cells. Wall of neck apex dark brown, 60-90 pm thick. composed of elongate, rectangular cells 7.1-14.5 x 0.9-2.6 pm with cell walls >0.5 pm thick that grade inwardly into compressed, thinner-walled, rectangular cells. Outer peridial layer of venter a textura intricata in surface view, composed of dark brown, septate, branched hyphae 2.6-3.5 pm diam with cell walls <0.5 pm thick, hyphae loosely arranged in indistinct radiate patches; internal peridial layer of Tenter a te-xrura angularis in surface view, composed ofcells 4.3-1 3.4 x 2-7.7 pm with cell walls <0.5 pm thick; at lower magnifications appearing pseudoparenchymatous. Munk pores abundant, 0.6-1.2 r m diam. Peridium of neck a te,utura porrecta to a t. pris?natica in surface view, composed of cells 2.9-4 pm diam with cell walls <O.5 pm thick. Asci eight-spored, clavate-cylindrical, long-stipitate, total length 1 19-1 68 pm. pars spor~fera 40-79 x 6.4-9.7 fim, with an indistinct, inamyloid apical ring. Ascospores suballantoid, (10.2-)11.3-14.8(-17.8) x 2.6-3.6(-4) pm, biseptate. composed of two brown cells of equal length with an apical germ pore, and a hyaline cell 0.8-1.7 x 1.7-2.3 pm, apically biseriatc bccoming uniseriate below. Apical and basal ascospores with germ pores often oriented toward one another, the remaining ascospores randomly oriented. Paraphyses filiform or filiform-ventricose, septate, 1.8-2.5 pm diam. sparse in mature ascomata. Anamorph not known. HABITAT. decorticated rotting wood of Plnus sp., On P. sylvestris L., and P. ponderosa Laws. HOLOTYPE. RUSSIA. Karelia (formerly Paanajarvi. Kuusamo. Finland), on dry wood of P. sylvestris, 7 July 1937, Matti Laurila (H, ex HPP 7128). ETYMOLOGY. Named in honour of the late, cminent pyrenomycete taxonomist Anders Munk. ADDITIONAL SPECIMENS EXAMINED. FINLAND. Fungi Fenniae no. 777, on Pinus sp., 31 Oct. 1868, P. Karsten (H); Vammala, T y r v s , Sept. 1859, P. Karsten (H 2896): Vammala, T y w E , 1859, P.Karsten (H 2895); Tammela, Mustiala. 2 1 Oct. 1867, P. Karszetz (H 2893. 2894). SWEDEN. Scleromyceti Suecia: no. 3 Sphaeria parallels Fr. (2nd ed.), pro parte. E. Af. Fries (UPS). USA. OREGON: Takilma, Siskiyou National Forest,

FIGS.1-7. Endo.xyla macrostoma. I. Asci, x 2400. 2. Ascospores. x 2400. 3. Abnormal ascospores, x 2400. 4. Longitudinal section through neck of ascoma, x 336. 5. Longitudinal section through venter of ascoma, x 336. 6. Habit sketch of ascomata, x 36. 7. Surface view of peridium of venter, x 1800.

on P. ponderosa, 1l Dec. 1925, coll. C. A. Brown no. 218, det. C. H. Kauffman, pro parte (MICH).
Endoxyla rnunkii closely resembles E . parallela. The dimensions of the ascospores, the linear arrangement of the ascomata, and the presence of a compact entostroma that overlies and often laterally encloses groups of ascomata are features shared by these two species. Also, E. rnunkii and E. parallela are the only species of the genus often associated with Scotiosphueria endo,u?,linae Sivanesan. Endoxyla mzlnkii is distinguished from E . parallela by the wall of the venter which is composed of polygonal, thick-walled cells, the pseudoparenchymatous appearance of its peridium at low magnification, and its biseptate ascospores that possess a hyaline cell. Endo.uyla operculata (Fr. : Fr.) Sacc., Syll. Fung. 1: 181. 1882. FIGS.12-1 5

= Sphaeria operculata Pers. 66 seriata Alb. & Schw., Conspect. Fung. Lusat., p. 40. 1805. Sphaeria operctllata Fr. : Fr., Syst. Mycol. 2(2): 479. 1823, non Persoon, Syn. Meth. Fung. p. 80. 1801. = Valsa operculata (Fr. : Fr.) Nitschke, Pyrenomyc. Germ. 1: 153. 1867. Anthostoma operculata (Fr. : Fr.) J. Schroeter, Krypt.-Fl. Schlesien 3(2): 432. 1897. = Ceratostoma operculata (Fr. : Fr.) Petrak, Fl. Boh. et Mor. Exs. Ser. ii, I Abt. Pilze, Lfg. xx, 968. 19 14 Vide Samuels, 1982). = Ceratostoma crassicoflis Kirschstein, Verh. Bot. Vereins Prov. Brandenburg 48: 52. 1906. = Endoxyla operculata (Fr. : Fr.) Sacc. var. compresslspora Kauffman, Pap. Michigan Acad. Sci. 11: 161. 1929.

Stromata poorly developed or absent, consisting oflight brown, septate hyphae that darken the wood around the ascomata. Ascomata immersed, solitary to gregarious, often in short linear groups, ampulliform, 540-950 x 350-680 pm, black. w ~ t hstout, cylindrical, separately erumpent necks that widen towards their apices, 140-490 x 1 10-3 10 pm; apices of necks plane or planoconvex, protruding slightly above the surface of the wood. Wall of venter dark brown. 14-28 pm thick; external layers composed of interwoven, tubular hyphae 1.4-4.7 pm diam: in-

ternal layers composed of polygonal cells 5.310.7 x 1.5-4.8 wm thatgrade inwardly into compressed, rectangular cells. Wall of neck dark brown, with external layers composed of globose-polygonal cells 2.5-3.9 pm diam that grade inwardly into compact rectangular-polygonal cells. Wall of neck apex dark brown, 50-85 pm thick, composed of rectangular cells 2.7-4.7 pm diam. Outer peridial layer of venter a te.rtura intricata in surface view, composed of dark brown, infrequently septate, branched hyphae 2.1-4.4 pm diam with cell walls <0.5 pm thick; hyphae arranged in radiate patches; internal peridial layer a textura angularls in surface view, composed of cells 5.9-14.4 x 3.1-8.6 pm with cell walls ~ 0 . pm thick; at lower magnifications 5 appearing areolate with regular, dark plates separated by bands of lighter cells. Munk pores numerous, 0.7-1 pm diam. Outer peridial layer of neck a textura porrecta to a t. intricata in surface view, composed of cells 2.4-3.9 pm diam with cell walls <0.5 pm thick. Asci eight-spored, clavate-cylindrical, long-stipitate, total length 109158 pm, pars sporfera 45.8-75.6 x 5.3-7.6 pm, with an indistinct, inamyloid apical ring. Ascospores ellipsoid-cylindrical, slightly curved, (8.0-)9.8-12.7(-14.6) x (2.1-)2.7-3.7(-4.9) pm, brown, with porate end somewhat acute, apically biseriate becoming uniseriate below, rarely unisenate. Apical and basal ascospores with germ pores often oriented toward one another, the remaining ascospores randomly oriented. Paraphyses filiform-ventricose. septate, l .8-3.6 pm diam, sparse in mature perithecia. Anamorph not known. HABITAT. decorticated wood of angiosperms and On gymnosperms. LECTOTYPE. SWEDEN. Scleromyceti Suecia: no. 268 Sphaeria operculata Fr. (6 A. S.) (1st ed.), E. .W. Fries (UPS). designated here. ADDITIONAL SPECIMENS EXAMINED. CANADA. ONTARIO: Haliburton Co.. Tory Hill, on Acer sp., 8 July 1953, R. F. Cain (TRTC 40040); Rondeau Prov. Park, Harris Trail, on rotten decorticated wood of Pinus sp., 20 Oct. 1987, W.A. Cntereiner (TRTC 5 1047). FINLAND. IOtkajoki, Juuma, Kuusamo, on rotten dry wood of P. sylvestris, 15 June 1938, Matti Laurila (H. ex HPP 7 129). FRANCE. Vosges, on wood of Pinus

FIGS.8-1 5. Endo.uyla munkii and Endoxyla operculata. 8-1 1 . Endoxvla munkii. 8. Habit sketch ofascomata, 9. Abnormal ascospores, x 2400. 10. Ascus, x 2400. 1I. Ascospores, x 2400. 12-1 5. Endoxyla operculata. 12. Habit sketch of ascomata, x 36. 13. Abnormal ascospores. x 2400. 14. Ascospores, x 2400. 15. Ascus, x 2400.
x 48.

sp., C. Montagne (two collections, G). GERMANY. portion on gymnosperm wood is Scleromyceti Fischerhaus, Grossbehnitz, on the wood of a rotten Sueciz no. 268 S. operculata ( I st ed.) (Holm and pine fence, 8 Oct. 1905. W Kirschstein (HOLOTYPE Nannfeldt, 1963; Shear, 1921). T h e identity of Ceratostorna crassrcollis. B). SWEDEN. Scleromyceti Suecia: no. 268 Sphaerra operculata Fr. (6 A. S.) (1st S. operclllata 66 seriata remains unclear. Fries ed.), E. .M. Fries (pH, UC): Scleromyceti Sueciz no. (1823) cited Scleromyceti Sueciz no. 268 as rep268 (2nd ed.), E. M. Fries (FH. ex herb. M. A. Curtis: resentative of S. operculata and listed pine a n d G. ex herb. A. P. de Candolle: K, ex herb. M. J . Berke- alder as its substrates. The example of this specley): e.u Scleromyceti Sueciz no. 268 (as no. 468) (edition unknown). von Hohnel slide collection. case VI- imen a t U P S is therefore designated as lectotype 3. A4080 (FH). USA. No locality. as E. tnacrostoma, (Art. 7.20). Dec. 1885, coll. I;. 0. Grover, det. R. Thaxter no. 379 Endo.q,la opercltlata var. cornpressispora, de(FH); North American Fungi no. 194 Sphueria par- scribed by Kauffman (1929) based o n specimens Fr., allel~ J. B. ENis (M; NY. ex herb. R. A. Harper); with highly compressed ascospores, is indistinFungi Columbiani no. 95 1 Endosyla parallela (Fr.), J. B. Ellis and E, hf. Everhart (NEB): e.u Fungi Columbi- guishable from E. operculufa. Ascomata from the ani no. 951. von Hohnel shde collection. case VI-3, holotype of var. cornpressispora are overmature; A408 1 (FH). NEWJERSEY: Newfield, on pine poles in asci are absent, a n d the ascospores are collapsed. grape trellis. 28 April 1878, J. B. Ellis (FH. e.u herb. T h e ascospores assume their normal shape when W. G. Farlow);Newfield. on Qitercuc sp., 22 Aug. 1888, J. 5.Ellis (NY); Newfield. on Kalr?7ra lat!folra L., 1 heated in lactophenol-cotton blue. Nov. 189 1 . J. B. Ellls (NY); Newfield, on rotten wood of Prnus sp., J. B. Ellis (NY. e.x herb. H. M. Richards): Endosj~laparallela (Fr. : Fr.) Sacc.. Syll. Fung. 1 : Newfield. on pine. J. B. Ellis (NY. e.u herb. J. B. Ellis). 181. 1882. FIGS. 16-19 OREGON: Takilma, Siskiyou Nat. Forest, on decorticated wood of Pinus sp.. 2 Dec. 1925, coll. C. .A. Rrow'n, E Sphaeria parallela Fr.. Kongl. Svenska Vetendet. C. H . Kauffman (HOLOTYPE and PARATYPE skapsakad. Handl., ser. 3, vol. 4: 155. 1816. E. operc~tlata var. cornpres~i.pora.MICH). = Sphaeria parallela Fr. : Fr., Syst. Mycol. 2(2): 373. 1823. = Falsa parallela (Fr. : Fr.) Nitschke. Pyrenomyc. T h e nomenclature of this species is quite inGerm. 1: 154. 1867. volved. Persoon described Sphaeriu opercltlata = Eufypa ? parallela (Fr. : Fr.) P. Karsten, Bd. and S. operculafa B aspera in 180 1. Albertini and Kann. Finl. Nat. Folk 26: 130. 1873. Schweinitz (1805) subsequentlq proposed three Euwpopsis parall~lu (Fr. : Fr.) P. Karsten. Meddeland. Soc. Fauna R. Fenn. 2: 182. 1878. Pers.: (7 sparsu o n ..llnzis varieties of S. opere~rluta -= Xjdosphaeria parallela (Fr. : Fr.) Cooke, Grevilsp., yy quirlcuncial~s n Fagus sp., and 66 seriafa o lea 7 : 86. 1879. o n ilhies. Fries (1823) recognized Sphaeria ell= Endosj~linaparall~la (Fr. : Fr.) Berlese, Icon. fypa var. aspera based o n S operculuta 3 uspera Fung. 3: 105. 1905. Per$. H e also clalmed that S operculata Pers. = Endo.nlina crocea Kirschst.. Ann. Mycol. 33: 2 14. 1935. included S. eutj'pa Fr. and the species for which = Endo.u.t~lina prni Sivanesan. Trans. Brit. Mycol. he adopted the name S. operculuta based o n AlSOC. 9: 117. 1977. 6 bertinl a n d Schweinitz's varieties tu sparsa and 66 seriata (Fries. 1823). Although Fries lists AlStromata poorly developed. evident as a darkbertini and Schweinitz as authorities for S. oper- ening of the surface of the wood and of the wood rularu In the lndlces of the SIsterlzu nly(o1ogzcurn around the ascomata, often as a discrete and (1823. p. 6 17. 1832. p. 170). thls specles should compact entostroma overlying and laterally enbe attributed t o Fries alone. closing groups of ascomata; composed of brown. Specimens of S. operculata and S. operculata septate hyphae 1.5-2.8 pm dlam. Ascomata Im$ asperu from Persoon's herbarium (L) represent mersed. rare11 solitary, often In regular or Irregmembers of the Diatrypaceae. There IS n o ma- ular l ~ n e a r groups 2-14 m m In length, ampulllterial corresponding to E . opercz~lafa. speci- form, 495-790 x 275-550 pm, black. with stout. A men of S. operculuta in Schweinitz's herbarium cylindrical, separately erumpent necks, 190-3 10 (PH) annotated "Niskcy A. S." includes Eut.~pa x 1 10-230 pm; apices of necks planoconvex or corn- pul\lnate. projecting above the surface of the lata (Pers. : Fr.) Tul. and Platj~stot)~urn pressurn (Pers. : Fr.) Trev, o n angiosperm wood wood. Wall of venter dark brown. 17-30 p m a n d E . operculara o n gymnosperm wood. Nei- thick: external layers composed of tubular hqther of the species o n angiosperm wood clearly phae 1.2-3.6 p m diam: Internal layers composed corresponds to the descriptions of u sparsa on of polygonal cells 4.4-8.2 x 1.7-3.3 pm with cell .-l/nus sp. o r yy q~rincuncialison Fhgtis sp. T h e walls > 0 . 5 pm thick that grade inwardly into

compressed, thinner-walled, narrowly rectangular cells. Wall of neck brown, with external layers composed of tubular hyphae and globose cells 1.2-3.6 pm diam that grade inwardly into interwoven tubular cells. Wall neck apex brown, 70-1 2 0 pm thick, composed of elongate, tubular cells 2.1-3.5 pm diam that grade inwardly into compressed, thinner-walled, rectangular cells. Outer peridial layer of venter a te-xtura intricata in surface view, composed of dark brown, septate, branched hyphae 2.1-4.6 fim diam with undulate cell walls < 0 . 5 pm thick; hyphae arranged in indistinct radiate patches; internal peridial layer a te.xtura angularis in surface view, composed of cells 4.9-1 5.1 x 3.2-8.7 pm; at lower magnifications appearing areolate with irregular, dark plates separated by bands of lighter cells. Munk pores 0.6-0.8 pm diam. Outer peridial layer of neck a te,utura porrecta to a f . intricata in surface view, composed of cells 1.7-3.4 pm diam with cell walls <0.5 pm thick. Asci eight-spored, clavate-cylindrical, long-stipitate, total length 105160 pm, pars spor~fera40.4-90 x 5.5-8.8 pm, with an indistinct, inamyloid apical ring. Ascospores suballantoid, (9.6-)I 1.O-14.6(-15.5) x 2.3-3.5(-4.2) pm, brown, with an apical germ pore, bicellular, septum central or not, apically biseriate becoming uniseriate below, without a fixed orientation within the asci. Paraphyses filiform, infrequently septate, 1.6-2.4 pm diam. absent in mature ascomata. Anamorph not known. HABITAT. decorticated rotting wood of Pinus sp., On P. ponderosa, P. sylvestris, and P. strobus L. LECTOTYPE. SWEDEN. Scleromyceti Suecia: no. parallela Fr. (1st ed.), E. M. Fries (UPS), 3 Sphaer~a designated here. ADDITIONAL SPECIMENS EXAMINED. CANADA. ONTARIO: Pinery Prov. Park, Pine Trail, on rotting decorticated wood of Pinus sp., 21 Oct. 1987, W. A. C'nterelner (TRTC 5 1040). NOVA Scorn: Halifax, April 1895, coll. (?) J. G. Hall (FH, ex herb. J. B. Ellis). FINLAND. Fungi Fenniae no. 777, on Pinus sp., 31 Oct. 1868, P. Karsfen (NY, ex herb. J. B. Ellis); Merimasku, on wood of Pinussp., 11 May 1865, P. Karsten (H 2892,2998). GERMANY. Wald am Gamensee bei Strausberg, 16 May 1926, coll. W. Krieger (HOLOTYPE Endoxylina crocea, MBB). SCOTLAND. Black Wood of Rannoch, Perthshire, on wood of Pinus sp., 17 Mar. 1976, B. J. Coppins no. 1579 (HOLOTYPE Endoxylina pini, IMI 202374a; ISOTYPE E V888); Black Wood of Rannoch, Perthshire, on wood of fallen Pinus sp.. 24 Sept. 1983, coll. B. J. Coppins (TOPOTYPE Endoxylinapini, TRTC, ex E); Mar Forest, 1882, R. K. Greville, under an unpublished name (E; K, ex herb. M. C. Cooke); Glen Affric, Pollon Buidhe. on

damp wood of Pinus sp., 20 June 1975, coll. et det. B. J. Coppins no. 1421 (E VC86). SWEDEN. Scleromyceti Suecia: no. 3 Sphaeria parallela Fr. (1st ed.), E. M. Fries (FH; PH); Scleromyceti Suecia: no. 3 (2nd ed.), E. M. Fries (E; FH, ex herb. M. A. Curtis, two collections; UPS two collections, one mixed); Sphaeria parallela Fr., ex herb. J. B. Ellis, ex herb. M. C. Cooke, received from E. M. Fries (NY). USA. North American Fungi no. 194. Sphaeria parallela Fr., J. B. Ellis (G; TRTC); Fungi Nova-Caesareenses no. 95 Sphaeria parallela Fr., 1876, J. B. Ellis (FH). NEW JERSEY: Newfield, on rotten pine, 20 July 1884, J. B. Ellis (FH, ex herb. W. G. Farlow); Newfield, J. B. Ellis (FH, ex herb. W. G. Farlow). M ~ s s ~ c ~ u s ~ r r s : Petersham Twp., Federation of Women's Clubs State Forest, 4 km W of Petersham, on rotting decorticated wood of P. strobus, 29 Aug. 1987, W. A. CJnfereiner(TRTC 5 104 1); Mt. Toby Forest, on rotting decorticated wood of P. strobus, 29 Aug. 1987, W.A. Untereiner(TRTC 5 1042, 5 1043): Mt. Toby Forest, on rotting decorticated wood of P. strobus. 27 Aug. 1987, W. A. Untereiner (TRTC Takilma, Siskiyou Nat. Forest, on P. 5 1044). OREGON: ponderosa, 1 1 Dec. 1925, coll. C. A. Brown, det. C . H. Kauffman (MICH, two collections).

There has been considerable confusion concerning ascospore septation in E . parallela. Uniseptate ascospores were first reported for Sphaerla parallela by Nitschke (1867) but this feature was not included by Fuckel (1871) in his diagnosis of Endoxyla. Karsten (1873) corroborated Nitschke's observation that the ascospores of this species were either aseptate or uniseptate. Cooke (1873) and Ellis and Everhart (1892) also described the ascospores as uniseptate. On the other hand, Currey (1858) and Saccardo (1882) reported the ascospores of S. parallela to be aseptate. Berlese (1905) described the collection of Scleromyceti Sueciie no. 3 that he studied as mixed; it included S . parallela with septate ascospores, which he transferred to Endoxylina, and Endoxyla operculata with aseptate ascospores. Von Hohnel (1915) described the ascospores of S. parallela as uniseptate but later suggested that they could remain aseptate or become uniseptate (von Hohnel, 1923). It is important to note that the descriptions of Currey (1858), Nitschke (1867), and Cooke (1873) are based either wholly or in part on Scleromyceti Suecia no. 3. It is possible that these collections, like the example examined by Berlese, included both E. operculata and E. parallela. In this study only one example of Scleromyceti Sueciie no. 3 (2nd ed., UPS) was a mixed collection, and it included E. parallela and E . m u n kii. All other examples included only E. paralIela. The existence of mixed collections initially

suggested that ascospore septation was a variable Scotiosphaeria is most closely allied to Poroscharacter. However, ascospores with a hyaline phaerella Miiller & Samuels, a genus described cell have not been observed in E. parallela and from decaying wood (Miiller and Samuels, 1982; septate ascospores have never been found in col- Romero and Samuels, 1991). Both genera are lections of E. operculata. The formation of septa characterized by their small, superficial ascomain the ascospores of species of Endoxyla is not ta, unitunicate asci with a n inamyloid apical ring delayed. Furthermore, the pattern of ascospore and brown, uniseptate ascospores with terminal septation within species of Endoxyla is consis- germ pores. Scotiosphaeria is apparently restricttent, even in abnormally enlarged o r misshapen ed to the necks of species of Endo.uyla and has ascospores. Species with ascospores that lack not been reported in the literature since its desepta o r a hyaline cell d o not form abnormal scription. ascospores with these features. Septa and a hyaline cell may, however, be absent in the abnorEndoxyla xanthostroma Untereiner, sp. nov. mal ascospores of species that normally possess FIGS. 20-23 ascospores with these features. The greatly enlarged and often multiseptate ascospores obStromata bene evoluta, denigrantia et elevantia suserved in some species of Endo.rj~lamay repre- perficiem ligni, entostroma compactum formantia tursent the fusion of ascospores within a single ascus. mas ascomatum circumdans; composita ex hyphis septatis brunneis 1.8-2.8 pm latis; infra ascomata The occurrence of Scotiosphaeria endoxj~linae entostroma diffusum luteum compositum ex hyphis in collections of E. parallela may have also con- septatis flavidis 1.8-2.8 pm latis, parietibus refringentributed to the confusion concerning ascospore tibus 0.5-1 pm latis, aggregata lutea formans. Ascomaseptation in this species. Scotiosphaeria endox- ta immersa, in turmis irregularibus 2-4.5 mm longis. j'linae possesses brown, uniseptate ascospores and ampulliformia, 450-630 x 245-310 fim, nigra, collis cylindricis discretim erumpentibus, 140-250 x 120small. dark ascomata and occurs o n the exposed 135 pm; apices plani, non protrudentes. Panes ventris necks of E. parallela and E. munkii. All speci- brunneus, 13-20 pm latus. stratum externum aspectu mens of Scleromyceti Suecia: no. 3 and numerous superficiali textura intricata. Asci octospori, clavatiother collections of E. parallela examined in this cylindrici, longipedicellati, 104-145 pm longi. pars sporifera 54-80 x 4.4-6.4 pm, annulo refringente inThe original de- amyloideo parvo. Ascosporae ellipsoideae-cylindricae. study include S . endod~.ylinae. scription of S. parallela (Fries, 1816) and Fries' 8.3-1 0.9 x 2.3-3.5 pm, bicellulares, compositae ex later description of this species (1823) clearly re- cellula brunnea, poro germinationis apicali praedita, fer to both fungi. The reports of Cooke (1873) et cellula hyalina 1-1.9 x 1.8-2.5 pm, oblique uniseriaand Nitschke (1867) also allude to the presence tae vel subinde biseriatae. Porus germinationis ascosporae apicalis typice apicem asci aspiciens. Paraphyses of S. endo.~ylinae on the necks of S. parallela. filiformes, septatae, 1.7-3.4 pm latae, sparsae. Habitat The association of Scotiosphaeria endo,~j~linae in ramis decorticatis putrifactis Pini. HOLOTYPUS: USA. MASSACHUSETTS: Petersham with species of Endoxyla was first reported by Kauffman (1929) who recognized that the asco- Twp.. Federation of Women's Clubs State Forest, 4 km W of Petersham, 29 Aug. 1987, W. A. Untereiner mata of two different fungi were present in a (TRTC 5 1039). single stroma. This species, the type of ScotioStromata well developed, evident as a n elesphaeria Sivanesan, was described from the necks of Endo,~ylinapini, a synonym of E. parallela vated darkening of the surface of the wood, con(Sivanesan, 1977). Although the taxonomic po- sisting of a compact entostroma that encloses sition of Scotiosphaeria is uncertain, the genus groups of ascomata; composed of brown o r dark should be placed in the Trichosphaeriaceae be- brown, septate hyphae 1.8-2.8 wm diam; below cause of its ascomatal morphology and unitun- the ascomata as a diffuse yellow entostroma comicate asci which possess a simple, inamyloid api- posed of septate, yellowish hyphae 1.8-2.8 k m cal ring. W i t h i n t h e Trichosphaeriaceae diam with refractive walls 0.5-1 pm thick that

FIGS. 16-23.

Endoxyla parallela and Endoxvla xanthostroma. 16-19. Endoxyla parallela. 16. Habit sketch

of ascomata, x 48. 17. Ascospores, x 2400. 18. Ascus, x 2400. 19. Abnormal ascospores, x 2400. 20-23. Endox?;laxanthostroma. 20. Habit sketch of ascomata, x 60. 2 1. Ascospores, x 2400. 22. Asci, x 2400. 23. Ab-

normal ascospores, x 2400.

form bright-yellow hyphal aggregates in ray cells ascospores. The prosenchymatous peridium, asof the wood. Ascomata Immersed, in irregular cus morphology and features of the ascospores groups 2-4.5 mm in length. ampulliform, 450- clearlq indicate, however, the close affinity of E. 630 x 245-3 10 pm, black, with cylindrical, sep- xarztho.strotna to the other members of this gearately erumpent necks. 140-250 x 120-1 35 pm; nus. apices of necks plane, confluent with the surface EXCLUDED A N D DOUBTFUL SPECIES of the stroma. Wall of venter brown. 13-20 bm thick; external layers composed of tubular hyUnless otherwise indicated, accepted names phae 2.1-4.7 pm diam; internal layers composed are preceded by an asterisk. of rectangular cells 4.1-1 2.8 x 1.5-2.8 pm that grade inwardly into compressed, thinner-walled E n d o s ~ ~ acericola Ell. & Ev., Proc. Acad. Nat. lu cells. Wall of neck brown. with external layers Sci. Philadelphia 46: 342. 1894. composed of ~ntenvo\en. tubular hyphae and This is Loph~ostoma yuadrrnucleaturn var. t n globose cells 1.1-4 5 pm dram that grade Inseptaturn (Peck) Chesters & Bell. wardly into rectangular cells. Wall of neck apex dark brown, 25-50 pm thick, composed of gloSPECIMEN EXAMINED: CANADA. ONT.~RIO: Granbose-polygonal cells 3 3-7.2 x 2.6-5.1 pm wlth ton, Jan. 1894, J Deumers no 2235 (HOLOTYPE, cell walls >0.5 pm thick that grade Inwardly Into NY). rectangular cells 1.2-3 4 pm dlam. Outer pendial Endo.q-la austrzaca Baumler. Ver. Zool.-Bot. Ges. layer of venter a teurtlra lrztrrcata In surface view, Wlen. 43: 277. 1893. composed of brown. septate, branched hyphae The type specimen was requested from B and 2.3-4.4 pm diam with cell walls <0.5 pm thick: hyphae arranged in indistinct radiate patches: W but could not be located. It was not possible internal layer a te2wturaangularis in surface view. to identify this species based on its description. composed of cells 6.3-1 1.4 x 2.9-7.4 pm; at lower magnifications appearing pseudoparen- Endo,syla uvocetia (Cooke & Ell.) Romero & Samuels. Sydowia 43: 23 I. 199 1. chymatous. Munk pores not o b s e r ~ e d .Outer peridial layer of neck a tcxiura porrcctu to a i = Sphaeria avoceitu Cooke and Ell.. Grcv~llea 8: intrrrata in surface view, cells indistinct. Asci I S . 1879. = Ccratostonza avocrtta (Cooke and Ell.) Sacc., eight-spored. clavate-cylindrical. long-stipitate, Syll Fung. 1: 216. 1881 total length 104-145 pm, pars sporifcra 54-80 x 4.4-6.4 pm, with a small, refractive. inamyloid Sphaeria avocetta resembles species of Endoapical ring. Ascospores ellipsoid-cylindrical. 8.3ryla In ~ t ~mmersed s ascomata and brown asco10.9 x 2.3-3.5 pm, bicellular, composed of a spores which possess an aplcal germ pore (Romebrown cell with an apical germ pore. and a hy- ro and Samuels. 1991) The loosening asci wlth aline cell 1-1.9 x 1.8-2.5 pm, obliquely unise- a draporthaceous aprcal nng, abundant paraphriate or occasionally biseriate. Germ pore of the 5 ses and absence of a stroma are, howm er, feaapical ascospore typically oriented toward the tures uncharacteristic of I: ndo.~l*/a suggested As ascus apex. Paraphyses filiform. septate. 1.7-3.4 by Romero and Samuels (1991) thrs species IS pm diam, sparse in mature ascomata. Anamorph better accommodated in the Dlaporthales not known. HA~ITAT decort~cated On rottlng wood of Pznus sp. HOLOTYPE USA. MASSACHUSETTS. Petersham Twp , Federation of Women's Clubs State Forest, 4 km W of Petersham, on decorticated rottlng branch of P~nussp. ( P srrobus 7) on the ground, 29 Aug 1987, R . C'ntererner (TRTC 5 1039) 1 ETYM~LOGY. wanthos (EavOoT) = yellow, In Greek, reference to the yellow stroma

Endos.vla cupparldis R. Rao. Sydowia 25: 54. 1971.

The type specimen, requested from the herbarlum of MSG College, Maharasthtra, Indla, E was not received. Based on its descr~ption, cappurldls appears to be a member of the Diatrq paceae

Erzdox~~la xanthostrorna is recognized by the dark, slightly elevated stroma that encloses groups of ascomata and yellow stroma beneath the ascomata, and by the uniseriate arrangement of the

Endox~,la ctrrhosa (Pers. : Fr.) v. Arx & Miiller. Beitr. Kryptogamenfl. Schweiz 11(1): 355. 1954.

= Sphaeria cirrhosa Pers., Syn. Meth. Fung. p. 59.

1801.

= Sphaeria cirrhosa Pers. :Fr., Syst. Mycol. 2(2):

475. 1823.

EXAMRUED:GERMANY. Westfalen, NienSPECIMEN berge bei Miinster, April 1865, T. Nitschke (HOLOTYPE, B).

= Ceratostoma cirrhosum (Pers. :Fr.) Fuckel, Jb.

* = Ceratostomella cirrhosa

Nassau. Ver. Naturk. 23-24: 127. 1870. (Pers. : Fr.) Sacc., Michelia 1: 370. 1878.

Endoxvla fraxini (Ell. & Ev.) Ell. & Ev., N . A m .

Pyrenomyc. p. 521. 1892.

N o specimens o f 5'. cirrhosa were found in Persoon's herbarium. E D: SPECIMENSX A M ~ EDENMARK. Sjaelland, Dyrehaven, 22 Nov. 1964, A. Munk (C). ITALY. Aug. 1878, P. '4. Saccardo (PAD). USA. INDIANA: Bloomington, 23 Aug. 1958, R. F. Cam (DAOM 136283).

Thyridariafraxini Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 42: 223. 1890.

T h i s is Cryptosphaeria eunomia. Pirozynski (1969) a n d Wehmeyer (1926b) discussed the syno n y m y o f this species. SPECIMENS EXAMINED: CANADA. ONTARIO: London, Jan. and Mar., 1890, J. Dearness no. 1371, 1496 (HOLOTYPE, FH). USA. MICHIGAN: Arbor, Oct. Ann 1923. L. E. Wehmeyer no. P234 (DAOM 122012).

Endoxyla comata Starback, Ark. Bot. 2: 19. 1904 (as Endoxylon cornatum).
T h i s i s Peroneutypa heteracantha (Sacc.) Berlese. SPECIMENXAMINED: E BRAZIL. Porto Alegre, 29 Sept. 192 1. K. Starback (HOLOTYPE, S).

Endo,xjda hvalostonza M u n k , Bot. Tidsskr. 61:

62. 1965. T h i s is Ceratostomella hyalostoma (Munk) Untereiner, comb. nov. SPECIMENS EXAMINED: DENMARK. Sjaelland, Ermelunden. 15 Dec. 1963, A. Munk (HOLOTYPE, C). SCOTLAND. Den of Airlie, 6 June 198 1, B. J. Copprns no. 4869 (E VC90).

Endo.x.vla eut,vpoides (Ell. & Ev.) Ell. & Ev., N.

A m . Pyrenomyc. p. 520. 1892.

= Thyrrdaria eut.vpoides Ell. & Ev., J. Mycol. 4:

78. 1888. T h i s is :Melanornr~iu fuscidulum Sacc. SPECIMEN EXAMINED: USA. LOUISIANA: Point a la Hache, 2 July 1888, A. B. Langlois no. 1377 (HOLOTYPE. NY).

Endo-xyla inusta (Cooke) Ell. & Ev., N. A m . Pyrenomyc. p. 520. 1892.

Sphaeria inusta Cooke, Grevillea 7: 52. 1878. = Kalmusia inusta (Cooke) Sacc., Syll. Fung. 2: 144. 1883. = Xvlosphacria inusta (Cooke) Cooke, Grevillea 17: 86. 1889.
T h i s is a species o f Karstenula Speg. SPECIMEN EXAMINED: USA. GEORGIA: Darien, 24 1 3. H. W. Ravenel (HOLOTYPE, K).

E n d ~ x ~ vexcelsior (Mouton) Munk, Bot. Tidsskr. la

61: 66. 1965.

* = Ceratostomella excelsiorMouton, Bull. Soc. Roy.

Bot. Belgique 36: 12. 1897. SPECIMENSEXAMINED: BELGIUM. Beaufay, 1897, V. .Wouton (HOLOTYPE, BR). DENMARK. Sjaelland, Lellinge, 14 May 1964, A. Munk (C).

Endoxyla laevirostris M u n k , Bot. Tidsskr. 61: 62. 1965, nom. illeg. Art. 63.1. = Endoxyla rostrata (Tode : Fr.) Munk, Dansk Bot. Ark. 17(1): 196. 1957, nom. inval. Art. 33.2. Sphaeria rostrata Tode, Fungi Mecklenb. Sel.

A4nthostoma.ferrugineumNitschke, Pyrenomyc. G e r m . 1: 118. 1867.

Shear (1938) treated this species a s Carnarops ferruginea (Nits.) Shear. Nannfeldt (1972) excluded this species from Camarops a n d suggested that i t m a y be related t o E. operculata. Anthostoma ferrugineum possesses a well-developed stroma, uniseptate, brown ascospores with a n apical germ pore a n d asci with a n inamyloid ring. It is close t o Pseudovalsariafoedens (P. Karsten) Spooner.

Ceratostoma rostratum (Tode : Fr.) Fuckel, Jb. Nassau. Ver. Naturk. 23-24: 127. 1870. * = Ceratostomella rostrata (Tode: Fr.) Sacc., Syll. Fung. 1: 408. 1882. = Ceratostomella ampullasca (Cooke) Sacc., Syll. Fung. 1: 409. 1882. Sphaeria ampullasca Cooke, Handb. Brit. Fungi 2: 876. 1871.

p. 14. 1791. 473. 1823.

= Sphaeria rostrata Tode : Fr., Syst. Mycol. 2(2):

M u n k (1965) established E. laevirostris t o solve the nomenclatural problems surrounding C. ros-

frafa and created a superfluous name. The type material for S . rostrata is lost and the identity of this species is unclear. A specimen under this name from Fries' herbarium represents a member of the Calosphaeriales. and Scleromyceti Sueciz no. 116 Sphaeria rostrata was identified as Cerafostornella investifu (Schw.) Starback. A specimen of C. ro.rtrutum from Fuckel's herbarium is Ceratosfotnella cirrhosu. Material from Saccardo's herbarium was not examined. T h e nomenclature of Crrafostornella rostrafa requires additional study.
EXAMINED: DENMARK. Jylland. StaksSPECIMENS rode near Juelsminde. 12 Oct. 1963. '1. .Mrtr~k(HOLOTYPE E. laevrrosfri~. ENGLAND. Shere, Feb. C). 1869. Dr. Capron (HOLOTYPE ('. atnpullasca. K). GERMANY. L. Fuckcl, as Cerarostofnarostratitrlz (G). SWEDEN. Scleromyceti Suecire no. 116 (1st ed.), E. .If. Frres (FH): Sm%land.Femsjo, E. .if. Frres (UPS).

do..;!sla. Ellis' exsiccata for this species (North American Fungi no. 90 and North American Fungi 2nd ser.. no. 1959) were not examined in this study. Endoxj~larnurzgrferue Hennings, Hedwigia 47:
258. 1908. This corresponds to Ezitjpa sptnosu (Pers. : Fr.) Tul. SPECIMEN EXAMINED: PHILIPPINES. Davao, Mindanao. 4 Apr. 1904. Cbpeland no. 825 (SYNTYPE, NY).

Endo.ujsla r)zrcrosporaFerraris. Malpighia 16: 449.

1902. The type specimen was requested from P A D but could not be located. Based on its d e s c r ~ p tion. E. tnicrospora may be a member of the Diatrypaceae.

Enclos~.la l ~ n e a f a (Alb. & Schw. : Fr.) Cooke. G r e + ~ l l e a 0: 82. 1892. 2

= Sphaeria lineata Alb. & Schw.. Consp. Fung. Lusat. p. 30. 1805. = Sphaeria lineata Alb. & Schw. : Fr., Syst. Mycol. 2(1): 273. 1822.
Nitschke. AdThis is a species ofA4nfhostotnu ditional material in Schweinitz's herbarium and material from Schweinitz a t K represent members of the Diatrypaceae. SPECIMENS EXAMINED: GERMANY. Niesky, as S . lrneata A. S. (PH). USA. MASSACHUSETTS and Salem PENNSYLVANIA: and Bethlehem. c . ~ herb. Schweinitz (K).

Etzdos1.1~ popull Romell. In Wlnter. Hedwlg~a 24: 263. 1885.

This is a species of C r ~ p t o . ~ p h u r r ~ a . SPECIMEN EXAMINED: SWEDEN. Uppsala. 1 Jan. 1885, L. Kot~~ell (HOLOTYPE. S 1588 1).

Endo.x\,la rosfruta (Tode : Fr.) Munk. Dansk Bot.

Ark. 17(1): 196. 1957. See Endosltla laevrrostrrs

Et1do.i-!.la \>estrtu (Sacc.) Munk. Bot. Tidsskr. 61:

64. 1965.

Lasrosphaeria luteohasis (Ell.) Ell. & Ev., N. Am.

Pyrenomyc. p. 147. 1892.

= Sphaeria luteohasis Ell., Bull. Torrey Bot. Club 6: 133. 1877. = Byssosphaeria luteohasis (Ell.)Cooke. Grevillea 15: 122. 1887. = Eutypa luteohasis (Ell.)Sacc.. Syll. Fung. 1 : 17 1. 1882.

Sphaeria luteohasis is described as possessing brown, uniseptate ascospores and superficial or subsuperficial ascomata surrounded by a yellow subiculum. A specimen annotated S. luteohasis in Ellis' hand a t F H (sub norn. Erzdo-~~.Ia (Kulmusia) parallela e x herb. W . G. Farlow) corresponds closely to this description. T h e ascospores of S. luteohasis are similar t o those of E . parallela. A s suggested by R a p p a z ( 1 9 8 7 ) Splzaeria luteohasis is likely a member of Etz-

= C'eratostomella vestrta Sacc., Michelia 1: 370. 1878. = Lerttorrzitella resrlta (Sacc.) Hohnel. Ann. Mycol. 3: 552. 1905. = Cerasto???is vestita (Sacc.)Clements, Gen. Fungi p. 259. 1931. = Ceratostornella vestita Sacc. var. ~arvicensrs ro\ e. G J. Bot. 23: 131. 1885. * = Cerarosror?~ellanvestrta (Schw.) Starback, Bih.
i Kongl. Svenska Vetensk.-Akad. Handl. 19(3) no.
2: 26. 1894. = Sphaeria investita Schw., Trans. Amer. Philos. Soc.. ser. 2. vol 4: 216. 1834. = Ceratostoma invest~rum(Schw.) Ell. & Ev. N . Am. Pyrenomyc. p. 193. 1892.
SPECIMENS EXAMINED: AUSTRIA. Pressbaum, Sattelberg. 28 June 1903. t- \.on flohnel3150 ( F H ) . DENMARK. Sjaelland, Dyrehaven, 3 Mar. 1965, ''1. .\funk (C). ENGLAND. Sutton. 25 Oct. 1884 (HOLOTYPE C'. resrita var. varvicensrs. K ) . ITALY. Sept. 1878. P. .4. Saccardo (HOLOTYPE C. resrita, PAD). USA.

PENNSYLVANIA: Bethlehem, L. Schweinitz (HOLOTYPE S. investita, PH).

E n d o x y l a yerhae Spegazzini, A n . Mus. Nac. Hist. N a t . B. Aires 17: 1 18. 1909.

T h e type is a very p o o r specimen representing a species o f Valsa SPECIMEN EXAMINED: PARAGUAY. San Pedro, Feb 1907. C. Spegazzrni (HOLOTYPE. LPS).
ACKNOWLEDGMENTS

I thank David Malloch and Greg Thorn for their useful suggestions and critical comments. Dr. Mark Finkelstein and Dr. Walter Cams assisted with the Latin diagnoses. The curators of the following herbaria are thanked for the loan of specimens: B, BR, C, DAOM. E, FH, G, H, IMI, K, LPS, M, MBB, MICH, NEB, NY, PAD, PH, S. TRTC, UC, and UPS. Special thanks are extended to Dr. Jean Boise Cargill (FH), Michelle Seidl (UC) and Dr. R. Moberg (UPS) for permitting me to study first edition material of Fries' Scleromyceti Suecic. This research was funded by a postgraduate scholarship to W.A.U. and an operating grant to Dr. David Malloch from the Natural Sciences and Engineering Research Council of Canada.
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