T A X O N O M Y AND M O R P H O L O G Y O F A N E W S P E C I E S OF HYPOCREA ON MARASMIUS SUSAN T. CAREY AND CLARK T. ROOERSON

Carey, Susan T. and Clark T. Rogerson (The New York Botanical Garden, Bronx, NY 10458). Taxonomy and morphology of a new species of Hypocrea on Marasmius. Brittonia 28: 381-389. 1976.--A new species of Hypocrea, It. avellanea Rogerson & Carey, parasitizing Marasmius subnudus (Ellis) Peck, is described. The development of its imperfect state, perithecia, and asci was studied in culture and compared with that of previously studied species of Hypocrea, Hypomyces and Nectria.

INTRODUCTION

Deformed basidiocarps of Marasmius subnudus (Ellis) Peck 1, collected over a period of many years, were partially or completely covered with either the imperfect or perfect state or both states of a new species of Hypocrea. H y p o e r e a a v e l l a n e a Rogerson & Carey, sp. nov. Status Verticillii mycelium bubalinum, matricem, Marasrniurn subnudum tegens et deformans, subiculum faciens. Perithecia globoso-ovata ad globoso-papillata, (100--)150--270(-310) X (100--) 120--200 #m, bubalina ad brunnea tempore maturescenti, de visu superficiali sub magnificatione inferiori apice atrobrunnea, parfirn vel omino in subiculo inclusa; periphysibus et paraphysibus apicalibus filamentosis atque hyalinis; asci octospori, unitunicafi, cylindrico-clavati (51-)57-87(-97) X 3-5 ~m; ascis apice simplex, parum incrassato, minute porato; ascosporae uniseriate, hyalinae, leves ad minute verrucosae, bicellulares; cellula superua plerumque globosa, interdum truncatosubglobosa, 2-4 X 2-3.5 /~m, inferna plerumque truncato-ellipsoidea, interdum truncato-subglobosa, raro globosa, 2.5-5 X 2-3.5/zm, deinde in duns cellulas decedentes. Mycelium thin at first, "light buff ''2 to "ochraceous buff," usually covering and deforming the host with a Verticillium state, becoming a subiculum; subiculum textura intricata (Fig. 1D) "avellaneous," "wood brown," to "buffy brown"; perithecia "Dresden brown" at maturity, globose-ovate to globose-papillate, (100-)150-270 (-310) X (100-) 120-200/zm, partially or completely embedded in subiculum (Fig. 1A), separate or densely compacted, apices conspicuously darker when viewed from above under low magnification; apices of mature perithecia turning violet-red in KOH; periphyses hyaline, filamentous, swollen at tips (Fig. 1B); perithecial wall yellow, textura angularis to textura prismatica, 10-18 /zm wide (Fig. 1C); apical paraphyses hyaline, filamentous, disintegrating as the asci form; asci 8-spored, unitunicate, cylindric-clavate ( 5 1 - ) 5 7 - 8 7 ( - 9 7 ) X 3-5 /zm (Fig. 1E); apex a simple pad pierced by a tiny pore at maturity (Fig. 1G); ascospores uniseriate, hyaline, smooth to minutely verrucose, two-celled, upper cell usually globose, sometimes truncate-subglobose, 2-4 X 2-3.5 #m, lower cell usually truncate-ellipsoid, sometimes truncate-subglobose, rarely globose, 2.5-5 X 2-3.5 /zm, disarticulafing into partascospores (Fig. 1E, F ) ; part-ascospores usually becoming ellipsoid after discharge (Fig. I F ) . Vertic,7lium imperfect state: phialophores up to 200/zm long and slender, hyaline, 5 /zm at the base tapering to the tip, with 1-6 whorls of phialides, 3-6 phialides per whorl; phialides broadest slightly above the base (ten-pin shaped), 1Collybia subnuda (Ellis ex Peck) Gilliam, Mycotaxon 4: 136-137. 1976.

Color names in quotation marks refer to Ridgway (1912). BRITTONIA28: 381-389. October-December, 1976.

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FIG. 1. ttypocrea avellanea. A, diagram of longitudinal section of perithecium with phialophores of Verticillium state on wall; B, perithecial neck with protruding ascus discharging ascospores; C, longitudinal section of perithecial wall; D, subiculum at surface of perithecial wall; E, an ascus containing eight ascospores; F, discharged part-ascospores; G, development of ascus apex; H-J, VerticiUium state from a single ascospore culture (69-77 1-6), H, phialophore; I, phialosporogenesis; J, released phialospores. tapering to a narrow tip, hyaline, 11.5-25 /zm long, 2.3-4 /zm at widest point; phialospore-heads hyaline, globose, 7-20 /zm; phialospores hyaline, ellipsoid, 3-9.2 • 1.5-5/zm. Host: Marasmius subnudus (Ellis) Peck. TYPE: U N I T E D STATES. MASSACHUSETTS: Franklin County, Leverett, Mt. T o b y State Forest, 27 Aug 1958, H. E. Bigelow & M. E. Bigelow ( M E B B 2471) (HOLOTYPE: N Y ; ISOW'ZPES: K, MASS). PARATYPES: U N I T E D STATES. CONNECTICUT: Sharon, 30 Jul 1969, Rogerson 69-77 & G. L. Smith, ( N Y ) . MASSACHUSETTS: Petersham, Harvard Forest, 21 Aug 1946, Rogerson s.n., immature (NY) ; Leverett, Mt. T o b y State Forest, 28 Sep 1958,

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FIc. 2. Marasrnius subnudus parasitized by Hypocrea avellanea (Rogerson 63-83).

M. E. B. Bigelow 2528 (NY, MASS), same locality, 13 Sep 1960, H. E. Bigelow et al. (M. E. B. Bigelow 2824) (NY, MASS), same locality, 25 Aug 1963, Rogerson 63-28, immature (NY), same locality, 25 Aug 1963, Rogerson 63-31 (NY), Savoy, Tannery Falls, 15 Sep 1960, H. E. Bigelow et al. (M. E. B. Bigelow 2833), immature (NY, MASS), Northfield, 2 Oct 1961, H. E. Bigelow et al. (M. E. B. Bigelow 3147) (MASS), near Cricket Hill, Conway State Forest, 24 Aug 1963, Rogerson 63-19, immature (NY), Ruggles Pond, Wendell State Forest, 16 Jun 1973, Carey 73-1 (NY). NEW YORK: Ithaca, Cascadilla Ravine, 19 Aug 1947, Rogerson 1601 (NY), Ithaca, Coy Glen, 19 Aug 1947, Rogerson 1605 (NY), Red Rock, 3 Sep 1958, S. J. Smith 25740 et al., (NY, NYS), Stony Point, 18 Oct 1959, Long (Rogerson 59-3), immature (NY), same locality, 1 Nov 1959, Long & Cage (Rogerson 59-94), immature (NY), 12 miles S. Kingston, near Esopus Gorge, 17 Sep 1960, Rogerson 60-219 (NY), Ladentown, Torn Mt., 2 Oct 1960, Long s.n., immature (NY), N. Oneonta, Winnie Hill, 21 Sep 1963, Rogerson 63-83 (Fig. 2) (NY), Bloomingdale, Saranac River, 11 Sep 1965, Rogerson 65-64, immature (NY), Wadhams, Plattsburg State College Twin Valleys Camp, 9 Sep 1967, Rogerson 67-139 & Carey (NY), near Wilton, Mt. McGregor, 10 Oct 1968, Rogerson 68-161 & S. J. Smith (NY), near Millbrook, Cary Arboretum, NYBG, 6 Sep 1975, Rogerson s.n., immature (NY). NORTH CAROLINA: Great Smoky Mts. National Park, Indian River, 14 Aug 1968, Rogerson s.n., immature (NY). Specimens with Verticillium state only: UNITED STATES. CONNECTICUT: Greenwich, 2 Oct 1974, Rogerson ,& Carey 74-14 (NY). MASSACHUSETTS: near Cricket Hill, Conway State Forest, 24 Aug 1963, Rogerson 63-48 (NY); Ruggles Pond, Wendell State Forest, 16 Jun 1973, Carey 73-2 (NY), Rutland State Park, 22 Jul I975, Kramer (Carey 75-1). NEW YORK: Waterford, 5 Aug I905, Burnham s.n., identified by C. H. Peck as Verticillium enecans Speg. (NYS); Vooreesville, Aug, C. H. Peck s.n., identified as V. enecans (NYS) ; NE. Tripoli, SW. Fort Ann, 27 Sep 1914, Burnham s.n. (NY); SE Ithaca, Coy Glen, 11 Aug 1948, Rogerson 2531 (NY); same locality, 19 Aug 1948, Rogerson 1606 (NY); same locality, 29 Sep 1951, Kor/s.n., (NY); same locality, 4 Sep 1952, Rogerson s.n. (NY); Red Rock, 3 Sep 1958, S. I. Smith 25772 et al. (NY, NYS); Mianus Gorge, 12 Aug 1960,

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Fi~s. 3-8. Hypocrea avellanea (Rogerson 66-77M). 3-4. Verticillium imperfect state. 3. X 1000. 4. Phase microscopy, X 2250. 5-8. Perithecial development. 5. J'oining of filamentous hypha with bulbous hypha in subiculum, X 2250. 6. Coil invested by surrounding hyphae, X 1125. 7. Cross section of young perithecium, X 1125. 8. Longitudinal section of perithecium with apical paraphyses, X 500.

Yarrow s.n. ( N Y ) ; Montauk, Hither Hills State Park, 23 Jul 1967, Rogerson 67-28 ( N Y ) ; Southfields, 2 Sep 1972, Stein (Rogerson 62-89) ( N Y ) . WISCONSIN: Univ. of Wisconsin Arboretum, 5 Sep 1953, Rogerson s.n. ( N Y ) . Asci from the fresh collection of Rogerson 69-77 were placed on corn meal agar medium (5% agar) in a Petri plate and single ascospores were dissected from them with the aid of a micromanipulator. Twenty-four ascospores, from three asci, were dissected; only five of these germinated, two from one ascus and three from another.

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The cultures of these five ascospores produced only the Verticillium state in Petri plates on corn meal, potato dextrose and oatmeal agar media (Ainsworth, 1961). No subiculum or perithecia formed in these cultures or in combinations of two of them on oatmeal agar medium. Perithecia were produced by a culture, formed from a combination of all five cultures of ascospores (69-77 1-5, 6; 69-77 IH-2, 5, 6) and by a culture obtained from the combined mycelia of many germinating ascospores (6977M). The perithecia were produced over a period of from two weeks to a month on oatmeal agar in Petri dishes incubated at room temperature. Blocks of the 69-77M culture, containing the developing perithecia, were cut out, fixed, embedded, sectioned and mounted by a previously described technique (Canham, 1969). The slides were stained in Ehrlich's hematoxylin (10 min), washed in two changes of 50% ethanol, brought through 70% ethanol, 1% fast green in 95% ethanol (5 rain), 95% ethanol (3 changes), and permanently mounted in Euparal. Clumps of asci from the same culture were removed from perithecia crushed on a slide, transferred to a drop of 2% orcein in solution of equal parts of acetic and propionic acids, flattened under a cover glass, sealed with nail polish, and examined the next day for meiotic figures and chromosomes. Cultures (69-77-1-6, 73-1M, 74-14M) of the Verticillium state of H. avellanea on potato dextrose agar medium, incubated in the dark at 20 C for 31 days (when growth reached the edges of the Petri plates), were white to "avellaneous" on the upper surface, and a "colonial buff" to "Natal brown" on the reverse. The aerial hyphae of these cultures produced hyaline phialophores, 11-138 /zm long x 3-5 /zm wide at the base, tapering toward the tip, at which there was a whorl of phialides. The phialophores had 1-6 whorls of phialides with 3-8 phialides in a whorl. The phialides were "ten-pin" shaped, hyaline, 12-30 long and 2-3 /zm at the widest point, usually bent upward at about a 45 degree angle, and were of different lengths at one whorl. The longest phialide was almost always uppermost. The phialophores frequently branched at a whorl (Fig. 1H; Fig. 3) producing a short branch with a whorl of phialides at its tip. The phialospores were hyaline, ellipsoid, smooth, 3-8 • 1-2.5/zm, and produced successively from the narrow tip of the phialide, where they formed clusters, 11-23/zm in diameter (Fig. l I - J , Fig. 4). Intercalary or terminal, hyaline, smooth, globose, 5-6 /zm, chlamydospores were rarely formed in older parts of the culture. Two types of hyphae, both of which stained deeply with cotton blue and orcein. were differentiated in the subiculum. One type of hypha, which was short and bulbous, was joined by the other type, which was long and filamentous (Fig. 5). No plasmogamy was observed. A perithecium was initiated around the joined hyphae, after they were enveloped with a coiled hypha. The coiled structure (Fig. 6) was quickly invested with hyphae from the surrounding subiculum and the base of the short, bulbous hypha. The developing perithecium became globose as its cells increased and formed tissues. In cross section (Fig. 7) the young perithecium (35/z diam) consisted of a large multinucleate cell in the center, probably the bulbous hypha, surrounded by smaller, multinucleate cells from the coil, in turn surrounded by several layers of small cells from the investing hyphae. Later, the centrum was composed of a septate, meandering hypha, with large multinucleate cells, embedded in smaller, thin-walled parenchymatous cells, and was enclosed by a perithecial wall, consisting of 6-8 layers of elongate cells with moderately thick cell walls. The parenchymatous cells of the centrum enlarged, their cell walls became very thin, and the cells disintegrated. In the upper part of the centrum a mass of deeply staining small cells developed, probably from a cell or several cells of the meandering hypha. At the same time, another meristem formed above the mass of small cells and produced

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FIos. 9-15. Hypocrea avellanea. (Rogerson 66-77M). 9. Neck of perithecium, X 1125. 10-15. Development of asci. 10. A crozier showing four nuclei, X 2250. 11. First meiotic division, late prophase, five chromosomes, X 2250. 12. Multinucleate cells found intermingled with young asci, X 2250. 13. Second meiotic division, metaphase, five chromosomes in each nucleus, X 1125. 14. Left ascus, metaphase of last mitotic division; right ascus, the resulting sixteen nuclei, X 1125. 15. Apices of asci, X 1125.

apical paraphyses (Fig. 8) that grew into the centrum, pushing the mass of small cells against the remains of the parenchymatous cells and meandering hypha. The small cells were spread out in a semicircle in front of the advancing apical paraphyses and when they reached the base of the centrum, these cells formed croziers. Wall ~:ells, at the morphological apex of the perithecium directly above the developing apical paraphyses, also produced hyphae. These hyphae developed in a double file, progressively outward toward the surface, separated at their tips on each side and became periphyses that lined the ostiolar canal (Fig. 9). The two nuclei of an ascogenous cell divided synchronously (Fig. 10) and a crozier formed. The two nuclei in the penultimate cell of the crozier fused, this cell elongated into an ascus, and the stages of meiosis ensued. At prophase I there were five bivalents (Fig. 1 [ ), a long one, a slightly shorter one attached to the nucleolus, two mediumsized ones, and a short one. Intermingled in the ascogenous cells, croziers and developing asci, cells with two to four nuclei (Fig. 12). These cells disappeared as the asci matured. At metaphase II, five chromosomes were counted in each of the two

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dividing nuclei (Fig. 13). After the two meiotic divisions occurred, there were two mitotic divisions within an ascus (Fig. 14). Ascospores were delimited around each of the eight pairs of nuclei, the products of the last mitotic division. Eight bicellular ascospores, each with uninucleate cells or part-ascospores, resulted. The partascospores of each ascospore separated at the median septum. The ascus apex was thin-walled during the stages of meiosis, became thicker during the subsequent mitosis, and when the part-ascospores were separate, consisted of a thin pad which was pierced in the center by a tiny pore (Fig. 15). No other apical structures were visible when the ascus was stained with Parker's blue-black Quink ink, fast green, erythrosin, cotton blue or iodine. DISCUSSION

Hypocrea avellanea belongs to that group of Hypocrea species that Petch (1937) placed in Protocrea, with perithecia immersed in subicula instead of, like most Hypocrea species, embedded in stromata. Doi (1972) accepted Protocrea with two species; P. farinosa (Berk. & Br.) Petch, which he described as having a mealy subiculum and a Cephalosporium imperfect state, and P. delicatula (Tut.) Petch, which has a spongy subiculum and a Verticillium imperfect state. Dingley (1952) and Rogerson (1970) did not think that the presence or absence of a stroma was sufficient basis for the separation of these Hypocrea species from the others and thus synonymized Protocrea under Hypocrea. Hypocrea species with subicula show affinities with Hypomyces, the species of which commonly have subicula, and a mycoparasitic habit. That there is a close relationship between Hypocrea and Hypomyces is further supported by the existence of other intermediate species with characteristics of both genera. Hypomyces papyraceus (Ellis & Holway) Seaver (1910) and Hypomyces tremellicola (Ell. & Ev.) Rogerson in Samuels (1976) resemble species of Hypocrea. Ellis and Holway (in Ellis and Everhart, 1886) placed H. papyraceus in Hypocrea because its two-celled ascospores, although they have a small apiculus at each end, a characteristic of Hypomyces species, do separate at maturity into part-ascospores, a characteristic of Hypocrea species. Moravec (1956) thought that Hypocrea stipata Fuckel, a synonym of H. papyraceus, was sufficiently different from Hypomyces and Hypocrea to serve as the basis of a new genus, A rachnocrea, which genus was accepted by Doi (1972), who added another species, A. scabrida Doi. Itypomyces papyraceus produces a very thin, white subiculum on decaying wood and fungi. It has not been grown in culture. Hypomyces tremellicola was first described by Ellis and Everhart (1892) as a Hypocrea, because it has small, spiny, non-apiculate, two-celled ascospores, but the two cells do not separate at maturity. Samuels (1976) studied//, tremellicola along with other Hypomyces species that show similarities to species of Nectria subg. ttyphonectrica, and concluded that it belonged in Hypomyces because its perithecia developed like other species of Hypomyces. Hypomyces tremellicola has a Verticillium imperfect state and produces a white subiculum on species of Crepidotus, Polyporus, and Pleurotus. The development of the perithecia of Hypocrea avellanea resembles the Nectria-type of Luttrell (1951) in the presence of: apical paraphyses, a true perithecial wall, a schizogenous ostiole with periphyses, and asci arising from croziers in a crescent-shaped layer at the base of the centrum. The parenchyma in the centrum shows affinities with the Diaporthe developmental type but is present only early in development and disintegrates with the formation of the apical paraphyses. It is not the same as the parenchyma of Nectria haematococca Berk & Br. which, according to Hanlin (1971) formed from the swelling of the cells of the apical paraphyses. Nor is it like the pseudoparenchyma of Neuronectria peziza (Tode ex Fr.) Munk [Nectria peziza (Tode

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ex Fr.) Ft. according to Samuels, 1976], described by Hanlin (1963a) as developing early in centrum formation from an apical meristem and as persisting until the asci formed. Centripetal paraphyses have been described in many of hypocrealean fungi (see Samuels, 1973), but were not observed in H. avellanea. The perithecial development of H. avellanea is intermediate between that which has been found in Hypomyces and Nectria. According to Samuels (1973) the Hypomyces-type is initiated in a sexually stimulated subiculum by an extensive, symmetrical coil that remains distinct throughout development; it has a centrum formed by centripetal paraphyses and apical paraphyses, and has a subhymenial parenchyma. The Nectria-type is initiated by a solitary, asymmetrical coil, usually does not have centripetal paraphyses and is characterized by a lysis of some cells; it frequently has apical paraphyses. The perithecia of Hypocrea avellanea are probably initiated in a sexually stimulated subiculum and the species is most likely heterothallic, since only mulfi-ascospore cultures produced perithecia. The initials are of two types and a short symmetrical coil develops. There is a lysis of cells in the centrum and apical paraphyses are present. Although the nuclear condition of the cells of the ascogenous system was not determined, the binucleate, trinucleate, and quadrinucleate cells intermingled with the asci, in squash mounts, are most likely part of this system. The binucleate cells are always smaller than the others (Fig. 12) ; perhaps they are formed by cytokinesis of a quadrinucleate cell, resulting in two binucleate cells. Hanlin (1961, 1963a,b, 1965, 1971) indicated that similar mulfinucleate cells, which he found in squash mounts of several hypocrealean fungi, formed short hyphal extensions into which two nuclei migrated and these produced croziers. The croziers of tt. avellanea form from binucleate cells. The life cycle of H. avellanea, as described here is incomplete, since the fungus ceased to produce perithecia in culture. However, in general, its characteristics conform to what has been found in other hypocrealean fungi. Other investigators have found variations among those species with the Nectria developmental type. These variations increase with each species studied. Four stromatic Hypocrea species have been examined: H. gelatinosa (Tode ex Ft.) Ft. (Vincens, 1917), H. spinulosa Fuckel [Doquet, 1957, as Creopus spinulosa (Fuckel) Moravec], H. schweinitzii (Fr.) Sacc. (Hanlin, 1965), and H. citrina (Fr.) Fr. (Canham, 1969), and among these there are similarities and differences. The perithecia of H. schweinitzii and H. spinulosa are initiated from coiled hyphae, those of H. gelatinosa and H. citrina have initials of two sizes and the coil is secondary, as in H. avellanea. The perithecial walls of H. gelatinosa and H. spinulosa develop from stromatic hyphae, that of H. schweinitzii develops from the base of the coil, and those of H. citrina and H. avellanea form from hyphae from the base of the coil and surrounding hyphae. The cells of the coil of H. schweinitzii are mulfinucleate, those of H. citrina are binucleate. The young perithecium of H. gelatinosa has a centrum of parenchyma and develops centripetal paraphyses later. The centra of H. spinulosa, tt. schweinitzii, and H. citrina have centripetal paraphyses, apical paraphyses, and a subhymenial parenchyma. Hypoerea schweinitzii has a haploid chromosome number of four (Hanlin, 1965), H. spinulosa probably has seven [Mathieson, 1952, as Chromocrea spinulosa (Fuckel) Petch]; H. citrina (Canham, 1969) and H. avellanea have five. ACKNOWLEDGMENTS

We wish to thank Dr. Margaret Bigelow and Dr. Howard Bigelow for the collections of ttypocrea avellanea, Dr. Gary Smith and Mr. Rupert Barneby for assistance with the Latin description, and Dr. Marjorie Anchel for the review of the manuscript.

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T h i s research was p a r t i a l l y s u p p o r t e d b y a General R e s e a r c h S u p p o r t G r a n t S01 R R 0 5 6 2 1 , f r o m the D i v i s i o n of R e s e a r c h Resources, N a t i o n a l I n s t i t u t e s of H e a l t h . LITERATURE CITED Ainsworth, G. C. 1961. Ainsworth and Bisby's Dictionary of the Fungi. C.M.I., Kew, Surrey. Canham, S. Carey 1969. Taxonomy and morphology of Hypocrea citrina. Mycologia 61: 315-331. Dingley, J . M . 1952. The Hypocreales of New Zealand III. The genus Hypocrea. Trans. Roy. Soc. New Zealand Bot. 79: 323-337. Doi, 3(. 1972. Revision of the Hypocreales with cultural observations IV. The genus Hypocrea and its allies in Japan (2) enumeration of the species. Bull. Natl. Sci. Mus. 15: 649-751. Doquet, G. 1957. Organog~nie du Creopus spinulosus (Fuck.) Moravec. Organog6nie eompar6e de quelques HypocrSales du mfime type. Bull. Soc. Mycol. France 73: 144-164. Ellis, J. B. & B. M. Everhart 1886. Synopsis of the North American Hypocreaceae with descriptions of the species. J. Mycol. 2 : 61-69. & 1892. The Nodth American Pyrenomycetes. Publ. by authors, Newfield, New Jersey, Hanlin, R . T . 1961. Studies in the genus Nectria. II. Morphology of N. gliocladioides. Amer. J. Bot. 48: 900-908. 1963a. Morphology of Neuronectria peziza. Amer. J. Bot. 50: 56-66. 1963b. Morphology of Hypomyces lactifluorum. Bot. Gaz. (Crawfordsville) 124: 395-404. 1965. Morphology of Hypocrea schweinitzii. Amer. J. Bot. 52: 570-579. 1971. Morphology of Nectria haematococca. Amer. J. Bot. 58: 105-116. Luttrell, E . S . 1951. Taxonomy of the Pyrenomycetes. Univ. Missouri Stud. Sci. Set. 24: 1-120. Mathieson, M. J. 1952. Ascospore dimorphism and mating type in Chromocrea spinulosa (Fucked Perch n. comb. Ann. Bot. (London) 16: 449-466. Moravee, Z. 1956. Arachnocrea, un genre nouveau de la famille des Nectriaceae. Bull. Soc. Mycol. France "/2: 160-166. Peteh, T. 1937. Notes on British Hypocreaceae III. J. Bot. 75: 217-231. Ridgway, R. 1912. Color standards and color nomenclature. Published by Ridgway, Washington, D.C. Rogerson, C. T. 1970. The hypocrealean fungi (Ascomycetes, Hypocreales). Mycologia 62: 865-910. Samuels, G. J. 1973. Perithecial development in Hypomyces aurantius. Amer. J. Bot. 60: 268-276. 1976. A revision of the fungi formerly classified as Nectria subgenus Hyphonectria. Mere. New York Bot. Gard. 26(3): 1-126. Seaver, F . J . 1910. The Hypocreales of North America III. Mycologia 2: 48-92. pl. 20, 21. Vineens, F. 1917. Rech~rches organog6nique sur quelques Hypocr~ales. Ph.D. Theses, Fac. Sci. Paris.