From Japan I

Home , Bonin Islands, Podospora appendiculata

J. Gen. Appl. Microbiol., 18, 433-454 (1972)

COPROPHILOUS PYRENOMYCETES FROM JAPAN I

KOUHEI FURUYA AND SHUN-ICHI UDAGAWA*

Fermentation Research Laboratories, Sankyo Co., Ltd., Hiro-machi 1-chome, Shinagawa-ku, Tokyo 140 and *Department of Microbiology , National Institute of Hygienic Sciences, Kamiyoga 1-chome, Setagaya-ku, Tokyo 158

(Received July 13, 1972)

For the purpose of these series of mycological survey, 220 dung samples of wild and domestic animals for determination of species of pyrenomycetous Ascomycetes were collected from various geographic regions of Japan, including Ryukyu and Bonin Islands. Fifteen species of Podospora (the Sor- dariaceae) from numerous collections are described and illustrated. Most of them were also obtained in living cultures. All species are new records in Japan.

Generally speaking, animal dungs contain very rich nutritive components which may serve as growth factors for various types of microorganisms. The coprophilous fungi, one of such dung inhabitants, comprise a special group made up of members of several classes ranging through Myxo- mycetes to Basidiomycetes. In their pioneering studies, MASSEE and SAL- MON (1, 2) emphasized that 187 genera and 757 species from coprophilous occurrence had been listed in SACCARDO's Sylloge Fungorum in the early of the present century. These fungi have long attracted many mycologists for more than a hundred years, and a considerably large number of taxonomic papers have been published on them from various areas of the world. Investigations on this fascinating group of fungi were relatively few in Japan. The most extensive work is that of TUBAKI (3), who isolated 16 species belonging to the Hyphomycetes from dung sources in Japan. SONEDA (4, 5), in more recent reports, also listed 22 species of coprophilous yeasts, In two investigations resulted in the primary work of the Japanese Mucorales, some dung-inhabiting fungi were described by Indoh (6, 7). These contributions, however, provided only a little knowledge of their geographical distribution, especially on the occurrence of Japanese coprophilous Ascomycetes. Two recent studies on the discomycete flora, initiated by OTANI and KANZAWA (8, 9), have led to a suggestion that a majority of the coprophilous Ascomycetes found in Japan are common among other

433 434 FURUYA and UDAGAWA VOL. 18 countries, though they are little localized geographically. The present paper is the first attempt to give a survey of the Japanese members of detached group of Ascomycetes usually referred to as the coprophilous Pyrenomycetes.

MATERIALS AND METHODS

A total of 220 dung samples were collected from different localities in Japan from September 1966 to November 1971. These localities included Ryu- kyu Islands (now Okinawa Prefecture) and Bonin Islands (now Ogasawara Is- lands, Tokyo). The collections comprised of the droppings of following wild and domestic Mammalias : Brown bear (Ursus arctos), covey (Ochotona hyperborea), deer (Cervus nippon), fox (Vulpes vulpes), Japanese hare (Lepus brachyurus), marten (Martes melampus), monkey (Macaca fuscata), raccoon dog (Nyctereutes procyonoides), squirrel (Sciurus vulgaris and Tamias sibiricus), cattle, horse, goat, sheep, and others. The samples of dung collected in the field were held in moist chambers at room temperature, observing frequently for a period of one to four weeks or longer for development of marked fungi. Isolation was usually attempted on several media such as Leonian's yeast extract agar, oatmeal agar (with wheat germ powder), cellulose agar, potato-carrot agar and potato-dextrose agar. With some exceptions, the fungi mentioned in these series may be grown with great difficulty on the above laboratory media. In most cases, therefore, microscopical observations were carried out using living materials on the dung. Lactophenol preparations and dried specimens of all species have been placed in the Mycological Collection of the National Institute of Hygienic Sciences, Tokyo.

RESULTS AND DESCRIPTIONS

Among the total collection of 52 species of Pyrenomycetes examined, the order of occurrence for the chief genera was Podospora, Sporormiella, Conio- chaeta, Delitschia, Sordaria, and Chaetomium. The selected publications used for most of the identifications were GRIFFITHS (10), CAIN (11, 12), MOREAU (13), BOEDIJN (14), MIRZA and CAIN (15), ARMED and CAIN (16), and LUNDQVIST (17). Thus, it is now possible to report that 15 species of Podospora are known to occur in Japan. The remainder will be described in subsequent papers.

Podospora appendiculata (Auersw.) Niessl, in Hedwigia, 22, 156 (1883) ; Mirza and Cain, in Can. J. Bot., 47, 2007 (1969) ; Lundqvist, in Symb. Bot. Upsal., 20, 129 (1972). (Fig. 1) Sordaria appendiculata Auersw., 1872. For additional synonymy see MIRZA and CAIN (15) and LUNDQVIST (17). Perithecia superficial, scattered or aggregated in small clusters, ovate to 1972 Coprophilous Pyrenomycetes I 435

Fig. 1. P, appendiculata (Auersw.) Niessl. a, Perithecia ; b, Ascus ; c, Ascus gores.

somewhat conical, 700-1,000 x 380-800 p, dark brown to nearly black, sparingly hairy, neck small, papilliform to broadly conical, usually 50-150>< 120-160 p, roughened with black papillae. Hairs short, straight or slightly flexuous, septate, 30-150 p long, 4-5 p wide and brown at the base, with rounded or sometimes pointed, hyaline tip. Peridium fairly thick, coriaceous, dark brown and opaque, consisting of irregular cells, measuring 11.5-22.5 in diam. Asci 8-spored, clavate, 180-300 x 25-45 p, contracted and narrowly rounded above, with small thickened ring at the apex, tapering below into a slender stipe, measuring 70-100 p long ; paraphyses hyaline, thread-like, evanescent. Ascospores biseriate in central portion, hyaline and clavate when young, becoming dark brown and ellipsoid at maturity, 25-30 x 12.5-15 p, rounded above, somewhat truncate at the base, with apical germinal pore ; primary appendage cylindrical to conical, 12.5-19x4.5-6.5 p, hyaline, fugacious ; lower secondary appendage lash-like, hyaline, gelatinous, 25-60 p in length, attached 436 FURUYA and UDAGAWA VOL. 18

to the end of primary appendage ; upper one lash-like, 60-100 p in length, hollow at the base, symmetrically attached at the apex of the ascospore. Cultures on oatmeal agar growing slowly, with a thin mycelial felt, more or less funiculose with the development of a few aerial hyphae, perithecia tardily produced, grayish brown to dark brown ; reverse dark brownish gray. Habitat : On hare dung, Fujiyoshida-shi, Yamanashi-ken, April 26, 1967, NHL 22385, Kazikazawa-machi, Minamikoma-gun, Yamanashi-ken, Nov. 4, 1966, Kamikawa-machi, Kamikawa-gun, Hokkaido, July 28, 1967, Sanada-machi, ChIsagata-gun, Nagano-ken, Aug. 20, 1967, Ebino-machi, Nishimorokata-gun, Miyazaki-ken, April 19, 1971, NHL 22542; and on deer dung, Nara-shi, Nara ken, Oct. 16, 1966, NHL 22543. P. appendiculata is characterized by large ovate perithecia with pseudo bombardioid peridium that is somewhat suggestive of members of the genus Cercophora.

Podospora australis (Speg.) Niessl, in Hedwigia, 22, 156 (1883) ; Mirza and Cain, in Can. J. Bot., 47, 2009 (1969) ; Lundqvist, in Symb. Bot. Upsal., 20, 203 (1972). (Fig. 2) Hypocopra australis Speg., 1880. For additional synonymy see MIRZA and CAIN (15) and LUNDQVIST (17). Perithecia scattered, superficial or partially immersed, dark brown to nearly black, pyriform, 900-1,600 x 600-700 p, densely covered on the exposed part with long, septate, branched, olivaceous brown hairs measuring 2.5-3 p in diam. ; neck black, papilliform to cylindrical, 280-700x180-280 p, covered with short, straight, septate, hyaline-tipped hairs measuring up to 55 p long and 4 p wide at base, or almost bare. Peridium dark olivaceous brown,, membranaceous to rather coriaceous. Asci 4-spored, cylindrical, 300-380 x 40 -45 p , tapering below into a slender stipe measuring 80-150 p in length ; paraphyses filiform to ventricose. Ascospores uniseriate, dark brown and opaque, ellipsoid, broadly rounded at the both ends, (52-)56-60(-70) x (28-) 32-34(-40) p ; exospore 1.5-2 p thick ; primary appendage reduced to a small basal apiculum, hyaline, 3.5-4 p wide at the base ; lower secondary appendage gelatinous, long, attached to the basal end of spore, completely covering the apiculum, appearing hollow and taeniate ; upper secondary appendage similar, 50-80 (-100) x 8-15 p, attached eccentrically ; germinal pore apical. Habitat : On goat dung, Sanada-machi, Chisagata-gun, Nagano-ken, Oct. . 3, 1969, NHL 22548. This species is quite distinctive and can be identified by its large ascospores with hyaline, small, basal apiculum and two taeniate gelatinous appendages.

Podospora communis (Speg.) Niessl, in Hedwigia, 22, 156 (1883) ; Mirza and Cain, in Can. J. Bot., 47, 2012 (1969) ; Lundqvist, in Symb. Bot. Upsal., 1972 Coprophilous Pyrenomycetes I 437-

Fig. 2. P. australis (Speg.) Niessl. a, Perithecium ; b, Ascus ; c, Ascospores.

20, 171 (1972). (Fig. 3) Hypocopra communis Speg., 1880. For additional synonymy see MIRzA and CAIN (15) and LUNDQVIST (17). Perithecia scattered, semi-immersed to nearly superficial, olivaceous brown to dark brown, pyriform, 650-820x325-400 u, covered on the basal part with pale olive, long, flexuous, septate hairs measuring 1.5-2 t wide near the base, sometimes nearly glabrous ; neck cylindrical, 170-400 x 100-135 ~c, roughened with small, black papillae. Perithecial peridium thin, mem-- branaceous, semitransparent, consisting of yellowish olive, angular cells. Asci 8-spored, clavate, 180-220x36-40 p, narrowly rounded above, with no discharged structure visible in apex, gradually contracted below into a short stipe up to 40 in length ; paraphyses ventricose, indistinct. Ascospores biseriate, dark olivaceous brown and opaque, ellipsoid, 28-36(-40) x 18-22 t ; primary appendage hyaline, long, cylindrical, 25-32 x 4-6 ~c, tipped at the lower end with 3 or 4, short, divergent, gelatinous secondary appendages ; apex of the spore with 4 similar secondary appendages ; germinal pore apical, small, about 1.5 in diam. Cultures on oatmeal agar growing rapidly, thin, loosely floccose, dark

. greenish gray, conidial structures limited in number ; perithecia lacking; reverse greenish black. Mycelium pale olive brown, branched, septate, 2-2.5 438 FURUYA and UDAGAWA VOL. 18

Fig. 3. P. communis (Speg.) Niessl. a, Ascus ; b, Ascospores. p in diam. Conidial structures scattered on the aerial mycelium, consisting of simple phialides. Phialides peg-like, small, about 4 x 3 p. Conidia (phialospores) ovate to subglobose, hyaline, 2-3(-3.5) p, with smooth walls. At 37°, little growth. Habitat : On goat dung, Mukojima and Chichijima, Ogasawara Isls., Tokyo, July 10-21, 1969, NHL 22551, Manazuru-machi, Ashigarashimo-gun, Kanagawa-ken, Oct. 1, 1966; on cattle dung, Sanada-machi, Chisagata-gun, Nagano-ken, Oct. 3, 1969; on horse dung, Nakatane-machi, Kumage-gun, Kagoshima-ken, April 16, 1971, NHL 22519. Notable variability in the size range of ascospores and their primary appendage within various collections of this species has been observed and discussed by LUNDQVIsT (17).

Podospora curvicolla (Winter) Niessl, in Hedwigia, 22, 156 (1883) ; Mirza and Cain, in Can. J. Bot., 47, 2013 (1969) ; Lundqvist, in Symb. Bot. Upsal., 20, 184 (1972). (Fig. 4) Sordaria curvicolla Winter, 1871. For additional synonymy see CAIN (I1) and LUNDQYIST (17). Perithecia scattered, partly immersed to nearly superficial, dark olivaceous brown to dark brown or nearly black, pyriform, 370-900 x 160-430 p, glabrous or covered sparingly with long, flexuous, septate, pale colored hairs measur- 1972 Coprophilous Pyrenomycetes I 439

Fig. 4. a-b. P. curvicolla (Winter) Niessl. a, Perithecium b, Ascospores. c-e. P. longicollis (Ames) Mirza et Cain. c, Perithecium ; d, Ascus ; e, Asco- spores.

ing 1.5-2 p in diam. ; neck long cylindrical, 200-700 x 65-95(-200) p, roughened with small black papillae, glabrous or ornamented with few tufts of long, septate, nearly straight hairs measuring 120-185 x 2 p. Perithecial peridium thin, membranaceous, semitransparent, consisting of pale olivaceous brown, angular cells, 5-10 p or more in diam. Asci 256-spored, broadly clavate, 120-200 x 70-100 p, contracted below into a short stipe, very deliquescent ; apical dehiscent structure not distinct. Paraphyses hyaline, tubular. Asco- spores multiseriate, olivaceous brown to dark brown, opaque, ellipsoid, 13- 17.5 x 8-12.5 p ; primary appendage hyaline, short clavate, 5-8 x 2.5-3 p, tipped with a long lash-like gelatinous secondary appendage, very fugacious ; upper secondary appendage similar, attached eccentrically to apex of the spore ; germinal pore apical, small, 1-2 p in diam. Cultures on potato-carrot agar spreading broadly, thin, vegetative mycelium mostly submerged, with few aerial hyphae appearing somewhat floccose, grayish olive green, perithecia abundantly produced, mostly buried in the substratum, olive black in color ; conidial structures not observed. At 37°, only very slight growth. Habitat : On deer dung, Ebino-machi, Nishimorokata-gun, Miyazaki-ken, 440 FURUYA and UDAGAWA VOL. 18

Fig. 5. P. curvula (De Bary) Niessl. a, Perithecium ; b, Cells of agglutinated hairs; c, Ascus ; d, Ascospores ; e, Conidial structures.

April 19, 1971, NHL 22522; on hare dung, Sobetsu-machi, Usu-gun, Hokkaido, Aug. 11, 1967, NHL 22553. In producing tufts of perithecial hairs, it is most likely to be confused with P, longicollis but the 256-spored asci and smaller ascospores are different.

Podospora curvula (De Bary) Niessl, in Hedwigia, 22, 156 (1883) ; Mirza and Cain, in Can. J. Bot., 47, 2014 (1969) ; Lundqvist, in Symb. Bot. Upsal., 20, 253 (1972) as Schizothecium conicum (Fuck.) Lundq. (Fig. 5) Sordaria curvula De Bary,1866. For additional synonymy see CAIN (11), MIRZA and CAIN (15), and LUNDQVIST (17). Perithecia scattered, superficial, dark olivaceous brown to dark brown, elongate to ovate or oblong, 500-1,265 x 275-500 I, covered on the basal part with clusters of agglutinated, pale yellowish brown, obliquely septate hairs consisting of swollen cells ; tufts up to 60 tc long ; neck nearly black, conical to short cylindrical, 200-400 x 130-220 fit, glabrous. Peridium of perithecium thin, membranaceous, semitransparent, consisting of rather large angular cells. Asci 8-spored, clavate, 140-190 x 24-32(-34) ,t, contracted above to narrowly rounded apex, with small indistinct ring-like structure, tapering below into a short stipe measuring 20-35 It in length, surrounded by hyaline paraphyses, at maturity deliquescent. Ascospores irregularly biseriate, sometimes uni- 1972 Coprophilous Pyrenomycetes I 441

senate, dark olivaceous brown to dark brown, opaque, ellipsoid, 25-28(-32 .5) x 14-18(-20) ii ; primary appendage cylindrical, hyaline, 8-11.5 x 2-2.5 p, attached to the base of ascospore ; lower secondary appendage indistinct, fugacious, lash-like, attached to the end of primary appendage ; upper secondary appendage hyaline, lash-like, eccentrically attached to the apex of ascospore, up to 30 x 2.5-4 tat the base ; a similar but shorter secondary appendage indistinct- ly produced near the base of the primary appendage ; germinal pore apical, 1.5-2 p in diam. Cultures on oatmeal agar growing rapidly, plane, lightly floccose, consisting of a rather thin mycelial felt, dark olive green, perithecia produced in limited numbers, buried in the mycelial felt ; reverse dark green. Mycelium hyaline to pale olivaceous brown, branched, septate, 1.5-3.5 in diam., often aggregated in fascicles. Conidial structures of phialospore-type. Phialides borne singly along aerial hyphae, sessile, hyaline, typically flask-shaped, 6-8 x 2.5-3 ii, with a distinct collarette at the tip. Conidia hyaline, subglobose or ovate, 2-2.5 tc in diam., smooth-walled. At 37°, growth is nil. Habitat : On hare dung, Otaru-shi, Hokkaido, July 1970, NHL 22518, Nayoro-shi, Hokkaido, Aug. 2, 1971, SSbetsu-machi, Usu-gun, Hokkaido, Aug. 11, 1967; on goat dung, Sapporo-shi, Hokkaido, July 4, 1970, NHL 22556; on sheep dung, Sapporo-shi, Hokkaido, July 26, 1967, NHL 22554. To accommodate a group of Podospora species characterized by the production of perithecia with tufts of agglutinated, inflated hairs and the presence of two-celled ascospores at a very early stage in their development, LUNDQVIST (17) used the generic name Schizothecium Corda in a limited sense. There are about 31 known species, namely "P. curvula group." However, the generic concept of MIRZA and CAIN (15) was followed in the present work, because the above segregation is not yet sufficiently well attempted to provide a preferable alternative name for the remainder Podo- spora outside Nordic. In addition, LUNDQVIST (17) did not mention the presence or absence of swollen peridial cells and hair tufts in his generic classification of related fungi such as Arnium or Cercophora, and possibly this has given rise to some doubts as to rearrangement within the genus Podo- spora.

Podospora dakotensis (Gritriths) Mirza et Cain, in Can. J. Bot., 47, 2016 (1969) ; Lundqvist, in Symb. Bot. Upsal., 20, 254 (1972) as Schizothecium dakotense (Gruff.) Lundq. (Fig. 6) Pleurage dakotensis Gruff., 1901; Philocopra dakotensis (Gruff.) Sacc.,1905. Perithecia scattered, nearly superficial, olivaceous brown, elongate-pyriform, 550-850 x 220-285 p, with the neck short, black, papilliform to somewhat elongate, 100-320 x 100-160 p, glabrous. Peridium of perithecium thin, membranaceous, olivaceous, semitransparent, upper part covered with clusters of agglutinated, subhyaline to pale brown hairs up to 50 p in length ; cells of 442 FURUYA and UDAGAWA VOL. 18

Fig. 6. P. dakotensis (Gruff.) Mirza et Cain. a, Perithecium ; b, Cells of agglutinated hairs; c, Ascus ; d, Ascospores. hairs thin-walled, swollen, rounded or irregularly angular. Asci 32-sporedy clavate, 200-240 x 36-45 p, broadly rounded above, with no distinct dehiscent structure visible in apex, contracted below into a short stipe up to 45-80 p in length ; paraphyses hyaline, irregularly swollen. Ascospores multiseriate,, dark olivaceous brown and opaque, ellipsoid, 19-23 x 12--1511; primary appendage short, cylindrical, 5-10 x 1-1.5 p ; upper secondary appendage long, lash- like, hyaline, gelatinous, 20-40 x 3 p, eccentric at apex of spore ; lower secondary appendage similar, attached to extremity of primary appendage ; germinal pore apical, 1.5 p in diam. Cultures on V-8 juice agar growing rapidly, dark olive gray, floccose,, consisting of a rather thick mycelial felt, black perithecia abundantly produced on the felt, commonly obscured by the overlying aerial growth ; conidial structures not observed. At 37°, growth slight with no perithecium production. Habitat : On hare dung, Ebino-machi, Nishimorokata-gun, Miyazaki-ken,, April 19, 1971, NHL 22560, Kirishima-cho, Aira-gun, Kagoshima-ken, Apr. 18, 1971, NHL 22562, Kochi-shi, Kochi-ken, March 1971. In some respects this species resembles P, glutinans but the asci are 32 spored.

Podospora decipiens (Winter ex Fuckel) Niessl, in H&wigia, 22, 156 (1883) ; Mirza and Cain, in Can, J. Bot. 47 2017 (1969) ; Lundqvist, in Symb. Bot. 1972 Coprophilous Pyrenomycetes I 443

Fig. 7. P. decipiens (Winter ex Fuckel) Niessl. a, Perithecium ; b, Ascus ; c, Ascospores.

Upsal., 20, 191 (1972). (Fig. 7) Sordaria decipiens Winter, 1873. For additional synonymy see CAIN (11), MIRZA and CAIN (15), and LUNDQVIST (17). Perithecia scattered, semi-immersed, olivaceous brown to dark brown, pyriform to elongate-pyriform, 650-950 x 250-470 fit, almost glabrous or covered on the basal part with a few, slender, flexuous, septate, pale brown, hyphal- like hairs measuring 1.5-2.5 u in diam. ; neck conical to cylindrical, 280-450 x 115-160 u, roughened above with black papillae. Peridium of perithecium. thin, pale yellowish olive to brown, semitransparent, membranaceous, consisting of rather, large, angular cells measuring 6-14 in diam. Asci 8-spored, clavate, 175-240 x 40-48 p, contracted and rounded above, with small ring like thickening in apex, tapering below into a short stipe ; paraphyses hyaline, ventricose. Ascospores biseriate, dark brown and opaque, elliptical, 34-44 x 18-22 ,cc, rounded at the ends ; primary appendage hyaline, cylindrical, 36-40 x 7-8 p, surrounded at the base by several short gelatinous secondary appendages ; apex of the spore with a lyre-shaped tuft of secondary appendages measuring 12-28 x 12-16 p ; germinal pore circular, about 2 e in diam., below apex of the spore, eccentric. Cultures on oatmeal agar spreading, floccose, dark yellowish brown, conidial structures and perithecia not produced ; reverse yellowish brown. At 37°, growing as at 25°, but dark olive gray in color. Habitat : On sheep dung, Sapporo-shi, Hokkaido, July 26, 1967, NHL 444 FURUYA and UDAGAWA VOL. 18

Fig. 8. P. fimbriata (Bayer) Cain. a, Perithecium ; b, Cells of agglutinated hairs; c, Ascus ; d, Ascospores.

22563; on cattle dung, Sanada-machi, Chisagata-gun, Nagano-ken, Oct. 3, 1969, NHL 22512; on hare dung, Sanada-machi, Chisagata-gun, Nagano-ken, Oct. 3, 1969, NHL 22564. In both the perithecia and the ascospores, this species is regarded as closely related to P. pleiospora, but separable from it primarily upon the basis of its smaller 8-spored asci and more larger ascospores.

Podospora fimbriata (Bayer) Cain, in Can. J. Bot., 40, 459 (1962) ; Mirza and Cain, in Can. J. Bot., 47, 2020 (1969). (Fig. 8) Sordaria fimbriata Bayer, 1924;? Sordaria vratislaviensis A. Schmidt, 1912; Bombardia lunata Zickler, 1934; Pleurage fimbriata (Bayer) Page, 1957. Perithecia superficial, scattered, dark olivaceous brown, ovoid to somewhat elongate, 350-550 x 200-240 p, covered on upper part with light brown, agglutinated hairs up to 60 p in length ; neck small, conical, up to 80 p in length. Distal cells of hairs usually fimbriate. Peridium semitransparent, membranaceous. Asci 8-spored, cylindrical, 110-150 x 9-10 p, narrowly rounded above, with an indistinct ring-like thickening in apex, tapering below into a stipe up to 45 p or more in length ; paraphyses thread-like, septate. Ascospores uniseriate, dark olive brown, inequilaterally ellipsoid, flattened on one side, 18- 20 x 8-9.5 p, more or less apiculate at both ends ; primary appendage hyaline, =clavate , 4-6 x 2 p, attached to base of spore ; secondary appendages not observed ; germinal pore apical, about 1 p in diam. 1972 Coprophilous Pyrenomycetes I 445

Mycelium yellowish brown, branched, 1-4 tc in diam., septate. Conidial structures abundantly produced, simple, of phialospore-type. Phialides cylindrical, 8-12 x 2.5-3 ~i, borne singly on the aerial hyphae, with a distinct funnel-like collarette measuring 1.5-2 i in length and 1.5 p in width. Conidia (phialospores) hyaline, subglobose to ovate, 1.5-3 ~i in long axis, smooth- walled. Cultures on oatmeal agar growing rapidly, floccose, olive black to grayish yellow brown, conidial structures abundantly produced, perithecia late developing ; reverse olive black. At 37°, growth is nil. Habitat : On goat dung, Kagoshima-shi, Kagoshima-ken, April 15, 1971, NHL 22514. P, fimbriata is easily recognized by its agglutinated hairs of the perithecium and its inequilaterally ellipsoid ascospores, from which the development of secondary appendages have not been observed.

Podospora fimiseda (Ces. et De Not.) Niessl, in Hedwigia, 22, 156 (1883) ; Mirza and Cain, in Can. J. Bot., 47, 2021 (1969) as P. fimicola Ces, in Rabenh. ; Lundqvist, in Symb. Bot. Upsal., 20, 135 (1972). (Fig. 9) Sordaria fimiseda Ces. et De Not., 1863. For additional synonymy see CAIN (11) and LUNDQVIST (17). Perithecia scattered, superficial, dark brown, pyriform, 900-1,400 x 500- 750 jc, covered on upper part (including neck) of the perithecium with separate, brown, straight, septate, cylindrical hairs measuring 30-80 x 3-4 and with a rounded, hyaline tip ; neck black, conical to subcylindrical, 400-500 x 200-250 p. Peridium brown, semitransparent, membranaceous. Asci 8-spored, clavate, 380-470 x 55-75 p, with a small indistinct apical ring ; stipe up to 160 p in length. Paraphyses abundant, filiform, septate. Ascospores biseriate, dark olivaceous brown, ellipsoid, 50-60 x 28-32 p, with a germinal pore at the apex measuring 4-5 p in diam. ; epispore thick ; primary appendage cylindrical- clavate, 36-48 x 5-6 p at base, swollen at the extremity up to 8-12 p in diam. ; lower secondary appendage lash-like, at the proximal end of primary appendage, 100-160 x 8-10 p ; upper secondary appendage attached eccentrically to the apex of ascospore, 120-180 x 16-18 p at base. Habitat : On cattle dung, Sanada-machi, Chisagata-gun, Nagano-ken, Oct. 3, 1969, NHL 22513. The characteristic perithecial hairs and large ascospores readily distinguish this species, which appears to be commonly associated with dung of domestic herbivorous animals.

Podospora glutinans (Cain) Cain, in Can. J. Bot., 40, 460 (1962) ; Mirza and Cain, in Can. J. Bot., 47, 2023 (1969) ; Lundqvist, in Symb. Bot. Upsal., 20, 254 (1972) as Schizothecium glutinans (Cain) Lundq. (Fig. 10) Sordaria glutinans Cain, 1934. 446 FURUYA and UDAGAWA VOL. 18

Fig. 9. P. fimiseda (Ces. et De Not.) Niessl. a, Perithecium ; b, Hairs; C, Ascus ; d, Ascospores.

Perithecia scattered, superficial, olivaceous brown to brown, pyriform, 560--850 x 300-400 p, covered on the basal part with long, flexuous, pale yellowish brown, sparingly septate hairs measuring 2-2.5 p wide near the base,, upper part covered with clusters of short, hyaline, agglutinated hairs about 40 p in length. Neck of perithecium papilliform to short cylindrical, 120-200 or more x 95-140 p, glabrous or roughened with black papillae. Perithecial peridium pale yellowish brown, membranaceous to slightly coriaceous, semitransparent. Asci 8-spored, cylindrical, 160-230 x 24--28 p, rounded above, no apical structure visible, tapering below into a short stipe up to 40 p in length ; distinct paraphyses absent. Ascospores obliquely uniseriate, olivaceous to dark brown and opaque, broadly ellipsoid, 28-30 X 18-22 p, rounded at both ends, with a short, hyaline, cylindrical primary appendage measuring 6-12 X 1.5-2 p ; secondary appendages hyaline, long, lash-like, one at the apex of the spore and one at the end of the primary appendage ; germinal pore apical, 2-3.5 in diam. 1972 Coprophilous Pyrenomycetes I 447

Fig. 10. P. glutinans (Cain) Cain. a, Perithecium b, Cells of agglutinated hairs ; c-d, Apical portion of ascus and ascospores.

Habitat : On hare dung, Sanada-machi, Chisagata-gun, Nagano-ken, Sept. 21, 1966, NHL 22565; on goat dung, Manazuru-machi, Ashigarashimo-gun, Kanagawa-ken, Oct. 1, 1966, NHL 22566. P, glutinans is distinguished from P. dakotensis by the formation of 8- spored asci and larger ascospores.

Podospora longicollis (Ames) Mirza et Cain, in Can. J. Bot., 47, 2028 (1969) (Fig. 4) Schizothecium longicolle Ames, 1951; Philocopra setosa (winter) Sacc. var. longicolle Marchal, 1884; Pleurage longicollis (Ames) Boedijn, 1962. Perithecia scattered, immersed to partly immersed, dark olivaceous brown to nearly black, pyriform, 500-950 x 285-520 p, glabrous or loosely covered on the exposed part with almost hyaline, septate, flexuous hyphae measuring 1.5-2 p in diam. ; neck dark olive and opaque, short cylindrical to cylindrical, 145-380 x 105-210 p, bearing a few tufts of long, olivaceous brown, septate hairs on the basal part, hairs up to 300-400 p long and 1.5-2.5 p wide at the base. Perithecial peridium thin, membranaceous, semitransparent. Asci multi- spored (about 512-spored), broadly clavate, 220-320 or more x95-145 p, with broadly rounded apex, ring-like structure not seen, contracted below into a short stipe measuring about 30 p in length, surrounded by hyaline, ventricose paraphyses, at maturity soon deliquescent. Ascospores in several series, ellipsoid, ranging from hyaline when young through olivaceous to dark olivaceous brown and opaque, 16-22 x 9-12 p ; primary appendage clavate, hyaline, 6-10 x 2.5-3 p ; secondary appendages lash-like, hyaline, respectively attached to the 448 FURUYA and UDAGAWA VOL. 18

apex of the spore-body and the end of the primary appendage, soon disappear- ing ; germinal pore circular, 1-1.5 in diam., apical. Cultures on potato-carrot agar growing rapidly, more or less cottony, grayish olive to dark olive gray, perithecia abundantly produced on a mycelial felt, overgrown by a loose network of aerial hyphae (excepting neck region) ; reverse dark olive to olive black. Mycelium pale olive, septate, branched, 1.5-4 j in diam. ; asexual state unknown. At 37°, growth is nil. Habitat : On goat dung, Mukojima, Ogasahara Isls., Tokyo, July 21, 1969, NHL 22499; on deer dung, Nishiumi-machi, Minamiuwa-gun, Ehime-ken, May 3, 1967, NHL 22567; on hare dung, Ebino-machi, Nishimorokata-gun, Miya- zaki-ken, Apr. 19, 1971, NHL 22498; Fujiyoshida-shi, Yamanashi-ken, Apr. 26, 1967, Sanada-machi, Chisagata-gun, Nagano-ken, Aug. 20, 1967, Kiyokawa- mura, Aiko-gun, Kanagawa-ken, Mar. 22, 1969, Kochi-shi, Kochi-ken, March 1971. P. longicollis is the most common species of the genus in Japan. It somewhat resembles P, setosa but differs in having tufted hairs of the perithecium and 512-spored asci.

Podospora myriaspora (Cr. et Cr.) Niessl, in Hedwigia, 22, 156 (1883) ; Lundqvist, in Symb. Bot. Upsal., 20, 200 (1972). (Fig. 11) Sordaria myriaspora Cr, et Cr., 1867; Philocopra myriaspora (Cr, et Cr.) Sacc., 1882; Pleurage myriaspora ("myriospora") (Cr. et Cr.) 0. Kuntze, 1898. Perithecia scattered, more or less semi-immersed, dark olivaceous brown, pyriform, 940-1,150 x 500-650 p, covered on the upper part with small tuber- cles and slender flexuous hairs ; neck conical to elongate, 300-650 x 180-300 p, roughened with black papillae. Hairs brown, septate, 2-2.5 p wide. Peridium thin, somewhat semitransparent, rather coriaceous. Asci mostly 64-spored (sometimes 32-spored), clavate to broadly clavate, 320-360 x 68-100 p, with small inconspicuous ring-like structure in apex, contracted below into a short stipe up to 100 p in length, at maturity deliquescent ; paraphyses hyaline, ventricose. Ascospores in several series, dark olivaceous brown and opaque, ellipsoid, (24-)27-32 x (15-)16-19 p ; primary appendage hyaline, cylindrical, 26-30(-50) x 5-6 p, surrounded at the base with 2 to 4 small secondary appendages ; upper secondary appendages agglutinated to form an apical tuft measuring 18-30(-45) x4-5 p ; germinal pore circular, 1.5-2 p in diam., eccentric near upper end of ascospore. Cultures on oatmeal agar growing rapidly, thin, floccose with few aerial growth, dull yellowish orange to olive, perithecia late produced ; mycelium hyaline, branched, septate, 1.5-4 p in diam., often aggregated in bundles ; conidial structures not seen. At 37°, little growth with no production of perithecia. Habitat : On goat dung, Sapporo-shi, Hokkaido, July 4, 1970, NHL 22496; on deer dung, Nara-shi, Nara-ken, Oct. 16, 1966, NHL 22572; on horse dung, 1972 Coprophilous Pyrenomycetes I 449

Fig. 11. P. myriaspora (Cr. et Cr.) Niessl. a, Perithecium ; b, Ascus and its apical portion ; c, Ascospores.

Nakatane-machi, Kumage-gun, Kagoshima-ken, Apr. 16, 1971, NHL 22576. P. myriaspora is distinguished from P, pleiospora (Winter) Niessl by the 64-spored asci and the smaller ascospores with slightly narrower primary appendage. In all other characters it is similar to the latter.

Podospora pauciseta (Ces.) Traverso, in Fl. Ital. Crypt. 1. Fungi 1, 431 (1907) ; Mirza and Cain, in Can. J. Bot., 47, 2005 (1969) as P. anserina (Ces. in Rabenh.) Niessl ; Lundqvist, in Symb. Bot. Upsal., 20, 147 (1972). (Fig. 12) Sphaeria pauciseta Ces. in Rabenh., 1852. For additional synonymy see MIRZA and CAIN (15) and LUNDQVIST (17). Perithecia scattered, nearly superficial, olivaceous brown to black, pyriform,. 500-700 x 250-350 i ; neck blackened with papillae, conical to subcylindrical, 180-300 x 120-160 u, often hairy at the base of neck ; hairs brown, straight, obscurely septate, up to 200-300 x 3-4 p at base, mostly adhering in fascicles forming a few tufts. Peridium brown, thin, membranaceous. Asci 4-spored, cylindrical to somewhat clavate, 160-200 x 22-28 p, with a short stipe up to 60 p in length ; apical structure not distinct ; paraphyses filiform to ventricose, septate. Ascospores obliquely uniseriate, ellipsoid, 34-38 x 20-22 p, becoming dark olivaceous brown and opaque ; primary appendage basal, cylindrical, 20- 24 x 4-5 p ; with two long secondary appendages lash-like, evanescent, one at the distal end of the primary appendage and the other at the apex of the 450 FURUYA and UDAGAWA VOL. 18

Fig. 12. P. Pauciseta (Ces.) Traverso. a, Perithecium ; b, Tufts of hairs; c, Ascus ; d, Ascospores. spores, the latter attached eccentrically and hollow at the base ; germinal pore apical, about 2 in diam. Cultures on potato-carrot agar spreading rapidly, floccose, dark gray, perithecia and conidial structures fairly abundantly produced ; reverse bluish black. Mycelium hyaline, branched, 2-8 in diam., septate. Conidial structures simple, of phialospore-type. Phialides small, peg-like to somewhat cylindrical, 2-6 x 1.5-2.5 i, with a collarette. Conidia (phialospores) hyaline, subglobose to ovate, 2-3 x 1.5-2.5 i, smooth-walled, accumulating at the collarette in a slime head. At 37°, growth is luxuriant but no perithecia produced. Habitat : On cattle dung, Sanada-machi, Chisagata-gun, Nagano-ken, Oct. 3, 1969, NHL 22574; on horse dung, Nakatane-machi, Kumage-gun, Kago- shima-ken, Apr. 16, 1971, NHL 22515. For this fungus, the name P, anserina is more widely known in the world but LUNDQVIST(17) adopted Traverso's treatment with detailed discussion of nomenclature.

Podospora setosa (Winter) Niessl, in Hedwigia, 22, 156 (1883); Mirza and Cain, in Can. J. Bot., 47, 2040 (1969); Lundqvist, in Symb. Bot. UPsal., 20, 178 (1972). (Fig. 13) Sordaria setosa Winter, 1873; Podospora tarvisina (Sacs.) Mirza et Cain, 1969. For additional synonymy see CAIN (11), MIRZA and CAIN (15), and 1972 Coprophilous Pyrenomycetes I 451

Fig. 13. P. setosa (Winter) Niessl. a, Ascus ; b, Ascospores.

LUNDQVIST (17). Perithecia scattered or densely aggregated, partially immersed to immersed, subspherical to pyriform, 700-950 x 450-650 p, base semitransparent at first, then becoming olivaceous to dark brown, often covered with light brownish hyphae, neck black, papilliform to cylindrical and somewhat curved, roughened with small papillae, 130-325 x 145-220 p, frequently provided with unevenly scattered, straight, continuous, smooth-walled, brown hairs measuring up to 160-200(-400) p long and (3-)4-5 p wide at the base, fading and tapering to rounded or collapsed tip. Peridium brown to dark brown, thin, membranaceous to coriaceous, consisting of irregular, pale olive brown cells measuring 4-15 p in diam. Asci 128-spored, fusiform to clavate, 240-320 x 40-60(-95) p, contracted and rounded above, each with a distinct apical thickening visible when young, gradually tapering below into a slender stipe, measuring up to 95 p long ; paraphyses filiform to ventricose, hyaline and septate. Ascospores in many series, forming an ellipsoidal mass, at first hyaline and clavate, then becoming olivaceous brown to dark brown and opaque, ellipsoid, 17.5-19(-22) x 11.5-14 p, rounded at the apex, slightly truncate at the base, with an apical germinal pore, measuring about 1.5-2 p in diam. Primary appendage cylindrical, hyaline, 8-12 x 2-3 p, attached to the lower end of spore. Secondary appendages very evanescent, gelatinous, lash-like, longer than spores, one attached eccentrically to the apex of spores and one to the end of the primary appendage. Cultures on potato-dextrose agar spreading broadly, consisting of a corn- 452 FURUYAand UDAGAWA VOL. 1$ pact, fairly thick felt, floccose, light gray to gray, perithecia slowly developing within a mycelial matrix ; reverse dark olive gray to nearly black. At 37°, poor growth. Habitat : On pheasant (Phasianus colchicus) dung, Kajikazawa-machi,. Minamikoma-gun, Yamanashi-ken, Nov. 4, 1966, NHL 22379 ; on deer dung,. Nara-shi, Nara-ken, Oct. 6, 1967, NHL 22384, Kyokawa-mura, Aiko-gun,. Kanagawa-ken, Oct. 1, 1970 ; on goat dung, Manazuru-machi, Ashigarashimo gun, Kanagawa-ken, Oct. 1, 1966, NHL 22579 ; on hare dung, Fujiyoshida-shi, Yamanashi-ken, Apr. 26, 1967, NHL 22578 ; Narugo-machi, Tamatsukuri-gun, Miyagi-ken, Nov. 8, 1967, NHL 22582, Sanada-machi, Chisagata-gun, Nagano ken, Sept. 21, 1966, Ito-shi, Shizuoka-ken, July 9, 1967, Sanda-shi, Hyogo-ken, Oct. 15, 1967. This species is one of the most common and widely distributed Podospora in Japan and was included in a complex of P, curvicolla from which it is readily distinguished by the character of perithecial hairs and somewhat larger ascospores.

Podospora tetraspora (Winter) Cain, in Can. J. Bot., 40, 460 (1962) ; Mirzaa and Cain, in Can. J. Bot., 47, 2042 (1969) ; Lundqvist, in Symb. Bot. Upsal., 20, 256 (1972) as Schizothecium tetrasporum (Wint.) Lundq. (Fig. 14) Sordaria tetraspora Winter, 1871. For additional synonymy see CAIN (11). Perithecia superficial, scattered, elongate-ovate or conical to pyriform, 450-700 x 200-300 p, light olivaceous below, black above, covered around base of neck with small clusters of agglutinated cells measuring about 7-10 x 5-7 p, neck small, papilliform or conical, black, 100-200 or more x 100-180 p. Perithecial peridium light olivaceous brown, thin, semitransparent, membrana ceous, consisting of irregular, rounded or elongate to angular, swollen cells measuring 7.5-16.5 x 6.5-14 p. Asci 4-spored (rarely 1- or 2-spored), cylindrical, 120-150 x 14-16 p, slightly narrowed above, rounded and with a distinct ring-like thickening in apex, tapering below into a slender stipe 50-70 p long paraphyses hyaline, varying from cylindrical to vesicular cells, soon deliquescent. Ascospores obliquely uniseriate, dark brown, ellipsoid, 17.5-22.5(-24) x 11.5-12.5(-15) p, acutely rounded at apex, truncate at base, with a circular apical germinal pore, measuring 1.5 p in diam. ; primary appendage hyaline, cylindrical, 10-17.5 x 2 p, fugacious ; secondary appendage lash-like, hyaline, gelatinous, eccentric at the apex of the spore ; lower secondary appendage similar, but attached at the end of the primary appendage. Cultures on potato-carrot agar growing rapidly, with abundant conidial structures, floccose, olive gray to brownish gray, late developing a few perithecia at the central area ; reverse dark olive gray to olive black. Mycelium hyaline to pale olivaceous brown, septate, branched, 1.5-4 p in diam., often aggregated in fascicles. Conidial structures consisting of single phialides borne 1972 Coprophilous Pyrenomycetes I 453

Fig. 14. P. tetraspora (Winter) Cain. a, Perithecium ; b, Cells of agglutinated hairs; c, Asci d, Ascospores ; e, Conidial structures. along the aerial hyphae, simple. Phialides flask-shaped with a swollen base, 6-10(-16) x 2.5-3 p, gradually tapering to a tip, with a distinct collarette. Conidia almost hyaline, globose to subglobose, 2-2.5 x 1.5-2 p, smooth-walled, accumulating in a small head at the apex of the phialide. At 37°, growth is nil. Habitat : On deer dung, Ebino-machi, Nishimorokata-gun, Miyazaki-ken, Apr. 19, 1971, NHL 22521, Kamikawa-machi, Kamikawa-gun, Hokkaido, Jan. 26, 1968; on hare dung, Fujiyoshida-shi, Yamanashi-ken, Apr. 26, 1967, NHL 22386, Sanada-machi, Chisagata-gun, Nagano-ken, Sept. 21, 1966, Ita-shi, Shizuoka-ken, July 9, 1967, NHL 22586, Ebetsu-shi, Hokkaido, July 25, 1967, NHL 22587, Kirishima-cho, Aira-gun, Kagoshima-ken, Apr. 18, 1971. This species, marked by elegant perithecia with numerous agglutinated hairs and 4-spored asci, is especially common and world-wide.

We wish to express our gratitude to Dr. H. Kurata for his continued interest and support. Thanks are also due to the numerous persons who assisted us in making field collections.

REFERENCES 1) G. MASSEEand ES. SALMON, Ann. Bot., 15, 313 (1901). 454 FURUYA and UDAGAWA VOL. 18

2) G. MASSEE and ES. SALMON, Ann. Bot., 16, 57 (1902). 3) K. TUBAKI, Nagaoa, 4, 1 (1954). 4) M. SONEDA, Nagaoa, 6, 1 (1959). 5) M. SONEDA, Trans. Mycol. Soc. Japan, 3, 36 (1962). 6) H. INDOH, Trans. Mycol. Soc. Japan, 3, 24 (1962). 7) H. INDOH, Trans. Mycol. Soc. Japan, 5, 60 (1965). 8) Y. OTANI and S. KANZAWA, Trans. Mycol. Soc. Japan, 10, 117 (1970). 9) Y. OTANI and S. KANZAWA, Trans. Mycol. Soc. Japan, 11, 43 (1970). 10) D. GRIFFITHS, Mem. Torrey Bot. Club, 11, 1 (1901). 11) R.F. CAIN, Univ. Toronto Stud. Biol. Ser., No. 38, 1 (1934). 12) R.F. CAIN, Can. J. Bot., 40, 447 (1962). 13) C. MOREAU, Encycl. Mycol., 25, 1 (1953). 14) K.B. BOEDIJN, Persoonia, 2, 305 (1962). 15) J.H. MIRZA and R.F. CAIN, Can. J. Bot., 47, 1999 (1969). 16) S.I. AHMED and R.F. CAIN, Can. J. Bot., 50, 419 (1972). 17) N. LUNDVIST, Symb. Bot. Upsal., 20, 1 (1972).