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Institute of Botany, Academia Sinica, Nankang, Taipei 115, Taiwan
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ABSTRACT |
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Penicilliopsis pseudocordyceps was collected from seeds of Diospyros discolor and is described as new. The anamorph produced in culture is Pseudocordyceps seminicola. The teleomorph was produced on oatmeal agar in 6 wk. Penicilliopsis clavariaeformis was also collected from the same seeds, yielding its Sarophorum palmicola anamorph in culture.
Key words: Eurotiales, Sarophorum palmicola, Trichocomaceae
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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who accepted only two species, i.e., P. clavariaeformis Solms-Laubach and P. africana Samson & Seifert. Samson and Seifert (1985)
also discussed four synnematous genera—Pseudocordyceps Hauman, Sarophorum Syd. & P. Syd., Stilbodendron Syd. & P. Syd., and Stilbothamnium Henn.—that were positively or circumstantially connected to species of Penicilliopsis. Sarophorum and Stilbodendron were the anamorphs of P. clavariaeformis and P. africana, respectively. Stilbothamnium had been observed to be associated with immature Penicilliopsis-like stromata. Pseudocordyceps was the only form-genus in the discussion that lacked evidence of a teleomorphic connection.
Pseudocordyceps, Sarophorum, and Stilbodendron have conidiogenous structures similar to those of Penicillium Link : Fr. but arranged on large synnemata. Despite their differences in synnematal gross morphology, i.e., acicular in Sarophorum, capitate in Pseudocordyceps, and tentaculate in Stilbodendron, a case could be made to lump these three genera if one intended to apply one generic name to the anamorphic Penicilliopsis in order to comply with the genus-for-genus concept. However, Stilbothamnium, which may have a Penicilliopsis connection, has an Aspergillus-like conidiogenous structure and, in fact, was treated as a subgenus of Aspergillus P. Mich. ex Link : Fr. by Samson and Seifert (1985)
. We thus feel that a formal decision should not be made until the teleomorph of Stilbothamnium is available for study.
Two synnematous fungi, Pseudocordyceps seminicola Hauman and Sarophorum palmicola (Henn.) Seifert & Samson, were collected from seeds of Diospyros discolor Willd. from southern Taiwan in September, 2000. On the seeds, two types of immature stromata were also found: one capitate and the other hemispherical. However, their connections with the two synnematous fungi had not been clear to us until the collecting site was revisited two months later and the mature stromata of two Penicilliopsis species were collected: the capitate ones belonging to P. clavariaeformis and the hemispherical ones to an undescribed species. Cultures obtained from ascospores of the undescribed Penicilliopsis yielded Pseudocordyceps seminicola, the generic name of which is used as the epithet for the holomorph described herein. Culturing of P. clavariaeformis confirmed that S. palmicola represents its anamorph; the teleomorph-anamorph connection reported previously was based on the occurrence of both morphs on the same seeds (Solms-Laubach 1887
, Samson and Seifert 1985
).
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MATERIALS AND METHODS |
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were followed. Isolates were also cultured on a medium containing oatmeal agar (OA) prepared by blending 37.5 g of Quaker instant oats in 1000 mL of water for 30 s. Following the addition of 12.5 g of Difco agar, the mixture was heated to boiling and autoclaved. Cultures on CYA, MEA, G25N, and OA were incubated at 25 C under 12 h fluorescent light. Inoculated plates of CYA were also incubated at 5 C and 37 C. Material mounted in water was observed by bright field and differential interference contrast (DIC) microscopy. Material for scanning electron microscopy (SEM) was subjected to critical point drying and coating with gold, and was then examined with a Zeiss DSM950 instrument. The color designations are from Rayner (1970)
. Cultures were deposited in the Culture Collection and Research Center (CCRC) in Taiwan. Terms for different parts of conidiophores are after Pitt (1979)
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TAXONOMY |
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Stromata gregaria, dispersa, hemisphaerica vel depresso-sphaerica, sessilia, 1–2 mm diam x 1–1.5 mm alta; superficies immatura saepe squamis ochraceis reticulatis tegens, demum denigrata et ochracea suffusa, fere laevis vel leviter rimosa; textura sub superficie peridio coriaceo faciens, ca 0.1 mm crassa, nigella, pigmentis armeniacis in KOH dissolutis. Ascostromata unilocula, 0.8–1.8 mm diam, primo coriacea, post demum pulveracea, alba in centro, extrorsus lutea. Asci octospori, globosi vel subglobosi, 20–25 µm diam, evanescentes. Ascosporae hyalinae, unicellulares, ellipsoideae, apicibus acutis vel angustatis, 15–18.5 x 7.5–9.5 µm, cristis 4–6 flexuosis, verruculosae inter cristas.
Stromata gregarious, scattered on seed, hemispherical to depressed-spherical, sessile, 1–2 mm diam x 1–1.5 mm high; surface often overlain with ochraceous reticulate cracks when immature, becoming blackish and tinged ochraceous, nearly smooth or slightly cracked; tissue immediately beneath surface forming a coriaceous peridium ca 0.1 mm thick, blackish, with KOH-extractable pigments Apricot (42). Ascostromata uniloculate, 0.8–1.8 mm diam, coriaceous initially, becoming powdery, white at center, grading into yellow outwards. Asci eight-spored, globose to subglobose, 20–25 µm diam, evanescent. Ascospores hyaline, one-celled, ellipsoid, with acute to narrowly rounded ends, 15–18.5 x 7.5–9.5 µm, with 4–6 flexuous crests, finely warted (tuberculate by SEM) between crests.
Colonies on CYA at 25 C reaching 4.6–4.8 cm diam in 7 d, velvety, azonate, with entire to slightly crenate, subsurface margins, Amber (47) to Ochreus (44), with synnemata produced occasionally in 2 wk but abundantly afterwards. Reverse uncolored. Sporulating regions on surface of colonies and on top of synnemata.
Colonies on MEA at 25 C reaching 4.6–5 cm diam in 7 d, similar to those on CYA.
Colonies on G25N at 25 C restricted, not growing much beyond the points of inoculation.
Colonies on OA at 25 C reaching 2.2–2.5 cm diam in 7 d, with diffuse margins, otherwise as those on CYA. Teleomorph produced in stromata in 6 wk, with morphological features as described for naturally-occurring material.
No growth on CYA at 5 C. Colonies on CYA at 37 C restricted, not growing much beyond the points of inoculation.
Anamorph in culture: Conidiogenous structure monoverticillate or infrequently biverticillate as defined in Pitt (1979)
, mononematous or synnematous, with synnemata capitate to cylindrical, Amber (47), up to 1.5 mm long x 0.1–0.3 mm broad. Conidiophores composed of unbranched to less frequently branched stipes, hyaline to subhyaline, smooth, vesiculate on top, 6.5–9 µm diam at stipes, 17–25 µm diam at vesicles. Phialides 5–12 in groups, hyaline to yellowish, cylindrical, tapering on top, with short collula, 25–40 x 6–8 µm. Conidia produced enteroblastically in basipetal sequence, hyaline to yellowish, roughened with reticulation or coil-like ornamentation, ellipsoid, 12–17 x 7–11 µm, slightly flattened at one or both ends.
Anamorph from naturally-occurring material: much as described in culture, but with conidiogenous structure mostly synnematous and more compacted, up to 7 mm long, 0.4–1 mm broad at top, 0.2–0.3 mm broad at stipe.
Specimens examined. TAIWAN. TAIWAN PROV.: Ping-tung Co., Heng-chun, Ken-ting, on seeds of Diospyros discolor half-buried in soil, 26 Nov 2000, Hsieh, H.-M. & Ju, Y.-M. 89112611 (CULTURED), Penicilliopsis pseudocordyceps (HOLOTYPE HAST, ISOTYPE WSP); TAIWAN PROV.: Ping-tung Co., Heng-chun, Ken-ting, on seeds of Diospyros discolor half-buried in soil, 16 Sep 2000, Hsieh, H.-M. & Ju, Y.-M. 89091649, Pseudocordyceps seminicola (HAST). ZAIRE. Bambesa, on seeds, no date, Vrydagh K2882, Pseudocordyceps seminicola (NEOTYPE BR).
Commentary.
Pseudocordyceps seminicola was previously represented only by several collections from Zaire, one of which was selected as the neotype by Samson and Seifert (1985)
. We have examined the neotype specimen; it contains only the anamorph without any trace of the teleomorph. Our Taiwan collection of P. seminicola is much the same as the neotype material but, in addition, contains immature stromata of P. pseudocordyceps. In the mature collection of P. pseudocordyceps, rusty brown, peg-like synnematous remnants of P. seminicola still can be seen among the stromata. Penicilliopsis pseudocordyceps differs from the other two species of Penicilliopsis in possessing a sessile stroma with a cracked surface that is particularly noticeable on immature stromata, larger ascospores, and orange-brown KOH-extractable pigments in the stromatal peridium. Features of the ascospores revealed by SEM also differ in these species. The ascospores of P. pseudocordyceps have entire crest edges and the surface between the crests is tuberculate (Fig. 12 ), whereas P. clavariaeformis and P. africana possess ascospores with dentate crest edges and echinulate spore surfaces (Fig. 22 and Samson and Seifert 1985
, respectively). The anamorph further separates P. pseudocordyceps from the other two Penicilliopsis species. The synnemata of P. pseudocordyceps are capitate, unbranched and have a determinate growth, while those of P. clavariaeformis and P. africana are filiform, branched or unbranched and have an indeterminate axial growth. Conidiophores of P. pseudocordyceps are monoverticillate, with the phialides directly borne on vesiculate stipes; those of P. clavariaeformis and P. africana, on the other hand, are biverticillate or terverticillate.
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Penicilliopsis clavariaeformis Solms-Laubach, Ann. Jard. Bot. Buitenzorg 6: 53. 1887. Figs. 18–27
Anamorph. Sarophorum palmicola (Henn.) Seifert & Samson, in Samson & Pitt, Advances in Penicillium and Aspergillus systematics: 403. 1985.
The teleomorph is much as described in Samson and Seifert (1985)
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Colonies on CYA at 25 C reaching 7 cm diam in 7 d, velvety, with diffuse, subsurface margins, Olivaceous Buff (89), with synnemata abundantly produced and arranged concentrically. Reverse Fulvous (43). Sporulating regions mainly on entire synnematous surface but also on colony surface.
Colonies on MEA at 25 C reaching 5.1–5.5 cm diam in 7 d, velvety, with crenate, subsurface margins, Amber (47) to Honey (64), with synnemata produced sparingly, with Pure Yellow (14) pigments diffused into media. Reverse Pure Yellow (14). Sporulating regions on surface of colonies and on entire surface of synnemata.
No growth on G25N at 25 C.
Colonies on OA at 25 C similar to those on CYA but with more abundant sporulation.
No growth on CYA at 5 C. Colonies on CYA at 37 C restricted, not growing much beyond the points of inoculation.
Anamorph in culture: Conidiogenous structure terverticillate or occasionally biverticillate as defined in Pitt (1979)
, mononematous or synnematous, with synnemata acicular, Amber (47) when fresh, becoming Olivaceous (48) when dried, flexuous, unbranched or infrequently branched, up to 2 cm long x 0.5–1.5 mm at base. Conidiophores composed of stipes, ramuli, and metulae; stipes hyaline to pale brown, cylindrical, smooth, branched or unbranched, 5.5–6.5 µm broad; ramuli 2–3 per stipe, pale brown, short-clavate or barrel-shaped, 10–16 x 7–9 µm; metulae 3–5 per ramulus, hyaline, clavate, smooth, slightly vesiculate on top, 13–18 x 7–8.5 µm. Phialides 5–10 per metula, hyaline to yellowish, ampuliform, 15–20 x 3.5–4.5 µm. Conidia produced enteroblastically in basipetal sequence, hyaline to yellowish, smooth to finely roughened, ellipsoid, 5.5–9.5 x 3.5–5 µm, slightly flattened at one or both ends.
Anamorph from naturally-occurring material: much as described in culture.
Specimens examined. TAIWAN. TAIWAN PROV.: Ping-tung Co., Heng-chun, Ken-ting, on seeds of Diospyros discolor half-buried in soil, 26 Nov 2000, Hsieh, H.-M. & Ju, Y.-M. 89112612 (CULTURED), Penicilliopsis clavariaeformis (HAST); TAIWAN PROV.: Ping-tung Co., Heng-chun, Ken-ting, on seeds of Diospyros discolor half-buried in soil, 16 Sep 2000, Hsieh, H.-M. & Ju, Y.-M. 89091650, Sarophorum palmicola (HAST).
Commentary.
The anamorph in general agrees with that described by Samson and Seifert (1985)
except for lacking the cylindrical conidia which they termed "B conidia." Samson and Seifert (1985)
stated that the cylindrical conidia were "usually not formed in culture," but we could not find this type of conidia in the Taiwan material from nature or in culture. The anamorph, Sarophorum palmicola, has been found fairly widespread in the tropics. It has been known previously from Indonesia, Brazil, Ivory Coast, Papua New Guinea, Singapore, and Malaysia. The teleomorph, however, is not as commonly found. Although the teleomorph-anamorph connection between P. clavariaeformis and S. palmicola has been known for a long time, we experimentally demonstrate the connection by culturing, i.e., with the S. palmicola anamorph produced from cultures initiated from ascospores of P. clavariaeformis.
Penicilliopsis africana highly resembles P. clavariaeformis in having stipitate stromata and bi- or terverticillate conidiophores, but differs primarily in having a smaller ascospore size range and tentaculate synnemata (Samson and Seifert 1985
).
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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Accepted for publication October 31, 2001.
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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Rayner RW., 1970 A mycological colour chart Kew: Commonwealth Mycological Institute. p 34, charts I and II
Samson RA, Seifert KA., 1985 The ascomycete genus Penicilliopsis and its anamorphs In: Samson RA, Pitt JI, eds. Advances in Penicillium and Aspergillus systematics. New York: Plenum Press. p 397–428
Solms-Laubach HG., 1887 Penicilliopsis clavariaeformis, ein neuer javanischer Ascomycet Ann Jard Bot Buitenzorg 6:53-72
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