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Institute of Agriculture and Forestry, University of Tsukuba, Tsukuba, Ibaraki 305-8572, Japan
Shun-ichi Udagawa
Nodai Research Institute, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan
Neale L. Bougher
CSIRO Forestry and Forest Products, P.O. Box 5 Wembley, Western Australia 6913, Australia
Akira Suzuki
Faculty of Education, Chiba University, Chiba 263-8522, Japan
Inez C. Tommerup
CSIRO Forestry and Forest Products, P.O. Box 5 Wembley, Western Australia 6913, Australia
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONTAXONOMYLITERATURE CITED |
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The genus Thecotheus is reported in Australia for the first time. A new species, Thecotheus urinamans is described and illustrated and included in a key to all known species of the genus. Critical macro- and micromorphological comparisons are presented to distinguish the new species from several closely related species, particularly the widespread fungus Thecotheus crustaceus. Thecotheus urinamans was growing on rotting, moist, plant litter from an experimental plot treated with urea (ammonia) in the indigenous jarrah (Eucalyptus marginata) forest of Western Australia.
Key words: ammonia fungi, Discomycetes, Pezizales, taxonomy
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONTAXONOMYLITERATURE CITED |
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, Eckblad 1968, Yao and Spooner 2000), North America (Kimbrough 1969), South America (Pardo et al 1997), the western Himalaya (Kaushal 1980), New Zealand (Bell 1983), East Africa (Krug and Khan 1987), Oman (Gené et al 1993) and Taiwan (Wang and Kimbrough 1993, Wang 1994), and three noncoprophilous species from Europe (Kimbrough 1969) and North America (Pfister 1972, 1981). A recent world monograph of the genus by Aas (1992) added another seven coprophilous species. The Western Australian species appears after urea or ammonia treatment, and its morphology differs from all previously described species of the genus. In this paper the fungus is described as a new species and a key to the currently known species of Thecotheus is provided. |
TAXONOMY |
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TOPABSTRACTINTRODUCTIONTAXONOMYLITERATURE CITED |
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Apothecia primo convexa, deinde concava et fulva aut cervina, 1.5–5 mm diam. Excipulum ectale textura globuloso vel angulari compositum; excipulum medullare textura globuloso. Asci octospori, apice iodo caerulescenti, "Congo red" aequabiliter tincti, cylindracei, 155–230 x 10–17.5 µm magni. Ascosporae ellipsoideae, 12–16.5 x 6–8 µm magnae, laeves, "Cotton blue" valde tinctae. Paraphyses filiformes ca 1 µm diam, flavidae, septatae in apice leviter vesiculares 2–7 µm diam.
Apothecia disk-shaped, dull orange-brown (Methuen 5D5–5D7, Kornerup and Wanscher 1978) to tan brown (6D6–6D7), convex and finally concave, sessile or subsessile, 1.5–5 mm in diam (Fig. 7). Ectal excipulum 127.5–175 µm thick, of textura globulosa to textura angularis, composed of polygonal cells measuring 5–25 µm in diam, pale salmon pink, with hyphoid hairs. Medullary excipulum 450–500 µm thick, of textura globulosa, composed of thin walled hyphae 5–5.5 µm in diam. Asci cylindrical, eight-spored, 155–230 x 10–17.5 µm, apex bluing in Melzer's reagent, staining uniformly in Congo red; operculum with a distinct thickened line of dehiscence (Fig. 8). A mucilaginous perisporic layer is present around immature ascospores (Fig. 9). Ascospores ellipsoid, 12–16.5 x 6–8 µm, smooth, with a thick outer layer staining intensely in cotton blue (Fig. 10). Paraphyses filiform, ca 1 µm in diam, pale yellow, septate, enlarged at the apex, 2–7 µm in diam.
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Habitat. On rotting, moist litter debris, such as leaves and twigs below the surface litter layer.
Specimens examined. Australia: Alcoa World Alumina Australia Mine Site F4517, Torrens Road, 10 km north of Dwellingup, in plant litter including rotting leaves and twigs of Eucalyptus marginata (Jarrah) and Eucalyptus calophylla (Marri) forest experimentally treated with urea, 26 Aug 1996, N. Bougher & J. Catchpole [HOLOTYPE: PERTH 06232116, ISOTYPE: E5792 CSIRO Forestry and Forest Products, Perth, Western Australia; Dwellingup, Western Australia, in plant litter of Eucalyptus marginata (Jarrah) and Eucalyptus calophylla (Marri) forest experimentally treated with urea: on partially decomposed plant litter including leaves and twigs, 20 Aug 1996, A. Suzuki & I. Tommerup E5783 (CSIRO); on fallen leaf of Banksia grandis, 26 Aug 1996, A. Suzuki E5791 (CSIRO); on partially decomposed leaf litter including leaves and twigs, 26 Aug 1996, N. Bougher E5793 (CSIRO).
This species is recognized by a pink, salmon coloration in the ectal excipulum of the apothecia and the bluing reaction of the ascus wall in iodine restricted to the apical zone (not the entire wall). These characters are typical of the genus Thecotheus. Thecotheus urinamans is distinguished from the other known species by its ascospore size. Among the noncoprophilous species of Thecotheus, T. pallens (Boud.) Kimbr. has smooth ascospores, but they are larger (35–45 x 15–20 µm) than those of T. urinamans. Thecotheus rivicola (Vacek) Kimbr. and Pfister differs in ascospore ornamentation and size from T. urinamans, while T. phycophilus Pfister differs in the number of ascospores in the ascus and in having two types of paraphyses. The asci of T. urinamans resemble those of the noncoprophilous group of Thecotheus species in that they are nonprotruding. Thecotheus crustaceus (Starb.) Aas and N. Lundq. is a coprophilous species with smooth ascospores. This species has been established as the new combination of Ascophanus crustaceus Starb. and is synonymized with T. agranulosus Kimbr. (Aas 1992). Thecotheus crustaceus is a widespread species and has a great amount of variation in apothecial, ascal, ascospore and paraphyses characters (Aas 1992). We examined the morphological characters of the holotype of T. crustaceus (F24444, S) and paratype (MICH 7171). In the type specimens of T. crustaceus, the apothecia were less than 1 mm diam with most apothecia less than 0.5 mm. According to the description of T. crustaceus (Aas 1992) the apothecia are 0.5–2 mm diam. However, our observation of the type specimens showed that apothecia were larger than 1 mm diam only if they were appressed. It was difficult to find a large apothecium that was more than 1 mm diam, even if moistened. We examined several apothecia that appeared to be more than 1 mm diam. In thin sections, these apothecia appeared to be a complex of two or three disks. The apothecia of this fungus tend to be aggregated or unified (like Orbilia spp. or Dermateaceae) when mature and the true size of individual apothecia is less than 1 mm. Our specimens of T. urinamans ranged in diam from 1.5 to 5 mm, so the apothecia sizes of these two species are quite different. Apothecia of the type specimens of T. crustaceus were not cup-shaped and the apothecia did not support subiculum-like hypha or protruding hypha from the ectal excipulum (outer side of apothecia). There were only minor differences in the size and shape of asci and paraphyses between T. urinamans and the various specimens of T. crustaceus (Table I). The average size of ascospores of T. urinamans (14.4 x 7.1 µm) differed from those of T. crustaceus (e.g., 17.0 x 8.6 µm for the holotype), but size ranges overlap (Table I). We also compared the sizes of immature ascospores that had not been released from asci and were embedded in cytoplasm. The size of immature ascospores of T. urinamans were smaller than those of T. crustaceus (Table I). Hymenial morphology of T. urinamans is similar to that of T. crustaceus, but the shape and size of the apothecia, the nonprotruding nature of asci and the range of ascospore sizes of T. urinamans represent significant differences from T. crustaceus. Thecotheus urinamans is similar to T. himalayensis Kaushal (FLAS F50920 and FLAS F50929) in ascospore size. However, T. himalayensis has verruculose ascospores and two types of paraphyses. Thecotheus africanus Khan and Krug also is similar to T. urinamans in size of ascospores but has projecting collarettes on the ascospores. Thecotheus species, particularly the coprophilous ones, generally are widespread in temperate to tropical regions of the world. However, the genus has not been reported from Australia. Two species, T. crustaceus (= T. agranulosus) and T. cinereus (Cr. and Cr.) Chen., have been reported from New Zealand (Bell 1983). Thecotheus cinereus also has smooth ascospores, ellipsoid to cylindrical-ellipsoid, but the spores are larger, (30–42 x 15–18 µm), than those of T. urinamans. Aas (1992) noted that there are two types of paraphyses in T. cinereus, whereas T. urinamans has one type of paraphyses.
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). However, some of these fungi have other habitats. For example, Amblyosporium botrytis Fres. (Pirozynski 1969, Sagara 1975), Ascobolus denudatus Fr. (van Brummelen 1967, Sagara 1975), and Coprinopsis cinerea Schaeff.: Fr. Redhead, Vilgalys and Moncalvo. (Sagara 1975, Orton and Watling 1979, Redhead et al 2001) have been reported to occur on both urea-treated soil and on dung. Laccaria bicolor (Mre.) Ort., Lactarius chrysorrheus Fr. and Rhizopogon succosus A. H. Smith (R. superiorensis A. H. Smith) occur on both urea-treated soil and on burnt soil (Sagara 1992). Some ammonia fungi have the capacity to occupy ecological niches indicative of pyrophilous fungi and coprophilous fungi, whereas other ammonia fungi do not appear to have such broad ecological capacity. We have observed T. urinamans only on forest plots treated with urea and, because the species could not be found on and/or around dung, we regard this species to be noncoprophilous.
The monograph on Thecotheus by Aas (1992) might not be readily available to most mycologists, as commented by Yao and Spooner (2000). They also said that three species, T. flavidus Wang and Kimbr., T. formosanus Wang and T. harasisus Gené, Elshafie and Guarro, published after Aas' monograph, are similar to some of the other known species. A revised key to the species of Thecotheus, including the new species, is presented below to aid further critical studies on the genus.
KEY TO THE SPECIES OF THECOTHEUS
1. On herbaceous, lignicolous or terricolous substrata . . . . . 2
1. On coprophilous substrata . . . . . 5
2. Ascospores apiculate; verruculose; 17–22 x 7–8(–9.5) µm . . . . . T. rivicola
2. Ascospores nonapiculate, smooth . . . . . 3
3. Asci usually four-spored; ascospores 30–36 x 15–16 µm; paraphyses of two types . . . . . T. phycophilus
3. Asci usually eight-spored; paraphyses of one type . . . . . 4
4. Ascospores 40–45 x 15–20 µm . . . . . T. pallens
4. Ascospores 12–16.5 x 6–8 µm . . . . . T. urinamans
5. Ascospores nonapiculate . . . . . 6
5. Ascospores apiculate . . . . . 13
6. Ascospores smooth . . . . . 7
6. Ascospores ornamented . . . . . 10
7. Paraphyses of one type . . . . . 8
7. Paraphyses of two types . . . . . 9
8. Ascospores less than 25 µm in length . . . . . T. crustaceus
8. Asci 16-spored, 32-spored or 64-spored; ascospores either 17–24.5 x 10–16 µm or 32–40 x 15–24 µm . . . . . T. pelletieri (Cr. and Cr.) Boud.
9. Ascospores ellipsoid, 28–32 (–36) x 14–16 µm . . . . . T. flavidus
9. Ascospores ellipsoid to cylindrical-ellipsoid, 30–42 x 15–18 µm . . . . . T. cinereus
10. Paraphyses strongly uncinate; ascospores regular ellipsoid; asci strangulate . . . . . 11
10. Paraphyses straight; ascospores slightly inequilateral, sub-ellipsoid; asci not strangulate . . . . . 12
11. Ascospores (13–) 14–17 x (6–) 7–9 (–9.5) µm . . . . . T. uncinatus Aas
11. Ascospores (20–) 22–26 (–28.5) x 11.5–16 µm . . . . . T. strangulates (Vel.) Aas and Lundq.
12. Ascospores (12–) 14–16 x (5.5–) 6–7 µm, ornamented with very small warts; paraphyses of two types . . . . . T. himalayensis
12. Ascospores 16–20 x 6–8 µm, ornamented with small extended warts to very fine reticulations; paraphyses of one type . . . . . T. viridescens E. Lundq.
13. Ascospores with projecting collarettes . . . . . 14
13. Ascospores without collarettes . . . . . 15
14. Apothecia yellowish; ascospores 12–15 x 7.5–9 µm . . . . . T. africanus
14. Apothecia white; ascospores 20–22 x 10–12 µm . . . . . T. perplexans (Faurel and Schotter) Krug and Khan
15. Ascospores smooth . . . . . 16
15. Ascospores warted . . . . . 17
16. Ascospores inequilateral, 13–16 x 7–8.5 µm; paraphyses of two types . . . . . T. inaequilateralis Aas
16. Ascospores symmetrical (16–) 17–21 (–22) x (7–) 8–10 (–11) µm . . . . . T. keithii (Phill.) Aas
17. Paraphyses of two types . . . . . 18
17. Paraphyses of one type . . . . . 19
18. Ascospores 18–20 x 7.5–9 µm; verruculose (warts less than 0.5 µm diam) . . . . . T. formosanus
18. Ascospores 23–28 x 12–13.5 µm; warty (warts 4.5–6.5 µm diam) . . . . . T. harasisus
19. Ascospores broad inequilateral; (14–) 16–19 x 7–9 (–10) µm; small punctate-like warts . . . . . T. biocellatus (Petrak) Aas
19. Ascospores longer than 25 µm, symmetrical . . . . . 20
20. Ascospores 25–30 x 12.5–14.5 µm; with ornamentations of minute granules . . . . . T. lundqvistii Aas
20. Ascospores (25–) 29–38 (–42) x (12–) 14–18 (–20) µm; with large isolated warts . . . . . T. holmskjoldii (E.C. Hansen) Chen.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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Accepted for publication March 11, 2003.
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONTAXONOMYLITERATURE CITED |
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Bell A., 1983 Dung fungi. Wellington,. New Zealand: Victoria Univ. Press. 88 p
Cannon PF, Hawksworth DL, Sherwood-Pike MA., 1985 The British Ascomycotina, an Annotated Checklist. Commonwealth Agricultural Bureaux, Slough
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Gené J, Elshafie AE, Guarro J., 1993 Two new coprophilous Pezizales from the Sultanate of Oman. Mycotaxon 46:275-284
Kaushal SC., 1980 A new species of Thecotheus (Pezizales) from the western Himalaya. Bot Not 133:319-321
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Orton PD, Watling R., 1979 Coprinaceae Part 1: Coprinus, British fungus flora Agarics and Boleti. In: Henderson DM, Orton PD, Watling R, eds. Royal Botanic Garden, Edinburgh
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Pfister DH., 1972 The psilopezizoid fungi. II. Thecotheus rivicola comb. nov. and other Iodophaneae (Pezizales) occurring on water-soaked wood. Bull Tor Bot Club 99:198-200
———. 1981 A new noncoprophilous species of Thecotheus, T. phycophilus. Mycologia 73:1001-1004
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Redhead SA, Vilgalys R, Monclvo J-M, Johnson J, Hopple JSJr., 2001 Coprinus Pers. and the disposition of Coprinus species sensu lato. Taxon 50:203-241
Sagara N., 1975 Ammonia fungi—a chemoecological grouping of terrestrial fungi. Contr Biol Lab Kyoto Univ 24:205-276, 7 pls
———. 1992 Experimental disturbances and epigeous fungi. In: Carroll GC, Wicklow DT, eds. The fungal community—its organization and role in the ecosystem. 2nd ed. New York: Mercel Dekker. p 427–454
van Brummelen J., 1967 A world-monograph of the genera Ascobolus and Saccobolus (Ascomycetes, Pezizales),. Persoonia, Suppl 1:1-260, 17 pls
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———, Kimbrough JW., 1993 A new species of Thecotheus (Pezizales) from Taiwan. Mycologia 85:1020-1022
Yao Y-J, Spooner BM., 2000 Notes on British species of Thecotheus (Ascobolaceae, Pezizales), with reference to other species of the genus. Kew Bulletin 55:451-457
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