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Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai, Thailand
P. Lumyong
Department of Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai, Thailand
S. R. Whitton
E. H. C. McKenzie
Landcare Research, Private Bag 92170, Auckland, New Zealand
K. D. Hyde
Centre for Research in Fungal Diversity, Department of Ecology & Biodiversity, The University of Hong Kong, Pokfulam Road, Hong Kong, China
| ABSTRACT |
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Linocarpon species are reported from Pandanaceae in Australia, Brunei, Hong Kong, Nepal, New Zealand, Philippines, Seychelles, Thailand and Vanuatu. Linocarpon lammiae sp. nov. were collected on decaying leaves of Pandanus tectorius in Hong Kong. Linocarpon siamensis sp. nov. and L. suthepensis sp. nov. were collected from decaying leaves of P. penetrans in Thailand. These taxa are described, illustrated and compared with Linocarpon species with similar ascospore morphology and dimensions. Included are a synoptic table, which compares the new species to similar known species, and a dichotomous key to species of Linocarpon known from members of the Pandanaceae.
Key words: new species, Pandanus, saprophytic fungi, systematics
| INTRODUCTION |
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Species of Linocarpon are distinguished among each other primarily through ascospore morphology and ascospore dimensions. Ascospore shape, the presence or absence of a gelatinous appendage and the morphology of appendages are of primary importance (Hyde 1992
, 1997
, Dulymamode et al 1998
, Poonyth et al 2000
). Linocarpon ascospores are aseptate, and might lack appendages, have a single appendage or have bipolar appendages.
In a review of scolecosporous genera of ascomycetes, Walker (1980)
discussed the current taxonomic status of species referred to Linocarpon and accepted five species, while L. appendiculatum K.D. Hyde, L. nipae (Henn.) K.D. Hyde and L. pandani were recorded from mangrove palm Nypa fruticans by Hyde (1988)
. Hyde (1992)
revised Linocarpon, accepting 10 species, and introduced L. bipolaris K.D. Hyde as well as L. longisporum K.D. Hyde. After this revision, other species were added; L. hamasporum (Ellis & Everh.) M.E. Barr (Barr 1993
); 11 new species and one new combination and key to all 23 accepted species (Hyde 1997
); four new species (Dulymamode et al 1998
); L. angustatum K.D. Hyde & Alias (Hyde and Alias 1999
); and four new species (Fröhlich and Hyde 2000
).
Nine species of Linocarpon have been described from pandanaceous substrates: L. appendisporum K.D. Hyde; L. breve K.D. Hyde; L. falciformisporum K.D. Hyde; L. fasciatum Dulymamode; P.F. Cannon & Peerally; L. pandani, L. pandanicola K.D. Hyde; L. spathulatum Dulymamode; P.F. Cannon & Peerally; L. sulcatum Dulymamode, P.F. Cannon & Peerally from Pandanus spp.; and L. freycinetiae (Rehm) K.D. Hyde from Freycinetia sp. (Hyde 1992
, 1997
, Dulymamode et al 1998
). Two other species, L. livistonae (Henn.) K.D. Hyde and L. elaeidis Petr., previously known from palms, also have been reported from Pandanus spp. (Hyde 1997
, Dulymamode et al 1998
). This research reports on seven species of Linocarpon, all on Pandanaceae, three of which are introduced as new to science.
| MATERIALS AND METHODS |
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25 C) to induce ascomata development. Care was taken not to put too many leaves in each container. All material was examined within a day and up to a month of collection. Material from fresh specimens were mounted in water, as in previous studies, and observed under differential interference microscopy. Measurements were made from discharged ascospores. | TAXONOMY |
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Type species: Linocarpon pandani (Syd. & P. Syd.) Syd. & P. Syd., Ann. Mycol. 15: 210. 1917.
Linospora pandani Syd. & P. Syd., Ann. Mycol. 11: 60. 1913.
Asci 79105 µm long, 911 µm wide at the widest point. Ascospores 27.536 x 35 µm.
Habitat/Distribution.
Known to inhabit decaying leaves of Pandanus sp. and P. nepalensis. Indonesia (Hyde 1997
), Nepal (current specimen).
Specimen examined. NEPAL, Pokhara, in forest around Lake Phewa, on decaying leaves of Pandanus nepalensis, 1 Oct 1997, S.R. Whitton HKU(M) 12710.
Notes.
Linocarpon falciformisporum is unique, being easily recognized by its short, cylindrical ascospores and a veil-like appendage at one end of the ascospore (Hyde 1997
, figure in Table I). The current specimen has ascospores that are slightly shorter than those reported by Hyde (1997)
for L. falciformisporum (3342 x 2.54.5 µm), but this variation is small and is not thought to be of taxonomic significance.
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Asci 72101 µm long, 7.510.5 µm wide at the widest point. Ascospores 3240 x 22.8 µm.
Habitat/Distribution.
Known to inhabit decaying leaves of Freycinetia sp. Philippines (Hyde 1997
, current specimen).
Specimen examined. PHILIPPINES, Luzon Island, Quezon Region, Los Ba
os, Mount Makiling, Baranggay Bagang Silang, on decaying leaves of Freycinetia sp., 22 Oct 1996, S.R. Whitton HKU(M) 4983.
Notes.
Linocarpon freycinetiae, described from Freycinetia from the Philippines, is characterized by short acicular ascospores. The current specimen differs from the holotype (Hyde 1997
) in having slightly longer ascospores (3240 x 22.8 µm vs. 2832 x 2.53 µm) and a distinct mucilaginous pad at the base of the ascospore. The mucilaginous appendage might not have preserved well in the old material. These differences are not significant enough to warrant specific recognition. Linocarpon freycinetiae is similar to L. falciformisporum in respect to ascospore size but differs in having slightly smaller ascospores (2832 x 2.53 µm vs. 3342 x 2.54.5 µm) and lacks the distinctive veil-like appendage found in freshly examined material of L. falciformisporum (Hyde 1997
).
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Etymology. Lammiae, referring to the type locality, Lamma Island, Hong Kong.
Ascomata forming beneath black (sometimes nonpigmented), raised, circular, shiny, dome-shaped areas (pseudostromata) on the host surface, 400675 µm diam, with a central, raised, black, shiny papilla, solitary or gregarious; pseudostromata pigmentation variable, sometimes black and shiny, sometimes nonpigmented; in some pseudostromata, the pigmentation is well defined with distinct edges, in others pigmentation is variable or indistinct. Ascomata 420485 µm wide, 190202 µm high, lenticular, immersed beneath pseudostromata. Ostiole central, surrounded by a dense area of black, stromatized tissue, which is surrounded by short, hyaline, thin-walled, septate, obtuse periphysoids. Peridium 414 µm thick, comprising 24 layers of thin-walled, brown to pale brown, flattened, elongate cells. Paraphyses 37 µm diam., irregularly cylindrical, flexuous, hyaline, smooth, septate, with a broadly rounded apex. Asci 97133 µm long, 13.526 µm wide, 8-spored, clavate, tapering to a rounded or truncate apex, tapering below to a narrow, obtuse base, thin-walled, with a refractive, J-cylindrical apical ring. Ascospores 5376 µm long, 4.57 µm wide, arranged in a single, overlapping fascicle, acicular, with a broadly rounded apex, tapering to an acute base, smooth, hyaline, aseptate, typically slightly curved, without visible gelatinous appendages.
Habitat/Distribution. Known to inhabit decaying leaves of Pandanus tectorius. Australia, Hong Kong.
Specimens examined. AUSTRALIA, QUEENSLAND, Brisbane, on decaying leaves of Pandanus tectorius, 5 Jan 1997, J.G. Wright HOLOTYPE: HKU(M) 12707. HONG KONG, Lamma Island, Hung Shing Yhe Beach, on decaying leaves of P. tectorius, 24 July 1997, S.R. Whitton HKU(M) 12706.
Notes.
Linocarpon lammiae is characterized by relatively short, acicular ascospores that have a broadly rounded apex, taper to an acute base, and have no gelatinous appendages (in fresh material). Other species of Linocarpon with short spores are L. freycinetiae, L. clavatum K.D. Hyde, L. falciformisporum, L. breve, L. palmetto (Ellis & Everh.) M.E. Barr and L. spathulatum. Linocarpon palmetto has filiform ascospores (5056 x 2.53.5 µm) and is a pathogen causing necrotic regions on leaves of Sabal palmetto. The ascospores of L. breve (3445 x 2.22.6 µm), L. falciformisporum (3342 x 2.54.5 µm) and L. freycinetiae (2832 x 2.53 µm) are filiform/cylindrical and shorter than those of L. lammiae (Hyde 1997
). In addition, L. falciformisporum has a veil-like appendage at one end of the ascospore. In L. clavatum the ascospores are widest at the center, rounded at the apex, truncate and appendaged at the base, and shorter (4151 x 45.5 µm) than those of L. lammiae. Linocarpon spathulatum has similarly shaped ascospores to those of L. lammiae, but they typically are slightly longer (6689 x 46 µm versus 5376 x 4.57 µm) and the base has a distinct gelatinous cap (Dulymamode et al 1998
). Based on ascospore morphology and dimensions, these specimens require specific recognition (see key).
Asci 85150 µm long, 713 µm wide at the widest point. Ascospores 5796 x 24 µm.
Habitat/Distribution.
Known to inhabit decaying leaves of Freycinetia banksii, Miscanthus sp., Pandanus sp., P. copelandii, P. furcatus, P. laevis, P. monticola, P. seychellarum, P. simplex, P. tectorius. Australia (current specimens), Brunei (current specimen), Hong Kong (current specimens), New Zealand (current specimen), Philippines (Hyde 1992
, current specimens), Seychelles (current specimen), Taiwan (Hyde 1992
), Vanuatu (current specimen).
Specimens examined. AUSTRALIA, NORTH QUEENSLAND, next to the road to Branston Beach from Cairns, on decaying leaves of Pandanus tectorius, 16 June 1996, S.R. Whitton HKU(M) 4977; HKU(M) 4996; 17 June 1996, HKU(M) 4978; North Queensland, Lacey Creek State Forest Park, on decaying leaves of P. monticola, 17 June 1996, S.R. Whitton HKU(M) 4979; HKU(M) 4982; North Queensland, in forest next to Lake Barrine, on decaying leaves of P. monticola, 18 June 1996, S.R. Whitton HKU(M) 4999. BRUNEI DARUSSALAM, Bandar Seri Begawan, in the mangroves at the mouth of Sungai Brunei, on decaying leaves of Pandanus sp., 28 Oct 1995, S.R. Whitton HKU(M) 12711. HONG KONG, Hong Kong Island, Mount Austin, on decaying leaves of P. furcatus, 7 June 1995, S.R. Whitton HKU(M) 4995; Hong Kong Island, The Peak, on decaying leaves of P. furcatus, 8 Aug 1995, S.R. Whitton HKU(M) 4997; Lamma Island, near Lamma Island Youth Hostel, on decaying leaves of P. tectorius, 28 Jan 1997, S.R. Whitton HKU(M) 5000; Lamma Island, Hung Shing Yhe Beach, on decaying leaves of P. tectorius, 24 July 1997, S.R. Whitton HKU(M) 14118; Hong Kong Island, Pokfulam, off Hatton Road in bush above Hong Kong University, on decaying leaves of P. furcatus, 7 Aug 1997, S.R. Whitton HKU(M) 12712. NEW ZEALAND, North Island, northern Hawke Bay, White Pine Bush, on decaying leaves of Freycinetia banksii, 30 Apr 1997, S.R. Whitton HKU(M) 12709. PHILIPPINES, Luzon Island, Quezon Region, Sinoloan, Baranggay Magsaysay U. P. Site, on decaying leaves of Pandanus sp., 21 Oct 1996, S.R. Whitton HKU(M) 4990; HKU(M) 4991; HKU(M) 4992; Luzon Island, Quezon Region, Baranggay Papalong Infanta, on decaying leaves of P. copelandii, 22 Oct 1996, S.R. Whitton HKU(M) 4988; HKU(M) 4989; HKU(M) 4993; Luzon Island, Quezon Region, Laguna, Baranggay Bakas Luisiana, on decaying leaves of P. simplex, 23 Oct 1996, S.R. Whitton HKU(M) 4986; HKU(M) 4987; Luzon Island, Quezon Region, Laguna, Baranggay Balaho Cavinti, on decaying leaves of P. simplex, 23 Oct 1996, S.R. Whitton HKU(M) 4994. SEYCHELLES, Mahe Island, La Reserve, on decaying leaves of P. seychellarum, 31 July 1996, K.D. Hyde HKU(M) 4984. VANUATU, Espiritu Santo Island, South Santo, on decaying leaves of Pandanus sp., 31 Oct 1996, E.H.C. McKenzie HKU(M) 4985.
Notes.
Linocarpon pandani is characterized by long, narrow ascospores (6280 x 24 µm) that have distinctive septum-like bands (closely packed guttules) and no mucilaginous appendages. Linocarpon pandanicola is similar to L. pandani in morphology and ascospore dimensions (72100 x 2.63.2 µm). The major morphological character distinguishing these two species is the presence of mucilaginous appendages in L. pandanicola (Hyde 1992
, 1997
). Many of these specimens have ascospores longer than that reported by Hyde (1992)
for L. pandani, but due to the lack of mucilaginous appendages it is considered more appropriate to place them in L. pandani than L. pandanicola.
Asci 100136 µm long, 89 µm wide at the widest point. Ascospores 7490 x 23 µm.
Habitat/Distribution.
Known to inhabit decaying leaves of Pandanus sp., P. furcatus and decaying petioles of Mauritia flexuosa. Brunei Darussalam (current specimen), Ecuador (Fröhlich & Hyde 2000
), Hong Kong (current specimen), Indonesia (Hyde 1997
).
Specimens examined. BRUNEI DARUSSALAM, Temburong, Batu Apoi Forest Reserve, Kuala Belalong Field Studies Centre, alongside Esu Stream, on decaying leaves of Pandanus sp., 25 Oct 1995, S.R. Whitton HKU(M) 4981. HONG KONG, New Territories, Tai Po Kau Country Park, on decaying leaves of P. furcatus, 1995, S.R. Whitton HKU(M) 4980.
Notes.
Linocarpon pandanicola is characterized by long, narrow ascospores (72100 x 2.63.2 µm) with small mucilaginous pads at each end (Hyde 1997
). Linocarpon pandani is similar to L. pandanicola but differs by having slightly shorter ascospores and no mucilaginous appendages (Hyde 1992
, 1997
). Both specimens collected in this study have mucilaginous appendages and thus are identified as L. pandanicola.
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Etymology. In reference to Siam (Thailand), where the specimens were collected.
Ascomata forming beneath black, raised, circular, shiny, dome-shaped areas (pseudostromata) on the host surface, 575825 x 650875 µm, with a minute central black dot, solitary. Pseudostromata pigmentation variable, sometimes being black and shiny, sometimes non-pigmented. Ascomata 850950 µm wide, 140240 µm high, hemispherical, with a flattened base. Ostiole central, raised. Peridium thin, comprising a few layers of brown-walled, compressed cells. Paraphyses 312 µm at the widest point, filamentous, smooth, septate, apex broadly rounded or obtuse, flexuous, hyaline, tapering toward the apex, surrounded by a gelatinous matrix. Asci 100156 x 7.79.3 µm, 8-spored, cylindrical, pedicellate, rounded or truncate apex, with a J- ring. Ascospores 5971 x 3.13.3 µm (
= 67.3 x 3.15 µm, n = 20), arranged in a single overlapping fascicle, filiform, hyaline, often slightly curved or straight, containing numerous refringent, septum-like bands, gelatinous cap-like or pad-like appendages at both ends (Figs. 1821).
Habitat/Distribution. Known to inhabit decaying leaves of Pandanus penetrans, Thailand.
Specimens examined. THAILAND, Chiang Mai, Doi Suthep-Pui National Park, on decaying leaves of Pandanus penetrans, 21 July 2001, S. Thongkantha PQ 010, HOLOTYPE: PDD 74300; 31 July 2001, S. Thongkantha PQ 010/1 PDD 74301.
Notes.
Linocarpon siamensis is characterized by filiform ascospores, with pad-like appendages at both ends. A similar species, L. appendisporum, is differentiated from L. siamensis by the morphology of the mucilaginous appendages and ascospore shape. In L. appendisporum the spore ends are truncate, while in L. siamensis they taper to a point (Hyde, 1997
). Linocarpon alpiniae K.D. Hyde, L. clavatum K.D. Hyde and L. falciformisporum have ascospores shaped similarly to those of L. siamensis but are shorter and slightly wider (L. alpiniae 5664 x 3.54.5 µm, L. clavatum 3342 x 2.54.5 µm, L. falciformisporum 4145 x 45.5 µm), and the latter two have a minute mucilaginous appendage, evident at only one end of the ascospore (Table I). Linocapon alpinae, which is most similar to L. siamensis, was observed as fresh material by Hyde (1997)
, and ascospores are narrower and taper to apiculate processes that lack appendages.
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Etymology. In reference to Doi Suthep Pui National Park, where the specimens were collected.
Ascomata forming beneath black, slightly raised, circular, grayish-black areas, 300485 µm diam (pseudostromata) on the host surface, with pronounced black papilla, typically gregarious. Ascomata 500650 µm wide, 162280 µm high, lenticular, black, developing deep inside leaf tissues. Ostiole central, raised. Peridium thin, comprising a few layers of brown-walled compressed cells. Paraphyses 4.67.7 µm at the widest point, filamentous, smooth, septate, obtuse, flexuous, hyaline, tapering toward the apex. Asci 7792.5 x 6.17.7 µm, 8-spored, cylindrical, pedicellate, apex rounded or truncate, with a J- apical ring. Ascospores 18.530.8 x 2.33.1 µm (
= 25.5 x 3 µm, n = 20), multiseriate, cylindrical, hyaline, straight or slightly curved, often containing numerous refringent septum-like bands, with a basal, arrow-like appendage.
Habitat/Distribution. Known to inhabit decaying leaves of Pandanus penetrans, Thailand.
Specimens examined. THAILAND: Chiang Mai, Doi Suthep Pui National Park, in rainforest, on dead leaves of Pandanus penetrans, 21 July 2001, S. Thongkantha PQ 011, HOLOTYPE: PDD 74602; 31 July 2001, S. Thongkantha PQ 011/1 PDD 74416.
Notes. Linocarpon suthepensis is characterized by short ascospores that have a rounded apex, with an arrow-like appendage at the base. Other species of Linocarpon with short ascospores and an appendage at only one end are L. clavatum and L. falciformisporum. Linocarpon suthepensis differs in having shorter ascospores (L. clavatum 3342 x 2.54.5 µm, L. falciformisporum 4145 x 45.5 µm) and for Linocarpon species, a unique appendage morphology. Appendages of L. clavatum and L. falciformisporum are mucilaginous and veil-like, respectively. Ascospores of L. suthepensis are shorter than L. appendisporum (6076 µm) and L. breve (3445 µm), which have appendages at both ends of the ascospores (Table I).
KEY TO SPECIES OF LINOCARPON KNOWN FROM THE PANDANACEAE
1. Ascospores swollen toward the apex, tapering to a narrow base . . . . . 2
1. Ascospores cylindrical or filiform . . . . . 4
2. Ascospores 76107 x 34 µm; basal appendage of ascospores 34.5 x 11.5 µm; asci 92170 x 1220 µm, cylindrical . . . . . L. sulcatum
2. Ascospores wider than 4 µm . . . . . 3
3. Ascospores 6689 x 46 µm, with short (6 x 2 µm) gelatinous appendage at base; asci 110170 x 1216 µm, cylindrical . . . . . L. spathulatum
3. Ascospores 5376 x 4.57 µm, without appendages; asci 97133 x 13.526 µm, clavate . . . . . L. lammiae
4. Ascospores <60 µm long . . . . . 5
4. Ascospores >60 µm long . . . . . 8
5. Appendages absent, ascospores 2832 x 2.53 µm; asci 5482 x 710 µm, cylindric-clavate . . . . . L. freycinetiae
5. Ascospores with appendages . . . . . 6
6. Appendage at each end collar-like, ascospores 3445 x 2.22.6 µm; asci 104138 x 4.56 µm, cylindrical . . . . . L. breve
6. Ascospore appendages only at basal end . . . . . 7
7. Ascospores 3342 x 2.54.5 µm; appendage mucilaginous, becoming sickle-shaped or veil-like in water; asci 112140 x 810 µm, cylindrical . . . . . L. falciformisporum
7. Ascospores 18.531 x 2.33.1 µm; appendage arrow-like; asci 7792.5 x 6.17.7 µm, cylindrical . . . . . L. suthepensis
8. Ascospore appendages absent . . . . . 9
8. Ascospore appendages present . . . . . 10
9. Ascospores 6280 x 24 µm; asci 100140 x 810 µm, cylindrical . . . . . L. pandani
9. Ascospores 84110 x 2.53.5 µm; asci 110178 x 814 µm, cylindrical . . . . . L. fasciatum
10. Ascospore appendages basal . . . . . 11
10. Ascospore appendages bipolar . . . . . 12
11. Ascospores 70104 x 1.62.3 µm; appendage mucilaginous; asci 100140 x 612 µm, cylindrical . . . . . L. livistonae
11. Ascospores 7297 x 34 µm; appendage mucilaginous; asci 116148 x 913 µm, cylindrical . . . . . L. elaeidis
12. Ascospore 72100 x 2.63.2 µm; appendages pad-like; asci 160190 x 810 µm, cylindrical . . . . . L. pandanicola
12. Ascospores typically shorter . . . . . 13
13. Ascospores 5971 x 3.13.3 µm; appendages pad-like; asci 100156 x 7.79.3 µm, cylindrical . . . . . L. siamensis
13. Ascospores 6076 x 2.74 µm, appendages gelatinous, spreading in water; asci 100150 x 810 µm, cylindrical . . . . . L. appendisporum
| ACKNOWLEDGMENTS |
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| FOOTNOTES |
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Accepted for publication August 31, 2002.
| LITERATURE CITED |
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Dulymamode R, Cannon PF, Peerally A., 1998 Fungi from Mauritius: Linocarpon species on Pandanus. Mycol Res 102:1331-1337
Fröhlich J, Hyde KD., 2000 Palm Microfungi. Fungal Diversity Research Series 3, Fungal Diversity Press, Hong Kong, 380 p
Hyde KD., 1988 The genus Linocarpon from the mangrove palm Nypa fruticans. Trans Mycol Soc Japan 29:339-350
. 1992 Fungi from palms. I. The genus Linocarpon, a revision. Sydowia 44:32-54
. 1997 Additions to the genus Linocarpon (Ascomycetes: Hyponectriaceae). Bot J Linn Soc 123:109-131
, Alias SA., 1999 Linocarpon angustatum sp. nov., and Neolinocarpon nypicola sp. nov. from petioles of Nypa fruticans, and a list of fungi from aerial parts of this host. Mycoscience 40:145-150
Poonyth AD, Hyde KD, Wong SW, Peerally A., 2000 Ultrastructure of asci and ascospore appendages in Linocarpon appendiculatum and L. nipae. Bot Mar 43:213-221
Walker J., 1980 Gaeumannomyces, Linocarpon, Ophiobolus and several other genera of scolecospored ascomycetes and Phialiophora conidial states, with a note on hyphopodia. Mycotaxon 11:1-129
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