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THE INTERNATIONAL JOURNAL OF FUNGAL TAXONOMY & NOMENCLATURE

Volume 96 : April-June 2006

CONTENTS

The genera Inocutis and Inonotus (Hymenochaetales) in Uruguay Sebastian Martinez 1 A new species and a new record of smut fungi from Northwestern China Shengrong Wang & Cuiyun Zeng 9 Additional and interesting lichenized and lichenicolous fungi from Turkey Mehmet Goékhan Halici, Mustafa Kocakaya & Ahmet Aksoy 13 Four species of Exobasidium from Korea Ji-Hyun Park, Kyung-Hee Kim, Kyun-Joon Lee & Seung-Kyu Lee 21 Annulate Pluteus species: a study of the genus Chamaeota in the United States Andrew M. Minnis, Walter J. Sundberg, Andrew S. Methven, Sedonia D. Sipes & Daniel L. Nickrent 31 Tylopilus microsporus, a new species from Southwest China S.-Z. Fu, Q.-B. Wang & Y.-J. Yao 41 An annotated checklist of Leccinum in China S.-Z. Fu, Q.-B. Wang & Y.-J. Yao 47 New species and new records of Graphis from India: transseptate species with completely carbonized exciples and norstictic acid Bharati Adawadkar & Urmila Makhija 51 Four species of Caloplaca (Teloschistaceae, lichenized Ascomycota) new to Poland Karina Wilk & Adam Flakus 61 The status of Psilocybe floridana (Strophariaceae) in Callistosporium

(Tricholomataceae) Etelvina Gandara & Gast6n Guzman 73 Three new species of Stemphyllum from China Yong Wang & Xiu-Guo Zhang 77 Caribbean heterobasidiomycetes: 2. Jamaica Peter Roberts 83

New taxa of the lichen genus Pertusaria (Ascomycota) from Thailand

Sureeporn Jariangprasert 109

Two new Cystoderma species from high Andean Ecuador Irja Saar & Thomas Lessee 123

Glomerospores: a new denomination for the spores of Glomeromycota, a group molecularly

distinct from the Zygomycota Bruno Tomio Goto & Leonor Costa Maia 129 The sequestrate genus Rhodactina (Boletales) in northern Thailand Zhu L. Yang, James M. Trappe, Manfred Binder, Rarunee Sanmee,

Pipob Lumyong & Saisamorn Lumyong 133

The Skibotn area in North Norway - an example of very high lichen species richness far to the north . Arve Elvebakk & Jarle W. Bjerke 141

_ Anew species of Didymium (Myxomycetes) with reticulate spores

Yuri K. Novozhilov & Inna V. Zemlyanskaya 147

[Content continues inside front cover] ISSN 0093-4666 MYXNAE 96: 1-364 (2006)

For subscription details, availability on microform, and availability of articles as photocopies or tear sheets, see back cover

[Content continued from front cover]

Anamorphic fungi from submerged leaves in Cuba: Brevicatenospora enteroproliferata gen.

et sp. nov. and Beltraniopsis aquatica sp. nov. Rafael F. Castaneda Ruiz, David W. Minter, Marc Stadler, Masatoshi Saikawa & Mayra Camino-Vilaré Leptosphaeria raphani does not occur on Draba aspera David L. Hawksworth

Drechslerella brochopaga, the anamorph of Orbilia (Hyalinia) orientalis Zefen Yu, Ying Zhang, Min Qiao, H.-O. Baral, E. Weber & Keqin Zhang A new species of Agyrium from Yunnan, China Wen-Ying Zhuang & Zhu-Liang Yang New species of Kananascus and its anamorph from Lithuania Au&ra Treigiené The genus Chlorophyllum (Basidiomycetes) in China Z.W. Ge & Zhu L. Yang Trechispora elongata species nova from North Europe

Otto Miettinen & Karl-Henrik Larsson ~

Two new species of Wrightoporia (Basidiomycota, Aphyllophorales) from southern China Yu-Cheng Dai & Bao-Kai Cui A new species of Oxyporus (Basidiomycota, Aphyllophorales) from Northwest China Bao-Kai Cui, Ming-Yun Huang & Yu-Cheng Dai Northern Antrodiella species: the identity of A. semisupina, and type studies of related taxa Otto Miettinen, Tuomo Niemela & Wjacheslav Spirin New records of Pluteus (Pluteaceae, Agaricales) from Brazil Felipe Wartchow, Vagner Gularte Cortez & Gilberto Coelho New species of marasmioid genera (Basidiomycetes, Tricholomataceae) from tropical Africa - VI. Marasmius cremeopileatus, a new species from Zimbabwe Vladimir Antonin & Cathy Sharp Myxomycete flora of Derebucak (Konya) and Akseki (Antalya) districts in Turkey Dursun Yagiz & Ahmet Afyon A taxonomical treatment of the North and Central American species in Lactarius sect. Deliciosi Jorinde Nuytinck, Steven L. Miller & Annemieke Verbeken A new species of Cephalotheca isolated from a Korean patient Takashi Yaguchi, Ayako Sano, Kyoko Yarita, Moo Kyu Suh, Kazuko Nishimura & Shun-ichi Udagawa A new species of Exobasidium (Exobasidiales) on Rhododendron from China

Zhenying Li & Lin Guo

Contribution to the study of Endogonales in Brazil: the first record of Peridiospora tatachia Bruno Tomio Goto & Leonor Costa Maia Book reviews and notices David L. Hawksworth

Indices & Information Nomenclatural novelties proposed in volume 96 Author index Reviewers Errata From the Editor-in-Chief

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MYCOTAXON

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MYCOTAXON

VOLUME NINETY-SIX TABLE OF CONTENTS

The genera Inocutis and Inonotus (Hymenochaetales) in Uruguay Sebastian Martinez A new species and a new record of smut fungi from Northwestern China Shengrong Wang & Cuiyun Zeng Additional and interesting lichenized and lichenicolous fungi from Turkey Mehmet Gokhan Halici, Mustafa Kocakaya & Ahmet Aksoy Four species of Exobasidium from Korea Ji-Hyun Park, Kyung-Hee Kim, Kyun-Joon Lee & Seung-Kyu Lee Annulate Pluteus species: a study of the genus Chamaeota in the United States Andrew M. Minnis, Walter J. Sundberg, Andrew S. Methven, Sedonia D. Sipes & Daniel L. Nickrent Tylopilus microsporus, a new species from Southwest China S.-Z. Fu, Q.-B. Wang & Y.-J. Yao An annotated checklist of Leccinum in China S.-Z. Fu, Q.-B. Wang & Y.-J. Yao New species and new records of Graphis from India: transseptate species with completely carbonized exciples and norstictic acid Bharati Adawadkar & Urmila Makhija Four species of Caloplaca (Teloschistaceae, lichenized Ascomycota) new to Poland Karina Wilk & Adam Flakus The status of Psilocybe floridana (Strophariaceae) in Callistosporium (Tricholomataceae) Etelvina Gandara & Gast6n Guzman Three new species of Stemphyllum from China Yong Wang & Xiu-Guo Zhang Caribbean heterobasidiomycetes: 2. Jamaica Peter Roberts New taxa of the lichen genus Pertusaria (Ascomycota) from Thailand Sureeporn Jariangprasert Two new Cystoderma species from high Andean Ecuador

Irja Saar & Thomas Lessge Glomerospores: a new denomination for the spores of Glomeromycota, a group molecularly distinct from the Zygomycota Bruno Tomio Goto & Leonor Costa Maia The sequestrate genus Rhodactina (Boletales) in northern Thailand Zhu L. Yang, James M. Trappe, Manfred Binder, Rarunee Sanmee, Pipob Lumyong & Saisamorn Lumyong The Skibotn area in North Norway - an example of very high lichen species richness far to the north Arve Elvebakk & Jarle W. Bjerke A new species of Didymium (Myxomycetes) with reticulate spores Yuri K. Novozhilov & Inna V. Zemlyanskaya Anamorphic fungi from submerged leaves in Cuba: Brevicatenospora enteroproliferata gen. et sp. nov. and Beltraniopsis aquatica sp. nov. Rafael F. Castafieda Ruiz, David W. Minter, Marc Stadler, Masatoshi Saikawa & Mayra Camino-Vilaré Leptosphaeria raphani does not occur on Draba aspera David L. Hawksworth

ili

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Drechslerella brochopaga, the anamorph of Orbilia (Hyalinia) orientalis Zefen Yu, Ying Zhang, Min Qiao, H.-O. Baral, E. Weber & Keqin Zhang A new species of Agyrium from Yunnan, China Wen-Ying Zhuang & Zhu-Liang Yang New species of Kananascus and its anamorph from Lithuania _AuSra Treigiené The genus Chlorophyllum (Basidiomycetes) inChina —_Z.W. Ge & Zhu L. Yang Trechispora elongata species nova from North Europe Otto Miettinen & Karl-Henrik Larsson Two new species of Wrightoporia (Basidiomycota, Aphyllophorales) from southern China Yu-Cheng Dai & Bao-Kai Cui A new species of Oxyporus (Basidiomycota, Aphyllophorales) from Northwest China Bao-Kai Cui, Ming- Yun Huang & Yu-Cheng Dai Northern Antrodiella species: the identity of A. semisupina, and type studies of related taxa Otto Miettinen, Tuomo Niemela & Wjacheslav Spirin New records of Pluteus (Pluteaceae, Agaricales) from Brazil Felipe Wartchow, Vagner Gularte Cortez & Gilberto Coelho New species of marasmioid genera (Basidiomycetes, Tricholomataceae) from tropical Africa - VI. Marasmius cremeopileatus, a new species

from Zimbabwe Vladimir Antonin & Cathy Sharp Myxomycete flora of Derebucak (Konya) and Akseki (Antalya) districts in Turkey Dursun Yagiz & Ahmet Afyon A taxonomical treatment of the North and Central American species in Lactarius sect. Deliciosi Jorinde Nuytinck, Steven L. Miller & Annemieke Verbeken

A new species of Cephalotheca isolated from a Korean patient Takashi Yaguchi, Ayako Sano, Kyoko Yarita, Moo Kyu Suh, Kazuko Nishimura & Shun-ichi Udagawa A new species of Exobasidium (Exobasidiales) on Rhododendron from China

Zhenying Li & Lin Guo

Contribution to the study of Endogonales in Brazil: the first record of Peridiospora tatachia Bruno Tomio Goto & Leonor Costa Maia Book reviews and notices David L. Hawksworth

Indices & Information Nomenclatural novelties proposed in volume 96 Author index Reviewers Errata From the Editor-in-Chief

PUBLICATION DATE FOR VOLUME NINETY-FIVE

MYCOTAXON for JANUARY-MARCH, VOLUME 95 (1-350 + I-VI) was issued on June 16, 2006

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Volume 96, pp. 1-8 April-June 2006

The genera /nocutis and Inonotus (Hymenochaetales) in Uruguay

SEBASTIAN MARTINEZ

sebamart@fing.edu.uy Laboratorio de Micologia Facultad de Ingenieria/ Ciencias, J. Herrera y Reissig 565 11300 Montevideo, Uruguay

Abstract—Species of Inocutis and Inonotus from Uruguay were studied. Inocutis texana comb. nov. and Inonotus ochroporus are recorded for the first time from Uruguay and compared with type and representative material. A complete description and illustration of both species are provided. Two species of Inocutis and three of Inonotus are known from Uruguay. A key for the five species is presented. The homogeneity and validity of the genus Inocutis is discussed. Sclerified hyphae from the rudimentary granular core of Inocutis jamaicensis is described for the first time.

Key Words—Basidiomycetes, Inonotus venezuelicus, polypores, taxonomy, wood-rot

Introduction

The genus Inonotus P. Karsten was proposed in 1879 to accommodate species of polypores with a pileate habit and pigmented basidiospores. Later, Donk (1933), emended the genus to encompass all the species with pigmented basidiospores and a brown context, characters present in I. cuticularis (Bull.) P. Karst., the type species (Ryvarden 1991). However, the concept of this genus was enlarged to include species with a great variation with regard to habit of the basidiocarp, shape of the basidiospores, type and pigmentation of the spore wall and presence or absence of setae. Characters that remained constant to the expanded species concept include the presence of a brown fibrous context, xanthochroic reaction of the basidiocarp when exposed to KOH or another alkali solution, and a monomitic hyphal system with simple septate hyphae.

Some characters are of limited value for delimiting polypore genera (Ryvarden 1991) and others intergrade with the closely related genus Phellinus Quélet. The distinction between the genera was reduced to the mitism of the hyphal system and the consistency and durability of the basidiocarps—dimitic (hard and perennial in Phellinus) versus monomitic, (soft and annual) in Inonotus. However, intermediate forms exist side by side in both genera (Corner 1991; Fiasson & Niemela 1984; Gilbertson & Ryvarden 1986, 1987; Larsen & Cobb-Poulle 1990).

There is growing morphological and molecular evidence to indicate that Inonotus is not a monophyletic genus. Some authors have divided Inonotus s.]. into more natural and smaller genera (Murrill 1904, Fiasson & Niemela 1984). Fiasson & Niemela (1984)

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created Inocutis to accommodate the European species of the Inonotus rheades group, or Inonotus sect. Phymatopilus (Donk 1974, Fiasson & Niemelad 1984). Based on the absence of setae and the presence of a marmorate core in the basidiocarps, Fiasson & Niemela (1984) transferred three European species of this group—I. dryophilus (Berk.) Murrill, I. rheades (Pers.) P. Karst. and I. tamaricis (Pat.) Maire—to Inocutis. In addition, these species of Inocutis have a unique pattern of pigments and sclerified hyphae in the marmorate core (Fiasson 1982, Gilbertson & Ryvarden 1986, Ryvarden & Gilbertson 1993). Recently, Wagner & Fischer (2001, 2002) showed that Inocutis is a well-supported monophyletic genus in the Hymenochaetales that encompasses the European I. rheades group along with Inonotus jamaicensis and Xanthochrous ludovicianus Pat., two American species. Gottlieb et al. (2002) arrived at the same conclusion working with internal transcribed spacer sequences.

Inonotus s. |. is poorly studied in Uruguay. Only two species are historically well known—lInonotus rickii (Pat.) D. A. Reid, found mainly on Acer and Platanus in Southern Uruguay (Bettucci & Guerrero 1971), and I. patouillardii (Rick) Imazeki, recorded on Celtis and Platanus (Gazzano 1998). Recently, Inocutis jamaicensis was identified as the causal agent of wood-rot and deformation of Eucalyptus stems in southeastern and northwestern Uruguay (Martinez 2005); it was misidentified as I. splitgerberi (Mont.) Ryvarden earlier (Martinez et al. 2002).

The main objective of the present work is to record for the first time from Uruguay two species of Inonotus s.l. recently collected, describe both species, to survey the genus for the country, and to contribute additional data to help circumscribe the genus Inocutis.

Materials and Methods

Uruguay specimens were compared with type and representative specimens from four herbaria—BAFC, BPI, MVHC and NY (Holmgren et al. 1990). Microscopic examinations were made from freehand sections mounted in 5% KOH solution and 1% phloxine, in Melzer reagent (IKI), and in 0.5% cotton blue (CB) in 25% lactophenol (Kirk et al 2001). Species were identified and compared using keys and descriptions from Pegler (1964), Urcelay & Rajchenberg (1999), Gottlieb et al. (2002) and Ryvarden (2004, 2005). Authors of fungal names are according to Kirk and Ansell (1992).

Results

Inocutis jamaicensis (Murrill) A. M. Gottlieb, J. E. Wright & Moncalvo, Myc. Prog. 1: 308. 2002. Fig. 1. Basionym: Inonotus jamaicensis Murrill, Bull. Torr. Bot. Club 31: 597. 1904. (NY!) Descriptions: Gilbertson & Ryvarden (1987), Gottlieb et al. (2002), Martinez (2005), Pegler (1964) and Rajchenberg & Wright (1998).

Remarks: For specimens studied see Martinez (2005). Inocutis jamaicensis is widely distributed in the Americas, and is reported to cause stem decay in Eucalyptus spp. in Uruguay (Martinez 2005). A character previously unreported is sclerified hyphae that are present in some sections of the context (Figure 1). Although sclerified hyphae are characteristic of the genus Inocutis, they are present only in some specimens and are easily

Figure 1. Microscopical photograph of sclerified hyphae from granular core of Inocutis jamaicensis. Scale bar = 10 um. :

broken in squash mounts. Gottlieb et al. (2002) reported the presence of a rudimentary granular core and black lines in the context. These black lines in the context can be easily verified in many specimens studied from Uruguay, but a rudimentary granular core was not observed previously (Martinez 2005). Both characters are correlated with age, habit of the basidiocarp and the way in which the specimens were collected. Most of the Uruguayan specimens come from standing eucalypt trees where the basidiocarps develop in a pileate fashion (Martinez 2005). In older specimens collected with bark, a thin zone of granulose context was observed at the basidiocarp-substrate interface. Sclerified hyphae were observed in the granular core. Thus the habit and age of the specimen has a direct bearing on whether a granular core is present and if sclerified hyphae are observed.

Inocutis texana (Murrill) Seb. Martinez, comb. nov. Fig. 2a-d. Basionym: Inonotus texanus Murrill, Bull. Torr. Bot. Club 31: 597. 1904. (NY!)

Basidiocarp annual, sessile, nodulose to ungulate, 40-70 x 25-45 x 15-50 mm. Pileus circular, glabrous, concentrically cracking in age, dark brown to almost black in old specimens. Hymenial surface creamish brown to dark yellowish brown. Context corky to fibrous, very thin, up to 5 mm thick, golden brown. Rudimentary granular core present in some specimens. When present, granular core containing sclerified hyphae, with short branches. Tubes brittle, ferruginous brown, up to 45 mm long. Pores irregular, smaller at margin, lacerate, 1-4/mm. Hyphal system monomitic, generative hyphae thin to thick walled, hyaline to reddish brown, 4-9 um diam., occasionally branched. Contextual hyphae mostly thick walled, yellowish to reddish brown, 4-10 um diam., ocasionally branched. Tramal and hymenial setae absent. Spores abundant, yellowish to golden reddish, subglobose, smooth, thick-walled, 8,0-9,0 x 6,0-7,0 um.

FIGURE 2. Microscopical characters of Inocutis texana (a-d) and Inonotus ochroporus (e-i). a. basidiospores; b. basidia; c. contextual and tramal hyphae; d. sclerified hyphae (a.-c. from MVHC 5259, d. from Holotype). e. tramal hyphae: f. basidiospores; g. hymenial setae; h. basidia; i. tramal setae. (All from MVHC 5302). Scale bars = 10 um.

5

Distribution: Argentina (Urcelay & Rajchenberg 1999, Gottlieb et al. 2002), U.S.A. (Pegler 1964, Gilbertson & Ryvarden 1986), Uruguay.

Substrates: Acacia, Diostea, Morus, Prosopis, Salix, Schinopsis (Gottlieb et al. 2002, Gilbertson & Ryvarden 1986, Pegler 1964, Urcelay & Rajchenberg 1999), Lithraea.

SPECIMENS EXAMINED: ARGENTINA. La Rioja Rio La Carpinteria, 1 km antes de Los Molinos, desde el S, on Acacia furcatispina, leg. O. Di Iorio, I-2001 (BAFC50971). CATAMARCA: San Pablo, Dpto. de Capayan, leg. V. Suarez & A. Gottlieb #53, 22-III-1995 (BAFC33667). SANTIAGO DEL EsTERO: Dpto. Copo, Reserva de Copo, leg. J. Protomastro, III-1986 (BAFC30686). San Luis, Carpinteria, on Prosopis sp., leg. O. Herrera, IV-1971 (BAFC50287).- URUGUAY. Dpto. LAVALLEJA: Minas, Paraje Campanero, on Lithraea sp., leg. G. Pérez, 18-IV-2003. (MVHC5259).- USA. Texas: Austin. on mesquite’, leg. Underwood, 24-XI-1891 (Inonotus texanus, Holotype NY!).

Remarks: This is the first report of I. texana for Uruguay. This species is well known from southwestern U.S.A. on Acacia sp. and Prosopis sp. (Fabaceae) (Gilbertson & Ryvarden 1986). Recently, Urcelay & Rajchenberg (1999) recorded this species from Argentina on Acacia sp., Prosopis sp. and Schinopsis sp. (Anacardiaceae) and Gottlieb et al. (2002) added Diostea sp. (Verbenaceae) to the host species. The Uruguayan collection is on Lithraea sp. (Anacardiaceae), and thus is able to collect this species on other substrata.

Gilbertson & Ryvarden (1986) observed a well developed granular core in this species, although Urcelay & Rajchenberg (1999) and Gottlieb et al. (2002) recorded only a rudimentary granular core in specimens from Argentina. A granular core was not observed in the Uruguayan specimen studied for it is composed of small fragments of a very young basidiocarp. The holotype is composed of small and blackened fragments with segments of different texture in the context. Under the microscope, sclerified hyphae were seen that were identical to those described by Gilbertson & Ryvarden (1986) (Figure 2d.). Sclerified hyphae were observed in some of the Argentinean specimens, but were absent in the Uruguayan specimen. Because I. texana shares with I. jamaicensis a similar hyphal system, granular core with sclerified hyphae, lack of setae, and spore type, its transfer to Inocutis is justified.

Inonotus ochroporus (Van der Byl) Pegler, Trans. Brit. Mycol. Soc. 47(2): 183. 1964. Fig. 2e-i. Basionym: Polyporus ochroporus Van der Byl., S. Afr. J. Sci. 18: 269. 1922.

Basidiocarp annual, dimidiate, sessile, 55-140 x 35-80 x 15-50 mm. Pileus circular, applanate to ungulate, single, glabrous, azonate, rusty brown to dark brown. Hymenial surface ochraceus to dark brown, blackish and radially cracked in old specimens. Context fibrous, soft, dark golden brown to chocolate brown, up to 30 mm thick. Tubes darker, up to 45 mm long. Pores irregular to angular, lacerated, 4-6/mm. Margin rounded to blunt. Hyphal system monomitic, generative hyphae thin-walled to thick-walled, hyaline to yellowish brown, 4-7 wm in diameter. Contextual hyphae thick-walled, dark reddish brown, 8-10 ym in diameter. Tramal setae brown, straight, lanceolate, mostly parallel to the tubes, 100-180 x 8-15 um. Hymenial setae rare, ventricose, 20-50 x 8-12 um. Basidia subglobose to clavate, 12-18 x 6-8 um. Spores ellipsoid, thick-walled, yellowish, 739-0.X 539-7 MM.

6

Distribution: Argentina (Gottlieb et al., 2002), Eastern Africa: Kenya, Tanzania, Uganda (Ryvarden & Johansen 1980), South Africa, West Pakistan (Pegler 1964), Uruguay. Substrates: Casuarina, Ficus, Lithraea, Salix (Gottlieb et al., 2002, Pegler 1964). SPECIMENS EXAMINED: ARGENTINA. CorriENTEs: Dpto. Capital, Riachuelo, leg. Schinini & Cristébal 12169, 27-VI-1974 (BAFC 50291). CHaco: San Bernardo, on Casuarina cunninghamiana, leg. Di Iorio (det. J. Wright & E. Blanchet), V-1995 (BAFC 33722). SAN Luts: Merlo, on Lithraea ternifolia, leg. C. Urcelay, XI-1998 (BAFC ex CORD). URUGUAY: Dpto. LAvALLEjA: Establecimiento “Santa Clara’, on living Lithraea sp., leg. S. Simeto & S. Martinez, 5-XI-2003 (MVHC 5302, 5303).

Remarks: This is the first record of I. ochroporus for Uruguay. I. ochroporus is similar to I. patouillardii, also present in Uruguay, but this last species has smaller spores measuring 5,0-7,0 x 3,5-5,0 um (Gilbertson & Ryvarden 1987). I. ochroporus is also reminiscent of I. quercustris M Blackw. & Gilb. The holotype of I. quercustris (M. Blackwell 1511 from BPI) differs in having slightly larger spores, absence of hymenial setae, and presence of conspicuous contextual hyphae with irregular wall thickening (see also Blackwell & Gilbertson 1985).

Inonotus venezuelicus Ryvarden, Mycotaxon 28: 529. 1987. SPECIMEN EXAMINED: VENEZUELA. Epo. Méripa: Laguna Negra, E. of Laguna Mucubaji, Parq. Nac. Sierra Nevada, near Apartaderos, on Polylepis sp., leg. K.P. Dumont, J.H. Haines, G.J. Samuels & A. Revas (Dumont-VE 2300), det. L. Ryvarden, 18-VII-1971 (isotype NY). Remarks: A complete description of this species is available in Ryvarden (1987, 2004). Inonotus venezuelicus was treated as a synonym of I. jamaicensis by Gottlieb et al. (2002). The isotype is a resupinate specimen, and although Ryvarden (1987, 2004) indicated the spore dimensions as 5,0-6,0 x 4,5-5,0 um, my measurements (n=30) are 5,8-7,5 x 4,6-5,8 um, slightly larger than those of I. jamaicensis. Thus, I believe this is a distinct species in the complex. Since only the type material is known, additional collections are needed to better characterize this species.

Key to the known species of Inocutis and Inonotus in Uruguay

la. Setae present in context and hymenium, granular core and sclerified hyphae absent sok MY Me ra lene yd ect Soe tke dag Sah Gree AEM me ea enc Na ect tee (Inonotus) 2

1b. Setae lacking in context and hymenium, rudimentary granular core with sclerified

hyphae present 04.24 AMeGt e. Sih ha Se eee oe cliente ctrnaeaneee (Inocutis) 4 2a. (1a) Chlamydospores present, basidiospores 6-8 x 4,5-7 um...... Inonotus rickii 2b. (1a), Chlamydospores absent j.. </.:.j:ssp aloo s/n cure ely ei ee mel ee eine 4 3a. (2b) Basidiospores ovoid 7,5-9,0 x 5,5-7,0 um .........+-- Inonotus ochroporus 3b. (2b) Basidiospores ellipsoid 5,0-7,0 x 3,5-5,0 um ........ Inonotus patouillardii

4a. (1b) Basidiocarps effusse-reflexed to triquetrous, basidiospores 5,5-7,0 x 4-5 um bit hse AUT 9th Ba SA SITS. APREN 1S eee Inocutis jamaicensis

4b. (1b) Basidiocarps dimidiate to ungulate, basidiospores 8,0-9,0 x 6,0-7,0 um. ..3 pete AT OMI ic Barca LC ade A ei RAG oye coh ies ee ae Cee Inocutis texana

Discussion

The genus Inonotus s.1. is poorly known in Uruguay with only two common species—I. rickii and I. patouillardii—reported for the country until recently/now. The genus has gained importance due to the great economic losses suffered by commercial Eucalyptus forests recently shown to be caused by Inocutis jamaicensis (Martinez 2005). The current intensive search to identify other species of Inonotus s.1. producing stem or wood rot of native or introduced trees of economic importance resulted in two new records for Uruguay: Inonotus ochroporus and Inocutis texana.

The genus Inocutis is well supported in molecular phylogenetic studies (Wagner & Fischer 2002, Gottlieb et al. 2002). Morphologically, the genus is characterized by sclerified hyphae in the context and a contextual granular core (although these structures were not observed in all the specimens studied) (Wagner & Fischer 2002). The genus also has a characteristic pigment pattern (Fiasson 1982).

This work reports for the first time the presence of sclerified hyphae in I. jamaicensis. Gilbertson & Ryvarden (1986) previously reported sclerified hyphae in I. texana. Because of its similarity to I. jamaicensis and the Inocutis rheades group, including the absence of setae, presence of a rudimentary granular core, presence of sclerified hyphae in the granular core and type of basidiospores, I. texanus is here transferred to Inocutis.

Acknowledgements

Dr. Karen Nakasone (Forest Products Laboratory, Madison, U.S.A.) and Dr. Mario Rajchenberg (CIEFAP, Chubut, Argentina) served as pre-submission reviewers. Specimens and information used in this study were loaned or put at my disposal by the curators of the following herbaria: BAFC, BPI and NY. Dra. Lina Bettucci (Montevideo, Uruguay) has suggested many improvements in a previous draft. Eufores SA and Proyecto PDT SC/OP/07/01 financially supported this work.

Literature cited

Bettucci L, Guerrero RT. 1971. Hongos xildfagos: Estudio de cultivos. Boletin de la Facultad de Agronomia (Montevideo) 118: 1-40.

Blackwell M, Gilbertson RL. 1985. A new species of Inonotus (Aphyllophorales, Hymenochaetaceae) on oak in Louisiana. Mycotaxon 23: 285-290.

Corner EJH. 1991. Ad Polyporaceas VII. The xanthochroic polypores. Nova Hedwigia. Heft 101. J Cramer: Berlin, Stuttgart (Germany). 175 pp

Donk MA. 1933. Revision der niederlandischen Homobasidiomycetae - Aphyllophoraceae II. Mededeelingen van het Botanisch Museum en Herbarium van de Rijks universiteit te Utrecht, 277p.

Donk MA. 1974. Check list of European Polypores. North Holland Publ. Co. Amsterdam, London, 469p.

Fiasson JL. 1982. Distribution of styrylpyrones in the basidiocarps of various Hymenochaetaceae. Biochemical Systematics and Ecology 10(4): 289-296.

Fiasson JL, Niemela T. 1984. The Hymenochaetales: a revision of the European poroid taxa. Karstenia 24: 14-28.

Gazzano S. 1998. Notas sobre Basidiomycetes xiléfilos del Uruguay. VIII. Registro de Aphyllophorales y sus sustratos arbéreos. Comunicaciones Botanicas del Museo de Historia Natural de Montevideo VI(109): 1-12.

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Gilbertson RL, Ryvarden L. 1986. North American Polypores Vol. I. Fungiflora: Oslo (Norway). 433 pp

Gilbertson RL, Ryvarden L. 1987. North American Polypores Vol. II. Fungiflora: Oslo (Norway). 452 pp

Gottlieb AM, Wright JE, Moncalvo JM. 2002. Inonotus s. ]. in Argentina - morphology, cultural characters and molecular analyses. Mycological Progress 1: 299-313.

Holmgren PK, Holmgren NH, Barnett LC. 1990. Index herbariorum part I. Herbaria of the world. 8th. edn. Regnum Vegetabile 120. New York Botanical Garden, New York. (http://www.nybg. org/bsci/ih/)

Kirk PM, Ansell AE. 1992. Authors of Fungal Names. Index of Fungi Supplement. CAB International: Wallingford (United Kingdom).

Kirk PM, Cannon PE, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9” Ed.. CAB International: Oxon (United Kingdom). 655 pp

Larsen MJ, Cobb-Poulle LA. 1990. Phellinus (Hymenochaetaceae). A survey of the world taxa. Synopsis Fungorum 3. Fungiflora: Oslo (Norway). 206 pp

Martinez S. 2005. Inocutis jamaicensis, the causal agent of eucalypt stem rot in Uruguay. Mycotaxon 91: 165-171.

Martinez S, Lupo S, Bettucci L. 2002. Inonotus splitbergeri (Mont.) Ryv. a stem pathogen of Eucalyptus globulus in Uruguay. Revista Brasileira de Fitopatologia 27: 420.

Murrill WA. 1904. The Polyporaceae of North America-IX. Inonotus, Sesia and monotypic genera. Bulletin of the Torrey Botanical Club 31: 593-610.

Overholts LO. 1953. The Polyporaceae of the United States, Alaska and Canada. Univ. Michigan Press: Ann Arbor (USA). 466 pp

Pegler DN. 1964. A survey of the genus Inonotus (Polyporaceae). Transactions of the British Mycological Society 47(2): 175-195.

Rajchenberg M, Wright JE. 1998. Two interesting polypore species (Hymenochaetales) from Argentina. Folia Cryptogamica Estonica 33: 119-122.

Ryvarden L. 1987. New and noteworthy Polypores from tropical America. Mycotaxon 28(2): 525- 541.

Ryvarden L. 1991. Genera of Polypores. Nomenclature and Taxonomy. Synopsis Fungorum 5. Fungiflora: Oslo (Norway). 363 pp

Ryvarden L. 2004. Neotropical Polypores part 1. Synopsis Fungorum 19. Fungiflora: Oslo (Norway). 229 pp

Ryvarden L. 2005. The genus Inonotus, a synopsis. Synopsis Fungorum 21. Fungiflora: Oslo (Norway). 149 pp

Ryvarden L, Johansen I. 1980. A preliminary polypore flora of East Africa. Fungiflora: Oslo (Norway). 636 pp

Ryvarden L, Gilbertson RL. 1993. European Polypores. Part 1. Synopsis Fungorum 6. Fungiflora: Oslo (Norway). 387 pp

Urcelay C, Rajchenberg M. 1999. Two North American Inonotus (Hymenochaetaceae, Aphyllophorales) found in Argentina. Mycotaxon 72: 417-422.

Wagner T, Fischer M. 2001. Natural groups and a revised system for the European poroid Hymenochaetales (Basidiomycota) supported by nLSU rDNA sequence data. Mycological Research 105: 773-782.

Wagner T, Fischer M. 2002. Proceedings towards a natural classification of the worldwide taxa Phellinus s.l. and Inonotus s.l., and phylogenetic relationships of allied genera. Mycologia 94: 998-1016.

MYCOTAZXON

Volume 96, pp. 9-12 April-June 2006

A new species and a new record of smut fungi from Northwestern China

SHENGRONG WANG! & CUIYUN ZENG?

' wangsr@gsau.edu.cn & * zengcy@st.gsau.edu.cn Department of Plant Pathology, College of Grassland Gansu Agricultural University, Lanzhou 730070, China

Abstract—A new species, Thecaphora oxytropis on Oxytropis ochrocephala, and a new record, Urocystis irregularis on a new host plant, Aconitum carmichaelii, are reported from Northwestern China.

Key words— Ustilaginales, taxonomy

During a mycological expedition to Tianzhu County of Gansu Province in Northwestern China in 2005, several interesting smut fungi were collected. Among them the authors identified a previously unknown species of Thecaphora on Oxytropis ochrocephala. According to Vanky (1991), eleven species of Thecaphora have been reported on Leguminosae world wide. None of them is on Oxytropis.

The authors compared the spore balls of the unknown Thecaphora sp. with those of other known species of Thecaphora on Leguminosae. Because no smut fungus known on Leguminosae has spore balls similar to those found on Oxytropis ochrocephala, we conclude that Thecaphora sp. on Oxytropis is a new species, and describe it as follows:

Thecaphora oxytropis S.R. Wang, sp. nov. Figs. 1-3 Sori in seminibus transformatis, in locum massa glomerulorum sporarum colore pallide- brunnea, granuloso-pulverea substituens. Glomeruli sporarum globosi, ellipsoidei vel leviter irregulares, 32.5-47.5 x 38.0-54.5 um, e sporis 4-21(-31) sat laxe connectis compositi. Sporae subpolyhedraliter irregulares, cuneiformis usque ellipsoidei, 16.5-22.0 x 14.0-17.0um, pallide brunneas, pariete in locis contactus levi, in locis liberibus superficiei rotundato et verrucoso.

Sori in seeds filling the pods with a light-brown, granular-powdery mass of spore balls. Spore balls globose, subglobose, ovoid, ellipsoidal or slightly irregular, 32.5-47.5 x 38.0-54.5 um, composed of 4-21(-31) easily separating ustilospores. Ustilospores subpolyhedral, cuneiform, ovoid, ellipsoidal, polyhedral or slightly irregular, 16.5-22.0 x 14.0-17.0um, yellowish to light yellowish-brown, wall smooth on the contact surfaces, coarsely and irregularly verrucose on the free surface. Germination unknown.

On Oxytropis ochrocephala Bunge (Leguminosae), China, Gansu: Tianzhu County,

Zhuaxi, alt. ca. 3100 m, 20 VIII 2005, coll. S.R. Wang & C.Y. Zeng. GAU 05011

(holotypus), HUV 21285 (isotypus). Known only from the type collection.

10

Thecaphora oxytropis and T: hedysari Vanky have loose spore balls, but they differ in that the former has smaller spore balls and fewer spores in a ball.

A species of Urocystis on Aconitum carmichaelii was also collected by the authors. After examination it turned out to be Urocystis irregularis. According to Vanky (1994), on Aconitum only one species of Urocystis is known, U. irregularis, occurring in Europe, Asia, and North America. It has not yet been reported from China nor anywhere previously on A. carmichaelii (Ling 1953, Guo 2000a,b, 2003, Guo & Zhang 2004, Guo & Wang 2005, Wang & Piepenbring 2002, Wang & Guo 2002, Vanky & Guo 1985, Kakishima et al. 2000). Thus, the collection represents a new record for China on a new host plant:

Urocystis irregularis (G. Winter) Savul. Bul. Sti. Acad. Rep. Populare Romane 3:220, 1951. Figs.4-6 = U. sorosporioides f. irregularis G. Winter, Hedwigia 19:2, 1880.

Sori on the abaxial side of leaves, on petioles and stems forming pustules, sometimes confluent, or in the veins as long swellings, at first covered by the epidermis subsequently rupturing to expose the black, powdery mass of spore balls. Spore balls subglobose to irregular, 26.0-37.5 x 21.0-28.5 um in diam., composed of 1-6 ustilospores, surrounded by an incomplete layer of peripheral sterile cells. Spores subglobose, elongated, irregular to slightly polyhedral, 13.5-18.0 x 9.5-13.0 um in diam., olive-brown to yellowish-brown, smooth or slightly granular, with irregular small thickenings on the free surface. Sterile cells globose, subglobose, ovoid to irregular, 8.5-12.0 x 4.0-8.5 um, light yellowish- brown, smooth, hyaline.

On Aconitum carmichaelii Debeaux (Ranunculaceae), China, Gansu: Tianzhu County, Tanshanling, alt. ca. 2890 m, 18 VIII 2005, coll. S.R. Wang & C.Y. Zeng. GAU 05015.

Acknowledgements

The authors would like to express their deep thanks to Dr. K. Turkington and Dr. D.J. Bing from Agriculture and Agri-Food Canada (Lacombe, Canada) for reading the manuscript and improving the English, to Dr. K. Vanky (Tiibingen, Germany) and Prof. M. Piepenbring (Frankfurt/M, Germany) for reading the manuscript and serving as pre-submission reviewers, to Fang Qiangen for identifying the host plants, to Xu Changlin for assistance in the field collection, and to Xu Bing and He Dongyun for assistance in the laboratory experiments. This research was partly supported by the Gansu Natural Science Foundation (No. ZR-97-041; No. 4RS045-A65-038).

Literature Cited

Guo L. 2000a. Flora Fungorum Sinicorum, Ustilaginaceae Vol. 12, Science Press, Beijing. 1-124.

Guo L. 2000b. Sporisorium pollinianum, a smut species new to China. Mycosystema 19(3): 420- 421.

Guo L. 2003. Smut fungi in Qinghai. Mycosystema 22 ( Suppl): 112-116.

Guo L, Zhang HC. 2004. A new species and two new records of Ustilaginomycetes from China. Mycotaxon 90: 387-390.

Guo L, Wang SR. 2005. A new species and a new record of Anthracoidea (Ustilaginales) from China. Mycotaxon 93: 159-162

Nt

Figs. 1-3. Thecaphora oxytropis on Oxytropis ochrocephala (holotypus). Fig. 1. Spore balls as seen by LM (light microscopy;). Fig. 2. Spore balls as seen by SEM (scanning electron microscopy). Fig. 3. Spore ball as seen by SEM. Figs. 4-6. Urocystis irregularis on Aconitum carmichaelii (GAU 05015).

Fig. 4. Spore balls as seen by LM. Fig. 5. Spore balls as seen by SEM. Fig. 6. A spore ball as seen by SEM Bars = 10 um

i

Kakishima M, Denchev CM, Zhou X, Zang M. 2000. Smut fungi from Yunnan province, China, collected in 1998. Bull. Nat. Sci. Mus., Tokyo, Ser. B, 26: 23-34.

Ling L. 1953. The Ustilaginales of China. Farlowia 4: 305-351.

Savulescu T.1951. Noutati din micoflora R.P. Romane. Bul.Sti.Acad. Rep. Populare Romane 3:211- 227,

Vanky K. 1991. Thecaphora (Ustilaginales) on Leguminosae. Trans. Mycol. Soc. Japan 32: 145-159.

Vanky K. 1994. European Smut Fungi. Gustav Fischer Verlag, Stuttgart, Jena, New York. 1-570.

Vanky K, Guo L. 1985. Ustilaginales from China. Acta Mycologica Sinica 5(Suppl): 234-235.

Wang SR, Piepenbring M. 2002. New species and new records of smut fungi from China. Mycological Progress 1(4): 399-418.

Wang SR, Guo L. 2002. A new record of smut fungi from China. Mycosystema 21(3): 452-453.

Winter G. 1880. Mycologische Notizen. Hedwigia 19:1-4.

MYCOTAXON

Volume 96, pp. 13-19 April-June 2006

Additional and interesting lichenized and lichenicolous fungi from Turkey

MEHMET GOKHAN HALICI

mghalici@erciyes.edu.tr Biology Department, Faculty of Science & Art, Erciyes University Kayseri, Turkey

MUSTAFA KOCAKAYA

kocakaya38@mynet.com Ahmet Gazi Ayhan Bulvari, Kazimkarabekir Mah Silvan Gegidi No:24 Kayseri, Turkey

AHMET AKSOY

aksoy@erciyes.edu.tr Biology Department, Faculty of Science & Art, Erciyes University Kayseri, Turkey

Abstract— 14 species of lichenized and lichenicolous fungi are reported from the Turkish provinces of Kayseri, Nigde, Adana, Yozgat and Mugla. Nine taxa, Anisomeridium viridescens, Cercidospora epicarphinea, Lichenodiplis lecanorae, Lichenothelia convexa, Pilophorus cereolus, Porpidia superba, Sagediopsis campsteriana, Toninia verrucarioides and Zwackhiomyces sphinctrinoides are new for Turkey. Comments on habitat and substrata and a short description are provided for each taxon.

Key words— Ascomycota, conidial fungi, lichens

Introduction

Serious lichenology in Turkey started in last two decades. A total of 360 papers actually refer to lichens from Turkey (John 2004). If floristic, these mostly focus on the lichens of the Black Sea region (e.g. John & Breuss 2004, Yildiz et al. 2002, Yazici1 1999, Ozdemir Tiirk 1997) and the Mediterranean region (e.g. John 2003, John & Nimis 1998, Nimis & John 1998, John, 1996), but there have been few lichenological studies in Central Anatolia (e.g. Halici et al. 2005, Karabulut & Tiirk 1998, Ozdemir 1991). This contribution reports further species as first records for Turkey or the province of Kayseri, Mugla or Yozgat.

Materials and Methods

The specimens on which this paper is based are all preserved in the herbarium of Erciyes University, Biology Department, Science & Art Faculty, Kayseri, Turkey. The accession numbers of the collections are given in parentheses after the locality details.

14

The specimens were first examined with a dissecting microscope, and then with an Olympus BH2 microscope fitted with Nomarski differential interference contrast. Specimens were examined in water, 10% KOH, Lugols iodine solution or lactofuchsin. Spore measurements were generally made in KOH, but in the case of thin walled spores these were also checked in water.

Species recorded

Anisomeridium viridescens (Coppins) R.C. Harris

A detailed description is provided by Purvis et al. (1992: 92, as Arthopyrenia viridescens).

The Turkish specimen was collected from bark of Cedrus libani associated with Caloplaca cerina var. cerina and C. flavorubescens in ancient woodland. ‘The thallus is lichenized with Trentopohlia. Ascomata are perithecia, involucrellum brown, K + green. Asci 8- spored. Ascospores colourless, 1-septate, the cells usually biguttulate, 12.5-14.5 x 4-5 tum.

New record for Turkey. Known also from the British Isles on smooth bark of Corylus in old woodlands (Purvis et al. 1992).

Kayser: Yahyali, Aladaglar Milli Parki, Hacer Ormanlari, 37° 48.152’ N, 35° 16.870’ E,

alt. 1690 m, on bark of Cedrus libani, 20 Sep.2004, leg. M. G. Halici & A. Aksoy, det. D.

L. Hawksworth (MGH 0.2275).

Bacidia laurocerasi (Delise ex Duby) Zahlbr. A detailed description is provided by Purvis et al. (1992: 107).

The Turkish specimen was collected from the bark of Pinus nigra growing with macrolichens such as Anaptychia ciliaris, Bryoria fuscescens and Parmeliopsis ambigua in ancient woodland. ‘Thallus very thin. Apothecia ~ 1 mm; numerous, blackish. Asci 8-spored. Ascospores 30-38 x 3-4 um, 8- to 13-septate at maturity, acicular.

In Turkey, previously recorded from Mugla (John 1996, Nimis & John 1998) and Sakarya (Szatala 1927 as B. endoleuca). New record for Kayseri province.

Known also from the British Isles, Europe, North America, Australia and New Zealand on + basic or nutrient rich bark of trunks and branches of trees and shrubs, usually in open situations or secondary woodland (Purvis et al. 1992).

KayseRI: Yahyali, Aladaglar Milli Parka, Hacer Ormanlani, 37’ 47.935’ N, 35° 18.136’ E,

alt. 1648 m, on bark of Pinus nigra, 20 Sep.2004, leg. M. G. Halici & A. Aksoy, det. D. L.

Hawksworth (MGH 0.2276).

Buellia erubescens Arnold A detailed description is provided by Purvis et al. (1992: 113).

The Turkish specimen, collected from + smooth bark of Abies in ancient woodland, was associated with Parmelina pastillifera, Anaptychia ciliaris and Lecidella elaeochroma. Thallus thin; whitish, K + red, PD + orange, C -. Apothecia ~ 1 mm, heavily pruinose when young. Asci 8-spored. Ascospores first colourless, then brown, 2-celled; 12.5-15 X5.6-7.5 pm.

15

Previously recorded from six provinces in Turkey (John & Nimis 1998, Huneck et al. 1992, Yildiz & John 2002, John & Breuss 2004, Steiner 1909, Ozdemir 1990), sometimes as B. zahlbruckneri. New record for Kayseri province.

Known also from the British Isles, western and central Europe, the Azores and North America, on + smooth bark (Purvis et al. 1992). KaysERI: Yahyali, Aladaglar Milli Parki, Hacer Ormanlani, 37° 47.935’ N, 35° 18.136’ E, alt. 1648 m, on bark of Abies, 20 Sep.2004, leg. M. G. Halici & A. Aksoy, det. D. L. Hawksworth (MGH 0.2277).

Cercidospora epicarphinea (Nyl.) Grube & Hafellner A detailed description is provided by Grube & Hafellner (1990). The Turkish specimen grows on the apothecia of an unidentified Caloplaca sp. and appears to suppress ascospore production in the host. Interascal tissues (hamathecium) present. Perithecia immersed, 150-180 um diam, peridial wall blue-green. Asci 8-spored. Ascospores 16.5-20 x 4.5-6 um, 1-septate, colourless. New record for Turkey. Known also from Spain and Canary Islands on Caloplaca carphinea and C. scoriophila (Grube & Hafellner 1990).

Kayseri: Erciyes Mountain, Exit of Hisarcik, Western part of the highway, 38°36’ N, 35°

30’ E, alt. 1540 m, on the apothecia of Caloplaca sp. on siliceous rocks, 25 Aug.2002, leg. M. G. Halici & M. Kocakaya, det. D. L. Hawksworth (MGH 0.2278).

Leptorhaphis atomaria (Ach.) Szatala A detailed description is provided by Aguirre-Hudson (1991: 99).

The Turkish specimen, collected from the bark of Populus nigra, was associated with Lecanora albella. Thallus smooth, very thin; weakly lichenized with Trentopohlia. Asci 8-spored; ascospores arranged in two bundles in the asci, colourless, 1-septate, filiform, crescent shaped, 20-22 x 2.5-4 um.

Previously recorded from Istanbul by Steiner (1889). New record for Kayseri province.

Widespread in temperate regions of Europe on bark of Fraxinus excelsior and also particularly on species of Populus, growing on the trunk, often mixed with lichens such as Xanthoria parietina and species of Caloplaca and Lecanora; perhaps facultatively lichenized with trentepohlioid algae or not, it has also been mentioned from Northern America (Aguirre-Hudson 1991).

KayserI: Yahyali, Aladaglar Milli Parki, Hacer Ormanlani, 37 47.935’ N, 35° 18.136’ E,

alt. 1648 m, on bark of Populus nigra, 20 Sep.2004, leg. M. G. Halici & A. Aksoy, det. D.

L. Hawksworth (MGH 0.2279).

Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw. A detailed description is provided by Hawksworth (1981).

The Turkish specimen, on the apothecia of Caloplaca holocarpa, was collected from the bark of Populus nigra; it seems to be commensalistic as there was no damage to the thallus, and ascospore production was not suppressed. Conidia pale brown to dark brown, arising singly, 1-septate, 4.5-5 x 2-2.5 um diam.

16

Commensalistic to parasitic on the apothecia, and more rarely the thalli, of Caloplaca, Lecanora and Pertusaria species, also on thalli of Evernia prunastri, Lecidea erratica and Schismatomma decolorans.

New record for Turkey. Widespread in Europe, the former U.S.S.R, and also known from Morocco (Hawksworth 1981).

KayseRI: Yahyali, Aladaglar Milli Parki, Hacer Ormanlari, 37° 47.906’ N, 35° 17.889’ E,

alt. 1595 m, on the apothecia of Caloplaca holocarpa on bark of Populus nigra, leg. M. G. Halici & A. Aksoy, det. D. L. Hawksworth, (MGH 0.2280).

Lichenostigma maureri Hafellner A detailed description is provided by Hafellner (1982).

The Turkish specimens, on Usnea subfloridana and Pseudevernia furfuracea, were collected from Pinus sylvestris. Commensalistic, since no damage observed in the host thallus. Ascomata internal structure paraplectenchymatic stromatic, non-aggregated. Ascospores brown, echinulate, upper cell wider; 10 x 5 um.

Previously recorded from Turkey by Hafellner (1982). New record for Mugla and Yozgat provinces.

Known also from Spain, the Canary Islands and Great Britain on Usnea species and Pseudevernia furfuracea (Hafellner 1982, Hawksworth 1982).

Muéza: Yaras Village, Picnic Area of Yaras Forest (20 km to the city center), 37 09.128” N, 28° 29.918’ E, alt. 750 m, on Pseudevernia furfuracea on Pinus sylvestris, 23 Aug.2004, leg. M. G. Halici & M. Kocakaya, det. D. L. Hawksworth, (MGH 0.2289).

YozGat: Akdagmaden District, Between Biiyiik Nalbant Mountain and Kizilkoca Village, Cat Ormanlari, 39°30’ N, 35° 59’ E, alt. 1780 m, on Usnea subfloridana on Pinus sylvestris, leg. M. G. Halici & M. Kocakaya, det. D. L. Hawksworth, (MGH 0.2281).

Lichenothelia convexa Henssen A detailed description is provided by Purvis et al. (1992: 359-360).

The Turkish specimen was collected from bare rock and sometimes on Aspicilia caesiocinerea. This material lacked ascospores, but has macroconidia, 10-20 um diam., agreeing with those illustrated and described by Hennsen (1987).

New record for Turkey. Known also from the British Isles, Germany, Sweden and North America on slate, intermixed with Rhizocarpon geographicum thalli, and on sarsens (Purvis et al., 1992).

ADANA: Aladaglar Milli Parki, Turhasan Mountain, 37° 45.280 N, 35° 18.028’ E, alt. 2000 m, on bare siliceous rocks and Aspicilia caesiocinerea, leg. M. G. Halici & A. Aksoy, det. D. L. Hawksworth, (MGH 0.2282).

Pilophorus cereolus (Ach.) Th.Fr. Detailed descriptions are provided by Jahns (1981) and Brodo et al. (2001).

The Turkish specimen was collected on calcareous soil. Primary thallus greyish, sorediate. No apothecia. Thallus and soralia K + yellow, PD -.

7

New record for Turkey. Known also from many localities in Europe and North America on weathered rocks or soil (Jahns, 1981, Brodo et al. 2001). NiGpE: Camardi, Aladaglar Milli Parki, Entrance of Demirkazik, 37° 51.050’ N, 35° 07.100’ E, alt. 1900 m, on calcareous soil, 29 Aug.2005, leg. M. G. Halici & M. Kocakaya, det. D. L. Hawksworth, (MGH 0.2283).

Porpidia superba (Korb.) Hertel & Knoph A detailed description is provided by Gowan & Ahti (1993: 69).

The Turkish specimen was collected from calcareous rocks. Thallus cream-white, thick, forming knob-like bullae, K-, C-. Apothecial disks brown, concave, heavily pruinose. True exciple well developed, black. Asci 8-spored. Ascospores colourless, simple; 15-22 x 9-10 um. New record for Turkey. Known also from Finland, Great Britain and Russia on calcareous and also on siliceous rocks, particularly schists (Gowan & Ahti 1993).

Kayseri: Yahyali, Aladaglar Milli Parki, Delikkaya, 37°54.948’ N, 35° 15.506’ E, alt. 2910

m, on calcareous rock, 3 Oct.2004, leg. M. G. Halici & A. Aksoy, det. D. L. Hawksworth,

(MGH 0.2284).

Rhizocarpon simillimum (Anzi) Lettau A detailed description is provided by Purvis et al. (1992: 541).

The Turkish specimen was collected from siliceous rocks. Thallus dark grey, areolate; areoles bullate. Asci 8-spored, ascospores brown, 1-septate; 12-13 x 6-7 um.

In Turkey, previously recorded from Mudanya (Verseghy 1982). New record for Kayseri province.

Known also from the British Isles and Norway on sunny, acid, + vertical outcrops (Purvis et al. 1992). KAYSERI: Erciyes Mountain, North of Perikartin (Northern slope of Erciyes Mountain),

38° 35’ N, 35° 27’ E, alt. 2300 m, on siliceous rock, 3 Oct.2004, leg. M. G. Halici & M. Kocakaya, det. D. L. Hawksworth, (MGH 0.2285).

Sagediopsis campsteriana (Linds.) D. Hawksw. & R. Sant. A detailed description is provided by Triebel (1993).

The Turkish specimen, on Aspicilia desertorum, was collected from calcareous rocks. Ascomata perithecia, arising singly, immersed at the base and appearing largely superficial, black, not shiny, 100-250 um diam; ostiolate, the ostiole not papilliform or raised in any way. Asci elongate-clavate, bitunicate in structure, discharge not seen, thickened at the apex, about 65-70 x 11-13 um, 8-spored. Ascospores overlapping and distichously arranged in the ascus, broadly fusiform, apices generally somewhat attenuated towards the apices but occasionally more rounded, colourless, (1-)3-septate, not constricted at the septum, the cells often with conspicuous rounded vacuoles, smooth walled, lacking a defined gelatinous sheath, 12-21 x 4.5-7 um, 1/b ratio 2-3.5.

New record for Turkey. Known also from Europe and North America on several Ochrolechia species and Lecanora hagenii (Triebel 1993).

18

KaysERI: Yahyali, Aladaglar Milli Parki, Gokoluk Mevkii, Dedegedigi gecidi, 37°55.392’ N, 35° 17.376’ E, alt. 2430 m, on Aspicilia desertorum on calcareous rocks, 2 Aug.2005, leg. M. G. Halici & A. Aksoy, det. D. L. Hawksworth, (MGH 0.2286).

Toninia verrucarioides (Nyl.) Timdal A detailed description is provided by Timdal (1991: 117).

The Turkish specimen was collected from calcareous rocks in a shaded habitat in a forest. Thallus squamulose. Asci 8-spored. Ascospores colourless, ellipsoid to bacilliform, 1- to 3-septate; 9.5-15 x 4-5 um.

New record for Turkey. Known from temperate and arctic parts of Europe and North

America on calciferous rocks, usually in fissures but also on smooth faces or on soil- covered rock (Timdal 1991).

MuGia: Yaras Village, Picnic Area of Yaras Forest (20 km to the city center), 37°09.128’ N, 28° 29.918’ E, alt. 750 m, on calcareous rock, 23 Aug.2004, leg. M. G. Halici & M. Kocakaya, det. D. L. Hawksworth, (MGH 0.2287).

Zwackhiomyces sphinctrinoides (Zwackh) Grube & Hafellner A detailed description is provided by Grube & Hafellner (1990).

The Turkish specimen, on Xanthoria elegans, was collected from calcareous rocks. Ascomata perithecia; pyriform, 150-250(-270) um diam; peridial wall brown with basal cells paler than the upper ones. Asci 8- spored. Ascospores colourless, 1-septate, verruculose, 18-20 x 6.5-9 um.

New record for Turkey. Widespread in Great Britain and Spain on Collema species, Clauzadea metzleri, Leptogium turgidum, Caloplaca variabilis, Solenopsora olivacea and other calcicolous lichens (Grube & Hafellner 1990).

Kayseri: Yahyali, Aladaglar Milli Park, Gékoluk Mevkii, Dedegedigi gecidi, 37°55.392’

N, 35° 17.376 E, alt. 2430 m, on Xanthoria elegans on calcareous rocks, leg. M. G. Halici

& A. Aksoy, det. D. L. Hawksworth, (MGH 0.2286).

Acknowledgements

We are grateful to Prof. David L. Hawksworth for the identification of the specimens, reviewing a draft of this paper, and for opening his laboratory, files and library to the first author. Dr Volker John and Prof. Mark R.D. Seaward are thanked for critically reviewing the paper and for most helpful suggestions for its improvement. This study was partly supported by TUBITAK.

Literature Cited

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Brodo M, Sharnoff SD, Sharnoff S. 2001. Lichens of North America. New Haven, Yale University Press.

Gowan SP, Ahti T. 1993. Status of the lichen genus Porpidia in eastern Fennoscandia. Ann. Bot. Fennici 30: 53-75.

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Grube M, Hafellner J. 1990. Studien an flechtenbewohnenden Pilzen der Sammelgattung Didymella (Ascomycetes, Dothideales). Nova Hedwigia 51: 283-360.

Hafellner J. 1982. Studien tiber lichenicole Pilze und Flechten II. Lichenostigma maureri gen. et spec. nov. ein in der Ostalpen haufiger lichenicoler Pilz (Ascomycetes, Arthoniales). Herzogia 6: 299-308.

Halici MG, John V, Aksoy A. 2005. Lichens of Erciyes Mountain (Kayseri, Turkey). Fl. Medit. 15: 567-580.

Hawksworth DL. 1981. The lichenicolous coelomycetes. Bull. Br. Mus. nat. Hist., Bot. 9: 1-98.

Hawksworth DL. 1982. Notes on British lichenicolous fungi IV. Notes R. bot. Gdn Edinb. 40: 375- SLI

Huneck §S, John V, Jakupovic J, Elix JA. 1992. Zur Chemie einiger Strauch- und Krustenflechten aus der Tiirkei. Herzogia 9: 489-492.

Jahns HM. 1981. The genus Pilophorus. Mycotaxon 13: 289-330.

John V. 1996. Preliminary catalogue of lichenized and lichenicolous fungi of Mediterranean Turkey. Bocconea 6: 173-216.

John V, Nimis PL. 1998. Lichen flora of Amanos Mountain and the province of Hatay. Tr. J. of Botany 22: 257-267.

John V. 2003. Flechten aus der Tirkei, von G.Ernst gesammelt, Herzogia 16: 167-171.

John V. 2004. Lichenological studies in Turkey and their relevance to environmental interpretation.- Abstract book, XI OPTIMA meeting, 5.-11.9.2004 Belgrad: 45.

John V, Breuss O. 2004. Flechten der dstlichen Schwarzmeer-Region in der Turkei (BLAM- Exkursion 1997). Herzogia 17: 137-156.

Karabulut B, Ozdemir Tiirk A. 1998. Lichens of the Aksehir District (Konya), Tr. J. of Botany 22: 191-198.

Nimis PL, John V. 1998. A Contribution to the Lichen Flora of Mediterranean Turkey. Cryptogamie, Bryol. Lichénol. 19(1): 35-58.

Ozdemir A. 1991. Eskisehir [li Likenleri. Tr. J. of Botany 15: 189-196.

Ozdemir Tirk A. 1997. A study on the lichen flora of Sinop and Kastamonu provinces. J. Fac. of Sci. Ege Univ. 20: 221-229.

Steiner J. 1899. Flechten. In: Fritisch C., Beitrag zur Flora von Konstantinopel. I. Kryptogamen, Denkschr.k. Akad. Wiss.,math-naturw.Cl. Wien 48: 222-238. Steiner J. 1909. Lichenes. In: Handel-Mazetti DHF von.: Ergebnisse einer bott. Reise in d. Pontische Randgebirge im Sandschak Trapezunt, etc.- Annal. Naturhist. Hofmus. Wien 23: 107-123. Szatala O. 1927. Lichenes Turcicae asiaticae a patre Prof. Stefano selinka in insula Burgas Adassi (Antigoni) lecti. Magy. Bot Lapok 26: 18-22.

Purvis OW, Coppins BJ, Hawksworth DL, James PW, Moore DM. 1992. The Lichen Flora of Great Britain and Ireland. London, Natural History Museum Publications.

Timdal E. 1991. A monograph of the genus Toninia (Lecideaceae, Ascomycetes). Opera Botanica 110: 1-137.

Triebel D. 1993. Notes on the genus Sagediopsis (Verrucariales, Adelococcaceae). Sendtnera 1: 273- 280.

Verseghy KP. 1982. Beitrage zur Kenntnis der tiirkischen Flechtenflora. Studia Botanica Hungarica 16: 53-65.

Yazici K. 1999. Lichen flora of Trabzon. Tr. J. of Botany 23: 97-112.

Yildiz A, John V, Yurdakulol E. 2002. Lichens from the Cangal Mountains (Sinop, Turkey). Cryptogamie, Mycologie 23(1): 81-88.

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MYCOTAXON

Volume 96, pp. 21-29 April-June 2006

Four species of Exobasidium from Korea

JI-HYUN PARK’, KYUNG-HEE KIM”*, KYUNG-JOON LEE’ & SEUNG-KyvU LEE?

"Department of Forest Sciences, Seoul National University Seoul 151-921, Korea

*kyung624@foa.go.kr "Department of Forest Environment, Korea Forest Research Institute Seoul 130-712, Korea

Abstract—Four species of Exobasidium collected in Korea are reported with descriptions and host plants. They are E. dubium, E. yoshinagai, E. japonicum and E. cylindrosporum. Of these, E. dubium, E. yoshinagai and E. cylindrosporum are newly added to the fungus flora of Korea.

Key words—Exobasidium diseases, Rhododendron

Introduction

The genus Exobasidium belongs to Basidiomycota and contains more than 100 described taxa (Alexopoulos et al. 1996, Nagao et al. 2004a). They are all plant pathogens causing leaf galls, blight, blisters, witches’ broom, dwarfing or abnormal bud proliferation in field trees and ornamentals (Coyier & Roane 1986; Ing 1998, 1999). The fungus flora in Korea is still very poorly known and has not been investigated in many regions of the country. In some phytopathological papers (cf. Korean Society of Plant Pathology 2004), two Exobasidium species parasitic on three host plants were listed and additional one unidentified species was recorded to be associated with leaf galls on Rhododendron schlippenbachii. Therefore, mycological studies for the Korean Exobasidium spp. are urgently needed for diagnosis of the diseases and precise identification of the fungus. In the present paper we described morphological characteristics and host plants of four Exobasidium species based on specimens collected in Korea.

Materials and Methods The materials listed in the present paper were collected from May to June, 2004 and were deposited in KFRI (Korea Forest Research Institute). Identification of each fungus was based on microscopic as well as cultural characteristics. Leaves with newly sporulating lesions were cut into small pieces and fixed to the inside of the lid of a

*Corresponding author.

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sterile Petri dish, poured with potato dextrose agar (PDA) acidified with 10% (v/v) lactic acid. The dish was kept at 23°C in the dark. The lid of the Petri dish was turned at two hours intervals. Basidiospores then fell down from the hymenium onto the agar surface. After microscopic examination, basidiospore was isolated from the dish, and 18 isolates were transferred to new PDA dishes for further growth. The cultures were incubated in an unlighted room at 23°C. Slides were prepared on Shear’s mounting fluid for morphological observation under light microscope. JSM-5410LV SEM was used for the observation of basidia and sterigmata. Additional anatomical observations using paraffin infiltration method were conducted to study the comparative anatomy of infected areas by Exobasidium species. Ezuka (1974a, b, 1990a, b, 1991a, b, 1992a, b, c), Ito (1955), Karatygin (2000), Karatygin & Huseyin (2002), Nannfeldt (1981), Nagao et al. (2001, 2003a, b, 2004a, b), Otani (1976), Sundstrém (1964) and Zhang et al. (1995) were consulted to aid in species identification.

Descriptions

1. Exobasidium dubium Racib., Kosmos 34: 1172 (1909). Fig. 1

DESCRIPTION: Germ tubes of the basidiospores emergent from both end cells at first then from each septal regions; conidia produced at the tip of germ tubes or laterally, budding to produce daughter cells polarly, developing into hyphae; colonies on PDA growing gradually, toa maximum 7 mm diam. in 21-day incubations, gelatinous, fixed on the agar surface, composed of branching and intricate hyphae, pseudohyphae and conidia; surface yellowish white to pale yellow, corrugate in center, not farinose by conidial formation; dark pigmentation absent.

Basidia 17-31 x 5.5-10.0 um, clavate to cylindrical, apices obtuse, (2-)3-4(-5)-spored, directly emergent from the host surface or through stomata, not fasciculate; sterigmata 3-7 um long, 1.5-2.0 um diam. at the base, outwardly emergent, tapering toward the tip; basidiospores 13-23 x 3.3-8.0 um, subcylindric to musiform, straight to slightly curved, becoming slender in center and septal regions, hyaline, smooth, unicellular when formed, becoming 1-4(-5)-septate when germinating; conidia 5-15 x 1.3-3.6 ym, linear, acicular, sometimes 1-septate.

SYMPTOMATIC CHARACTERISTICS: Flat lesions on leaves, up to 15 mm across, usually 1 to 5 lesions on one leaf, circular to irregular in outline, sometimes confluent to develop larger, pale yellow to yellow above, becoming reddish or brownish in center, sometimes with conspicuous red margin above; the lower leaf surface of the infected area eventually with a thin white layer of hymenium; mycelial mat localized between the lower epidermis and the first layer of spongy mesophyll; infected parts not thickened; hypertrophy and hyperplasia absent.

Hapsirat: On living leaves of Rhododendron yedoense f. poukhanense (H. Lév.) M. Sugim. and R. yedoense Maxim. f. yedoense.

SPECIMENS EXAMINED: On R. y. f. poukhanense, KFRI2237, 2238, 2239, 2240, 2241, 2242

(3 June, 2004, Seoul), KFRI2243, 2244, 2245, 2246 (5 June, 2004, Pocheon-si), KFRI2247

(24 June, 2004, Seoul); On R. y. f. yedoense, KFRI2248 (3 June, 2004, Seoul), KFRI2249

(24 June, 2004, Seoul).

25

Fig. 1. Exobasidium dubium. A. A colony on PDA, 16 weeks, KFRI2244. B. Germination of basidiospores on PDA, Arrow shows a long germ tube, KFRI2237. C. A basidium with four immature basidiospores, Arrows show sterigmata, KFRI 2243. D. Three basidiospores before septation and two spores becoming 3-septate, KFRI2249. E. Conidia, KFRI2249. EF. Flat lesions with conspicuous red margin on adaxial leaf surface in a naturally infected host. G. A cross section of healthy part of infected leaf, KFRI2249. H. A cross section of infected part, Arrows show mycelial mats in abaxial side, KFRI2249. Scale bars: B = 10 um, C = 5 um, D-E = 10 um, G-H = 20 um.

Notes: This is the first record on this species in Korea. This fungus was recorded on R. luteum from Europe and on Rhododendron sp. from Turkey (Nannfeldt 1981, Karatygin & Huseyin 2002). Recently, Nagao et al. (2003a) discussed this fungus on leaves of R. y. f. yedoense in Japan. The present study adds R. y. f. poukhanense and R. y. £. yedoense as new host records to Korea. Although the Korean specimens did not show colonies with pale pink described by Nagao et al. (2003a), the other taxonomic characteristics of this species are in close agreement with the previous records.

2. Exobasidium yoshinagai Henn., Bot. Jahrb. 31: 736 (1902). Fig. 2

DESCRIPTION: Germ tubes of the basidiospores emergent from each cell, developing into long and branched hyphae; conidia produced at the tip of germ tubes or laterally, budding to produce daughter cells polarly, developing into hyphae; colonies on PDA growing gradually, to a maximum 4 mm diam. in 21-day incubations, gelatinous, fixed on the agar surface, composed of branching and intricate hyphae, pseudohyphae and conidia; surface yellowish white, slightly corrugate in center, not farinose by conidial formation; dark pigmentation absent.

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Fig. 2. Exobasidium yoshinagai. A. A colony on PDA, 16 weeks, KFRI2250. B. A basidium with four immature basidiospores, Arrows show sterigmata, KFRI2252. C. Strongly geniculate basidiospores towards the hilum before septation, KFRI2252. D. Flat lesions without conspicuous red margin on adaxial leaf surface in a naturally infected host. E. A cross section of healthy part of infected leaf, KFRI2250. EF. A cross section of infected part, Arrows show mycelial mats in abaxial side, KFRI2250. Scale bars: B = 5 um, C = 10 um, E-F = 20 wm.

Basidia 20-34 x 5.4-9.1 um, clavate to cylindrical, apices obtuse, 2-4-spored, directly emergent from the host surface or through stomata, not fasciculate; sterigmata 3-8 um long, 1.5-2.1 um diam. at the base, outwardly emergent, tapering toward the tip; basidiospores 13-22 x 4.2-6.5 um, cylindrical, slightly curved, strongly geniculate towards the hilum, hyaline, smooth, unicellular when formed, becoming (1-)3-septate when germinating; conidia 5-10 x 1.4-2.9 um, linear to acicular.

SYMPTOMATIC CHARACTERISTICS: Flat lesions on leaves, up to 49 lesions under 5 mm across on one leaf, sometimes 1 to 4 lesions ranging from 7 to 14 mm across on one leaf, circular in outline, sometimes confluent to develop larger, pale yellow to yellow above, without conspicuous margin above; the lower leaf surface of the infected area eventually with a thin white layer of hymenium; mycelial mats localized in the abaxial side of the lesions; infected parts not thickened; hypertrophy and hyperplasia absent.

Hasitat: On living leaves of Rhododendron schlippenbachii Maxim.

SPECIMENS EXAMINED: On R. schlippenbachii, KFRI2250, 2251 (3 June, 2004, Seoul), KFRI2252 (5 June, 2004, Pocheon-si).

25

Notes: Earlier literature (Ezuka 1991a) gave relatively long measurements of basidia, in the range of 50-70 um. In the Korean materials, they are much shorter (20-34 um). It may be due to the different criteria for measurement of the length of basidia that develop from mycelium beneath the epidermis. In addition, Ezuka (1991a) commented that the number of leaf lesions by this fungus was under ten on one leaf, but E. yoshinagai was occasionally found to form abundant small lesions (under 5 mm) on one leaf of R. schlippenbachii in this study. The other characteristics of taxonomic value match well each other. This is the first record on the fungus in Korea. Ezuka (1974a, 1991a) reported this species as a causal agent of the leaf blisters of R. dilatatum, R. kiyosumense, R. reticulatum, R. wadanum, R. viscistylum var. glaucum and R. weyrichii from Japan. Rhododendron schlippenbachii was recognized as a new host record to Korea.

3. Exobasidium japonicum Shirai, Bot. Mag. Tokyo 10: 52-53 (1896). Fig. 3

DESCRIPTION: Germ tubes of the basidiospores emergent from both polar cells, sometimes from septal regions; conidia produced at the tip of germ tubes or laterally, budding to produce daughter cells polarly, developing into hyphae; colonies on PDA growing gradually, toa maximum 8 mm diam. in 21-day incubations, sometimes heaping up to form a mound, gelatinous, fixed on the agar surface, composed of branching and intricate hyphae, pseudohyphae and conidia; surface pastel yellow to pale yellow but sometimes pale orange or pale red to greyish red, slightly corrugate around the center, sometimes farinose by conidial formation; sometimes dark pigmentation present. Basidia 29-50 x 3.5-8.5 um, clavate to cylindrical, apices obtuse, 2-4-spored, directly emergent from the host surface or through stomata; sterigmata 3-7 um long, 1.5-3.0 jum diam. at the base, outwardly emergent, tapering toward the tip; basidiospores 10-19 x 3.0-5.9 um, musiform, straight to slightly curved, hyaline, smooth, unicellular when formed, becoming 1(-2)-septate when germinating; conidia 6-15 x 1.2-2.0 um, linear.

SYMPTOMATIC CHARACTERISTICS: Pocket like depressions in leaves, concave above and convex below, | to 5 galls on one leaf, sometimes covering the entire leaf, light green above, sometimes becoming reddish; both adaxial surface and abaxial surface of the infected area eventually covered with white powdery hymenium; mycelial mats located under both epidermal layers; infected parts 2-9 times thicker than healthy parts; hypertrophy and hyperplasia occurred; cell differentiation absent in the infected tissues except for the epidermis and vascular bundle; cell contents lost in the infected parts.

Hapsirar: On living leaves of Rhododendron yedoense f. poukhanense and R. yedoense f. yedoense. SPECIMENS EXAMINED: On R. y. f. poukhanense, KFRI2253, 2254, 2255 (3 June, 2004, Seoul), KFRI2256 (5 June, 2004, Pocheon-si); On R. y. f. yedoense, KFRI2257, 2258 (3 June, 2004, Seoul), KFRI2259 (5 June, 2004, Pocheon-si), KFRI2260 (24 June, 2004, Seoul).

Notes: Chung et al. (1977) and Korean Society of Plant Protection (1986) listed this fungus in Korea without morphological description on R. schlippenbachii and R. mucronulatum, respectively. However, R. schlippenbachii and R. mucronulatum have wrongly been identified as hosts of E. japonicum instead of R. y. f. poukhanense in Korea. There is no Exobasidium species causing such pocket-like leaf galls on those species and

26

Fig. 3. Exobasidium japonicum. A. A colony on PDA, 16 weeks, KFRI2256. B. A colony on PDA exuding dark pigment, 16 weeks, KFRI2260. C. Basidiospore germination of mycelial type on PDA, Arrow shows a long germ tube, KFRI2255. D. A Basidium with four immature basidiospores, Arrows show sterigmata, KFRI2260. E. Musiform basidiospores before septation, KFRI2256. EF. Conidia, KFRI2256. G. Pocket-like depressions on a naturally infected host. H. A cross section of healthy part of infected leaf, KFRI2260. I. A cross section of infected part with hypertrophy and hyperplasia, Arrows show mycelial mats in both adaxial and abaxial sides, KFRI2260.

Scale bars: C = 10 um, D = 5 um, E-F = 10 um, H = 20 um, I = 40 um.

related sections, Sciadorhodion and Rhodorastum. Therefore, this fungus is considered as a new record in Korea. The Korean materials have shorter basidia than those of previous records (Ito 1955, Nannfeldt 1981, Ezuka 1990b). Nannfeldt (1981) commented that infected parts were eventually hymenium-clad and chalky white on the morphological underside. However, white hymenium-clad occurred on both sides of leaf galls in Ezuka’s study and present study. The other characteristics of taxonomic value for this species are generally in agreement with the previous records. R. indicum, R. kaempferi, R. lateritium, R. linearifolium, R. mucronatum, R. obtusum, R. pulchrum, and R. simsii were reported from Japan and Europe (Ito 1955, Nannfeldt 1981, Ezuka 1990b). R. y. f. poukhanense and R. y. f. yedoense were recognized as new hosts to Korea.

4. Exobasidium cylindrosporum Ezuka, Trans. Mycol. Soc. Japan 31: 451 (1990). Fig. 4

DESCRIPTION: Germ tubes of the basidiospores emergent from each cell, small and short; conidia produced at the tip of germ tubes, budding to produce daughter cells polarly; colonies on PDA growing gradually, toa maximum 11 mm diam. in 21-day incubations, gelatinous, fixed on the agar surface, composed of hyphae and conidia; surface white to orange white, corrugate narrowly around the center; dark pigmentation absent.

27

Fig. 4. Exobasidium cylindrosporum, KFRI2261. A. A colony on PDA, 16 weeks. B. Basidiospore germination of conidial type on PDA, Arrow shows a short germ tube. C. A basidium with six immature basidiospores, Arrows show sterigmata. D. Long and slenderly musiform basidiospores before septation. E. Conidia. F. Shallow leaf depressions on a naturally infected host. G. A cross section of healthy part of infected leaf. H. A cross section of infected part in which hypertrophy without hyperplasia occurred, Arrows show mycelial mats in both adaxial and abaxial sides.

Scale bars: B = 10 um, C = 3 um, D = 10 um, E= 10 um, G-H = 20 um.

Basidia 20-36 x 5.0-6.9 um, clavate to cylindrical, apices obtuse, 4-6-spored, directly emergent from the host surface or through stomata; sterigmata 4-6 um long, 1.7-2.7 um diam. at the base, outwardly emergent, tapering toward the tip; basidiospores 19-24 x 2.9-4.8 tum, slenderly and long cylindrical, slightly curved, hyaline, smooth, unicellular when formed, becoming 1-3-septate when germinating; conidia 7-21 x 1.7-2.6 um, linear.

SYMPTOMATIC CHARACTERISTICS: Shallow depressions on leaves, 4 to 11 mm diam. across, with red margin above, usually concave above and convex below, mostly circular in outline, sometimes confluent to develop larger and irregular distortion; 1 to 3 shallow depressions on one leaf; both adaxial and abaxial sides of leaf galls eventually covered with white powdery hymenium; mycelial mats concentrated under both epidermal layers; infected parts 3-7 times thicker than healthy parts; hypertrophy occurred without hyperplasia; cell differentiation absent in the infected tissues except for the epidermis and vascular bundle; cell contents lost in the infected parts.

Hasirat: On living leaves of Rhododendron tashiroi Maxim.

SPECIMENS EXAMINED: On R. tashiroi, KFRI2261 (16 June, 2004, Suwon-si).

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Nores: This is the first record on the fungus in Korea. The Korean collection is in general in accordance with the description by Ezuka (1990b) except for the basidia that were measured longer in the literature. This fungus was recorded from Japan on leaves of R. hortense, R. macrosepalum, R. mucronatum and R. oomurasaki (Ezuka 1974b, 1990b). The present study adds Rhododendron tashiroi as a new host in Korea.

Acknowledgements

The authors express sincere thanks to Dr. Makoto Kakishima at University of Tsukuba, Japan, and Dr. Igor V. Karatygin at Komarov Botanical Institute of Russian Academy Sciences, for critical comments on the manuscript and for acting as pre-submission reviewers.

Literature cited

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Chung BJ, Lee YH, Lee EK. 1977. Survey on the occurrence of plant diseases in some important agricultural crops. Res. Rep. Plant Prot. Res. and Train. 7: 21. (in Korean).

Coyier DL, Roane MK. 1986. Compendium of rhododendron and azalea diseases. APS press: St. Paul, MN (USA). 88 pp.

Ezuka A. 1974a. A taxonomic examination of Exobasidium in Japan. Ann. Phytopath. Soc. Japan 40(2): 147.

Ezuka A. 1974b. A taxonomic examination of Exobasidium in Japan. Ann. Phytopath. Soc. Japan 40(3): 185-186.

Ezuka A. 1990a. Notes on some species of Exobasidium in Japan (I). Trans. Mycol. Soc. Japan 31: 375-388.

Ezuka A. 1990b. Notes on some species of Exobasidium in Japan (II). Trans. Mycol. Soc. Japan 31: 439-455.

Ezuka A. 1991a. Notes on some species of Exobasidium in Japan (III). Trans. Mycol. Soc. Japan 32: 71-86.

Ezuka A. 1991b. Notes on some species of Exobasidium in Japan (IV). Trans. Mycol. Soc. Japan 32: 169-185.

Ezuka A. 1992a. Exobasidium species in Japan (1). Forest Pests 41(1): 3-9.

Ezuka A. 1992b. Exobasidium species in Japan (2). Forest Pests 41(2): 2-9.

Ezuka A. 1992c. Exobasidium species in Japan (3). Forest Pests 41(3): 2-9.

Ing B. 1998. Exobasidium in the British Isles. The Mycologist 12(2): 80-82.

Ing B. 1999. Exobasidium in Scotland. Bot. J. Scotl. 51(2): 221-225.

Ito S. 1955. Mycological flora of Japan. Vol II. Yokendo: Tokyo (Japan). 450 pp.

Karatygin IV. 2000. Order Exobasidiales: position in class Ustilaginomycetes and key for the identification of species from genus Exobasidium. Mikologiia I Fitopathologiia 34(6): 23-31.

Karatygin IV, Huseyin E. 2002. A rare species from the genus Exobasidium on Rhododendron. Mikologiia I Fitopathologiia 36(4): 13-15.

Korean Society of Plant Pathology. 2004. List of plant diseases in Korea. 4th edition. Jeong-haeng Pub. Co.: Seoul (Korea). 779 pp.

Korean Society of Plant Protection. 1986. A list of plant diseases, insect pests, and weeds in Korea. 2nd edition. Korean Society of Plant Protection: Seoul (Korea). 633 pp. (in Korean).

Nagao H, Ezuka A, Ohkubo H, Kakishima M. 2001. A new species of Exobasidium causing witches’ broom on Rhododendron wadanum. Mycoscience 42: 549-554.

Zo

Nagao H, Akimoto M, Kishi K, Ezuka A, Kakishima M. 2003a. Exobasidium dubium and E. miyabei sp. nov. causing Exobasidium leaf blisters on Rhododendron spp. in Japan. Mycoscience 44: 1-9.

Nagao H, Ogawa §, Sato T, Kakishima M. 2003b. Exobasidium symploci-japonicae var. carpogenum var. nov. causing Exobasidium fruit deformation on Symplocos lucida in Japan. Mycoscience 44: 369-375.

Nagao H, Sato T, Kakishima M. 2004a. Three species of Exobasidium causing Exobasidium leaf blight on subgenus Hymenanthes, Rhododendron spp., in Japan. Mycoscience 45: 85-95.

Nagao H, Kurogi S, Sato T, Kakishima M. 2004b. Taxonomy of Exobasidium otanianum causing Exobasidium leaf blight on Rhododendron species in Japan. Mycoscience 45: 245-250.

Nannfeldt JA. 1981. Exobasidium, a taxonomic reassessment applied to the European species. Symb. Bot. Upsal. 23(2): 1-72.

Otani Y. 1976. A new species of Exobasidium collected in Iriomote-island, Okinawa. Trans. Mycol. Soc. Japan 17: 355-357.

Sundstrém KR. 1964. Studies of the physiology, morphology and serology of Exobasidium. Symb. Bot. Upsal. 18: 1-89.

Zhang X, Arai K, Sakoda T, Iwai H. 1995. A new variety of Exobasidium otanianum isolated from Rhododendron dilatatum var. satsumense. Trans. Mycol. Soc. Japan 36: 97-102.

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MYCOTAXON

Volume 96, pp. 31-39 April-June 2006

Annulate Pluteus species, a study of the genus Chamaeota in the United States

ANDREW M. MINNIs’, WALTER J. SUNDBERG’, ANDREW S. METHVEN?’, SEDONIA D. SIPES’, & DANIEL L. NICKRENT’

*drew3@siu.edu ‘Department of Plant Biology, Southern Illinois University at Carbondale Carbondale, IL USA 62901-6509

*Department of Biological Sciences, Eastern Illinois University Charleston IL 61920-3099

Abstract—Chamaeota is a rare agaric genus traditionally classified in the Pluteaceae. The present work offers a treatment of the two species described from the United States. Historical records, morphological and anatomical examinations of type and other collections, and molecular data from nuclear LSU rDNA sequences were studied. Typifications of Annularia mammillata and A. sphaerospora are offered for nomenclatural stability. Chamaeota sphaerospora is synonymized with C. mammillata and the new combination Pluteus mammillatus is proposed.

Key words—Agaricales, taxonomy, Hispidoderma, lectotype, epitype

Introduction

The genus Chamaeota (W.G. Sm.) Earle, typified by C. xanthogramma (Ces.) Earle, a poorly known fungus, is typically classified in the Pluteaceae Kotl. & Pouzar (Singer 1986) because the better known species have pink basidiospores, free lamellae, a convergent lamellar trama, and an annulus. The genus shares the first three characters with Pluteus Fr. and Volvariella Speg., the other members of the family, but differs in the last (Singer 1986). Recent molecular evidence, however, suggests that Pluteus and Volvariella together do not form a clade (Moncalvo et al. 2002). The relationship of Chamaeota to other agarics and its potential use as an outgroup for Pluteus was unknown.

Chamaeota is a small genus consisting of approximately ten species worldwide. Species of Chamaeota have been infrequently reported in the United States from a limited number of sites (Longyear 1902, Peck 1906, Murrill 1917, Murrill 1943, Kauffman 1918, Beardslee 1939). Species delineation remains unclear and several authors (Kauffman 1918, Beardslee 1939, Murrill 1943, Singer 1978) have commented on the need to clarify the range of variation and boundaries of individual species. Two species, Chamaeota sphaerospora and Chamaeota mammillata are the only species originally described from the United States. Their interpretation is made difficult by the lack of explicitly designated type specimens. The present paper is offered to address these problems in the United States.

32

Materials and methods

Molecular Study

Genomic DNA was isolated from the ASM 7916 collection identified by classical morphology and anatomy as C. sphaerospora using a version of the method described by Nickrent (1994) scaled down to use less material. Universal primers routinely used in Vilgalys’ laboratory at Duke University (http://www.biology.duke.edu/fungi/mycolab/ primers.htm) were employed to amplify (5.8SR, LR7) and to obtain (LR5, LR16, LROR, LR3R) a partial nuclear LSU rDNA sequence using standard protocols. The resulting sequence was subsequently aligned manually to the LSU sequence alignment used in the analysis of Moncalvo et al. (2002). A heuristic parsimony analysis similar to that of Moncalvo et al. (2002) was performed to place the sequence on a fungal tree (results not shown). The above analysis placed the sequence within a Pluteus clade. A reduced dataset containing 21 taxa was constructed to demonstrate the position of Chamaeota within the Pluteus clade. Limacella illinita (Fr.) Maire, Volvariella volvacea (Bull.) Singer, and Auricularia polytricha (Mont.) Sacc. were chosen as outgroup taxa because of their positions outside of the Pluteus clade. A branch-and-bound analysis was performed with the following parameters: gaps treated as missing data, multistate taxa interpreted as uncertainty, addition sequence furthest, branches collapsed if maximum branch length is zero, MulTrees option in effect, characters weighted using the Moncalvo et al. (2002) stepmatrix. Branch robustness was evaluated with 1000 bootstrap replications using identical search parameters.

Morphological & anatomical study

Fresh collections identified as Chamaeota sphaerospora were made, documented, and preserved with standard methods (Smith 1949). Color terms are from Kornerup and Wanscher (1978). Standard techniques were used to prepare material for anatomical study (Smith 1949, Largent et al. 1977). Dried material was wetted with 95% ethanol for 1 min, soaked in water for 1 min, blotted to remove excess water, and sectioned by hand. Material was mounted and viewed in 3% KOH or Melzer’s reagent. Basidiospore dimensions of the newly designated lectotype of Annularia sphaerospora are based on observation of 30 basidiospores. All other microscopic dimensions are based on observation of at least 10 structures. Length to width ratios are reported as Q. Mean values for length, width, and Q are designated as L™, W™, and Q™ respectively. Anatomical data for the lectotype of A. sphaerospora are presented separately in the species description because of the importance of this collection. Data from other specimens are presented as a composite description under “additional data” where they add to or differ from the type material and original circumscription. Herbarium acronyms are from Holmgren and Holmgren (1998 onwards).

Results

The nuclear LSU rDNA dataset analyzed included 781 characters. Of these, 74 were parsimony-informative. The maximum parsimony analysis produced one shortest tree of 990.3 steps (Fig. 1). The consistency (CI), homoplasy (HI), retention (RI), and rescaled consistency (RC) indices were 0.7157, 0.2843, 0.7698, and 0.5509, respectively.

33

Pluteus petasatus (AF042611)

i Pluteus primus (AF042610)

Pluteus brunneoradiatus (AF261567) 2 soitnse Pluteus pallidus (AF261569)

Pluteus cervinus (AF261570)

Pluteus pouzarianus (AF261568)

Pluteus salicinus (AF261573)

Chamaeota sphaerosporal

67 Pluteus mammillatus (DQ45 1549) _ Pluteus ephebeus (AF261574)

Pluteus sp. (AF042612)

o Pluteus sp. (AF261576)

: Pluteus aurantiorugosus (AF261579) Pluteus romellii (AF261575)

Pluteus admirabilis (AF261577)

Hs} ike Pluteus admirabilis (AF261578)

Pluteus chrysophlebius (AF261581) Pluteus umbrosus (AF261580)

100 Pluteus pellitus (AF261571)

Pluteus atromarginatus (AF261572) *Limacella illinata (AF261439)

on *Volvariella volvacea (AF261531)

*Auricularia polytricha (AF261554)

Fig. 1. The single most parsimonious tree based on partial nuclear LSU rDNA sequences. Bootstrap values greater than 50% are indicated above branches. Genbank accession numbers for each sample are shown. All sequences originated in the Moncalvo et al. (2002) study except Pluteus sp. (AF042612) from Moncalvo et al. (2000) and Chamaeota sphaerospora/Pluteus mammillatus from this work. Asterisks denote outgroup taxa.

34

Pluteus forms a clade (84% BS) with the sequence of the ASM 7916 collection of C. sphaerospora embedded within it. There is moderate bootstrap support (67% BS) for a sister relationship of the ASM 7916 sequence to the P. ephebeus clade.

Typification

Chamaeota mammillata (Longyear) Murrill, N. Amer. Fl. 10: 139. 1917. = Annularia mammillata Longyear, Rep. Mich. Acad. Sci. 3: 59. 1902. Type: Pl. I, Fig. 4 in Longyear, Rep. Mich. Acad. Sci. 3: 58. 1902—Lectotype [icon.] designated here. Type: Annularia mammillata Longyear, Pluteus mammillatus Minnis et al., ASM 7916 (EIU)—Epitype designated here.

When Longyear (1902) named the species, he based it on a single basidiocarp collected from Greeneville, Michigan on a decaying log in woods and did not explicitly designate a type. The original material mentioned in the protologue was not found at Michigan State University, formerly the Michigan Agricultural College (Heather Hallen and Alan Fryday, pers. comm.). Likewise, it is not present at the New York Botanical Garden (NY) (Barbara Thiers, pers. comm.) or the University of Michigan Herbarium (MICH).It is apparently lost. No other authentic material or other specimens are available for study. Since it is allowable to select an illustration as a lectotype, we chose the figure from Longyear’s original circumscription. See Fig. 2 for a reproduction of this image.In order to facilitate a more precise taxonomic interpretation, a collection used in this study was chosen as the epitype. All of the specimen data on the herbarium box are given above.

Chamaeota sphaerospora (Peck) Kauffman, The Agaricaceae of Michigan 534. 1918.

= Annularia sphaerospora Peck, Bull. Torrey Bot. Club 33: 216. 1906. Type: Annularia

sphaerospora Peck, Loc. Detroit, Mich, Leg. O.E. Fischer, Aug. ‘05 (NYS)—Lectotype designated here.

In the protologue, Peck gave O.E. Fischer, a physician, credit for collecting the material used to circumscribe the species. However, unpublished correspondence between Peck and Fischer indicate that a Mr. Burrows of 50 Hazelwood Ave. (Detroit?) was the collector. Fischer sent one of the two collections given to him by Mr. Burrows to Peck. This material mentioned in the protologue was apparently used by Peck to write the circumscription, but he did not clearly designate a type. All of the type specimen data present on the specimen label are given above. An isolectotype not mentioned in the original species description is located in the University of Michigan Herbarium (MICH). According to the herbarium collection manager, Patricia Rogers (pers. comm.), Dr. Pierre-Arthur Moreau noticed some Hebeloma material mixed in with this collection. We have not examined this collection.

Taxonomy

Pluteus mammillatus (Longyear) Minnis, Sundb., & Methven, comb. nov. Figs. 2-7. = Annularia mammillata Longyear, Rep. Mich. Acad. Sci. 3: 59. 1902 (basionym). = Chamaeota mammillata (Longyear) Murrill, N. Amer. Fl. 10: 139. 1917. = Annularia sphaerospora Peck, Bull. Torrey Bot. Club 33: 216. 1906. =Chamaeota sphaerospora (Peck) Kauffman, The Agaricaceae of Michigan 534. 1918.

35

Data based on study of the lectotype of Annularia sphaerospora:

Basidiospores globose to subglobose to short-ellipsoid to slightly ovate in face and profile views, circular in end-view, 5.3-7.0 x 4.8-6.6 um, Q = 1.0-1.18 (L™ = 6.1 um, W™ = 5.5 um, Q™= 1.11), apiculate, smooth, walls slightly thickened, subhyaline to pale yellow in KOH, with a single large oil drop inside or several smaller drops resulting in a granular appearance. Basidia clavate, tetrasterigmate, 20-23 x 7-9 um without sterigmata, walls thin and hyaline in KOH. Pleurocystidia abundant, broadly to narrowly fusoid-ventricose with pedicels and short necks with obtuse apices, 37- 61 x 14-24 um, walls thin and hyaline in KOH, no apparent contents. Cheilocystidia abundant, vesiculate to clavate to fusoid-ventricose, 31-58 x 11-22 um, walls thin and hyaline in KOH, no apparent contents. Lamellar trama convergent. Subhymenium cellular. Pileipellis more or less a cutis with bundles of erect to appressed hyphae that are more abundant towards the disc, hyphae thin-walled occasionally containing light brown intracellular pigment. Context composed of 3-15 um wide hyphae, thin-walled, hyaline in KOH, no apparent contents. Stipitipellis a cutis of cylindrical hyphae, 3-8 um wide, thin-walled, hyaline in KOH, no apparent contents. Clamp connections absent. All anatomical features inamyloid in Melzer’s reagent.

Additional data:

Pileus 0.9-7 cm in diameter, initially conic-campanulate becoming convex-plane, low narrow to broad umbo present or not; margin entire, slightly striate, occasionally appendiculate, initially incurved then becoming decurved; surface dry, radially fibrillose with fibrillose scales on disc, fibrils becoming smaller and more separated towards the margin, fibrils light brown to brown (6D7-D6, 6E8-E6); ground color deep yellow to orange yellow (4A-8-A6 to 4B8-B7). Lamellae free, close, thin, broad, not marginate, white becoming grayish red (7B3) in age. Lamellulae in two tiers. Stipe 0.2-8 cm x 1-14 mm, equal or narrowly clavate, terete, whitish to pale yellow (3A3), surface dry, longitudinally fibrillose or streaked, fibrils concolorous with those on pileus, basal tomentum white. Annulus found on lower 1/2 to 1/3 of stipe, white to yellowish white (4A2,3A2), thin, membranous, sheathing with flaring apex, occasionally evanescent. Flesh of pileus and stipe solid, white. Odor and taste not distinctive.

Basidiospores brownish pink in mass, 4.8-7.0 x 3.1-6.6 um, Q = 1.08-1.88 (L™ = 6.1 um, W" = 5.3 um, Q™= 1.17). Basidia 17-29 x 6-11 um, usually with granular contents. Pleurocystidia rarely broadly clavate, 39-99 x 8-30 um. Cheilocystidia 30-86 x 8-24 um. Pileipellis hyphae often containing brown intracellular pigment, 5-13 um wide. Stipitipellis hyphae 4-18 um wide, with or without brown intracellular pigment. Caulocystidia consist of cylindrical hyphae with obtuse apices, 5-9 um wide, thin- walled, with brown intracellular pigment. Annulus composed of filamentous hyphae, 3-10 um wide, thin-walled, hyaline in KOH, no apparent contents.

Specimens examined—UNITED STATES. Fioripa: Seminole Co., Oviedo,

22.XI1.1935, coll. HC Beardslee Jr. (35135, MICH as Chamaeota pusilla); 9.X1.1936, coll. HC Beardslee Jr. (35135, MICH as Chamaeota fenzlii); Highlands Hammock State Park near Sebring, 1942, coll. R Singer, F244a (F). MICHIGAN: Wayne Co., Detroit, VIII.1904, coll. RH Stevens (F as Annularia fenzlii). Missouri: Wayne Co., Mingo National Wildlife Refuge near Puxico, scattered on lignicolous substrates, 20.IX.2003, coll. AM Minnis, Minnis 3-09-20-1 (SIU) & Minnis 3-09-20-2 (SIU); 17.[X.1994, coll. AS Methven, ASM

36

Figs. 2-6. Pluteus mammillatus. Fig. 2. Lectotype illustration of Annularia mammillata from Longyear (1906). Fig. 3. Basidiospores (A. sphaerospora lectotype). Fig. 4. Pleurocystida (A. sphaerospora lectotype). Fig. 5. Cheilocystidia (A. sphaerospora lectotype). Fig. 6. Pileipellis (Minnis 5-09-20-1). Scale bars for Figs. 3-6 = 10 um.

an

Fig. 7. Pluteus mammillatus. Basidiocarps of two collections, Minnis 3-09-20-1 and Minnis 3-09- 20-2.

7916, DQ451549 (designated epitype of A. mammillata, EIU); 20.1X.2005 coll. AM

Minnis, Minnis 5-09-20-1 (SIU). Comments—This species is easily recognized in the field by its yellowish pileus, free, dirty-pink lamellae, annulus, and occurrence on decaying deciduous substrates. Microscopically, there is considerable variation in pleurocystidial shape. This variation around the fusoid-ventricose type is present within and among individual basidiocarps. In our experience, this variation is not unusual in Pluteus. Kauffman (1918) did not note the presence of pleurocystidia in Chamaeota sphaerospora. However, they are present in all of the collections we have examined. Hesler noted the presence of pleurocystidia on an annotation label for the lectotype of A. sphaerospora. Despite the misleading species epithet of C. sphaerospora, the spores are not always globose.

Murrill (1917) considered C. mammillata to be a synonym of C. sphaerospora. Kauffman (1918) and Singer (1978) recognized the two as distinct species. Considering the range of variation we have seen in C. sphaerospora, Longyear’s description (1902) is a match and the two species are synonyms. See discussion for justification of the transfer to Pluteus.

Pluteus mammillatus (designated above) is rarely collected in the United States. Even so, it appears to be widespread in the Eastern United States. It has also been reported from Argentina (Moreno and Alberté 1996). Wright and Alberté (2002) included a photograph with their description of an Argentinean collection that appears to be P mammillatus.We have not examined any of the South American collections.

Chamaeota fenzlii (Schulzer) Singer, a European species, is similar. According to Singer's description, it differs from C. sphaerospora by possessing yellow-marginate lamellae (1978).

Discussion

Singer (1986) discussed the similarities of Chamaeota to Pluteus section Hispidoderma Fayod. This section is characterized by a pileipellis consisting entirely of elongated

38

elements and thin-walled cystidia. The species of Chamaeota noted by Singer differ primarily by the presence of a partial veil. For convenience in matters of identification, Singer (1986) felt that it was acceptable to maintain the genus as separate at that time despite the number of other agarics where the presence or absence of a veil does not warrant genus level distinction. In contrast, in his seminal work on Pluteus in South America, Singer (1958) noted the presence ofa rudimentary volva in Pluteus stephanobasis Singer and stated that it was acceptable to maintain this species in the genus instead of placing it in Volvariella. Characters that are not found in Volvariella including metuloid pleurocystidia and the lack of a natural transition from a rudimentary volva to the more complex volvas found in Volvariella supported his view (Singer 1958).

Based on the molecular evidence presented in this study, Chamacota is not a good outgroup for Pluteus because it is in the Pluteus clade. The partial veil of C. mammillata is a derived character within this Pluteus lineage. Of the taxa examined, C. mammillata is most closely related to Pluteus ephebeus (Fr.) Gillet, a member of Pluteus section Hispidoderma. We feel that the partial veil does not warrant exclusion of C. mammillata from the genus Pluteus. Future descriptions of the genus should be modified to potentially include the presence of a partial veil.

Interestingly, Chamaeota tropica Pegler and Chamaeota subolivascens Courtec. are described as having cellular pileipelli (Pegler 1983, Courtecuisse 1991). This feature is also found in Pluteus section Celluloderma Fayod. It would be interesting to know if these species are also derived from within the Pluteus lineage. If so, a partial veil is likely to have been derived more than one time in the genus. Future studies on the other species of Chamaeota including the type of the genus, C. xanthogramma, should be performed. It is our belief that the genus will be rendered obsolete.

Acknowledgements

We thank Dr. Else C. Vellinga and Dr. Scott A. Redhead of the University of California, Berkeley and the Eastern Cereal and Oilseed Research Centre, respectively, for their presubmission reviews of the manuscript. We also acknowlege the New York State Museum (NYS), the University of Michigan Herbarium (MICH) and the Field Museum of Natural History (F) for the loans of collections used in this study. Dr. Raymond Stotler and Dr. Dale H. Vitt are credited for advice on nomenclature. Finally, our gratitude is extended to the Missouri Mycological Society for their support and comraderie.Financial support for this work was provided by the Alexander H. and Helen V. Smith Research Fund.

Literature cited

Beardslee HC. 1939. New and interesting fungi. Mycologia 26: 253-260.

Courtecuisse R. 1991. Eléments pour un inventaire mycologique des environs du saut Pararé (Arataye) et linselberg des Nouragues (Guyane francaise). V. Pluteaceae. (Pluteales, Basidiomycota). Crypt. Bot. 2:136-152.

Holmgren PK, Holmgren NH. 1998 onwards (continuously updated). Index Herbariorum. New York Botanical Garden. http://sciweb.nybg.org/science2/IndexHerbariorum.asp

Kauffman CH. 1918. The Agaricaceae of Michigan. Vol. 1. Wynkoop Hallenback Crawford Co., State Printers, Lansing, Michigan. 924 p.

Kornerup A, Wanscher JH. 1978. Methuen Handbook of Color. 3"! ed. Eyre Methuen, London. 252 p.

oo

Largent D, Johnson D, Watling R. 1977. How to Identify Mushrooms to Genus III: Microscopic Features. Mad River Press, Eureka, California. 148 p.

Longyear BO. 1902. New species of Michigan fungi. Rep. Mich. Acad. Sci. 3: 57-60.

Moncalvo JM, Lutzoni FE, Rehner SA, Johnson J, Vilgalys R. 2000. Phylogenetic relationships of agaric fungi based on nuclear large subunit ribosomal DNA sequences. Syst. Biol. 49: 278-305.

Moncalvo JM, Vilgalys R, Redhead SA, Johnson JE, James TY, Aime MC, Hofstetter V, Verduin SJW, Larsson E, Baroni TJ, Thorn RG, Jacobsson S, Clémencon H, Miller Jr. OK. 2002. One hundred and seventeen clades of Euagarics. Mol. Phyl. Evol. 23: 357-400.

Moreno G, Albertdé E. 1996. Agaricales sensu lato de Argentina. I. Cryptogamie, Mycol. 17: 61-84.

Murrill WA. 1917. (Agaricales), Agaricaceae (pars), Agariceae (pars). N. Amer. Fl. 10: 77-144.

Murrill WA. 1943. Some Southern novelties. Mycologia 35: 422-433.

Nickrent DL. 1994. From field to film: Rapid sequencing methods for field collected plant species. BioTechniques 16: 470-475.

Peck CH. 1906. New species of fungi. Bull. Torrey Bot. Club 33: 213-221.

Pegler DN. 1983. Agaric Flora of the Lesser Antilles. Her Majesty’s Stationary Office, London. 668 p27 pl

Singer R. 1958. Contribution toward a monograph of the genus Pluteus, especially those of the east slope of the Andes and Brazil. Lloydia 21: 195-299.

Singer R. 1978. Keys for the identification of the species of Agaricales II. Sydowia 31: 193-237.

Singer R. 1986. The Agaricales in Modern Taxonomy. 4" ed. Koeltz Scientific Books, Koenigstein. 981 p.

Smith AH. 1949. Mushrooms in Their Natural Habitats. Sawyers, Portland. 626 p.

Wright JE, Alberto E. 2002. Hongos. Guia de la Regidn Pampeana. 1. Hongos con Laminillas. L.O.L.A., Buenos Aires. 279 p.

MYCOTAXON

Volume 96, pp. 41-46 April-June 2006

Tylopilus microsporus, a new species from Southwest China

SZ FUS2sO2B. WANG Si Ye) Y AOe

yaoyj@sun.im.ac.cn 'Key Laboratory of Systematic Mycology and Lichenology Institute of Microbiology, Chinese Academy of Sciences Beijing 100080, China

*Mycology Section, Royal Botanic Gardens, Kew Richmond, Surrey TW9 3AB, UK

3Graduate School of Chinese Academy of Sciences, China y.

Abstract—A new species, Tylopilus microsporus, collected from Sichuan and Yunnan, China, is described and illustrated. The new species is characterized by pale violet to violet pileus, paler purple to purplish brown stipe, flesh color to pale purplish red hymenophore and small spores. The differences between this species and its related taxa, such as T. plumbeoviolaceus, T. rubrobrunneus, T: plumbeoviolaceoides and T.: violatinctus, are discussed.

Key words—Boletaceae, boletes, taxonomy

Introduction

Tylopilus P. Karst. is a cosmopolitan genus in Boletales with about 75 species worldwide (Kirk et al. 2001). Some species in the genus are edible but others are considered mildly poisonous (Singer 1986). Singer et al. (1991) considered tropical Asia and Central America as the distribution centres of Tylopilus, but its species are widespread, e.g. North America (Smith & Thiers 1971, Bessette et al. 2000), Asia (Corner 1972, Nagasawa 1997, Li & Song 2000), Mexico and Central America (Singer et al. 1991) and Australasia (Pegler & Young 1981, Li & Watling 1999, Watling 2001). Some uncertainties remain as to the taxonomic position of the genus largely owing to poor generic definitions. The genus was originally limited to T! felleus (Bull.) P. Karst., and defined as being gymnocarpic, ectomycorrhizal and producing pinkish, light greyish brown or pinkish orange-brown basidiospores not exceeding 15 um in length by Pegler & Young (1981), who also placed the genus within Strobilomycetaceae E.-J. Gilbert. This taxonomic treatment was accepted by Hawksworth et al. (1995). However, the genus is also characterized by having pinkish, pinkish orange, reddish brown, to purplish brown, amber brown or rusty brown basidiospores, lacking a veil and generally having pallid tubes when

* Author for correspondence.

42

young (Smith & Thiers 1971, Wolfe, 1979, Bessette et al. 2000), and placed in Boletaceae Chevall. (Watling 1970, Smith & Thiers 1971, Singer 1986, Kirk et al. 2001).

There is no major molecular study on the relationships within Tylopilus and between Tylopilus and Strobilomyces Berk. Tylopilus s. |. may not be monophyletic, but a clade of Tylopilus spp. including T: felleus is the sister-group to Leccinum, while Strobilomyces spp. are at the base of the Boletaceae (Binder 1999). The narrow generic concept proposed by Pegler & Young (1981) may be better to maintain the monophyletic position for Tylopilus, but it is difficult to define Tylopilus and to justify it as a natural genus in Boletaceae or in Strobilomycetaceae because of the simplicity and plasticity of morphological and anatomical characters and the lack of molecular data.

In China, species of the genus have been reported from various areas but mainly from the South and the South-west (Chiu 1948, Zang 1986, Ying & Zang 1994, Li & Song 2000), especially in Guangdong, Sichuan and Yunnan provinces. The early Chinese records of Tylopilus were mostly given as Boletus L. A total of some 30 taxa, including species and varieties, has so far been reported (Tai 1979, Zang 1986, Ying & Zang 1994, Yuan & Sun 1995, Shao & Xiang 1997, Chen & Peng 1998, Mao 1998, 2000, Zhang 1999, Li & Song 2000, Li et al. 2002, Wang et al. 2004) and a check-list of the genus in China with a review of all taxa reported is currently underway (Wang et al., in preparation).

In this paper, a new species of Tylopilus is proposed and illustrated, based on several collections made from Yunnan and Sichuan during an investigation into the diversity of macrofungi in the South-west of China.

Materials and Methods

All the specimens used in this study were collected from Yunnan and Sichuan, China, and are deposited in the Herbarium Mycologicum, Academia Sinica (HMAS). The macro-characters described here were based on fresh collections. The collections were dried by using an electric air-ventilation drier. Micro-characters were examined in a 5% KOH solution in laboratory from dried specimens. The line drawings of basidiospores and pleurocystidia were made with the assistance of a camera lucida. The sizes of basidiospores are based on at least 30 measurement of each mature specimen.

Taxonomy Tylopilus microsporus S.Z. Fu, Q.B. Wang & Y.J. Yao sp. nov. Figs 1 & 2 Etymology: The epithet refers to the comparatively small basidiospores found in this

species.

Pileus hemisphericus, convexus vel subconvexus, 6.0-12.0 cm latus, subamethystinus, amethystinus vel brunneo-amethystinus, subtomentosus vel pruinosus. Contextus 1.0-2.0 cm crassus ad centrum, cretaceus, constans; odor indistinctus, sapor amarus. Tubuli ad 5.0-10.0 mm longi, depressi ad stipitem, juvenes albi, maturi constanter albi vel parum subrosei; pori subangulares vel rotundi, 3-4 per mm, cretacei primum, deinde carnei vel pallide purpureo-coccinei, maturitate parum brunnei. Stipes cylindricus, 5.0-14.0 cm longus, 1.0-3.5 cm crassus ad centrum, subaequalis vel saepe leniter amplificatus deorsum, non reticulatus, leniter striatus, cretaceus vel brunneus, apice parum amethystinus, juventute deorsum pallide brunneo-amethystinus, amethystinus vel brunneo-amethystinus, maturitate pallide brunneus vel brunneus, basi albus, solidus. Basidiosporae (6.5-)7.0-

43

9.0(-10.0) x 3.0-4.0(-5.0) um, subfusoideae, glabrae, pallide subroseo-brunneae. Basidia 22-24 x 8.0-9.0 um, clavata. Cystidia 37-58 x 8.0-12.0 um, clavata vel fusoidea infra inflata, in solutione KOH intus pallide aurantiaca. Trama hymenophori divergens. Hyphae hyalinae, fibulis destitutis.

Pileus 6.0-12.0 cm in diam., at first hemispherical, becoming convex, subconvex to nearly applanate at maturity; margin incurved at first, becoming decurved with a narrow band of sterile tissue; surface dry, subtomentose or more or less pruinose, pale violet to violet when young, becoming pale violet, purple pink, with brown tints when mature, then pale brownish with greyish violet tints when old, often pale yellow brown at centre and with pale violet tints at margin; unchanging or slightly deeper on bruising. Context 1.0-2.0 cm thick at centre, white, unchanging when bruised, odour not distinctive, taste very bitter. Tubes 5.0-10.0 mm long, white when young, and remaining so when mature or slightly pinkish, unchanging when bruised, adnate to slightly depressed around the stipe; pores nearly round, 3-4 pores per mm, white at first, becoming flesh-coloured to purplish pink to pale violet with slightly brown tints when mature, unchanging when bruised. Stipe 5.0-14.0 cm long, 1.0-3.5 cm thick, sub-cylindric, always enlarging downward, non-reticulate, slightly striate or subglabrous, white or brownish with pale purple to violet tints at the apex, pale brownish lilac to lilac or brownish lilac downwards when young, becoming brownish to brown, with white mycelium at the base, solid; context white, unchanging when bruised. Basidiospores (6.5-)7.0-9.0(-10.0) x 3.0- 4.0(-5.0) um, subfusoid, distally expanding, smooth, thin-walled, pale pinkish brown containing one or two oil-droplets. Basidia 22-24 x 8.0-9.0 um, clavate, 2- or 4-spored. Pleurocystidia 37-58 x 8.0-12.0 um, copious, clavate to fusoid, inflated below, with pale golden yellow contents in KOH. Hymenophoral trama of Boletus-type, typically divergent from a central strand, hyphae hyaline in KOH, thin-walled. Pileipellis an interwoven trichodermium, slightly brown in KOH. Stipitipellis a trichodermium, with some short, narrowly ventricose dermatocystidia 35-60 x 6.5-12.0 um. Clamp-connections absent.

Specimens examined: China: Yunnan, Kunming, Heping Village Market, 31 Aug. 2002, Q. B. Wang No. 85, HMAS 79720; Simao, Simao Market, 10 Aug. 2003, Q. B. Wang No. 190, HMAS 84730 (HOLOTYPE). Sichuan, Dechang County, Badong Town, Gaofeng, 1900 m alt., solitary to gregarious on the ground, in mixed woods, 16 Aug. 2003, Q. B. Wang & S. Z. Fu No. 234, HMAS 84743; Dechang County, Badong Town, Yinjiashan, 2200 m alt., solitary on the ground, in mixed woods, 14 Aug. 2003, S. Z. Fu No. 12, HMAS 79471; Dechang County, Nanshan Town, Datianwan Village, 1700 m alt., solitary on the ground, under the mixed woods of Quercus sp. and Pinus sp., 15 Aug. 2003, S. Z. Fu No. 32, HMAS 79468; same locality and same date, solitary to gregarious on the ground, S. Z. Fu No. 33, HMAS 84745; same locality and same date, gregarious, S. Z. Fu No. 34, HMAS 79458.

Habit and habitat: Solitary to gregarious on the ground in mixed woods with Quercus sp. and Pinus sp.

Distribution: China: Yunnan and Sichuan.

This species is mainly characterized by the pale violet to violet pileus, pale purple hymenophore, brownish stipe and small spores, (6.5-)7.0-9.0(-10.0) x 3.0-4.0(-5.0) um. These characters, together with the very bitter context, may be confused with other closely related species in the field, such as T: plumbeoviolaceoides T.H. Li et al., T.

44

Figs 1 & 2. Tylopilus microsporus (HMAS 84730, Holotype). Fig. 1. Photograph of basidiomata. Fig. 2. Line drawing of microscopic characters. A. Pleurocystidia; B. Basidiospores.

plumbeoviolaceus (Snell & E.A. Dick) Singer, T. rubrobrunneus Mazzer & A.H. Sm. and T. violatinctus T.J. Baroni & Both, which have a vinaceous or purple brown pileus and very bitter context.

Tylopilus microsporus differs from T. rubrobrunneus and T. plumbeoviolaceus in having smaller spores. The spores of T: rubrobrunneus are 10-14 x 3.5-4.5 um (Smith & Thiers 1971, Bessette et al. 2000) or 12-14 x 4-5 um (Grund & Harrison 1976), larger than those of T: microsporus. Furthermore, the pileus of T: rubrobrunneus is purple- brown, dark reddish brown, dull brown or cinnamon in age (Bessette et al. 2000), while the pileus of T. microsporus is pale violet, purple pink, with brown tints. Although spores of T. plumbeoviolaceus were measured as 7-9 x 2.8-4 um (mostly 8-9 x 3 um) by Snell & Dick (1941), Singer (1947) considered the spore size of 9.5-14 x 3-3.8 um (most frequently 10.2-12 x 3.3-3.5 um) was reasonably constant because of slow maturing of the spores in this species. Singer et al. (1991) gave the same size of spores for collections from Mexico. The spore size of 10-13 x 3-4 um was also provided for T. plumbeoviolaceus by both Smith & Thiers (1971) and Bessette et al. (2000), and 11-14 x 4-5 um by Grund & Harrison (1976). All these spore measurements are larger than those in T: microsporus. Further, the pileus of T. plumbeoviolaceus is quite violaceous when young (Snell & Dick, 1941), contrasting pale violet to violet when young in T! microsporus.

Tylopilus plumbeoviolaceoides was described from Guandong, China, by Li et al. (2002). It has a dark violaceous, brown-vinaceous to purple-chestnut pileus, which is subviscid to viscid, and the pileipellis is an ixotrichodermium or a trichodermium with some gelatinized material; whilst the pileus of T: microsporus is dry, pale violet to violet when young and becoming pale violet, purple pink, with brown tints in age, and the pileipellis is an interwoven trichodermium without gelatinized material. The stipe of T. plumbeoviolaceoides is concolorous with the pileus to more pinkish chestnut, compared with the brownish stipe of T. microsporus. Further, the spores of T: plumbeoviolaceoides, (7.5-)8.5-10.5(-12) x (2.5-)3-3.8(-4.2) um (Li et al. 2002), are longer than those of T. microsporus. The fungus illustrated for T. plumbeoviolaceoides by Wang et al. (2004) from Yunnan is the same as T: microsporus described here (Wang, pers. comm.).

Tylopilus violatinctus, hitherto known only from North America, is another similar species, but its pileus becomes rusty violet to dark violet and stipe turns typically to dull yellow to yellow brown when bruised, and its hymenophore becomes pinkish flesh-color

45

and finally deep cocoa-brown when mature (Bessette et al. 2000); contrasting to the unchanging, or just deeper, pileus and the unchanging stipe when bruised, and the pale violet, purple pink, with brown tints hymenophore when mature in T’ microsporus.

The complex of Tylopilus species with purplish-tinted basidiomata is difficult to determine by macro-characters alone. There might be many species involved in this complex. Further studies, especially DNA sequence analyses, are needed in order to understand the complex.

Acknowledgements

The authors are grateful to Drs H. Knudsen and T.-H. Li for serving as pre-submission reviewers and for their valuable comments and suggestions, and to Prof. J.-Y. Zhuang for assistance with the Latin diagnosis. We are thankful to Ms. X. H. Wang, Profs P.-G. Liu and Zhu-L. Yang, and Mr. F.-Q. Yu for their kind help in the field work. This study is supported by the Programme for Key International Science and Technology Cooperation Project of P. R. China (2001CB711103) from Ministry of Science and Technology, the National Science Fund for Distinguished Young Scholars (30025002) from National Natural Science Foundation of China, the Key Research Direction of Innovation Programme (KSCX2-SW-101C) and the scheme of Introduction of Overseas Outstanding Talents, operated by the Chinese Academy of Sciences.

Literature Cited

Bessette AE, Roody WC, Bessette AR. 2000. North American Boletes, a Color Guide to the Fleshy Pored Mushrooms. Syracuse University Press, Syracuse, USA.

Binder M. 1999. Zur Molekularen Systematik der Boletales: Boletineae und Sclerodermatineae subordo nov. PH.D. thesis. University of Regensburg, Germany.

Chen C-M, Peng J-J. 1998. Four Tylopilus new to Taiwan. Fung. Sci. 13: 11-16.

Chiu W-F. 1948. The boletes of Yunnan. Mycologia 40: 199-231.

Corner EJH. 1972. Boletus in Malaysia. Government Printer, Singapore.

Grund DW, Harrison KA. 1976. Nova Scotian Boletes. J. Cramer, Vaduz, Liechtenstein, Germany.

Hawksworth DL, Kirk PM, Sutton BC, Pegler DN. 1995. Ainsworth & Bisby’s Dictionary of the Fungi. 8th ed. CAB International, Wallingford, UK.

Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the Fungi. 9th ed. CAB International, Wallingford, UK.

Li T-H, Song B. 2000. Chinese boletes: a comparison of boreal and tropical elements. In Whalley AJS. Tropical Mycology 2000, the Millennium Meeting on Tropical Mycology (Main Meeting 2000). 1-9p. British Mycological Society & Liverpool John Moores University, Liverpool, UK.

Li T-H, Watling R. 1999. New taxa and combinations of Australian Boletes. Edinb. J. Bot. 56: 143- 148.

Li T-H, Song B, Shen Y-H. 2002. A new species of Tylopilus from Guandong. Mycosystema 21: 3-5.

Mao X-L. 1998. The Economic Fungi in China. Science Press, Beijing, China (in Chinese).

Mao X-L. 2000. The Macrofungi of China. Henan Science and Technology Press, Zhengzhou, China. (in Chinese).

Nagasawa E. 1997. A preliminary checklist of the Japanese Agaricales. I. the Boletineae. Rep. Tottori Mycol. Inst. 35: 39-78.

Pegler DN, Young TWK. 1981. A natural arrangement of the Boletales, with reference to spore morphology. Trans. Br. Mycol. Soc. 76: 103-146.

46

Shao L-P, Xiang C-T. 1997. Forest Mushrooms of China. Northeast Forestry University Press, Haerbin, China. (in Chinese).

Singer R. 1947. The Boletoideae of Florida. The Boletineae of Florida with notes on extralimital species III. Amer. Mid]. Nat. 37: 1-135.

Singer R. 1986. The Agaricales in Modern Taxonomy. 4th ed. Koeltz Scientific Books, Koenigstein, Germany.

Singer R, Garcia J, Gomez LD. 1991. The Boletineae of Mexico and Central America. III. Beihefte zur Nova Hedwigia Heft 102: 1-99.

Smith AH, Thiers HD. 1971. The Boletes of Michigan. The University of Michigan Press, Ann Arbor, USA.

Snell WH, Dick EA. 1941. Notes on boletes. VI. Mycologia 33: 23-37.

Tai F-L. 1979. Sylloge Fungorum Sinicorum. Science Press, Beijing, China. (in Chinese).

Teng S-C. 1936. Additional fungi from China. III. Sinensia 8: 529-569.

Teng S-C. 1963. Fungi of China. Sciences Press, Beijing, China. (in Chinese).

Wang X-H, Liu P-G, Yu F-Q. 2004. Color Altas of Wild Commercial Mushrooms in Yunnan. Yunnan Science and Technology Press, Kunming, China. (in Chinese).

Watling R. 1970. British Fungus Flora - Agarics and Boleti. I. Boletaceae, CU ai aR Paxillaceae. Her Majesty’s Stationery Office, Edinburgh, UK.

Watling R. 2001. Australian boletes: their diversity and possible origins. Aust. Syst. Bot. 14: 407- 416.

Wolfe JRCB. 1979. Austroboletus and Tylopilus Subgenus Porphyrellus with Emphasis on the North American Taxa. J. Cramer, Vaduz, Germany.

Ying J-Z, Zang M (eds). 1994. Economic Macrofungi from Southwestern China. Science Press, Beijing, China. (in Chinese).

Yuan M-S, Shun P-Q. 1995. Sichuan Mushrooms. Sichuan Science and Technology Press, Chengdu, China. (in Chinese).

Zang M. 1986. Notes on the Boletales from Eastern Himalayas and adjacent areas of China. Acta Bot. Yunnan. 8: 1-22. (in Chinese).

Zhang G-Y. 1999. Illustrations of China Popular Edible Mushrooms. Yunnan Science & Technology Press, Kunming, China. (in Chinese).

MYCOTAXON

Volume 96, pp. 47-50 April-June 2006

An annotated checklist of Leccinum in China

S.-Z. FU, Q.-B. WANG® & Y.-J. YAO"

yaoyj@sun.im.ac.cn 'Key Laboratory of Systematic Mycology and Lichenology Institute of Microbiology, Chinese Academy of Sciences Beijing 100080, China

*Graduate School of Chinese Academy of Sciences Beijing 100049, China

Abstract—Species of the genus Leccinum reported from China are reviewed and annotated with a summary of their current research status. A total of 32 names have been included from an extensive literature research. Among these names, two are regarded as synonyms of other species of the genus and another one transferred to Tylopilus. The taxonomic status of three names, the generic placement of two names and the application of one name to Chinese material are controversial. Among the remaining 23 names, seven were reported without specimen support and 16 with specimen citation. It is also found that the descriptions of six of these taxa from China differ from those from Europe and elsewhere. Specimen citation, morphology description and species identification of the Chinese records are also discussed. The full checklist is posted at http://www.mycotaxon.com/resources/weblists.html.

Key words—boletes, distribution, nomenclature

Leccinum Gray is a bolete genus with about 75 accepted species (Kirk et al. 2001) among some 282 proposed names (http://www.indexfungorum.org/Names/Names.asp, March 2006). The genus is worldwide, but distributed mainly in the northern temperate zone (Kirk et al. 2001). Species of Leccinum are ectomycorrhizal (Singer 1975) and play important roles in both ecology and economy. Ectomycorrhizal basidiomycetes, including the boletes, constitute an important component of forest fungal communities (Dahlberg et al. 1997, Baxter & Dighton 2001, Yan & Yao 2003) and forest ecosystems. It has been shown that ectomycorrhizal fungal diversity can determine plant productivity (Jonsson et al. 2001) and that ectomycorrhizal fungi benefit their hosts by enhancing mineral uptake, thus contributing to the recycling of nutrients in the ecosystem (Dames et al. 1999). Furthermore, all the known species of Leccinum seem to be edible; some are used for food and sold in markets as fresh, dried, salted or pickled products (Singer 1975). Leccinum extremiorientale, for example, is often found in the wild mushroom market in Kunming, Yunnan, China.

Species of Leccinum in China have been the subject of modern scientific studies since the early 20" century but such studies were infrequent until the 1980s. Baccarini

* Author for correspondence

48

(1905) reported L. scabrum from Qinling Mountains, Shanxi, and the same species was subsequently reported some 30 years later as Boletus scaber from Jiangsu (Teng 1936). Subsequently, two more Leccinum species were reported from Yunnan (Keissler & Lohwag 1937). Chiu (1948) recorded three Leccinum species in Boletus L. in his comprehensive study of boletes in the south-west of China, mainly Yunnan province. A further species was added by Teng (1963). Ina summary of all known Chinese fungi, Tai (1979) included six species of Leccinum. Zang carried out further research on boletes in the south-west of China from the 1980s and reported six Leccinum species, including one new to science (Zang 1986). Subsequently, 15 species were reported from the same region by Ying & Zang (1994). Some additional species of Leccinum were also included in local lists of fungi. In total, 32 names of Leccinum have been reported from China. They are listed in Table 1, with references and notes on their current status.

Among the 32 names in Table 1, two are regarded as synonyms of other species of the genus (L. nigrescens and L. rufum) and one (L. eximium) is transferred to Tylopilus. The taxonomic status of three (L. crocipodium, L. holopus and L. versipelle), the generic placement of two (L. chromapes and L. subglabripes), and the name application to Chinese material of one (L. rugosiceps) are controversial because of the simplicity and plasticity of morphological and anatomical characters.

Among the remaining 23 names, seven were reported without specimen support, ie. L. duriusculum, L. intusrubens, L. oxydabile, L. quercinum, L. subleucophaeum, L. subradicatum and L. variicolor, and 16 with specimen citation, i.e. L. albellum, L. ambiguum, L. atrostipitatum, L. aurantiacum, L. extremiorientale, L. griseum, L, hortonii, L. insigne, L. olivaceopallidum, L. potteri, L. roseofractum, L. rubropunctum, L. rubrum, L. scabrum, L. subgranulosum and L. subleucophaeum var. minimum. The descriptions of some of these taxa from China are different from those of the same taxa from Europe and elsewhere, i.e. L. olivaceopallidum, L. oxydabile, L. quercinum, L. rugosiceps, L. roseofractum and L. rubropunctum. A revision of these taxa is required to verify their occurrence in China.

Acknowledgements

The authors are grateful to Drs P. Roberts and T.-H. Li for serving as pre-submission reviewers and for their valuable comments and suggestions. This study is supported by the Programme for Key International Science and Technology Cooperation Project of P. R. China (2001CB711103) from Ministry of Science and Technology, the National Science Fund for Distinguished Young Scholars (30025002) from National Natural Science Foundation of China, the Key Research Direction of Innovation Programme (KSCX2-SW-101C) and the scheme of Introduction of Overseas Outstanding Talents, operated by the Chinese Academy of Sciences.

Literature Cited

AS [Mycological and Lichenological Expedition to Shennongjia, Academia Sinica] 1989. Fungi and Lichens of Shennongjia. World Publishing Corp., Beijing, China. (in Chinese).

Baccarini P. 1905. Funghi dello Schensi septentrionale raccolti dal Padre Giuseppe Giraldi. Appunti Annali Nuovo Giorno Botanica Italiana 22 (nuova serie): 689-698.

Baxter JW, Dighton J. 2001. Ectomycorrhizal diversity alters growth and nutrient acquisition of grey birch (Betula populifolia) seedlings in host-symbiont culture conditions. New Phytologist 152: 139-149.

Table 1. Species of Leccinum in China

Name

L. albellum (Peck) Singer

Place of publication

Mycologia 37: 799 (1945)

Report from China

Yeh & Chen(1982)

49

AAA HE NS be Pa Oe ALB IG TOMS Ce SIN SEE aa he ck iy ees abla an eg Uk Beal 1937) ee erate eed? & Lohwag

L. duriusculum (Schulzer ex

1

eT ety dex wxéuihaly eed deine ees tes cere: cu Fuge rs ted Vandel Ges RMI Ct Stage aie 6 wild pede OL) t sre cot et F = Tylopilus L. eximium (Peck) Singer Persoonia 7: 319 (1973) Chiu (1948) eximius PA me im he debra. Arie FOR. fk a ein, ACE lle >” Ne plepten( Peck Singers. L. extremiorientale (Lar.N. Agar. Mod. Tax. 2nd Ed.: 744 i AS (1989) Vv fee, V; ASSINEVA). SINGER. ere DIOR rcsessrinin ining gage pagan titiiiieriiinnnninninin L. griseum (Quel.) Singer Die Rohrlinge II: 89 (1967) ee wis V He rolopee (lon Watling Trans. Br. Mycol. Soc. 43: 692 eon ; Ce on hea Res oro yee nee Vase ae oe ; Ree Es, ede Prono CoNaegseiaret! tertile OS} ow. Stet eens Je Mee CL OOEY MED EIU See ek” Aiba: cape cen Yeh & Chen L. insigne A.H. Sm.et al. Michigan Bot. 5: 160 (1966) (1981) V L. intusrubens (Corner) Hoil. Nordic J] Bot. 2: 270 (1982) Wu et al. (1997) erect Richon ec Kozs) Amer, Midl. Nat. 37: 116 (1947) Chiu (1957) a aaa sa Inger in er TR pp Gon og geen es ca ge ecg ee al eer ag crocipodium........ 3 poe fe RS Ot Michigan Bot. 6: 125 (1967) Bi et al. (1997) # L. oxydabile (Singer) Singer Amer. Midl. Nat. 37: 116 (1947) | Mao (2000) et _, potteri AH. Sm.etabe Michigan Bot. 5: 1381966), Dai & Li(1994) L. quercinum E.E. Green & Notes R. bot. Gdn. Edinb. 29: e a Utunwtatiateles(D, xia whavorI51 C968\K waycateiuiel levab uh Saea ai mal ae PLE neler, A Notes. R. bot. Gdn. Edinb. 28: : eee on arts ns SIG (IDGR iene ren. reed, ENE. Cee el Cen L. rubropunctum (Peck) Singer Amer. Midl. Nat. 37: 117 (1947) Chiu (1948) # L. rubrum M. Zang Bee Ee eat Zang (1986) V L. rufum (Schaeff.) Kreisel Boletus Schr Reihe 1: 30 (1984) Mao (2000) ins L. rugosiceps (Peck) Singer Mycologia 37: 799 (1945) ponieee Hen if ssc ood ee Ce eh Be, oe reer a Tie Re a ae at. Arr. Brit. Pl. (London) 1: ie et ON Yn thoes GAG (ARDY Userart abtes Sosle BAC CATAL (1205) esl eae pele, L. subglabripes (Peck) Singer Mycologia 37: 799 (1945) Tai (1936-1937) t We elec a Pr gure LU mal agg Ving & Zang LY ER te tet ee ts, 4 ee ee ee a eee eee pee Bs ea Mycologia 52: 453 (1960) Li & Song (2000) —* ee eee ene stoner nnnrnentitcnannecann ite a rater ian, Mepemunienc’® Marcols5in«3: 701,199) peli AMC aI Stilt L. subradicatum Hongo Mem. Shiga Univ. 23: 41 (1973) Mao (2000) : Ee variicolor Watling ee a ee L. versipelle (Fr.) Snell Lloydia 7: 58 (1944) Liu (1991) t

\VRecognizable species. {Controversial in taxonomic status, generic placement and application of name to Chinese material. #To be confirmed; records different in description from the western literature. *To be confirmed; records lacking specimen support.

50

Bi Z-S, Li T-H, Zheng G-Y, Li C. 1984. Basidiomycetes from Dinghu Mountain of China. III. Some species of Boletaceae (2). Acta Mycologica Sinica 3: 199-206. (in Chinese).

Bi Z-S, Li T-H, Zhang W-M, Song B. 1997. A Preliminary Agaric Flora of Hainan Province. Guangdong Higher Education Press, Guangzhou, China. (in Chinese).

Chiu W-F. 1948. The boletes of Yunnan. Mycologia 40: 199-231.

Chiu W-F. 1957. Atlas of the Yunnan Boletes. Science Press, Beijing, China. (in Chinese).

Dahlberg A, Jonsson L, Nylund JE. 1997. Species diversity and distribution of biomass above and below ground among ectomycorrhizal fungi in an old-growth Norway spruce forest in south Sweden. Canadian Journal of Botany 75: 1323-1335.

Dai X-C, Li T-H. 1994. Macrofungus Flora of Ganzi, Sichuan Province. Sichuan Technology and Science Press, Chengdu, China. (in Chinese).

Dames JF, Straker CJ, Scholes MC. 1999. Ecological and anatomical characterization of some Pinus patula ectomycorrhizas from Mpumalanga, South Africa. Mycorrhiza 9: 9-24.

Jonsson L, Nilsson MC, Wardle DA, Zackrisson O. 2001. Context dependent effects of ectomycorrhizal species richness on tree seedling productivity. Oikos 93: 353-364.

Keissler K, Lohwag H. 1937. Fungi in Handel-Mazzetti. Symbolae Sinicae 2: 1-79.

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MYCOTAXON

Volume 96, pp. 51-60 April-June 2006

New species and new records of Graphis from India: transseptate species with completely carbonized exciples and norstictic acid

BHARATI ADAWADKAR & URMILA MAKHYJA

UV_Makhija@hotmail.com Agharkar Research Institute G.G. Agarkar Road, Pune - 411004, India

Abstract—The present paper gives an account of ten species of the genus Graphis. G. albidofarinacea, G. cinnamomea, G. colliculoides, G. kollaimalaiensis, G. nigrocarpa, G. nilgiriensis, and G. verruciformis are described as new to science. Graphis flavens, G. induta, and G. marginata are recorded for the first time from India. The two species G. flavens and G. nilgiriensis without norstictic acid are described in an appendix.

Keywords—lichenized fungi, ascomycetes, taxonomy, Graphidaceae

Introduction

The Graphidaceae is a large and widely distributed group of mainly tropical lichens with about 918 species worldwide (Kirk et al. 2001). The clear concepts of the genera (and several species) are available in the recent treatment of Graphidaceae (Staiger 2002).

Awasthi (1965), in his “Catalogue of lichens from India, Nepal, Pakistan and Ceylon,’ enumerated 36 species of Graphis Adans., which were recorded in the nineteenth and the early twentieth centuries by various lichenologists—viz. Bélanger (1838), Leighton (1869), Stirton (1879), Nylander (1867, 1891, 1900), Jatta (1905) and Rasanen (1952)— based on the collections of European botanists or naturalists from India.

Lichenological studies in India were resumed beginning in the early 1940s by Dr. D.D. Awasthi, Dr. P.G. Patwardhan, and their associates. Subsequently 33 new species were recorded and published in various scattered publications by Indian lichenologists [Awasthi & Singh (1975) Awasthi & Singh (1977), Patwardhan & Kulkarni (1976, 1979), Kulkarni (1977), Nagarkar & Patwardhan (1982), Singh & Sinha (1994) and Makhija et al. (1992)].

Awasthi (2000) lists 74 species of Graphis so far known from India in his “Lichenology in Indian subcontinent”. We are currently undertaking a detailed survey of this family, which is particularly well represented in India.

As a result of a major revision of the spore genus Graphis and studies of recent collections of this genus from India we have discovered approximately 145 species that can be transferred to different genera following the new system set forth by Staiger (2002). Consequently, the transseptate species in the genera —namely, Acanthothecis,

a2

Dyplolabia, Fissurina, Graphis, Hemithecium, Platythecium and Thalloloma are described elsewhere (Makhija & Adawadkar 2003, 2005a,b; Makhija et al. 2005; Adawadkar & Makhija 2004, 2005).

In the present account we describe ten species in the genus Graphis with completely carbonized exciple and colourless, transseptate ascospores from India. Graphis flavens, G. induta, G. marginata are recorded for the first time from India, and G. albidofarinacea, G. cinnamomea, G. colliculoides, G. kollaimalaiensis, G. nigrocarpa, G. nilgiriensis, and G. verruciformis are new to science. The two species G. flavens and G. nilgiriensis without norstictic acid are described in an appendix but are not included in the key.

Materials and Methods

In the present work chemical data has been obtained by thin-layer chromatography using methods standardized for lichen products (Culberson & Kristinsson 1970, Culberson 1972, White & James 1985). All specimens were examined under UV light (365 nm).

We have also consulted morphological and chemical data obtained from the re- examination of type or authentic material of several taxa in the Graphidaceae together with photographs of type specimens from the late Dr. Mason Hale, who provided materials for our studies during his several visits to our laboratory at ARI.

Key to the Indian Graphis species with norstictic acid, completely carbonized exciples and transseptate ascospores

VaAscomataiconcolorous with thethalliis) 42, eas ee eet en se eee ee 2 Th Ascomata dark OV Wiehe cere cee ee 6 ei oe iy oite oo ce encinclec a eyecare eee ee ener 4

2a. Ascomata short, up to 5 mm long. Thallus pale grayish brown to brownish gray, thick, warty; ascomata 1-5 mm long, concolorous, simple to branched, immersed; disc narrow; ascospores hyaline, 8-11-transseptate, 33-63 x 4-8 um; norstictic acid present............ G. induta

2b; Ascomata long “note shane tim long, tr eee os 9 Se aera oa a tcalerae 3

3a. Thallus pale grayish white, rough, warty; ascomata 4-8 mm long, simple to branched, immersed to semi-immersed, terminally acute to subacute; disc white pruinose;

ascospores hyaline, 12-14-transseptate, 55-71 x 6-8 um; norstictic acid present ho pene Re eR: thee Sec ia me Ee unis ies eee G. albidofarinacea

3b. Ascomata concolorous with the thallus.

Thallus greenish gray, glossy, uneven; ascomata 1-8 mm long, concolorous, simple to rarely branched; disc white pruinose; ascospores hyaline, 18—21-transseptate, 96-105 x 10-12 um; norstictic and con-norstictic acids present ... G. marginata

4a. Salazinic acid present. Thallus greenishyellowto yellowish brown, finely cracked; ascomata 0.3-8 mm long, black, simple to rarely branched, immersed, terminally acute to subacute; disc cinnamon coloured in wet condition; ascospores 7-15-transseptate, 21-42 x 5-8 (un; norsticticand/salazinic.acids presenti: 917, Blli2. HOY. Sey. G. cinnamomea

AlyeSalazinice CIC ADSENIE. usc cree ee ree ee Te te cena ae Eee ean aaa aee ee 5

bawmscospores upto 50 fim long © 2 eee seid ie ames ae Stele Oe ed Greet? 6 Sbs Ascospores more,tham 50 (Um One. «asi qaikee Gran Saey ucies ia elisae esd <Slarake> 6: 7

6a. Thallus dull greenish gray, warty; ascomata 1-4 mm long, black; ascospores 6-12- transseptate, 37-50 x 4-6 um; norstictic acid present ......... G. verruciformis

6b. Thallus grayish, evanescent; ascomata 0.5-2 mm long, black, simple to branched, semi-emergent, ends round; ascospores 5-9-transseptate, 21-27 x 4-6 um; norstictic. acidiprésentssiyitta hss) oh) art emetiepres ley fa tae < G. colliculoides

7a. Thallus green, flaking; ascomata 1-5 mm long, black, simple, triradiately to sparsely and irregularly branched; ascospores 9-13-transseptate, 50-71 x 8-12 um; MORStiCtiGAcic. Presehtats dismay aa seer, bees ice te D G. kollaimalaiensis

7b. Thallus greenish to yellowish buff; ascomata 1-10 mm long, black; exciple present below; ascospores 8-14-transseptate, 33-63 x 4-6 um; norstictic acid present me lore |) PPAR, eee ens Tet a SPE RR nce etnt gee te TLL OCMI PO

Taxonomic descriptions

Graphis albidofarinacea Adaw. & Makhija sp. nov. Figure 1

Similis Graphis immersa Fink sed thallo farinaceo, ascomata albo pruinoso et ascosporis majoribus differt.

Etymology: from the Latin albidus, whitish and farinosus, mealy; a reference to whitish, effuse thallus.

Holotype-India, Tamil Nadu, Kodaikanal, 24.9.1973, PG. Patwardhan & M.B. Nagarkar, 73.2254; holotype: AMH.

Thallus whitish grayish-green, rough, unevenly thickened, warty, somewhat effuse to granular. Ascomata lirelline, 4-8 mm long, 0.2-0.3 mm broad, simple to branched, irregularly spread, immersed to slightly emergent, terminally acute to obtuse. Disc black when exposed, white pruinose. Exciple present at the base, convergent, covered by a thalline margin up to the top, not striate, laterally to completely carbonized. Epithecium hyaline, thin. Hymenium hyaline, not inspersed, 138-232 um high and 168-315 um broad, I-, KI-. Hypothecium orange yellow, thick, 42-63 um high. Paraphyses simple, long, thin. Asci 6-8 sporate. Ascospores hyaline, 12-14-transseptate, ellipsoidal, 55-71 x 6-8 um, I + blue.

Chemistry- Norstictic acid present.

Specimens examined-Tamil Nadu: Kodaikanal, Munnar road, PG. Patwardhan & M.B.

Nagarkar, 73.2248, 73.2253; Kodaikanal, C.R. Kulkarni & PD. Badhe, 73.2247. 73.2249; PG. Patwardhan & C.R. Kulkarni, 73.2251 (AMH).

Remarks-The new species Graphis albidofarinacea is comparable to G. immersa in the nature of the exciple, but differs from the latter species in having white mealy thallus, ascomata covered with white pruina and large ascospores. Ascospores are 38-45 um long in G. immersa.

Graphis albidofarinacea closely resembles an Australian species G. stenospora var. deficiens A.W. Archer and G. catherinae A.W. Archer but they have no lichen substances in their thalli.

54

Graphis cinnamomea Adaw. & Makhija sp nov. Figure 2 Similis Graphis adpressa Vain. sed acida norsticticum et salazincium continens differt.

Etymology: from the Latin cinnamomeus, mixed with yellowish and red; a reference to disc colour.

Holotype-India, Tamil Nadu, Kodaikanal, near Golf Club, 24.1.1975, PG. Patwardhan & A.V. Prabhu, 75.184; holotype:AMH.

Thallus greenish yellow to yellowish brown, thick, delimited by thin, black hypothallus. Ascomata lirelline, black, 0.3-8 mm long and 0.1-0.2 mm broad, simple to branched, immersed, terminally obtuse. Disc narrow, black, cinnamon coloured when wet, pruinose. Exciple present at the base, convergent, covered by a thalline margin up to the top, not striate, laterally to completely carbonized. Epithecium hyaline to light brown, thin, K-. Hymenium hyaline, not inspersed, 52-87 um high and 105-157 um broad, I-, KI-. Hypothecium hyaline to light orange, 17-21 um thick. Asci 8 sporate. Ascospores hyaline, 7-15-transseptate, ellipsoidal, 21- 42 x 5-8 um, I+ blue.

Chemistry- Norstictic and salazinic acids present.

Specimens examined-Tamil Nadu: Sim’s Park, Coonoor, PD. Badhe & C.R. Kulkarni, 73.814; Niligiri hills, Korakundha, PG. Patwardhan 73.1473 (AMH).

Remarks-Graphis cinnamomea is characterized by mostly simple ascomata with cinnamon coloured disc covered by white pruina, the not striate, laterally to completely carbonized exciple and the presence of norstictic and salazinic acids. The new species is distinguished from the externally similar G. adpressa by the absence of lichen substances and larger ascospores of 46-58 x 12-5 um in that species. The combination of norstictic and salazinic acids is rather rare in Graphis.

The most closely related Graphis emersa Mill. Arg., a species from Queensland, differs from the new species in having only norstictic acid, while the new species has salazinic acid in addition to the norstictic acid in its thallus.

Graphis colliculoides Adaw. & Makhija sp. nov. Figure 3

Similis Platythecium colliculosum (Mont.) Staiger sed excipulo omnino carbonaceo et acidum norsticticum continens differt.

Etymology: from the species name colliculosum.

Holotype-India, Tamil Nadu, Devicolam, 25.1.1976, PG. Patwardhan, 76.701; holotype: AMH.

Thallus grayish green, thin, mostly evanescent, sparsely cracked with age, delimited by thin, black hypothallus. Ascomata lirelline, 0.5-2 mm long, 0.1-0.2 mm broad, black, simple to branched, semiemergent, terminally round. Disc black, narrow to moderately broad, epruinose. Exciple convergent, covered by a crystal studded thalline margin up to the top, not striate, completely carbonized. Epithecium hyaline to light brown, K-. thin. Hymenium hyaline, not inspersed, 96-105 um high and 147-155 um broad, I-, KI-. Hypothecium hyaline to pale, thin. Asci 8 sporate. Ascospores hyaline, 5-9- transseptate, ellipsoidal, 21-27 x 4-6 um, I + blue.

Chemistry- Norstictic acid present. Remarks-The present new species resembles Platythecium colliculosum in its appearance.

a,

However, Platythecium colliculosum has non carbonized exciple and has salazinic and protocetraric acids in its thallus.

It is somewhat similar to Graphis centrifuga Rasanen and G. semiaperta Mill. Arg. with respect to the characters of ascospores and the chemistry but G. centrifuga has the hymenium inspersed and has large lirellae arranged in well defined stellate clusters, where as G. semiaperta has mostly simple lirellae and a black epithecium.

Graphis induta Miill. Arg., Hedwigia 30: 185, 1891. Figure 9

Thallus buff with pinkish tinge, pale grayish brown to brownish gray, thick, distinctly warty, cracked. Ascomata lirelline, 1-5 mm long, 0.1 mm broad, concolorous with the thallus, simple to branched, immersed, crowded, terminally acute to obtuse. Disc blackish, covered by white pruina. Exciple present at the base, convergent, covered by a thalline margin up to the top, not striate, completely carbonized. Epithecium hyaline, thin, K-. Hymenium hyaline, not inspersed, 120-150 um high and 170-200 um broad, I-, KI-. Hypothecium hyaline, thin. Asci 4-6 sporate. Ascospores hyaline, (4-) 8-11- transseptate, ellipsoidal, 33-63 x 4-8 um, I + blue.

Chemistry- Norstictic acid present.

Specimens examined- Tamil Nadu: Kollaimalai, M.B. Nagarkar & P.G. Patwardhan 85.1625, 85.1626 (AMH).

Remarks-Graphis induta can easily be distinguished by its distinctly warty, buff coloured thallus with pinkish tinge; concolorous, crowded ascomata; disc black, white pruinose; not striate, completely carbonized exciple; (4-) 8-11 -transseptate ascospores of 33-63 x 4-8 um.

The species resembles Graphis anguilliformis Taylor but that is distinguished from the present species by the tall, heavily carbonized, robust exciple, larger (75-110 um long) ascospores and absence of lichen substances in G. anguilliformis.

Graphis kollaimalaiensis Adaw. & Makhija sp. nov. Figure 6

Similis Graphis turgidula var. norstictica A.W. Archer sed lirellis longiorobus et ascosporis locularibus minores differt.

Etymology: from the Latin ensis, a place of origin, and Kollaimalai, the type locality.

Holotype-India, Tamil Nadu, Kollaimalai, 15.10.1985, M.B. Nagarkar & PG. Patwardhan, 85.1526; holotype:AMH.

Thallus green, flaking, cracked, thick. Ascomata lirelline, 1-5 mm long, 0.1 mm broad, black, simple to sparsely branched, straight to curved, flat to slightly raised above, scattered, terminally obtuse. Disc narrow to slightly broad, dark blackish brown. Exciple present at the base, convergent, covered by a thalline margin up to the top, not striate, completely carbonized. Epithecium hyaline, thin. Hymenium hyaline, not inspersed, 150-160 um high and 200-210 um broad, I-, KI-. Hypothecium hyaline, thin. Asci 8 sporate. Ascospores hyaline, 9-13-transseptate, ellipsoidal, 50-71 x 8-12 um, I + blue.

Chemistry- Norstictic acid present.

Specimens examined-Tamil Nadu: Kollaimalai, PG. Patwardhan & M.B. Nagarkar 85.1525, 85.1636, 85.1637 (AMH).

56

Remarks-The present new species seems to be similar to Graphis tenuescens Nyl. from which it differs in having 8 sporate asci. In G. tenuescens the asci are 1-2 sporate.

It can easily be distinguished from the somewhat similar species Graphis leucoparypha Kremp. by its thick, flaking thallus, much conspicuous lirellae and with only 9-13- transseptate ascospores. Graphis leucoparypha has (15-) 16-18 (-20) locular ascospores and has only 0.5-2(-3) mm long lirellae.

Graphis marginata Raddi, Memor. Soc. Ital. Sc. 18: 344, 1820. Figure 5

Thallus greenish gray, thick, uneven, cracked, glossy. Ascomata lirelline, 1-8 mm long, 0.1-0.2 mm broad, concolorous, simple to rarely branched, immersed to slightly raised, scattered, terminally round. Disc very narrow, black, covered by white pruina. Exciple present at the base, convergent, covered by a thalline margin up to the top, not striate, slightly irregular outline, thick, completely carbonized. Epithecium brownish, thin. Hymenium light orange yellowish tinged, 176-189 um high and 163-168 um broad, not inspersed, I-, KI-. Hypothecium pale yellowish, 6-8 um high. Asci 6-8 sporate. Ascospores hyaline, 18-21-transseptate, ellipsoidal, 96-105 x 10-12 um, I + blue.

Chemistry- Norstictic and con-norstictic acids present.

Specimens examined-Tamil Nadu: Anamalai hills, Topslip, M.B. Nagarkar & U.V. Makhija, 82.142, 82.173 (AMH).

Remarks-Graphis marginata resembles G. congesta in respect of external morphology and exciple nature. However, G. congesta has much smaller ascospores of 50-75 x 9-15 um in size and only norstictic acid in its thallus. G. marginata was previously reported from Brazil and is here reported for the first time from India. In India this species has been collected in semi-evergreen forest at higher elevation.

Graphis nigrocarpa Adaw. & Makhija sp. nov. Figure 7 Similis Graphis kollaimalaiensis sp. nov. sed lirellis longioribus et ascosporis angustibus differt.

Etymology: from the Latin niger, black and from Greek karpos, fruit, a reference to black lirellae.

Holotype-India, Tamil Nadu, Munnar to Kodai, 24.1.1976, M.B. Nagarkar & K.D. Gole, 76.623; holotype: AMH.

Thallus dull yellow, yellowish buff, unevenly thickned, delimited by thin black hypothallus. Ascomata lirelline, 1-10 mm long and 0.1-0.2 mm broad, black, simple to irregularly branched, flexuose, immersed to slightly raised, terminally acute to subacute. Disc narrow, indistinct. Exciple present at the base, convergent, covered by a thalline margin up to the top, crystals studded, not striate, completely carbonized. Epithecium hyaline, thin, K-. Hymenium hyaline, clear to sometimes inspersed with crystals, 126- 168 um high and 84-168 um broad, I-, KI-. Hypothecium hyaline, 6-8 um high. Asci 4-6 sporate. Ascospores hyaline, 8-14-transseptate, ellipsoidal, 33-63 x 4-6 um, I + blue.

Chemistry- Norstictic acid present.

Specimens examined-Tamil Nadu: Munnar to Kodai, M.B. Nagarakar & K.D. Gole, 76.586, 76.587, 76.113 (AMH).

57

Figures1-10. Habit. 1. Graphis albidofarinacea, 2. G. cinnamomea, 3. G. colliculoides, 4. G. flavens, 5. G. marginata, 6. G. kollaimalaiensis, 7. G. nigrocarpa, 8. G. nilgiriensis, 9. G. induta, 10. G. verruciformis. Bar = 2mm

58

Remarks-Graphis nigrocarpa differs from G. kollaimalaiensis in having much larger (up to 10 mm long) irregularly branched ascomata and has narrower ascospores. This species has been collected in montane forest.

Graphis nigrocarpa can easily be separated from G. rustica Kremp. by the presence of stictic acid, slightly larger ascospores (66-76 x 9-10 um) and more number of locules (16-18 locules) of the ascospores in G. rustica.

The new species seems to be very similar to Graphis catherinae but that species lacks lichen substances.

Graphis verruciformis Adaw. & Makhija sp. nov. Figure 10 Similis Graphis congesta (Fée) Miill. Arg. sed ascosporis minoribus differt. Etymology: from the Latin verruca, wart; a reference to warty thallus.

Holotype-India, Kerala, near Chinnar, Munnar to Udumalpet, 19.10.1985, PG. Patwardhan & M.B. Nagarkar, 85.1756; holotype:AMH.

Thallus dull grayish green, smooth to thick, finely and distinctly cracked, rough, distinctly warty. Ascomata lirelline, 1-4 mm long, 0.1 mm broad, black, simple, semi- emergent, scattered, terminally acute to obtuse. Disc narrow to moderately broad, dark blackish brown, pruinose. Exciple present at the base, convergent, covered by a thalline margin up to the top, not striate, completely carbonized. Epithecium hyaline, thin, K-. Hymenium hyaline, not inspersed, 110-130 um high and 150-190 um broad, I-, KI-. Hypothecium hyaline, thin. Asci 8 sporate. Ascospores hyaline, 6-12-transseptate, ellipsoidal, 37-50 um x 4-6 um, I + blue.

Chemistry- Norstictic acid present.

Remarks-Graphis verruciformis differs from the closely related G. congesta in having smaller ascospores of 37-50 um x 4-6 um. In Graphis congesta ascospores are rather larger of 50-75 um long.

Appendix

Graphis flavens Mill. Arg., Flora 85: 334, 1882. Figure 4

Thallus yellowish green, uneven, cracked, delimited by thin, black hypothallus. Ascomata lirelline, 1-2.5 mm long, narrow, mostly simple to occasionally branched, concolorous with the thallus, scattered, immersed to semiemergent, irregularly curved, flexuose, terminally obtuse. Disc narrow, black, epruinose. Exciple present at the base, convergent, covered by a thalline margin upto the top, crystal studded, not striate, completely carbonized. Epithecium brownish, thin, K-. Hymenium hyaline, not inspersed, 97-105 um high and 100-105 um broad, I-, KI-. Hypothecium hyaline, 8-12 um thick. Asci 8 sporate. Ascospores hyaline, 14-18-transseptate, ellipsoidal, 29-63 x 8-12 um, I+ blue.

Chemistry- No lichen substances present.

Specimens examined-Tamil Nadu: Agasthi hills, Upper Kodayar, on road side tree, on Kakachi road, elev. approx. 1400 m., PG. Patwardhan & P.K. Sethy, 84.76, near Chinnar, Munnar-Udumalpet, M.B. Nagarkar & P.G. Patwardhan, 85.1751 (AMH).

59

Remarks-Graphis flavens, a species previously reported from Java, is reported for the first time from India. This species has been collected in undisturbed forest at higher elevation, in montane forest.

Graphis nilgiriensis Adaw. & Makhija sp. nov. Figure 8

Similis Graphis proserpens Vain. sed excipulo omnino carbonaceo differt et acidum sticticum continens.

Etymology: from the Latin ensis, a place of origin, and Nilgiri, the type locality.

Holotype-India, Tamil Nadu, Nilgiri hills, Naduvattam, in Shola forest, app. 5000 ft., 25.2.1978, PG. Patwardhan, 78.109; holotype:AMH.

Thallus yellowish green, thick, smooth, glossy, delimited by thin, black hypothallus. Ascomata lirelline, 1-5(-7) mm long and 0.1 mm broad, simple to branched, black, raised, scattered, narrow, terminally acute to subacute. Disc narrow to somewhat broader. Exciple convergent, covered by a thick thalline margin till the top, striate, 5-6 striae on each side, striae fused, much expanded at the apical region, completely carbonized. Epithecium hyaline, thin. Hymenium hyaline, not inspersed, 60-70 um high and 120- 149 um broad, I-, KI-. Hypothecium indistinct. Asci 8 sporate. Ascospores hyaline, 9-13-transseptate, ellipsoidal, 38-42 x 4-6 um, I+ blue.

Chemistry- Stictic acid present.

Remarks-Graphis nilgiriensis is characterized by its stramineous thallus; conspicuous, black, narrow, semi-emergent ascomata; striate, completely carbonized exciple; 38-42 um long ascospores and presence of stictic acid. It resembles to Graphis proserpens in gross morphology, but differs from the latter in having completely carbonized exciple and fewer striae. G. proserpens has partially carbonized and multistraite exciple. G. persulcata Stirt., which resembles closely to the present species, however, lacks lichen substances. This lichen has been collected from evergreen montane forest.

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Makhija U, Adawadkar B. 2005b. Some new species of Graphis (Lichenised Ascomycota) from the Andaman and Nicobar Islands of India. Mycotaxon 91: 369-379.

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Graphis. Biovigyanam 8: 125-131.

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MYCOTAXON

Volume 96, pp. 61-71 April-June 2006

Four species of Caloplaca (Teloschistaceae, lichenized Ascomycota) new to Poland

KARINA WILK! & ADAM FLAKUS?

'palka@ib-pan.krakow.pl *flakus@ib-pan.krakow.pl Laboratory of Lichenology, W. Szafer Institute of Botany Polish Academy of Sciences, Lubicz 46, PL-31-512 Krakow, Poland

Abstract The species Caloplaca conciliascens, C. obliterans, C. percrocata and C. vitellinaria are reported as new to the lichen biota of Poland. All of them were collected in the Polish Carpathians, mainly in the Tatra mountains. Detailed taxonomic descriptions and comments are provided.

Key words lichenized fungi, taxonomy, geographical distribution

Introduction

The lichen genus Caloplaca Th. Fr. has been the subject of taxonomic surveys by many authors. Most recently European species of the genus were studied in detail by Giralt et al. (1992), Laundon (1992a), Nimis (1992), Roux & Navarro-Rosinés (1992), Sochting (1994), Navarro-Rosinés & Roux (1995), Navarro-Rosinés & Hladun (1996), Sochting & Stordeur (2001) and Khodosovtsev (2001, 2002), Arup (2006). North American Caloplaca species have been investigated by Arup (1992ab, 1993ab, 1994, 1995ab), Wetmore (1994, 1996a, 2001, 2003, 2004) and Wetmore & Karnefelt (1998, 1999). Antarctic species are also fairly well known (e.g., Sachting & @vstedal 1992; Olech & Sechting 1993; Sochting & Olech 1995, 2000). Polish species of the genus have not been investigated in detail, however, and are in need of taxonomic revision. Currently 67 taxa have been reported from Poland (Nowak & Tobolewski 1975, Faltynowicz 2003), approximately 50 of which occur in the Polish Carpathians (Bielczyk 2003). New records of Caloplaca for Poland have recently been supplied by Kossowska (1999), Zalewska (2000), Bielczyk (2003), Sparrius (2003), and Ceynowa-Gieldon & Adamska (2005).

The paper is based mainly on collections made by the second author during fieldwork in the Polish Tatra Mts. in 2002-2004. The study area plays an important role for the preservation of biological diversity including lichens, not only in Poland but also on a European scale. Despite the long tradition of lichenological research in the Polish Tatra Mts., many species are still being discovered from the area (e.g., Bielczyk 2003, Czarnota 2004, Flakus 2004, Krzewicka 2004, Kukwa 2004, Flakus & Bielczyk 2006, Sliwa 2006).

The present paper contributes to the study of the genus Caloplaca in Poland and expands the knowledge of lichen diversity in the Polish Carpathians. Detailed taxonomic descriptions and comments are presented below.

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Taxonomic Descriptions

Caloplaca conciliascens (Nyl.) Zahlbr. FIGURE la Catal. Lich. Univ. 7, p. 109 (1931)

Thallus small, indistinct, areolate, greyish white. Apothecia sessile, clustered, angular, 0.5-1.0 mm diam. Disc flat to slightly convex, brownish red. Margin persistent, thin, slightly flexuose, black. Parathecium without algae, inside colourless, outside blackish green. Epihymenium brownish golden; hymenium hyaline, 60-75 um; hypothecium hyaline, with oil drops, ca. 80 um; paraphyses simple or with some branches in upper parts, tips with 1-2 slightly swollen cells. Spores 8 per ascus, 10-14 x 5-7 um, isthmus 3-4(-5) um.

Spot test reactions: thallus K-, cN+ weakly violet (lecidea green); apothecial margin K+ violet red, cN+ distinctly violet (lecidea green), H+ intensifying green; epihymenium K+ violet red; upper parts of paraphyses H+ weakly greenish.

Ecology At the collection site, C. conciliascens grew on mylonitized granite rock in shady place in the subnival belt of Tatra Mts. It was accompanied by Lecanora cenisia Ach.

Geographical distribution The species is known from Austria (Wunder 1974, Hafellner & Turk 2001), Italy (Nimis 1993) and Turkey (John 1996).

Comments Caloplaca conciliascens is characterized by a reddish black apothecial disc and brownish black margin. Wunder (1974) treated this taxon in the study on saxicolous species of Caloplaca with black coloured apothecia. The group of lichens is not homogeneous and this particular species has been considered as a member of C. ferruginea group (Wunder 1974, Karnefelt 1989). Wetmore (1996a) discussed C. conciliascens in his taxonomic treatment of C. siderites group in North and Central America. He concluded that C. conciliascens, C. exsecuta (Nyl.) Dalla Torre & Sarnth., C. concilians (Nyl.) H. Olivier, C. litoricola Brodo and C. britzelmayri (Hue) Zahlbr. are probably closely related and form one distinct group characterized by presence of “dark apothecial margin and lecidea green instead of thalloidima green” (Wetmore 1996a).

Caloplaca conciliascens is very similar to C. exsecuta, which is a fairly common species distinguished by a very thin, sometimes almost disappearing thallus, flat to strongly convex apothecia, higher hymenium (65-135 tm) and a hypothecium (50-125 um), which is always brownish coloured with numerous oil drops and wider spores (5.5-8.0 um) (Wunder 1974).

Other related species are: C. concilians that differs from C. conciliascens in having abundant algae in the apothecial margin, C. britzelmayri with dirty yellowish apothecium disc, larger spores that occupies calcareous rocks, and C. litoricola, which besides distinctive characters of the apothecia is a maritime species known from western coast of North America (Wetmore 1996a,b). Clauzade & Roux (1985) regarded the taxon as closely related to C. conversa (Kremp.) Jatta. Caloplaca conversa in comparison with C. conciliascens produces smaller apothecia of different anatomy - margins usually lecanorine, with distinct oval cells, never black or greenish coloured (Wunder 1974). The two species differ also due to their chemistry (see Wetmore 1996a).

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SPECIMEN EXAMINED POLAND, WESTERN CARPATHIANS, Tatry Wysokie Mts., Ciemnosmreczynska Przelaczka pass, N aspect, slope 30°, mylonite area, subnival belt, alt. 2115 m, 49°11'21"N, 20°02’59”E, on mylonite rock, 7 August 2004, A. Flakus 3046 (KRAM).

Caloplaca obliterans (Ny1.) Blomb. & Forssell FIGURE 1b Points-Forteckning, p. 69 (1880)

Thallus small, more or less irregularly shaped in outline, areolate, pale to dark orange. Areoles irregularly shaped, often minutely lobed, plane to convex, scattered or huddled in groups. Hypothallus thin, concolorous with thallus, characterized by dendritic extensions to rock cracks. Soralia in the centre of thallus, concave and eroded, yellowish orange. Apothecia numerous in the centre of thallus, partially innate or adnate, small, up to 0.5 mm diam. Disc flat to slightly convex, orange, surrounded by thin, concolorous or slightly lighter margin. Amphithecium with algae. Epihymenium brownish golden; hymenium hyaline, 50-65 um; hypothecium hyaline, 40-50 um; paraphyses with some branches, tips with 1-2 swollen cells, up to 5 um diam. Spores 10-11.5 x 4 um, isthmus 2-3 jm.

Spot test reactions: thallus K+ violet red; apothecial margin cN-; epihymenium K+ violet red.

Ecology At the collection site the species occupied a typical niche, viz.: an overhanging granite wall, in moist and shady place. It was accompanied by Buellia alboatra (Hoftm.) Th. Fr., Lecanora dispersa (Pers.) Sommerf., Pseudosagedia chlorotica (Ach.) Hafellner & Kalb and Xanthoria elegans (Link.) Th. Fr. The species is characteristic for the silicate community Caloplacetum obliterantis Wirth 1972 (see Wirth 1972, Obermayer 1993).

Geographical distribution According to Wirth (1995) the species occurs in Europe in boreal to montane and high montane zones of temperate zone. It was reported from Iceland (Orange 1990), Norway, Sweden, Finland, Russian Fennoscandia (Nordin 1972), United Kingdom (Wade 1965, Laundon 1992b, Coppins 2002), Denmark (Sochting & Alstrup 2002), Czech Republic (Vézda & Liska 1999), Slovakia (Pisut 1995, Pisut et al. 1996, Lisicka 2005), Ukraine (Kopachevskaya 1986, Oksner 1993), Germany (Wirth 1995), Austria (Obermayer 1993), Romania (Kondratyuk et al. 2003), Hungary (Verseghy 1971), Bulgaria (Mayrhofer et al. 2005), Italy (Nimis 1993), France (Harmand 1913, Ozenda & Clauzade 1970), Portugal (Llimona & Hladum 2001) and Spain (Etayo 1990). Besides Europe it occurs in Southwest Asia in Israel (Alon & Galun 1971) and Jordan (El-Oqlah 1992), in North America (Esslinger 1997, Fryday et al. 2001) and in Australia (Kalb 1996).

Comments Caloplaca obliterans, as a minutely lobate species containing anthraquinones, was included by Nordin (1972) in the Caloplaca subgen. Gasparrinia. More recently, Wetmore & Karnefelt (1998) carried out a revision of the lobate and subfruticose species of Caloplaca in North and Central America. The authors discussed the species in the paper but did not find any specimens of that taxon in the studied material.

According to Wetmore & Karnefelt (1998), C. obliterans should not be included in subgen. Gasparrinia as it does not develop distinct elongated lobes.

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Caloplaca obliterans resembles C. cirrochroa (Ach.) Th. Fr., C. proteus Poelt and C. decipiens (Arnold) Blomb. & Forssell, but those species occur on calcareous substrate. There are also some morphological differences between the taxa: C. cirrochroa and C. proteus, have a thallus consisting of more convex and longer lobes in comparison with C. obliterans and C. decipiens has rather lip-shaped or capitate soralia.

Moreover, Arup (1993a, 1995a) compared C. obliterans with two other sorediate species, i.e. C. flavogranulosa Arup and C. citrina (Hoftm.) Th. Fr. However, C. obliterans is well distinguishable due to the dark orange colour of thallus, concave and eroded soralia as well as distinct, dendritic hypothallus (Arup 1993a, 1995a).

Caloplaca obliterans has been reported from many European countries where it is usually collected in sterile form. The apothecia, which develop very rarely, are small, only up to 0.5 mm diam. Our collection of C. obliterans has a lower hymenium and hypothecium when compared to those provided by Alon & Galun (1971) - 70-90 um and ca. 90 um, respectively. Moreover, it is worth noting that our specimen is. partly fertile and so has more aggregated areoles lacking soralia and a poorly developed hypothallus.

SPECIMEN EXAMINED POLAND, WESTERN CARPATHIANS, Tatry Wysokie Mts., Dolina Rybiego Potoku valley, Mokra Wanta, in moist and shaded place, WWN aspect, alt. 1750 m, on overhanging granite wall, 17 August 2002, A. Flakus 83 (KRAM).

REFERENCE MATERIAL EXAMINED J. J. Havaas, Lich. Norv. Occid. 218 (KRAM).

Caloplaca percrocata (Arnold) J. Steiner FIGURE Ic Halacsy in Denkschrift. Math.-naturw. Classe Kais. Akad. Wiss. Wien, vol. LXI, p. 523 (1894)

Thallus thick or thin, areolate, whitish grey, sometimes with a tinge of bluish or brownish. Areoles irregular, with uneven surface, subfarinose, plane to convex, up to 3 mm diam. Hypothallus thin, dark grey and rather rarely visible. Apothecia numerous in the centre of thallus, aggregated in groups, 0.4 to 1.3 mm diam, often angular to flexuose, sessile from the beginning. Disc flat, dark, brownish orange to brownish red, contrasting with pale, yellowish orange, thick margin. Thalline margin pale yellow, soon excluded (only sometimes good visible). Parathecium with radiating hyphae; amphithecium with numerous algae. Epihymenium brownish golden; hymenium hyaline, up to 100 um; hypothecium hyaline, with oil droplets, ca. 60 um; paraphyses with some branches. Spores 15-18(-20) x 8-10(-12.5) um, isthmus 3-4(-5) um.

Spot test reactions: thallus K-, C-, cN+ weakly violet (thalloidima green); epihymenium K+ violet red. The thallus contains the anthraquinone 7-chloroemodin (detected by U. Sochting).

Ecology At the collection site the lichen occurred on base-rich siliceous shale on steep slopes in exposed places in the alpine belt. It was accompanied by Aspicilia sp., Eiglera flavida (Hepp) Hafellner, Polyblastia cupularis A. Massal. and Thelidium sp.

Geographical distribution The species was reported from Norway (Santesson et

al. 2004), Austria (Poelt 1960, Hoffmann et al. 1998, Tiirk & Berger 1999), Hungary (Verseghy 1994), Bulgaria (Mayrhofer et al. 2005), Ukraine (Kopachevskaya 1986,

65

ese

Figs. 1 A-D. Species of Caloplaca. A: C. conciliascens (A. Flakus 3046 KRAM)); B: C. obliterans (A. Flakus 83 KRAM); C: C. percrocata (A. Flakus 151 KRAM). D: C. vitellinaria (A. Flakus 5120 KRAM). Scale bar = 1 mm.

Oksner 1993), Croatia (KuSan 1953), Italy (Nimis 1993), France (Ozenda & Clauzade 1970, Roux 1976, Houmeau & Roux 1991), Spain (Egea 1984) and Portugal (Llimona & Hladun 2001). Additionally, the species was reported by Kondryatuk et al. (2004) from France, Byelorussia, Turkey and North Africa.

Comments According to Magnusson (1944a) C. percrocata belongs to the C. ferruginea group which consists of taxa closely related to C. ferruginea (Huds.) ‘Th. Fr. and united by ferrugineous coloured apothecia. Moreover, the species of the C. ferruginea group are characterized by containing the chemosyndromes based on 7-chloroemodin (see Sochting & Tonsberg 1997, Elix et al. 2000). This group is rather large and includes species representing all ecological groups of lichens, for example corticolous, lignicolous, muscicolous and saxicolous (Magnusson 1944a,b, Clauzade & Roux 1985, Roux & Navarro-Rosinés 1992).

More recently Wetmore (1996a) established a more restricted C. siderites group in which C. percrocata was included. This group consists of exclusively saxicolous species characterized by grey, whitish or brown thalli that lack anthraquinones with apothecia that are red or orange and do contain anthraquinones. Furthermore, Wetmore (1996a) reported the non-crystallizing pigment thalloidima green in most of the treated species, including C. percrocata. This pigment reacts with K, cN, 10% N, C and concentrated HCl giving a violet colour.

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Caloplaca percrocata is a rather characteristic taxon due to the prominent apothecia and the usually distinct areolate thallus. It could be confused with C. teicholyta (Ach.) J. Steiner, which has a scurfy-granular to sorediate surface of thallus (not at all or weakly areolate), uniformly red coloured apothecia with usually well visible white thalline margin.

Caloplaca erythrocarpa (Pers.) Zwackh differs in producing smaller and also uniformly red apothecia.

The species C. atroflava (Turner) Mong. and C. submergenda (Nyl.) H. Olivier may also be considered similar to C. percrocata but they differ in ecological preferences (both are acidophilous and hygrophilous) and some morphological details. Caloplaca atroflava forms dark grey thallus and apothecia without any thalline margin while C. submergenda has apothecia with persistent, greyish thalline margins and produces considerably smaller spores (Clauzade & Roux 1987).

Another acidophilous species, C. limitosa (Nyl.) H. Olivier, was also considered as possible to be mistaken with C. percrocata (see Nimis 1993). A detailed description of C. percrocata is available in several papers, e.g. Magnusson (1944a), Oksner (1993), Wetmore (1996a) and Kondryatuk et al. (2004).

SPECIMENS EXAMINED POLAND, Western Carpathians, Tatry Zachodnie Mts., Twardy Uplaz, N slope below Ciemniak Mt., alt. 1900 m, 49°14°12”N 19°54’12”E, on siliceous shale contains calcium, 30 August 2002, A. Flakus 151 (C, KRAM, MIN); and 9 July 2004, A. Flakus 2251 with K. Palka & B. Cykowska (KRAM; the duplicates of the specimen will be distributed by Wetmore in Telosch. Exs. Fasc. 4).

REFERENCE MATERIAL EXAMINED Poelt, Lich. Alp. no. 50 (KRAM).

Caloplaca vitellinaria Szatala FIGURE ld Ann. Hist.-Nat. Mus. Natl. Hungarici, ser. nov. 7, p. 276 (1956)

Thallus absent or very scanty, pale orange, over the thallus or apothecia of the host Candelariella vitellina (Hoffm.) Mull. Arg. Apothecia numerous, usually grouped and then angular, up to 1 mm diam. Disc flat to slightly convex, orange. Margin thin, paler than disc, flush and only in young apothecia slightly raised. Parathecium with radiating hyphae; amphithecium with numerous algae. Epihymenium brownish golden; hymenium hyaline, 60-70 um; hypothecium hyaline, ca. 30 um; paraphyses with branches, tips with 1-2 swollen cells, up to 5 um diam.; spores 9-14 x 5-8 um, isthmus (3-)4-5 um.

Spot test reactions: thallus K+ violet red; epihymenium K+ violet red; hymenium I+ blue; hypothecium, parathecium and amphithecium I-.

Ecology At the collection site the lichen colonized the thallus of Candelariella vitellina on sandstone and shale containing calcium in lower montane and alpine belts of Polish West Carpathians. It was accompanied by Acarospora sp., Lecanora dispersa and Lecidella stigmatea (Ach.) Hertel & Leuckert.

Geographical distribution The species is known from Slovakia (Pisut et al. 1996, Lisicka 2005), Austria (Hafellner & Tiirk 2001), Hungary (Szatala 1956), the Iberian Peninsula (Llimona & Hladun 2001) and Turkey (John 1996).

Comments Caloplaca vitellinaria is a parasitic species occurring on the thallus of Candelariella spp., e.g. C. vitellina, C. coralliza (Nyl.) H. Magn. (Szatala 1956, Poelt

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1969, Wirth 1995, Hafellner 2001). Caloplaca grimmiae (Nyl.) H. Olivier is another taxon colonising Candelariella spp., but it differs significantly by the colour of apothecia, which have reddish brown discs with blackish brown margins.

Caloplaca vitellinaria was treated by Clauzade & Roux (1985) in connection with the C. ferruginea group and was accepted by the authors at variety level of C. holocarpa (Hoffm.) A.E. Wade. The primary differences between C. vitellinaria and C. holocarpa apparently are their ecology, although there are some other distinguishing characters such as size and arrangement of apothecia (bigger and aggregated in case of C. vitellinaria).

Caloplaca vitellinaria is similar to C. insularis Poelt but the latter taxon has better developed thallus and smaller apothecia (up to 0.8 mm diam.) with medulla reacting distinctly I+ violet. Moreover, C. insularis is parasitic on the Aspicilia spp., i.e. A. candida (L.) Korb. (Poelt 1960), A. polychroma Anzi (Clauzade & Roux 1985).

It is worth noting that in the specimens of C. vitellinaria examined, both collected by us and A. Vézda, a considerably low hymenium (ca. 80 um) was observed compared to the original description where it reaches ca. 100-125 um (Szatala 1956).

SPECIMENS EXAMINED POLAND, Western Carpathians: Beskid Slaski Mts., Pasmo Baraniej Gory range, Pietraszyna village, Koczy Zamek Mt., in open, sunny place, alt. 847 m, 49°32’54”N 18°58732”E, on calcareous sandstone, 9 September 2005, K. Wilk 3838 (KRAM)); Tatry Zachodnie Mts., Twardy Uplaz, N slope below Ciemniak Mt., alt. 1900 m, 49°14'14”N 19°54'13”E, on siliceous shale contains calcium, 30 July 2005, A. Flakus 5120 (KRAM).

REFERENCE MATERIAL EXAMINED HUNGARY, VEspREm: supra balneas Tihany prope lacum Balaton, alt. 220 m, 17 August 1964, A. Vézda (KRAM-L-28050).

Acknowledgements

We sincerely thank C. Roux (Mirabeau, France) and U. Sochting (Copenhagen, Denmark) for peer reviewing the manuscript. We are grateful to L. Sliwa (Krakow, Poland) for valuable comments on the text and to C. Roux for important information on France bibliography. U. Sochting and C. Wetmore (Minnesota, U.S.A.) are acknowledged for confirming C. percrocata. Financial support for the study was provided to the second author by Polish Ministry of Education and Sciences for the years 2005-2007 (MEIN, grant no. 2 P04G 078 29).

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MYCOTA XON

Volume 96, pp. 73-76 April-June 2006

The status of Psilocybe floridana (Strophariaceae) in Callistosporium (Tricholomataceae)

ETELVINA GANDARA & GASTON GUZMAN

gandara@ecologia.edu.mx guzmang@ecologia.edu.mx Instituto de Ecologia Apartado Postal 63, Xalapa 91000, Veracruz, Mexico

Abstract— Evidence is presented indicating that the type specimen of Psilocybe floridana is the same type of Callistosporium psilocybe and that both names are synonyms of C. luteoolivaceum.

Keywords— synonymy, intracellular pigment

Introduction

During the study of type specimens of Psilocybe (Fr.) P. Kumm. for a second edition of the monograph on the genus Psilocybe (Guzman 1983), we found that the type of P. floridana (Murrill 1945) does not belong to Psilocybe, as Guzman (1983) previously stated. Guzman had previously considered that Murrill’s fungus was a Psathyrella sp. However, the intracellular pigment in the spores, basidia and hyphae, lack of clamp connections, and lack of a germ pore on the spores for P. floridana indicates this species is actually a Callistosporium Singer (1944). Notwithstanding, Singer (1944) had not considered Murrill’s species when he, together with Murrill (Singer 1944), described C. psilocybe. Singer (1946, 1970) maintained this position in subsequent papers. Interestingly, the specimens used for the description of C. psilocybe are the same ones that were used for the description of Psilocybe floridana. Singer (1946) considered C. psilocybe as a synonym of C. luteoolivaceum. In this paper we discuss the synonymy of Psilocybe floridana with Callistosporium luteoolivaceum.

Materials and methods

Available specimens of Psilocybe floridana and Callistosporium luteoolivaceum from FH, FLAS and MICH were studied. Microscopical study was made with sections of each basidioma mounted in 5% KOH, 3% NH sO and 1% Congo red (previously treated with 96% alcohol for rehydration). Spore shape was based on the quotient (Q), obtained from the length : width ratio following Bas (1969).

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Both Psilocybe floridana and Callistosporium psilocybe have the same holotype and synonyms of C. Juteoolivaceum

Callistosporium luteoolivaceum (Berk. & M.A. Curtis) Singer, Lloydia 9: 117, 1946. Figs. 1-6 = Agaricus luteoolivaceum Berk. & M.A. Curtis, Ann. Mag. Nat. Hist. III 4: 286, 1859. = Collybia luteoolivaceum (Berk. & M.A. Curtis) Sacc., Syll. Fung. 5: 215, 1887. = Callistosporium psilocybe Murrill & Singer, in Singer, Mycologia 36: 363, 1944. = Psilocybe floridana Murrill, Proc. Fl. Acad. Sci. 7: 126, 1945. Non Stropharia floridana Murrill, Mycologia 35: 536, 1943, according to the study of the type in FLAS: Murrill 8738 (FLAS-10170). For other synonyms, see Redhead (1982)

For a complete description, see Singer (1970). However, as the basidiomata show a wide variation in form and color, we present here some additional morphological features of the pileus, based on a study of the type of Psilocybe floridana, which includes a color drawing by Murrill at FLAS. The pileus is subcampanulate or convex to plane-convex, finally subumbilicate or umbilicate. All the basidiomata are yellowish- melleous to dark vinaceous brown as dried. The following microscopic features were observed: spores (4-) 4.5-5.5 (-6) x 3-4.5 um, Q = (1.1-) 1.2-1.6, broadly ellipsoid to ellipsoid, occasionally subglobose, without germ pore, thin-walled, smooth, hyaline or with a brownish vinaceous content, inamyloid. Basidia 20-28 (-30) x 4.5-7 um, 4- spored, cylindric-clavate, flexuous, thin-walled, hyaline or with same pigment as the spores. Pleurocystidia absent. Cheilocystidia absent. Subhymenium not differentiated or composed of parallel hyphae, 1.5-2 (-3) um wide, thin-walled, hyaline or brownish vinaceous from intracellular pigment. Hymenophoral trama regular, hyphae 3-7 um wide, thin-walled, hyaline or with brownish vinaceous intracellular pigment. Pileipellis with hyphae radially arranged, repent, (3.5-) 5-7 (-10) um wide, thin-walled, hyaline or with brownish vinaceous intracellular pigment. Stipitipellis a cutis, occasionally disrupted by small clusters of ascending hyphae, 3-7 um wide, thin-walled, hyaline. Clamp connections absent.

Habitat and Distribution- According to Singer (1944, 1946, 1970), Murrill (1945), Dennis (1951), Redhead (1982), Pegler (1983), Moser (1984) and Hongo (1989), this fungus is reported from North America, South America, the Caribbean, Europe and Asia. However, it seems that the concept of C. Iuteoolivaceum accepted by those authors encompasses several species, both temperate and tropical. A critical revision of some synonyms is necessary.

This fungus is reported on wood (logs, trunks and stumps) or rotting wood, frequently covered by mosses, rarely in soil, in temperate, subtropical and tropical forests. Pulido & Boekhout (1989) cited P. aff. floridana from Colombia, in high mountains.

Discussion—This collybioid fungus was considered as a Collybia by Saccardo (1987) and also by Murrill in the herbarium in the holotype of Psilocybe floridana. The basidiomata are also similar to a Psilocybe or Psathyrella. However, the intracellular pigment in spores, basidia and hyphae is the main feature to consider the species as Callistosporium, following Singer (1944, 1946, 1970), and as discussed by Redhead (1982). The thin-walled, ellipsoid and hyaline to pigmented spores and the absence of

75

Figs. 1-6. Callistosporium luteoolivaceum. 1, 2, 4: spores. 3, 5: basidia, 6: stipitipellis (1: holotype; 2, 3: holotype of both C. psilocybe and Psilocybe floridana; 4, 5, 6: Singer F-2651). Scale bar 10 um.

a germ pore are important features. However, Singer (1970) described the spores of Callistosporium luteoolivaceum having a germ pore. Material from the Czech Republic in F (Singer & Pouzar C-5546, F-1018787) was identified by Singer as Psilocybe floridana but it is in fact P laetissima Hauskn. & Singer (!). The confusion on the holotypes of both Psilocybe floridana and Callistosporium psilocybe, was because both Murrill (1944) and Singer (1944) published descriptions at the same time, based on the same herbarium material from Juniper Springs, in Marion Co., Florida, gathered by Murrill

76

on October 13, 1940 and recorded in FLAS as 19323. Halling (1986) listed the type of P. floridana: “Marion Co., Juniper Springs, 13 Oct 1940, Murrill, F-19323” and for C. psilocybe: “Juniper Springs, Oct, Murrill (Holotype: FH!; isotype: FLAS)” based on the protologues for both species together with data attached to specimens. In both descriptions of P. floridana and C. psilocybe it is noted that the fungus grows on an oak log in October, although Singer did not mention the year.

Studied Material. UNITED STATES, Florida: Marion Co., Juniper Springs, Oct. 13, 1940, Murrill sn. (FLAS-19323, HOLOTYPE of Psilocybe floridana and ISOTYPE of Callistosporium psilocybe; FH, HOLOTYPE of C. psilocybe; MICH-12136, ISOTYPE of Psilocybe floridana); Gainesville, Jun. 8, 1943, Singer F-2651 (FH, as C. psilocybe).

Additional Material Examined. North Carolina: Curtis Herb., 1856, Sprague 962 (FH- 5728, HOLOTYPE of Agaricus luteoolivaceum).

Acknowledgments

We are grateful to Dr. Roy Halling (New York Botanic Garden) and Dr. Timothy J. Baroni (State University of New York) for their critical review. Dr. Dennis E. Desjardin (San Francisco State University) and Dr. Laura Guzman-Davalos (Universidad de Guadalajara, Mexico) gave useful comments to a preliminary version. Also we express our thanks to Curators of F. FH, FLAS and MICH for loaned important specimens and to Virginia Ramirez-Cruz for her help in some microscopic observations. Guzman acknowledgments the support of his institution to his research. We also thank Lorelei Norvell and Shaun Pennycook their editorial support.

Literature cited

Bas C. 1969. Morphology and subdivision of Amanita and a monograph of its section Lepidella. Persoonia 5: 285-579,

Dennis RWG. 1951. Agaricaceae of Trinidad and Venezuela. Leucosporae I. Trans. _ Brit. Mycol. Soc. 34: 411-480.

Guzman G. 1983. The Genus Psilocybe. Beih. Nova Hedwigia 74, Cramer, Vaduz.

Halling RE. 1986. An Annotated Index to Species and Infraspecific Taxa of Agaricales and Boletales Described by William A. Murrill. Mem. New York Botanical Garden 40, New York.

Hongo T. 1989. Notulae Mycologicae 17. IN: Selected Mycological Papers of Dr. T. Hongo. Shiga Univ., Otsu.

Moser M. 1984. Notes on the genus Callistosporium. In: La Famiglia delle Tricholomataceae. Centro Studi per la Flora Mediterranea, Borgo val di Taro.

Murrill WA. 1945 (‘1944’). New Florida Fungi. Procc. Florida Acad. Sc. 7: 107-127.

Pegler DN. 1983. Agaric flora of the Lesser Antilles. Kew Bull. Addit. Ser. 9, Her Majestys’s Stationery Office, London.

Pulido M, Boekhout T. 1989. Distribution of macrofungi along the Parque Los Nevados transect. IN: Van der Hammen, T., S. Diaz-Piedrahita and V.J. Alvarez (Eds.), La Cordillera Central Colombiana. Transecto Parque Los Nevados II, Cramer, Berlin.

Redhead SA. 1982. The systematics of Callistosporium luteoolivaceum. Sydowia 35: 223-235.

Saccardo PA. 1987. Sylloge Fungorum 5, Patavia (ed. 1944, Brothers, Ann Arbor).

Singer R. 1944. New genera of fungi. Mycologia 36: 358-368.

Singer R. 1946. Type Studies in Agaricales II. Lloydia 9: 114-131.

Singer R. 1970. A monograph of the subtribe Omphalinae. IN: Omphalinae (Clitocybeae, Tricholomataceae, Basidiomycetes). Flora Neotropica 3: 1-84.

MYCOTAXON

Volume 96, pp. 77-81 April-June 2006 Three new species of Stemphylium from China

YONG WANG & XIU-GUO ZHANG *

zhxg@sdau.edu.cn Department of Plant Pathology, Shandong Agricultural University Taian, 271018, China

Abstract—Three new species of Stemphylium from China are described and illustrated: S. pruni, S. plantaginis and S. capsici. The type specimens are deposited in the Herbarium of the Department of Plant Pathology of Shandong Agricultural University (HSAUP).

Key words—hyphomycetes, dematiaceous

Introduction

Until now, about 150 Stemphylium taxa have been reported in the world, but only four new species have previously been described from China (Zhang & Zhang 2003). During the past few years, we have obtained numerous isolates of Stemphylium spp. from a large number of leaf spots derived from different plants. However, most isolates belonged to a few well-known species of Stemphylium such as S. botryosum, S. solani and S. lycopersici. This study presents a survey of Stemphylium species isolated from necrotic leafspots on Prunus persica, Plantago major, and Capsicum annuum in China.

Materials and Methods

Specimens of leaves with leafspots were collected from various zones in China during 2002-2005. All specimens were examined in the laboratory and then deposited in the herbarium of the Department of Plant Pathology, Shandong Agricultural University (HSAUP). Morphological comparisons of sporulation elements of Stemphylium spp. were based on cultures that developed under standardized conditions (Simmons 2001): potato-carrot agar (PCA) in plastic petri dishes, at ambient room temperatures 28°, and examined within 3-4 weeks. Microscope slides of superficial colony growth were prepared in a lactophenol and glycerin solution. The slides were examined and camera lucida drawings were prepared to show the most diagnostic characters. The taxonomic determinations were made from descriptions in the literature.

*Corresponding author

78

Taxonomic Description

Stemphylium pruni Yong Wang bis & X.G. Zhang, sp. nov. FIGURE 1

Ex culturis in agaro ‘potato-carrot’ descripta. Coloniae effusae, pallide brunneae vel medio- brunneae. Mycelium superficiale, hyphae ramosae, septatae, pallide brunneae, laeves, 3.5-4.5 um latae. Conidiophora solitaria, laevia, recta vel curvata, nonramosa vel raro ramosa, pallide brunnea, cylindrica, 4-7-septata, 87-134x3.5-4.5 um, ad apicem in saepissime 1 proliferationis usque 6.0-7.5 um, laevi. Conidia singularia in apice conidiophori et dein proliferationis cujusque successivae oriunda, pallide brunneae, oblonga-ellipsoidea vel oblonga, ad apicem subtruncata, ad basim rotundata vel subtruncata, recta, laevia, 1-3-transverse septata, 0-1 longitudinaliter septata, ad 1-2 distincte constricta, 17-44x 11.5-24 um.

Holotype: on leaves of Prunus persica (L.) Batsch, Arboretum of Guiyang, Guizhou Province, China. Aug. 16. 2003, Y. Wang, HSAUPIII 0159.

Etymology: in reference to the host, Prunus persica

W 77 OG

Fig. 1 Conidia and conidiophores of S. pruni

Colonies on PCA effuse, pale brown to medium brown. Mycelium superficial, hyphae branched, septate, pale brown, smooth, 3.5-4.5 um wide. Conidiophores solitary, unbranched or occasionally branched, straight or slightly curved, light brown, cylindrical, swollen at the apex 6.0-7.5 um diam and distinctly flared, 4-7-septate, 87- 134x3.5-4.5 um, smooth or occasionally echinulate near the proliferation pore. Conidia

79

develop singly through a wide pore at the apex of each conidiophore, pale brown, oblong-elliposoid or oblong, subtruncate at the apex, rounded or subtruncate at the base, straight, smooth-walled, with 1-3 transverse septa and 0-1 longitudinal septum, and always with 1-2 distinct constrictions at the transverse septa, 17-44x11.5-24 um (av. 34.5x18 um), L/W ratio is 1.5-2.4 (av. 1.9).

The conidia of this species are similar to those of S. nabarii (Sarwar & Srinath 1966). However, the conidial size range and L/W ratio (av. 35x18 um, L/W=1.5-2.4) are smaller than those of S. nabarii (av. 44x14 um, L/W=2.8-3.3). Some conidia of the new taxon have a longitudinal septum, while those of S. nabarii have no longitudinal septa. In addition, the conidiophores of the new taxon (87-134 um) are longer than those of S. nabarii (48-77 um).

Stemphylium plantaginis Yong Wang bis & X.G. Zhang, sp. nov. FIGURE 2

Ex culturis in agaro ‘potato-carrot’ descripta. Coloniae effusae, pallide brunneae vel brunneae. Mycelium superficiale ex hyphis ramosis, septatis, pallide brunneis, laevibus, 3.5-4.5 um crassis compositum. Conidiophora singula, recta vel curvata, nonramosa vel raro ramosa, pallide brunneae, cylindrica, 7-9-septata, 48-108x4.0-6.5 um, ad apicem in saepissime 2 proliferationis usque 6.5-12.0 ym, laevibus. Conidia singularia in apice conidiophori et dein proliferationis cujusque successivae oriunda, brunneae, ovoidea, late ellipsoidea vel oblonga-ellipsoidea, ad apicem vel basim subtruncata, recta vel curvata, laevia, 1-2-transverse septata, 0-1 longitudinaliter septata, 26-54x 16-26 um.

W 77 OG

Fig. 2 Conidia and conidiophores of S. plantaginis

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Holotype: on leaves of Plantago major L., Arboretum of Moutain Tai, Shandong Province, China. Oct. 5. 2003, Y. Wang, HSAUP III, 0532.

Etymology: in reference to the host, Plantago major

Colonies on PCA effuse, light brown to brown. Mycelium superficial, composed of branched, septate, pale brown, smooth, hyphae 3.5-4.5 um wide. Conidiophores solitary, simple or occasionally branched, straight or slightly curved, light brown, cylindrical, 7- 9-septate, 48-108x6.5-4.0 um, swollen at the apex 6.5-12.0 um diam and distinctly flared, smooth. Conidia develop singly through a wide pore at the apex of each conidiophore, pale brown, ovoid, broadly-ellipsoid or oblong-ellipsoid, subtruncate at the apex and at the base, straight or slightly curved, smooth-walled, with 1-2 transverse septa, and 0-1 longitudinal septum, 26-54x16-26 um (av. 37.5x21.5 um), L/W ratio is 1.3-2.5 (av. Oy

The conidia of S. plantaginis are similar in shape to those of S. vesicarium (Simmons 1969), but the two species can be separated by conidia septation and wall ornamentation. The conidia in the new taxon generally have 1-2 transverse septa while those of S. vesicarium have 1-5 septa. In addition, the conidia of the new taxon are also smooth- walled with 0-1 longitudinal septum, as opposed to those of S. vesicarium that are conspicuously and densely verrucose at all ages and have 1-2 (-3) longitudinal septa.

Stemphylium capsici Yong Wang bis & X.G. Zhang, sp. nov. FIGURE 3 Ex culturis in agaro ‘potato-carrot’ descripta. Coloniae effusae, medio-brunneae vel brunneae. Mycelium superficiale ex hyphis ramosis, septatis, medio-brunneis, laevibus, 4.0-5.0 um crassis compositum. Conidiophora singula, recta vel curvata, simplicia vel raro ramosa, medio-brunneae vel brunneae, cylindrica, 1-7-septata, 22-58x4.5-8.5 um, ad apicem in saepissime 1 proliferationis usque 7.0-9.0 um, inflato, laevis. Conidia singula in apice conidiophori et in proliferationis cujusque successivae oriunda, medio-brunneae vel brunneae, ovoidea, oblonga-ellipsoidea, ad apice rotundata vel subtruncata, ad basim subtruncata, recta vel curvata, 1-3 transverse septata, 0-2 longitudinaliter vel 2-5-oblique septata, distincte constricta ad septa transversa, 14-33x9.0-10.5 um, laevia.

Holotype: on leaves of Capsicum annuum L., Arboretum of Dali, Yunnan Province, China. Aug. 5. 2002, X.G. Zhang, HSAUPIII 1012

Etymology: in reference to the host, Capsicum annuum

Colonies on PCA spreading flatly, medium brown to brown. Mycelium supercial, composed of branched, septate, medium brown, smooth, hyphae 4.0-5.0 um wide. Conidiophores simple or occasionally branched, straight or curved, medium brown to brown, cylindrical except at the apical proliferations which are swollen or distinctly flared, 1-7-septate, 22-58x4.5-8.5 um, flared apically up to 7.0-9.0 um, smooth. Conidia developing singly through a wide pore at the apex of each conidiophore, medium brown to brown, ovoid or oblong-ellipsoid, rounded or subtruncate at the apex, and subtruncate at the base, straight or slightly curved, smooth-walled, with 1-3 transverse septa, and 0-2 longitudinal or 2-5 oblique septa, 14-33x7.5-12.5 um (av. 20.0x11.5 um ), L/W ratio is 1.3-2.5 (av. 1.9).

The new species differs from S. allii-cepae and S. lactucae (Zhang & Zhang 2003) in the smaller size range of its conidia (14-33x7.5-12.5 um). In addition, the mature conidia

81

U7 0G

Fig. 3 Conidia and conidiophores of S. capsici

of S. allii-cepae and S. lactucae have 1-3 and 1 distinctly consticted transverse septa respectively, while conidia of the new taxon are not distinctly constricted transverse septa. On the other hand, the conidia of the new taxon have 2-5 oblique septa, more than either S. allii-cepae or S. lactucae. The apical conidium cell of the new taxon is also less acute than that of both well-known species.

Acknowledgments

The authors are indebted to W.B. Kendrick, N.R. ONeill and G. Okada for serving as pre-submission reviewers and for their valuable comments and suggestions. This project was supported by the National Science Foundation of China (no.30570006).

Literature Cited

Sarwar M, Srinath KV. 1966. A new species of Stemphylium on Chrysanthemum cinerariaefolium Vis. Mycopath. et Mycologia Applicata 29 (3-4): 320-322.

Simmons EG. 1969. Perfect states of Stemphylium. Mycologia 61:1-26.

Simmons EG. 2001. Perfect states of Stemphylium IV. Harvard Papers in Botany 6 (1): 199-208.

Zhang XG, Zhang TY. 2003. Taxonomic studies of Stemphylium from China. Mycotaxon 85: 247-252.

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Volume 96, pp. 83-107 April-June 2006

Caribbean heterobasidiomycetes: 2. Jamaica

PETER ROBERTS

p.roberts@rbgkew.org.uk Mycology Section, Royal Botanic Gardens Kew, Surrey TW9 3AB, UK

Abstract - Fifty-five species of heterobasidioid fungi belonging to the orders Atractiellales, Auriculariales, Ceratobasidiales, Dacrymycetales, Exidiales, Platygloeales, Septobasidiales, Tremellales, and Tulasnellales are described or reported from Jamaica. The new species Exidia aeruginosa and Achroomyces dennisii are described and the new combination Exidiopsis scutelliformis proposed, with E. fuliginea as a synonym. Heterochaete verruculosa sensu Bodman (et auct. seq.) is considered a mix of H. verruculosa sensu stricto and Exidia japonica. Twenty-six of the 55 species (47%) are newly recorded from Jamaica; 19 of them (34%) are also new records for the Caribbean islands.

Key words - cloud forest, endoparasites, Basidiomycetes, Urediniomycetes

The majority of species recorded in this paper were collected by the author and colleagues in June 1999 as part of the US National Science Foundation, Biotic Surveys and Inventories Program for the Greater Antilles.

The sites surveyed were almost all within the Holywell Park area of the Blue Mountains at an altitude of around 1000 m. The park is in the county of Surrey, mostly in St Andrew parish, but with one trail (Oatley Mountain) extending into Portland parish. Vegetation is mainly secondary cloud forest, with a mix of deciduous trees, Podocarpus, and a few planted Pinus caribaea. The nearby gardens at Cinchona (c. 1000 m) and upland valley woodland at Clydesdale (c. 800 m), both with some planted Eucalyptus, were also visited. Most of the heterobasidiomycete collections are held in the mycological herbarium [K(M)] of the Royal Botanic Gardens, Kew, but some additional collections, forwarded by Dr Karen Nakasone, are held at the Center for Forest Mycology Research (CFMR), Wisconsin, USA. Duplicate specimens are deposited in the herbarium of the Institute of Jamaica (IJ).

Previous collections from Jamaica held at Kew are also recorded. These include specimens of Septobasidium species from the Blue Mountains collected in 1926 by J.N. Couch (Professor of Botany at the University of North Carolina), specimens from the American botanist and collector Charles Orcutt (who moved to Jamaica in the 1920s), material sent to Kew by E.B. Martyn (plant pathologist in Jamaica from 1940 - 1950), and specimens collected by R.W.G. Dennis (Royal Botanic Gardens, Kew) in 1949-50. Also included are references to Jamaican heterobasidiomycetes in other publications, notably

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the paper on Jamaican Septobasidiales by Couch (1929), the booklet on neotropical heterobasidiomycetes by Lowy (1971), and the checklist of Caribbean fungi by Minter et al. (2001).

A previous paper on Caribbean heterobasidiomycetes (Roberts 2003b) dealt with species from the Dominican Republic.

Atractiellales

Hobsonia mirabilis (Peck) Linder, Ann. Missouri Bot. Gard. 16: 340 (1929)

SPECIMENS EXAMINED - SuRREy: St Andrew: CASTLETON BOTANICAL GARDENS, on dead palm leaf, 16 Dec. 1949, Dennis 250B, K(M) 85760; HoLywELL PARK (PATH TO CABINS), on dead Hedychium coronarium stems, 10 June 1999, Roberts GA541, K(M) 63247; MorcE’s GAP, on dead H. coronarium stems, 25 Dec. 1949, Dennis 250C, K(M) 85759.

Hobsonia mirabilis produces viscid-gelatinous, hyaline to whitish fruitbodies on a variety of woody or herbaceous substrata. It has long been classed as an anamorphic ascomycete, but recent molecular research (Sikaroodi et al. 2001) has suggested that Hobsonia is related to the genus Helicogloea. The species was illustrated from Venezuela by Roberts (2003a) and recorded from the Dominican Republic by Roberts (2003b). It has not previously been reported from Jamaica.

Auriculariales

Auricularia cornea Ehrenb., in Nees von Esenbeck, Horae Phys. Berol.: 91 (1820) Auricularia polytricha (Mont.) Sacc., in Berlese & Saccardo, Atti. R. Inst. Veneto VI. 3: 722 (1885)

SPECIMENS EXAMINED - SURREY: Portland: Hoc HOLE To CarT FALLS VIEW, on dead trunk, 22 Dec. 1973, B.D. Morley & C. Whitefoord 772 (as A. polytricha), K(M) 61992; HOLYWELL PARK, OATLEY MOUNTAIN TRAIL, on log, 10 June 1999, Roberts GA561, K(M) 63248; Nanny Town, on wood, 22 Dec. 1973, B.D. Morley & C. Whitefoord 716 (as A. polytricha), K(M) 61991; Port ANTONIO (8 MI FROM), WINNIFRED GUEST HOUSE, on log, 16 Mar. 1946, A.M. Barry F54, K(M) 61990; St Andrew: CLYDESDALE, on wood, 16 June 1999, D.J. Lodge (Roberts GA717), K(M) 63295; KINGSTON, on wood, June 1999, comm. Roberts, K(M) 121795; St Thomas: ARNTULLY, Apr. 1928, Orcutt 5595 (as A. auricula), K(M) 85308.

Auricularia cornea is a common and conspicuous pantropical species, distinguished by its smooth hymenium and hirsute, but unzoned pileus. The species was recorded from the Dominican Republic by Roberts (2003b) and has previously been reported from Jamaica (parishes of Manchester, Portland, St Andrew, St Ann, St Thomas, and Trelawny) in Minter et al. (2001).

Auricularia delicata (Fr.) Henn., in Bresadola et al., Bot. Jahrb. Syst. 17: 492 (1893)

SPECIMEN EXAMINED - JAMAICA. Unlocalized, Mr Poore (as Tremella auricula; det. M.J. Berkeley as Exidia fuscosuccinea), K(M) 51755.

Auricularia delicata is a common pantropical species characterized by its distinctly reticulate hymenium. The species was recorded from the Dominican Republic by

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Roberts (2003b) and has previously been reported from Jamaica (Portland, St Andrew, St Catherine, St Elizabeth, St Thomas, and Westmorland) in Minter et al. (2001).

Auricularia fuscosuccinea (Mont.) Henn., Bot. Jahrb. Syst. 17: 19 (1893)

SPECIMENS EXAMINED - JAMAICA. Unlocalized, on tree, 1928, Orcutt 4231 (as A. tenuis), K(M) 44591; SURREY: Portland: CoRN Puss GAP (BETWEEN PORTLAND AND ST THOMAS), on wood, 21 Feb. 1946, C.B. Lewis F44, K(M) 51767.

Auricularia fuscosuccinea is a common tropical species, distinguished by its smooth, purplish brown basidiomes, without a macroscopically hirsute pileus. The species was recorded from the Dominican Republic by Roberts (2003b) and previously reported from Jamaica (Portland, St Andrew, and St Catherine) by Lowy (1971).

Auricularia mesenterica (Dicks.) Pers., Myc. Eur. 1: 97 (1822)

Auricularia mesenterica is a cosmopolitan species, distinguished by its effuso-reflexed basidiomes, with distinctly zoned and hirsute pilei and smooth to reticulate hymenia. The species was reported from Jamaica (Manchester, St Andrew, and Trelawny) by Lowy (1971).

Ceratobasidiales

Ceratobasidium noxium (Donk) P. Roberts, Rhizoctonia-forming fungi: 48 (1999) Koleroga noxia Donk, Fungus 28: 35 (1958) Pellicularia koleroga Cooke, Grevillea 4: 116 (1876), p.p. [nom. conf.]

SPECIMENS EXAMINED - JAMAICA. Unlocalized, on living Citrus maxima leaves, July 1948, Martyn P/P 340, K(M) 52849; MipDLESEX: Manchester: NEAR MANDEVILLE, on living Coffea leaf, Oct. 1892, anon. (as Pellicularia koleroga), K(M) 29392; St Catherine: LINSTEAD, on living Mangifera indica leaves, 27 Sep. 1967, EF Thompson CB 1120, K(M) 32933.

Ceratobasidium noxium is a widespread tropical species, originally described from India, but with few recent records. It forms web-blights on living leaves of trees and shrubs, causing the disease variously called ‘black rot; ‘koleroga, or ‘candelillo’ in coffee. The species was previously reported from Jamaica in Minter et al. (2001) as Pellicularia koleroga, a nomen confusum. Ceratobasidium noxium was redescribed and illustrated in Roberts (1999).

Thanatephorus cucumeris (A.B. Frank) Donk, Reinwardtia 3: 376 (1956) [anamorph] Rhizoctonia solani J.G. Kihn, in Die Krankheiten der Kulturgewachse: 224 (1858)

Thanatephorus cucumeris, mostly found in its anamorphic (mycelial) state, is a cosmopolitan plant pathogen on an extremely wide range of hosts. It was reported in Minter et al. (2001) from the parishes of St Andrew, St Catherine, and St Thomas in Jamaica, and was previously recorded from the Dominican Republic by Roberts (2003b). The species was redescribed and illustrated, with synonyms listed, in Roberts (1999).

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Dacrymycetales

Calocera cornea (Batsch) Fr., Stirp. Agri. Fems. 5: 67 (1827)

Originally described from Germany, Calocera cornea is a cosmopolitan species distinguished by its gregarious, yellowish, subulate-cylindrical, gelatinous basidiomes up to 10 mm high, its unclamped hyphae, and its comparatively small, single-septate basidiospores. It was reported by Lowy (1971) from St Andrew and St Thomas parishes.

Cerinomyces crustulinus (Bourdot & Galzin) G.W. Martin, Mycologia 41: 85 (1949)

Fig. 1 Basidiomes discoid at first, c. 1 mm across, then coalescing and becoming smooth, effused, gelatinous, 20 mm or more across, translucent tawny when fresh, drying similar. The whitish, fimbriate subiculum is visible at margins.

Hyphae hyaline, 4 - 6 um wide in context, walls thickly gelatinized in KOH, with clamp- connexions; hyphidia conspicuous, branched; basidia dacrymycetoid, c. 30 um long; basidiospores cylindrical (Q = 3.0 - 3.7), 14 - 15.5 x 3.5 - 5 um, allantoid, aseptate.

SPECIMEN EXAMINED - SuRREyY: St Andrew: HOLYWELL PARK, FAIRY GLADE TRAIL, on wood, 13 June 1999, Roberts GA649, K(M) 116646.

Following McNabb (1964) and subsequent authors, this effused species accords quite well with Cerinomyces crustulinus as originally described from France and recorded from North America. The presence of branched hyphidia is a distinguishing feature of the species. The Jamaican collection differs, if at all, from McNabb’s description in having larger spores (McNabb gave 9 - 11 (- 12) x 3 - 4 um as the spore measurement) and context hyphae with gelatinized walls. Cerinomyces crustulinus has not previously been recorded from the Caribbean.

Dacrymyces capitatus Schwein., Trans. Am. Phil. Soc. II, 4: 186 (1832)

SPECIMEN EXAMINED - SuRREY: St Andrew: HOLYWELL PARK, WATERFALL TRAIL, on wood, 8 June 1999, K. Nakasone (Roberts GA521), K(M) 98265.

The Jamaican collection falls within a rather difficult group of unclamped Dacrymyces species with 1-3-septate basidiospores. It is distinguished mainly by its distinctly discoid basidiomes attached to the substratum at a point. The species was originally described from North America and is cosmopolitan, though not previously recorded from the Caribbean.

Dacrymyces chrysospermus Berk. & M.A. Curtis, in Berkeley, Grevillea 2: 20 (1873)

SPECIMEN EXAMINED - SuRREyY: St Andrew: CLYDESDALE, on decorticated log, 16 June 1999, Roberts GA716, K(M) 98264.

Dacrymyces chrysospermus is distinguished by its coalescing, cerebriform basidiomes, its unclamped hyphae, and its 7-septate basidiospores. The species was originally described from North America and appears to be cosmopolitan. It has not, however, previously been reported from the Caribbean.

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Fig. 1. Cerinomyces crustulinus (Jamaica, K(M) 116646). Mature and immature basidia with basal clamp-connexions; aseptate basidiospores; branched hyphidia; thick-walled context hyphae.

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Dacrymyces falcatus Brasf., Lloydia 1: 154 (1938)

Originally described from Panama, the species is distinguished by its clampless hyphae and basidiospores with 15 or more septa. It was reported by Lowy (1971) for Portland parish.

Dacrymyces minor Peck, Rep. New York Mus. Nat. Hist. 30: 49 (1878)

SPECIMEN EXAMINED - SuRREY: St Andrew: HOLYWELL PARK, WATERFALL TRAIL, on fallen decorticated stick, 14 June 1999, Roberts GA668, K(M) 98261.

The collection comprises a swarm of small, discoid, dull orange basidiomes having unclamped hyphae and 1 3-septate basidiospores, the septa slightly thickened. As such, it seems best referred to Dacrymyces minor, originally described from North America and reported from the Dominican Republic by Roberts (2003b). The species has not previously been recorded from Jamaica.

Dacryopinax elegans (Berk. & M.A. Curtis) G.W. Martin, Lloydia 11: 116 (1948)

Dacryopinax elegans is a spathulate species with 3-septate basidiospores, originally described from the southern United States. It was reported by Lowy (1971) for Clarendon parish.

Dacryopinax spathularia (Schwein.) G.W. Martin, Lloydia 11: 116 (1948) SPECIMENS EXAMINED - MipDLESEx: St Mary: ORANGE RIVER EXPERIMENTAL STATION, Jan. 1948, Martyn P/P 290, K(M) 52664; SuRREyY: St Andrew: CLYDESDALE, on fallen wood, 16 June 1999, D.J. Lodge (Roberts GA718), K(M) 114754; St Thomas: BaTH, on wooden gate, Mar. 1946, A.M. Barry F36, K(M) 52663.

This is a ubiquitous warm temperate and tropical species, producing conspicuous, spathulate basidiomes. It was originally described from North America and has previously been reported from the parishes of Portland, St Andrew, and St Thomas in Minter et al. (2001). It was reported from the Dominican Republic in Roberts (2003b).

Guepiniopsis buccina (Pers.) L.L. Kenn., Mycologia 50: 888 (1958)

SPECIMENS EXAMINED - SuRREY: St Andrew: Morce’s Gap, on wood, 10 June 1888, W. Fawcett (as Guepinia pezizaeformis), K(M) 8698; St HELEN GaP TO Monkey HILt, on wood, 1 Sep. 1957, A.L. Welden 878, K(M) 8699.

Guepiniopsis buccina is a cosmopolitan species producing trumpet-like basidiomes (cup-shaped with a long stipe). The collections cited above were previously reported by McNabb (1965).

Exidiales

Basidiodendron cinereum (Bres.) Luck-Allen, Can. J. Bot. 41: 1043 (1963)

SPECIMENS EXAMINED - SuRREY: St Andrew: HOLYWELL PARK, WaG WATER RIVER TRAIL, on dead Hedychium coronarium stem, 11 June 1999, Roberts GA600, K(M) 63274; same location and date, on Pinus caribaea stick, Roberts GA610, K(M) 114690;

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HOLYWELL PaRK, WATERFALL TRAIL, on fallen wood, 14 June 1999, Roberts GA675, K(M) 63320.

Basidiodendron cinereum is a cosmopolitan species or species complex, originally described from Italy. Like all Basidiodendron species, the basidiomes are normally thin, effused, greyish and farinaceous-ceraceous, often drying ochraceous and finely reticulate. The specimen K(M) 63274 is typical, having a thin farinaceous basidiome with abundant gloeocystidia under the microscope and basidiospores measuring 9.5 - 11 x 5.5 - 7 um. The specimens K(M) 63274 and K(M) 114690 are microscopically similar, but have thicker, ceraceous basidiomes, possibly representing more mature specimens.

An additional collection (Surrey: St Andrew: HoLYwELL PARK, WATERFALL TRAIL, on twig, 14 June 1999, Roberts GA689, K(M) 116886) has a thin, pale greyish, farinaceous basidiome and appears similar to a Basidiodendron species, but lacks gloeocystidia. The basidiospores are close to those of B. cinereum, measuring 11.5 - 13.5 x 6.5 - 7.5 um. The specimen is tentatively referred to B. cinereum, but may well represent a distinct species.

Basidiodendron cinereum has not previously been reported from the Caribbean.

Basidiodendron eyrei (Wakef.) Luck-Allen, Can. J. Bot. 41: 1034 (1963)

SPECIMENS EXAMINED - SurREyY: St Andrew: HOLYWELL Park, FAIRY GLADE TRAIL, on fallen wood, 13 June 1999, Roberts GA652, K(M) 63324; same location, on hardwood twigs, 15 June 1999, Nakasone 150383, CFMR; HOLYWELL PARK, WATERFALL TRAIL, on dead tree fern stems, 8 June 1999, Roberts GA505, K(M) 63398; same location, on fallen wood, 14 June 1999, Roberts GA664, K(M) 63319.

Basidiodendron eyrei is a common, cosmopolitan species, originally described from England. It is distinguished by its comparatively small subglobose to globose basidiospores (with Q frequently <1.0, hence with the apiculus on one of the wider sides) and the formation of conspicuous ‘acropetal basidial stalks’ in mature specimens. Basidiospores are typically 3.5 - 5 um across, but in some of the collections cited above they are slightly larger, up to 5.5 - 6 um across. The species has not previously been reported from Jamaica.

Basidiodendron fulvum (Massee) Ginns, Opera Botanica 61: 54 (1982)

SPECIMENS EXAMINED - SurREy: St Andrew: HOLYWELL PARK, OATLEY MOUNTAIN TRAIL, on fallen wood, 15 June 1999, Roberts GA694, K(M) 63321; CINCHONA, TRAIL TO Morce’s Gap, on decayed hardwood, 12 June 1999, Nakasone 150309, CFMR and K(M) 116728; USA. NEw JERSEY. NEWFIELD, on Magnolia sp., undated, J.B. Ellis 3399, HOLOTYPE, K(M) 28713.

Basidiodendron fulvum, described from the USA (type cited above), is similar to B. eyrei but is distinguished by its smaller basidia and basidiospores. It was recorded from Venezuela in Roberts (2003a), but has not previously been reported from the Caribbean.

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Basidiodendron radians (Rick) P. Roberts, Kew Bull. 56: 170 (2001) Basidiodendron nodosum Luck- Allen, Can. J. Bot. 41: 1045 (1963)

SPECIMENS EXAMINED - BRAZIL. R10 GRANDE DO SUL: SAO LEOPOLDO, on wood, 1930, J. Rick 47, HOLOTYPE, K(M) 56863; CANADA. Ontario: Algoma Dist.: LITTLE WHITE RIVER, on decayed Pinus wood, 14 Sep. 1956, R.E Cain, HOLOTYPE of Basidiodendron nodosum, TRTC 33408; JAMAICA. SURREY: Portland: HoLtyweELL PARK, OATLEY MounraiN TRAIL, on rotten fallen wood, 10 June 1999, Roberts GA574, K(M) 116642; St Andrew: CINCHONA, on rotten Podocarpus wood, 9 June 1999, Roberts GA538, K(M) 113669; HOLYWELL PARK, FAIRY GLADE TRAIL, on decorticated hardwood, 15 June 1999, Nakasone 150380, CFMR.

Basidiodendron radians is similar to B. cinereum (above), but differs in its subglobose to ellipsoid basidiospores (Q = 1.1 - 1.4). The species was originally described from Brazil (the synonym B. nodosum from Canada) but appears to be cosmopolitan. Type specimens have been examined and are cited above. It was redescribed and illustrated from the Balearic Islands by Roberts (1996, as B. nodosum). Though apparently common, the species has not previously been reported from the Caribbean.

The specimen K(M) 116642 is extremely thin and unusual in having swollen elements, some with yellowish contents, which are branched and appear intermediate between hyphidia and gloeocystidia.

Basidiodendron spinosum (L.S. Olive) Wojewoda, Mala Flora Grzybow II: 91 (1981)

SPECIMEN EXAMINED - SurREy: St Andrew: HOLYWELL PARK, WATERFALL TRAIL, on fallen wood, 8 June 1999, Roberts GA513, K(M) 63259.

Basidiodendron spinosum is a cosmopolitan species, recognized microscopically by its distinctly warted basidiospores. It was originally described from Tahiti and has been redescribed and illustrated from Costa Rica by Kisimova-Horovitz et al. (1997) and noted from Venezuela by Roberts (2003a). It has not previously been reported from the Caribbean.

Ductifera sucina (Moller) K. Wells, Mycologia 50: 413 (1958)

Originally described from Brazil, Ductifera sucina was redescribed and reported from Jamaica by Wells (1957) and Lowy (1971), based on a collection from Morce’s Gap (St Andrew). The species forms pallid, pustular to cerebriform basidiomes with gloeocystidia.

Eichleriella alliciens (Berk. & Cooke) Burt, Ann. Missouri Bot. Gard. 2: 746 (1915) Exidiopsis alliciens (Berk. & Cooke) K. Wells, Mycologia 53: 354 (1962 [‘1961’])

Originally described from Brazil, Eichleriella alliciens was reported from Jamaica by Wells (1961, as Exidiopsis alliciens). The species forms ochraceous, effused, stereoid basidiomes and appears to be cosmopolitan.

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Eichleriella leveilleanum (Berk. & M.A. Curtis) Burt, Ann. Missouri Bot. Gard. 2: 744 (1915)

Originally described from the United States, Eichleriella leveilleanum was reported from Jamaica by Wells (1961). The species forms reddish, effused, stereoid basidiomes and appears to be restricted to the Americas.

Exidia aeruginosa P. Roberts sp. nov. Fig. 2 Basidiomata gelatinosa, pustulata vel lobata, stipitata, usque ad 10 mm lata, areolata, non setulosa, grisea vel brunnea, subter aeruginosa; hyphae 1 - 2.5 um latae, fibulatae; hyphidia ramosa; basidia tremelloidea, quadricellularia, oblongea, 23 - 25 x 12 - 13 um; basidiosporae cylindraceae, suballantoideae, 16 - 22 x 7.5 - 9.5 um. Holotype: JAMAICA. Surrey: St Andrew: HOLYWELL PARK, WATERFALL TRAIL, on dead attached deciduous twigs, 14 June 1999, Roberts GA670, K(M) 116842.

Basidiomes small, gelatinous, pustular to lobate, stipitate, up to 10 mm across, strongly areolate and appearing (overall) dull grey to brown in the field. Under lens, areolate to reticulate with distinct, rather roughened ridges, hymenial surface between ridges smooth (without hyphal pegs), brown to rusty brown at surface, bluish green below. When dried, pustular, dark greyish brown.

Hymenium comprising a surface layer of basidia and hyphidia in a brown gelatinous matrix, arising from a loosely gelatinized (not agglutinated), bluish-green context; hyphae hyaline, narrow, 1 - 2.5 um wide, highly branched and irregular, thin-walled, with conspicuous clamp-connexions; hyphidia finely branched; cystidia absent; basidia tremelloid, 4-celled, oblong (Q = 1.7 - 1.9), 23 - 25 x 12 - 13 um, immature basidia narrow and often fusiform; sterigmata elongated and sinuous; basidiospores (not clearly seen) apparently cylindrical (Q = 2.0 - 2.5), weakly allantoid, 16 -