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Studies in neotropical polypores 15: new and interesting species from Guyana

     Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061

Terry W. Henkel

     Department of Biological Sciences, Humboldt State University, Arcata, California 95521

Leif Ryvarden

     Department of Botany, University of Oslo, P.O. Box 1045, Blindern, N-0316 Oslo, Norway

	ABSTRACT

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY A PRELIMINARY CHECKLIST OF... LITERATURE CITED

 

During fieldwork in Guyana several unusual and distinctive taxa of polypores were collected, three of which are described here as new. The first of these, Amauroderma coltricioides is the first species known in the Ganodermataceae with smooth basidiospores. Coltricia verrucata and Coltriciella navispora also are described as new, and a key to the neotropical species of Coltricia is provided. Finally, a checklist of 73 poroid fungi from Guyana is given, of which 29 are new records for the country.

Key words: Ganodermataceae, Guayana Highlands, Hymenochaetaceae, neotropics, Polyporaceae

	INTRODUCTION

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY A PRELIMINARY CHECKLIST OF... LITERATURE CITED

 
Wakefield (1934) reported 55 species of poroid, aphyllophoralean taxa, including many that now are recognized synonyms, found in now-former British Guiana. By including additional scattered information from previous publications, Wakefield's account can be looked on as a preliminary checklist of the polypores of the country. Although Dennis (1970) also provided some information on polypores in Guyana, this was taken directly from Wakefields publication, and no new records for the country were provided. As far as we know, no other accounts have been published on the polypores of Guyana since Wakefield's time, and this mycota remains poorly characterized.

This study reports on polypores collected from an area of extensive, primary tropical forest in the Pakaraima Mountains of west-central Guyana near the common border with Venezuela and Brazil (general area: 5° 16' N, 59° 54' W). The Pakaraima Mountains form the eastern extension of the Guayana Highlands, a region of unique floristic composition (Berry et al 1995). The collection area reported here, in the upper Potaro River drainage system, is characterized by unusual pockets of forest heavily dominated by the ectomycorrhizal arborescent legume Dicymbe corymbosa Spruce ex Benth. (Caesalpiniaceae), sharply delimited from surrounding mixed rain forest (Henkel 2001). Recent efforts to document the macrofungi of these Dicymbe forests have revealed a rich mycota including many new and unusual taxa (Henkel 1999, Henkel et al 2000, Miller et al 2001, 2002, Matheny et al 2003). This study is the first to focus on the Aphyllophoralean component of this ecosystem. In this paper we: (i) describe three new species of polypores from the Pakaraima Mountains of Guyana; (ii) provide a key to the neotropical species of Coltricia S.F. Gray; and (iii) give a preliminary checklist of the polypores of Guyana, based on a taxonomic update of Wakefield's (1934) list and on recent collections made in the Pakaraima mountains that include 29 new records for the country.

	MATERIALS AND METHODS

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY A PRELIMINARY CHECKLIST OF... LITERATURE CITED

 
All collecting was done during the rainy season (May–July) in the Pakaraima Mountains, upper Potaro River, 20 km east of Mount Ayanganna, near the confluence of the Potaro River and Alukyadongbaru Creek. Coordinates of collecting sites were obtained with a hand-held GPS device wherever the canopy cover permitted. Herbaria designations are according to Holmgren et al (1990).

Basidiomata were examined in the field for their fresh characteristics. Color characteristics are described subjectively in general color terms and assigned alphanumeric color designations from Kornerup and Wanscher (1981). Basidiomata were dried slowly over charcoal and subsequently placed in containers with silica gel to prevent spoilage in the humid conditions.

All measurements of microscopic structures were made in mounts of 3% KOH; Melzer's reagent was used to determine amyloid reactions. Mounts were examined with a Zeiss phase contrast microscope, and measurements were made on a Compaq Deskpro computer screen via a Sony video camera attached to the microscope using Lucia Image Archive version 4.51 (Laboratory Imaging, Prague, Czech Republic). A minimum of 20 basidiospores was measured for each specimen and the two largest and two smallest measurements were eliminated before the average values were calculated. Scanning electron micrographs were produced with a Joel model JSM 6400 scanning electron microscope and images were developed electronically with analySIS version 3.1 (Soft Imaging System, Münster, Germany).

	TAXONOMY

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY A PRELIMINARY CHECKLIST OF... LITERATURE CITED

 
GANODERMATACEAE

Amauroderma coltricioides Henkel, Aime et Ryvarden, sp. nov. Figs. 1–3, 9–10

View larger version (47K): [in this window] [in a new window]   FIGS. 1–3. Amauroderma coltricioides (ISOTYPE). 1. Arboriform and unbranched skeletal hyphae. 2. Basidiospores. 3. Generative hypha. FIGS. 4–6. Coltricia verrucata (ISOTYPE). 4. Basidiome. 5. Basidiospores. 6. Ornamented generative hyphae from the context. FIGS. 7–8. Coltriciella navispora (ISOTYPE). 7. Ornamented basidiospores. 8. Generative hypha

View larger version (153K): [in this window] [in a new window]   FIGS. 9–12. Scanning electron micrographs. 9, 10. Amauroderma coltricioides (ISOTYPE), basidiospores. 11. Coltricia verrucata (ISOTYPE), ornamented generative hyphae from context. 12. Coltriciella navispora (ISOTYPE), basidiospores.

Fructificatio stipitate, pileus brunneus, 2–10 cm, pori 7–8 per mm, systema hyphale dimiticum, hyphae generatoriae fibulatae, hyphae skeletoriae arboriforme, basidiosporae laeves globosae, 6.5–7 µm.

Basidiomata annual, laterally to centrally stipitate. Pileus circular to semicircular or flabellate in young specimens, flat to infundibuliform, up to 10 cm in diam, up to 3 mm thick in center; margin lobed, incised to entire, sharp and mostly deflexed when dry; pileus surface finely appressed velutinate, shiny with numerous distinct to indistinct concentric zones, brown to deep reddish brown 6D–F8, the latter especially in mature specimens. Pore surface reddish brown with olivaceous overtones 4B–D5, bruising black where injured in immature basidiomata; pores round to angular, 7–8 per mm, barely visible to the naked eye; tube layer up to 2 mm thick, concolorous with the pore surface. Context up to 1 mm thick, fibrous and rusty to reddish brown 7E8, exuding a watery reddish-black latex when injured in immature basidiomata. Stipe cylindrical, finely velutinate, rusty to deep reddish brown 5D8, up to 5 cm long, 4–8 mm in diam, expanded toward the pore layer and then up to 1.5 cm in diam in the sterile part, dense and homogenous in section. Odor and taste none.

Hyphal system dimitic; generative hyphae with clamps, 3–5 µm in diameter, difficult to observe in dried specimens, hyaline and richly branched; arboriform skeletal hyphae present and dominating in the basidiome, yellow to brown, thick-walled to almost solid, apically moderately branched, up to 10 µm wide in the basal stems, and these up to 300 µm long without branching, thus fragments might be mistaken for true skeletal hyphae. Cystidia or other sterile hymenial elements absent. Basidiospores globose, smooth, thick-walled, pale rusty brown, nonamyloid in Melzers reagent, 6.5–7 µm in diameter.

Specimens examined. GUYANA. PAKARAIMA MOUNTAINS: Upper Potaro River, 20 km east of Mount Ayanganna, near confluence of Potaro River and Alukyadongbaru Creek, near Dicymbe plot 1 on "Benny's Ridge", N 5° 16.580', W 59° 54.966', approximately 850 m elev, near base of dead Tachigali rusbyi Harms, 24 Jul 2000, Henkel 7873 (HOLOTYPE BRG, ISOTYPE O); near Dicymbe plot 1, on buttresses of T. rusbyi, 12 June 2000, Aime 1211 (VPI,O); Dicymbe plot 1, gregarious at base of T. rusbyi, 19 Jun 2002, Aime 2081 (VPI).

Etymology. Coltricioides, Latin, resembling Coltricia.

Habit, habitat and distribution. Gregarious to occasionally caespitose on the ground around standing dead trees of Tachigali rusbyi (Caesalpiniaceae) or fruiting occasionally from lower buttresses of these trees. Currently observed only around T. rusbyi trees in forests otherwise dominated by Dicymbe corymbosa. Known only from the type collection locale.

Comments. This taxon presents a perplexing mixture of characters. It macroscopically resembles a Coltricia species but is easily distinguished from that genus microscopically by its hyphal system, which is monomitic in Coltricia. Amauroderma includes species with brown stipitate basidiomata and a dimitic hyphal system composed of clamped generative hyphae and arboriform skeletal hyphae, as found in A. coltricioides. However, all previously described species in Amauroderma, and in the Ganodermataceae as a whole, possess double-walled basidiospores with an ornamented endosporium, while the present species has smooth spores (Figs. 2, 9–10). Because of these strikingly different basidiospores the authors originally considered erecting a new genus within the Ganodermataceae to accommodate this species. However, subsequent phylogenetic analysis of internal transcribed spacer region DNA sequences (unpubl.) show this taxon to be a member of Amauroderma, where we now place it.

Amauroderma coltricioides tends to fruit in large troops, which can consist of up to 200 or more individual fruiting bodies in various stages of development around and on the lower buttresses of Tachigali rusbyi. Immature basidiomata with undeveloped pores exude a watery rusty-black latex that is absent in mature specimens. One collection (Aime 1211) consists entirely of immature basidiomata. While no intact, measurable basidia were observed in these collections, basidia were present, and abundant basidiospores, some still attached to sterigmata (Fig. 10), were evident within the pores.

HYMENOCHAETACEAE

Coltricia verrucata Aime, Henkel et Ryvarden, sp. nov. Figs. 4–6, 11

Fructificatio stipitata, pileus brunneus, pori angulatis 4–5 per mm, brunneus, systema hyphale monomiticum, hyphae generatoriae afibulatae, verrucosae, basidiosporae ellipsoideae, laeves, pallide flavae, 7–9 x 5–6 µm.

Basidiomata annual, centrally stipitate. Pileus circular, infundibuliform, 7–15 mm in diameter, deep reddish brown 8F6, striate with bristle-like bundles of hyphae, erect in the center, more flattened toward the margin, each bundle up to 6 mm long, projecting beyond margin edge; margin thin, partly incised, deflexed when dry. Pore surface reddish brown 5F5; pores thin-walled, angular, 2–5 per mm. Context reddish brown, fibrous, up to 4 mm thick. Stipe 8–30 x 0.5–2 mm, broader at base, dark reddish- to blackish-brown and velutinous. Odor and taste indistinct.

Hyphal system monomitic; generative hyphae with simple septa, thin-walled to thick-walled, strongly verrucose and golden to rusty brown throughout, (5) 8–13 (16) µm in diam. Basidia barrel-shaped 18–20 x 7–10 µm with 4 sterigmata. Cystidia, setae or other sterile hymenial elements absent. Basidiospores ellipsoid, smooth, thick-walled, golden yellow and negative in Melzers reagent, 7–9 x 5–6 µm in diameter.

Specimens examined. GUYANA. PAKARAIMA MOUNTAINS: Upper Potaro River, 20 km east of Mount Ayanganna, near confluence of Potaro River and Alukyadongbaru Creek, near Ayanganna airstrip No. 1 in Dicymbe corymbosa-dominated forest, general area N 5° 16', W 59° 54', approximately 650 m elev. Gregarious over base of decorticated tree stump, 17 May 2000, Aime 962 (HOLOTYPE BRG, ISOTYPE O); near base camp, on humic mat on D. corymbosa trunk, 29 Jun 2002 Aime 2160 (VPI).

Etymology. Verrucata, Latin, referring to the tuberculate ornamentation of the generative hyphae.

Habit, habitat and distribution. Scattered to gregarious in accumulated humicolous layer covering tree trunks and stumps in forests dominated by Dicymbe corymbosa. Known only from the type locality but possibly overlooked in other places due to its size and cryptic coloring.

Comments. This species is made conspicuous by its strongly striate pileus composed of bundles of short, erect bristles in a central depression that become appressed like combed hairs toward the margin (Fig. 4). The tuberculate ornamentation of the hyphae (Figs. 6, 11) is highly characteristic as well, being present on almost every hypha except those close to the basidia. A key to the neotropical species of Coltricia is given below; a full description of each taxon can be found in the reference provided after its name.

KEY TO THE NEOTROPICAL SPECIES OF COLTRICIA S.F. GRAY

1. Setae or setal hyphae present  . . . . . C. hamata (Romell) Ryvarden (see Ryvarden and Johansen 1980:109)

1. Setae or setal hyphae absent . . . . . 2

     2. Basidiospores longer than 6 µm . . . . . 3

     2. Basidiospores shorter than 6 µm . . . . . 6

3. Basidiospores cylindrical, basidiomata on burnt wood or fire places . . . . . C. focicola (Berk. & M.A. Curtis) Murrill (see Gilbertson and Ryvarden 1986:204)

3. Basidiospores ellipsoid, substrate different . . . . . 4

     4. Basidiospores 9–14 µm long, pores 1–3 mm wide . . . . . C. montagnei (Fr.) Murrill (see Gilbertson and Ryvarden 1986:207)

     4. Basidiospores 6–10 µm long, 2–5 pores per mm . . . . . 5

5. Basidiomata usually 3–10 cm in diameter, upper surface shiny and multizonate, hyphae smooth . . . . . C. cinnamomea (Pers.) Murrill (see Gilbertson and Ryvarden 1986:203)

5. Basidiomata < 2 cm wide, pileus striate, hyphae strongly verrucate  . . . . . C. verrucata Aime et al

     6. Basidiospores globose, margin ciliate  . . . . . C. barbata Ryvarden & Meijer (see Ryvarden and Meijer 2002:46)

     6. Basidiospores ellipsoid, no cilia along the margin . . . . . 7

7. Basidiospores 2.5–3 µm wide, pores 6–8 mm, context duplex  . . . . . C. fonsecoensis Cooke & Bonar (see Cooke and Bonar 1961)

7. Basidiospores 3–3.5 µm wide, pores 2–6 per mm, context homogenous . . . . . 8

     8. Pores 4–6 per mm, on the ground in Argentina and Paraguay . . . . . C. stuckertiana (Speg.) Rajchenb. & Wright (see Rajchenberg and Wright 1998:119)

     8. Pores 2–4 per mm, on burnt ground in Venezuela  . . . . . C. pyrophila (Wakef.) Ryvarden (see Ryvarden & Johansen 1980:111)

Coltriciella navispora Henkel, Aime et Ryvarden, sp. nov. Figs. 7–8, 12.

Coltriciella oblectabilis (Lloyd) Kotlaba et al aemulans, differt sporae navisporae.

Basidiome annual, laterally stipitate. Pileus slightly infundibuliform, 0.5–2 cm in diameter, up to 2 mm thick in center; margin thin, deflexed when dry; pileus surface dull, finely appressed velutinate, rusty-brown 6D8 to 7F8, weakly concentrically zonate, no cuticle present. Pore surface concolorous with pileus; pores angular, shallow, 1–2 per mm; tubes concolorous, up to 1 mm deep. Context concolorous with pileus, homogeneous, about 1 mm thick. Stipe up to 1 cm long, 1–2 mm in diameter, velutinate, more distinctly so than the pileus, concolorous with pileus. Odor and taste indistinct.

Hyphal system monomitic, with simple septate generative hyphae, golden-yellow to light rusty-brown, moderately branched, 5–8 µm wide on the pileus and with distinct wide lumen, 3–6 µm in the trama and with slightly thicker walls, moderately branched. Basidia broadly clavate, 15–20 x 8–12 µm with four sterigmata. Cystidia or other sterile hymenial elements absent. Basidiospores navicular, golden-yellowish to light rusty-brown, finely verrucose and somewhat thick-walled, negative in Melzer's reagent, 10–12 x 4–5 µm.

Specimens examined. GUYANA. PAKARAIMA MOUNTAINS: Upper Potaro River, 20 km east of Mount Ayanganna, near confluence of Potaro River and Alukyadongbaru Creek, Dicymbe plot 3, general area N 5° 16', W 59° 54', 13 Jul 2000, Henkel 7576 (HOLOTYPE BRG, ISOTYPE O); plots 2, N 5° 16.463', W 59° 54.731', approximately 845 m elev, 2 Jun 2000, Aime 1112 (VPI); near Dicymbe plot 1 on "Benny's ridge" in Dicymbe corymbosa-dominated forest, N 5° 16.580', W 59° 54.966', approximately 850 m elev. 18 Jun 2000, Aime 1268 (VPI); near Dicymbe plot 2, fruiting from humus layer between coppices of D. corymbosa tree, 29 Jun 2002, Aime 2156 (VPI).

Etymology. Navispora, Latin, referring to the navicular basidiospores.

Habit, habitat, and distribution. Scattered in sheltered areas in the hollows of well-decorticated trees or on the underside of fallen trunks in forests dominated by Dicymbe corymbosa. Known only from the type locality but easily overlooked due to its size and cryptic fruiting habit.

Comments. Coltriciella is a small genus separated from the more common and widespread Coltricia by having verrucose spores. It includes resupinate, pendant and stipitate species. Of the latter type there are only two other described species, C. oblectablis and the east Asian species C. pusilla (Imazeki & Yasuda) E. Corner. Coltriciella navispora macroscopically resembles C. oblectablis. However, the two species are readily distinguished by their basidiospores, those of the latter being oblong ellipsoid and shorter (7–10 x 4–5 µm) (see Gilbertson and Ryvarden 1986:213 for a description of C. oblectablis). Likewise, the basidiospores of C. pusilla are ellipsoid and smaller (7.5–8.5 x 4.5–5 µm) than those of C. navispora (see Nunez & Ryvarden 2000:57 for a description of C. pusilla). In fact, no other species of Coltriciella is known to possess spores greater than 10 µm in length, making C. navispora a distinct species within the genus.

	A PRELIMINARY CHECKLIST OF THE POLYPOROID SPECIES OF GUYANA

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In the following list, those species that are new records for the country are marked with an asterix (*). New taxa are indicated in bold type.

GANODERMATACEAE

*Amauroderma boleticeum (Pat. & Gaillard) Torrend

Amauroderma camerarium (Berk.) Furtado

*Amauroderma coltricioides Henkel et al

*Amauroderma corneri Gulaid & Ryvarden

Amauroderma exile (Berk.) Torrend

*Amauroderma gusmanianum Torrend

Amauroderma partitum (Berk.) Wakef.

*Amauroderma praetervisum (Pat.) Torrend

Amauroderma schomburgkii (Mont. & Berk.) Torrend

*Amauroderma trichodermatum Furtado

Ganoderma australe (Fr.) Pat.

Ganoderma nitidum Murrill

Ganoderma oerstedii (Fr.) Torrend

Ganoderma opacum (Berk. & Mont.) Pat.

*Ganoderma resinaceum Boud.

*Ganoderma stipitatum Murrill

Haddowia longipes (Lév.) Steyaert

HYMENOCHAETACEAE

*Coltricia montagnei (Fr.) Murrill

*Coltricia verrucata Aime et al

*Coltriciella navispora Henkel et al

*Coltriciella oblectabilis (Lloyd) Kotl., Pouzar & Ryvarden

Phellinus gilvus (Schwein.) Pat.

Phellinus roseo-cinereus (Murrill) D. A. Reid

Phellinus senex (Nees & Mont.) Imazeki

Phylloporia spathulata (Hook.) Ryvarden

POLYPORACEAE

*Antrodiella hydrophila (Berk.) Ryvarden

Antrodiella liebmannii (Fr.) Ryvarden

*Ceriporia spissa (Schwein. : Fr.) Rajchenb.

*Coriolopsis rigida (Berk. & Mont.) Murrill

Daedalea sprucei Berk.

Datronia caperata (Berk.) Ryvarden

Dichomitus cavernulosus (Berk.) Masuka & Ryvarden

Earliella scabrosa (Pers.) Gilb. & Ryvarden

Flabellophora obovata (Jungh.) Nunez & Ryvarden

Fomes fasciatus (Swartz : Fr.) Cooke

Fomitella supina (Swartz : Fr.) Murrill

Fomitopsis nivosa (Berk.) Gilb. & Ryvarden

*Gloeophyllum striatum (Fr.) Murrill

Gloeoporus thelephoroides (Hook.) G. Cunn.

Hexagonia hydnoides (Fr.) Fidalgo

Laetiporus sulphureus (Bull. : Fr.) Murrill

Lentinus crinitus (L. : Fr.) Fr.

*Microporellus dealbatus (Berk. & Curt.) Murrill

Nigroporus vinosus (Berk.) Murrill

*Perenniporia inflexibilis (Berk.) Ryvarden

Perenniporia martius (Berk.) Ryvarden

*Polyporus arcularius Fr.

*Polyporus dictyopus Mont.

Polyporus guianensis Mont.

Polyporus leprieurii Mont.

*Polyporus puttemannsii Henn.

Polyporus rhizomorphus Mont.

*Porogramme albocincta (Cooke & Massee) J. Lowe

*Porogramme graphica (Bres.) Pat.

Pycnoporus sanguineus (Fr.) Murrill

*Rigidoporus lineatus (Pers.) Ryvarden

Rigidoporus microporus (Fr.) Overeem

*Rigidoporus ulmarius (Fr.) Imazeki

Rigidoporus vinctus (Berk.) Ryvarden

Tinctoporellus epimiltinus (Berk. & Broome) Ryvarden

*Trametes elegans (Fr.) Fr.

*Trametes lactinea Berk.

Trametes membranacea (Swartz : Fr.) Kreisel

*Trametes menziezii (Berk.) Ryvarden

Trametes modesta (Fr.) Ryvarden

*Trametes pavonia (Hook.) Ryvarden

Trametes villosa (Fr.) Kreisel

Trichaptum byssogenus (Jungh.) Ryvarden

Trichaptum griseo-fuscus (Mont.) Ryvarden & Iturriaga

Trichaptum perrottetii (Lév.) Ryvarden

Trichaptum sector (Fr.) Kreisel

Tyromyces albogilvus (Berk. & M.A. Curtis) Ryvarden & Sjekk.

Wrightoporia tropicalis (Cooke) Ryvarden

	ACKNOWLEDGMENTS

 
This work was made possible by grants from the National Geographic Society's Committee for Research and Exploration and the Smithsonian Institution's Biological Diversity of the Guianas Program to TWH, and an Explorers Club, Washington Group Exploration and Field Research Grant to MCA. Mimi Chin, Leonard Williams, Christopher Andrew, Benny Cheong and Steve Miller provided field assistance in Guyana. We are grateful to Jean-Marc Moncalvo for sharing his database of Ganodermataceae sequences and to Robert Gilbertson and one other reviewer for their valuable comments on this manuscript. This paper is No. 74 in the Smithsonian Institution's Biological Diversity of the Guianas Program publication series.

	FOOTNOTES

 
¹ Corresponding author. Current addresses: USDA-ARS, Rm 319, B011A, Systematic Botany and Mycology Lab, 10300 Baltimore Ave., Beltsville, MD 20705. E-Mail: cathie{at}nt.ars-grin.gov

Accepted for publication February 3, 2003.

	LITERATURE CITED

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY A PRELIMINARY CHECKLIST OF... LITERATURE CITED

 
Berry PE, Holst BK, Yatskievych K, eds 1995 Flora of the Venezuelan Guayana. Vol.1. Introduction. St. Louis: Missouri Botanical Garden Press. 320 p

Cooke WB, Bonar L., 1961 Additional fungi from Galapagos Islands and other Pacific coastal islands collected during the Templeton Crocker expedition, 1932. Occas. Papers Calif Acad Sci 29:1-5

Dennis RWG., 1970 Fungus flora of Venezuela and adjacent countries. London: HMSO. 531 p

Gilbertson RL, Ryvarden L., 1986 North American polypores. Vol. 1. Oslo: Fungiflora. 431 p

Henkel TW., 1999 New taxa and distribution records for Tylopilus from Dicymbe forests of Guyana. Mycologia 91:655-665

———. 2001 Systematics and ecology of ectomycorrhizal fungi associated with Dicymbe in Guyana [PhD Dissertation]. Durham, North Carolina: Duke University. 259 p

———, Aime MC, Miller SL., 2000 Systematics of pleurotoid Russulaceae from Guyana and Japan, with notes on their ectomycorrhizal status. Mycologia 92:1119-1132

Holmgren PK, Holmgren NH, Barnett LC, eds 1990 Index herbariorum part 1. The Herbaria of the World. 8th ed. New York: New York Botanical Gardens. 693 p

Kornerup A, Wanscher JH., 1981 Methuen handbook of colour. 3rd ed. London: Eyre Methuen. 252 p

Matheny PB, Aime MC, Henkel TW., 2003 New species of Inocybe from Dicymbe forests in Guyana, South America. Mycol Res (In press)

Miller OK, Henkel TW, James TY, Miller SL., 2001 Pseudotulostoma, a remarkable new volvate genus in the Elaphomycetaceae from Guyana. Mycol Res 105:1268-1272

Miller SL, Aime MC, Henkel TW., 2002 Russulaceae of the Pakaraima Mountains of Guyana. I. New species of pleurotoid Lactarius. Mycologia 94:545-553[Abstract/Free Full Text]

Nunez M, Ryvarden L., 2000 East Asian polypores. Vol. 1. Synopsis Fung 13:1-168

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

Ryvarden L, Johansen I., 1980 A preliminary polypore flora of East Africa. Oslo: Fungiflora. 636 p

Ryvarden L, Meijer ARA., 2002 Studies in Neotropical polypores 14—new species from the state of Parana, Brazil. Synopsis Fung 15:34-69

Wakefield EM., 1934 XXX—contributions to the flora of tropical America: XXI. Fungi collected in British Guiana, chiefly by the Oxford University Expedition, 1929. Kew Bulletin Misc. Info 1934:238-258

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