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Home , Atheniella, Clitocybula, Hydropus, Megacollybia, Mycena, Mycenaceae

A TAXONOMIC INVESTIGATION OF OF SAO TOME AND PRINCIPE

A thesis submitted to the faculty of $ *■; San Francisco State University ^ & In partial fulfillment of 7j o \% the requirements for ft I CL. the Degree

Master of Science

In

Biology: Ecology, Evolution and Conservation Biology

by

Alexandra Christine Cooper

San Francisco, California

May 2018 Copyright by Alexandra Christine Cooper 2018 CERTIFICATION OF APPROVAL

I certify that I have read A TAXONOMIC INVESTIGATION OF MYCENA OF SAO

TOME AND PRINCIPE by Alexandra Christine Cooper, and that in my opinion this work meets the criteria for approving a thesis submitted in partial fulfillment of the requirement for the degree Master of Science in Biology: Ecology, Evolution, and

Conservation Biology at San Francisco State University.

San Francisco State University

Thomas Parker, Ph.D. Professor of Biology San Francisco State University

Brian Perry, Ph.D. Associate Professor of Biology California State University East Bay A TAXONOMIC INVESTIGATION OF MYCENA OF SAO TOME AND PRINCIPE

Alexandra Christine Cooper San Francisco, California 2018

Knowledge of the diversity of the fungi from the Gulf of Guinea islands is very limited due to the fact that until recently, there have been no biotic surveys of the mycota of the region. In April of2006 and 2008, expeditions were led to document the diversity of plants, amphibians, marine invertebrates and macrofungi on the West African islands of Sao Tome and Principe. This project aims to further document the diversity of macrofungi on the island by focusing on 24 mycenoid fungi collected from the expedition. Nineteen distinct are recognized, of which nine are new to science. The other ten species are new distribution records for the islands. These data suggest that there is a high level of endemism in the saprotrophic macrofungi of ST&P. Molecular sequence data indicate that the currently accepted infrageneric classification of Mycena recognizes many non-monophyletic groups and is in need of revision. ACKNOWLEDGMENTS

I would like to express my gratitude and appreciation to my advisor Dr. Dennis

Desjardin and committee members Dr. Brian Perry and Dr. Thomas Parker. Dr. Desjardin spent many hours sharing his knowledge of Mycena, going through the literature, and working with me to review and improve my project. Also, Dr. Brian Perry for the knowledge he shared with me about DNA sequencing, phylogenetic analyses, and patience he had with me throughout this research. This project would not have been possible without the guidance and direction received by all of them.

Additionally, I would like to thank the following groups that provided financial support for this project: Advancing Science in America (ARCS), Sally Casanova Pre-

Doctoral Scholarship, The Emily M. Hewitt Memorial Scholarship, Mycological Society of San Francisco, and the Sonoma Mycological Society. The support from these groups greatly contributed to the costs associated with this research and made this project finish in a timely manner. TABLE OF CONTENTS

List of Figures...... viii

Introduction...... 1

Methods...... 3

Results...... 6

Key for Identification of Mycena from Sao Tome and Principe ...... 10

Taxonomy...... 13

Mycean Section Calodontes...... 13

Mycean Section Exornatae...... 17

Mycean Section Fragilipedes...... 20

Mycean Section Granuliferae...... 22

Mycean Section Lactipedes...... 24

Mycean Section Longisetae...... 26

Mycean Section Mycena...... 28

Mycean Section Polydelphia...... 31

Mycean Section Sacchariferae...... 33

Mycean Section uncertain...... 41

Clitocybula...... 43

Filoboletus...... 45

Hydropus...... 48

“”...... 53

Excluded Species...... 55

Literature Cited...... 56 LIST OF FIGURES

Figures Page

1. Maximum Likelihood phylogeny of (nLSU)...... 64 2. Maximum Likelihood phylogeny of Mycenoid Fungi (ITS)...... 66 3. Maximum Likelihood phylogeny of Hydropoid Fungi (ITS)...... 68 4. Mycena brunneoviolacea basidiomes...... 69 5. Mycena brunneoviolacea micromorphology...... 70 6. Mycena aff. holoporphyra micromorphology...... 71 7. Mycena aff. discobasis basidiomes...... 72 8. Mycena aff. discobasis micromorphology...... 73 9. Mycena solis basidiomes...... 74 10. Mycena solis micromorphology...... 75 11. Mycena antennae micromorphology...... 76 12. basidiomes...... 77 13. Mycena galopus micromorphology...... 78 14. Mycena breviseta micromorphology...... 79 15. Mycenaphaeonox basidiomes...... 80 16. Mycena phaeonox micromorphology...... 81 17. Mycena longinqua basidiomes...... 82 18. Mycena longinqua micromorphology...... 83 19. Mycena alphitophora basidiomes...... 84 20. Mycena alphitophora micromorphology...... 85 21. Mycena discogena micromorphology...... 86 22. Mycena lasiopus micromorphology...... 87 23. Mycena oboensis micromorphology...... 88 24. Mycena lamprospora basidiomes...... 89 vii 25. Mycena lamprospora micromorphology...... 90 26. intervenosa basidiomes...... 91 27. Clitocybula intervenosa micromorphology...... 92 28. Filoboletuspallescens basidiomes...... 93 29. Filoboletus pallescens micromorphology...... 94 30. globosporus basidiomes...... 95 31. Hydropus globosporus micromorphology...... 96 32. Hydropus murinus basidiomes...... 97 33. Hydropus murinus micromorphology...... 98 34. “Trogia delicata” micromorphology...... 99

viii 1

Introduction

Scientists today know very little about the biodiversity of fungi, with the global estimate of only 3-8% of species being described to date (Hawksworth and Lucking 2017). One reason they are not well documented worldwide is due to their transient nature and difficulties that have traditionally been associated with their identification. Lack of accessible monographs and long-term studies have hindered the ability to estimate fungal diversity and their evolutionary relationships (Perry 2002, Mueller et al. 2007). There is more known about the diversity of macrofungi from North America and Europe compared to lesser studied regions in the world, such as Africa, but even there, our knowledge is incomplete. No accurate estimates exist of the fungal diversity of Africa, South America, south and southeastern Asia, and other understudied regions. Accordingly, both regional and site-based studies of macrofungal diversity are needed to determine their levels of endemism worldwide (Mueller et al. 2007). One of the lesser-documented groups of macrofungi is the basidiomycetous Mycena (Pers.) Roussel. Worldwide in distribution, Mycena is reported from many temperate and tropical regions in the Northern and Southern Hemispheres (Maas Geesteranus 1992a, Perry 2002, Robich 2003). This genus includes saprotrophic, pathogenic, and endophytic species that grow on decaying plant material such as leaves, twigs, cones, and wood. Mycena species are recognized morphologically by a combination of characters. They are small to medium sized, fragile and often inconspicuous. The cap () is conical to campanulate (bell-shaped) and typically pellucid-striate. The gills (lamellae) are usually attached or sometimes free from a central, hollow stalk (). Their are white to pale cream in deposit, globose to ellipsoid or oblong, smooth, and stain blue in Melzer’s reagent (amyloid). Most species form cystidia on their gill edges (cheilocystidia), and may or may not have them on the gill faces (pleurocystidia) and stipe surface (caulocystidia). The anatomy of the pileus surface (pileipellis) and stipe surface (stipitipellis) are variable and taxonomically informative. Species of Mycena are character-rich but often difficult to 2 identify accurately based solely on morphological features, but the use of molecular sequence data, especially the variable internal transcribed spacer (ITS) region, is helpful in delimiting taxa. Sao Tome and Principe (ST&P) are volcanic islands located off the coast of western Africa. These oceanic islands originated 13,000,000 years ago, are not part of the Cameroon line in the Gulf of Guinea, and were never connected to continental Africa by a land bridge (Desjardin & Perry 2009). Principe is geologically the oldest and the most northerly of the chain and much smaller than the geologically younger Sao Tome. The islands are separated from each other and from the West African coastlines of Equatorial Guinea and Gabon by ocean depths of 3000 m (Drewes & Wilkinson 2004). The biodiversity of many groups of organisms on ST&P are well-known and show significant levels of endemism (Desjardin & Perry 2009). However, knowledge of the diversity of fungi from the Gulf of Guinea islands is very limited because until recently there have been no biotic surveys of the mycota of the region (Desjardin & Perry 2009, 2015a, 2015b, 2016, 2017, 2018). An early account of the fungi on ST&P is that of Bresadola and Roumeguere (1890) who documented mostly polypores along with some , clavarioid fungi, gasteromycetes, stereoid fungi, jelly fungi, ascomycetes, and myxomycetes. To date, only three species of Mycena have been reported from ST&P. Bresadola and Roumeguere (1890) reported (Paulet) Quel, from Sao Tome (currently accepted in sect. Mycena). Coutinho (1925) reported 76 species of fungi from the island of Sao Tome, of which two belong to Mycena: (Bull.) Gramberg., a member of sect. Calodontes with lilac-rose pileus and venose- reticulate lamellae; and an illegitimate new species described as Agaricus (Mycena) rufescens Cout. (non Agaricus rufescens J.F. Gmel 1792), currently recognized as a nomen dubium. None of these three taxa were encountered during the course of this project. In April 2006 and 2008, expeditions were led by scientists from the California Academy of Sciences and mycologists from San Francisco State University to document the diversity of plants, amphibians, marine invertebrates and macrofungi on the West 3

African islands of ST&P. Six papers have been published so far documenting the basidiomycetous macrofungi of the islands, based on specimens collected during these expeditions (Desjardin & Perry 2009, 2015a, 2015b, 2016, 2017, 2018). This project adds to our understanding of the diversity of macrofungi from ST&P by focusing on documenting species of mycenoid fungi, primarily belonging to the genus Mycena and allied genera.

Methods

Twenty-four specimens of Mycena and allied genera were collected from ST&P in 2008, dried and deposited in the Harry D. Thiers Herbarium at San Francisco State University (SFSU). Macromorphological data and photographs from fresh material were recorded by Desjardin and Perry at the time of the collection, following the protocols of Largent (1986). Photographs were taken with Nikon D200 dSLR camera outfitted with a Micro-Nikkor 60mm lens. Color terms are those of Komerup and Wanscher (1978). All specimens were dried in a Nesco® food dehydrator, packed in ziplock bags, and hand carried back to the U.S. by Desjardin and Perry. Microscopic data were derived from each of the dried specimens using an Olympus CH30 compound microscope and Leica Zoom 2000 dissecting scope. Radial and scalp sections were made of the pileus and stipe surfaces, and cross-sections were made of the lamellae to examine cell-types and tissue organization. All material was rehydrated in ethanol followed by distilled water and potassium hydroxide (KOH). All materials were also stained with Melzer’s reagent to test for amyloid, dextrinoid, or negative reactions, and with Congo Red to make hyphal walls more apparent and to assist with drawing tissues. All pertinent cell-types were illustrated at a minimum of 1000X with the aid of a drawing tube. Measurements of basidia, cystidia, and hyphae were made from the broadest portion and any apical or lateral appendages observed were measured independently. Basidiospores were measured in profile, with a minimum of 20 4 measured per specimen. Statistical analyses were performed for the basidiospores as follows: xm, the arithmetic mean of the length by the spore width (± standard deviation) from n spores measured in a single specimen; xmr, the range of spore means, and Xmm, the mean of spore means (± SD) when more than one specimen is available; Q, the quotient of spore length by spore width in any one spore, indicated as a range of variation in n spores measured; Qm, the mean of Q-values in a single specimen; Qmr, the range of Qm values and Qmm, the mean of Qm values where more than one specimen is available; n, the number of spores measured per specimen; s, the number of specimens involved. All cited specimens are deposited in the H.D. Thiers Herbarium, Dept, of Biology, San Francisco State University (SFSU). Genomic DNA was extracted from dried material using the Extract-N-Amp Plant PCR Kit (Sigma-Aldrich, St. Louis, MO) following the manufacturer instructions. If poor quality sequences were obtained, DNA was re-extracted using E.Z.N.A. Forensic DNA Extraction Kit and E.Z.N.A. Fungal DNA Mini Kit (Omega-Bio-tek Inc. Norcross, GA) following the manufacturer protocol. For all specimens, the nuclear ribosomal internal transcribed spacer region (ITS) was amplified using primers ITS1-F and ITS4 (Gardes and Bruns 1993, White et al. 1990). The nuclear ribosomal large subunit (nLSU) was amplified using primers LROR and LR7, LR5 or LR16 (Moncalvo et al. 2002). The amplified products were then cleaned using the Exo-SAPit kit (Affymetrix, Santa Clara, CA) and sent to Elim Biopharmaceuticals (Hayward, CA) for sequencing with the same primers that were used during the PCR amplification process. All ITS and LSU sequences generated were deposited in GenBank. Sequences were edited and assembled with the program Geneious (Biomatters Ltd., Auckland, New Zealand). Collection BAP 634 Mycena lamprospora, did not yield a useable high-quality ITS sequence due to multiple indels and therefore is not included in the ITS analyses. Additionally, the collections Hydropus murinus (BAP 657), Mycena brunneoviolacea (BAP 594), Clitocybula intervenosa (BAP 588), Mycena lasiopus (BAP 635), and Mycena aff holoporphyra (DED 8334), did not yield useable high-quality nLSU sequences due to indels and therefore are not included in the nLSU analyses. 5

Mesquite (Maddison and Maddison 2015) was used to align sequences of mycenoid fungi from ST&P with a broader sampling of mycenoid taxa downloaded from GenBank. Parsimony analyses were completed in PAUP* 4.0a 161 (Swofford 2002) and consisted of a heuristic search with 500 stepwise addition replicates, TBR branch swapping, equal weighting of all characters and holding no more than 50 topologies per replicate. The remaining phylogenetic analyses were performed within the CIPRES Science Gateway (Miller et al. 2010). Maximum likelihood analyses (Felsenstein 1981) were conducted using RAxML 8.2.10 (Stamatakis 2014) under the GTRGAMMA model and consisted of 100 alternative runs using the default parameters, with node support estimated by 100 RAxML bootstrap replicates. Bayesian analyses of the nLSU dataset used Metropolis Coupled MCMC methods as implemented in MrBayes 3.2.6 (Huelsenbeck and Ronquist 2001, Ronquist and Huelsenbeck 2003) under a GTR+G model sequence of evolution. The analysis consisted of two parallel searches, run for 90 million generations and initiated with random starting trees, temperature set to 0.05 and 2 swaps per generation. Default settings were used in MrBayes to set unconstrained branch lengths and uninformative topology priors. Eight chains were sampled every 30000 generations for a total of 3,000 trees each, sampled from the posterior distribution. Those trees sampled prior to the runs reaching an average standard deviation of split frequencies of 0.027 were discarded as the burn-in, while the remaining trees were used to calculate the posterior probabilities of the individual . Bayesian analyses of both ITS datasets were performed using Metropolis Coupled MCMC methods as implemented in MrBayes under a HKY+G model sequence of evolution. The ITS Mycenoid dataset consisted of two parallel searches, run for 90 million generations and initiated with random starting trees, and default settings for the remaining criteria to set unconstrained branch lengths and uninformative topology priors. Eight chains were sampled every 10,000 generations for a total of 9,000 trees each, sampled from the posterior distribution. Those trees sampled prior to the runs reaching an average standard deviation of split frequencies of 0.01 were discarded as the burn-in, while the remaining trees were used to calculate the posterior probabilities of the individual clades. The ITS Hydropoid dataset consisted of two 6 parallel searches, run for 10 million generations and initiated with random starting trees, and default settings for the remaining criteria to set unconstrained branch lengths and uninformative topology priors. Eight chains were sampled every 1,000 generations for a total of 10,000 trees each, sampled from the posterior distribution. Those trees sampled prior to the runs reaching an average standard deviation of split frequencies of 0.01 were discarded as the bum-in, while the remaining trees were used to calculate the posterior probabilities of the individual clades. The aligned ITS and LSU tree files have been deposited in TreeBase (submission #22780).

Results

Twenty-four specimens of mycenoid fungi were collected on the islands of Sao Tome and Principe. Based on a combination of morphological and molecular characters these represent 19 species belonging to two lineages, the and the hydropoid . Molecular sequences of the nuclear large subunit (nLSU) were generated to inform which lineage each specimen belonged to, while internal transcribed spacer (ITS) sequences, the fungal bar code region, were used to aid in delimiting species. Phylogenetic analyses were performed separately on nLSU and ITS sequences. The backbone for our nLSU phylogeny is based on 175 sequences of Agaricales from Matheny et al. (2006) “Major clades of Agaricales: a multilocus phylogenetic overview.” Fibulorhizoctonia sp. was chosen as the outgroup based on results from their study. A total of 565 characters were deemed to ambiguous to align and were excluded prior to analysis. The nLSU dataset consists of 1,454 characters from 193 ingroup taxa and contains 615 variable positions of which 447 are parsimony informative. Parsimony analysis retained 150 trees that were equally parsimonious (best score = 5,910). Maximum likelihood analyses recovered a single topology that is presented here (Fig. 1). Bayesian analyses reached an average standard deviation of split frequencies of 0.0174 7

after 90 million generations, and the first 1,500 trees sampled from the posterior distribution were excluded as the burn-in. Two ITS datasets were used to further analyze the relationships amongst the ST&P specimens and a broader sampling of mycenoid species, and to aid in species delimitation. The Mycenaceae dataset includes 18 ITS sequences of ST&P collections of Mycena and Filoboletus that were generated for this project, and a total of 46 sequences of Mycena, , Filoboletus and that were available from GenBank. brunneocaperata J.A. Cooper was chosen as the outgroup because it belongs to the clitocyboid clade which is sister to the Mycenaceae in our nLSU analysis. A total of 112 characters were deemed to ambiguous to align and were excluded prior to analysis. This ITS dataset consists of 1,030 characters from 63 ingroup taxa and contains 734 variable positions of which 466 are parsimony informative. Parsimony analysis retained 9 trees that were equally parsimonious (best score = 3,122). Maximum likelihood analyses recovered a single topology that is presented here (Fig. 2). Bayesian analyses reached an average standard deviation of split frequencies of 0.01 after 43.955 million generations, and the first 4,500 trees sampled from the posterior distribution were excluded as the burn-in. The second ITS dataset combines sequences of species belonging to the hydropoid clade. This dataset includes a total of 37 sequences of species in the genera Clitocybula, Hydropus, Megacollybia, , Pseudoclitocybe, , and Mycena sect. Radiatae (M auricoma). Five of these sequences were generated for this project, while 32 sequences were available from GenBank and used in previous research (Latha et al. 2015, Dutta et al. 2017). gibba (Pers.) Harmaja was chosen as the outgroup based on results from the Latha et al. (2015) study. A total of 134 characters were deemed to ambiguous to align and were excluded prior to analysis. The hydropoid clade ITS dataset consists of 886 characters from 36 ingroup taxa and contains 585 variable positions of which 416 are parsimony informative. Parsimony analysis retained 8 trees that were equally parsimonious (best score = 2,107). Maximum likelihood analyses recovered a single topology that is represented here (Fig. 8

3). Bayesian analyses reached an average standard deviation of split frequencies of 0.01 after 840,000 generations, and the first 2,500 trees sampled from the posterior distribution were excluded as the burn-in. Sequences of the nLSU gene region were used to help place the mycenoid fungi of ST&P into their appropriate genera. The nLSU phylogeny (Fig. 1) indicates that the ST&P species belong to two different clades, the Mycenaceae clade and hydropoid clade. The Mycenaceae clade (98% BS, 1.0 PP support) contains 13 species of Mycena and one Filoboletus from ST&P, and is segregated into two well-supported clades. The topology of our nLSU phylogenetic tree is similar to that of Matheny et al. (2006) and places the Mycenaceae in the Tricholomatoid clade along with the Clitocybeae, , Entolomataceae and Lyophyllaceae. The hydropoid clade has low support in our nLSU phylogeny. It contains one Clitocybula sequence, one Hydropus sequence and "Trogia delicata" (of Mycena sect. Radiatae) from ST&P. The hydropoid clade is sister to a clade containing the Omphalotaceae, , Cyphellaceae and Physalacriaceae, consistent with the results of Matheny et al. (2006), but with low support. Within the hydropoid clade neither Clitocybula nor Hydropus form monophyletic lineages. Sequences of the ITS gene region were used to help delimit species and suggest relationships within a broad sampling of mycenoid species. Based on our ITS phylogeny of the Mycenaceae taxa, the genus Mycena, with M. galericulata as the type species, is paraphyletic if one accepts Filoboletus, Favolaschia, and Panellus as distinct genera. In addition, most of the currently accepted infrageneric taxa are not monophyletic, with a few exceptions. Section Calodontes in our small sample set of nine species forms a well-supported clade (97% BS, 1.0 PP). Four species within sect. Sacchariferae form a well-supported clade (92% BS, 1.0 PP), although excluding M. adscendens which is the type species for this section. The poroid mycenoid taxa, belonging in Filoboletus and Favolaschia, cluster together with 99% BS and 1.0 PP support. The ITS phylogeny of the hydropoid clade (Fig. 3) shows a similar topology to that published previously (Latha et al. 2015, Dutta et al. 2017). This tree includes two 9 species of Hydropus, one Clitocybula and "Trogia delicata" from ST&P, and clearly indicates that each represents a distinct . Both Clitocybula and Hydropus represent non-monophyletic groups."Trogia delicata", which based on morphology belongs to Mycena sect. Radiatae, is distant from Mycena auricoma, a species currently accepted in that section. The former species placement within a group of Clitocybula is probably due to long-branch attraction. 10

Artificial Key to the Mycenoid species from Sao Tome and Principe

1. Pileosetae present...... Mycena breviseta 1’. Pileosetae absent...... 2

2. Stipe covered by a thick, hyaline glutinous layer...... Mycena lamprospora 2’. Stipe lacking a glutinous layer...... 3

3. Pileus surface initially with sugar-like granules; pileipellis with numerous acanthocysts ...... 4 (section Sacchariferae) 3’. Pileus surface lacking sugar-like granules; pileipellis lacking acanthocysts...... 7

4. Pileipellis with thick-walled cherocytes...... 5 4’. Pileipellis lacking cherocytes...... 6

5. Cherocytes pear-shaped; cheilocystidia absent; pileus marginal cystidia present...... Mycena oboensis 5’. Cherocytes irregularly-shaped; cheilocystidia present; pileus marginal cystidia absent ...... Mycena lasiopus

6. Caulocystidia densely spinulose; stipe lacking a basal disc Mycena alphitophora 6’. Caulocystidia smooth; stipe arising from a small basal disc Mycena discogena

7. Caulocystidia irregularly cylindrical to filamentous with apical finger-like projections (antenna-like)...... 8 7’. Caulocystidia a different shape...... 9

8. Basidiospores amyloid; basidia 2-spored; tramal tissues dextrinoid; clamp connections absent...... Mycena longinqua 11

8’. Basidiospores inamyloid; basidia 4-spored; tramal tissues inamyloid; clamp connections present...... Mycena antennae

9. Pileipellis hyphae with finger-like diverticulate projections...... 10 9’. Pileipellis hyphae non-diverticulate...... 13

10. Lamellae subporoid to reticulate...... Mycena aff. holoporphyra 10’. Lamellae non-intervenose...... 11

11. Stipe arising from a small basal disc; pileipellis an ixolattice of gelatinized hyphae and differentiated terminal cells...... Mycena aff. discobasis 11’. Stipe lacking a basal disc; pileipellis non-gelatinous, terminal cells undifferentiated . 12

12. Pileus yellow to yellowish grey; cheilocystidia of 2 distinct types: 1) clavate to narrowly lageniform or fusoid, many with apical papilla; and 2) clavate to narrowly lageniform, fusoid or cylindrical with few to numerous finger-like projections...... Mycena solis 12’. Pileus dark brown to light brown or brownish grey; cheilocystidia of 1 type, fusoid to narrowly lageniform, acute, occasionally with one or more apical projections...... Mycena galopus

13. Basidiospores inamyloid, globose to subglobose; basidia 2-spored or rarely 3-spored ...... Hydropus globospora 13'. Basidiospores amyloid, subglobose to ellipsoid; basidia 4-spored, seldom 2-spored ...... 14

14. Pleurocystidia present 15 14’. Pleurocystidia absent 16 12

15. Pileus 4-11 mm diam, dark brown to greyish brown with dark brown granules; lamellae decurrent...... Hydropus murinus 15’. Pileus 15-31 mm diam, dark violet-brown, glabrous; lamellae ascending-adnate with a short decurrent tooth...... Mycena brunneoviolacea

16. Pileus white with disc and striae greyish turquiose; lamellae free, non-intervenose ...... “Trogia delicata” 16'. Pileus brown to brownish grey or dark greyish black; lamellae adnexed to ascending- adnate or decurrent, strongly intervenose to reticulate-subporoid...... 17

17. Cheilocystidia absent; lamellae decurrent...... Clitocybula intervenosa 17'. Cheilocystidia present; lamellae adnexed to ascending-adnate...... 18

18. Pileus pustulate-tesselate, brown to greyish brown with grey margin; lamellae reticulate-subporoid; caulocystidia conspicuous...... Filoboletus pallescens 18'. Pileus glabrous to felt-like, dark greyish black fading to ash grey; lamellae strongly intervenose; caulocystidia rare...... Mycena phaeonox 13

Taxonomy

Mycena Section Calodontes Maas. Geest.

1. Mycena brunneoviolacea A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Figs. 4, 5) MycoBank no.: MB 825510 Holotype:—AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00°16.557\ E06°36.326’, 11 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 594 (SFSU). Etymology:—brunneo- = brown, -violacea = violet - referring to the pileus and stipe color. Diagnosis - Pileus 15-31 mm diam, initially broadly campanulate, becoming broadly convex to planar, margin pellucid-striate or striate, eroding in age; surface moist, glabrous, hygrophanous, disc dark brown (5-6F6-7) or dark violet-brown (9-10F4-5), fading to greyish brown (5-6E4-5) away from disc, pale grey to pale cream-grey, ash- grey or greyish ruby (12C-D3-4) with lilac tones towards the margin. Context thin, concolorous with pileus. Lamellae ascending-adnate to adnexed with a short decurrent tooth, subdistant with 1-2 series of lamellulae, non-intervenose or dorsally weakly intervenose, white to pale cream-grey, pale brownish grey or pale grey; edges entire, concolorous. Stipe 40-65 x 1.5-2.5 (apex) x 3-5 (base) mm, central, terete or compressed, broadened towards the base, hollow; surface moist to dry, smooth, glabrous or minutely pruinose, greyish brown (8E-F3-4) to brownish grey (4-5D-E4) or pale grey with hints of greyish ruby (12C-D3—4), base with white tomentum. Odor indistinct. Basidiospores 5.6-8.0 x 3.2^1.8 pm [xmr= 6.5-7.3 x 3.8-4.4 pm, xmm= 6.93 ± 0.6 x 3.88 ± 0.1 pm, Q = 1.3-2.3, Qmr= 1.73-1.86, Qmm= 1.80 ± 0.1, n = 20, s = 2], ellipsoid to oblong, smooth, hyaline, amyloid, thin-walled. Basidia 17.6-25 x 4.8-6.4 pm, clavate, 4-spored, sterigmata up to 6.4 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Cheilocystidia 32-72 x 9.6-18.4 pm, narrowly lageniform to lageniform, occasionally fusoid-strangulate, hyaline, inamyloid, thin-walled. 14

Pleurocystidia common, similar to cheilocystidia. Pileipellis a cutis; hyphae 2-8 pm diam, repent, cylindrical, non-diverticulate, hyaline, inamyloid, thin-walled, non- gelatinous. Hypodermium composed of inflated cells up to 38 pm diam, globose to ovoid, smooth, hyaline, dextrinoid, thin-walled; interspersed with numerous laticiferous hyphae up to 4.0 pm diam with knob-like projections. Lamellar regular; hyphae 2.5-19 pm diam, hyaline, dextrinoid, non-gelatinous, thin-walled; interspersed with numerous laticiferous hyphae up to 3.2 pm diam with knob-like projections. Stipitipellis a cutis; cortical hyphae 4.0-8.0 pm diam, parallel, cylindrical, hyaline, dextrinoid, thin- walled, non-gelatinous; medullary hyphae 6.4-12.0 pm diam, smooth, dextrinoid, thin- walled; interspersed with numerous laticiferous hyphae up to 8.0 pm diam with knob-like projections. Caulocystidia rare or scattered, 36-80 x 4.8-10.4 pm, subcylindrical to fusoid, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Gregarious and cespitose on wood in upland forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00°16.557’, E06°36.326’, 11 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 594 (Holotype, SFSU); same location, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 656 (SFSU). Notes - Mycena brunneoviolacea is distinguished by medium-sized basidiomes with greyish brown to violet-brown, striate pileus, white to pale grey lamellae, a glabrous to minutely pruinose, hollow, brownish grey stipe with a hint of greyish ruby, an indistinct odor, ellipsoid basidiospores with mean 6.9 x 3.9 pm, lageniform to fusoid- strangulate hymenial cystidia and caulocystidia, a cutis-type pileipellis of non- diverticulate hyphae, and growth on decayed wood. In combination, these features indicate placement in sect. Calodontes. Pairwise comparisons of aligned, overlapping ITS sequences of Mycena brunneoviolacea (BAP 594) with the top 10 BLAST results show 94.5-95.4% similarity to several north temperate species of sect. Calodontes (M diosma Krieglst. & Schwobel, M. pura (Pers.) P. Kumm.), and 88.4-95.1% similarity to several Malaysian species of 15 sect. Calodontes (M cahaya A.L.C. Chew & Desjardin, M. sinar A.L.C. Chew & Desjardin, M. seminau A.L.C. Chew & Desjardin). As reported above, the closest similarity to M. brunneoviolacea within sequenced members of sect. Calodontes is 95.4% (>4.5% difference). In a three-gene phylogeny of the complex, Harder et al. (2013) reported that a 1.5-3% ITS sequence difference threshold level underestimated diversity within the group. Hence, we are recognizing the Sao Tome taxon as a distinct species. Mycena brunneoviolacea is morphologically similar to the Malaysian species noted above, which all differ in forming bioluminescent basidiomes with a white or beige stipes and growth on decomposing leaves. In addition, M. seminau and M. sinar lack pleurocystidia. Mycena griseoviolacea Metrod, described from Madagascar (Metrod 1949), differs in forming much broader basidiospores (5.5-5.7^im) and longer hymenial cystidia (80-120 (im). Two lignicolous species described from , viz., M. clarkeana Grgur. and M. subcorticalis Grgur., differ in basidiomes coloration (reddish brown to brick red) and larger basidiospores (means 9.9-10 x 6.3-6.7 pm) (Grgurinovic 2003). In the ITS phylogeny (Fig. 2), M. brunneoviolacea falls into the sect. Calodontes clade, sister to M. pearsoniana Dennis ex Singer.

2. Mycena aff. holoporphyra (Berk. & M.A. Curtis) Singer, Sydowia 15: 64. 1962. (Fig. 6) Basionym: Agaricus holoporphyrus Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10(45): 284 (1868) 1869. Homotypic synonyms: Clitocybe holoporphyra (Berk. & M.A. Curtis) Sacc. Syll. fung. (Abellini) 5: 196. 1887. Dictyoploca holoporphyra (Berk. & M.A. Curtis) Dennis, Trans. Brit. Mycol. Soc. 34: 475. 1951. 16

Pileus 11-19 mm diam, broadly convex to campanulate with raised disc, becoming broadly convex to planar, margin pellucid-reticulate to tesselate, split in age; surface moist, glabrous, hygrophanous, dull, disc dark brown (7-8F4-8) with margin pale reddish brown (8E4). Lamellae horizontal, adnate with a decurrent tooth to arcuate, distant to subdistant with 1-2 series of lamellulae, subporoid to reticulate with shorter cross lamellae, non-marginate, greyish brown (8D-E3). Stipe 20-30 x 1.5-2 mm, central, terete or flattened towards base, hollow; surface moist to dry, glabrous, dull, dark brown (8F4-5). Odor and taste not recorded. Basidiospores 6.4-8.0 x 4.0-4.8 pm [xm= 7.36 ± 0.49 x 4.28 ± 0.39 pm, Q = 1.5- 2, Qm= 1.73 ± 0.17, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin-walled. Basidia 20-25 x 7.2-8.0 pm, clavate, 4-spored, rarely 2-spored, sterigmata up to 8.0 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge sterile. Cheilocystidia 34-45 x 8.0-8.8 pm, fusoid to lageniform, hyaline, inamyloid, thin- walled. Pleurocystidia absent. Pileipellis a cutis; hyphae 1.6-4.8 pm diam, repent, cylindrical, smooth or sparsely diverticulate, hyaline, inamyloid, thin-walled, non- gelatinous; diverticula 1-2 x 0.5-1 pm, cylindrical. Hypodermium composed of inflated cells up to 19.2 pm diam, globose to ovoid, smooth, hyaline, dextrinoid, thin-walled. Lamellar trama regular; hyphae 2.5-8.8 pm diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 1.6-6.4 pm diam, parallel, cylindrical, hyaline, dextrinoid, thin-walled, non-gelatinous; medullary hyphae 8-20 pm diam, smooth, dextrinoid, thin-walled. Caulocystidia common at apex, scattered elsewhere, 30- 67 x 5.6-11.2 pm, subcylindrical to fusoid, occasionally subcapitate or with one or more broad knobs, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Solitary in leaf mulch or soil in primary forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00° 16.557’, E06°36.326’, 25 April 2008, coll. by D.E. Desjardin, DED 8334 (SFSU). 17

Notes - Mycena aff. holoporphyra is characterized by a convex to campanulate, reticulate-tesselate, dark brown pileus, arcuate, subporoid to reticulate, greyish brown lamellae, a dark brown stipe, basidiospores with mean 7.4 x 4.3 |im, fusoid to lageniform cheilocystidia, no pleurocystidia, a cutis-type pileipellis of smooth or sparsely diverticulate, non-gelatinous hyphae, subcylindrical to fusoid caulocystidia, and growth in leaf mulch in primary forest. In combination, these features indicate placement in sect. Calodontes. The New World M. holoporphyra differs from the Sao Tome specimen in forming deep violaceous to vinaceous basidiomes and growing on wood (Pegler 1983). In all other macro- and micromorphological features they are indistinguishable. Until more material becomes available for the dark brown form described herein, we recognize it as Mycena aff. holoporphyra. Pairwise comparisons of aligned, overlapping ITS sequences of the Sao Tome specimen (DED 8334) with the top ten BLAST results show 93.9-94.3% similarity to several sequences of M. pearsoniana (JN182201, FN394615, FN394616, FN394612), a member of sect. Calodontes. No sequences of New World M. holoporphyra are available for comparison. In the ITS phylogeny (Fig. 2), Mycena aff. holoporphyra falls into the clade with other members of sect. Calodontes, sister to the bioluminescent Malaysian M. cahaya.

Mycena Section Exornatae Maas Geest.

3. Mycena aff. discobasis Metrod, Les Mycenes de Madagascar: 131. 1949. (Figs. 7, 8) Pileus 13-18 mm diam, broadly convex to plano-convex, centrally depressed, striate to depression; surface glutinous to viscid, glabrous, shiny, disc pale greyish white to light yellowish brown (5D5) or with a brown ring, pale watery whitish grey or white away from disk. Context <1 mm thick, concolorous with pileus. Lamellae free to subfree, close with 1-2 series of lamellulae, moderately broad (1-1.5 mm), white. Stipe 15-19 x 18

1-2 mm, central, terete, cylindrical, hollow, arising from a small (3 mm diam), pale yellowish brown (C5D5) basal disc; surface dry, pruinose overall, white or watery white grey. Odor none. Basidiospores (6.4-) 7-8 (-9.5) x (3.2-) 4.0-5.1 (-5.6) pm [xmr= 7.3 x 3.9—4.6 pm, xmm = 7.3 ± 0 x 4.22 ± 0.5, Q = 1.4-2.3, Qmr= 1.61-1.88, Qmm= 1.75 ± 0.2, n = 20- 30, s = 2], ellipsoid, smooth, hyaline, weakly amyloid, thin-walled. Basidia 12.8-26 x 5.6-8.8 pm, clavate, 4-spored, rarely 2-spored, sterigmata up to 4.8 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge sterile. Cheilocystidia numerous, 38-69 x 11-18 pm, broadly clavate to fusiform, often mucronate, hyaline, inamyloid. Pleurocystidia absent. Pileipellis an ixolattice of hyphae and differentiated terminal cells; hyphae 0.8-4 pm diam, irregular, cylindrical, smooth, embedded in a gel layer up to 140 pm thick, hyaline, inamyloid, thin-walled. Terminal cells numerous; 31- 78 x 4.0-14 pm, clavate, densely spinulose, embedded in upper portion of gel layer, hyaline, inamyloid, thin-walled; spinulae 0.8-7.2 x 0.5-1.0 pm, cylindrical. Pileus marginal cells not differentiated from terminal cells. Hypodermium composed of inflated cells up to 25 pm diam, globose to ovoid, smooth, hyaline, dextrinoid, thin-walled. Hymenophoral trama regular to subregular; hyphae 1.6-23 pm diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 1.6-11.2 pm diam, parallel, cylindrical, hyaline, inamyloid, thin-walled; medullary hyphae 7.2-26 pm diam, smooth, dextrinoid, thin-walled, with rare laticiferous hyphae up to 4 pm diam with knob-like projections. Caulocystidia abundant; 49-142 x 9.6-22 pm, fusiform to narrowly cylindrical, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Solitary on decaying dicot wood in secondary upland and primary forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, N00° 16.557’, E06°36.326’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 658 (SFSU); same location, 11 April 2008, coll. by D.E. Desjardin, DED 8211 (SFSU). 19

Notes - Mycena discobasis, described from Madagascar, is distinguished by a glutinous, striate, plano-convex-depressed, white pileus with greyish white to yellowish brown disc and striate, subtree lamellae, a dry, pruinose, white stipe that arises from a well-developed basal disc, basidiospores with mean 7.3 x 4.2 |am, broadly clavate to fusiform cheilocystidia, no pleurocystidia, an ixo-lattice-type pileipellis with clavate, densely spinulose terminal cells, no differentiated pileus marginal cells, fusiform caulocystidia, and growth on wood. The specimens from Sao Tome differ from the protologue and from a description of material from Sao Paulo, Brazil determined as M. discobasis, in forming smaller basidiospores. Metrod (1949) reported the basidiospores of Madagascar material as 7.5-9.5 x 6.5 ^im, while Desjardin et al. (2007) reported them from Brazil material as 9-10.5 x 6-7.5 |am. Neither of the latter papers mentioned differentiated pileus marginal cells. (Berk. & M.A. Curtis) Sacc. and M. margarita (Murrill) Murrill, two bioluminescent members of sect. Exornatae, differ subtly from M. discobasis in presence of distinctive pileus marginal cystidia and ITS sequence data. Desjardin et al. (2007) report M. discobasis from Brazil as bioluminescent, whereas the Sao Tome specimens were not bioluminescent. Pairwise comparisons of aligned, overlapping ITS sequences of Mycena aff. discobasis (BAP 658, DED 8211) with the top 10 BLAST results indicate 93.4-94.0% similarity to ten sequences of M. chlorophos. ITS sequences of M. margarita are not available for comparison. In the ITS phylogeny (Fig. 2), Mycena aff. discobasis is sister to M. chlorophos with 100% BS and 1.0 PP support. 20

Mycena Section Fragilipedes (Fr.) Quel.

4. Mycena solis A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Figs. 9, 10) MycoBank no.: MB 825511 Holotype:—AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00° 16.557’, E06°36.326\ 11 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 592 (SFSU). Etymology:—solis = sun - referring to the sun yellow color of the pileus surface. Diagnosis - Pileus 8-14 mm diam, hemispherical to parabolic when young, expanding to obtusely conical or campanulate, margin decurved to upturned, striate to sulcate; surface moist, glabrous, disc and striations disc and striations yellowish brown to brownish yellow (4-5C5-6), yellowish grey (3—4B5-6) towards the margin, pale grey at margin and between striations. Context thin; concolorous with the pileus. Lamellae ascending-adnate with a short decurrent tooth, subdistant (12-14) with 1-2 series of lamellulae, pale greyish white to pale cream; edges smooth, concolorous. Stipe 30^18 x 1-1.5 (apex) x 2-2.5 (base) mm, central, terete, subcylindrical or enlarged downward, inflated and twisted in larger specimens; surface dry, polished, smooth, glabrous, yellowish brown (3—4B-C5-6), some with pale grey apex, some with brown tones at the base, attached to substrate with scant, short, white, non-radiating fibrils. Odor and taste indistinct. Basidiospores 7.2-8.8 (-9.6) x 4.0-4.8 |am [xm= 8.05 ± 0.7 x 4.36 ± 0.41 (am, Q = 1.50-2.20, Qm= 1.86 ± 0.12, n = 22, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 19-24 x 4.8-5.6 (am, clavate, 4-spored, sterigmata up to 8.8 (am long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge sterile. Cheilocystidia of 2 types; 1) 30-59 x 7.2-10.4 |am, clavate to narrowly lageniform or fusoid, many with an apical papilla, hyaline, inamyloid, thin-walled, 2) 30—49 x 7.2-10.4 (am, clavate to narrowly lageniform, fusoid or cylindrical, with few to numerous finger-like projections, hyaline, inamyloid, thin-walled; projections 1.6-8 x 0.8-1.6 |am, cylindrical. Pleurocystidia abundant, similar to type 1 cheilocystidia. Pileipellis a cutis; hyphae 0.8- 21

2.4 (am diam, repent, cylindrical, irregularly diverticulate with finger-like projections, hyaline, inamyloid, thin-walled, non-gelatinous; diverticula 1.6—4 x 0.8-1.6 |am, cylindrical. Hypodermium of inflated hyphae up to 37 (am diam, ovoid to globose, hyaline, dextrinoid, thin-walled. Hymenophoral trama subregular to irregular; hyphae 3.2-8.8 (am diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 1.4-2.4 |am diam, parallel, cylindrical, diverticulate with finger-like projections, hyaline, dextrinoid, thin-walled, diverticula 2.4—4 x 0.8-2.4 (am, cylindrical; medullary hyphae 6.4—24 (am diam, smooth, dextrinoid, thin-walled; interspersed with laticiferous hyphae up to 8.8 (am diam with knob-like projections 1.6-4 x 4-6 (am. Caulocystidia absent. Clamp connections present in all tissues. Habitat and known distribution - Gregarious to scattered on woody debris in upper elevation forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00° 16.557’, E06°36.326\ 11 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 592 (Holotype, SFSU). Notes - Mycena solis is distinguished by a small, sulcate, campanulate, yellowish brown pileus, ascending-adnate, subdistant, pallid, non-marginate lamellae, a glabrous, yellowish brown stipe, amyloid basidiospores with mean 8 x 4.4 (am, two types of relatively narrow (7.2-10.4 |am) cheilocystidia, clavate to lageniform or fusoid pleurocystidia, diverticulate pileipellis and stipitipellis hyphae, no differentiated caulocystidia, and growth on woody debris. In combination, these features indicate placement in sect. Fragilipedes, allied with a few yellow-pigmented species, such as M. citrinomarginata Gillet, M. junquillina Dennis, and M. luteola Maas Geest. & de Meijer. Mycena citrinomarginata, a north temperate species, differs in forming yellow-marginate lamellae, differentiated caulocystidia, and no pleurocystidia (Maas Geesteranus 1988a). Mycena junquillina forms basidiomes with sulphur yellow lamellae, a shorter (5-15 mm), xanthine orange, pruinose stipe, basidiospores with mean 7.4 x 5 (am, more consistently fusoid hymenial cystidia, and hair-like caulocystidia (Pegler 1983). Mycena luteola, described from Parana, Brazil, forms much smaller (1.5—4 mm diam) pileus, a pale straw- 22 colored, puberulous stipe that arises from a patch of radiating fibrils, broader cheilocystidia (10-15 |j.m), stipitipellis hyphae with broader and longer diverticula, some thick-walled hair-like caulocystidia, thick-walled (-2 (im) stipe medullary hyphae, and growth on leaves (Maas Geesteranus and de Meijer 1997). The ITS sequence of M. solis (BAP 592) was rather short (380 BP) and of poor quality. In pairwise comparisons with the top 10 BLAST results, M. solis showed 97.7% similarity to Mycena cf. sanguinolenta (JX434650), and 96.3% similarity to nine uncultured Mycena clones. In the ITS phylogeny (Fig. 2), M. solis is sister to M. sanguinolenta in a well-supported clade (97% BS, 1.0 PP).

Mycena Section Granuliferae Maas Geest. & de Meijer

5. Mycena antennae A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Fig. 11) MycoBank no.: MB 825512 Holotype:—AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00° 16.557’, E06°36.326’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 660 (SFSU). Etymology:—referring to the antenna-shaped caulocystidia and the site where the holotype was collected, Macambrara radio antenna area. Diagnosis - Pileus 2-3 mm diam, broadly convex, margin fimbriate, pellucid- striate; surface glabrous, disc with tiny dark brown spots, off-white elsewhere. Lamellae ascending, adnexed, subdistant with 1-2 series of lamellulae, white; edges concolorous. Stipe 7-22 x <0.5 mm, central, terete, cylindrical, arising from a small basal disc (0.5 mm diam) densely covered with short white hairs; surface minutely white-pubescent, off- white, developing brown tones in age. Odor none; taste indistinct. Basidiospores (7.2-) 8.8-10.5 x 3.2-4.8 (j.m [xm= 9.16 ± 0.71 x 4.52 ± 0.47 )im, Q = 1.67-2.75, Qm = 2.05 ± 0.1, n = 20, s = 1], elongated-ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 11.2-15 x 8.8-10.4 (am, clavate, 4-spored, sterigmataup to 3.2 (im long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge fertile. 23

Cheilocystidia abundant, scattered; 8.8-27 x 5.8-14 |im, clavate, apically spinulose, hyaline, inamyloid, thin-walled; spinulae 1.6-5.6 x <0.8 (am. Pleurocystidia absent. Pileipellis a cutis; hyphae 7.2-15 pm diam, repent, cylindrical, densely spinulose, hyaline, inamyloid, thin-walled, non-gelatinous; spinulae 0.5-2 x 0.5 pm, cylindrical. Hypodermium undifferentiated. Pileus trama hyphae 5-25 pm diam, cylindrical to slightly inflated, hyaline, inamyloid, thin-walled. Hymenophoral trama subregular; hyphae 2.4-5.6 pm diam, hyaline, inamyloid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 2.4-5.6 pm diam, parallel, cylindrical, smooth, hyaline, dextrinoid, thin-walled; medullary hyphae 8.8-22 pm diam, smooth, dextrinoid, thin-walled, numerous laticiferous hyphae up to 4 pm diam with occasional knob-like projections 5.6- 6.4 x 1.6-3.2 pm. Caulocystidia numerous; 8.0-114 x 1.6-6.4 pm, irregularly cylindrical with a broader base, seldom geniculate, with apical finger-like projections, hyaline, inamyloid, thin-walled; apical projections 1.6-11.2 x 0.8-2.4 pm. Clamp connections present in all tissues. Habitat and known distribution - Gregarious on sticks in upland secondary forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00°16.557’, E06°36.326’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 660 (Holotype, SFSU). Notes - Mycena antennae is characterized by a unique combination of features, including: a tiny, glabrous, white pileus with dark brown spots on the disc, ascending, adnexed white lamellae, a pubescent white stipe that arises from a small basal disc, inamyloid, elongate-ellipsoid basidiospores with mean 9.2 x 4.5 pm, apically spinulose, clavate cheilocystidia, no pleurocystidia, a cutis-type pileipellis of densely spinulose hyphae, non-dextrinoid pileipellis and lamellar trama tissues, smooth stipitipellis cortical hyphae, irregularly cylindrical caulocystidia with apical diverticula, and growth on sticks. The closest currently accepted infrageneric taxon in which Mycena antennae could be grouped is Mycena sect. Granuliferae Maas Geest. & de Meijer (1997). The two known members of this section, viz., M. granulifera Maas Geest. & de Meijer and M. sertipes 24

Maas Geest. & de Meijer, differ in growing on leaves, having much narrower basidiospores (2.7-3.6 pm, Q = 3.2), adnate to arcuate lamellae, and different caulocystidia. Mycena antennae shows many similarities to M. longinqua of sect. Polyadelphia, especially the unusual caulocystidia, but the latter species has amyloid basidiospores, 2-spored basidia, dextrinoid tramal tissues, and lacks clamp connections. Pairwise comparisons of aligned, overlapping ITS sequences of Mycena antennae (BAP 660) with the top 10 BLAST results indicate 92.6% similarity to a sequence of (DQ490645) and 91.8-92% similarity to three sequences of M. illuminans. In the ITS phylogeny (Fig.2), M. antennae belongs in a heterogeneous clade with members of sects. Exornatae, Polyadelphia, Longisetae and Sacchariferae.

Mycena Section Lactipedes (Fr.) Quel.

6. Mycena galopus (Pers.) P. Kumm., Ftihr. Pilzk. (Zerbst): 108. 1871. (Figs. 12, 13) Pileus 7-13 mm diam, parabolic with upturned margin to campanulate, striate; surface moist, glabrous, disc and striations dark brown (5-6F6-7), fading to light brown (3-4E5-6), pale brownish grey (4C-D3^1) between striations and towards the pale grey margin. Context thin; concolorous with the pileus. Lamellae ascending-adnate with a short decurrent tooth, subdistant (13-14) with 1-2 series of lamellulae, pale grey with upper portion taking on darker tones; edges concolorous. Stipe 35-60 x 1-1.5 mm, central, terete, enlarged slightly towards base, hollow; surface smooth, glabrous, apex pale grey, greyish brown below (3-4D-E4-5), base with white tomentum; latex not observed. Odor and taste indistinct. Basidiospores 8.0-8.8 x (4.0-) 4.8-5.6 pm [xm= 8.32 ± 0.40 x 4.88 ± 0.36 pm, Q = 1.57-2.00, Qm= 1.71 ± 0.09, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 25-31 x 6.4-8.8 pm, clavate, 4-spored, rarely 2-spored, sterigmata up to 9.6 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Cheilocystidia abundant, 57-80 x 11.2-15 pm, fusoid to narrowly lageniform, acute, occasionally with 1 25 or more apical projections, hyaline, inamyloid, thin-walled. Pleurocystidia common, similar to cheilocystidia, projecting up to 56 (am beyond basidioles. Pileipellis a cutis; hyphae 0.8-2.4 |am diam, repent, cylindrical, irregularly diverticulate with finger-like projections, hyaline, inamyloid, thin-walled, non-gelatinous; diverticula 6.4-22 x 1.6-2.4 |am, cylindrical. Hypodermium of inflated hyphae up to 37 |am diam, ovoid to globose, hyaline, dextrinoid, thin-walled. Hymenophoral trama regular to subregular; hyphae 4.0- 9.6 (am diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 2.4-6.4 jam diam, parallel, cylindrical, diverticulate with finger-like projections, hyaline, inamyloid, thin-walled; diverticula 2.4-8.8 x 1.6-2.4 (am, cylindrical; medullary hyphae 2.4-8 |am diam, smooth, dextrinoid, thin-walled; interspersed with laticiferous hyphae up to 7.2 (am diam with knob-like projections, occasionally with brown granular contents. Caulocystidia absent. Clamp connections present in all tissues. Habitat and known distribution - Gregarious and scattered on soil and woody debris in upland forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1300 m, N00°16.557\ E06°36.326\ 11 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 593 (SFSU). Notes - The Sao Tome specimen of Mycena galopus is characterized by a brown to pale brownish grey, striate pileus, pale grey, ascending-adnate lamellae, a glabrous, greyish brown stipe, an indistinct odor and taste, basidiospores with mean 8.3 x 4.9 (am, acutely fusoid cheilocystidia and pleurocystidia, occasionally with one or more apical projections, a cutis-type pileipellis with long diverticula, clamp connections, diverticulate stipe cortical hyphae, no caulocystidia and growth on woody debris. White latex was not observed in the rather old basidiomes of the single collection, although morphology and ITS data indicate this specimen represents M. galopus. In addition, the basidiospores were significantly shorter than those reported from European material, viz., 8.0-8.8 x (4.0-) 4.8-5.6 (am versus 9.5-14 x 5-6.3 (am (Maas Geesteranus 1988a, Robich 2003). The Sao Tome material would key to (Fr.) Kummer, as reported from 26

East Africa by Pegler (1977) except for the lack of a strong alkaline or nitrous odor, and the much larger hymenial cystidia in Mycena galopus. The Sao Tome specimen shows similarities to M. subcana A.H. Sm., a species described from western North America on , but the latter forms larger (10-30 mm diam), dark grey pileus without brown tones, and much smaller and less projecting hymenial cystidia (only 19—42 pm long) (Smith 1947, Maas Geesteranus 1988b). Pairwise comparisons of aligned, overlapping ITS sequences of the Sao Tome specimen of M. galopus (BAP 593) showed 99% similarity to four sequences of European M. galopus, forming a well-supported clade (99% BS, 1.0 PP) in the ITS phylogeny (Fig. 2).

Mycena Section Longisetae A.H. Sm. ex Maas Geest.

7. Mycena breviseta Hohn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1, 118: 283. 1909. (Fig. 14) Pileus 1.5-2 mm diam, convex, margin becoming eroded or fimbricate in age, striate; surface covered with long, hyaline setae up to 0.4 mm long, greyish white, developing brown tones in age. Lamellae ascending to adnate, subdistant with 1 series of lamellulae, white. Stipe 10-17 x <0.5 mm, central, terete, cylindrical, hollow, arising from a small (<1 mm diam), yellowish brown basal disc; surface covered with short, minute hairs, greyish white, developing yellowish brown tones with moisture loss. Odor none and taste not recorded. Basidiospores 8.0-9.6 x 4.0^.8 pm [xm= 8.68 ± 0.54 x 4.42 ± 0.4 pm, Q = 1.67- 2.20, Qm= 1.97 ± 0.09, n = 20, s = 1], ellipsoid, smooth, hyaline, weakly amyloid, thin- walled. Basidia 12-16.8 x 7.2-9.6 pm, clavate, 4-spored, rarely 2-spored, sterigmata up to 4.8 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Cheilocystidia absent. Pleurocystidia absent. Pileipellis a cutis with differentiated terminal cells, marginal cystidia and pileosetae; hyphae 1.6-5.6 pm diam, repent, cylindrical, smooth, 27 hyaline, inamyloid, thin-walled. Terminal cells numerous; 16-31 x 10.4-18.5 |am, clavate to broadly clavate, densely spinulose, hyaline, inamyloid, thin-walled; spinulae <0.8-2.4 x <0.8 (am. Marginal cystidia 15.2-34 x 8.0-18 (am, broadly clavate to clavate, densely spinulose, hyaline, inamyloid, thin-walled; spinulae <0.8-1.6 x <0.8 |am. Pileosetae scattered; 165-365 x 8-11.2 |am, narrowly cylindrical to acicular, hyaline, inamyloid, thick-walled (up to 4.0 (am). Hypodermium composed of inflated cells up to 22 (am diam, occasionally globose to ovoid, smooth, hyaline, inamyloid, thin-walled. Hymenophoral trama regular to subregular; hyphae 1.6-4.8 |am diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis with caulocystidia; cortical hyphae 2.4- 5.6 (am diam, parallel, cylindrical, hyaline, dextrinoid, thin-walled; medullary hyphae 16- 22 (am diam, smooth, dextrinoid, thin-walled. Caulocystidia abundant; 30-111 x 4.8-15 (am, narrowly cylindrical to aculeate, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues except stipe tissues. Habitat and known distribution - Gregarious on sticks in secondary upland forest. (Principe). Material examined- AFRICA. Principe, Ro9a Pico Papagaio, N01°37.182’, E07°23.474’, 20 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 633 (SFSU). Notes - Mycena breviseta, described from Java, forms tiny basidiomes with greyish white, striate pileus with scattered hyaline setae, subdistant lamellae, a filiform, hispidulose, greyish white stipe, weakly amyloid basidiospores with mean 8.7 x 4.4 (am, no cheilocystidia or pleurocystidia, a cutis-type pileipellis with clavate to broadly clavate, densely spinulose terminal cells and marginal cystidia, hyaline pileosetae up to 365 (am long, aculeate caulocystidia, and growth on woody sticks. In combination, these features indicate placement in sect. Longisetae A.H. Sm. ex Maas Geest. Desjardin and Horak (2002) studied the holotype specimen, reported that no intact basidiomes were present, and provided a translation of the protologue. The material from Principe matches nicely with the limited data provided by Desjardin and Horak. Mycena longiseta is similar, but forms much longer pileosetae (up to 1000 |am), smaller 28 basidiospores (mean 7.3 x 3.7 |im) and marginal cystidia with smooth apices (Desjardin et al. 2002, Desjardin & Horak 2002). Pairwise comparisons of aligned, overlapping ITS sequences of Mycena breviseta (BAP 633) with the top 10 BLAST results indicate 89.5-91.9% similarity to four sequences determined as Mycena sp. from Norway (KU861567, KU861566, KU861565) and (AB512340). ITS sequences from other members of sect. Longisetae are unavailable for comparison. Mycena breviseta is on a rather long branch sister to M. stylobates in the ITS phylogeny (Fig. 2) with 82% BS and 1.0 PP support.

Mycena Section Mycena (Pers.) Roussel

8. Mycena phaeonox A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Figs. 15, 16) MycoBank no.: MB 825513 Holotype:—AFRICA. Sao Tome, Parque Nacional Obo, Lagoa Amelia Trail N00°16.922’, E06°36.062’, 14 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 615 (SFSU). Etymology:—phaeo- = dark, -nox = night - referring to the dark pileus pigmentation. Diagnosis - Pileus 7-30 mm diam, parabolic to campanulate, expanding to broadly campanulate with a flattened umbo, margin striate, becoming undulating to eroded in age; surface moist to dry, glabrous, felt-like with a 10x hand lens, disc dark greyish black initially, fading to dark ash grey (6-7F1-3) then eventually paler ash grey overall (6-7E1-2), hygrophanous from disc outwards, becoming pale grey as it dries Context thin, concolorous with pileus except white under disc. Lamellae ascending- adnate with a decurrent tooth, subdistant with 2 series of lamellulae, strongly interwoven, pale ash grey (6-7E1-2); edges paler grey, entire. Stipe 16-35 x 1-3.5 mm, central, terete, cylindrical, rarely flattened in larger specimens, hollow; surface dry, polished, minutely white-pruinose, pale grey, developing brown tones towards the base; base with white to yellow tomentum at point of attachment. Odor and taste not indistinct. 29

Basidiospores 6.4-8.0 (-8.8) x 4.0-5.6 pm [xm= 7.45 ± 0.72 x 4.79 ± 0.55 pm, Q = 1.29-2.20, Qm- 1.57 ± 0.15, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 20-36 x 5.6-8 pm, clavate, 4-spored, rarely 2-spored, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge sterile. Cheilocystidia numerous, 30-86 x 9.6-22 pm, clavate to subcylindrical, smooth or with one or more knobby apical projections, hyaline, inamyloid, thin-walled; projections 2.4-7.2 x 2-7 pm, cylindrical to knob-like. Pleurocystidia absent. Pileipellis a cutis lacking pileocystidia; hyphae 4-13 pm diam, repent, cylindrical, smooth, hyaline, inamyloid, thin-walled, non-gelatinous; terminal cells undifferentiated, repent. Hypodermium composed of inflated hyphae up to 34 pm diam, ovoid to ellipsoid, with brown vacuolar pigments, dextrinoid, thin-walled, with rare laticiferous hyphae 2-6 pm diam with knob-like projections. Hymenophoral trama appearing sarcodimitic; many hyphae 2.4-6.5, thin-walled or some thick-walled (- 1 pm), interwoven among inflated to fusiform hyphae up to 36 pm diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 0.8-8.8 pm diam, parallel, cylindrical, smooth, hyaline, dextrinoid, thin-walled; medullary hyphae 4- 22 pm diam, smooth, hyaline, dextrinoid, thin-walled, some thick-walled (-0.8 pm), occasional laticiferous hyphae 4.8-8 pm diam with knob-like projections. Caulocystidia absent or rare, as undifferentiated, terminal cells or intercalary projections, solitary or in small clusters, erect to suberect, 24-130 x 4-7.2 pm, cylindrical, obtuse, rarely with a few knob-like outgrowths, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Gregarious on rotting wood in upland forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Parque Nacional Obo, Lagoa Amelia Trail N00° 16.922’, E06°36.062’, 14 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 615 (Holotype, SFSU). Notes - Mycena phaeonox is distinguished by a campanulate, striate, dark grey pileus, ash grey, strongly interwoven lamellae, a minutely pruinose, pale grey stipe, amyloid basidiospores with mean 7.5 x 4.8 pm, broadly clavate, hyaline cheilocystidia 30 with a few knobby projections, no pleurocystidia, sarcodimitic hymenophoral trama tissue, a cutis-type pileipellis of smooth, repent hyphae lacking pileocystidia, dextrinoid tissues, few, cylindrical caulocystidia, and growth on woody debris. In combination, these features indicate placement in sect. Mycena. It shows similarities to M. maculata P. Karst, but the latter has basidiomes that become spotted reddish in age (maculate), less intervenose lamellae, larger basidiospores (8.1-9.8 x 5 2 -6 3 (im), and more coarsely diverticulate cheilocystidia (Maas Geesteranus 1985). The poorly known Mycena plectophylla var. umbrina (Dennis) Dennis has similar macromorphology, but forms smaller basidiospores (6-7 x 3-3.5 |im), has a fertile lamellar edge lacking cheilocystidia, and grows on litter (as Dictyoploca, Dennis 1951, 1970). Mycena phaeonox is similar to what Pegler (1977) called Mycena parabolica (Fr.) Quel., a problematic European species, and we follow Maas Geesteranus (1988a: 238) in recognizing Pegler’s East African taxon as different from the European taxon. Several Trogia sensu Comer (1966, 1991) species show affinities to Mycena phaeonox. Trogia subdistans Comer differs in a plano-umbilicate pileus, decurrent lamellae, clavate to ventricose cheilocystidia with a filiform apical projection, and pileocystidia and caulocystidia similar to the cheilocystidia (Comer 1966). Trogia guadalupensis (Heim) Comer shows similar macromorphology, but has inamyloid, subglobose basidiospores, clavate to subventricose cheilocystidia lacking knobby projections, lacks a hypodermium, and has pileocystidia and caulocystidia similar to the cheilocystidia (Comer 1991). Trogia atropruinosa Comer forms shorter and much narrower basidiospores (5.5-6.3 x 2.7-3 (im), clavate to ventricose cheilocystidia with a filiform apical projection and dark contents resulting in a dark-pruinose lamellar edge, and distinctive pigmented pileocystidia and caulocystidia (Comer 1966). Pairwise comparisons of aligned, overlapping ITS sequences of the Sao Tome specimen (BAP 615) with the top ten BLAST results show 94.7-95.9% similarity to eight sequences of uncultured mycorrhizal . Mycena phaeonox is sister to M. plumbea P. Karst., with limited support in the ITS phylogeny (Fig. 2), somewhat distant from other members of sect. Mycena. 31

Mycena Section Polyadelphia Singer ex Maas Geest.

9. Mycena longinqua A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Fig. 17, 18) MycoBank no.: MB 825514 Holotype:—AFRICA. Principe, Dois Irmeos area around 100 meters, N01°34.89’, E07°25.548’, 23 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 648 (SFSU). Etymology:—longinquus = outlying - referring to the outlier position of the species within section Polyadelphia, because of the non-diverticulate stipitipellis cortical hyphae and unusual caulocystidia. Diagnosis - Pileus 1.5-3 mm diam, paraboloid, margin entire, striate; surface glabrous, hygrophanous, dark grey initially fading to brownish grey with age, margin pale grey. Lamellae ascending, adnate, subdistant with 2 series of lamellulae, pale grey; edges paler. Stipe 3-8 x <0.5 mm, central, terete, cylindrical, arising from a small basal disc, covered with short white hairs; surface glabrous, smooth, pearlescent, pale whitish grey. Odor none; taste indistinct. Basidiospores 6.0-7.2 x 4.0^.8 pm [xm= 6.94 ± 0.42 x 4.40 ± 0.41 pm, Q = 1.33-1.80, Qm= 1.59 ± 0.21, n = 20, s = 1], broadly ellipsoid, smooth, hyaline, amyloid, thin-walled. Basidia 12.8-17 x 5.6-7.2 pm, clavate, 2-spored, sterigmataup to 3.2 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge mostly sterile. Cheilocystidia abundant; 18-25 x 12-20 pm, clavate, apically to entirely spinulose, hyaline, inamyloid, thin-walled; spinulae <0.8-1.6 x <0.8 pm, cylindrical. Pleurocystidia absent. Pileipellis a cutis with terminal cells; hyphae 4-16 pm diam, repent, cylindrical, densely spinulose, hyaline, inamyloid, thin-walled, non-gelatinous; spinulae <0.8-1.6 x <0.8 pm, cylindrical. Terminal cells 34-76 x 8.0-24 pm, broadly clavate, densely spinulose. Hypodermium of inflated hyphae up to 29 pm diam, ovoid, hyaline, dextrinoid, thin-walled. Hymenophoral trama subregular to irregular; hyphae 1.6-8 pm diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 1.6-4.8 pm diam, parallel, cylindrical, smooth or with sparse diverticula, hyaline, dextrinoid, thin-walled; medullary hyphae 8.8-19 pm diam, smooth, dextrinoid, thin- 32 walled. Caulocystidia numerous, scattered; 29-90 x 6-11 (base) x 1.6-2.4 (filamentous apex) pm, irregularly cylindrical to filamentous with a broader base, with apical finger- like projections, hyaline, inamyloid, thin-walled; apical projections 0.8-3.2 x <0.8 pm. Clamp connections not observed in all tissues. Habitat and known distribution - Gregarious on coconut hulls in secondary, coastal forest. (Principe). Material examined- AFRICA. Principe, Dois Irmeos area around 100 meters, N01°34.89’, E07°25.548\ 23 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 648 (Holotype, SFSU). Notes - Mycena longinqua is characterized by a tiny, glabrous, striate, greyish brown pileus, ascending-adnate lamellae, a glabrous, whitish grey stipe that arises from a small basal disc, broadly ellipsoid, amyloid basidiospores with mean 7 x 4.4 pm, 2- spored basidia, very broadly clavate, densely spinulose cheilocystidia, no pleurocystidia, dextrinoid tissues, a cutis-type pileipellis of densely spinulose hyphae with distinctively clavate terminal cells, no clamp connections, unusual caulocystidia with a swollen base and apically diverticulate filamentous appendage, and growth on coconut debris. In combination, these features indicate placement in sect. Polyadelphia. Mycena longinqua differs from all described temperate species (e.g., M. polyadelpha (Lasch) Kiihner, M. alniphila Robich, M. capillaris (Schumach.) P. Kumm.) in forming smooth (non- diverticulate) stipitipellis cortical hyphae and unusual caulocystidia with filamentous appendage (Maas Geesteranus 1986, Robich 2003). Mycena tuberifera Maas Geest. & de Meijer, described from Parana State, Brazil, differs mainly in forming larger basidiospores (8.9-10.3 x 4.5-4.9 pm) from 4-spored basidia, globose, non-diverticulate caulocystidia, and growth on leaves of Araucaria angustifolia (Maas Geesteranus and de Meijer 1997). Pairwise comparisons of aligned, overlapping ITS sequences of Mycena longinqua (BAP 648) with the top 10 BLAST results indicate 94.2% similar to M cyanorhiza Quel. (JF908385) and 93.6% similarity to M. stylobates (Pers.) P. Kumm. (JF908439). In the ITS phylogeny (Fig.2), M. longinqua belongs in a heterogeneous clade 33 with members of sects. Exornatae, Polyadelphia, Granuliferae, Longisetae and Sacchariferae.

Mycena Section Sacchariferae Kuhn, ex Singer

10. Mycena alphitophora (Berk.) Sacc., Syllog. Fung. (Abellini) 5: 290. 1887. (Figs. 19, 20) Basionym: Agaricus alphtophorus Berk., J. Linn. Soc., Bot. 15: 48. 1876. (1877). Heterotypic synonyms: Mycena osmundicola J.E. Lange, Dansk Bot. Arkiv. 1(5): 35. 1914. Refer to Desjardin (1995) for additional synonyms. Pileus 2-8 mm diam, ovoid to obtusely conical, convex or hemispherical, striate; surface covered with clusters of detersile, sugar-like granules, pure white overall. Context paper thin, translucent, white. Lamellae ascending, shallowly adnexed, subdistant with 1 series of lamellulae, narrow, non-marginate, white. Stipe 10-15 x 0.5-1 mm, central, terete, cylindrical with a slightly enlarged base, lacking a basal disc, translucent; surface dull, dry, hispid above, strigose at base, pure white. Odor and taste not recorded. Basidiospores 6.4-8.0 x 4.0^1.8 pm [xm= 7.32 ± 0.54 x 4.56 ± 0.38 pm, Q = 1.33-1.80, Qm= 1.61 ± 0.07, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 11.2-14.4 x 7.2-11.2 pm, clavate, 4-spored, sterigmata up to 3.2 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Cheilocystidia numerous, 13.6— 23 x 8.8-10.4 pm, clavate, densely spinulose over upper half, hyaline, inamyloid, thin- walled; spinulae <0.8-1.6 x <0.8 pm, cylindrical. Pleurocystidia absent. Pileipellis a cutis with acanthocysts; hyphae 4.0-8.8 pm diam, repent, cylindrical, smooth, hyaline, dextrinoid, thin-walled, non-gelatinous. Acanthocysts common, 17.6-30 x 13.6-29 pm, globose, densely spinulose, thin-walled, hyaline, inamyloid; spinulae <0.8-1.6 x <0.8- 1.6 pm. Marginal cystidia uncommon, 23-56 x 13.6-16.8 pm, fusoid to clavate, densely spinulose, hyaline, inamyloid, thin-walled; spinulae 0.8-2.4 x 0.8-1.6 pm. Hypodermium 34 composed of inflated hyphae up to 35 (am diam, globose to ovoid, hyaline, dextrinoid, thin-walled. Hymenophoral trama subregular; hyphae 1.6-7.2 jam diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 1.6-7.2 jam diam, parallel, cylindrical, smooth or rarely with knob-like diverticula, hyaline, dextrinoid, thin-walled; medullary hyphae 9.6-17.6 (am diam, smooth, hyaline, dextrinoid, thin-walled, occasional laticiferous hyphae with knob-like projections 1.6—4.8 |am diam. Caulocystidia abundant; 11-130 x 5.6-12.8 |am, cylindrical to fusoid, densely spinulose, hyaline, inamyloid, thin-walled; spinulae 0.8-2.4 x 0.8-1.6 (am. Clamp connections present in all tissues. Habitat and known distribution - Gregarious, lignicolous on undetermined dicot branches in primary forest. Widespread globally; Africa (Kenya, Sao Tome, Tanzania). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, elevation 1100 m, N00°16.554\ E06°36.326\ 11 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 591(SFSU); same location, 25 April 2006, coll. by D.E. Desjardin (material lost in transit). Notes - Mycena alphitophora is a commonly encountered, widespread member of sect. Sacchariferae. The Sao Tome specimen matches nicely material reported by Maas Geesteranus (1983), Desjardin (1995), and Pegler (1977; as M. osmundicola, from East Africa). It is characterized by small, white basidiomes with the pileus covered with sugar­ like granules, shallowly adnexed lamellae, a hispid stipe, basidiospores in the range 6.4- 10 x 4.0-5.5 (am, 4-spored basidia, clavate, spinulose cheilocystidia, no pleurocystidia, a pileipellis with numerous globose, spinulose acanthocysts, and cylindrical caulocystidia covered with spinulae. Pairwise comparisons of aligned, overlapping ITS sequences of the Sao Tome specimen (BAP 591) with the top ten BLAST results show 90-93.6% similarity to ten sequences of undetermined, uncultured fungus clones. Few sequences of members of sect. Sacchariferae are available for comparison, and no sequences of M. alphitophora from Europe are available in GenBank. A sequence determined as M. alphitophora (MH136831) from China is nearly identical to a sequence determined as M. corynephora 35

(JF908369) from Italy, and together are sister to several members of sect. Sacchariferae from ST&P, including M. alphitophora, which is on a long branch (Fig. 2).

11. Mycena discogena Singer, Ark. Bot. 4(9): 388. 1959. (Fig. 21) Pileus 3-4 mm diam, convex, striate to sulcate; surface dull, dry, glabrous, disc with brownish tones, pale ash white elsewhere. Lamellae ascending, forming a pseudocollarium, subdistant with 2 series of lamellulae, white. Stipe 20-22 x 0.5 mm, central, terete, cylindrical, arising from a small, white tomentose basal disc; surface dull, dry, minutely white-puberulous, ash white, developing brown tones towards the base. Odor and taste not recorded. Basidiospores (6.4-) 7.2-8.5 x 4.0-4.8 ^m [xm= 7.40 ± 0.44 x 4.36 ± 0.41 |j.m, Q = 1.50-2.00, Qm- 1.71 ± 0.19, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 17.6-19.2 x 6.4-8 |j.m, clavate, 4-spored, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge sterile. Cheilocystidia numerous, 19.2-26.4 x 9.6-12 (am, clavate, spinulose over upper half, hyaline, inamyloid; spinulae 0.8-1.6 x 0.8-1.6 |am. Pleurocystidia absent. Pileipellis a hymeniform layer of acanthocysts; 17.6-26 x 10.4-28 |am, globose to broadly clavate, densely spinulose, hyaline, inamyloid to weakly dextrinoid, thin-walled; spinulae 0.8 x <0.8 (am Hypodermium composed of inflated hyphae up to 33 (im diam, ovoid to globose, hyaline, dextrinoid, thin-walled. Hymenophoral trama regular to subregular; hyphae 6.4-23 jxm diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 2.4-8.8 |xm diam, parallel, cylindrical, smooth, hyaline, dextrinoid, thin-walled; medullary hyphae 13.6-23 (im diam, smooth, hyaline, dextrinoid, thin-walled, occasional laticiferous hyphae 2.4-4 (im diam with knob-like projections. Caulocystidia abundant, scattered to clustered, 25- 96 x 6.4-17 |xm, cylindrical to fusoid-acuminate, smooth, hyaline, inamyloid, thin- walled. Basal disc cystidia 36-90 x 6.5-20 fim, clavate to fusoid or cylindrical, narrowing to a acute tip, non-diverticulate, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. 36

Habitat and known distribution - Gregarious on sticks in secondary coastal forest. (Principe). Material examined- AFRICA. Principe, Dois Irmaos area, N 0r34.889’, E07°25.548’, 23 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 649 (SFSU). Notes - Mycena discogena is distinguished by a small, greyish brown to ash white, striate pileus, white, pseudocollariate, subdistant lamellae, a white, puberulous stipe that arises from a small basal disc, basidiospores with mean 7.4 x 4.4 (am, clavate, spinulose cheilocystidia, no pleurocystidia, a hymeniform pileipellis of spinulose globose acanthocysts, non-spinulose, fusoid-acuminate caulocystidia and basal disc cystidia, and growth on sticks. In combination, these features indicate placement in sect. Sacchariferae, stirps Adscendens. The species was described from material collected on the Juan Fernandez Islands, Chile, and the protologue differs from the material from Principe only in forming slightly broader basidiospores with mean 8.3 x 5.3 nm (Singer 1959, Desjardin 1995). Pairwise comparisons of aligned, overlapping ITS sequences of the Principe specimen (BAP 649) with the top ten BLAST results show 92.3-93.1% similarity to seven uncultured Mycena clones (but only 61% query coverage). Mycena discogena falls in a heterogeneous clade with members of sects. Exornatae, Polyadelphia, Granuliferae, Longisetae and Sacchariferae in the ITS phylogeny (Fig. 2).

12. Mycena lasiopus Maas Geest. & de Meijer, Persoonia 17(1): 37. 1998. (Fig. 22) Pileus up to 3mm diam, broadly convex to paraboloid, striatulate; surface minutely pruinose, covered with dark brown to black granules, background pale grey. Lamellae ascending-adnate, subdistant (10-11) with 2 series of lamellulae, white. Stipe 5-7 x 0.5 mm, central, cylindrical, covered with scattered pubescence, white, arising from a white, striate basal disc and white tomentum.

Basidiospores 8.0-9.6 (-10.5) x 4.0-5.6 |im [xmr= 9.0-9.2 x 4.5-5.0 ^m, xmm = 9.06 ± 0.1 x 4.77 ± 0.4, Q = 1.6-2.4, Qmr = 1.79-2.04, Qmm = 1.91 ± 0.2, n = 20, 21, s = 2], 37 ellipsoid, smooth, hyaline, weakly amyloid, thin-walled. Basidia 8.0-16.8 x 6.4-8.8 pm, broadly clavate, 4-spored, sterigmata up to 2.4 pm long, thin-walled, hyaline, inamyloid. Basidioles clavate. Cheilocystidia numerous, 12.8-29.6 x 5.6-10.4 (am, clavate, densely spinulose over upper half, hyaline, inamyloid; spinulae <0.8—4 x 0.8-1.6 pm, cylindrical. Pleurocystidia absent. Pileipellis a cutis with acanthocysts and cherocytes; hyphae 0.8- 4.0 pm diam, repent, cylindrical, with brownish contents, smooth, hyaline, dextrinoid, thin-walled, non-gelatinous. Acanthocysts common, 15.2-26.4 x 14.4-23.2 pm, globose, densely spinulose, thin-walled, with homogeneous brown contents, hyaline, inamyloid; spinulae <0.8-1.6 x <0.8 pm. Cherocytes uncommon, 23.2-28.8 x 13.6-21.6 pm, irregular in shape, densely spinulose, thick-walled (1.6-4.0 pm), with homogeneous brown contents, hyaline, inamyloid; spinulae <0.8-1.6 x <0.8 pm, cylindrical. Hypodermium composed of parallel hyphae up to 19.2 pm diam. Hymenophoral trama regular; hyphae 2.4-19.2 pm diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 2.4-5.6 pm diam, parallel, cylindrical, hyaline, dextrinoid, thin-walled; medullary hyphae 7.2-17.6 pm diam, smooth, hyaline, dextrinoid, thin-walled, occasional laticiferous hyphae with knob-like projections 2.4-3.2 pm long. Caulocystidia abundant; 23.2-600 x 5.6-10.4 pm, cylindrical for most of length, gradually narrowing toward the broadly rounded apex, densely spinulose at base with less spinulae at apex, hyaline, inamyloid, thin-walled; spinulae <0.8-2.4 x <0.8 pm, cylindrical. Basal disc with cherocytes and acanthocysts as on pileus surface and cystidia like the caulocystidia. Clamp connections present in all tissues. Habitat and known distribution - Gregarious on woody debris in forest with cacao or on sticks in secondary, upland forest, Africa (Principe, Sao Tome). Material examined- AFRICA. Principe, along trail to Ro?a Pico Papagaio, N01°37.182’, E07°23.474’, 21 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 635 (SFSU). Sao Tome, along main road (EN2) on south side of island approx. 200 m east of 38 km marker, N00°08.500’, E06°39.560’, 13 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 603 (SFSU). 38

Notes - Mycena lasiopus, described from Parana State, Brazil, is distinguished by tiny basidiomes with grey pileus covered with dark brown granules, subdistant white lamellae, a white, pubescent stipe arising from a striate basal disc, basidiospores with mean 9 x 4.8 jam (Q = 1.9), densely spinulose, clavate cheilocystidia, a pileipellis with brown, densely spinulose, globose, thin-walled acanthocysts and irregular-shaped, thick- walled cherocytes, very long, cylindrical, densely spinulose caulocystidia, and growth on woody debris. The specimens from Sao Tome and Principe are indistinguishable from the original material from Brazil (Maas Geesteranus and de Meijer 1998). The species belongs in sect. Sacchariferae, stirps Amparoina sensu Desjardin (1995). Pairwise, aligned sequences of M. lasiopus (BAP 603, BAP 635) with the top 10 BLAST results show 88.3-89.2% similarity to M. corynephora Maas Geest. (JF908366, JF908367, JF908368, JF908369), a north temperate member of sect. Sacchariferae, stirps Alphitophora. Unfortunately, few sequences of members of sect. Saccharifera are available for comparison. Mycena corynephora differs in forming pure white pileus lacking cherocytes, a stipe that lacks a basal disc, and globose to subglobose basidiospores (Q = 1.0-1.1) (Maas Geesteranus 1983). In the ITS phylogeny (Fig. 2), M. lasiopus clusters with two additional members of sect. Sacchariferae from ST&P (M. oboensis, M. alphitophora) in a clade with other members of the section (92% BS, 1.0 PP support).

13. Mycena oboensis A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Fig. 23) MycoBank no.: MB 825515 Holotype:—AFRICA. Sao Tome, Parque Nacional Obo, Lagoa Amelia Trail, N00°16.922’, E06°36.062’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 669 (SFSU). Etymology:—referring to the site where the holotype was collected, Parque Nacional Obo. 39

Diagnosis - Pileus 1.5-3 mm diam, initially paraboloid, becoming plane with a central depression, striate; surface dull, dry, minutely granulose, disc with black pruinae, less pruinose towards the margin, white. Lamellae adnate with a short decurrent tooth, subdistant with 1 series of lamellulae, broad, white. Stipe 8-14 x <0.5 mm, central, terete, cylindrical, arising from a small, pubescent basal disc; surface dull, dry, pubescent, white. Odor and taste not recorded. Basidiospores (6.4-) 7.2-8.8 x (4.0-) 4.8-5.6 |am [xm= 7.92 ± 0.73 x 4.84 ± 0.41 Hm, Q = 1.33-1.83, Qm= 1.64 ± 0.12, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin-walled. Basidia 11.2-16.8 x 7.2-10.4 |am, clavate, 2-spored, hyaline, inamyloid, thin-walled. Basidioles clavate. Cheilocystidia absent. Pleurocystidia absent. Pileipellis a cutis with acanthocysts and cherocytes; hyphae 2.4-4.0 (xm diam, repent, cylindrical, smooth, hyaline, dextrinoid, thin-walled, non-gelatinous. Cherocytes scattered, 21-51 x 18-39 fxm, globose to ovoid or pyriform, with knob-like projections over the upper half, apically thick-walled (3-10 (-23) ^im), walls hyaline, contents greyish brown; projections 1.5-5 x 1.5-3.2 (am, solid. Acanthocysts common, 15.2-39 x 11.2-24 |j.m, globose, densely spinulose, thin-walled, hyaline, inamyloid; spinulae <0.8-1.6 x <0.8 (am. Marginal cystidia common, 16-53 x 10.4-14.4 nm, clavate to fusoid, densely spinulose on the upper half or apex, thin-walled, hyaline, inamyloid; spinulae <0.8-1.6 x <0.8 |im. Hypodermium composed of inflated hyphae up to 21 ^m diam, hyaline, inamyloid, thin- walled. Hymenophoral trama hyphae 2.4-8.8 |im diam, hyaline, dextrinoid, non- gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 2.4-7.2 |xm diam, parallel, cylindrical, hyaline, dextrinoid, thin-walled; medullary hyphae 9.6-29 (am diam, smooth, hyaline, dextrinoid, thin-walled. Caulocystidia abundant; 23-79 x 5.6-10.4 (am, cylindrical to fusoid, densely spinulose, hyaline, inamyloid, thin-walled; spinulae <1.6- 2.4 x <0.8-1.6 |am. Clamp connections present in all tissues. Habitat and known distribution - Gregarious on decaying seed in upland forest. (Sao Tome). 40

Material examined- AFRICA. Sao Tome, Parque Nacional Obo, Lagoa Amelia Trail, N00° 16.922’, E06°36.062’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 669 (Holotype, SFSU). Notes - Mycena oboensis is distinguished by tiny basidiomes with a white, striate pileus with scattered black pruinae, white, adnate lamellae with a decurrent tooth, a white, pubescent stipe arising from a small basal disc, basidiospores with mean 7.9 x 4.8 (im (Q = 1.6), 2-spored basidia, an absence of cheilocystidia and pleurocystidia, pileipellis with acanthocysts, very thick-walled cherocytes with brown contents and knobby projections, distinctive marginal cystidia, and cylindrical caulocystidia covered with spinulae. The morphology of Mycena oboensis is nearly indistinguishable from M. lasiopus, but the latter has a more grey pileus, shorter stipe (up to 7 mm), grows on sticks, has longer basidiospores with mean 9 x 4.8 pm (Q = 1.9), spinulose cheilocystidia, irregular-shaped cherocytes with tiny spinulae, lacks distinctive marginal cystidia, and has much longer caulocystidia with apically more sparse spinulae. Pairwise comparisons of aligned, overlapping ITS sequences of the Sao Tome specimen (BAP 669) with the top ten BLAST results show 88.2% similarity to four sequences determined as M. corynephora, a European species in sect. Sacchariferae. Interestingly, the ITS sequence of Mycena oboensis is only 71.2% similar to that of M. lasiopus because of multiple insertions in M. oboensis. In the ITS phylogeny (Fig. 2), M. oboensis clusters with two additional members of sect. Sacchariferae from ST&P (M lasiopus, M. alphitophora) in a clade with other members of the section (92% BS, 1.0 PP support). 41

Mycena section uncertain

14. Mycena lamprospora (Comer) E. Horak, Ber. Schweiz. Bot. Ges. 88(12): 22.1978 (Figs. 24, 25) Basionym: Mycena rorida var. lamprospora Comer, Mycologia 42(3): 427. 1950. = Roridomyces lamprosporus (Corner) Rexer, Die Gattung Mycena s.L, Studien zu Ihrer Anatomie, Morphologie und Systematik (Tubingen): 140. 1994. Pileus 4-6 mm diam, hemispherical with a flattened disc, margin undulate, shallowly striate; surface moist, disc appressed fibrillose with short brown fibrils and pale brown tones, glabrous and off-white at margin. Lamellae decurrent, subdistant (14-15) with 1-2 series of lamellulae, white, edges concolorous. Stipe 15-23 x 0.5-1 mm, central, terete, cylindrical; surface with a thick, hyaline, glutinous layer, apex with minute white granules, pearlescent, greyish white, base with yellowish brown tones. Odor none; taste not recorded. Basidiospores 6.4-8.8 x 3.2-4.8 fxm [xm = 7.48 ± 0.66 x 3.84 ± 0.56 |am, Q = 1.60-2.50, Qm= 1.98 ± 0.38, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 8.8-20.8 x 4-7.2 (im, clavate, 4-spored, sterigmata up to 3.2 |am long, hyaline, inamyloid, thin-walled. Basidioles clavate to lageniform. Lamellar edge sterile. Cheilocystidia of 2 types: 1) 27.2—40.8 x 8.8-10.4 (a.m, clavate to subclavate, hyaline, inamyloid, thin-walled; 2) 18.4-25.6 x 5.6-6.4 (im, clavate to subclavate with a few irregular apical projections, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis an interrupted cutis to trichoderm with irregular or sphaeropedunculate terminal cells; hyphae 2.5-5 |im diam, cylindrical or irregular, smooth or with scattered projections, hyaline or with brown contents, inamyloid, thin-walled. Terminal cells of 2 types: 1) 20-60 x 4-6 |j.m, irregularly cylindrical with scattered projections, with brown contents, inamyloid, thin-walled; 2) 24—44 x 8.8-16.8 (am, sphaeropedunculate, hyaline, inamyloid, thin-walled. Hypodermium composed of inflated cells up to 25 (am diam, globose to ovoid, smooth, hyaline, dextrinoid, thin-walled. Hymenophoral trama subregular; hyphae 2.4-5.6 (im diam, hyaline, dextrinoid, non-gelatinous, thin-walled. 42

Stipitipellis a cutis with erect caulocystidia; cortical hyphae 1.6—4.8 pm diam, parallel, cylindrical, hyaline, dextrinoid, thin-walled; medullary hyphae 5.6-15.2 pm diam, smooth, dextrinoid, thin-walled; terminal cells diverticulate. Caulocystidia abundant; 31- 42 x 6.4-9.6 pm, clavate or subcapitate, with occasional scattered projections, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Gregarious, inserted on leaves in secondary upland forest. (Principe). Material examined- AFRICA. Principe, Ro?a Pico Papagaio, N 0r37.182’, E07°23.474\ 20 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 634 (SFSU). Notes - The morphology of the Principe specimen matches quite closely that of the protologue of Mycena rorida var. lamprospora (Corner 1950), based on material collected in Malaysia, with one exception. The Malaysian type is reported to have a palisade-type pileipellis of clavate-pyriform-pedicellate cells, whereas the West African material has an interrupted cutis to trichodermium with erect terminal cells similar to those described from the holotype. Our material is also nearly indistinguishable from Horak’s (1978) description of M. lamprospora as represented in the southern hemisphere, differing only in pileipellis anatomy. Because of the reported hymeniform pileipellis in M. lamprospora, the species was transferred to the genus Roridomyces, based on R. rorida (Fr.) Rexer, a genus distinguished from Mycena by hymeniform versus cutis-type pileipellis, respectively. Both Comer (1950) and Horak (1978) reported the spores as bioluminescent, a feature not observed in the Principe specimen. Unfortunately, the sequence data (ITS, LSU) indicate that what we have determined as matching M. lamprospora is distantly related to R. rorida, suggesting that M. lamprospora does not belong in Roridomyces. Alternatively, the Principe specimen, which is morphologically indistinguishable from M. lamprospora, does not represent M. lamprospora. We were able to acquire only 150 base pairs of the ITS region of Mycena lamprospora (BAP 634) and consequently we did not include it in the ITS phylogeny. The nLSU sequence of M. lamprospora is 92.3% similar with R. rorida, and falls into a poorly supported clade with Panellus stypticus, Cruentomycena viscidocruenta and 43

Roridomyces rorida. Mycena lamprospora was not sister to Roridomyces rorida in the tree, indicating that they may not be congeneric.

Clitocybula (Singer) Singer ex Metrod

15. Clitocybula intervenosa A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Figs. 26, 27) MycoBank no.: MB 825516 Holotype:—AFRICA, Sao Tome, Parque Nacional Obo, Lagoa Amelia Trail between N00°17.112’, E06°35.967’ and N00° 16.922’, E06°36.062’, 14 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 613 (SFSU). Etymology:—intervenosa - referring to the strongly intervenose lamellae. Diagnosis - Pileus 6-25 mm diam, broadly convex to campanulate, umbilicate or with a central depression, margin striate, becoming upturned, flared and eroding in age; surface glabrous, moist, watery brownish grey (7-8C-D3), hygrophanous, becoming pallid grey overall with moisture loss. Context thin, concolorous with pileus. Lamellae decurrent to deeply decurrent, close to subdistant (12-21) with 1-2 series of lamellulae, highly intervenose to forked, narrow, pale grey. Stipe 8-30 x 1-2.5 mm, central, terete or flattened, cylindrical or broadened towards base, occasional tapered in mid-section, solid; surface smooth, glabrous, pale grey, some with pale brown tones towards base, base with or without white . Odor none. Taste indistinct. Basidiospores 5.6-7.2 (-8.8) x (3.2-) 4.0-4.8 (-5.6) fxm [xmr= 6.5-6.8 x 4.4^1.5

|xm, X m m = 6.65 ± 0.2 x 4.47 ± 0.1, Q = 1.2-2.3, Qmr= 1.48-1.52, Qmm= 1.50 ± 0, n = 19- 20, s = 2], ellipsoid to lacrymoid, smooth, hyaline, amyloid, thin-walled. Basidia 21.6- 38.4 x 5.6-7.2 |xm, clavate, 4-spored, sterigmata up to 5.6 |xm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge fertile. Cheilocystidia absent or undifferentiated. Pleurocystidia absent. Pileipellis a cutis; hyphae 0.8-12.0 (am diam, repent, cylindrical, non-diverticulate, hyaline, inamyloid, thin-walled, non-gelatinous. Hypodermium composed of inflated cells up to 16.0 |_im diam, smooth, hyaline, 44 inamyloid to weakly dextrinoid, thin-walled, rare to numerous laticiferous hyphae up to 4.0 pm diam with knob-like projections 4.8-10.4 x 2.4-3.2 pm. Hymenophoral trama subregular; hyphae 2.0-12.8 pm diam, hyaline, weakly dextrinoid, non-gelatinous, thin- walled. Stipitipellis a cutis; cortical hyphae 2.4-16.0 pm diam, parallel, cylindrical, hyaline, dextrinoid, thin-walled; medullary hyphae 2.4-16.8 pm diam, smooth, dextrinoid, thin-walled, occasional laticiferous hyphae up to 6.4 pm diam with knob-like projections. Caulocystidia abundant; 8.0-36.0 x 6.4-15.2 pm, versiform, fusoid-clavate or irregular in outline, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Densely cespitose on decaying in primary forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Bom Sucesso Botanical Garden, N00°17.317\ E06°36.746’, 11 April 2008, coll. by B.A. Perry, BAP 588 (SFSU). Sao Tome, Parque Nacional Obo, Lagoa Amelia Trail between N00°17.112’, E06°35.967’ and N00'T6.922’, E06°36.062’, 14 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 613 (Holotype, SFSU). Notes - Clitocybula intervenosa is characterized by convex-umbilicate, striate, pale brownish grey pileus, pale grey, decurrent, strongly intervenose lamellae, amyloid, ellipsoid basidiospores with mean 6.6 x 4.5 pm, an absence of cheilocystidia, small, versiform caulocystidia, and cespitose habit on decaying hardwood in primary forest. Clitocybula intervenosa shows some similarities to C. omphaliformis Pegler, described from Tanzania, but the latter forms more crowded, non-intervenose lamellae, smaller subglobose basidiospores (mean 5 x 3.5 pm), cylindrical to lageniform cheilocystidia, and larger caulocystidia (Pegler 1977). Clitocybula grisella (G. Stev. & G.M. Taylor) E. Horak, described from New Zealand, differs in forming adnate to sinuate, non-intervenose lamellae, subglobose basidiospores 4.5-7.0 pm diam, cylindrical cheilocystidia, relatively large pileocystidia, and growth on wood of Nothofagus (Stevenson 1965). Clitocybula lenta (Maire) Malen?on & Bertault, described from 45

Mauretania, has a dirty brown, fibrillose-rimose pileus, reminiscent of an Inocybe, non- intervenose lamellae, and clavate to fusiform cheilocystidia (Maire 1928). Pairwise comparisons of aligned, overlapping ITS sequences of Clitocybula intervenosa (BAP 588, BAP 613) with the top 10 BLAST results show 95.9% similarity to two sequences determined as Clitocybula aff. lacerata (KY242502, FJ596916). Clitocybula lacerata (Scop.) Metrod, a common north temperate species, shares size and lack of cheilocystidia, but differs from C. intervenosa in forming larger basidiomes with a non-striate, radially fibrillose pileus 20-60 mm diam, close, non-intervenose lamellae, and cylindrical to clavate pileocystidia (Bigelow 1973). Clitocybula intervenosa is sister to C. lacerata in the ITS phylogeny (Fig. 2) with low support.

Filoboletus Henn.

16. Filoboletus pallescens (Boedijn) Maas Geest., Proc. K. Ned. Akad. Wet., Ser. C, Biol. Med. Sci. 95(2): 269. 1992. (Fig. 28,29) Basionym: Poromycenapallescens Boedijn, Bull. Jard. Bot. Buitenz., 3 Ser. 16(4): 398. 1940. Pileus 7-20 mm diam, initially parabolic, becoming broadly campanulate to broadly obtusely conical or convex, margins inrolled, pellucid-striate to short-striatulate, pustulate-tesselate; surface moist, glabrous, hygrophanous, disc brown (6-7E4-5) or brownish grey (6-7C-D-3), margin grey (7D2), fading overall with moisture loss to brownish grey (6-7C-D-3). Context 1 mm thick, watery-white to grey. Lamellae ascending-adnate to adnexed, horizontal, distant (15-18) with 2-3 series of lamellulae, broad (~3 mm), reticulate to subporoid, pores round to angular, cross lamellae much shallower, pale greyish white with brown tones in upper portion. Stipe 21—40 x 2—4.5 mm, central, cylindrical, terete or flattened, base enlarged, hollow, pliant; surface dull, 46 dry, appressed-pubescent to minutely white-pruinose overall, ornamentation very dense towards base, white to pale greyish white overall. Odor none. Taste indistinct. Basidiospores 6.4-8.0 x 4.8-6.4 pm [xmr= 7.0-7.7 x 5.2-5.5 pm, xmm= 7.34 ± 0.5 x 5.38 ± 0.2, Q = 1.1-1.7, Qmr= 1.28-1.47, Qmm= 1.37 ± 0.1, n = 20, s = 2], subglobose to broadly ellipsoid, smooth, hyaline, amyloid, thin-walled. Basidia 25.6-34.4 x 4.8-7.2 pm, clavate, 4-spored, sterigmata up to 10.4 (am long, thin-walled, hyaline, inamyloid. Basidioles clavate. Lamellar edge sterile. Cheilocystidia 30-102 x 6.4-15.2 (am, fusoid to subclavate, occasionally fusoid-strangulate, rarely with a single apical projection, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Pileipellis a cutis; hyphae 2.4-9.6 (am diam, repent, cylindrical, rarely diverticulate, hyaline or with a few brown incrustations, inamyloid or weakly dextrinoid, thin-walled, non-gelatinous; diverticula 1.6-7.2 x 0.8-3.2 (am, knob-like. Hypodermium undifferentiated. Pileus trama loosely interwoven; hyphae 3-20 (am diam, non-diverticulate, hyaline, weakly dextrinoid, thin-walled, non-gelatinous. Lamellar trama regular; hyphae 2.4-24 pm diam, cylindrical to inflated, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 2.4-5.6 pm diam, parallel, cylindrical, occasional cylindrical diverticula, hyaline, dextrinoid, thin-walled; medullary hyphae 8.8-26.4 pm diam, smooth, dextrinoid, thin- walled, rare laticiferous hyphae up to 8.8 pm diam with knob-like projections. Caulocystidia abundant; 32.8-105 x 6.4-12.0 pm, fusoid-clavate with obtuse diverticula, hyaline, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Gregarious to scattered or solitary on fallen rotten dicot wood in coastal palm grove. (Principe). Material examined- AFRICA. Principe, Bom Bom, N 0r41.559’, E07°24.17r, 22 April 2008, coll. by D.E. Desjardin, DED 8303 (SFSU). Principe, Dois Irmeos , N 0r35.728’, E07°25.263’, 23 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 654 (SFSU). Notes - Filoboletus pallescens is characterized by a pustulate-tesselate, brownish grey pileus, subporoid lamellae, a white to pale greyish white, pruinose stipe, subglobose to broadly ellipsoid basidiospores with mean 7.3 x 5.4 pm, fusoid to subclavate 47 cheilocystidia, no pleurocystidia, a cutis-type pileipellis with none to few diverticula, and numerous fusoid-clavate caulocystidia. The protologue indicates that the pileus is campanulate and in age convex often with a small umbo, pale grey when fresh and white when dried. The material from Principe lacks an umbo and is more deeply pigmented, otherwise the basidiomes are similar. The micromorphology reported by Boedijn (1940) matches nicely with our material. Maas Geesteranus (1992) reported that the hymenophore of F. pallescens was luminous. The material from Principe was not luminescent. This species is very closely allied with F. manipularis (Berk.) Singer, a strongly bioluminescent taxon widely distributed in East Asia. The latter differs, however, in forming cheilocystidia with diverticulate projections, and a pileipellis with more densely diverticulate hyphae (Corner 1954, Pegler 1986). Filoboletus lachiwalensis Maas Geest., described from India, differs in forming more crowded (16-30), decurrent lamellae, inamyloid lamellar trama, cheilocystidia with numerous apical diverticula, and gelatinized pileipellis and stipitipellis (Maas Geesteranus 1992b). The New World F. gracilis (Klotzsch ex Berk.) Singer differs in forming light pinkish cinnamon, conical pileus, much narrower lamellar tubes (0.5-1 mm), utriform to lecythiform, subcapitate cheilocystidia, a well-developed hypodermium, and smaller, oblong-ellipsoid to pyriform caulocystidia (Pegler 1983). Pairwise comparisons of aligned, overlapping ITS sequences of Filoboletus pallescens (BAP 654, DED 8303) with the top 10 BLAST results show 93.6-94.0% similarity to six sequences of F. manipularis from Vietnam, and is sister to F. manipularis in the ITS phylogeny (Fig. 2) with 90% BS and 0.93 PP support. 48

Hydropus Kuhner ex Singer

17. Hydropus globosporus A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Figs. 30, 31) MycoBank no.: MB 825517 Holotype:—AFRICA. Sao Tome, Macambrara radio antenna area, above 1300 meters, N00°16.557’, E06°36.326’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 661 (SFSU). Etymology:—referring to the globose basidiospores. Diagnosis - Pileus 5-11 mm diam, hemispherical with flattened apex to convex, centrally depressed to umbilicate, striate to sulcate; surface glabrous, moist, brown to dark brown (6E-F4-5) when young, fading to greyish brown (4-5E4-5) in age, central depression remaining dark. Context thin, white. Lamellae adnate with decurrent tooth to decurrent, distant (9-11) with 1-2 series of lamellulae, broad (up to 4 mm), non- intervenose, white, edges concolorous. Stipe 6-10 x 0.75-1.5 mm, central, terete, tapered towards base, hollow; surface minutely pruinose, with white pruinae at apex and dark brown pruinae below, otherwise white, developing greyish brown tones towards the base. Basidiomes not blackening. Odor indistinct; taste not recorded. Basidiospores (4.8-) 5.6-7.2 (-8.0) x (4.8-) 5.6-6.4 |im [xm= 6.04 ± 0.73 x 5.67 ± 0.42 (im, Q = 0.86-1.25, Qm= 1.06 ± 0.07, n = 25, s = 1], globose to subglobose with a prominent apiculus, smooth, hyaline, inamyloid, thin-walled. Basidia 18.4-21 x 4.8-5.6 (im, clavate, 2-spored, rarely 3-spored, sterigmata up to 7.2 |im long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge sterile. Cheilocystidia 42-66 x 8.4-10 (im, subclavate to cylindrical, hyaline, inamyloid, thin-walled. Pleurocystidia common, 30-40 (-62) x 6.5-9 (im, similar to cheilocystidia. Pileipellis a cutis; hyphae 4-10 (im diam, cylindrical to inflated, smooth, hyaline or with contents brown, inamyloid, thin-walled; terminal cells undifferentiated of seldom clavate, repent to suberect. Hypodermium undifferentiated. Pileus trama hyphae 6-28 |iim diam, cylindrical to inflated, with brown contents, inamyloid, thin-walled. Hymenophoral trama regular to subregular; hyphae 3- 20 (-32) (im diam, cylindrical to inflated, hyaline, inamyloid, non-gelatinous, thin- 49 walled. Stipitipellis a cutis with caulocystidia; cortical hyphae 3.2-4.8 (am diam, parallel, cylindrical, smooth, hyaline, inamyloid, thin-walled; medullary hyphae 7.2-18 |am diam, smooth, inamyloid to weakly dextrinoid, thin-walled. Caulocystidia abundant; 8-38 x 6.4-13 |am, cylindrical to subclavate or geniculate, few intercalary, hyaline or occasionally with brown contents, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Gregarious on decaying wood in secondary upland forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, above 1300 meters, N00° 16.557’, E06°36.326’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 661 (Holotype, SFSU). Notes - Hydropus globosporus is distinguished by small basidiomes with brown, sulcate, convex-umbilicate pileus, distant, broad, subdecurrent white lamellae, a white, brown-pruinose stipe, inamyloid, globose to subglobose basidiospores with mean 6 x 5.7 (am, 2-spored basidia, subclavate to cylindrical cheilocystidia and caulocystidia, a cutis- type pileipellis of smooth hyphae with brown cytoplasmic contents, cylindrical to geniculate, hyaline or brown caulocystidia, and lignicolous habit. In combination, these features indicate placement in sect. Floccipedes subsect. Floccipedes. Within this infrageneric group, the globose basidiospores suggest our Sao Tome material is close to H. floccipes (Fr.) Singer, however the latter species forms campanulate-subumbonate to convex-papillate pileus, adnate to adnexed, close to crowded lamellae, a much larger stipe, 4-spored basidia, more ampullaceous hymenial cystidia (20-100 x 7-15 (am), and differentiated pileipellis terminal cells (Singer 1982, Laessoe 2008). Comer (1966) accepted numerous Hydropus in the genus Trogia. If keyed there, Hydropus globosporus comes closest to Trogia mycenoides Comer, but the latter forms a conical pileus, adnexed, subclose (17-22) lamellae, ellipsoid basidiospores (6-8 x 4-5 |am), 4-spored basidia, and ventricose hymenial cystidia (Comer 1966). Pairwise comparisons of aligned, overlapping ITS sequences of Hydropus globosporus (BAP 661) with the top 10 BLAST results indicate 87.4-89.2% similarity to 50 several undetermined Agaricales. The closest determined species was Hydropus funebris (Speg.) Singer, with 87.1% similarity, and the two species are sister in the ITS phylogeny (Fig. 2) with high support (100% BS, 1.0 PP).

18. Hydropus murinus A.C. Cooper, Desjardin & B.A. Perry, sp. nov. (Figs. 32, 33) MycoBank no.: MB 825518 Holotype:—AFRICA. Sao Tome, Macambrara radio antenna area, above 1300 meters, N00°16.557’, E06°36.326\ 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 657 (SFSU). Etymology:—murinus = mouse grey - referring to the mouse gray color and mouse hair-like texture of the pileus surface. Diagnosis - Pileus 4-11 mm diam, broadly paraboloid to convex when young, becoming plano-convex and centrally depressed to umbilicate in age, margin entire, striatulate; surface dull, dry, minutely pruinose with dark brown granules and very short hairs, disc dark brown (6-7F7-8), fading to light brown, margin greyish brown, fading to greyish white. Lamellae decurrent, close to subdistant with 2 series of lamellulae, off- white, developing brown tones when drying, edges concolorous. Stipe 7-16 x 0.5-1 mm, central, terete, cylindrical, hollow; surface minutely pruinose, with dark brown granules and short brown hairs, greyish white to pale yellowish brown, darkening in age from base upwards. Odor none; taste not recorded. Basidiospores 7.2-8.0 x 4.0-4.8 pm [xm- 7.72 ± 0.39 x 4.44 ± 0.41 pm, Q = 1.50-2.00, Qm= 1.75 ± 0.12, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 17.6-35 x 5.6-7.2 pm, clavate, 4-spored, sterigmata up to 5.6 pm long, hyaline, inamyloid, thin-walled. Basidioles clavate. Lamellar edge sterile. Cheilocystidia 42-81 x 4.8-10.4 pm, fusoid to subclavate or cylindrical, occasionally strangulate, hyaline, inamyloid, thin-walled. Pleurocystidia uncommon, narrowly fusoid, similar to cheilocystidia. Pileipellis a cutis with differentiated terminal cells; hyphae 7.2-10.4 pm diam, cylindrical, smooth, hyaline or with brown contents, inamyloid, thin-walled. 51

Terminal cells repent to erect, 23-55 x 8.8-14.4 pm, subclavate to fusoid, with brown contents, inamyloid, thin-walled. Hypodermium absent. Hymenophoral trama regular to subregular; hyphae 4-18 pm diam, hyaline, inamyloid, non-gelatinous, thin-walled. Stipitipellis a cutis with caulocystidia; cortical hyphae 3.2-9.6 pm diam, parallel, cylindrical, hyaline, inamyloid, thin-walled; medullary hyphae 5.6-11.2 pm diam, smooth, inamyloid to weakly dextrinoid, thin-walled. Caulocystidia abundant; 26-67 x 5.6-13 pm, fusoid to cylindrical, occasionally geniculate, hyaline or occasionally with brown contents, inamyloid, thin-walled. Clamp connections present in all tissues. Habitat and known distribution - Gregarious on decaying wood in secondary upland forest. (Sao Tome). Material examined- AFRICA. Sao Tome, Macambrara radio antenna area, above 1300 meters, N00° 16.557’, E06°36.326’, 25 April 2008, coll. by D.E. Desjardin and B.A. Perry, BAP 657 (Holotype, SFSU). Notes - Hydropus murinus is distinguished by convex-umbilicate, pruinose, dark brown to grey pileus 4-11 mm diam, decurrent, off-white lamellae, a hollow, pruinose, greyish white stipe 7-16 x 0.5-1 mm, amyloid basidiospores with mean 7.7 x 4.4 pm, 4- spored basidia, fusoid to cylindrical cheilocystidia and pleurocystidia, a cutis-type pileipellis with numerous subclavate to fusoid, brown terminal cells, fusoid to cylindrical caulocystidia, and lignicolous habit. In combination, these features indicate placement in sect. Hydropus, subsect. Marginelli Singer (1982). The Sao Tome taxon is morphologically closest to H. cavipes (Pat. & Gallard) Dennis and Hydropus atropruinosus (Corner) Singer. Hydropus cavipes, described from Venezuela, forms larger basidiomes with olive brown, translucent striate pileus 15-70 mm diam, adnate lamellae, a stipe 17-60 x 2-8 mm, smaller basidiospores with mean 6 x 4 pm, and broader hymenial cystidia (up to 18 pm diam) (Singer 1982, Pegler 1983). Hydropus atropruinosus, described from Brazil, forms much smaller basidiospores (5.5- 6.3 x 2.7-3 pm, and ventricose to clavate cheilocystidia with a narrow apical prolongation up to 35 pm long (Comer 1966, Singer 1982). 52

Pairwise comparisons of aligned, overlapping ITS sequences of Hydropus murinus (BAP 657) with the top 10 BLAST results indicate 99.7% similarity to two sequences from Australia determined as Hydropus sp. (KPO12822, KP012912). These results suggest that there occurs a species in Australia that is conspecific with our material from Sao Tome. Cooke (1892) and others (cited in May and Wood 1997) reported Hydropus putredinus (Berk. & M.A. Curtis) Pegler (as Marasmius putredinus) from Australia. Pegler (1987), who studied the holotype specimen of M putredinus, a species described from Cuba, noted the close similarity of H. putredinus with H. cavipes, but indicated that H putredinus had inamyloid basidiospores. Hydropus putredinus differs from Hydropus murinus in forming larger basidiomes (pileus 12-25 mm diam) with a solid stipe and smaller inamyloid basidiospores (mean 6 x 3.7 pm) (Pegler 1987). In the ITS phylogeny, H. murinus is sister to H. marginellus plus H atramentosus with high support (100% BS, 1.0 PP). 53

19. “Trogia delicata” Comer, Gdns. Bull. Singapore. Suppl. 2: 26. 1991. (Fig. 34) Pileus 22-38 mm diam, obtusely conical to piano-conical with flattened disc, sulcate-plicate to disc; surface dull, dry, silky-glabrous, translucent, white with disc and central half of striate greyish turquoise (23E5-6). Context very thin. Lamellae ascending, free, subdistant with no lamellulae or a few scattered short lamellulae, narrow (2-2.5 mm), greyish turquoise near the stipe, white at margin. Stipe 35-55 x 2-3 mm, central, terete, cylindrical with a slightly enlarged base, fragile, translucent, hollow; surface dull, dry, apex pruinose, base with a few annular granulose-pruinose zones, white overall. Odor none; taste indistinct. Basidiospores 6.4-8.0 x 3.2-4.8 (^m [xm= 7.08 ± 0.39 x 4.04 ± 0.48 |^m, Q = 1.50-2.25, Qm= 1.78 ± 0.29, n = 20, s = 1], ellipsoid, smooth, hyaline, amyloid, thin- walled. Basidia 11.2-23 x 7.2-8.8 (am, clavate, 4-spored and 2-spored, sterigmata up to 4 |xm long, hyaline, inamyloid, thin-walled, clamped. Basidioles clavate. Lamellar edge sterile. Cheilocystidia clavate, undifferentiated from basidioles. Pleurocystidia absent. Pileipellis a cutis; hyphae 3.2-13 |^m diam, repent, cylindrical, non-diverticulate, hyaline, inamyloid, thin-walled, non-gelatinous. Terminal cells uncommon, 21-57 x 6.4-9.6 fim, cylindrical with occasional projections 2.4^1 x 2.4-3.2 (xm, hyaline, inamyloid, thin- walled. Hypodermium composed of inflated cells up to 56 |^m diam, smooth, hyaline, inamyloid to weakly dextrinoid, thin-walled, numerous laticiferous hyphae up to 3.2 |j.m diam with knob-like projections. Hymenophoral trama irregular; hyphae 3.2-20 |im diam, hyaline, dextrinoid, non-gelatinous, thin-walled. Stipitipellis a cutis; cortical hyphae 3.2-8.8 |nm diam, parallel, cylindrical, smooth, hyaline, inamyloid, thin-walled; medullary hyphae 4.8-11.2 (im diam, smooth, dextrinoid, thin-walled, numerous branching laticiferous hyphae up to 13 (am diam with occasional knob-like projections. Caulocystidia scattered, 26-33 x 7.2-8.8 ^m, clavate to irregularly cylindrical, with occasional projections, hyaline, inamyloid, thin-walled. Clamp connections present but inconstant in all tissues. Habitat and known distribution - Solitary on rotten wood in primary forest. (Sao Tome). 54

Material examined- AFRICA. Sao Tome, along trail between Bom Sucesso and Lagoa Amelia at 1300 meters, N00°17.317’, E06°36.746’, 14 April 2008, coll. by D.E. Desjardin and B.A. Perry, DED 8235 (SFSU). Notes - The material from Sao Tome was rather old and in fair condition but matches nicely with the protologue of Trogia delicata (Corner 1991), described from Borneo, with the exception of basidiospores being broader and tending towards subglobose in the Borneo material. The species is delicate, with paper-thin tissues, characterized by a white pileus with greyish turquoise disc (in Sao Tome specimen), free lamellae, a white, fragile stipe, 2- and 4-spored basidia, undifferentiated cheilocystidia, a cutis-type pileipellis, clavate caulocystidia, clamp connections, and solitary growth on wood. The amyloid basidiospores, dextrinoid tissues, plicate pileus reminiscent of a Coprinus, free lamellae and inconstant clamp connections indicate placement in Mycena sect. Radiatae Singer (Desjardin and Hemmes 2001). A formal transfer of “T. delicata” to Mycena, which we believe is inappropriate, would require establishment of a new name (non Mycena delicata Aravind. & Manim.) and examination of more material from Borneo. Molecular data indicate that selected members of Mycena sect. Radiatae do not belong in the genus Mycena, but rather belong to the hydropoid clade. It is premature to establish a new genus for these taxa until more phenetically similar species are included in molecular analyses. Pairwise comparisons of aligned, overlapping ITS sequences of “Trogia delicata" (DED 8235) with the top 10 BLAST results show 95.8-96.3% similarity to three sequences of Clitocybula lacerata. Unfortunately, the quality of the sequence of Sao Tome material was poor. A molecular phylogeny of the Agaricales (Matheny et al. 2006) places Clitocybula and Mycena sect. Radiatae (represented by M. auricoma Takahashi) in the hydropoid clade. In the ITS phylogeny, “Trogia delicata” is on a very long branch embedded amongst several Clitocybula species and distant from a second member of Mycena sect. Radiatae (M auricoma). 55

Excluded Species

The following species of Mycena reported from Sao Tome and Principe represent excluded taxa.

Mycena tintinnabulum (Paulet) Quel. Bresadola and Roumeguere (1890) reported Mycena tintinnabulum from Sao Tome, without description. We have not encountered this species in ST&P.

Mycena rosea (Bull.) Gramberg. Coutinho (1925) reported Mycena rosea from Sao Tome (as Agaricus {Mycena) roseus). The species, a member of sect. Calodontes, was reported to have a lilac-rose pileus and venose-reticulate lamellae. We have not encountered this species in ST&P.

Agaricus (Mycena) rufescens Cout. [nom. illeg., non Agaricus rufescens. J.F. Gmel. 1792.] Agaricus rufescens was described as a new species by Coutinho (1925). The illegitimate taxon is likely not a Mycena. It was described to have an umbonate, pale rufescent, minutely squamulose pileus 25^40 mm diam, flesh-red lamellae, a rufescent stipe 30-60 x 3 mm, and lignicolous habit. We recognize this taxon as a nomen dubium. 56

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Mycena alphitophora BAP591

■Roridomyces rorida

Mycenaceae

Fistulina pallida Porodiscutus pendulus

Tricholomopsis decora

- Xeromphatina campaneUa ------CamaropbyBus canescens - Volvariella volvacea - gruberi

~Pieurotus tuberregium

Figure 1: continued from below. 65

■Flammuiina velutipes CyHndrobaskiium laeve Cyptotram asprata

Hydropoid clade

Strobilomyces Soccopus

- Hericium amencanum - Stereum hirsutum -Fibulorhizoctonia sp.

Figure 1: Maximum likelihood phylogeny of Agaricales based on nLSU sequence data (-InL = 29077.132125). Sequences of species from Sao Tome and Principe are indicated in bold type. Values separated by / refer to ML bootstrap proportions and Bayesian posterior probabilities. Only values greater than 70/0.70 are shown. Nodes receiving support values greater than 90/0.95 are highlighted in bold. 66

-Mycena aff. holoporphyra DED8334 Mycena cahaya KF537248

Mycena brunneoviolacea BAP656 Mycena brunneoviolacea BAP594 — Mycena pearsoniana JN182200

- r Mycena pura EU517504 ' Mycena diosma JF908417 92/1.0N Mycena sinar var tangkaisinar KF537251 '— Mycena seminau KF537252

Mycena rosella KX449424 Mycena roseila JF908488 — Mycena pura JN182202 r Mycena aff discobasis DED8211 I Mycena aff discobasis BAP658 -A3.77 — Mycena chlorophos LC274964 -/0 84 Mycena longinqua BAP648 85/1.0 Mycena antennae BAP660 ' 0.8 - Mycena discogena BAP649 64/0.99 — Mycena cyanohiza JF908385

- Mycena amicta KJ705188 ------Mycena breviseta BAP633 JF908439 KT900141

______i Mycena lasiopus BAP603 ‘ Mycena lasiopus BAP635 ------Mycena oboensis BAP669 ------Mycena alphitophora BAP591 Mycena alphitophora MH136831 -/0-81 Mycena corynepbora JF908369 -Roridomyces rorida FJ596761 - Panellus stypticus AF289071 - Mycena capillaris JF908443 A> / - Mycena alexandri KT900144 k 71/- 0.2

Figure 2: continued from below. 67

Filoboletus manipuiaris KF746992 90/0.93 \ Filoboletus manipuiaris KY349090

J Filoboletus pallescens DED8303 99/1.0 Filoboletus pallescens BAP654

' Favolaschia dnnabarina KF727415 Favolaschia caiocera KX447791

— MF686517

Mycena galopus BAP593 Mycena galopus FR846482 r Mycena galopus KU516420 99/1.0 Mycena dtrinomarginata JF908416 r l Mycena albidoaquosa JF908490

Mycena polyadelpha JF908456 97/1.0 / j— Mycena sangumoienta JF908370 L Mycena solis BAP592 71/0.96 Mycena seynesii JF908470 rI MycenaM supina JF908389

L JF908462

Mycena piumbea DQ494677 L Mycena phaeonox BAP615

- Mycena abramsii JF908400

Mycena aurantiomarginata JF908479

------ KJ705183 - JF908434

- JF908442

Mycena epipterygia JF908468

— Mycena subcana JF908457

Mycena dura JF908448

— Mycena oiida KF499358

- Mycena capillaripes JF9Q8489

- Chtocybe brunneocaperata NR_137868 82

Figure 2: Maximum likelihood phylogeny of Mycenoid fungi based on ITS sequence data (-InL = 13908.844047). Sequences of species from Sao Tome and Principe are indicated in bold type. Values separated by / refer to ML bootstrap proportions and Bayesian posterior probabilities. Only values greater than 70/0.70 are shown. Nodes receiving support values greater than 90/0.95 are highlighted in bold. 68

Clitocybula intervenosa BAP613 HClitocybula intervenosa BAP588 I Clitocybula lacerata HM191747 I Clitocybula lacerata HM191746 //■ “Trogia delicata” DED8235 -/0.85 - Clitocybula oculus DQ192178 r Clitocybula familia KM406970 -/0.9 ' Clitocybula familia JF730328 — Hydropus atramentosus JF908050 -/0.86 Hydro pus marginellus EU669314 Hydropus murinus BAP657 - Megacollybia clitocyboidea NR_119690 Megacollybia platyphylla AF498289 98/0.9 — Clitocybula atroalba DQ192179 -Hydropus funebris JQ694112 Hydropus globosporus BAP661

95/ 1.0 j Clitocybula albida MG250189 I Clitocybula albida MG250188 99/0.97 clitocybula flavoaurantia HM191744 1 ClitocybulaClit flavoaurantia HM191745

91/ 1.0 93/ 1.0 Clitocybula flavoaurantia HM191743 - Mycena auricoma DQ490647 ------Clitocybula sulcata KR029720 Atheniella aurantiidisca DQ490646 Atheniella amabilissima DQ490644 j Hydropus cfscabripes DQ404389 - Hydropus trichoderma JF908049

70/0.95 Clitocybula lignicola HM191731 _ _ _ Clitocybula lignicola HM191733 Clitocybula lignicola HM191732

-/1.Q ■ Pseudoclitocybe cyathiformis HM191730 . Pseudoclitocybe cyathiformis HM191729 l Pseudoomphalina kalchbrenneri HM191752 pseudoomphalina pachyphylla HM 191749 ( 0/0.91 I , Omphalina mutila FJ770394 ' Omphalina mutila FJ770399 - Infundlbulicybe gibba GU188436 0.3

Figure 3: Maximum likelihood phylogeny of Hydropoid fungi based on ITS sequence data (-InL = 9400.411582). Sequences of species from Sao Tome and Principe are indicated in bold type. Values separated by / refer to ML bootstrap proportions and Bayesian posterior probabilities. Only values greater than 70/0.70 are shown. Nodes receiving support values greater than 90/0.95 are highlighted in bold. 69

Figure 4: Basidiomes of Mycena brunneoviolacea (BAP 594, Holotype). Scale bar = 10 mm. Figure 5: Mycena brunneoviolacea (BAP 594, Holotype). a. Basidiospores. b. Basidia and basidiole. e. Cheilocystidia and pleurocystidia. d. Caulocystidia. Scale bar = 10 |xm. Figure 6: Mycena aff holoporphyra (DED 8334). a. Basidiospores. b. Basidia. c. Cheilocystidia. d. Pileipellis hyphae. e. Caulocystidia. Scale bar = 10 |am. 72

Figure 7: Basidiomes of Mycena aff discobasis (DED 8211). Scale bar =10 mm. 73

Figure 8: Mycena aff. discobasis (DED 8211). a. Basidiospores. b. Basidia. c. Cheilocystidia. d. Pileipellis hyphae. e. Caulocystidia. Scale bar = 10 |nm. Figure 9: Basidiomes of Mycena solis (BAP 592, Holotype). Scale bar =10 mm. Figure 10: Mycena solis (BAP 592, Holotype). a. Basidiospores. b. Basidia. c. Cheilocystidia of two types, d. Pileipellis hyphae. e. Stipitipellis hyphae. .Celcsii.d iepli yhe .Cuoytda cl a 10 = bar Scale Caulocystidia. e. hyphae. Pileipellis d. Cheilocystidia. c. Figure 11: Figure a Mycena antennae Mycena

c< < «■ (BAP 660, Holotype). a. Basidiospores. b. Basidia. Basidia. b. Basidiospores. a. Holotype). 660, (BAP c \xm. 77

Figure 12: Basidiomes of Mycena galopus (BAP 593). Scale bar = 10 mm. Figure 13: Mycena galopus (BAP 593). a. Basidiospores. b. Basidia. c. Cheilocystidia. d. Pileipellis hyphae. e. Stipitipellis hyphae. Scale bar =10 (j.m. 79

Figure 14: Mycena breviseta (BAP 633). a. Basidiospores. b. Basidia. c. Pileipellis hyphae. d. Pileus marginal cystidia. e. Pileosetae. f. Caulocystidia. Scale bar = 10 [xm. 80

Figure 15: Basidiomes of Mycena phaeonox (BAP 615, Holotype). Scale bar = 10 mm. Figure 16: Mycenaphaeonox (BAP 615, Holotype). a. Basidiospores. b. Basidia and basidiole. e. Cheilocystidia. d. Caulocystidia. Scale bar = 10 |im. Figure 17: Basidiomes of Mycena longinqua (BAP 648, Holotype). Scale bar = 1 mm. 83

OO

Figure 18: Mycena longinqua (BAP 648, Holotype). a. Basidiospores. b. Basidia. e. Cheilocystidia. d. Pileipellis hyphae. e. Caulocystidia. Scale bar = 10 (am. 84

Figure 19: Basidiomes of Mycena alphitophora (BAP 591). Scale bar = 10 mm. 85

Figure 20: Mycena alphitophora (BAP 591). a. Basidiospores. b. Basidia. c. Cheilocystidia. d. Acanthocysts. e. Pileus marginal cystidia. f. Caulocystidia. Scale bar =10 (xm. 86

Figure 21: Mycena discogena (BAP 649). a. Basidiospores. b. Basidia. c. Cheilocystidia. d. Acanthocysts. e. Caulocystidia. f. Basal disc cystidia. Scale bar =10 (am. 87

Figure 22: Mycena lasiopus (BAP 603). a. Basidiospores. b. Basidia and basidiole. e. Cheilocystidia. d. Acanthocysts. e. Cherocytes. f. Caulocystidia. Scale bar = 10 pm. 88

Figure 23: Mycena oboensis (BAP 669, Holotype). a. Basidiospores. b. Basidia. c. Acanthocysts. d. Cherocytes. e. Pileus marginal cystidia. f. Caulocystidia. Scale bar =10 |im. Figure 24: Basidiomes of Mycena lamprospora (BAP 634). Scale bar = 1 mm. 90

Figure 25: Mycena lamprospora (BAP 634). a. Basidiospores. b. Basidia and basidiole. c. Cheilocystidia of two types, d. Pileipellis terminal cells of two types, e. Caulocystidia. Scale bar =10 |_im. 91

Figure 26: Basidiomes of Clitocybula intervenosa a. BAP 588. b. BAP 613 (Holotype). Scale bar =10 mm. Figure 27: Clitocybula intervenosa (BAP 613 Holotype a-c, BAP 588 d-f). a,d. Basidiospores. b,e. Basidia and basidioles. c,f. Caulocystidia. Scale bar = 10 (xm. 93

Figure 28: Basidiomes of Filoboletuspallescens a. BAP 654. b. DED 8303 Scale bar = 10 mm. Figure 29: Filoboletuspallescens (DED 8303). a. Basidiospores. b. Basidia. e. Cheilocystidia. d. Caulocystidia. Scale bar =10 (am. 95

Figure 30: Basidiomes of Hydropus globosporus (BAP 661, Holotype). Scale bar =10 mm. Figure 31: Hydropus globosporus (BAP 661, Holotype). a. Basidiospores. b. Basidia. e. Cheilocystidia and pleurocystidia. d. Caulocystidia. Scale bar =10 (am. 97

Figure 32: Basidiomes of Hydropus murinus (BAP 657, Holotype). Scale bar = 10 mm. 98

Figure 33: Hydropus murinus (BAP 657, Holotype). a. Basidiospores. b. Basidia. c. Cheilocystidia and pleurocystidia. d. Pileipellis terminal cells, e. Caulocystidia. Scale bar =10 |xm. Figure 34: “Trogia delicata” (DED 8235). a. Basidiospores. b. Basidia and basidiole. c. Pileipellis hyphae. e. Caulocystidia. Scale bar =10 pm.