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Favolaschia species (Agaricales, Basidiomycota) from Ecuador and Panama

February 2013
Nova Hedwigia 96(1-2):117-165

February 2013
96(1-2):117-165

DOI:10.1127/0029-5035/2012/0070

Authors:

Thomas Læssøe

University of Copenhagen

Roland Kirschner

National Taiwan University

Meike Piepenbring

Goethe-Universität Frankfurt am Main

Based on recent collections from Ecuador and Panama and on type studies, 16 species of Favolaschia (Mycenaceae, Agaricales), including one new to science, are presented with detailed descriptions and illustrations. Favolaschia macropora is described as new from a bamboo species from Panama. Favolaschia cinnabarina and F. gaillardii probably are synonyms. Six species are recorded from Ecuador and seven from Panama with F. andina, F. aurantiaca, and F. gaillardii being new to Ecuador and F. andina, F. aurantiaca, F. cinnabarina and F. heliconiae to Panama. A synoptic key is presented containing data from this study and the literature. New DNA sequences (nuc LSU rDNA) from five species from Panama are used in molecular-phylogenetic analyses. Standards for future species descriptions are outlined. Problems with literature, concepts on species diversity, and distinguishing characteristics are discussed.

Favolaschia andina. (A) Optical longitudinal section through the pileus (KG0025). Scale bar = 10 mm. (B) Optical longitudinal section through a pore edge (TL-10053). Scale bare = 50 µm. (C) Spores in lateral and front view, one germinating (fourth from above). Top down: 1, 2 (KG0028), 3-5 (AAU 59886), 6 (KG0029). Scale bar = 10 µm. (D) Subhymenial hyphae (TL-10053). Scale bar = 10 µm. (E) Basidia. Top down: 1 (KG0025), 2 (TL-10053), 3 (AAU 59886). Scale bar = 10 µm.

…

Favolaschia andina. (A) Pileipellis (TL-10053). Scale bar = 50 µm. (B) Acanthocystida. Left: three-dimensional view, right: optical longitudinal view. From left to right: 1 (KG0028), 2-4 (AAU 59886), 5 (KG0029). Scale bar = 10 µm. (C) Basidia and spores of the type material (B7022). Scale bar = 10 µm. (D) Acanthocystida and gloeocystidia of the type material (B7022). Scale bar = 10 µm.

…

Favolaschia aurantiaca. (A) Optical longitudinal section through the basidioma (KG0022). Scale bar = 5 mm. (B) Optical longitudinal section through a pore edge with acanthocystida (drawing constructed from both KG0013 and KG0020). Scale bar = 50 µm. (C) Optical longitudinal section through a pore edge with acanthocystida and gloeocystidia (drawing constructed from both KG0013 and KG0020). Scale bar = 50 µm. (D) Subhymenial hyphae with basidia. From left to right: 1 (KG0013), 2 (KG0020). Scale bar = 10 µm. (E) Spores and basidium (KG0013, KG0020). Scale bar = 10 µm.

…

Favolaschia aurantiaca. (A) Optical longitudinal section through the pileipellis (KG0013). Scale bar = 50 µm. (B) Acanthocystida. Left: three-dimensional view (KG0013). Scale bar = 10 µm. (C) Gloeocystidia (KG0013). Scale bar = 10 µm.

…

+13

Favolaschia cinnabarina. Basidium and spores (AGo101). Scale bar = 10 µm.

…

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Nova Hedwigia Vol. 96 issue 1–2, 117–165

published online November 8, 2012 Article

Germany. DOI: 10.1127/0029-5035/2012/0070 0029-5035/2012/0070 $ 12.25

Favolaschia species (Agaricales, Basidiomycota) from

Ecuador and Panama

Kerstin Gillen1,*, Thomas Læssøe², Roland Kirschner3 and Meike

Piepenbring1

1 Department of Mycology, Institute for Ecology, Evolution & Diversity, J.W. Goethe-

Universität, Siesmayerstr. 70, 60323 Frankfurt am Main, Germany

² Department of Biology, University of Copenhagen, Universitetsparken 15, 2100

København Ø, Denmark

3 Department of Life Sciences, National Central University, Jhongli City, 320 Taoyuan

County, Taiwan

With 22 gures and 2 tables

Gillen, K., T. Læssøe, R. Kirschner & M. Piepenbring 2012: Favolaschia species (Agaricales,

Basidiomycota) from Panama and Ecuador. – Nova Hedwigia 96: 117–165.

Abstract: Based on recent collections from Ecuador and Panama and on type studies, 16 species of

Favolaschia (Mycenaceae, Agaricales), including one new to science, are presented with detailed

descriptions and illustrations. Favolaschia macropora is described as new from a bamboo species

from Panama. Favolaschia cinnabarina and F. gaillardii probably are synonyms. Six species are

recorded from Ecuador and seven from Panama with F. andina, F. aurantiaca, and F. gaillardii being

new to Ecuador and F. andina, F. aurantiaca, F. cinnabarina and F. heliconiae to Panama. A synoptic

key is presented containing data from this study and the literature. New DNA sequences (nuc LSU

rDNA) from ﬁve species from Panama are used in molecular-phylogenetic analyses. Standards for

future species descriptions are outlined. Problems with literature, concepts on species diversity, and

distinguishing characteristics are discussed.

Key words: Andes, Choco, Neotropics, taxonomy.

Introduction

Favolaschia species are worldwide distributed (excluding Antarctica), with a high

diversity in Latin America. According to Singer (1974), the highest species diversity

is in lowland warm-temperate to subtropical and tropical zones, however, the species

occurring in tropical montane zones are insufﬁciently investigated. From Ecuador,

*Corresponding author; email: kerstin.gillen@slu.se

118

species occurring up to the timber line at approx. 4000 m asl are reported. Apart from

that, several species are reported from temperate lowland zones, e.g. New Zealand

(Johnston et al. 2006), Japan (Kobayasi 1952), and recently Italy (Vizzini & Zotti

2002, Vizzini et al. 2008).

The species described in this paper have been collected in Panama (mainly July 2009)

and in Ecuador (different periods 1983–2005). Both Ecuador and Panama are known as

hot spots regarding vascular plants (e.g. Mutke & Barthlott 2005). The total number of

fungal species worldwide is estimated at 1.5 millions (Hawksworth 1991) with a large

number of undescribed species presumed in tropical areas, however, until now only ca

100,000 species have been described (Kirk et al. 2008). The number of fungal species

for Panama is estimated at about 50,000 with only about 1,800 species recorded in

2006 (Piepenbring 2007), resp. ca 100,000 for Ecuador with 3,766 species recorded

(Læssøe & Petersen 2008a).

The genus Favolaschia (Pat.) Pat., belonging to the family Mycenaceae (Moncalvo

et al. 2002, Bodensteiner et al. 2004), comprises currently 145 species names (http://

www.indexfungorum.org), but it is unknown how many species are represented by

them. Singer (1974) in his monograph recognized 51 species and several subspecies

and varieties. Generic characters are the porioid hymenophore, amyloid spores (e.g.

Patouillard 1887, 1900, Singer 1945), and the frequent presence of gloeocystidia

and acanthocystida (terminal, mostly swollen tips of hyphae covered by outgrowths;

Clémençon 1997) as well as the usually gelatinous trama. In the past gross morphology

has been overemphasized for classiﬁcations with the delimitation in relation to Panellus

P.Karst. being particularly deﬁcient. Favolaschia minima (Jungh.) Kuntze has recently

been transferred to Panellus minimus (Jungh.) P.R.Johnst. & Moncalvo, based on ITS

sequence data (Johnston et al. 2006). This species presents all gross morphological

characters representative for Favolaschia sp., however, true gloeocystidia and

acanthocystida are lacking.

A saprobic lifestyle is presumed and basidiomata are mostly found on decaying parts

of di- and monocotyledonous plants, as well as ferns. It is insufﬁciently investigated

how far substrate specificity pertains on species level. Singer (1974) assumed high

substrate specificity, whereas the so far best investigated species F. calocera has been

reported from a range of different substrates (Vizzini et al. 2008). Jonsson & Nylund

(1979) observed that F. dybowskyana (Singer) Singer forms a "mycorrhiza" with the

protocorms of the Japanese orchid Bletilla striata (Thunb.) Rchb.f.

A good overview on the history of the genus, including the most important references

is given by Johnston et al. (2006). Favolaschia was ﬁrst deﬁned as a section of the

by now invalid genus Laschia (Patouillard 1887) and subsequently raised to genus

level (Patouillard & Lagerheim 1892). Subsequent important publications are the

monographs by Singer (1945 and 1974) and Pegler’s (1987) revision of the Cuban

material described by Berkeley & Curtis (1868).

So far, no identiﬁcation key containing all published species names of Favolaschia

exists. With the construction of a synoptic key, it is for the ﬁrst time possible to include

all described species; hence, Favolaschia species can be keyed out correctly by the

use of the contemporary existing knowledge.

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Material and methods

Favolaschia specimens were collected in montane rainforest areas at altitudes from ca 1200–2200 m

asl. in the province Chiriquí, Western Panama, with the main ﬁeldwork carried out in July 2009 by

the corresponding author and others. Material from Ecuador was collected at several sites within

the "fungal biodiversity project in Ecuador" and earlier projects (T.Læssøe and others), at altitudes

from 250 to 3800 m asl. The Panamanian specimens are deposited at the National Herbarium of

the Universidad de Panamá (PMA), with duplicates at the Botanische Staatssammlung München,

Germany (M), and the Ecuadorian ones at Universidad Católica (QCA) and Herbario Nacional

(QCNE) in Quito, Ecuador, with duplicates at Statens Naturhistoriske Museum Copenhagen, Botanisk

Museum, Denmark (C) with some specimens recently transferred from the University of Aarhus

(AAU) to C. Type material for comparative studies was obtained from Field Museum of Natural

History, Chicago (F), Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard,

Cambridge (FH), Royal Botanic Gardens, Kew (K), and New York Botanical Garden (NY).

For light microscopy, material was mounted in 10% KOH. If possible, measurements of at least 50

spores and basidia have been made (concerning collected material from Ecuador and Panama), resp.

25 spores and basidia (concerning type material), if possible. The measurements are given as extreme

values in parentheses and means ± standard deviation, with n = number of measured cells, and c =

number of investigated collections. For the spores, the q value (length/width) was calculated. Iodine

reactions were tested with Melzer’s reagent (recipe: 100 g chloral hydrate, 5 g potassium iodide, 1.5 g

iodine, 100 ml distilled water). Optical longitudinal sections with a thickness of 15–20 µm were made

with help of a cryostate (Leica CM1510). Shapes of spores, acanthocystida and gloeocystidia were

given in accordance with the terminology in Flora Agaricina Neerlandica (Noordeloos et al. 1988).

Comprehensive species descriptions containing full details of macroscopic (pileus, hymenophore,

pores, stipe) and microscopic (spores, basidia, acanthocystida, gloeocystidia, and hyphae of the trama)

characteristics were created, regarding both features of fresh and dried material (if both available).

For DNA isolation, two methods were used. Selected freshly collected material from Panama were

bruised with a sterilized razor blade, a drop of sterilized water was added, and eventually transferred

to FTA Cards ("FTA Starter Pack", Whatman Schleicher and Schuell). FTA Cards contain chemicals

that lyse cells, denature proteins and protect nucleic acids from nucleases, oxidative and UV damage.

Here, they were chosen to avoid the process of isolating DNA in the tropics and hence to minimize

contamination of the target material. The following steps were completed according to manufacturer’s

instructions.

Selected dried material from Ecuador was blended with sterilized glass balls and shredded for 2×

1.5 min at a frequence of 30 Hz with a RETSCH Mixer Mill. The following steps were completed

according to V.07.06, "peqGOLD Fungal DNA Mini isolation protocols, C: Kurzprotokoll". For

ampliﬁcation of the DNA by PCR, the SAWADY Taq DNA polymerase kit and peqGOLD dNTP-Mix

(PEQLAB) kit were used. A reaction mix containing reaction buffer Y, enhancer solution P, dNTP

mix, Taq polymerase, distilled autoclaved water, and the primer pairs NL1 and NL4, resp. ITS1/

ITS1f and ITS4 (Euroﬁns MWG GmbH Ebersberg) was made. 45 µl were added to each FTA Card

sample, resp. to 5 µl of each isolated DNA sample. The used thermocycler program was TOUCH

60 (Weiß et al. 1998). Agarose gelelectrophoresis using GelRed was performed to analyse the PCR

products. PCR products were puriﬁed using the PeqLab E.Z.N.A.® Cycle-Pure kit. Sequencing of

dsDNA was done by Scientiﬁc Research & Development GmbH (Oberursel, Germany). Sequences

were edited with CodonCode Aligner version 1.2.2 (2002–2003, CodonCode Corporation). An

alignment was produced with the default options of MEGA version 4.1 (1993-2008) using partial

DNA sequences of the gene coding for the nuclear large subunit rRNA (LSU rDNA) deposited in

GenBank (accession numbers listed in Table 1) without manual editing. MEGA version 4.1 was used

to perform a neighbour-joining (NJ) analysis with 1000 replicates. The optimal tree was chosen.

The synoptic key was constructed with DELTA (Dallwitz 1974, 1980, Dallwitz et al. 1993). All

published Favolaschia species names were included. Species of which type material was investigated

in the context of the present study were marked with the appendix "(type!)". For each species name,

characters were entered according to own investigation and literature. For unknown characters all

possible character states were entered. If in the literature a single value is stated (e.g. measurements

120

of basidia and spores), this value was for the present interpolated ± 5 µm and will be interpolated

more exactly in the key when more information becomes available. Deﬁnitions and illustrations

were included into the key. DELTA Intkey is used to display the key. The key was uploaded to the

website http://species-id.net/wiki/key_to_Favolaschia_(Kerstin_Gilen).

Results

Species descriptions

Favolaschia alsophilae Singer, Beihefte Nova Hedwigia 50:102–103 (1974) Figs 1A, 2

Type: HoloType: Colombia, departamento Valle del Cauca, Dagua road near km 25, walking

up to 1900 m alt., 28.IV.1968, R.Singer, B6443 (F!).

Pileus 0.5–1.5 × 1–2 mm (n=10, c=1), convex, pale brown to grey, glabrous, margin

crenulated, outline of pores visible on upper surface and colour of depressions slightly

Table 1. GenBank accession numbers of species used in the molecular-phylogenetic analysis.

Species Collection

number GenBank

accession no. Country Reference

Favolaschia andina KG0025 HM246679 Panama own material

Favolaschia aurantiaca KG0013 HM246676 Panama own material

Favolaschia cf. calocera JM98/186 AF261418 China (Yunnan) Moncalvo et al. 2002

Favolaschia calocera PDD71528 AY572007 New Zealand Bodensteiner et al. 2004

Favolaschia calocera JM98/372 AF261419 Thailand Moncalvo et al. 2002

Favolaschia cinnabarina AF261416 AF261416 not stated Moncalvo et al. 2002

Favolaschia dealbata KG0015 HM246677 Panama own material

Favolaschia heliconiae KG0026 HM246680 Panama own material

Favolaschia macropora KG0027 HM246682 Panama own material

Favolaschia peziziformis PDD 67440 DQ026246 Japan Bodensteiner et al. 2004

Favolaschia cf. sprucei TH6418 AY572008 Guyana Johnston et al. 2006

Mycena crocata GLM 45968 AY207241 not stated Walther et al. 2005

Mycena galericulata GLM 45970 AY207251 not stated Walther et al. 2005

Poromycena manipularis JM98/217 AF261423 not stated Moncalvo et al. 2002

Fig. 1: Basidiomata of the described Favolaschia species. (A) F. alsophilae. Type material (B6443).

(B) F. andina.Type material (B7022). (C) F. aurantiaca.KG0013. (D) F. dealbata. KG0015.

(E) F. fendleri. Type material (B4118). (F) F. gaillardii. TL-8224. (G) F. heliconiae. KG0011.

(H) F. macropora. Type material (KG0027). (I) F. pantherina. Type material (M1003). (J) F. pterigena

var. boliviana. Type material (B817). (K) F. roseogrisea. Type material (B6035). (L) F. sabalensis

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var. geonematis. Type material (B7441). (M) F. teapae. Type material (M8798). (N) F. violascens

Type material (B1227). Scale bars = 1mm, except (G) = 10 mm.

122

darker than the colour of pores. Hymenophore consisting of 2–8 pores; pore edges

with same colour and surface as pileus, mostly thin. Pores 0.3–0.9 × 0.2–0.6 mm

(n=15, c=1), mostly round. Stipe absent or rudimentary (up to 1 mm long) and laterally

attached to the pileus.

Fig. 2: Favolaschia alsophilae (type material, B6443). (A) Basidiomata on Alsophilae sp. Scale

bar = 1 mm. (B) Optical longitudinal section through a basidioma. Scale bar = 1 mm. (C) Spores

and basidia. Scale bar = 50 µm. (D) Left: acanthocystida, scale bar = 10 µm. Right: gloeoplerous

hyphae, scale bar = 50 µm.

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Spores (7–)8–10(–11) × (5–)5.5–7.5(–8) µm, Q=(1.2–)1.3–1.5(–1.6) (n=25, c=2),

amygdaliform or ellipsoid, rarely oblong or broadly ellipsoid, smooth, thin-walled,

multiguttulate or without guttules, weakly amyloid. Basidia (27–)30–37(–42) ×

(7–)7.5–9.5(–10) µm (n=25, c=2), cylindric or clavate, 4-spored. Acanthocystida

numerous on pore edges, scattered in pileipellis, oblong to broadly clavate, (17–)19–

32.5(–40) × (9–)10–13.5(–15) µm (n=15, c=2), with numerous rod-shaped outgrowths

(up to 5 µm in length) covering the distal part or the entire cell, wall thickened below

the ornamented part, colourless. Gloeocystidia absent. Gloeoplerous hyphae few

scattered in the trama, with numerous small or slightly bigger droplets (greenish grey

in H2O and KOH), vermiform, (40–)62–141(–170) × 4–7(–8) µm (n=15, c=2), thin-

walled. Trama not or slightly gelatinous, hyphae irregularely arranged in pileus trama,

± parallel in pore walls, thin-walled.

SubSTraTe and HabiTaT: On dead fronds and stems of tree fern (Alsophila sp.) in

mountain rainforests (alt. ca 1900-2400 m asl).

Known diSTribuTion: Colombia (Singer 1974), Ecuador (new record).

SpeCimen examined: eCuador, prov. Napo, Cuyuja, South of river, pasture with Alnus sp.

along river, 2400 m asl, on petioles of tree fern fronds in Chusquea scrub, 04.V.2002,

T.Læssøe, TL-9480 (QCA, C).

noTeS: Favolaschia alsophilae is characterized by the few large pores and the absence of

gloeocystidia. The species is quite similar to the newly described species F. macropora,

but differs by the presence of clamps.

Favolaschia andina Singer, Beihefte Nova Hedwigia 50: 102 (1974) Figs 1B, 3, 4

Type: HoloType: Colombia, departamento Cundinamarca, páramo Chisaca-Sumapás, on twig of a

dicotyledonous plant, 8.V.1968, R.Singer, B7022 (F!).

Pileus up to 10 mm diam. (fresh) and (0.8–)0.9–2(–3) × 1.5–3(–4.5) mm (when

dry) (n=15, c=3), semiglobose to applanate, pale mouse grey (fresh), beige to brown

(when dry), glabrous or rarely slightly pulverulent; outline of pores visible on sterile

carpophore surface (fresh and when dry); margin crenulate, straight or involute.

Hymenophore consisting of 10–60 pores; two pores/mm (fresh), pore edges with

the same colour and surface as pileus or slightly paler, mostly thin and often torn.

Pores 0.2–0.3(–0.4) × 0.1–0.2(–0.3) mm (when dry) (n=15, c=3), irregular or radially

elongated. Stipe absent or rudimentary.

Spores (5.5–)7.5–9.5(–10.5) × (4.5–)5.5–7.5(–8.5) µm, Q=(1.1–)1.2–1.5 (n=50, c=4)

[type (8–)9–10.5(–11) × (6–)6.5–8 µm, Q=1.3–1.5(–1.7), n=25], amygdaliform or

broadly ellipsoid to ellipsoid, smooth, thin-walled, multiguttulate or with a single

large droplet, amyloid. Basidia (17–)21.5–33(–40) × (7.5–)8.5–10.5(–12) µm (n=51,

c=4) [type (25–)26.5–33(–40) × 9–10.5(–12) µm, n=25], cylindric to clavate, 4-spored.

Acanthocystida numerous on pore edges and pileipellis, often breaking off during

drying process and forming the pulverulent covering of the pileus, of two types:

1. globose to clavate, (13–)17–26.5(–32) × (11–)13.5–23(–29) µm (n=30, c=3)

[type (19–)20–24(–25) × (16–)17–20.5(–22) µm, n=15], with numerous rod-shaped

outgrowths densely covering the entire cell or restricted to the apex, wall thickened

below the ornamented part; 2. oblong, not broader than trama hyphae, wall not

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thickened, colourless. Gloeocystidia with numerous small droplets (greenish grey

in H2O and KOH), thin-walled, of two types: 1. vermiform, ± parallel to pileipellis,

(60–)62–152(–190) × (6–)6.5–9(–10) µm (n=15, c=3) [type (65–)70–140(–170) ×

(4–)5–7.5(–9) µm, n=15]; 2. shaped as basidia, scattered among the basidia, not present

Fig. 3: Favolaschia andina. (A) Optical longitudinal section through the pileus (KG0025). Scale

bar = 10 mm. (B) Optical longitudinal section through a pore edge (TL-10053). Scale bare = 50 µm.

3–5 (AAU 59886), 6 (KG0029). Scale bar = 10 µm. (D) Subhymenial hyphae (TL-10053). Scale bar

= 10 µm. (E) Basidia. Top down: 1 (KG0025), 2 (TL-10053), 3 (AAU 59886). Scale bar = 10 µm.

125

in all basidiomata, (25–)34–52(–56) × (6–)7–10(–11) µm (n=15, c=3). Gloeoplerous

hyphae scarce and absent in some basidiomata, contents thromboplerous (pale grey in

H2O and KOH). Trama slightly or rarely very gelatinous, hyphae irregularely arranged

in pileus trama, ± parallel in pore wall and stipe trama, thin-walled.

Fig. 4: Favolaschia andina. (A) Pileipellis (TL-10053). Scale bar = 50 µm. (B) Acanthocystida. Left:

three-dimensional view, right: optical longitudinal view. From left to right: 1 (KG0028), 2–4 (AAU

59886), 5 (KG0029). Scale bar = 10 µm. (C) Basidia and spores of the type material (B7022). Scale

bar = 10 µm. (D) Acanthocystida and gloeocystidia of the type material (B7022). Scale bar = 10 µm.

126

SubSTraTe and HabiTaT: On rotting wood of dicotyledonous trees or shrubs or herbaceous

stems in mountain or cloud forests (alt. of investigated specimens ca 1200-3800 m asl).

Known diSTribuTion: Colombia (Singer 1974), Ecuador, and Panama (new records).

SpeCimenS examined: eCuador, prov. Carchi, El Angel, Páramo del Angel, ca 3200 m asl, on

attached Polylepis twigs, 14.II.2003, T.Læssøe & J.Salazar, TL-10053 (QCA, C). – Prov. Napo,

western slope west of Papallacta, 0°19'S/78°10'W, Asteraceae shrub and grass paramo, 3800 m asl,

on thin herbaceous stems in dense moss forest, 01.VII.1985, T.Læssøe, AAU 59886 (C). panama,

prov. Chiriquí, Lagunas de Volcán, 1200-1300 m asl, on rotting dicotyledonous twigs, 22.VII.2009,

K.Gillen KG0025, KG0028 & KG0029 (PMA, M).

noTeS: F. andina is characterized by the globose acanthocystida and the presence

of gloeocystidia among the basidia. However, according to the existent species

descriptions, this species shares many characters with F. subceracea, F. pygmaea, and

F. dumontii. Since good illustrations of these species are lacking and no type material

was investigated for these three species, it can not be clariﬁed whether the three names

refer to the same species.

Favolaschia aurantiaca Singer, Beihefte Nova Hedwigia 50: 101 (1974) Figs 1C, 5, 6

Type: HoloType: bolivia, departamento Pando, prov. Manuripi, Conquista, on Bambusa

sp., 25.III.1956, R. Singer, B2199 (F 95091!).

Pileus 1.3–8 × 0.9–8 mm (when fresh) and 1–5.5 × 0.8–3.7 mm (n=15, c=3) (when dry),

semiglobose or rarely applanate, brownish grey to light grey, rarely white (fresh) and

brown to light yellow (when dry), glabrous and gelatinous; outline of pores visible on

sterile carpophore surface and colour of depressions slightly darker than colour of pores;

margin crenulate, straight or involute (when dry). Hymenophore consisting of 10–100

pores; pore edges white to pale beige (fresh and dry), up to 1 mm in width (n=15, c=3)

(fresh), thinner when dry, glabrous (fresh and dry) or sometimes slightly pulverulent.

Pores 0.1–0.4 mm diam. (fresh) (n=15, c=3), irregular or radially elongated, rarely

round. Stipe mostly present, 1.5–5 × 0.4–1 mm (fresh) and 1.5–4 × 0.2–0.7 mm (when

dry) (n=15, c=3); concolourous with pileus, often slightly translucent (fresh), glabrous

(fresh and when dry) or slightly pulverulent (when dry), laterally attached.

Spores (6.5–)7–8.5(–9) × (4–)4.5–5.5(–6.5) µm, Q=(1.2–)1.3–1.7(–2) (n=50, c=3)

[type 7–8.5(–9) × (5–)5.5-7 µm, Q=1.1–1.5, n=15], ellipsoid to oblong, smooth,

thin-walled, multiguttulate or rarely without guttules, weakly amyloid. Basidia

(20–)20.5–27.5(–30) × (6.5–)7.5–11.5(–14) µm (n=15, c=3) [type (20–)24–34(–39)

× (7–)8–10(–11) µm, n=15], cylindric, ellipsoid or clavate, 4-spored. Acanthocystida

numerous on pore edges, in the pileipellis and stipe surface, often breaking off during

drying process and forming the pulverulent covering of the pileus, clavate to ellipsoid,

(25–)35–56.5(–39) × (7–)10.5–18.5(–25) µm (n=25, c=3) [type (11–)13–20.5(–25)

× (7–)8.5–14(–15) µm, n=15], with numerous rod-shaped outgrowths (up to 6 µm in

length) covering the entire cell, wall thickened below the ornamented part, colourless.

Gloeocystidia mostly numerous in pileipellis and stipe margin, few or absent on pore

edges, with numerous small or slightly bigger droplets (greenish grey in H2O and KOH),

thin-walled, long clavate or ellipsoid, (16.5–)35–90.5(–130) × (5–)5.5–17(–27.5) µm

(n=25, c=3) [type (20–)24.5–34.5(–39) × (11–)14–22(–26) µm, n=15]. Gloeoplerous

hyphae scattered in trama of pileus, pore edges and stipe, with same contents as

127

gloeocystidia. Trama very gelatinous, hyphae irregularely arranged in pileus trama,

± parallel in pore wall and stipe trama, thin-walled, some with a small interspace

between the hypha and the clamp.

Fig. 5: Favolaschia aurantiaca. (A) Optical longitudinal section through the basidioma (KG0022).

Scale bar = 5 mm. (B) Optical longitudinal section through a pore edge with acanthocystida

(drawing constructed from both KG0013 and KG0020). Scale bar = 50 µm. (C) Optical longitudinal

section through a pore edge with acanthocystida and gloeocystidia (drawing constructed from both

KG0013 and KG0020). Scale bar = 50 µm. (D) Subhymenial hyphae with basidia. From left to right:

1 (KG0013), 2 (KG0020). Scale bar = 10 µm. (E) Spores and basidium (KG0013, KG0020). Scale

bar = 10 µm.

128

SubSTraTe and HabiTaT: On decaying stems of Chusquea sp. in mountain rainforests

(alt. of investigated specimens from ca 1500–3500 m asl).

Known diSTribuTion: Argentina (Singer 1974), Bolivia (Singer 1974), Ecuador, and

Panama (new records).

Fig. 6: Favolaschia aurantiaca. (A) Optical longitudinal section through the pileipellis (KG0013).

Scale bar = 50 µm. (B) Acanthocystida. Left: three-dimensional view (KG0013). Scale bar = 10 µm.

129

SpeCimenS examined: eCuador, prov. Orellana, Añangu, South of Río Napo, 0°31'S/76°23'W,

260–350 m asl, tropical rain forest (terra ﬁrme), 20.VI.1983, T.Læssøe, AAU 44782 (C). – 95 km

downstream from Coca, 0°32'S/76°23'W, 300 m asl, lowland rain forest, 19.VI.1985, T.Læssøe,

AAU 59581A (C). – 300 m asl, lowland rain forest, 19.VI.1985, T.Læssøe, AAU 59640 (C). –

Prov. Napo, Cascadas de San Rafael, "Coca Falls", large waterfall on Río Coca, between El Chaco

and San Pedro de los Cofanes, 0°6'05"S/77°35'07"W, 1250 m asl, 21.VII.1985, T.Læssøe, AAU

59856 (C). – Prov. Pichincha, Yanacocha, Volcan Pichincha, 0°7'S/78°34'W, 3500 m asl, mountain

forest with thick epiphyte cover and grassland with horses, on dead mossy stem from bambusoid

grass, 5.VIII.1985, T.Læssøe, AAU 59909 (C) – Prov. Pichincha, Mindo, along stream just above

Casa Amarilla, 0°03'S/78°47'W, ca 1550 asl, on trunk of Chusquea sp., 20.III.2002, T.Læssøe &

J.H.Petersen, TL-9031 (QCA, C). – Prov. Loja, Parque Nacional Podocarpus, Cajanuma, 2800 m asl,

on Chusquea sp., 31.III.2002, T.Læssøe, TL-9147 (QCA, C). –. abundant on site of Chusquea twigs,

31.III.2002, T.Læssøe, TL-9151 (QCA, C). – Prov. Carchi, between Tuﬁño and Maldonado on the

Tulcán-Maldonado road, 2500–2800 m asl, cloud forest, on Chusquea twigs, 16.II.2003, T.Læssøe

& J.Salazar, TL-10115 (QCA, C). – Prov. Pichincha, via Calacali-Nono, 3 km towards Nono, Finca

Tara, ca 2800 m asl, grassland and dense shrubby wet forest, abundant on Chusquea and Rubus

twigs, 22.II.2003, T.Læssøe, J.Salazar, A.Calco & T.Sanjuan, TL-10187 (QCA, C). – panama, prov.

Chiriquí, Parque Internacional La Amistad, Sendero del Cerro Picacho, N08°53/4'W082°37,6', ca

2500 m asl, on rotting stems of Chusquea sp., 30.VIII.2008, R.Mangelsdorff, RMP 633 (M, PMA).

Sendero del Cerro Picacho, ca 2060–2100 m asl, on rotting stems of Chusquea sp., 18.VII.2009,

K.Gillen, M.Piepenbring & A.Gockele, KG0013, KG0014, KG0019 & KG0022. Sendero El Retoño,

ca 2110 m asl, on rotting stems of Chusquea sp., 26.V.2009, M.Piepenbring, M 4781 (M, PMA).

noTeS: Favolaschia aurantiaca is characterized by the very gelatinous trama and the

occurrence on decaying Chusquea stems. The spores measured in the type collection are

1-2 µm broader than those from the collections from Ecuador and Panama. However,

the basidiomata of the type collection did not contain many mature basidia and spores,

and the range of values of measurements probably is thereby distorted. Acanthocystida

and gloeocystidia in the here investigated specimens slightly differ regarding their size

and shape depending on the developmental stage of the basidioma.

Fries (1888) described F. testudinella (R.E.Fr.) Kuntze (as Laschia testudinella R.E.Fr.)

from a green house in Göteborg, Sweden. The basidiomata illustrated look like those

of F. aurantiaca, and the substrate is an unidentiﬁed bamboo species. A description

of the microscopic characters does not exist. Though loan requests have been made

to the herbaria of Göteborg (GB), Naturhistoriska Riksmuseet Stockholm (S), and

Evolutionsmuseet Uppsala (UPS), the type collection of F. testudinella could not be

found.

Favolaschia cinnabarina (Berk. & M.A.Curtis) Pat., Essai Taxonomique sur les familles

et les genres des Hyménomycetes (Lons-le-Saunier): 141 (190) Fig. 7

≡ Laschia cinnabarina Berk. & M.A.Curtis, Journal of the Linnean Society 10(46): 322 (1868)

Type: HoloType: Cuba, on twig of dicotyledonous plant, K Wright Plantae Cubenses, without

number, n.v.

Macroscopic and microscopic features as in F. gaillardii. Differences: spores (7.5)8–

9(–10) × (5.5–)6–7.5(–8) µm, Q=(1.1–)1.2–1.4(–1.5) (n=25, c=2), broadly ellipsoid

to ellipsoid. Basidia (24–)25–35(–41) × (7–)8–9.5(–10) µm (n=13, c=1), rarely with

basal clamps. Acanthocystida (9–)13.5–20.5(–24) × (9–)12–17(–18.5) µm (n=15,

c=1). Gloeocystidia (17.5–)20.5–36(–41) × (9.5–)12.5–19.5(–20) µm (n=15, c=1).

Trama hyphae rarely clamped.

130

SubSTraTe and HabiTaT: As in F. gaillardii.

Known diSTribuTion: Bolivia, Brazil, Colombia (resp. Singer 1974), Cuba (Berkeley

& Curtis 1868), Guyana (Singer 1974), Ecuador (Singer 1974, 1988), Panama (own

collections and Singer 1974), U.S.A. (Florida) (Pegler 1987), and Venezuela (Dennis

1961, Singer 1974).

SpeCimenS examined: eCuador, prov. Mazan, on dead twig, 28.IX.1987, V.Fleming 277,

K(M): 156855 (by Fleming annotated/identiﬁed as F. gaillardii). PANAMA, prov. Chiriquí,

Boquete, Cerro Palo Alto, Río Cristal Mountain Lodge, 1600–1800 m asl, on dead twig,

28.VI.2009, A.Gockele, AGo101 (M).

noTeS: See notes on F. gaillardii.

Favolaschia dealbata Singer, Mycologia 47(5): 763 (1955) Figs 1D, 8, 9

TypeS: HoloType: panama, Casita Alta, on Bambuseae ("spring bamboo"), 31.VII.1952, Martin &

Welden, 8070 (F).

epiType (deSignaTed Here): panama, prov. Chiriquí, Parque Internacional La Amistad, Sendero

de Cerro Picacho, 2060–2100 m asl, on decaying stems of Chusquea sp., 18.VII.2009,

K.Gillen, M.Piepenbring & A.Gockele, KG0015 (PMA!). Isoepitype (M!).

Pileus 1.3–9 × 1.2–6.7 mm (n=15, c=3), semiglobose to applanate, orange (fresh)

and orange, orange red, light yellow or pale brown (when dry) (colour caused by the

numerous gloeocystidia under the surface), glabrous; outline of pores visible on sterile

carpophore surface; margin crenulate, straight or involute. Hymenophore consisting

of 15–120 pores; pore edges with the same colour and surface as pileus, mostly very

thin and often torn. Pores 0.2–1 × 0.2–0.9 mm (n=15, c=3), mostly irregular or rarely

round. Stipe present or absent, up to 8 × 0.5–1 mm (fresh) and up to 6 × 0.2–0.3 mm

(when dry) (n=15, c=3); concolourous with pileus or rarely paler, glabrous (fresh and

when dry) or slightly pulverulent (when dry), laterally attached. In rare cases one stipe

develops two pilei.

Spores (7–)8–9.5(–10.5) × (6–)6.5–8(–9) µm, Q=1.1–1.3(–1.5) (n=50, c=3), amygdali-

form or broadly ellipsoid to subglobose (rarely ellipsoid), smooth, thin-walled,

multiguttulate, weakly amyloid. Basidia (26–)27–35(–40) × (7–)8–11(–14.5) µm

(n=32, c=2), cylindric to clavate, 4-spored, no basal clamps observed. Acanthocystida

numerous on pore edges and pileipellis, few on the stipe surface, globose to broadly

ellipsoid or rarely oblong, (15–)18–27(–32.5) × (13–)15.5–22(–26) µm (n=35, c=2),

with numerous rod-shaped outgrowths (up to 3 µm long) restricted to apex zone or

Fig. 7: Favolaschia cinnabarina. Basidium and spores (AGo101). Scale bar = 10 µm.

131

rarely covering the whole cell, wall thickened below the ornamented part, colourless

or rare in dissolved contents (greenish in H2O and KOH). Gloeocystidia numerous in

pileipellis and fewer on pore edges, rarely in pileus and stipe trama, with numerous

small or slightly bigger droplets (orange, greenish yellow or rarely colourless in H2O

and KOH), thin-walled, clavate, (14.5–)17.5–37(–47) × (7–)10–19(–27.5) µm (n=40,

c=2). Gloeoplerous hyphae scattered in trama of the pore edges, pileus and stipe,

Fig. 8: Favolaschia dealbata. (A) Optical longitudinal section through the pileus (KG0017). Scale

bar = 0.5 mm. (B) Pore edge (KG0017). Scale bar = 10 µm. (C) Subhymenial hyphae with basidia

(KG0015). Scale bar = 10 µm. (D) Basidia (TL-9151). Scale bar = 10 µm. (E) Spores in lateral and

front view including a germinating spore (fourth from left) (TL-9151). Scale bar = 10 µm.

132

contents thromboplerous (pale grey in H2O and KOH). Trama slightly or rarely very

gelatinous, hyphae irregularely arranged in pileus trama, ± parallel in pore wall and

stipe trama, thin-walled.

SubSTraTe and HabiTaT: On decaying stems of Chusquea sp. in mountain rainforests

(alt. of investigated specimens from ca 2000–2800 m asl).

Known diSTribuTion: Ecuador (new record) and Panama (Singer 1955).

Fig. 9: Favolaschia dealbata. (A) Optical longitudinal section through the pileipellis (KG0017). Scale

bar = 50 µm. (B) Gloeocystidia (TL-9151). Scale bar = 10 µm. (C) Acanthocystida. Left: optical

longitudinal sections, right: three-dimensional views (KG0015). Scale bar = 10 µm.

133

SpeCimenS examined: eCuador, prov. Loja, Parque Nacional Podocarpus, Cajanuma, 2800 m asl, on

Chusquea sp., abundant on site, 31.III.2002, T.Læssøe, TL-9151 (QCA, C). – panama, prov. Chiriquí,

Parque Internacional La Amistad, Sendero de Cerro Picacho, ca 2060–2100 m asl, on decaying

stems of Chusquea sp., 18.VII.2009, K.Gillen, M.Piepenbring & A.Gockele, KG0017 & KG0020

(PMA, M). – Ca 2110 m asl, on decaying stems of Chusquea sp., 26.V.2009, M.Piepenbring, M4780

(PMA, M). – Parque Nacional Volcán Barú, Sendero Los Quetzales, ca 2500 m asl, 15.IX.2005,

M.Piepenbring, M3560 (PMA, M).

noTeS: The type collection from Panama, which should be deposited in Field Museum

of Natural History, Chicago (F), could not be found (last request in August 2010),

and hence, could not be investigated. In the original description, Singer (1974) only

described dried material, and illustrations are lacking. The colour of the dried specimens

is said to be white, after soaking in water honey to cinnamon brown. The data obtained

from the proposed epitype and other cited material corresponds well with the meagre

data from Singer (1974).

Thus, a new epitype for F. dealbata is proposed here: the collection KG0015, as

sequences have been obtained from this specimen and data on fresh material exist.

Favolaschia fendleri Singer, Beihefte Nova Hedwigia 50: 101 (1974) Figs 1E, 10

Type: HoloType: brazil, São Paulo, Reserva Florestal do Horto Florestal, Serra de Cantareira,

17.XII.1964, Singer & Furtado, B4118 (F!).

Pileus 1–2 × 1–2.5 mm (n=8) (when dry), plano-convex to convex, pale brown to

brown, mostly glabrous, margin mostly even or weakly crenulated and sometimes

involute, mostly applanate, brownish orange, glabrous, outline of pores visible on

sterile carpophore surface. Hymenophore consisting of 15–20 pores; pore edges with

same colour and surface as pileus, mostly thin. Pores 0.1–0.3 × 0.1–0.5 mm (n=8) of

irregular shape. Stipe present and up to 1–2 mm long or absent.

Spores (8–)8.5–10 × (4–)4.5–5.5(–6) µm, Q=(1.5–)1.6–2.1(–2.5) (n=25), oblong,

amygdaliform or rarely ellipsoid, smooth, mostly without guttules or rarely

multiguttulate, weakly amyloid. Basidia (20–)21–25.5(–30) × 7–8.5(–9) µm (n=25),

cylindric or clavate, 4-spored. Acanthocystida numerous in pileipellis and on pore

edges, globose to oblong, (12–)13.5–23.5(–30) × (9–)10.5–16(–19) µm (n=15), with

numerous rod-shaped outgrowths covering the whole or distal two thirds of the cell,

wall thickened below the ornamented part, colourless. Gloeocystidia of two types:

1. numerous in pileipellis, with numerous small or slightly bigger droplets (greenish

yellow in H2O and KOH), globose to broadly clavate, (25–)30–40.5(–45) × (18–)

20–30.5(–35) µm (n=15); 2. scattered among acanthocystida in pileipellis, oblong to

clavate, contents isomorphous, (28–)29.5–37.5(–41) × (9–)11.5–15(–16) µm (n=15).

Gloeoplerous hyphae absent. Trama not or slightly gelatinous, hyphae irregularely

arranged in pileus trama, ± parallel in trama of pore walls and stipe, walls thin- to

medium thick.

SubSTraTe and HabiTaT: On dead parts of monocotyledonous plants in rainforests.

Known diSTribuTion: Brazil, Columbia, and Honduras (all Singer 1974).

noTeS: The basidiomata of the type material of F. fendleri resemble those of F. auran-

tica. Minor differences are the size of the basidiomata, the number of the pores, the

134

1–2 µm longer spores, the less gelatinous trama and the slightly broader gloeocystidia

of F. fendleri. F. aurantiaca has been found on Chusquea sp., the substrate where the

basidiomata of F. fendleri have been found has not been identiﬁed. Possibly F. auran-

tiaca and F. fendleri are synonyms, with F. fendleri being the valid name.

Fig. 10: Favolaschia fendleri (type material, B4118). (A) Basidiomata. Scale bar = 1 mm. (B)

Optical longitudinal section through a basidioma. Scale bar = 1 mm. (C) Spores and basidium. Scale

bar = 10 µm. (D) Gloeocystidia of the pileus trama. Scale bar =10 µm. (E) Acanthocystida. Scale

bar = 10 µm. (F) Gloeocystidia of the pileipellis. Scale bar = 50 µm.

135

Favolaschia gaillardii (Pat.) Pat. [as "gaillardia"], Essai Taxonomique sur les familles

et les genres des Hyménomycetes (Lons-le-Saunier): 141 (1900) Figs 1F, 11, 12

≡ Laschia gaillardi Pat., Journal de Botanique 1: 218 (1887)

Type: HoloType: venezuela, "Puerto Perico", on twig of dicot. plant, 08.V.1887, A.Gaillard

(FH 284654!).

Pileus (0.7–)1–1.1(–2.7) × (0.9–)2.1–3.5(–5.5) mm (n=15, c=3) (when dry), mostly

applanate, yellow, orange or reddish orange (when fresh) and orange, reddish orange,

pinkish, yellow or pale yellow (colour caused by numerous small orange dots on the

surface); margin mostly crenulate and sometimes involute. Hymenophore consisting

of 6–30 pores; pore edges with same colour and surface as pileus, mostly very thin.

Pores (0.25–)0.26–0.44(–0.55) × (0.15–)0.24–0.58(–0.7) mm (n=15, c=3), mostly

irregular. Stipe absent or rudimentary.

Spores (6.5–)7–9.5(–11.5) × (4–)4.5–6.5 µm, Q=(1.2–)1.4–1.7(–2.1) (n=65, c=6) [type

(6–)6.5–8 × 4–5.5(–6) µm, Q=(1.2–)1.3–1.6, n=8], amygdaliform, rarely ellipsoid or

oblong, smooth, thin-walled, multiguttulate or with one single guttule (greenish grey in

H2O and KOH), weakly amyloid. Basidia (15–)20.5–29(–35) × (5–)6.5–8.5(–12.5) µm

(n=55, c=6) [type (20–)21.5–27.5(–31) × (5–)6.5–8.5(–9) µm, n=16], cylindric or

clavate, mostly with two large sterigmata, rarely four (these mostly long and thin).

Acanthocystida numerous on pore edges and pileipellis, forming the orange dots on

the pileus surface, globose to broadly ellipsoid, (14–)16.5–32.5(–42.5) × (12.5–)13.5–26

(–34) µm (n=35, c=4) [type (13–)13.5–24(–33) × (10–)11–15.5(–17) µm, n=15], with

numerous rod-shaped outgrowths (up to 3 mm in length) covering the entire cell, wall

thickened below the ornamented part, orange or rarely colourless. Gloeocystidia

numerous in pileipellis, few on pore edges, rarely scattered among basidia, with numerous

small or slightly bigger droplets (greenish yellow in H2O and KOH), thin-walled, of two

types: 1. broadly clavate, (19–)22–36.5(–51) × (12.5–)15–20(–21.5) µm (n=25, c=3) [type

(27–)29–42.5(–45) × (18–)18.5–23(–24) µm, n=10], present in all basidiomata, 2. long

and vermiform, (20–)56–117(–120) × 5–10.5(–17.5) µm (n=11, c=1) [type (19–)26–

37.5(–40) × 10–11.5(–12) µm, n=12], present in most of the examined basidiomata.

Gloeoplerous hyphae not observed. Trama not or slightly gelatinous, hyphae irregularely

arranged in pileus trama, ± parallel in pore wall trama, thin-walled.

SubSTraT e and HabiTaT: Favolaschia gaillardii is found on rotting wood of

dicotyledonous plants, mostly twigs on the ground, in lowland and mountain rainforests

(alt. of investigated specimens from ca 250–4000 m asl).

Known diSTribuTion: Brazil (Singer 1974), Ecuador, Panama, Peru (new records), and

Venezuela (Patouillard 1887).

SpeCimenS examined: eCuador, prov. Orellana, Añangu, south of Río Napo, 0°31'S/76°23'W,

260–350 m asl, tropical rain forest (terra ﬁrme), on small sticks, 2.II.1983, T.Læssøe, AAU 43102

(C). – Pompeya, Parque Nacional Yasuni, Estación Cientiﬁca Yasuni, 0°40'26"S/76°23'50"W, 300 m

asl, "terra ﬁrme", on dicotyledonous twigs, 9.III.2002, J.H.Petersen, TL-8224 (QCA, C). – On twig

(dicotyledonous, woody), A.C.Steenkjær Hastrup, TL-8661 (QCA, C). – panama, prov. Chiriquí,

Boquete, forest above Finca Lerida, 1400–1500 m asl, on dead dicotyledonous twigs on the ground,

15.VII.2009, K.Gillen & A.Gockele, KG0007, KG0008, KG0009, and KG0010 (PMA, M). – peru,

Huánuco, Tingo Maria, grounds of the Universidad Agraria (UNAS), 09°15'S/76°W, 641 m asl, on

bark of dicotyledonous twigs, 6.VII.1987, T.Læssøe, TL-P-200 (C).

136

noTeS: F. gaillardii is the type species of the genus and was described in 1887 by

Patouillard (as Laschia gaillardii). However, the species is rather simlar to F. cinna-

barina (Berk. & M.A.Curtis) Pat. described in 1868 (Berkeley & Curtis) (as Laschia

Fig. 11: Favolaschia gaillardii. (A) Optical longitudinal section through the basidioma (TL-8224).

Scale bar = 1 mm. (B) Optical longitudinal section through a pore edge (TL-8224). After embedding

the section into KOH solution, the gloeocystidia bursted. Scale bar = 50 µm. (C) Subhymenial hyphae

with immature basidia (KG0008, KG0009). Scale bar = 10 µm. (D) Basidia. Top down: TL-P-200,

TL-8224, TL-8661, and KG0008). Scale bar = 10 µm. (E) Spores in lateral and front view. From left

to right: 1-2 (TL-8224), 3-4 (TL-P-200), 5 (KG0008). Scale bar = 10 µm.

137

cinnabarina). Singer (1974), who investigated the type material from Venezuela, notes

the slightly narrower basidia and spores as the only distinguishing character, but the

measuring method is critical. However, Singer mentions narrow spores (5.5–7.5 µm)

Fig. 12: Favolaschia gaillardii. (A) Optical longitudinal section through the pileipellis (TL-8224).

Scale bar = 50 µm. (B) Acanthocystida with dissolved content, shown as dots (KG0008). Left:

three-dimensional view, right: optical longitudinal sections. Scale bar = 10 µm. (C) Gloeocystidia.

Top: gloeocystidia of the pore edges and pileipellis (AAU 43102, KG0008, TL-8224). Down

left: gloeocystidium of the hymenium (burst in KOH solution) (TL-8224), down right: oblong

gloeocystidium of the pileipellis (KG0008). Scale bar = 10 µm.

138

produced by 4-spored basidia and broader spores (6.7–11 µm) produced by two-spored

basidia. This measuring method has to be considered critically, as it includes immature

spores still attached to basidia, which causes impreciseness in the measurements.

Rick (1938) suggested that F. gaillardii could be a variety of F. cinnabarina. Further

morphological and anatomic studies on additional material combined with a molecular-

phylogenetic investigation is recommended. It was not possible to extract DNA from

the Panamanian and Ecuadorian material. The valid name would be F. cinnabarina if

only one species can be recognized.

Favolaschia heliconiae Singer, Beihefte Nova Hedwigia 50: 101 (1974) Figs 1G, 13, 14

Type: HoloType: venezuela, prov. Sucre, trail from Los Pocitos to Río Grande Arriba, on petioles

of Heliconia sp., 13.VII.1972, Dumont et al., VE1548 (NY 775546!).

= Favolaschia oligogloea Singer, Beihefte Nova Hedwigia 50: 103 (1974), syn. nov.

Type: HoloType: eCuador, prov. Napo, Lago Agrio, on monocotyledonous culms and stems,

gregarious, 8.V.1973, Singer B 7278 (F!).

Pileus 0.7–2 mm diam. (when fresh) and 0.5–1.5 × 0.8–2 mm (when dry) (n=10,

c=3), applanate to plano-convex, white (when fresh) and white to pale yellow (when

dry), glabrous (when fresh) and glabrous or slightly pulverulent (when dry); margin

crenulated; outline of pores visible on sterile carpophore surface. Hymenophore

consisting of 10–25 pores; pore edges with same colour and surface as pileus, up to

0.25 mm broad. Pores 0.3–0.6 mm in diam. (n=10, c=3), round, radially elongated, or

irregular. Stipe present, 0.8–2 × 0.2–0.3 mm (when fresh) and 0.5–2 × 0.2 mm (when

dry) (n=10, c=3), with same colour and surface as pileus, laterally attached to the pileus.

Spores (5–)5.5–8.5(–11) × (2.5–)3–5(–6) µm, Q=(1.3–)1.6–2.1(–2.3) (n=50, c=4) [type

(5–)6.5–8(–9) × (3–)4–5.5(–6) µm, n=50], oblong to phaseoliform, rarely ellipsoid

or amygdaliform, smooth, thin-walled, colourless or multiguttulate (greenish grey

H2O and KOH), weakly amyloid. Basidia (15–)18.5–24(–29) × (5.5–)7–9.5(–11.5)

µm (n=50, c=4) [type (14–)17.5–21(–23) × (6–)7–8.5(–9) µm, n=50], cylindric or

ellipsoid, four- or rarely two-spored. Acanthocystida numerous on pore edges and

numerous to few in pileipellis and stipe margin, often breaking off during drying

process and forming the pulverulent covering of the pileus, globose, ellipsoid or

clavate, few oblong, sometimes branched, (8–)15–28(–42) × (7–)10.5–18.5(–24) µm

(n=35, c=3) [type (14–)17–22.5(–25) × (7–)8–11(–12) µm, n=30], with numerous

rod-shaped outgrowths (up to 5 µm in length) covering the entire cell, wall thickened

below the ornamented part, colourless or orange. Gloeocystidia few to numerous in

pileipellis and stipe margin, very few or absent on pore edges, with numerous small or

slightly bigger droplets (yellowish in H2O and KOH), thin-walled, ellipsoid or clavate,

(15–)24.5–60(–70) × (6.5–)7.5–11.5(–14) µm (n=20, c=2) [type (80–)86–114(–120) ×

(4–)6.5–9.5(–10) µm, n=15]. Gloeoplerous hyphae scattered in the trama of the pileus,

pore walls, and stipe, contents thromboplerous (pale grey in H2O and KOH). Trama

not or slightly gelatinous, hyphae irregularely arranged in pileus trama, ± parallel in

pore wall and stipe trama, thin-walled.

SubSTraTe and HabiTaT: On rotting petioles of Heliconia sp. in lowland and mountain

rain forests (alt. of investigated specimens from ca 250–2000 m asl).

139

Known diSTribuTion: Ecuador (own collections, Singer 1974), Panama (new record),

and Venezuela (Singer 1974).

SpeCimenS examined: eCuador, prov. Orellana, Añangu, south of Río Napo, hunting trail E of CEPE

building, 0°31'S/76°23'W, 260–350 m asl, tropical rain forest (terra ﬁrme), on rotten leaves of

Heliconia sp., 02.II.1983, T.Læssøe, AAU 43112 (C). – panama, prov. Bocas del Toro, El Valle,

Finca Celestine, 8°47,1'N/82°11,3'W, ca 600 m asl, on rotting petioles of Heliconia sp., 17.XII.2004,

R.Mangelsdorff, RMP2163 (PMA, M). – Prov. Chiriquí, Lagunas de Volcán, ca 1300 m asl, on rotting

Fig. 13: Favolaschia heliconiae. (A) Optical longitudinal section through the basidioma (KG0011).

Scale bar = 1 mm. (B) Pore edge (KG0011). Scale bar = 10 µm. (C) Subhymenial hyphae and basidia

(AAU 43112). Scale bar = 10 µm. (D) Basidia (AAU 43112). Scale bar = 5 µm. (E) Spores in lateral

and front view. From left to right: 1 (KG0030), 2–7 (AAU 43112). Scale bar = 5 µm.

140

petioles of Heliconia sp., 22.VII.2009, K.Gillen, KG0026 (PMA, M). – Parque Internacional La

Amistad, Sendero de Cerro Picacho, ca 2000 m asl, on rotting petioles of Heliconia sp., 18.VII.2009,

K.Gillen, KG0011 and KG0012 (PMA, M). – Sendero El Retoño, ca 2000 m asl, on rotting petioles

of Heliconia sp., 28.VII.2009, K.Gillen, KG0030 (PMA, M).

noTeS: The basidiomata of the type collections of F. heliconiae and F. oligogloea are

morphologically similar, the only differing character being the slightly shorter stipes

of the basidiomata of F. oligogloea, which probably is due to ontogenetic factors.

Fig. 14: Favolaschia heliconiae. (A) Optical longitudinal section through the pileipellis (KG0011).

Scale bar = 5 µm. (B) Acanthocystida (AAU 43112). Left: optical longitudinal sections, right: three-

dimensional views. Scale bar = 5 µm. (C) Stipe margin (KG0011). Scale bar = 10 µm.

141

Thus, they represent with great likelihood the same species. As both species names

have been published in the same document (Singer 1974), the page number has been

used as a crucial criterion, and thus F. heliconiae is the valid name.

Favolaschia macropora K.Gillen, sp. nov. – MycoBank 563466 Figs 1H, 15

Pileus ad 1.7 mm diam., semiglobosus, pallide brunneus ad pallide griseo-olivaceus, 2–6 poris.

Stipes absens. Sporae (7–)7.5–9(–9.5) × (4.5–)5–6.5(–7) µm, amygdaliformes vel ellipsoideae,

amyloideae. Basidia 4 sterigmatibus. Acanthocystida ad os pori et pileipellem insidentia, oblonga,

(11.5–)15.5–27.5(–35) × (5–)5.5–8(–10) µm; gloeocystidia absentia; ﬁbulae absentes.

TypeS: panama, prov. Chiriquí, Lagunas de Volcán, ca 1300 m asl, on decaying stem of a bamboo

species, 22.VII.2009, K.Gillen, M.Piepenbring and students of UNACHI, KG0027 (PMA holotype!,

M isotype!).

eTymology: The epitheton refers to the unusually large pores.

Observations are based on dried material, as the basidiomata were found under rather

dry conditions in the ﬁeld.

Pileus (0.4–)0.7–1.3(–1.4) × (0.7–)0.9–1.4(–1.7) mm (n=15, c=1), semiglobose, pale

brown to pale grey brown, glabrous or rarely slightly pulverulent; margin crenulate and

straight. Hymenophore consisting of 2–6 pores; pore edges with the same colour and

surface as pileus, thin to medium broad. Pores (0.3–)0.35–0.65(–0.8) × (0.2–)0.25–0.4

(-0.5) mm (n=15, c=1), mostly round. Stipe absent.

Spores (7–)7.5–9(–9.5) × (4.5–)5–6.5(–7) µm, Q=(1.2–)1.3–1.6(–1.8) (n=25, c=1),

amygdaliform to ellipsoid, rarely oblong or broadly ellipsoid, smooth, thin-walled,

multiguttulate or with a single large droplet, amyloid. Basidia (24–)29–39.5(–46) ×

(7–)8.5–10.5(–11) µm (n=25, c=1), cylindric to clavate, 4-spored. Acanthocystida

numerous on pore edges and pileipellis, oblong, (11.5–)15.5–27.5(–35) × (5–)5.5–8

(–10) µm (n=15, c=1), with numerous rod-shaped outgrowths covering the entire cell,

wall thin, colourless. Gloeocystidia absent. Gloeoplerous hyphae few, scattered in

trama and the pore edges, contents thromboplerous (pale grey in H2O and KOH). Trama

not or slightly gelatinous, hyphae irregularely arranged in pileus trama, ± parallel in

pore wall and stipe trama, thin-walled.

All septa without clamps.

SubSTraTe and HabiTaT: On decaying stem of a bamboo species in rain forest (alt. of

investigated specimen ca 1300 m asl).

Known diSTribuTion: Panama (only from type collection).

noTeS: This species is characterized by basidiomata with a few large pores and the

absence of clamps and gloeocystidia. The species otherwise resembles the clamped

F. alsophilae Singer, both in morphology (the few large pores) and anatomy (spore size,

basidium size). Differing characters are the lacking clamps and the slightly narrower

acanthocystida in F. macropora [(5–)5.5–8(–10) µm; F. alsophilae: (9–)10–13.5(–15) µm].

As there are no sequence data available for F. alsophila, a molecular-phylogenetic

relationship could not be inferred. However, ITS and LSU sequences of F. macropora

were obtained in this study.

142

Favolaschia pantherina Singer, Beihefte Nova Hedwigia 50: 103 (1974) Figs 1I, 16

Type: HoloType: argenTi na, Misiones, Igazu, Refugio Yaguareté, on Chusquea sp.,

15.IV.1957, R.Singer, M1003 (F!).

Fig. 15: Favolaschia macropora (type material, KG0027). (A) Optical longitudinal section through

the pileus. Scale bar = 10 mm. (B) Section of the pileipellis and hymenium. Scale bar = 50 µm. (C)

Acanthocystida: optical longitudinal section (left) and three-dimensional view (right). Scale bar =

10 µm. (D) Spores in lateral and front view (right). Scale bar = 10 µm. (E) Basidia. In the middle:

with young spores. Scale bar = 10 µm. (F) Hyphae. Clamps are not present. Scale bar = 10 µm.

143

Pileus 0.5–2 × 1–2 mm (n=7), plano-convex to convex, pale brown to pale grey,

glabrous or slightly pulverulent, margin crenulated and sometimes involute, outline

of pores visible on sterile carpophore surface, colour of depressions slightly darker

than colour of pores. Hymenophore consisting of 25–45 pores; pore edges with same

Fig. 16: Favolaschia pantherinae (type material, M1003). (A) Optical longitudinal section through

a basidioma. Scale bar = 1 mm. (B) Basidia and spores. Scale bar = 10 µm. (C) Acanthocystida.

Scale bar = 10 µm.

144

colour and surface as pileus, mostly thin. Pores 0.3–0.9 × 0.2–0.6 mm (n=15), round

or radially elongated. Stipe present and up to 1 mm long or absent.

Spores (8–)8.5–10(–11) × (6–)6.5–8.5(–9) µm, Q=1.1–1.4(–1.5) (n=25), broadly

ellipsoid to ellipsoid, smooth, thin-walled, colourless, rarely multiguttulate, amyloid.

Basidia (19–)22–30(–35) × (9–)10–14.5(–16) µm (n=25), ellipsoid to cylindric,

4-spored. Acanthocystida numerous in pileipellis, on pore edges and stipe margin,

often breaking off during drying process and forming the pulverulent covering of

the pileus, ellipsoid or clavate, often very long, 22–75(–130) × (8–)8.5–11(–12) µm

(n=15), with numerous rod-shaped outgrowths covering the entire cell, colourless.

Gloeocystidia and gloeoplerous hyphae absent. Trama slightly gelatinous, hyphae

irregularely arranged in pileus trama, ± parallel in trama of pore walls and stipe, thin-

walled.

SubSTraTe and HabiTaT: On Chusquea sp. in rainforests.

Known diSTribuTion: Argentina (Singer 1974).

noTeS: Favolaschia pantherina is characterized by the often rather long acanthocystida

and the absence of gloeocystidia.

Favolaschia pterigena var. boliviana Singer, Beihefte Nova Hedwigia 50: 101 (1974)

Figs 1J, 17

Type: HoloType: bolivia, departamento La Paz, Nor-Yungas, Carmen Pampa, on petioles of a tree

fern, 1.II.1956, R.Singer, B817 (F!).

Pileus 1–1.5 mm diam. (n=10), plano-convex, pale brown to greyish brown, glabrous

or slightly pulverulent, margin crenulated and sometimes involute, outline of pores

visible on sterile carpophore surface. Hymenophore consisting of 15–35 pores; pore

edges with same colour and surface as pileus, mostly thin. Pores 0.1–0.3 mm in diam.

(n=15), of irregular shape. Stipe present and up to 2 mm long, mostly glabrous, laterally

attached to the pileus.

Spores (6–)7–8.5(–9) × (4–)4.5–6.5(–7) µm, Q=(1.2–)1.3–1.6(–1.8) (n=25), ellipsoid to

amygdaliform, rarely broadly ellipsoid or oblong, smooth, thin-walled, multiguttulate

or rarely without guttules, amyloid. Basidia (19–)20.5–26.5(–30) × (6–)7–9(–10) µm

(n=25), ellipsoid to cylindric, 4-spored. Acanthocystida numerous in pileipellis and

on pore edges, scattered on stipe margin, often breaking off during drying process and

forming the pulverulent covering of the pileus, broadly ellipsoid or globose, (15–)18–

30(–34) × (6–)7–17.5(–20) µm (n=15), with numerous rod-shaped outgrowths covering

the entire cell, colourless. Gloeocystidia numerous on pileipellis, and scattered among

basidia, with numerous small or slightly bigger droplets (greenish grey in H2O and

KOH), vermiform, (60–)75–135(–160) × (6–)7–10 µm (n=15), thin-walled. Trama

slightly gelatinous, hyphae irregularely arranged in pileus trama, ± parallel in trama

of pore walls and stipe, thin-walled.

SubSTraTe and HabiTaT: On dead petioles of a tree fern (species not identiﬁed), known

altitude ca 2000 m asl.

Known diSTribuTion: Bolivia and Venezuela (both Singer 1974).

145

noTeS: Favolaschia pterigena var. boliviana is characterized by numerous acantho-

cystida in the pileipellis and on the pore edges and by the many scattered vermiform

gloeocystidia in the pileipellis, the trama, and among the basidia.

Fig. 17: Favolaschia pterigena var. boliviana (type material, B817). (A) Optical longitudinal section

through a basidioma. Scale bar = 1 mm. (B) Basidia and spores. Scale bar = 10 µm. (C) Acanthocystida.

Scale bar = 10 µm. (D) Gloeocystidia. Scale bar = 10 µm.

146

Favolaschia roseogrisea Singer, Beihefte Nova Hedwigia 50: 103 (1974) Figs 1K, 18

Type: HoloType: Columbia, departamento Tolima, Ruta de Cajamarca a Calana, on bamboo,

11.IV.1968, R.Singer, B6035 (F!).

Pileus 0.5–1.5 × 0.5–2 mm diam. (n=10), applanate to convex, pale brown, but colour

often hidden under the pulverulent covering, margin mostly crenulated and sometimes

involute. Hymenophore consisting of 20–45 pores; pore edges with same colour and

surface as pileus, medium-thick to thin. Pores 0.1–0.3 mm in diam. (n=15), of irregular

shape. Stipe rudimentary and up to 0.5 mm long or absent.

Spores (8–)8.5–10 × (6–)6.5–8 µm, Q=(1.1–)1.2–1.4(–1.5) (n=25), broadly ellipsoid to

ellipsoid, smooth and thin-walled, multiguttulate, amyloid. Basidia (24–)27–33.5(–42)

× (7–)8.5–10.5(–12) µm (n=25), ellipsoid to cylindric, 4-spored, sterigmata often long.

Acanthocystida numerous in pileipellis, on pore edges and stipe margin, often breaking

off during drying process and forming the pulverulent covering of the pileus, ellipsoid or

clavate, (18–)21–36.5(–40) × (8–)8.5–12.5(–15) µm (n=15), with numerous rod-shaped

outgrowths covering the distal third of the cell, colourless. Gloeocystidia scattered on

pileipellis, pore wall trama and stipe margin, with numerous small or slightly bigger

droplets (light grey in H2O and KOH), vermiform, (50–)65.5–132(–150) × 4–7(–8) µm

(n=15), thin-walled. Trama slightly gelatinous, hyphae irregularely arranged in pileus

trama, ± parallel in trama of pore walls and stipe, thin-walled.

Most cells clamped.

SubSTraTe and HabiTaT: On dead stems of Bambuseae (Guadua angustifolia Kunth).

Known diSTribuTion: Columbia and Trinidad (all Singer 1974).

noTeS: Favolaschia roseogrisea is characterized by the frequent presence of a more

pulverulent covering of the pileipellis and the pore edges than the other examined

specimens as well as by unsually long sterigmata. An interesting character is the

inconsistent presence of clamps.

Favolaschia sabalensis var. geonematis Singer, Beihefte Nova Hedwigia 50: 101

(1974) Figs 1L, 19

Type: HoloType: eCuador, prov. Napo, Shushuﬁndi, on Geonoma sp., 15.V.1973, R.Singer, B7441

(F!).

Pileus 0.5–1 mm in diam. (n=8), plano-convex to convex, beige to pale brown, glabrous,

margin crenulated and not involute, applanate to convex, pale brown, but colour often

hidden under the pulverulent covering, margin mostly crenulated and sometimes

involute, outline of pores visible on sterile carpophore surface. Hymenophore

consisting of 3–10 pores; pore edges with same colour and surface as pileus, and thin.

Pores 0.2–0.4 × 0.2–0.6 mm (n=15), round or of irregular shape. Stipe rudimentary

and up to 0.5 mm long or absent.

Spores 11–13.5(–15) × 9–11(–12) µm, Q=1.2–1.3 (n=9), broadly ellipsoid, smooth and

medium thick-walled, without guttules or multiguttulate. Basidia 34–38 × 13–14 µm

(n=4), mostly clavate, two-spored. Acanthocystida scattered in pileipellis and mostly

numerous on pore edges, oblong, (20–)21.5–37.5(–41) × (4–)6–9 µm (n=15), with

147

numerous rod-shaped outgrowths covering the entire cell, colourless. Gloeocystidia

numerous in pileipellis and on pore edges, with numerous small or slightly bigger

droplets (greenish grey in H2O and KOH), broadly clavate or ellipsoid, 28–56(–80)

× (16–)17.5–26(–32) µm (n=15), thin-walled. Trama slightly gelatinous, hyphae

irregularely arranged in pileus trama, more or less parallel in trama of pore walls and

stipe, thin-walled.

Fig. 18: Favolaschia roseogrisea (type material, 6035). (A) Sterile surface of basidiomata. Scale bar

= 1 mm. (B) Spores and basidia. Scale bar = 50 µm. (C) Acanthocystida. (D) Gloeocystidium. (E).

Hyphae. Scale bars (C, D, E) = 50 µm.

148

SubSTraTe and HabiTaT: On stem of Geonoma sp. in rain forests.

Known diSTribuTion: Ecuador (Singer 1974).

noTeS: Only few spores (9) and mature basidia (4) have been found in the material.

Due to the small number of spores it could not be observed whether they are amyloid

Fig. 19: Favolaschia sabalensis var. geonematis (type material, B7441). (A) Optical longitudinal

section through a basidioma. Scale bar = 1 mm. (B) Spores and basidia. Scale bar = 10 µm.

149

or not since the colour change of Favolaschia spores in Melzer’s reagence is mostly

faint. Two-spored basidia are an infrequent character among species of the genus

Favolaschia.

Favolaschia teapae Singer, Beihefte Nova Hedwigia 50: 102 (1974) Figs 1M, 20

Type: HoloType: mexiCo, Río Teapa, between Ixtacomitán and Solusuchiapa, on dead twigs of a

dicotyledonous plant, 1.VIII.1969, R.Singer, M8798 (F!)

Pileus 1–3 × 1.5–3.5 mm (n=9), applanate to plano-convex, pale orange to pale brown,

glabrous, margin mostly crenulate. Hymenophore consisting of 30–70 pores; pore

edges with same colour and surface as pileus, thin. Pores 0.1–0.5 × 0.1–0.4 mm (n=15),

round or of irregular shape. Stipe absent.

Spores 7–8.5(–9) × 4–4.5(–5) µm, Q=1.8–2 (n=4), oblong, smooth, colourless. Basidia

20–25(–26) × 7–8.5(–9) µm (n=10), clavate to cylindric, 4-spored. Acanthocystida

mostly numerous in pileipellis and on pore edges, distinct stipe, globose to clavate,

(14–)16–23.5(–28) × (14–)16.5–21.5(–24) µm (n=15), with numerous rod-shaped

outgrowths covering the entire cell (apart from stipe), wall thickened below the

ornamented part, colourless. Gloeocystidia scattered in the trama of the pileus and

pore walls, with numerous small or slightly bigger droplets, vermiform, (70–)75–130

(–150) × 4–6(–7) µm (n=15), thin-walled. Trama not or slightly gelatinous, hyphae

irregularely arranged in pileus trama, more or less parallel in trama of pore walls and

stipe, thin-walled.

SubSTraTe and HabiTaT: On dead twigs of dicotyledonous plants.

Known diSTribuTion: Bolivia, Mexico, and Venezuela (all Singer 1974).

noTeS: Only few spores (4) and mature basidia (10) have been found in the material.

Due to the small number of spores it could not be observed whether they are amyloid

or not. The investigated basidiomata resemble those of F. andina. However, the few

observed spores are smaller and the Q value is higher.

Favolaschia violascens Singer, Beihefte Nova Hedwigia 50: 103–104 (1974) Figs 1N, 21

Type: HoloType: bolivia, departamento La Paz, Nor-Yungas, Carmen Pampa, on twigs of a

dicotyledonous plant, 17.II.1956, R.Singer, B1227 (F!).

Pileus 0.5–1.5 × 0.5–2 mm (n=10), applanate to plano-convex, pale brown to greyish

brown, but fruit body colour often hidden under a slightly pulverulent covering, margin

mostly involute. Hymenophore consisting of 15–100 pores; pore edges with same

colour and surface as pileus, medium-thick. Pores 0.1–0.3 × 0.1–0.2 mm (n=15), round

or rarely of irregular shape. Stipe present and up to 1 mm long or absent.

Spores (8–)8.5–10.5(–12) × (4–)5–6.5(–8) µm, Q=(1.4–)1.6–2(–2.5) (n=15),

oblong, rarely ellipsoid, smooth, thin- to slightly thicker walled, without guttules or

multiguttulate (greenish grey in H2O and KOH). Basidia 23–35 × 5–9 µm (n=5), clavate

or rarely cylindric, two-spored. Acanthocystida numerous in pileipellis and on pore

edges, often breaking off during drying process and forming the pulverulent covering

of the pileus, globose to broadly clavate, (11–)11.5–19(–24) × (7–)9.5–15(–16) µm

150

(n=15), with numerous rod-shaped outgrowths covering the whole cell, wall thickened

below the ornamented part, colourless, amyloid. Gloeocystidia numerous in pileipellis,

stipe surface and on pore edges, scattered in the trama of the pileus and stipe, with

numerous small or slightly bigger droplets (greenish grey in H2O and KOH), clavate to

vermiform, (80–)95–145(–155) × (8–)9.5–13.5(–14) µm (n=15), thin-walled. Trama

Fig. 20: Favolaschia teapae (type material, M8798). (A) Optical longitudinal section through a

basidioma. Scale bar = 1 mm. (B) Spores and basidia. Scale bar = 10 µm. (C) Acanthocystida. Scale

bar = 10 µm. (D) Gloeocystidia. Scale bar = 10 µm.

151

slightly to very gelatinous, hyphae irregularly arranged in pileus trama, more or less

parallel in trama of pore walls and stipe, thin to slightly thicker walled.

SubSTraTe and HabiTaT: On dead twigs of dicotyledonous plants.

Known diSTribuTion: Bolivia (Singer 1974).

Fig. 21: Favolaschia violascens (type material, B1227). (A) Optical longitudinal section through

a basidioma. Scale bar = 0.5 mm. (B) Spores and basidia. Scale bar = 10 µm. (C) Acanthocystida.

Scale bar = 10 µm. (D) Gloeocystidia. Scale bar = 50 µm.

152

noTeS: Only few spores (15) and mature basidia (5) have been found in the investigated

specimens. It was not possible to observe whether the spores are amyloid or not.

Molecular phylogeny

A neighbour joining analysis was conducted, including the LSU rDNA sequences

obtained in the context of the present study from ﬁve species from Panama (Favolaschia

andina, F. aurantiaca, F. dealbata, F. heliconiae, and F. macropora) and ten LSU rDNA

sequences selected by a BLAST similarity search in GenBank with a similarity of >95%

(F. calocera R.Heim, F. cinnabarina, F. peziziformis, F. cf. sprucei (Berk.) Singer,

Mycena crocata (Schrad.) P.Kumm., M. galericulata (Scop.) Gray, and Poromycena

manipulata (Berk.) R.Heim). Fig. 22 shows the optimal tree. However, it is important

to mention that this is a very preliminary data set with mainly low bootstrap values

due to the inclusion of only a small number of the known species of Favolaschia.

Mycena crocata and M. galericulata were chosen as an outgroup. The sequences

were aligned in 414 positions. The sequences obtained from the material from

Panama form a clade with the Favolaschia sequences from GenBank. Though this

is a preliminary analysis, the following subgroups can be distinguished: "group 1"

comprising F. aurantiaca and F. cinnabarina (though showing only few morphological

and anatomic similarities) and "group 2" comprising F. dealbata and F. calocera (both

species characterized by the orange colour and the arrangement of the gloeocystidia

in the pileipellis); however, this group is not well supported. F. macropora is placed

within the Favolaschia clade, though morphological and anatomical similarity to the

other included species is low (combination of characteristics: few large pores of the

basidiomata, absence of both clamps and gloeocystidia).

Fig. 22: Molecular-phylogenetic tree with LSU rDNA sequences. Mycena crocata and M. galericulata

were chosen as outgroup. The obtained sequences of the collected material from Panama are indicated

with bold letters. The numbers after the species names indicate the GenBank accession numbers of

the sequences. Only bootstrap values > 50% are shown.

153

Synoptic key

A key which for the ﬁrst time comprises available data on the published taxa names of

the genus Favolaschia (see Table 2) was created based on the own investigations and

an intensive study of the literature. Additionally, it contains comprehensive species

descriptions and illustrations of 14 investigated type specimens. The key as well as

a manual are provided on the website http://species-id.net/wiki/key_to_Favolaschia_

(Kerstin_Gilen) and can be used with the program DELTA (Dallwitz 1974 and 1980,

Dallwitz et al. 1993).

Analytical key

The present chapter includes a dichotomous key to the species documented in

the present publication, i.e. 13 of 145 known species of Favolaschia. Most of the

remaining species are incompletely described in the literature, so they cannot be

included in a dichotomous key. More specimens, especially type material, have to be

investigated in detail in order to establish a more comprehensive dichotomous key. For

a synoptic key including all species names see http://www.mycokey.com/Favolaschia/

FavolaschiaStart.html

1. Gloeocystidia absent ................................................................................................................. 2

Gloeocystidia present................................................................................................................ 3

2. Hyphae with clamps ................................................................................................................. 4

Hyphae without clamps ........................................................................................ F. macropora

4. Fruitbodies on tree ferns and with ≤ 10 pores ..................................................... F. pantherinae

Fruitbodies on other substrates and with > 10 pores ............................................. F. alsophilae

3. Majority of basidia two-spored ................................................................................................. 5

Majority of basidia four-spored ................................................................................................ 7

5. Fruitbodies at least in dried condition very pulverulent ........................................ F. violascens

Fruitbodies in dried condition not or only slightly pulverulent ................................................ 6

6. Spore width (5.5–)6–7.5(–8) µm ........................................................................ F. cinnabarina

Spore width (4–)4.5–6.5 µm .................................................................................... F. gaillardii

7. Fruitbodies on Heliconia sp. or Chusquea sp. .......................................................................... 8

Fruitbodies on other substrates ............................................................................................... 10

8. On Heliconia sp. .................................................................................................... F. heliconiae

On Chusquea sp. ....................................................................................................................... 9

9. Fruitbodies in fresh condition orange, not or little gelatinous .................................. F. dealbata

Fruitbodies in fresh condition white to brown, rather gelatinous ......................... F. aurantiaca

10. Spore length ≥10 µm .................................................................... F. sabalensis var. geonematis

Spore length <10 µm ............................................................................................................... 11

11. Fruitbodies on tree ferns ................................................................... F. pterigena var. boliviana

Fruitbodies on other substrates ............................................................................................... 12

12. Gloeocystidia present among the basidia, fruitbodies glabrous or only slightly pulverulent .......

..................................................................................................................................... F. andina

Gloeocystidia present, but not among the basidia, fruitbodies very pulverulent . F. roseogrisea

154

Table 2. Synoptic key: the comprised species names, known distributions and references.

Species name and synonyms Collection site(s) Reference(s)

Favolaschia agaricina (Mont.)

Kuntze, Exidia

agaricina Mont.

Brazil Bresadola 1896, Kuntze 1898

Favolaschia alsophilae Singer Colombia, Ecuador this work, Singer 1974

Favolaschia amoene-rosea Henn.

[as 'amoene rosea'] Argentina, Bolivia,

Brazil Hein 1988, Hennings 1904, Saccardo

1905, Singer 1950, 1974

Favolaschia andina Singer Colombia, Ecuador,

Panama this work, Singer 1974

Favolaschia antarctica (Speg.)

Kuntze, Laschia antarctica Speg. Argentina, Ecuador,

South Georgia,

Venezuela

Dennis & James 1960, Dennis 1970,

Kuntze 1898, Pegler et al. 1980, Singer

1974, Spegazzini 1887, Læssøe

& Petersen 2008

Favolaschia aulaxina (Mont.)

Singer, Agaricus aulaxinus Mont. Chile Montagne 1850

Favolaschia aurantiaca Singer Argentina, Bolivia,

Ecuador, Panama this work, Singer 1974

Favolaschia auriscalpium (Mont.)

Henn., Laschia auriscalpium Mont. Cameroon, Cuba,

French Guiana,

Venezuela

Berkeley & Curtis 1868, Dennis 1970,

Hennings 1897a, Pegler 1987, Singer

1974

Favolaschia austrocyatheae

P.R.Johnst. New Zealand Johnston et al. 2006

Favolaschia baumanniana

Henn. Togo Hein 1988, Hennings 1897b

Favolaschia bibundensis Henn. Cameroon Hein 1988, Hennings 1897a

Favolaschia bispora Holterm. Indonesia, Sri Lanka Holtermann 1898, Willis 1907–10

Favolaschia brasiliensis Henn. Brazil Hein 1988, Hennings 1897c

Favolaschia caerulescens (Berk.

& M.A.Curtis) Kuntze, Laschia

caerulescens Berk. & M.A.Curtis

Cuba Berkeley & Curtis 1868, Kuntze 1898,

Pegler 1987

Favolaschia caespitosa (Berk.)

Kuntze, Favolus caespitosus Berk.

[as 'caespitosa']

Australia Berkeley & Broome 1873, Kuntze

1898

Favolaschia cagnii Mattir. Uganda Kobayasi 1981, Mattirolo 1908–9

Favolaschia calamicola Henn. &

E.Nyman Indonesia Hein 1988, Hennings 1899, Saccardo

1902

Favolaschia calocera R.Heim,

F. calocera var. calocera R.Heim,

F. calocera R.Heim var. claudopus

Singer

Australia, China,

Italy, Kenya,

Madagascar, New

Zealand, Norfolk

Island, Réunion

Island, Thailand

e.g. Heim 1945, 1966, Johnston et al.

2006, Singer 1974, Vizzini & Zotti

2002, Vizzini et al. 2008

Favolaschia cantharelloides Pat. Java Patouillard 1897

Favolaschia celebensis (Pat.)

Kuntze, Laschia celebensis Pat. Java Kuntze 1898, Patouillard 1887

155

Favolaschia cinereopruinosa

(Kalchbr.) Kuntze, Laschia

cinereopruinosa Kalchbr.

Australia Kalchbrenner 1876, Kuntze 1898

Favolaschia cinnabarina (Berk.

& M.A.Curtis) Pat., Laschia

cinnabarina Berk. & M.A.Curtis

Bolivia, Brazil,

Colombia, Cuba,

Ecuador, Guyana,

Panama, U.S.A.,

Venezuela

e.g. this work, Berkeley & Curtis 1868,

Dennis 1961, 1970, Pegler 1987, Rick

1938 Singer 1945, 1974, 1988

Favolaschia citrina Kobayasi Papua New Guinea Kobayasi 1981

Favolaschia citrinella Henn. Cameroon, Republic

of Congo Hein 1988, Hennings 1897b, Saccardo

1902

Favolaschia clypeata (Pat.) Kuntze,

Laschia clypeata Pat. Venezuela Kuntze 1898, Patouillard 1887

Favolaschia congolensis

(De Seynes) Pat. Madagascar De Seynes 1897, Patouillard 1928

Favolaschia curtisii (Berk.) Kuntze,

Laschia curtisii Berk. Cuba Berkeley & Curtis 1868, Kuntze 1898,

Pegler 1987

Favolaschia cyatheae P.R.Johnst. New Zealand Johnston et al. 2006

Favolaschia dealbata Singer Ecuador, Panama this work, Singer 1974

Favolaschia decurrens (Berk. &

M.A.Curtis ex Cooke) Kuntze,

Laschia decurrens Berk. &

M.A.Curtis ex Cooke

Venezuela Cooke 1891, Kuntze 1898

Favolaschia dumontii Singer Venezuela Singer 1974

Favolaschia dybowskyana (Singer)

Singer, Favolaschia thwaitesii

subsp. dybowskyana Singer

Kenya, Mexico,

Republic of Congo,

Uganda, Zambia

Jonsson & Nylund 1979, Pegler &

Rayner 1969, Singer 1945, Singer

1974

Favolaschia echinata Singer Argentina, Venezuela Singer 1950, 1974

Favolaschia fendleri Singer Belize, Brazil,

Colombia, Honduras e.g. this work, Singer 1974

Favolaschia ﬁlopes Singer &

O.Fidalgo Brazil, Venezuela Singer 1974, Singer & Fidalgo 1965

Favolaschia ﬂabellulum (Berk.

& M.A.Curtis ex Cooke) Kuntze,

Laschia ﬂabellulum Berk. &

M.A.Curtis ex Cooke

Venezuela Cooke 1891

Favolaschia lava (Bres.) Singer,

Laschia lava Bres. Argentina, Bolivia,

Brazil, Panama

Trinidad and Tobago,

Venezuela

Bresadola 1896, Dennis 1970, Singer

1950, 1974

Favolaschia frieseana Henn. Cameroon Hein 1988, Hennings 1897a

Favolaschia fujisanensis Kobayasi Japan Kobayasi 1952

Favolaschia furfurella Singer Venezuela Singer 1974

Favolaschia gaillardii (Pat.) Pat.

[as 'gaillardi'], Laschia gaillardii

Pat.

Brazil, Ecuador,

Panama, Peru,

Venezuela

e.g. this work, Dennis 1961, 1970,

Patouillard 1887, Singer 1945, 1974,

Læssøe & Petersen 2008b

Favolaschia goetzei Henn. Malawi Hein 1988, Hennings 1900, Saccardo

1902

156

Favolaschia grandiuscula Syd. New Guinea Sydow & Sydow 1916

Favolaschia helena Otieno Kenya Otieno 1964

Favolaschia heliconiae Singer,

F. oligogloeoa Singer Ecuador, Panama,

Venezuela this work, Singer 1974, Læssøe &

Petersen 2008b

Favolaschia holtermannii Henn. [as

'holtermanni'] Indonesia e.g. Holtermann 1898

Favolaschia infundibuliformis

(Berk.) Kuntze, Laschia

infundibuliformis Berk.

Brazil Berkeley 1842, Kuntze 1898

Favolaschia intermedia (Berk.

& M.A.Curtis) Singer, Laschia

intermedia Berk. & M.A.Curtis,

Favolaschia intermedia subsp.

intermedia Singer

Brazil, Colombia,

Cuba, Venezuela Berkeley & Curtis 1868, Pegler 1987,

Singer 1969, 1974

Favolaschia intermedia subsp.

pulverulenta var. dennisiana Singer Venezuela Singer 1974

Favolaschia intermedia subsp.

pulverulenta var. dicotyledonea

Singer

Bolivia, Trinidad and

Tobago, Venezuela Singer 1974

Favolaschia intermedia subsp.

pulverulenta var. pulverulenta

Singer

Brazil, Venezuela Singer 1974

Favolaschia intermedia subsp.

pulverulenta var. tetraspora Singer Venezuela Singer 1974

Favolaschia javanica Holterm. Indonesia, Java Holtermann 1898, Kobayasi 1981

Favolaschia keniae Otieno Kenya Otieno 1964

Favolaschia kimakiensis Otieno Kenya Otieno 1964

Favolaschia lamellosa (Pat. &

Gaillard) Kuntze, Laschia lamellosa

Pat. & Gaillard

Venezuela Patouillard & Gaillard 1888, Saccardo

1891

Favolaschia lateritia Henn. Cameroon Hein 1988, Hennings 1897a, Singer

1974

Favolaschia lauterbachii (Henn.)

Kuntze, Laschia lauterbachii Henn. Papua New Guinea Hennings 1894, Kuntze 1898

Favolaschia ledermannii Syd. New Guinea Sydow & Sydow 1916

Favolaschia longipes (Berk.)

Kuntze, Laschia longipes Berk. South America Berkeley 1878

Favolaschia lurida (Ces.) Kuntze Borneo, Sri Lanka De Cesati 1879

Favolaschia magniﬁca Henn. Brazil, Venezuela Hein 1988, Hennings 1897c

Favolaschia macropora K.Gillen Panama this work

Favolaschia mainsii Singer Belize Singer 1974

Favolaschia manjaniana Otieno Kenya Otieno 1964

Favolaschia meridae Singer Venezuela Singer 1974

Favolaschia micropus (Berk.)

Kuntze, Laschia micropus Berk. Australia Berkeley & Curtis 1873

157

Favolaschia moelleri (Bres.) Singer,

Laschia moelleri Bres. Bolivia, Brazil,

Colombia, Ecuador,

Peru

Bresadola 1896, Singer 1969, 1974,

Læssøe & Petersen 2008b

Favolaschia montana Singer Argentina Singer 1974

Favolaschia nigrostriata Henn. &

E.Nyman Indonesia, Java Hein 1988, Hennings 1899, Saccardo

1902

Favolaschia nipponica Kobayasi Japan Kobayasi 1952

Favolaschia oligopora Singer Colombia, Ecuador Singer 1974, Læssøe & Petersen 2008b

Favolaschia pantherina Singer Argentina this work, Singer 1974

Favolaschia papuana Kobayasi Papua New Guinea Kobayasi 1981

Favolaschia papulata (Mont.)

Singer, Laschia papulata Mont. Chile Montagne 1850, Singer 1969

Favolaschia pegleri Parmasto Russia Parmasto 1999

Favolaschia peziziformis (Berk.

& M.A.Curtis) Kuntze, Laschia

peziziformis Berk. & M.A.Curtis

[as 'pezizaeformis']

Japan, Venezuela Eaton et al. 1858, Kobayasi 1952,

Kuntze 1898, Singer 1945

Favolaschia pezizoidea (Berk.

& M.A.Curtis) Kuntze, Laschia

pezizoidea Berk. & M.A.Curtis

Cuba, Ecuador Berkeley & Curtis 1868, Kuntze 1898,

Læssøe & Petersen 2008b, Singer

1945, 1974

Favolaschia phyllostachydis

Imazeki & Kobayasi Japan Kobayasi 1952

Favolaschia pterigena Singer,

F. pterigena var. pterigena Singer Argentina Singer 1950, 1974

Favolaschia pterigena var. bolivi-

ana Singer Bolivia, Venezuela this work, Singer 1974

Favolaschia pterigena var. purpu-

rea Singer Bolivia Singer 1974

Favolaschia puberula Singer Venezuela Singer 1974

Favolaschia puiggarii (Speg.)

Singer, Favolus puiggarii Speg. Brazil Singer 1950, 1974, Spegazzini 1887

Favolaschia pulverulenta Henn.,

F. intermedia subsp. pulverulenta

(Henn.) Singer, Laschia

pulverulenta (Henn.) Sacc.

Brazil, Venezuela e.g. Hein 1988, Hennings 1897c,

Singer 1974

Favolaschia purpurea (Berk. &

M.A.Curtis) Kuntze, Laschia

purpurea Berk. & M.A.Curtis

Cuba Berkeley & Curtis 1868, Kuntze 1898,

Pegler 1987

Favolaschia pustulata (Berk. &

Broome) Kuntze, Laschia pustulata

Berk. & Broome

Sri Lanka Berkeley & Broome 1873, Kuntze

1898

Favolaschia pustulosa (Jungh.)

Kuntze Borneo, China,

Java, Malaysia, New

Zealand, Papua New

Guinea, Philippines,

Samoa

e.g. Johnston et al. 2006, Junghuhn

1838, Kuntze 1898, Singer 1945, 1974

Favolaschia pygmaea (Speg.)

Singer, Favolus pygmaeus Speg. Argentina, Brazil,

Colombia, Trinidad

and Tobago

Dennis 1970, Singer 1950,

1974, Spegazzini 1887

158

Favolaschia reticulata (Berk. &

Cooke) Kuntze, Laschia reticulata

Berk. & Cooke

Brazil Berkeley & Cooke 1876

Favolaschia rosea Henn. Togo Hein 1988, Hennings 1897b

Favolaschia roseogrisea Singer Colombia, Trinidad

and Tobago this work, Singer 1974

Favolaschia rubra (Bres.) Kuntze,

Laschia rubra Bres. Bolivia, Brazil,

Ecuador, Guyana,

Panama

Bresadola 1896, Kuntze 1898, Læssøe

& Petersen 2008b, Singer 1945, 1974,

1988

Favolaschia sabalensis (Charles)

Singer, Laschia sabalensis Charles,

Favolaschia sabalensis var.

sabalensis (Charles) Singer

Cuba, Panama,

Trinidad and

Tobago, U.S.A.

Charles 1942, Dennis 1970, Singer

1945, 1974

Favolaschia sabalensis var.

geonematis Singer Ecuador this work, Singer 1974, 1988

Favolaschia sacchalinensis

Parmasto ex Singer Russia e.g. Singer 1974

Favolaschia saccharina Pat.,

F. intermedia subsp. saccharina

(Pat.) Singer, Laschia saccharina

(Pat.) Sacc. & P. Syd.

Brazil, Ecuador,

Trinidad and Tobago,

Venezuela

Dennis 1952, 1961, 1970,

Patouillard 1900, Singer 1945 and

1974

Favolaschia sanguinea Henn. Cameroon Hein 1988, Hennings 1901, Saccardo

1905

Favolaschia selloana Henn. Bolivia, Brazil,

Colombia Hein 1988, Hennings

1897c, Singer 1945, 1974

Favolaschia singeriana Dennis,

F. intermedia subsp. singeriana

(Dennis) Singer

Venezuela Dennis 1952, 1970, Singer 1974

Favolaschia sprucei (Berk.) Singer,

Favolus sprucei Berk. Brazil, Colombia,

Costa Rica, Panama,

Venezuela

Dennis 1961, 1970, Gómez 1983,

Singer 1945, 1974

Favolaschia staudtii Henn. Cameroon Hein 1988, Hennings 1897b

Favolaschia subamyloidea Singer Colombia, Venezuela Singer 1974

Favolaschia subcaerulea (Berk.

& M.A. Curtis) Kuntze, Laschia

subcaerulea Berk. & M.A. Curtis

Cuba Berkeley & Curtis 1868, Pegler 1987

Favolaschia subceracea (Henn.)

Donk, Cyphella subceracea Henn. Argentina, Bolivia,

Brazil, Venezuela Donk 1959, Hennings 1897c, Singer

1974

Favolaschia teapae Singer Bolivia, Mexico,

Venezuela Singer 1974

Favolaschia testudinella (R.E.Fr.)

Kuntze, Laschia testudinella R.E.Fr. Sweden (hothouse) Fries (1888)

Favolaschia thwaitesii (Berk. &

Broome) Kuntze, Laschia thwaitesii

Berk. & Broome

Sri Lanka Singer 1974

Favolaschia tonkinensis (Fr.)

Singer, Laschia tonkinensis Pat. Cameroon, Kenya,

Philippines,

Switzerland

(hothouse), Uganda,

Vietnam, Zaire

e.g. Pegler & Rayner 1969, Singer

1945, 1974, Wilhelm 2009

159

Discussion

According to Index Fungorum 145 Favolaschia species names have been published.

However, presumably this value is too high. Several collections, that were described

as different species belong with great likelihood to the same species, as in the case of

F. heliconiae and F. oligogloea. This happens due to the fact that new material was

not compared to type material and as the knowledge on distinguishing characters was

lacking to a great extent. Several specimens were described as species belonging to the

now invalid genus Laschia. Kuntze (1898) transferred them without closer investigation

to Favolaschia, hence, several of those species probably belong to other genera than

Favolaschia. Concurrently, species richness presumably is greater than presently

known. Large parts of the tropical areas, where a diversity hot spot of the species is

assumed, presently is only poorly investigated. During ﬁeldwork in Panama, which

was carried out during four weeks only, one new species was found.

Species delimitation within the genus is difﬁcult and presently insufﬁciently clariﬁed.

The combination of the aspects discussed in the following allows however a rather

accurate delimitation. Several species are stated to have a high substrate specificity

(i.e. F. heliconiae), others have been found on diverse substrate types (i.e. the well

Favolaschia torrendii (Lloyd)

Singer, Laschia torrendii Lloyd Brazil Lloyd 1919, Singer 1974

Favolaschia valparadisiensis Henn. Chile Hein 1988, Hennings 1900, Saccardo

1902

Favolaschia varariotecta Singer Colombia, Ecuador,

Grenada, Panama,

Puerto Rico,

Trinidad and Tobago,

Venezuela

Dennis 1952, 1970, Lodge et

al., Læssøe & Petersen 2008b,

Singer 1950, 1974

Favolaschia violascens Singer Bolivia this work, Singer 1974

Favolaschia viridula (Berk. &

M.A.Curtis ex Cooke) Kuntze,

Laschia viridula (Berk. &

M.A.Curtis) Cooke

Venezuela e.g. Cooke 1891

Favolaschia volkensii (Bres.)

Henn., Laschia volkensii Bres. Tanzania Hennings 1900, Singer 1974

Favolaschia vulensis Otieno Kenya Otieno 1964

Favolaschia zenkeri Henn., Laschia

zenkeri (Henn.) Sacc. & Trotter Cameroon e.g. Hennings 1904

Favolaschia zenkeriana Henn. Bolivia, Cameroon,

Liberia Hein 1988, Hennings 1904,

Singer 1945, 1974

Panellus minimus (Jungh.)

P.R.Johnst. & Moncalvo,

Favolaschia minima (Jungh.)

Kuntze, Polyporus minimus Jungh.,

Favolaschia calamicola Henn. &

E.Nyman)

Indonesia, Java,

New Zealand,

Sri Lanka, U.S.A.

Desjardin 1993, Johnston

et al. 2006, Singer 1945

160

investigated species F. calocera). However, mostly the number of investigated

specimens per species is rather low, so speciﬁcities to a certain substrate are doubtful.

Morphological and anatomical aspects used for species delimitation should always

be treated in combination. Characters important to distinguish species of Favolaschia

are basidioma colour and consistency, number of the pores, spore size, number of the

sterigmata per basidium, the presence of clamps, as well as shape, size, and position of

acanthocystida and gloeocystidia. However, single characters can have rather differing

shapes in young and mature basidiomata.

For an improved investigation of the genus it is neccessary to investigate the type

specimens of all the species described and to comprise the data in well-structured,

detailed and illustrated species descriptions as they are given in this work and in earlier

publications, e.g. in Dennis (1952), Parmasto (1999), and Pegler (1987). However,

illustrations showing the arrangements of the different cell types in the basidioma

rarely have been given in earlier publications. In the future the avaible data should be

complemented with respect to macroscopic and microscopic characters of fresh and

dried specimens, substrate, habitat (vegetation type, season, altitude), DNA sequence

data, cultures, and enzyme testing. Morphological and anatomical characters and

data obtained from molecular-phylogenetic investigations supplement each other, as

molecular-phylogenetic trees can only be interpreted correctly based on morphology

and anatomy, while molecular phylogeny shows which morphological and anatomical

characters are reliable to deﬁne species limitations and systematic relationships. The

molecular-phylogenetic investigation shows that the species of which sequences

were obtained for the ﬁrst time belong to the genus Favolaschia. The subsections

Auriscalpium and Dealbatae erected by Singer (1974) apparantly are paraphyletic.

Presently, 47 sequences of Favolaschia species are available in GenBank. Of these,

eight sequences were obtained from material which has only been identiﬁed on genus

level. It is dubious whether the specimens identiﬁed on species level are identiﬁed

correctly. The lack of reliable data shows again how essential it is to investigate type

material both morphologically, anatomically and molecular-phylogenetically. However,

due to the high age of most type specimens (50 to 150 years) and scarce quantity of

the basidiomata, it is difﬁcult to extract DNA. So more sequences of freshly collected

material should be obtained. Numerous further sequences of correctly identiﬁed species

are necessary to make reliable statements of molecular-phylogenetic relationships.

Most publications on Favolaschia species from the end of the 19th/beginning of the

20th century contribute knowledge only on macroscopical characters and contain no

or scarce illustrations. It is therefore to be expected that many superﬂuous names

were introduced. Singer (1974), surprisingly, treated only ﬁve names as synonyms.

The largest work on the genus is the monograph by Singer (1974). However, Singer

overlooked several publications that include Favolaschia names: Fidalgo & Kauffmann-

Fidalgo (1962), Hennings (1897b), Holtermann (1898), Kalchbrenner (1876), Kuntze

(1898), and Otieno (1964). Furthermore, illustrations are lacking or poor, and species

descriptions have mostly only scarce indications on the number of measured structures.

Singer attributes high value to characters such as substrate speciﬁcity, basidioma colour,

and size and shape of certain cell types, but his material was not of a quality to support

his conclusions, and for this reason his keys do not work well.

161

The synoptic key available in the internet is the ﬁrst identiﬁcation key comprising the

available data on all published Favolaschia species names as well as comprehensive

data (including illustrations) on the 14 investigated type specimens. Synoptic keys hold

advantages compared to dichotomous keys. Characters with lacking data can be skipped

without the too early exclusion of species. The key is user-friendly. As all characters

are weighted equally, it is easy to extend it without reorganization. The provision via

the internet makes frequent updates possible. In addition, the key can be used as a

database, and single species can be compared directly with each other. However, the

frequent scarce species descriptions in older publications reduce the quality of the key,

as characters that are important for species delimitations are unknown.

Good identiﬁcation tools have not only an important impact on fundamental research,

but also on applied sciences. Anke et al. (1995, 2004), Nicholas et al. (1997),

Kornsakulkarn et al. (2010), and Wood et al. (1996) found that several Favolaschia

species produce substances with antimycotic or antimalarial effects: favolon,

laschiatron, oudemansines, and strobilurines. However, the investigated species often

were only identiﬁed on genus level, or the identiﬁcation on species level is doubtful,

as good identiﬁcation tools were lacking. Therefore, the published data cannot easily

be reproduced. This clearly highlights the importance of fundamental research for

applied sciences.

Acknowledgements

We thank the curators of the herbaria of the Field Museum of Natural History, Chicago (F), Farlow

Reference Library and Herbarium of Cryptogamic Botany, Harvard, Cambridge (FH), Royal Botanic

Gardens, Kew (K), Statens Naturhistoriske Museum, Botanisk Museum, København (C), and New

York Botanical Garden (NY) for answering requests and loans of specimens. Thanks to the DAAD,

Autoridad Nacional del Ambiente in Panama, and the Universidad Autónoma de Chiriquí who

make the ﬁeldwork in Panama possible. Special thanks to Jens H. Petersen and all collaborators on

the "Fungi of Ecuador" project as well as the students of the UNACHI who helped conducting the

ﬁeldwork. Thanks to Mike J.Dallwitz (DELTA) for his friendly email support. TL acknowledges for

support from the Danish government RUF grant 91056 (ﬁeldwork in Ecuador 2002-2005).

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Three new species of Favolaschia (Mycenaceae, Agaricales) from South China

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Full-text available

Apr 2024

The genus Favolaschia within the family Mycenaceae is characterised by the gelatinous basidiomata with poroid hymenophore and most species inhabit monocotyledonous plants. In this study, many samples covering a wide geographic range in China were examined morphologically and phylogenetically using concatenated ITS1-5.8S-ITS2-nLSU sequence data. Three new species clustering in Favolaschia sect. Anechinus, namely Favolaschia imbricata, F. miscanthi and F. sinarundinariae, are described. Favolaschia imbricata is characterised by imbricate basidiomata with pale grey to greyish colour when fresh and broadly ellipsoid basidiospores measuring 7–9 × 5–6.8 µm; F. miscanthi is characterised by satin white basidiomata when fresh, broadly ellipsoid basidiospores measuring 7.5–10 × 5.5–7 µm and inhabit rotten Miscanthus; F. sinarundinariae is characterised by greyish-white basidiomata when fresh, dark grey near the base upon drying, broadly ellipsoid to subglobose basidiospores measuring 7–9 × 5–7 µm and inhabit dead Sinarundinaria. The differences amongst the new species and their morphologically similar and phylogenetically related species are discussed. In addition, an updated key to 19 species of Favolaschia found in China is provided.

Favolaschia roldana (Agaricales: Mycenaceae), una especie nueva para México

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Full-text available

Dec 2014

Favolaschia roldana is described and illustrated as a new species. It was collected in the south of Distrito Federal, in Abies religiosa with Quercus and Pinus forest and Quercus-Pinus forest. It is characterized by the presence of larger basidia than those found in other species, acanthocysts in all hymenial surface and by growing on dry branches of Roldana angulifolia.

Taxonomy and Phylogeny of the Favolaschia calocera Complex (Mycenaceae) with Descriptions of Four New Species

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Oct 2021

Favolaschia calocera was originally described from Madagascar, and reported to have a worldwide distribution. In the current study, samples of the Favolaschia calocera from Central America, Australia, China, Kenya, Italy, New Zealand, and Thailand were analyzed by using both morphological and molecular methods. Phylogenetic analyses were based on the internal transcribed spacer (ITS) dataset, and the combined five-locus dataset of ITS, large subunit nuclear ribosomal RNA gene (nLSU), the small subunit mitochondrial rRNA gene (mt-SSU), the small subunit of nuclear ribosomal RNA gene (nu-SSU), and the translation elongation factor 1α (TEF1). Our study proves that Favolaschia calocera is a species complex, and six species are recognized in the complex including four new species. Three new species F. brevibasidiata, F. brevistipitata, and F. longistipitata from China; and one new species F. minutissima from Asia. In addition, Favolaschia claudopus (Singer) Q.Y. Zhang & C. Dai, earlier treated as a variety of Favolaschia calocera R. Heim, were raised to species rank. Illustrated descriptions of these five new taxa are given. An identification key and a comparison of the characteristics of species in the Favolaschia calocera complex are provided.

Collybiopsis bambusicola sp. nov. in Yunnan Province, Southwestern China

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Jan 2023

Collybiopsis bambusicola sp. nov. is described and illustrated from Yunnan Province, Southwestern China, based on morphology and molecular phylogenetic analyses. It is characterized by its minute, cream to cinnamon basidiomata, short, eccentric stipe, forked lamellae, nearly poroid hymenophore which are radially arranged from the base of the stipe, 2–spored basidia, oblong ellipsoid basidiospores (6–8.3 × 3–4.3 μm), and gregarious on dead bamboo. In ITS + nLSU based phylogeny, the new species formed a distinct lineage nested within Collybiopsis ramealis complex clade.

Bioluminescent Mushrooms: Boon for Environmental Health

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Jan 2022

Mushrooms contain secondary metabolites in their fruit bodies, cultured mycelium, and cultured broth which are higher Basidiomycetes and Ascomycetes. Mushrooms have been used in many sides of human activity for many years. Bioluminescence mushrooms gained more attention. Researchers focused on the ecological role of bioluminescence mushrooms as attracting insects for spore dispersal, which may be beneficial for fungi growing on the forest floor where wind flow is low. In this review, we will focus on some famous bioluminescent mushrooms, their ecology, bioluminescence mechanisms and their ecological benefits.

The case of the missing mushroom: a novel bioluminescent species discovered within Favolaschia in southwestern China

Article

Full-text available

Feb 2022

Seven species of bioluminescent mushrooms belonging to seven genera have been described in the tropical rainforests of China. This study contributes the eighth species, found growing on decaying bamboo in Xishuangbanna Tropical Botanical Garden (XTBG). Morphological characteristics and phylogenetic analyses using internal transcribed spacer (ITS) and ribosomal large subunit (LSU) gene regions placed the species within the genus Favolaschia. Comprehensive morphological descriptions, micro and macro photographs, and a phylogenetic tree showing the placement of the new species are provided. This is the second report of bioluminescent Favolaschia in China.

Highlights on Unique Orange Pore Cap Mushroom Favolaschia Sp. and Beech Orange Mushroom Cyttaria sp. and Their Biological Activities

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Full-text available

Aug 2021
J PHARM SCI-US

Fungi exist everywhere around us. However, edible Mushrooms represent promising source for novel compounds with potential uses in different biotechnological fields. Many mushrooms were sources of novel compounds showing biological actions such as antimicrobial, anticancer, antiinflammatory activities. Likewise, mushrooms are macrofungi that exist everywhere around us. They have significant roles in human life as source of nutrition and bioactive compounds. Many mushrooms have been reported as promising biotechnological tools for production of secondary metabolites of various biological activities. Favolaschia sp.; Cyttaria sp. mushrooms are especially interesting due to the rareness of Basidiomycetes. In this review, the ecology, occurrence, of unique Favolaschia sp.; Cyttaria sp. mushrooms were described, and their biological activities.

A new Mycena species with blue basidiomata and porioid hymenophore from Taiwan

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Sep 2018

A small blue stipitate-pileate mushroom with porioid hymenophore was found on dead Calamus sp. (Arecaceae) in Taiwan. Microscopic characteristics indicated a placement within the Mycenaceae. ITS as well as partial LSU ribosomal RNA gene sequences analyses showed a close relationship with Mycena illuminans within the Mycenaceae. A new species with completely blue basidiomata, Mycena indigotica, is proposed based on morphological and molecular evidence.

KEANEKARAGAMAN JAMUR DI KAWASAN PT BADAK NGL

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Mar 2023

Dalam rangka pelestarian keanekaragaman hayati di sekitar perusahaan, PT Badak NGL telah melakukan beberapa langkah penting untuk meningkatkan kesadaran terhadap keberadaan biota dan makhluk hidup asli Kalimantan, yaitu dengan melakukan kegiatan edukasi serta penyebaran informasi. Pemahaman terhadap keanekaragaman hayati dapat dimulai dari beberapa aspek, salah satunya adalah tentang kehidupan dan kelimpahan jamur di sekitar wilayah operasional PT Badak NGL. Jamur merupakan cendawan berukuran makroskopik, yang ternyata dapat ditemukan di berbagai area di PT Badak NGL. Keberadaan jamur menandakan ekosistem yang baik serta lingkungan yang masih lestari. Selain sebagai pengurai zat-zat organik, jamur juga dapat menjadi sumber pakan alami bagi mamalia yang berada di sekitar kawasan konservasi PT Badak NGL. Diharapkan dengan adanya buku ini, pembaca dapat memperoleh wawasan baru terkait keanekaragaman jamur di PT Badak NGL, meningkatkan penjagaan ekosistem dan habitat jamur, serta mendapatkan ilmu terkait pemanfaatan jamur yang aman untuk dikonsumsi.

Notes, outline and divergence times of Basidiomycota

Article

Nov 2019

The Basidiomycota constitutes a major phylum of the kingdom Fungi and is second in species numbers to the Ascomycota. The present work provides an overview of all validly published, currently used basidiomycete genera to date in a single document. An outline of all genera of Basidiomycota is provided, which includes 1928 currently used genera names, with 1263 synonyms, which are distributed in 241 families, 68 orders, 18 classes and four subphyla. We provide brief notes for each accepted genus including information on classification, number of accepted species, type species, life mode, habitat, distribution, and sequence information. Furthermore, three phylogenetic analyses with combined LSU, SSU, 5.8s, rpb1, rpb2, and ef1 datasets for the subphyla Agaricomycotina, Pucciniomycotina and Ustilaginomycotina are conducted, respectively. Divergence time estimates are provided to the family level with 632 species from 62 orders, 168 families and 605 genera. Our study indicates that the divergence times of the subphyla in Basidiomycota are 406–430 Mya, classes are 211–383 Mya, and orders are 99–323 Mya, which are largely consistent with previous studies. In this study, all phylogenetically supported families were dated, with the families of Agaricomycotina diverging from 27–178 Mya, Pucciniomycotina from 85–222 Mya, and Ustilaginomycotina from 79–177 Mya. Divergence times as additional criterion in ranking provide additional evidence to resolve taxonomic problems in the Basidiomycota taxonomic system, and also provide a better understanding of their phylogeny and evolution.