Phytotaxa 331 (2): 273–280 ISSN 1179-3155 (print edition)

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Article PHYTOTAXA
Copyright © 2017 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.331.2.11

Trogia benghalensis (Marasmiaceae, Basidiomycota), a new species from India

ARUN KUMAR DUTTA1,2, SUDESHNA NANDI1, ENTAJ TARAFDER1,2, RIMPA SIKDER1, ANIRBAN ROY2 &
KRISHNENDU ACHARYA1*
1
Molecular and Applied Mycology and Plant Pathology Laboratory, Department of Botany, University College of Science and Agricul-
ture, University of Calcutta, Kolkata - 700019, West Bengal, India.
2
West Bengal Biodiversity Board, Poura Bhawan, FD-415A, Bidhan Nagar, Kolkata, West Bengal 700106, India.
*Corresponding author: krish_paper@yahoo.com

Abstract

A new species of Trogia is described from West Bengal, India based on both morphological characteristics and phyloge-
netic data. Analysis of the nuclear ribosomal internal transcribed spacer region sequence data suggests that T. benghalensis
is phylogenetically distinct from its closely related species. Comprehensive description based on macro- and microscopic
characteristics, colour photographs and comparisons with morphologically similar and phylogenetically related species are
provided.

Key words: Agaricales; nrITS; taxonomy; West Bengal

Introduction

The genus Trogia (Marasmiaceae, Agaricales, Basidiomycota) was established by Elias Magnus Fries in 1835 with
Trogia montagnei as the type species. Characteristic features of the genus include clitocyboid to omphalinoid habit,
tough texture and ability of reviving in situ (Kumar & Manimohan 2009). They are mostly saprophytic in nature and
predominantly grows on dead woods or on other plant debris (Corner 1991).
The first worldwide treatment of the genus was published by Corner (1966) employing a broad generic concept
where he recognized 56 species relying on the presence of sarcodimitic tramal structure. Based on the Singer’s concept
(Singer 1975), Corner’s accepted species could be segregated into not less than four genera [viz. Gerronema Singer,
Hydropus Kühner ex Singer, Hemimycena Singer, Mycena (Pers.) Roussel etc.]. However, Singer (1986) in his book
“The Agaricales in Modern Taxonomy” strongly opposed the generic circumscription as mentioned by Corner (1966)
and accepted only three species belonging to the genus Trogia. He employed a combination of features like an easily
reviving basidiomata, the presence of narrow, mostly furcated lamellae consisting of interwoven hyphal arrangement,
and characteristically presence of pigmented trichodermial epicutis for demarcating the genus from its close relatives
like Gerronema, Clitocybe (Fr.) Staude, and Neoclitocybe Singer. Redhead (1987) found Corner’s circumscription of
the genus solely based on the presence of sarcodimitic tissues to be taxonomically insignificant as the feature is shared
by quite a number of agaric genera. So, Redhead (1987) adopted the family Xerulaceae Jülich for members containing
sarcodimitic tissues. Later, Corner (1991) defended his concept of Trogia and recorded altogether 90 species from
Asia, Australasia and Neotropics.
Currently, there are approximately 93 well accepted species belonging to the genus (www.mycobank.org), but
most of the previous studies were solely based on the morphological observations and thus, lack molecular data (Kumar
& Manimohan 2009). The type species of the genus, T. montagnei, originally described from southern India, is poorly
known and the type specimen also remains untraceable. So, the taxonomic circumscription of Trogia still remains
unknown (Wilson & Desjardin 2005, Kumar & Manimohan 2009, Yang et al. 2012). For demonstrating the relationship
of gymnopoid and marasmioid fungi, Wilson & Desjardin (2005) for the first time included Trogia infundibuliformis
Berk. & Broome in their phylogenetic analyses where the taxon formed a sister group to the marasmioid clade with
weak statistical support. Almost seven years later, Yang et al. (2012) while describing Trogia venenata showed the
placement of the genus together with other genera like Megacollybia Kotl. & Pouzar, Gerronema, Clitocybula (Singer)

Accepted by Samantha Karunarathna:22 Nov. 2017; published: 12 Dec. 2017 273

Singer ex Métrod, Porotheleum Fr. and Calyptella Quél. in a strongly supported clade based on the combined ITS and
nrLSU rDNA dataset.
To date, India is represented only by five species of Trogia (Manjula 1983, Natarajan et al. 2005, Kumar &
Manimohan 2009). Keeping in view the wide range of phyto-topographical features, presence of amiable micro-
climate admixed with favorable ecological conditions, the total reported number of species from India seems to be
an underestimate. During a routine expedition in the state West Bengal, one interesting Trogia was found possessing
characteristics distinct from those found in any previously described species, and is described here as a new species
based on morphological and molecular (nuclear ribosomal internal transcribed spacer) sequence data.

Material and methods

Morphological protocols:—Basidiomata were collected from West Bengal during the field trips in the year 2015–
2016. The morphological and ecological features of collected specimens were noted in the field. Colour codes and
terms follow Kornerup & Wanscher (1978). The terms used to describe lamellae spacing refer to the number of lamellae
that run from the stipe to the pileus margin (L), and do not include the lamellulae whose spacing is indicated by the
number of series present (l).
Microscopic features were obtained from dried material by mounting free-hand sections of basidiomata in 5 %
KOH, Melzer’s reagent and Congo Red. Specimens were then examined with a Carl Zeiss AX10 Imager A1 phase
contrast microscope under different lights and photographed with a digital camera using maximum optical zoom. In the
spore statistics: values in parentheses indicate the minimum and maximum measured values, Xmr, denotes the range of
spore means, Xmm indicate the mean of spore means (± SD); Q value, the quotient of spore length by spore width in any
one spore, indicated as a range of variation in n spores measured; Qmr, the range of mean of Q-values and Qmm, the mean
of Q values for two collected specimens (± SD). Voucher specimens have been deposited in the Calcutta University
Herbarium (CUH).
DNA extraction, PCR amplification, and sequencing:—Genomic DNA was extracted from dried basidiomata
following the protocol as described by Dutta et al. (2015). PCR primers ITS1 and ITS4 (White et al. 1990) were used
for amplification of the nuclear ribosomal internal transcribed spacer (nrITS) region. PCR cycling started with 4 min at
94 °C, and was followed by 30 cycles consisting of denaturation for 30 s at 94 °C, annealing for 30 s at 55 °C, extension
for 1 min at 72 °C, and a final elongation step of 5 min at 72 °C. The PCR products were purified using the QIAquick®
Gel Extraction Kit (QIAGEN, Germany). Cycle sequencing was performed on amplified products using nrITS primers
mentioned above. Nucleotide sequences were read using an automated DNA sequencing on ABI3730xl DNA Analyzer
(Applied Biosystems, USA). The newly generated sequences were edited manually using BioEdit sequence alignment
editor version 7.0.9.0 (Tom Hall, Ibis Biosciences, Carlsbad, USA) and deposited in GenBank (www.ncbi.nlm.nih.
gov).
Dataset representation:—The edited sequence was then used in BLAST searches across the GenBank database
to determine the most closely related taxa for molecular identification. A dataset was generated from the highest scored
hits most relevant for identification, as well as from samples previously used in the phylogenetic reconstruction of
Trogia (Yang et al. 2012).
Altogether, forty-six nrITS sequences representing 33 species were used in the analyses, of which 43 were retrieved
from GenBank and three sequences were generated for this study. Gymnopus contrarius (Peck) Halling was selected as
outgroup for rooting purposes following Yang et al. (2012).
Sequence alignment and phylogenetic analysis:—The nrITS dataset was aligned using ClustalX (Thompson et
al. 1997), and then imported to MEGA v. 6.0 (Tamura et al. 2013) for manual check and adjustment where necessary.
The appropriate model of evolution for maximum likelihood (ML) and Bayesian analyses (BA) was determined
using jModeltest 2.1.6 (Darriba et al. 2012). Based on the Bayesian information criterion (BIC), the GTR+I+G
(19167.019891) model was selected for the nrITS data.
ML bootstrapping analysis was performed with RAxML ¬ 8.2.4, using the parameters specified by jModeltest on
the Cyberinfrastructure for Phylogenetic Research (CIPRES) web portal (https://www.phylo.org/portal2/) (Miller et al.
2009) with bootstrap statistics calculated from 1000 bootstrap replicates.
Bayesian phylogenetic analysis (BA) was carried out using MrBayes v. 3.2.2 (Ronquist et al. 2012). This program
performs a Bayesian Inference (BI) of the phylogeny, using Metropolis-coupled Markov chain Monte Carlo method
(Geyer 1991). For a given data set, the GTR+I+G model was employed. Markov chains ran for 106 generations, saving

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a tree every 100th generation. Default settings in MrBayes were used for the incremental heating scheme for the chains
(3 heated and 1 cold chain), unconstrained branch length (unconstrained: exponential (10.0)), and uninformative
topology (uniform) priors. The trees in a burn-in period (25%) were excluded, and the 50% majority-rule consensus
tree of the remaining trees was used to obtain estimates of the posterior probabilities (PPs) of each clade.

Results

Taxonomy

Trogia benghalensis K. Acharya & A.K. Dutta, sp. nov., Figures 1 & 2
MycoBank MB 822864

FIGURE 1. Trogia benghalensis (CUH AM031). Field photograph of the basidiomata . Scale Bar: 10 mm.

Etymology:—The specific epithet ‘benghalensis’ refers to West Bengal, India, the region where the type species was
discovered.
Diagnosis:—Pileus small to medium sized (8–23 mm), greyish yellow to light orange; lamellae subdistant to
moderately close; stipe central, greyish yellow to dull yellow; basidiospores 5–8 × 5–7.5 μm (Q mean of 1.1), globose
to subglobose or broadly ellipsoid; basidia 2–4 spored; cheilocystidia 21–36 × 8–14 μm, clavate to subclavate, crowded;
pleurocystidia, pileocystidia and caulocystidia absent.
Holotype:—INDIA. West Bengal: Kolkata, Salt Lake, Central park, 22°35’18.3”N, 88°25’02.8”E, 6 m elev., 09
July 2015, K. Acharya & A. K. Dutta, CUH AM031.
Description:—Pileus 8–23 mm diam., infundibuliform, yellowish white (2A2) to greyish yellow (1-2B3, 3-4B4)
or dull yellow (3B3) to light orange (5A4), unchanging on bruising, becoming translucent with KOH, glabrous, margin
striate to sometimes eroded , often so deeply rimos that the entire pileus divides into two to three halves; context thin
(< 1 mm), yellowish white (4A2), becoming translucent with KOH. Lamellae up to 1–2 mm broad, decurrent, at first
white, becoming concolorous to the pileus in age, subdistant to moderately close (L = 22–28, l = 1–2), even to slightly

TROGIA BENGHALENSIS (MARASMIACEAE, BASIDIOMYCOTA) Phytotaxa 331 (2) © 2017 Magnolia Press • 275
wavy or eroded, concolorous. Stipe 12–22 × 1–3 mm, central to slightly eccentric, cylindrical, curved, gradually
tapered towards base, slightly bulbous at the point of attachment to the substrate, fibrous, hollow, surface greyish
yellow (1-2B3, 3-4B4) to dull yellow (3B3), glabrous. Odour and taste indistinct. Spore-print white.

FIGURE 2. Microscopic features of Trogia benghalensis (CUH AM031, holotype). a. Basidiospores; b. Basidia and basidioles; c.
Cheilocystidia; d. Hyphal arrangement in the pileipellis. Scale Bars: a–c = 5 μm; d = 10 μm.

276 • Phytotaxa 331 (2) © 2017 Magnolia Press DUTTA ET AL.

FIGURE 3. Phylogram generated from maximum likelihood (RAxML) analysis using a GTR+I+G model of nucleotide substitution (-lnL
= 9257.028122) based on nrITS sequence data. Numbers to the left of / are Maximum likelihood bootstrap (MLBS) support, and those to
the right indicate the Bayesian posterior probabilities (PP, in bold). MLBS values ≥ 50% and PP > 0.50 are shown. The scale bar represents
the expected changes per site. The newly generated sequences are indicated by an asterisk (*), and the newly described taxon is placed in
bold font to highlight its phylogenetic position in the tree. GenBank accession numbers for all of the sequences are provided in the tree
with their country of origin.

Basidiospores (5–)6.5–7.5(–8) × 5–6(–7.5) μm [Xmr = 6.8–7.2 × 6.5–6.9 μm, Xmm = 6.9 ± 0.8 × 6.7 ± 0.7 μm, Qmm
= 1.1 ± 0.04, Qmr = 1–1.2, n = 20 spores, s = 2 specimens], globose to subglobose or broadly ellipsoid, smooth, hyaline,
inamyloid, non-metachromatic in cresyl blue, truncate with prominent hilar appendage (1–1.5 μm long), I-guttate
when viewed with KOH, thin-walled. Basidia 25–36 × 7–9.5 μm, clavate to broadly clavate, hyaline, thin-walled,
2–4 spored; sterigmata 2.5–4(–7) μm long, cylindrical. Basidioles 30–34 × 7–9 μm, cylindrical to clavate, hyaline,
oil granules present when viewed with KOH, thin-walled. Pleurocystidia absent. Lamellae edge sterile, crowded
with cystidia. Cheilocystidia (21–)25–29(–36) × (7–)10–12(–14) μm, clavate to subclavate, hyaline, moderately
thick-walled. Lamellar trama hyphae interwoven, slender, 3–10 μm broad, filamentous, hyaline, inamyloid, thin- to
slightly thick-walled. Pileipellis an epicutis, hyphae 3–10 μm broad, cylindrical, hyaline to pale yellow when viewed
with KOH, inamyloid, non-gelatinous, thin-walled. Pileocystidia absent. Pileus trama hyphae interwoven, 3–15

TROGIA BENGHALENSIS (MARASMIACEAE, BASIDIOMYCOTA) Phytotaxa 331 (2) © 2017 Magnolia Press • 277
μm broad, non-gelatinous, hyaline, inamyloid, thin-walled. Stipe trama hyphae interwoven, 3.5–4.5(–6) μm broad,
cylindrical, non-gelatinous, hyaline, inamyloid, thin-walled. Stipitipellis hyphae 3–4(–5) μm broad, cylindrical, often
branched, non-gelatinous, hyaline to pale yellow, inamyloid, thin-walled. Caulocystidia absent. Clamp connections
present in all the tissues.
Habit and habitat:—Solitary to gregarious, on dead and decayed wood of Mangifera indica L.
Additional specimen examined:—INDIA. West Bengal: Kolkata, Salt Lake, Central park, 22°35’20.0” N,
88°24’59.4” E, 5 m elev., 09 July 2016, A. K. Dutta & K. Acharya, CUH AM122.
Molecular phylogeny:—An alignment of 712 nucleotides in the final dataset was produced and analyzed. Each of
the ML analysis iterations recovered a single tree of which the likelihood values did not differ significantly. Bayesian
analyses reached a standard deviation of split frequencies of 0.004 after one million generations. The trees generated
using the ML and Bayesian analyses (BA) were identical in topology. Therefore, only the phylogenetic tree generated
using ML analysis has been presented here (Figure 3; -lnL = 9257.028122). Figure 3 highlights branch support values
recovered by ML and posterior probabilities (PP) derived from the Bayesian majority-rule consensus tree (MLBS: ≥
50%, PP: ≥ 0.50).

Discussion

In the nrITS tree (Figure 3), three included species of Trogia (viz. T. benghalensis, T. infundibuliformis and T. venenata)
cluster together in a clade with 0.97 PP and 68% MLBS support. This clade is close to species of Megacollybia,
Gerronema and Clitocybula (76% BS, 0.97 PP). The newly described taxon is sister to Trogia venenata with moderate
support values (79% BS, 0.96 PP). Morphologically, T. venenata differs from T. benghalensis by its pleurotoid habit;
pinkish, larger pileus (up to 60 mm diam.); solid, pinkish to brownish stipe; more slender (3.5–5.5 μm, Q mean of
1.59), ellipsoid to pip-shaped basidiospores; presence of pileocystidia; and presence of Rameales-like outgrowth on
the stipe surface (Yang et al. 2012). Trogia infundibuliformis, originally described from Sri Lanka, has spathulate to
flabellate, pale pink to purplish brown pileus; distant lamellae (l = 1–3); differently sized (6.3–8.5 × 3.5–4.5 μm, Q
mean of 1.75) ovoid to ellipsoid basidiospores; absence of cheilocystidia; and presence of nodulose to diverticulate
outgrowth of the pileipellis hyphae (Pegler 1986, Kumar & Manimohan 2009, Senthilarasu 2014).
Considering the overall morphology, the newly described taxon seems close to Trogia icterina (Singer) Corner,
Gerronema citrinum (Corner) Pegler, Trogia subglobospora Corner, Trogia holochlora (Berk. & Broome) Corner, and
Trogia straminea Corner. However, Trogia icterina, originally described from Florida and subsequently reported in
Europe, North America, Brazil and China, primarily differs by its sulphureous to mustard yellow pileus, somewhat
differently sized basidiospores (8–9 × 6–7 μm, Corner 1966; 7–9.5 × 5–7 μm diam., Zhishu et al. 1993), and the
presence of subcylindrical to obtuse pileocystidia and caulocystidia (Singer 1948, Corner 1966, Zhishu et al. 1993).
‘Gerronema citrinum’ has much larger pileus (up to 40 mm diam.) which is dull sulphur yellow with greyish brown
towards center, broader lamellae (up to 4 mm) that is often accompanied by slightly intervenose nature, much longer
stipe (30–50 mm), and less broader (5.5–7.5 × 3–4 μm, Q mean of 1.83), oblong ellipsoid basidiospores (Pegler 1983,
Corner 1991). Trogia subglobospora differs by its fuscous fuliginous to greyish pileus that contains furfuraceous
squammules towards center, poroid reticulate lamellae with 2–4 series of lamellulae, much longer (up to 50 mm),
pruinose stipe, smaller (4–4.7 × 3–3.5 μm), amyloid basidiospores, and presence of caulocystidia (Corner 1991).
Trogia holochlora has pastel yellow to greenish yellow pileus that becomes reddish on drying or with alkali, more
slender (6.5–8.5 × 3.5–5.3 μm) ellipsoid basidiospores, and fertile lamellae edge (Pegler 1977, Corner 1991). Trogia
straminea Corner has considerably larger basidiome (pileus 20–50 mm, stipe 30–60 mm), pruinose stipe surface,
crowded lamellae (L = 16–22, l = 3–4), absence of cheilocystidia and presence of caulocystidia (Corner 1991).

Acknowledgements

Authors gratefully acknowledge the Department of Environment, Government of West Bengal, India for providing
financial assistance. The first author (AKD) is thankful to Vladimír Antonín (Brno, Czech Republic) for providing
necessary literatures on the allied taxa.

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