A new species of Pluteus sect. Celluloderma from Kerala State, India

Abstract

A new species of Pluteus has been identified from Kerala State, India based on morphological and molecular (nrITS) characters. This new species, here described as Pluteus
fuscopunctatus, has medium-sized basidiomata, brown pruinose pileus and stipe, globose to subglobose basidiospores, fusiform cheilocystidia, broadly lageniform pleurocystidia, pileipellis of a transitional mixed type epithelium, abundant versiform
caulocystidia and lack clamp connections in all tissues. Phylogenetic analysis confirmed the novelty of the species and its placement in Pluteus sect. Celluloderma.

Full text

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NORDIC JOURNAL OF
BOTANY
Nordic Journal of Botany
Page 1 of 9
Subject Editor: Klaus Høiland
Editor-in-Chief: Torbjörn Tyler
Accepted 24 April 2023
doi: 10.1111/njb.03974
00
1–9
Published 3 June 2023
2023: e03974
© 2023 Nordic Society Oikos. Published by John Wiley & Sons Ltd
A new species of Pluteus has been identified from Kerala State, India based on morpho-
logical and molecular (nrITS) characters. is new species, here described as Pluteus
fuscopunctatus, has medium-sized basidiomata, brown pruinose pileus and stipe, glo-
bose to subglobose basidiospores, fusiform cheilocystidia, broadly lageniform pleu-
rocystidia, pileipellis of a transitional mixed type epithelium, abundant versiform
caulocystidia and lack clamp connections in all tissues. Phylogenetic analysis con-
firmed the novelty of the species and its placement in Pluteus sect. Celluloderma.
Keywords: Hispidoderma, phylogeny, Pluteaceae, tropical, Western Ghats
Introduction
e species-rich genus Pluteus Fr. is widespread from the artic to the tropical habitats
and encompasses ca 500 species (Justo et al. 2011a, b, 2014, He et al. 2019). Most
species of this genus are found on well-rotten wood and are characterized by basidi-
omata with free lamellae, pinkish brown, inamyloid non-ornamented basidiospores,
and convergent hymenophoral trama (Singer 1986). Primarily based on the charac-
teristics of the hymenial cystidia and the pileipellis, three sections have traditionally
been recognized in the genus, viz. Pluteus sect. Pluteus, P. sect. Celluloderma Fayod and
P. sect. Hispidoderma Fayod (Singer 1959, 1986) and these have later been supported
by molecular phylogenetic studies with some minor rearrangements (Menolli et al.
2010, Justo et al. 2011a, b). Pluteus sect. Celluloderma, as recognized by Singer (1959,
1986) includes species with non-metuloidal pleurocystidia and pileipellis as an epithe-
lial hymeniderm with or without cystidioid elements.
Previous taxonomic and molecular studies on Pluteus of Kerala State resulted in
the discovery of 23 species (Pradeep et al. 2002, 2012, Pradeep and Vrinda 2005a, b,
2006, 2008, Farook et al. 2013, Keerthi and Pradeep 2022) and indicate high diver-
sity of this genus in the region. During our continuing study on the genus, an inter-
esting Pluteus was collected repeatedly from the forests of Jawaharlal Nehru Tropical
Botanic Garden and Research Institute (JNTBGRI) campus. Detailed morphological
and molecular studies revealed that the species is distinct from the known species of
A new species of Pluteus sect. Celluloderma from Kerala State,
India
V. Keerthi1,2 and C. K. Pradeep ✉1
1Microbiology Division, Jawaharlal Nehru Tropical Botanic Garden and Research Institute, Palode, iruvananthapuram, India
2Research center, University of Kerala, iruvananthapuram, India
Correspondence: C. K. Pradeep (pradeeptbgri@gmail.com)
Research article
9

Page 2 of 9
Pluteus sect. Celluloderma and presented here as a new species
based on morphological and molecular data.
Material and methods
Collection site and sampling
Fresh specimens of Pluteus (five collections) were collected
during the monsoon months of 2022 from the forests of
JNTBGRI. e JNTBGRI campus is located in the southern
part of Kerala State (southern India) and the forests are part
of the Western Ghats. e region receives an average rain-
fall of 2500 mm per annum and the temperature varies from
20–37°C. e humidity varies from 74–89% and the climate
is tropical, maritime, and monsoonal.
Morphological study
Conventional morphology-based taxonomic methods
were employed for this study (Pradeep et al. 2012). e
macromorphological description of Pluteus fuscopunctatus
was based on fresh specimens collected from Kerala State.
Photographs of fresh specimens were taken in the field along
with other ecological information like habit, habitat, odor
and forest type. Color codes used in the description are from
Kornerup and Wanscher (1978). Basidiomata were dried in
a hot-air oven (40–50°C) overnight for herbarium preserva-
tion. Micro-morphological studies were based on thin free
hand sections of dried material from different parts of the
dried basidioma. e sections were rehydrated with 3% aque-
ous KOH and stained with 1% aqueous solution of Congo
red. Microcharacters were examined, photographed and
measured under an Olympus CX43 optical microscope at
600× magnification. Microphotographs were made with a
Magcam DC10 digital camera attached to the same micro-
scope. irty basidiospores per collection were measured for
length and width. Basidiospore statistical dimensions include
the mean for the length (avL) and the width (avW), the
spore quotient (Q, length/width ratio) and its mean value
(Qm). e holotype and all additional specimens examined
are deposited at the Mycological Herbarium of JNTBGRI,
iruvananthapuram [TBGT(M)].
DNA extraction, PCR amplification and sequencing
Genomic DNA was extracted from fresh specimens
[TBGT(M)18903, TBGT(M)18924] following protocols
in Izumitsu et al. (2012). e nuclear ribosomal Internal
Transcribed Spacer region (nrITS) was amplified and
sequenced from the new species. PCR reactions were per-
formed with the primer pair ITS1 and ITS4 (White et al.
1990). e protocols for PCR amplification and sequencing
followed Kumar et al. (2018a). e newly generated sequences
are deposited in GenBank (www.ncbi.nlm.nih.gov/genbank)
with accession numbers (OQ252937, OQ252936 nrITS).
e sequences of 96 Pluteus taxa retrieved from GenBank
based on a BLAST search and that obtained in this study are
listed in Table 1.
Sequence alignment and phylogenetic analyses
For phylogenetic analysis, two ITS sequences were newly
generated from two collections [TBGT(M)18903,
TBGT(M)18924]. e nrITS sequences of the new species,
along with related species of Pluteus (Pradeep et al. 2012,
Kaygusuz et al. 2021a, b) retrieved from GenBank based on
BLAST search and an outgroup were aligned using MAFFT
web tool (www.ebi.ac.uk/Tools/msa/mafft) with default set-
tings. e final aligned data matrix of nrITS sequences from
98 taxa, including Volvopluteus earlei (Murrill) Vizzini, Contu
and Justo as the outgroup, was imported into BioEdit ver.
7.2.6.1 (Hall 1999) for manual adjustment. Volvopluteus earlei
was selected as the outgroup following Justo et al. (2011a).
Maximum likelihood (ML) analysis was performed with
MEGA X (Kumar et al. 2018b) with 1000 rapid bootstrap
replicates. HKY + G was selected as the best substitution model
as per BIC score, by running modeltest in the same software.
Bootstrap values ≥ 60% (Jeewon and Hyde 2016) were con-
sidered significant. e phylogram inferred from the ML anal-
ysis was visualised with MEGA X (Kumar et al. 2018b).
Results
Molecular phylogeny
e phylogeny inferred by ML analysis of the data matrix
of nrITS sequences is shown in Fig. 1, where seven clades
are observed. e new species forms a subclade with Pluteus
podospilloides E.F.Malysheva and O.V.Morozova from
Vietnam (MT611236) along with two other collections
from China (KU382739, KU382738) in the podospileus
clade. e nrITS sequences of P. fuscopunctatus from India
are distinct from all other sequences of Pluteus species in the
podospileus clade as revealed by BLAST search and phylo-
genetic analyses.
Taxonomy
Pluteus fuscopunctatus C.K.Pradeep and V.Keerthi, sp.
nov. Fig. 2, 3
A species distinguished from the closely related Southeast
Asian species Pluteus podospilloides by its larger basidiomata
(22–60 mm), nonmarginate lamellar edges, larger basidio-
spores (5.6–6.0 × 5.2–6.0 µm), and larger hymenial cystidia.
MycoBank: MB847208
GenBank: OQ252937 (holotype: ITS).
Type
India, Kerala State, iruvananthapuram district, Palode,
JNTBGRI campus, 08°75’N, 77°02’E, 150 m a.s.l., 13 July
2022, Keerthi (holotype: TBGT(M)18903).

Page 3 of 9
Table 1. List of taxa, specimen, GenBank accession numbers of ITS sequences and geographic origin used in the molecular analysis. Type
specimens are identified with a “T”.
Taxon Specimen GenBank Geographic origin
Pluteus albotomentosus T LE289394 KM658284 Russia
P. anatolicus T OKA-TR202 MT523034 Turkey
P. aurantiorugosus ILLS42433 HM562081 USA
P. aurantiorugosus va r. aurantiovelatus TOAVPP212 HQ654908 Italy
P. aureovenatus FK1699 KM983702 Brazil
P. c.f. aureovenatus SP394388 HM562160 Brazil
P. beniensis RSPF0299 JQ065029 Brazil
P. brunneocrinitus T MC4535 KM983692 Brazil
P. brunneosquamulosus T K12794 JN603204 India
P. cebolinhae T CAC39 KM983713 Brazil
P. chrysophaeus LE 312949 KX216316 Russia
P. chrysophlebius SF12 HM562182 USA
P. cinereofuscus AJ229 HM562108 Portugal
P. aff. cinereofuscus LE 303665 KX216324 Russia
P. crenulatus T AJ353 JN603202 Dominican Republic
P. crinitus T FK2064 KM983691 Brazil
P. cutefractus T MCVE30110 MN264751 Solvenia
P. diettrichii JLS1624 HM562143 Spain
P. diptychocystis NMJ184 KM983674 Brazil
P. diverticulatus T 0092579 FJ375247 France
P. dominicanus var. hyalinus SP393695 FJ816665 Brazil
P. eliae Isaacs 2460 HM562076 USA
P. eludens T 50497 NR 119873 Portugal
P. ephebeus 10151 JF908621 Italy
P. aff. ephebeus SF18 JQ065025 USA
P. c.f. eugraptus TNSF12042 HM562116 Japan
P. exiguus LE 226543 FJ774083 Russia
P. extremiorientalis T LE 262872 NR 153249 Russia
P. fastigiatus NKI12 KM983678 Brazil
P. fenzlii PSG 587 MF356564 Slovakia
P. floccipes T BRNM LN794642 Czech Republic
P. fluminensis SP393711 FJ816664 Brazil
P. fuligineovenosus SP393705 FJ816662 Brazil
P. fuscopunctatus T TBGT(M)18903 OQ252937 India
P. fuscopunctatus TBGT(M)18924 OQ252936 India
P. galeroides 886 JF908609 Italy
P. globiger ICN 139025 JQ065030 Brazil
P. halonatus T FK2084 KM983680 Brazil
P. hirtellus T SFSU DED 8259 MG968804 Sao Tome
P. hispidilacteus PDD 108758 MN738667 New Zealand
P. hispidulopsis T NMJ179 KM983684 Brazil
P. hispidulus A1882 KM983681 Spain
P. hispidulus var. cephalocystis ARAN8200509 KM983695 Spain
P. horakianus T OR-2009a FJ375251 Mexico
P. hubeiensis HMJAU 45200 MH167353 China
P. iguazuensis NKI10 KM983704 Brazil
P. inconspicuus T PDD72485 NR 166354 New Zealand
P. insidiosus BRNM788198 MN597451 Czech Republic
P. aff.insidiosis BRNM788196 MN597450 Czech Republic
P. jamaicensis SP393706 FJ816657 Brazil
P. c.f. jamaicensis SP416738 KM983709 Brazil
P. karstedtiae T FK1700 KM983682 Brazil
P. keselakii T BRNM 817402 MN250223 Slovakia
P. liliputianus T LE 312868 MK982288 Russia
P. ludwigii T MCVE 30136 NR 164496 Solvenia
P. mammillatus EIUASM 7916 NR 119874 USA
P. multiformis T AH40107 NR 119877 Spain
P. nanus UC1859980 KF306030 USA
P. nanus PD96725 MN738643 New Zealand
P. c.f. nanus AJ 348 JN603203 Spain
(Continued)

Page 4 of 9
Etymology
e epithet fuscopunctatus (Latin) refers to the brown pru-
inose pileus of the new species.
Description
Basidiomata medium-sized, moderately fleshy. Pileus 22–60
mm in diam., applanate, convex to plano-convex with a small
obtuse umbo when young, becoming depressed at centre
with an uplifted margin in old basidiomata; surface brown
to teak brown (6F4/6F5) with dark brown disc, finely pru-
inose throughout, denser at disc, rarely splitting radially to
expose the white context below; margin straight, entire to
incised, dry, nonhygrophanous. Lamellae 5–7 mm, orange
white (5A2–6A2), crowded with lamellulae of different
lengths; edge concolorous to sides, whitish-flocculose dentic-
ulate when fresh. Stipe 30–55 × 3–6 mm, central to excen-
tric, cylindric, curved and twisted in some, stuffed, solid at
extreme base, equal or tapering towards apex with a slightly
broad base; surface white with dark brown pruinae through-
out. Context white, 1.5 mm thick, soft. Mycelial codons and
mycelial mat absent. Odor nil.
Basidiospores 5.6–6.0 × 5.2–6.0 µm (avL = 5.90 ± 0.17,
avW = 5.84 ± 0.20 µm); Q = 1.00–1.07, Qm = 1.02, glo-
bose to subglobose, hyaline, smooth, thick-walled. Basidia
9.0–40.0 × 4.8–7.2 µm, clavate to cylindro-clavate, four-
spored, smooth, thin-walled, hyaline. Lamella edge sterile
with crowded cheilocystidia. Cheilocystidia 30.0–159.9
× 11.4–32.5 µm, abundant, narrowly fusiform to fusi-
form with obtuse to subobtuse apices, thin-walled, hyaline.
Pleurocystidia scattered, moderately abundant, 42.0–91.0 ×
15.0–28.8 µm, broadly lageniform, non-metuloidal, thin-
walled, hyaline. Hymenophoral trama inversely bilateral with
convergent hyphae, 1.2–28.8 µm wide, thin-walled, hyaline.
Subhymenium pseudoparenchymatous. Pileal trama com-
posed of interwoven, inflated hyphae, 3.9–39.0 µm wide,
thin-walled, hyaline. Pileipellis appearing like a trichoderm
Taxon Specimen GenBank Geographic origin
P. nanus f.griseopus BRNM751771 LN866291 Czech Republic
P. necopinatus T FK1701 KM983693 Brazil
P. nevadiensis T ORA-2010 GU551942 Mexico
P. pallescens AJ214 HM562056 Spain
P. pallidus T Smith62487 HM562193 USA
P. paucicystidiatus SP394383 HM562173 Brazil
P. phlebophorus AJ228 HM562138 Spain
P. aff. phlebophorus SF16 HM562186 USA
P. podospileus AJ617 KM983686 USA
P. podospileus AJ782 KM983687 USA
P. podospileus LE 303682 KX216331 Russia
P. aff. podospileus GM1524 KM983690 England
P. podospilloides T LE 313230 MT611236 Vietnam
P. poliocnemis GM2190 KM983715 Spain
P. riberaltensis va r.conquistensis SP393704 HM562162 Brazil
P. rimosoaffinis SP416740 KM983706 Brazil
P. romellii AJ864 KM983700 USA
P. romellii f.albidus MCVE 28336 KM035790 Italy
P. rugosidiscus BRNM 761706 MH010876 Slovakia
P. sapiicola SP394382 HM562148 Brazil
P. seticeps SF24 HM562192 USA
P. seticeps S.D. Russell MycoMap6309 MK560139 USA
P. aff. seticeps PDD: 107338 MN738664 New Zealand
P. spurius T LE 312866 MK982290 Russia
P. squarrosus T GDGM 42302 MK791273 China
P. stenotrichus T AJ352 JN603201 Dominican Republic
P. sublaevigatus SP393694 FJ816667 Brazil
P. tenebromarginatus T GC17102401 MT079860 France
P. terricola JAC13711 MN738648 New Zealand
P. terricola PDD 106511 MN738653 New Zealand
P. thomsonii 155 FJ375253 Spain
P. tomentosulus MO 93719 KM983672 USA
P. tomentosulus f. brunneus T LE 312908 KX216307 Russia
Pluteus sp. JLF 1767 MK634597 USA
Pluteus sp. PDD 110518 MN738674 New Zealand
Pluteus sp.GDGM 41576 KU382738 China
Pluteus sp.GDGM 42371 KU382739 China
Volvopluteus earlei MA22816 HM562204 Spain
Table 1. Continued.

Page 5 of 9
with a transition towards an epithelium. Pileipellis mainly
composed of two types of elements: 1) long cylindrical nar-
row elements with obtuse to subacute apices, 15.6–221.0 ×
6.5–48.1 µm, which are more clustered near the disc and
elsewhere in tufts, containing brown intracellular pigments
and 2) globose, subglobose, clavate to subclavate cells,
19.5–65.0 × 6.5–52.0 µm, with brown intracellular con-
tents. Stipitipellis a cutis composed of parallel hyphae, 2.6–
32.5 µm wide, thin-walled, hyaline. Caulocystidia scattered
throughout the surface of the stipe, 13.0–149.5 × 5.2–39.0
Figure 1. Maximum likelihood tree generated from nrITS sequence data. Bootstrap values are indicated above/below branches. Pluteus
fuscopunctatus sp. nov. from India are indicated in bold.

Page 6 of 9
µm, versiform, narrowly fusiform, clavate, cylindro-clavate to
subclavate with obtuse, subobtuse to subacute, apices or with
a narrow neck, thin-walled with brown intracellular contents.
Clamp connections absent. Oleiferous hyphae present.
Habitat and phenology
Solitary, paired to scattered on dead decaying tree stumps or
on soil in tropical evergreen forest, Kerala State, India. July,
October, November.
Additional specimens examined (paratypes)
India, Kerala State, iruvananthapuram district, Palode,
JNTBGRI campus, 21 July 2022, Keerthi TBGT(M)18923;
ibid., 22 July 2022, Keerthi TBGT(M)18924; ibid., 27 Oct.
2022, Keerthi TBGT(M)18975; ibid., 1 Nov. 2022, Keerthi
TBGT(M) 18985.
Notes
Pluteus fuscopunctatus is characterized by a set of features
such as medium-sized basidiomata, convex to plano-
convex, depressed, pruinose, brown pileus, nonmarginate
lamellar edges, white stipe with brown pruinae, globose to
subglobose basidiospores, fusiform cheilocystidia, broadly
lageniform pleurocystidia, pileipellis as a transitional
trichoderm towards an epithelium, presence of abun-
dant caulocystidia, terrestrial and lignicolous habitat and
lack of clamp connections. e nrITS based phylogeny
together with nonmetuloidal pleurocystidia and pileipellis
Figure 2. Pluteus fuscopunctatus. [TBGT(M)18903, holotype], (a–b) habit in-situ, (c) basidiomata with pink lamellae, (d) sectional view of
basidioma, (e) basidia, (f) basidiospores. Scale bars: (a–d) = 10 mm, (e–f) = 10 μm.

Page 7 of 9
composed of epithelioid hymeniform layer interrupted
by elongate cystidioid structures supports the placement
of P. fuscopunctatus in Pluteus sect. Celluloderma. Singer
(1956, 1959, 1986) subdivided Pluteus sect. Celluloderma
into Pluteus subsect. Mixtini Singer, characterised by
elongated elements in the pileipellis and Pluteus subsect.
Eucellulodermini Singer characterized by the lack of these
elements. e new species thus belongs to subsect. Mixtini
due to the presence of nonmetuloidal pleurocystidia and a
transitional pileipellis containing both elongate and epi-
thelial elements.
Molecular analyses based on nrITS (Fig. 1) shows that
P. fuscopunctatus is placed within the podospileus clade
and is closely related to P. podospilloides E.F.Malysheva
and O.V.Morozova (MT611236) from Vietnam and two
other collections of Pluteus sp. from China (KU382738,
KU382739). Pluteus podospilloides, originally described from
Vietnam (Malysheva et al. 2020) is similar in some macro-
and microscopic features, especially in the nature of hymenial
cystidia, pileipellis, presence of caulocystidia and clampless
hyphae, but P. podospilloides differs markedly in its smaller
basidiomata (9–12 mm), marginate lamellar edges, smaller
Figure 3. Pluteus fuscopunctatus. [TBGT(M)18903, holotype]. (a) lamella edge with cheilocystidia, (b) cheilocystidia, (c, d) pleurocystidia,
(e) pileipellis, (f) caulocystidia. Scale bars: (a–f) = 20 µm.

Page 8 of 9
basidiospores (4.5–5.5 × 4.0–5.0 µm), larger hymenial
cystidia with brown intracellular contents. Moreover, the
BLASTn results using the nrITS (641 bp) sequence of P. fus-
copunctatus showed only 92.61% sequence identity with P.
podospilloides and differ at 36 positions. Pluteus fuscopunctatus
is also closely related to two other collections of Pluteus spe-
cies from China (KU382738, KU382739) in the podospi-
leus clade. However, the taxonomic details are not available
of these species for morphological comparison. Furthermore,
the sequences of these two species indicate only 93.53% simi-
larity with P. fuscopunctatus and differ at 39 positions which
indicate that they represent different species.
Some of the other related species of the podospileus
clade include Pluteus podospileus Sacc. and Cub., P. cute-
fractus Ferisin, Dovana and Justo, P. necopinatus Menolli
and Capelari, P. seticeps (G.F.Atk.) Singer, P. brunneocrini-
tus Menolli, Justo and Capelari and P. crinitus Menolli and
Capelari. Pluteus podospileus, though morphologically some-
what similar to the present species, differs by its small (9–34
mm) pileus with velvety to pilose scaly surface, larger (5.5–
7.5 × 4.5–6.0 µm), subglobose to broadly ellipsoid basidio-
spores, smaller hymenial cystidia and lack of caulocystidia
(Orton 1986, Vellinga 1990). Pluteus cutefractus, originally
described from Slovenia differs mainly by its cracked pileus,
large, globose to broadly ellipsoid basidiospores (4.9–7.1 ×
5.0–5.8 µm), small clavate to broadly clavate hymenial cys-
tidia and habitat on underground twigs (Song et al. 2019).
Pluteus necopinatus is distinguished mainly by its small,
punctate fibrillose pileus (14 mm), sulcate-striate margin,
slightly smaller basidiospores and lack of pleurocystidia
and caulocystidia (Menolli et al. 2015). Pluteus brunneo-
crinitus and P. crinitus were originally described from Brazil
(Menolli et al. 2015) and are distinct by their smaller basidi-
omata, smaller basidiospores, lack of pleurocystidia and
predominantly clavate and smaller cheilocystidia. Pluteus
seticeps can be separated by its small fragile basidiomata,
small, ovate to subglobose basidiospores and small, differ-
ently shaped hymenial cystidia (Homola 1972, Minnis and
Sundberg 2010).
e recently described Pluteus species in sect. Celluloderma
with mixed epithelium from Kerala includes Pluteus brun-
neosquamulosus C.K.Pradeep and K.B.Vrinda (Pradeep et al.
2012) and P. delicatulus C.K.Pradeep and K.B.Vrinda
(Pradeep and Vrinda 2006). Pluteus brunneosquamulosus
is distinct by its brown squamulose pileus with distinctive
concentric cracking, silky white stipe, subglobose to broadly
ellipsoid basidiospores, cheilocystidia with a long flexuous
neck and lageniform to narrowly utriform caulocystidia.
Pluteus delicatulus can be distinguished from P. fuscopuncta-
tus by its small, delicate basidiomes (7–10 mm), white stipe,
slightly smaller basidiospores, clavate to vesiculose hymenial
cystidia and lack of caulocystidia.
Funding – VK acknowledges financial support from the CSIR-
UGC, India in the form of Senior Research Fellowship (UGC-Ref.
No.: 950/(CSIR-UGC NET JUNE 2019).
Permits – e authors are thankful to the Director, JNTBGRI for
facilities and also the principal chief conservator of forests, Govt. of
Kerala for granting permission to collect specimens from the forests
of Kerala.
Author contributions
V. Keerthi: Conceptualization (equal); Data curation
(equal); Formal analysis (equal); Funding acquisition
(lead); Investigation (equal); Methodology (equal); Project
administration (supporting); Resources (lead); Supervision
(supporting); Validation (equal); Writing – original draft
(equal); Writing – review and editing (supporting). C. K.
Pradeep: Conceptualization (equal); Data curation (equal);
Formal analysis (equal); Funding acquisition (support-
ing); Investigation (equal); Methodology (equal); Project
administration (lead); Resources (equal); Supervision (lead);
Validation (lead); Writing – original draft (lead); Writing –
review and editing (lead).
Data availability statement
e newly generated sequences are deposited in GenBank
(www.ncbi.nlm.nih.gov/genbank) with accession numbers
(OQ252937, OQ252936 nrITS). All other data are available
from the Dryad Digital Repository: https://doi.org/10.5061/
dryad.n02v6wx2k (Keerthi and Pradeep 2023).
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