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Trechisporales emended with a segregation of Sistotremastrales ord. nov. (Basidiomycota)

November 2022
mycosphere 13(1):862-954

November 2022
13(1):862-954

DOI:10.5943/mycosphere/13/1/11

Authors:

Shi-Liang Liu

Chinese Academy of Sciences

Shuang-Hui He

Beijing Forestry University

Tom W. May

Royal Botanic Gardens Victoria

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Trechisporales, typified by Trechispora and equivalent to Hydnodontales, is a recently introduced order within Agaricomycetes. This order only comprises one family Hydnodontaceae and 16 independent genera, but the relationships among these genera are not fully clarified. Here, via a wider sampling especially from Asia Pacific, careful morphological examinations and comprehensive multilocus-based phylogenetic analyses, the classification of Trechisporales is emended. Sertulicium and Sistotremastrum are segregated from Trechisporales, and placed in the new family Sistotremastraceae within the new order Sistotremastrales. A new genus Allotrechispora segregated from Trechispora is introduced within Hydnodontaceae, Trechisporales, and Boidinella, Litschauerella and Sphaerobasidium are excluded from Trechisporales. Brief summaries to genera accepted in Sistotremastrales and Trechisporales, and a key to all 12 genera accepted in Trechisporales are provided. In addition, Tomentella and Murrilloporus, potential synonyms of Trechispora, are excluded from Trechisporales and of uncertain position, respectively. At the species level, 19 new species are described with one from Allotrechispora, one from Fibrodontia, one from Subulicystidium and 16 from Trechispora, and seven new combinations are proposed with two for Allotrechispora and five for Trechispora. In addition, Trechispora yunnanensis is excluded from Trechispora. A key to all 87 species accepted in Trechispora is provided. In conclusion, an emended classification of Trechisporales within Agaricomycetes is constructed, which will help to further clarify species diversity and explore trait evolution within Trechisporales.

Phylogenetic position of Sistotremastrales and Trechisporales within the Agaricomycetes inferred from the combined dataset of ITS, nrLSU, tef1-α and rpb2 regions. The topology is generated by the maximum likelihood algorithm. Bootstrap values and Bayesian

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Phylogenetic relationship among genera within Sistotremastrales and Trechisporales inferred from the combined dataset of ITS, nrLSU, tef1-α, rpb2 and mtSSU regions. The topology

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Phylogenetic relationship among species of Fibrodontia inferred from the combined dataset of ITS and nrLSU regions. The topology is generated by the maximum likelihood algorithm. Bootstrap values and Bayesian posterior probabilities, when simultaneously above 50% and 0.8, respectively, are labelled at the nodes. The newly described species are in boldface.

Basidiomes of Allotrechispora gatesiae (LWZ 20180515-18, holotype). Scale bars: a = 1 cm, b = 2 mm.

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Submitted 29 June 2022, Accepted 4 October 2022, Published 4 November 2022

Corresponding Author: Li-Wei Zhou – e-mail – liwei_zhou1982@im.ac.cn 862

Trechisporales emended with a segregation of Sistotremastrales ord.

nov. (Basidiomycota)

Liu SL1, He SH2, Wang XW1,3, May TW4, He G5,6, Chen SL6 and Zhou LW1,*

1State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, P.R.

China

2School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, P.R. China

3University of Chinese Academy of Sciences, Beijing 100049, P.R. China

4Royal Botanic Gardens Victoria, Birdwood Avenue, Melbourne 3004, Australia

5School of Pharmacy and Life Science, Jiujiang University, Jiujiang 332005, P.R. China

6College of Life Sciences, Nanjing Normal University, Nanjing 210023, P.R. China

Liu SL, He SH, Wang XW, May TW, He G, Chen SL, Zhou LW 2022 – Trechisporales emended

with a segregation of Sistotremastrales ord. nov. (Basidiomycota). Mycosphere 13(1), 862–954,

Doi 10.5943/mycosphere/13/1/11

Abstract

Trechisporales, typified by Trechispora and equivalent to Hydnodontales, is a recently

introduced order within Agaricomycetes. This order only comprises one family Hydnodontaceae

and 16 independent genera, but the relationships among these genera are not fully clarified. Here,

via a wider sampling especially from Asia Pacific, careful morphological examinations and

comprehensive multilocus-based phylogenetic analyses, the classification of Trechisporales is

emended. Sertulicium and Sistotremastrum are segregated from Trechisporales, and placed in the

new family Sistotremastraceae within the new order Sistotremastrales. A new genus

Allotrechispora segregated from Trechispora is introduced within Hydnodontaceae,

Trechisporales, and Boidinella, Litschauerella and Sphaerobasidium are excluded from

Trechisporales. Brief summaries to genera accepted in Sistotremastrales and Trechisporales, and a

key to all 12 genera accepted in Trechisporales are provided. In addition, Tomentella and

Murrilloporus, potential synonyms of Trechispora, are excluded from Trechisporales and of

uncertain position, respectively. At the species level, 19 new species are described with one from

Allotrechispora, one from Fibrodontia, one from Subulicystidium and 16 from Trechispora, and

seven new combinations are proposed with two for Allotrechispora and five for Trechispora. In

addition, Trechispora yunnanensis is excluded from Trechispora. A key to all 87 species accepted

in Trechispora is provided. In conclusion, an emended classification of Trechisporales within

Agaricomycetes is constructed, which will help to further clarify species diversity and explore trait

evolution within Trechisporales.

Keywords – 29 new taxa – Hydnodontaceae – Hydnodontales – macrofungi – Sistotremastraceae –

Taxonomy

Introduction

Trechisporales typified by Trechispora was newly introduced by Hibbett et al. (2007). An

earlier order name Hydnodontales is equivalent to Trechisporales (Jülich 1981), but Hibbett et al.

(2007) stated that Trechispora, a synonym with priority over Hydnodon (Ryvarden 2002), is the

most species-rich genus in this order and thus proposed the new order name. Although not

Mycosphere 13(1): 862–954 (2022) www.mycosphere.org ISSN 2077 7019

Article

Doi 10.5943/mycosphere/13/1/11

863

recommended (Rec. 16A.1 of the Shenzhen Code), the proposal of Trechisporales is permissible

according to Art. 11.10 of the Shenzhen Code (Turland et al. 2018). Since then, almost all authors

has abandoned the order name Hydnodontales.

Trechisporales is a taxon-poor order compared with most other orders within

Agaricomycetes, Basidiomycota (Wijayawardene et al. 2022b). For now, one family

Hydnodontaceae and 14 independent genera, viz. Brevicellicium, Brevicellopsis, Dextrinocystis,

Fibrodontia, Litschauerella, Luellia, Porpomyces, Pteridomyces, Sertulicium, Sistotremastrum,

Subulicystidium, Suillosporium, Trechispora and Tubulicium are accepted in Trechisporales in

studies specific to the order (Larsson 2007, Hjortstam & Ryvarden 2008, de Meiras-Ottoni et al.

2021, Spirin et al. 2021). It is worth mentioning that although two monotypic genera

Dextrinodontia and Fibriciellum were also listed as members of Trechisporales (Larsson 2007,

Hibbett et al. 2014, Wijayawardene et al. 2022a), their type species were previously combined to

Trechispora (Larsson 1992). Therefore, Dextrinodontia and Fibriciellum are actually later

synonyms of Trechispora. In addition, some recent synopses place further genera in

Hydnodontaceae, Trechisporales, including Boidinella (Kirk 2019) and Sphaerobasidium (He et al.

2019, Kirk 2019, Wijayawardene et al. 2022a), which bring the genus number to 16. However, the

taxonomic position of Boidinella, Brevicellopsis, Litschauerella and Sphaerobasidium within

Trechisporales has never been confirmed from a phylogenetic perspective. Morphologically,

Trechisporales is highly diverse: with stipitate, clavarioid or resupinate basidiomes; smooth,

grandinioid, odontioid, hydnoid or poroid hymenophores; two, four, six or eight sterigmata on

basidia; smooth or variously ornamented basidiospores; but all species of Trechisporales bear a

mono- or dimitic hyphal system with clamp connections. Most known species of Trechisporales

may be saprotrophs on wood, while some species are considered to be ectomycorrhizal fungi or at

least have a plant biotrophic lifestyle (Vanegas-León et al. 2019). For example, of the 672

nucleotide sequences of Trechispora in GenBank (https://www.ncbi.nlm.nih.gov/genbank/; access

on 2 June 2022), 258 (more than one third) are named as ‘uncultured Trechispora’ that were

generated mainly from rhizosphere and soil.

Within Trechisporales, Sistotremastrum and its recently segregated genus Sertulicium are

long known to be a separate lineage from Hydnodontaceae that accommodates all other genera in

the order (Larsson 2007). However, the Sistotremastrum lineage was designated as the

“Sistotremastrum family” and a new formal family name was not introduced (Larsson 2007). This

taxonomic treatment is mainly due either to restricted taxa sampled in phylogenetic analysis

(Larsson 2007) or to lack of reliable phylogenetic support (Spirin et al. 2021). Indeed, all available

phylogenetic studies on members of Trechisporales are based on only ITS and/or nrLSU regions

(e.g. Larsson et al. 2004, Liu et al. 2019, Spirin et al. 2021). These two regions are generally not

suitable for resolving fungal relationships at family and higher-taxonomic ranks when sampling

taxa comprehensively.

At the species level, numerous new species of Trechisporales have been recently described

worldwide in most of the accepted genera, such as Brevicellicium (Telleria et al. 2013a),

Dextrinocystis (Liu et al. 2019), Fibrodontia (Yurchenko & Wu 2014, Liu et al. 2021), Porpomyces

(Wu et al. 2015, Spirin et al. 2021), Sertulicium (Spirin et al. 2021), Sistotremastrum (Gruhn &

Alvarado 2021, Spirin et al. 2021), Subulicystidium (Volobuev 2016, Ordynets et al. 2018, Liu et

al. 2019), Trechispora (Ordynets et al. 2015, Chikowski et al. 2020, de Meiras-Ottoni et al. 2021,

Liu et al. 2022) and Tubulicium (Liu et al. 2019, Ushijima et al. 2019). However, without the help

of a well-defined phylogeny across Trechisporales, the taxonomic placement of certain new species

may be inappropriate even based on molecular phylogenetic analyses (Telleria et al. 2013b, 2014,

Xu et al. 2019, Furtado et al. 2021, Zong et al. 2021). Therefore, an emended classification of

Trechisporales is urgently needed to provide a backbone for clarifying the species diversity and

their appropriate placement within this order.

Via sampling a wider range of taxa especially from Asia Pacific, the current study aims to

construct a multilocus-based phylogenetic frame of Trechisporales within Agaricomycetes.

864

Accordingly, one new order, one new family, one new genus and 19 new species are described, and

the taxonomic position of certain previously known genera and species are adjusted.

Materials & Methods

Material deposition

Specimens studied are preserved at the Fungarium, Institute of Microbiology, Chinese

Academy of Sciences (HMAS), Beijing, China, the herbarium of Institute of Microbiology, Beijing

Forestry University (BJFC), Beijing, China, the herbarium of the Institute of Applied Ecology,

Chinese Academy of Sciences (IFP), Shenyang, China, and the National Herbarium of Victoria

(MEL), Melbourne, Australia.

Morphological examination

Macromorphological characters of basidiomes were examined with the aid of a Leica M 125

stereomicroscope (Wetzlar, Germany) at magnifications up to 100 times. Special color terms

followed Anonymous (1969). The microscopic procedure followed Wang et al. (2021). Specimen

sections were separately mounted in Cotton Blue, Melzer’s reagent and 5% potassium hydroxide.

Micromorphological characters were examined with an Olympus BX43 light microscope (Tokyo,

Japan) at magnifications up to 1000 times. All measurements were taken from the sections mounted

in Cotton Blue. When presenting the variation in the size of the basidiospores, 5% of measurements

were excluded from each end of the range and are given in parentheses. Unless specified, all

measurements exclude the ornamentations of basidiospores. In the morphological descriptions, L

stands for the length of arithmetic average of all measured basidiospores, W for the width of

arithmetic average of all measured basidiospores, Q for variation in the ratio of L to W among the

studied specimens, and n (a/b) for number of basidiospores (a) measured from given number of

specimens (b). Drawings were made with the aid of a drawing tube. The ornamentations of

basidiospores were examined with a Hitachi SU8010 scanning election microscope (Tokyo, Japan).

The sections from hymenophores of basidiomes were sprayed with gold and platinum using Leica

EM ACE600 (Wetzlar, Germany).

Molecular sequencing

Total DNA was extracted from basidiomes of dry specimens as templates for subsequent

PCR amplifications using the CTAB rapid plant genome extraction kit (Aidlab Biotechnologies

Co., Ltd, Beijing, China) according to the manufacturer’s instructions. The ITS, nrLSU, tef1-α,

rpb2 and mtSSU regions were amplified with primer pairs ITS5/ITS4 (White et al. 1990),

LR0R/LR7 (Vilgalys & Hester 1990), 983F/1567R (Rehner & Buckley 2005), RPB2-f5F/RPB2-

b7.1R (Liu et al. 1999, Matheny 2005) and MS1/MS2 (White et al. 1990), respectively, using

2×EasyTaq® PCR SuperMix (TransGen Biotech Co., Ltd, Beijing, China). The PCR procedure was

as follows: for ITS and tef1-α, initial denaturation at 95°C for 3 min, followed by 35 cycles at 94°C

for 40 s, 54°C for 45 s and 72°C for 1 min, and a final extension of 72°C for 10 min; for nrLSU,

initial denaturation at 94°C for 1 min, followed by 34 cycles at 94°C for 30 s, 50°C for 1 min, 72

°C for 1.5 min, and a final extension of 72°C for 10 min; for rpb2, initial denaturation at 94 °C for

2 min, followed by 10 cycles at 94°C for 45 s, 60°C for 45 s (minus 1°C per cycle) and 72 °C for

1.5 min, then followed by 36 cycles at 94 °C for 45 s, 53 °C for 1 min and 72°C for 1.5 min, and a

final extension of 72°C for 10 min; for mtSSU, initial denaturation at 95°C for 3 min, followed by

35 cycles at 94°C for 40 s, 53°C for 45 s and 72°C for 1 min, and a final extension of 72°C for 10

min. The PCR products were sequenced with the same primers used in PCR amplification at the

Beijing Genomics Institute, China. All newly generated sequences were submitted to GenBank

(Table 1).

865

Phylogenetic analysis

Besides the newly generated sequences for this study, additional related sequences were downloaded from GenBank (Table 1) and incorporated

together for phylogenetic analyses. Five datasets were employed to explore the relationships among members of Trechisporales. The combined dataset

of ITS, nrLSU, tef1-α and rpb2 regions (1) was used to clarify the phylogenetic consistency of members of Trechisporales within Agaricomycetes.

Besides representatives from the genera of Trechisporales, species from additional 20 orders within Agaricomycetes were also included as the ingroup

taxa, while two species from Tremellomycetes were selected as the outgroup taxa (Table 1). The combined dataset of ITS, nrLSU, tef1-α, rpb2 and

mtSSU regions (2) was used to further clarify the phylogenetic relationship among genera within Trechisporales. The representatives from the genera

of Trechisporales were included as the ingroup taxa, while two species from Polyporales were selected as the outgroup taxa (Table 1) according to the

topology inferred from the dataset (1). Three combined datasets of ITS and nrLSU regions were used to explore the relationships among species within

the genera Fibrodontia (3), Subulicystidium (4) and Trechispora (5), respectively. Besides the species from each target genus listed in Table 1, two

species from Dextrinocystis and Tubulicium were also included as the ingroup taxa, and a species from Porpomyces was selected as the outgroup taxon

according to the topology inferred from the dataset (2). Regarding the dataset (5) for Trechispora, sequences from named and unnamed species of

Scytinopogon listed in Table 1 were also included as the ingroup taxa.

Table 1 Species and sequences used in the phylogenetic analyses.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Agaricomycetes

Agaricales

Lepiota cristata

ZRL20151133 #

China

LT716026

KY418841

KY419048

KY418992

Marasmius oreades

ZRL2015086 #

China

LT716048

KY418864

KY419066

KY419010

Psathyrella candolleana

ZRL20151400 #

China

LT716063

KY418879

KY419075

KY419024

Amylocorticiales

Amylocorticium

cebennense

HHB-2808 #

USA

GU187505

GU187561

GU187675

GU187770

Plicaturopsis crispa

MR00464 #

LR694209

LR694187

LR694225

LR694281

Atheliales

Athelia arachnoidea

CBS 418.72 #

Netherlands

GU187504

GU187557

GU187672

GU187769

Byssocorticium

atrovirens

BS1710033 #

Sweden

LR694198

LR694175

LR694214

LR694271

Piloderma fallax

S-12 #

Finland

GU187535

GU187591

GU187738

GU187797

Auriculariales

Auricularia heimuer

Xiaoheimao #

China

LT716074

KY418890

KY419083

KY419035

Exidia crenata

PBM2527 #

DQ241774

AY700191

DQ408144

Boletales

Coniophora arida

FP104367 #

USA

GU187510

GU187573

GU187684

GU187775

Gomphidius roseus

MB 95-038 #

Germany

DQ534570

DQ534669

GU187702

GU187818

Gyrodontium sacchari

MUCL40589 #

French Guiana

GU187522

GU187579

GU187703

GU187764

Cantharellales

Clavulina sp.

AFTOL-667 #

DQ202266

AY745694

DQ028589

DQ366286

Hydnum albomagnum

AFTOL-471 #

USA

DQ218305

AY700199

DQ234568

DQ234553

Sistotrema confluens

AFTOL-613 #

DQ267125

AY647214

DQ381837

Corticiales

Punctularia

strigosozonata

AFTOL-1248 #

DQ398958

AF518642

DQ408147

DQ381843

866

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Vuillemenia comedens

AFTOL-1247 #

DQ398959

AF518666

DQ381844

Geastrales

Geastrum recolligens

OSC41996 #

DQ218486

DQ219230

DQ219052

Pyrenogaster pityophilus

OSC59743 #

DQ218519

DQ219232

DQ219057

Gloeophyllales

Gloeophyllum trabeum

1320 #

USA

HM536094

HM536067

HM536113

HM536112

Heliocybe sulcata

IBUG-9930 #

Mexico

HM536095

HM536069

HM536115

HM536114

Gomphales

Clavariadelphus

truncatus

OSC67280 #

AY574649

DQ219240

DQ219064

Kavinia alboviridis

O102140 #

AY574692

DQ219250

DQ219073

Hymenochaetales

Fomitiporia aethiopica

MUCL 44777 #

Ethiopia

NR137575

NG059421

GU461893

JQ087956

Rigidoporus corticola

ZRL20151459 #

China

LT716075

KY418899

KY419087

KY419038

Peniophorella

praetermissa

AFTOL-ID 518 #

AY854081

AY700185

AY787221

Hysterangiales

Aroramyces

gelatinosporus

H4010 #

China: Hunan

DQ218524

DQ219118

DQ218941

Chondrogaster

pachysporus

OSC49298 #

DQ218538

DQ219136

DQ218958

Jaapiales

Jaapia argillacea

CBS 252.74 #

Netherlands

GU187524

GU187581

GU187711

GU187788

Lepidostromatales

Lepidostroma vilgalysii

RV-MX16 #

Brazil

JN698907

JN698908

Sulzbacheromyces

caatingae

Sulzbacher 1479 #

Mexico

KC170320

KC170318

Phallales

Dictyophora duplicata

OSC38819 #

DQ218481

DQ219265

DQ219087

Phallus hadriani

AFTOL-683 #

DQ404385

AY885165

DQ435792

DQ408114

Polyporales

Neofavolus alveolaris

Dai 11290 * #

China: Hainan

KU189768

KU189799

KU189913

KU189982

KU189949

Polyporus squamosus

Cui 10595 * #

China: Sichuan

KU189778

KU189809

KU189925

KU189988

KU189960

Climacodon

septentrionalis

ZW #

AY854082

AY684165

AY885151

AY780941

Phlebia radiate

AFTOL-484 #

AY854087

AF287885

AY885156

AY218502

Russulales

Bondarzewia montana

AFTOL-ID 452 #

Canada

DQ200923

DQ234539

DQ059044

AY218474

Heterobasidion annosum

AFTOL-ID 470 #

DQ206988

AF287866

DQ028584

AH013701

Lactifluus deceptivus

AFTOL-ID 682 #

USA

AY854089

AY631899

AY885158

AY803749

Sebacinales

Tremellodendron

pallidum

AFTOL- 699 #

DQ411526

AY745701

DQ029196

DQ408132

Piriformospora indica

AFTOL-612 #

DQ411527

AY293202

AJ249911

DQ408131

Sistotremastrales

Sertulicium

lateclavigerum

Spirin 13457 *

Slovenia

MW049161

Sertulicium guttuliferum

He 3338 * #

China: Yunnan

MK204540

MK204552

Sertulicium jacksonii

Svantesson 699 *

Norway

MN937562

867

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Sertulicium

niveocremeum

Larsson 13727 * #

France

MN937563

Sertulicium vernale

Söderholm 3886 *

Finland

MT002311

MT664174

Sistotremastrum

aculeatum

Miettinen 10380.1 *

China: Yunnan

MN991176

MW045423

Sistotremastrum

aculeicrepitans

Larsson 16097 *

Brazil

MN937564

Sistotremastrum

confusum

Larsson 16004 *

Brazil

MN937567

Sistotremastrum

denticulatum

Motato-Vásquez 894 *

Brazil

MN954694

MW045424

Sistotremastrum

fibrillosum

GG GUY12-180 *

MG913222

MG913208

Sistotremastrum

geminum

Miettinen 14333 *

MN991177

Sistotremastrum

induratum

Spirin 8598 *

USA

MT002324

MT664173

Sistotremastrum mendax

Larsson 12022 *

Norway

MN937570

Sistotremastrum rigidum

Motato-Vásquez 833 *

Brazil

MN954693

MW045435

Sistotremastrum

suecicum

KHL-11849 * #

Sweden

MN937571

Sistotremastrum vigilans

Fonneland 2011-78 *

Norway

MN937572

Sistotremastrum sp.

LWZ 20171015-32 *

Vietnam

OM523376

OM339204

Sistotremastrum sp.

LWZ 20191107-25 * #

China: Yunnan

MW477771

MW474864

MW478703

MW478712

OM422784

Sistotremastrum sp.

LWZ 20191207-26 * #

Malaysia

OM523377

OM339205

OM416796

OM416816

OM422785

Stereopsidales

Stereopsis radicans

OLR45395 #

Belize

KC203496

KC203516

KC203502

Stereopsis globose

KHL 12592 #

Costa Rica

KC203495

KC203515

KC203501

Thelephorales

Boletopsis leucomelaena

PBM2678 #

USA

DQ484064

DQ154112

GU187763

GU187820

Thelephora ganbajun

ZRL20151295 #

China

LT716082

KY418908

KY419093

KY419043

Trechisporales

Allotrechispora

daweishanensis

CLZhao 17860 *

China: Yunnan

MW302337

MW293866

Allotrechispora gatesiae

LWZ 20180515-18 * #

Australia

OM523378

OM339206

OM416817

Allotrechispora gatesiae

LWZ 20180515-20 *

Australia

OM523379

OM339207

OM416797

Allotrechispora xantha

CLZhao 2632 *

China: Yunnan

MW302339

MW293868

Brevicellicium

atlanticum

LISU 178566 *

Portugal

HE963773

HE963774

Brevicellicium sp.

LWZ 20190809-10b * #

China: Shandong

OM339208

OM416798

OM416818

OM422800

Brevicellicium sp.

LWZ 20190918-13 *

China: Sichuan

OM523380

OM416799

OM422792

868

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Dextrinocystis

calamicola

He 5693 *

China: Fujian

MK204533

MK204546

OM416800

Dextrinocystis

calamicola

He 5701 * #

China: Fujian

MK204534

MK204547

OM416801

OM416819

Fibrodontia alba

EYu 110703-25

China: Taiwan

KC928274

KC928275

Fibrodontia alba

He 3392

China: Yunnan

OM523381

OM339209

Fibrodontia alba

He 3432

China: Yunnan

OM523382

OM339210

Fibrodontia alba

He 3475

China: Yunnan

OM523383

Fibrodontia alba

He 3501

China: Yunnan

OM523384

OM339211

Fibrodontia alba

He 4243

China: Jiangxi

OM523385

OM339212

Fibrodontia alba

He 4255

China: Jiangxi

OM523386

OM339213

Fibrodontia alba

He 4380

China: Jiangxi

OM523387

OM339214

Fibrodontia alba

He 4761

China: Guangxi

MK204529

MK204541

Fibrodontia alba

He 5953

China: Jiangxi

OM523388

OM339215

Fibrodontia alba

He 5954a

China: Jiangxi

OM523389

OM339216

Fibrodontia alba

LWZ 20170820-34 * #

China: Hubei

MT802102

MT802108

MW478698

MW478706

OM422802

Fibrodontia alba

LWZ 20170820-39

China: Hubei

OM523390

OM339217

Fibrodontia alba

LWZ 20170820-40

China: Hubei

OM523391

OM339218

Fibrodontia alba

LWZ 20180415-18

Malaysia

OM523392

OM339219

Fibrodontia alba

LWZ 20180923-2

China: Yunnan

OM523393

Fibrodontia alba

LWZ 20180923-20

China: Yunnan

OM523394

OM339220

Fibrodontia alba

LWZ 20180923-4

China: Yunnan

MT802107

MT802101

Fibrodontia alba

LWZ 20191207-1 *

Malaysia

OM523395

OM339221

OM416802

OM416820

OM422803

Fibrodontia alba

TNM F24944

China: Taiwan

NR153983

NG060401

Fibrodontia alba

Yuan 1491

China: Yunnan

OM523396

Fibrodontia

austrosinensis

He 6283 *

China: Yunnan

MT802110

MT802104

MW478699

MW478710

Fibrodontia

austrosinensis

LWZ 20190820-11b *

China: Sichuan

MT802111

MT802105

MW478700

MW478709

Fibrodontia

austrosinensis

He 3453

China: Yunnan

MT802109

MT802103

Fibrodontia brevidens

He 3559 *

China: Hainan

MK204528

MW478701

MW478707

OM422791

Fibrodontia brevidens

Wu 9807-16

KC928276

KC928277

Fibrodontia gossypina

AFTOL-ID 599

DQ249274

AY646100

Fibrodontia subalba

Dai 15931

China: Xinjiang

MT802106

MT802100

Fibrodontia

subaustrosinensis

He 6033

China: Hainan

OM523397

OM339222

869

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Fibrodontia

subaustrosinensis

He 6279

China: Yunnan

OM523398

OM339223

Luellia cystidiata

JHP-09.455 * #

Portugal

MW371211

Luellia recondita

O-F-253622 *

Norway

UDB038222

Porpomyces mucidus

Dai 12692 *

Czech Republic

KT157833

KT157838

Porpomyces submucidus

Cui 5183

China: Hainan

KT152143

KT152145

Porpomyces submucidus

Dai 13708 * #

China: Hainan

KT152144

KT152146

MW478702

Pteridomyces galzinii

Bernicchia 8122 *

Italy

MN937559

Pteridomyces galzinii

GB0150230 * #

Estonia

LR694210

LR694188

LR694226

LR694282

Scytinopogon

angulisporus

TENN-F-066226

USA

MK278574

Scytinopogon

angulisporus

TFB13611

USA

JQ684661

Scytinopogon sp.

BAB5120

India

KT804576

Scytinopogon sp.

MEL:2382675

Australia

KP013038

Scytinopogon sp.

MEL:2382987

Australia

KP012842

Scytinopogon sp.

MEL:2382992

Australia

KP012847

Scytinopogon sp.

MEL2382623

Australia

KP012986

Scytinopogon sp.

MEL2382744

Australia

KP012927

Subulicystidium

acerosum

He 3804

China: Guizhou

MK204539

MK204543

Subulicystidium boidinii

KHL 12830

Costa Rica

MH041537

MH041570

Subulicystidium

brachysporum

He 2207

USA

MK204532

MK204549

Subulicystidium

brachysporum

KHL 16100

Brazil

MH000599

Subulicystidium daii

LWZ 20170820-35 *

China: Hubei

OM523399

OM339224

OM422786

Subulicystidium daii

Xiong 221

China: Guangxi

OM523400

Subulicystidium

fusisporum

KHL 10360

Puerto Rico

MH041535

MH041567

Subulicystidium

grandisporum

506781

Costa Rica

MH041547

MH041592

Subulicystidium

harpagum

L 1726a

Reunion

MH041532

MH041588

Subulicystidium

inornatum

KHL 10444

Puerto Rico

MH041558

MH041569

Subulicystidium

longisporum

KHL 14229

Sweden

MH000601

870

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Subulicystidium

meridense

Hjm 16400

Brazil

MH041538

MH041604

Subulicystidium nikau

L 1296

Reunion

MH041513

MH041565

Subulicystidium

obtusisporum

Piepenbrink & Lotz-

Winter W213-3-I O

Germany

MH041521

MH041566

Subulicystidium

parvisporum

L 0140

Reunion

MH041529

MH041590

Subulicystidium

perlongisporum

KHL 16062

Brazil

MH000600

Subulicystidium

perlongisporum

TU 124388

Italy

UDB028355

Subulicystidium

rarocrystallinum

918488

Colombia

MH041512

MH041564

Subulicystidium

robustius

KHL 10813

Jamaica

MH041514

MH041608

Subulicystidium

tedersooi

TU 110894

Vietnam

UDB014161

Subulicystidium tropicum

He 3583

China: Hainan

Subulicystidium tropicum

He 3968 * #

China: Hainan

MK204531

MK204544

MW478711

OM422787

Subulicystidium tropicum

LWZ 20180411-4 *

Malaysia

OM523401

OM339225

OM422788

Subulicystidium sp.

LWZ 20170816-7 *

China: Hubei

OM523402

OM416803

Subulicystidium sp.

LWZ 20180804-5 *

China: Beijing

OM523403

OM339226

OM416804

Subulicystidium sp.

LWZ 20190816-24a

China: Sichuan

OM523404

Suillosporium

cystidiatum

Spirin 3830 * #

Russia

MN937573

Trechispora alnicola

AFTOL-ID 665

DQ411529

AY635768

Trechispora araneosa

KHL 8570

Sweden

AF347084

Trechispora bambusicola

CLZhao 3302

China: Yunnan

MW544021

MW520171

Trechispora bambusicola

CLZhao 3305

China: Yunnan

MW544022

MW520172

Trechispora bambusicola

He 3381

China: Yunnan

OM523405

OM339227

Trechispora bambusicola

LWZ 20191107-5

China: Yunnan

OM523406

OM339228

Trechispora bispora

CBS 142.63

Australia

MH858241

MH869842

Trechispora

candidissima

Dai 7092

China: Jilin

OM523407

OM339229

Trechispora caucasica

O-F-253764

Sweden

UDB038261

Trechispora

caulocystidiata

FLOR 56314

Brazil

MK458772

871

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora

chaibuxiensis

He 5072

China: Hubei

OM523408

OM339230

Trechispora

chaibuxiensis

LWZ 20170814-34

China: Hubei

OM523409

OM339231

Trechispora

chaibuxiensis

LWZ 20170814-35

China: Hubei

OM523410

OM339232

Trechispora

chaibuxiensis

LWZ 20170814-36

China: Hubei

OM523411

OM339233

Trechispora

chaibuxiensis

LWZ 20170814-42

China: Hubei

OM523412

OM339234

Trechispora chartacea

FLOR 56185

Brazil

MK458775

Trechispora cohaerens

TUF115568

Estonia

UDB016421

Trechispora cf.

cohaerens

UC2022832

USA

KP814538

Trechispora confinis

KHL 11064

Sweden

AF347081

Trechispora confinis

LWZ 20200809-30b

China: Sichuan

OM523413

Trechispora confinis

LWZ 20210920-23b

China: Hubei

OM523414

OM339235

Trechispora confinis

SFC20180710-18

South Korea

MK992834

Trechispora confinis

SFC20180710-23

South Korea

MK992839

Trechispora constricta

Dai 10488

China: Jiangxi

OM523415

Trechispora constricta

Dai 10534

China: Jiangxi

OM523416

Trechispora constricta

He 5899

China:

Guangdong

OM523417

OM339236

Trechispora constricta

LWZ 20210924-30a

China: Henan

OM523418

OM339237

Trechispora copiosa

AMO423

Brazil

MN701014

MN687972

Trechispora copiosa

AMO453

Brazil

MN701018

MN687975

Trechispora crystallina

LWZ 20170729-2

China: Inner

Mongolia

OM523419

OM339238

Trechispora crystallina

LWZ 20171013-7

Vietnam

OM523420

OM339239

Trechispora cyatheae

FR 0219442

France: La

Réunion

UDB024014

UDB024015

Trechispora cyatheae

FR 0219443

France: La

Réunion

UDB024016

UDB024017

Trechispora cyatheae

FR 0219446

France: La

Réunion

UDB024020

UDB024021

Trechispora

damansaraensis

He 6415

Malaysia

OM523421

OM339240

872

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora

damansaraensis

LWZ 20180417-26

Malaysia

OM339241

Trechispora dealbata

FLOR 56182

Brazil

MK458776

Trechispora dealbata

FLOR 56183

Brazil

MK458777

Trechispora dentata

Dai 15277

China: Hainan

OM523505

Trechispora dentata

Dai 22565

China

OK298491

OM049408

Trechispora dimitiella

Dai 17772

Singapore

OM523450

Trechispora dimitiella

Dai 17891

Singapore

OM523451

Trechispora dimitiella

Dai 21181

China

OK298493

OK298949

Trechispora dimitiella

Dai 21931

China

OK298492

OK298948

Trechispora

echinocristallina

FR 0219445

France: La

Réunion

UDB024018

UDB024019

Trechispora

echinocristallina

TUF 110414

Papua New

Guinea

UDB013050

Trechispora echinospora

E09/60-06

Equatorial Guinea

JX392847

JX392848

Trechispora echinospora

E11/37-03

Equatorial Guinea

JX392845

JX392846

Trechispora echinospora

E11/37-05

Equatorial Guinea

JX392849

Trechispora echinospora

E11/37-10

Equatorial Guinea

JX392850

JX392851

Trechispora echinospora

E11/37-11

Equatorial Guinea

JX392852

Trechispora echinospora

E11/37-12

Equatorial Guinea

JX392853

JX392854

Trechispora farinacea

KHL 8451

AF347082

Trechispora farinacea

KHL 8454

AF347083

Trechispora farinacea

KHL 8793

Sweden

AF347089

Trechispora farinacea

MA-Fungi 79474

JX392855

Trechispora farinacea

TUB 011825

Germany

EU909231

Trechispora fimbriata

CLZhao 4154

China: Yunnan

MW544023

MW520173

Trechispora fimbriata

Cui 7962

China: Yunnan

OM523422

Trechispora fimbriata

Dai 17612

China: Yunnan

OM523423

OM339242

Trechispora fimbriata

He 4873

China: Guangxi

OM523424

OM339243

Trechispora fimbriata

He 6134

China: Yunnan

OM523425

OM339244

Trechispora fimbriata

Xiong 21

China: Hunan

OM523426

Trechispora fissurata

CLZhao 4571

China: Yunnan

MW544027

MW520177

Trechispora fissurata

He 6190

China: Yunnan

OM523427

OM339245

Trechispora fissurata

He 6322

China: Yunnan

OM523428

OM339246

Trechispora fissurata

LWZ 20171015-11

Vietnam

OM523429

OM339247

Trechispora fissurata

LWZ 20171015-16

Vietnam

OM523430

OM339248

Trechispora fissurata

LWZ 20171015-35 *

Vietnam

OM523431

OM339249

OM416821

OM422793

Trechispora fissurata

LWZ 20171015-4

Vietnam

OM523432

OM339250

873

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora fissurata

LWZ 20180922-51

China: Yunnan

OM523433

OM339251

Trechispora fissurata

LWZ 20191110-9

China: Yunnan

OM523434

OM339252

Trechispora foetida

FLOR 56315

Brazil

MK458769

Trechispora fragilis

Dai 20535

China

OK298494

OK298950

Trechispora gelatinosa

AMO1139

Brazil

MN701021

MN687978

Trechispora gelatinosa

AMO824

Brazil

MN701020

MN687977

Trechispora gracilis

LWZ 20170814-17

China: Hubei

OM523435

OM339253

Trechispora gracilis

LWZ 20210626-5b

China: Jiangxi

OM523436

OM339254

Trechispora gracilis

LWZ 20210919-9a

China: Hubei

OM523437

OM339255

Trechispora gracilis

LWZ 20210922-7b

China: Hubei

OM523438

OM339256

Trechispora havencampii

SFSU DED8300 *

Africa

NR154418

NG059993

Trechispora hondurensis

HONDURAS19-F016a

Honduras

MT571523

MT636540

Trechispora hondurensis

HONDURAS19-F016b

Honduras

MT636541

Trechispora

hymenocystis

Dai 2247

Finland

OM523439

Trechispora

hymenocystis

KHL 16444

Norway

MT816397

Trechispora

hymenocystis

KHL 8795

Sweden

AF347090

Trechispora incisa

EH 24/98

AF347085

Trechispora incisa

GB-0090521

Sweden

KU747093

Trechispora incisa

GB-0090648

Sweden

KU747095

KU747087

Trechispora incisa

GB-0105521

Sweden

KU747086

Trechispora incisa

GB-0105526

Sweden

KU747094

Trechispora incisa

He 5008 * #

China: Hebei

OM523440

OM339257

OM416805

OM416822

Trechispora invisitata

UC2022935

USA

KP814182

Trechispora invisitata

UC2023088

USA

KP814425

Trechispora kavinioides

KGN 981002

Norway

AF347086

Trechispora laevis

TUF115551

Estonia

UDB016406

Trechispora laevispora

Dai 21655

China

OK298495

OM108710

Trechispora larssonii

He 5450

China: Guizhou

OM523441

OM339258

Trechispora larssonii

LWZ 20190817-11a *

China: Sichuan

OM523442

OM339259

OM416823

OM422795

Trechispora larssonii

LWZ 20200818-10b

China: Sichuan

OM339260

Trechispora latehypha

He 3924

China: Hainan

OM523443

OM339261

Trechispora latehypha

He 4472

China: Fujian

OM523444

Trechispora latehypha

He 5438 *

China: Guizhou

OM523445

OM416806

OM422798

Trechispora latehypha

He 5848

Sri Lanka

OM523446

OM339262

Trechispora latehypha

LWZ 20170611-16

China: Hainan

OM523447

OM339263

874

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora longiramosa

HG 140168

China: Guizhou

OM523448

OM339264

Trechispora longiramosa

CH 19233

China: Guizhou

OM523449

Trechispora malayana

Dai 17876

Singapore

OM523452

OM339265

Trechispora malayana

He 4156

Thailand

OM523453

OM339266

Trechispora mellina

URM85756

Brazil

MH280000

Trechispora microspora

O-F-253725

Sweden

UDB038247

Trechispora minispora

AM170

Mexico

MK328885

MK328894

Trechispora minispora

AM176

Mexico

MK328886

MK328895

Trechispora mollis

URM 85884

Brazil

MK514945

MH280003

Trechispora mollis

URM 85885

Brazil

MT423667

Trechispora mollusca

CBS 439.48

Canada

MH856428

Trechispora mollusca

Cui 2455

China: Qinghai

OM339267

Trechispora mollusca

Dai 1931

Estonia

OM523454

Trechispora mollusca

Dai 6174

China: Anhui

OM339268

Trechispora mollusca

Dai 6191

China: Anhui

OM523455

OM339269

Trechispora mollusca

Dai 7097

China: Jilin

OM523456

Trechispora mollusca

Dai 11085

China: Inner

Mongolia

OM523457

OM339270

Trechispora mollusca

Dai 11157

China: Inner

Mongolia

OM523458

OM339271

Trechispora mollusca

Dai 13289

China: Gansu

OM523459

OM339272

Trechispora mollusca

DLL2010-077

USA

JQ673209

Trechispora mollusca

DLL2011-186

USA

KJ140681

Trechispora mollusca

Li 1449

China: Hubei

OM523460

Trechispora nivea

GB-0087593

Sweden

KU747088

Trechispora nivea

GB-0102694

Sweden

KU747089

Trechispora nivea

LWZ 20180804-3

China: Beijing

OM523461

OM339273

Trechispora nivea

MA-Fungi 74044

JX392832

JX392833

Trechispora nivea

MA-Fungi 76253

JX392837

Trechispora nivea

MA-Fungi 76254

JX392834

Trechispora nivea

MA-Fungi 76257

JX392826

JX392827

Trechispora nivea

MA-Fungi 82479

JX392828

Trechispora nivea

MA-Fungi 82481

JX392831

Trechispora nivea

MA-Fungi 82483

JX392838

Trechispora nivea

O 90120

Tanzania

KU747103

KU747092

Trechispora cf. nivea

F-506673

Venezuela

KU747091

Trechispora pallescens

FLOR 56184

Brazil

MK458767

875

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora pallescens

FLOR 56188

Brazil

MK458774

Trechispora pallescens

He 5192

Vietnam

MK204553

Trechispora pallescens

SC1 *

MZ518207

MZ518091

Trechispora aff.

pallescens

AM21

Mexico

MK328883

MK328893

Trechispora aff.

pallescens

RL115

Mexico

MK328887

MK328896

Trechispora aff.

pallescens

RL132

Mexico

MK328889

MK328898

Trechispora aff.

pallescens

RL133

Mexico

MK328890

MK328899

Trechispora papillosa

AMO713

Brazil

MN701022

MN687979

Trechispora papillosa

AMO795

Brazil

MN701023

MN687981

Trechispora regularis

KHL 10881

Jamaica

AF347087

Trechispora rigida

URM 85754

Brazil

MT406381

MH279999

Trechispora robusta

FLOR 56179

Brazil

MK458770

Trechispora robusta

FLOR 56190

Brazil

MK458768

Trechispora scaber

FLOR 56189

Brazil

MK458773

Trechispora sinensis

Dai 7227

China: Fujian

OM523462

Trechispora sinensis

Dai 11239

China: Jiangsu

OM523463

Trechispora sinensis

He 3714

China: Jilin

OM523464

OM339274

Trechispora sinensis

He 4314

China: Jiangxi

OM523465

OM339275

Trechispora sinensis

He 4668

China: Liaoning

OM523466

Trechispora sinensis

He 4698

China: Guangxi

OM523467

OM339276

Trechispora sinensis

He 5446

China: Guizhou

OM523468

OM339277

Trechispora sinensis

He 5491

China: Chongqing

OM523469

OM339278

Trechispora sinensis

He 5649

China: Hunan

OM523470

OM339279

Trechispora sinensis

He 5652

China: Hunan

OM523471

OM339280

Trechispora sinensis

He 5898

China:

Guangdong

OM523472

OM339281

Trechispora sinensis

LWZ 20170805-14

China: Liaoning

OM523473

Trechispora sinensis

LWZ 20170814-11

China: Hubei

OM523474

OM339282

Trechispora sinensis

LWZ 20170814-27

China: Hubei

OM523475

OM339283

Trechispora sinensis

LWZ 20170814-28

China: Hubei

OM523476

OM339284

Trechispora sinensis

LWZ 20170815-38 *

China: Hubei

OM523477

OM339285

OM416807

Trechispora sinensis

LWZ 20170816-16

China: Hubei

OM523478

OM339286

Trechispora sinensis

LWZ 20170816-35 *

China: Hubei

OM523479

OM339287

OM416808

OM422796

Trechispora sinensis

LWZ 20170817-5

China: Hunan

OM523480

OM339288

876

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora sinensis

LWZ 20170909-11

China: Beijing

OM523481

OM339289

Trechispora sinensis

LWZ 20180804-19

China: Beijing

OM523482

OM339290

Trechispora sinensis

LWZ 20180804-20 *

China: Beijing

OM523483

OM339291

OM416809

Trechispora sinensis

LWZ 20210923-15b

China: Henan

OM523484

OM339292

Trechispora sinensis

LWZ 20210925-13b

China: Henan

OM523485

OM339293

Trechispora sinensis

LWZ 20210925-3a

China: Henan

OM523486

OM339294

Trechispora sinensis

LWZ 20210928-7

China: Guizhou

OM523487

OM339295

Trechispora sinensis

LWZ 20210928-9

China: Guizhou

OM523488

OM339296

Trechispora sinensis

Wei 7909

China: Beijing

OM523489

OM339297

Trechispora stevensonii

KHL 14654

Norway

MH290762

Trechispora stevensonii

MA-Fungi 70645

JX392843

JX392844

Trechispora stevensonii

MA-Fungi 70669

JX392841

JX392842

Trechispora stevensonii

TU 115499

Estonia

UDB016467

Trechispora subfissurata

He 3907

China: Hainan

OM523490

OM339298

Trechispora subfissurata

LWZ 20190613-48

China:

Guangdong

OM523491

Trechispora

subhymenocystis

LWZ 20190818-29b *

China: Sichuan

OM523492

OM339299

OM416824

OM422794

Trechispora

subhymenocystis

LWZ 20190818-32b

China: Sichuan

OM339300

Trechispora subsinensis

He 4122

Thailand

OM523493

OM339301

Trechispora subsinensis

He 4125

Thailand

OM523494

OM339302

Trechispora subsinensis

He 5894

China:

Guangdong

OM523495

OM339303

Trechispora subsinensis

LWZ 20190611-19

China:

Guangdong

OM523496

Trechispora subsinensis

LWZ 20190611-9

China:

Guangdong

OM523497

OM339304

Trechispora

subsphaerospora

KHL 8511

Sweden

AF347080

Trechispora taiwanensis

He 4571 *

China: Taiwan

OM523498

OM339305

OM416825

OM422799

Trechispora taiwanensis

He 4574

China: Taiwan

OM339306

Trechispora termitophila

AMO390

Brazil

MN701024

MN687982

Trechispora termitophila

AMO396

Brazil

MN701025

MN687983

Trechispora termitophila

AMO893

Brazil

MN701026

MN687984

Trechispora thailandica

He 4101 *

Thailand

OM523499

OM339307

OM416810

Trechispora thailandica

He 4112

Thailand

OM523500

OM339308

Trechispora thailandica

He 4114

Thailand

OM523501

OM339309

877

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora thelephora

URM 85757

Brazil

MH280001

Trechispora thelephora

URM85758

Brazil

MH280002

Trechispora thelephora

UTC 252606

Belize

HM104485

Trechispora torrendii

KHL 15465

Brazil

MH290759

Trechispora torrendii

URM85886

Brazil

MK515148

MH280004

Trechispora tropica

LWZ 20170613-14

China: Hainan

OM523502

OM339310

Trechispora tropica

LWZ 20170613-16

China: Hainan

OM523503

OM339311

Trechispora tropica

LWZ 20171015-22

Vietnam

OM523504

OM339312

Trechispora sp.

AMO799

Brazil

MN701008

MN687969

Trechispora sp.

Dai 16179

China: Hainan

OM523506

OM339313

Trechispora sp.

Dai 17433

Brazil

OM523507

OM339314

Trechispora sp.

Dai 18781

Australia

OM523508

OM339315

Trechispora sp.

Dai 22173

China

OK298496

OK298951

Trechispora sp.

Dai 22174

China

OK298497

OK298952

Trechispora sp.

F909645

Sweden

JX392817

JX392818

Trechispora sp.

He 3431

China: Yunnan

OM523509

OM339316

Trechispora sp.

He 3984

China: Hainan

OM523510

OM339317

Trechispora sp.

He 3996

China: Hainan

OM523511

Trechispora sp.

He 4503

China: Fujian

OM523512

OM339318

Trechispora sp.

He 4641

China: Taiwan

OM523513

OM339319

Trechispora sp.

He 5812

Sri Lanka

OM523514

OM339320

Trechispora sp.

He 6400

Malaysia

OM523515

OM339321

Trechispora sp.

HG 19350

China: Yunnan

OM523516

Trechispora sp.

KHL 10715

AF347088

Trechispora sp.

KHL 16968

Brazil

MH290763

Trechispora sp.

LWZ 20170805-15

China: Liaoning

OM523517

Trechispora sp.

LWZ 20170815-20 *

China: Hubei

OM523518

OM339322

OM416811

OM422804

Trechispora sp.

LWZ 20171015-17

Vietnam

OM523519

OM339323

Trechispora sp.

LWZ 20180512-12 *

Australia

OM523520

OM339324

OM416812

Trechispora sp.

LWZ 20180513-8

Australia

OM523521

OM339325

Trechispora sp.

LWZ 20180517-43 *

Australia

OM523522

OM339326

OM416813

OM416826

Trechispora sp.

LWZ 20180517-44 *

Australia

OM523523

OM339327

OM416827

OM422797

Trechispora sp.

LWZ 20180517-45

Australia

OM523524

OM339328

Trechispora sp.

LWZ 20190816-39a *

China: Sichuan

OM523525

OM339329

OM416814

Trechispora sp.

LWZ 20191206-27

Malaysia

OM523526

OM339330

Trechispora sp.

LWZ 20191208-10

Malaysia

OM523527

Trechispora sp.

LWZ 20200921-33a

China: Sichuan

OM523528

OM339331

Trechispora sp.

LWZ 20210918-10a

China: Hubei

OM523529

OM339332

878

Table 1 Continued.

Class/Order

Species

Vouchers

Country

ITS

nrLSU

tef1-α

rpb2

mtSSU

Trechispora sp.

LWZ 20210921-7a

China: Hubei

OM523530

OM339333

Trechispora sp.

NCC16

Brazil

MN701007

MN687968

Trechispora sp.

SP48

Brazil

MN701005

MN687965

Trechispora sp.

Yuan 6129

China: Guangxi

OM523531

Trechispora sp.

ZP-1029

China

OM523532

Trechispora sp.

ZP-3658

China

OM523533

Tubulicium bambusicola

He 4776 *

China: Guizhou

MK204536

MK204551

OM422789

Tubulicium

raphidisporum

He 3191 *

China: Yunnan

OM523534

OM339334

OM422801

Tubulicium sp.

LWZ 20180414-5 * #

Malaysia

OM523535

OM339335

OM416815

OM416828

OM422790

Tremellomycetes

Tremellales

Bullera alba

CBS 501 #

AF444368

AF075500

KF037016

KF036745

Dioszegia antarctica

CBS 10920 #

DQ402529

FJ640575

KF037129

KF036858

# The vouchers are used in the phylogenetic analysis of Agaricomycetes in Figs 1, 2

* The vouchers are used in the phylogenetic analysis of genera within Sistotremastrumales and Trechisporales in Fig. 3

Newly generated sequences are in bold.

Of the five datasets, each gene region was separately aligned using MAFFT 7.110 (Katoh & Standley 2013) with the G-INS-i strategy (Katoh et

al. 2005) and then accordingly concatenated to five alignments (Supplementary files 1–5). The best-fit evolutionary models for the five alignments

were separately estimated using jModelTest 2.1.10 (Darriba et al. 2012, Guindon & Gascuel 2003). Following these models, maximum likelihood

(ML) and Bayesian inference (BI) algorithms were used to performed phylogenetic analyses. ML algorithm was conducted using raxmlGUI 2.0 (Edler

et al. 2021, Stamatakis 2014) and bootstrap (BS) replicates were determined under the auto FC option (Pattengale et al. 2010). BI algorithm was

conducted using MrBayes 3.2.7a (Ronquist et al. 2012). Two independent runs were employed, each with four chains and starting from random trees.

Trees were sampled every 1000th generation. The first 25% of sampled trees were discarded as burn-in, while the other 75% of trees were used to

construct a 50% majority consensus tree and for calculating Bayesian posterior probabilities (BPPs). Chain convergence was judged using Tracer 1.7

(Rambaut et al. 2018).

The alignment resulted from the dataset (1) was also subjected to a molecular clock analysis using BEAST 2.6.3 (Bouckaert et al. 2019).

Following Wang et al. (2021), the divergence time and the corresponding credibility intervals were estimated under the lognormal relaxed molecular

clock and the Yule speciation prior set. The following time points were set for calibration: 90 million years ago (Mya) as the minimum age of

Agaricales by Archaeomarasmius leggetti, a fossil agaricoid species preserved in a Dominican amber (Hibbett et al. 1997); 113 Mya as the minimum

age of Hymenochaetales by Quatsinoporites cranhamii, a fossil poroid species collected from Apple Bay on Vancouver Island (Smith et al. 2004); and

290 Mya as the mean age of Agaricomycetes by the analyses of genome data (Floudas et al. 2012). Accordingly, the offset ages for Agaricales and

Hymenochaetales were, respectively, set as 90 Mya and 113 Mya with a gamma distribution prior (alpha = 20, beta = 1), while the mean age for

Agaricomycetes was set as 290 Mya with a normal distribution prior (sigma = 1). After 200 million generations, the first 10% of the trees sampled

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every 1000th generation were removed as burn-in. Chain convergence of the resulting log file was

judged using Tracer 1.7 (Rambaut et al. 2018). A maximum-clade-credibility tree was summarized

using TreeAnnotator 2.6.3 incorporated into BEAST 2.6.3 (Bouckaert et al. 2019).

Results

Molecular phylogeny

In this study, 160 ITS, 132 nrLSU, 20 tef1-α, 13 rpb2 and 21 mtSSU sequences were newly

generated from 174 specimens (Table 1).

The combined dataset of ITS, nrLSU, tef1-α and rpb2 regions (1) comprised 65 samples and

resulted in an alignment of 3842 characters with GTR + I + G as the best-fit evolutionary model. In

ML algorithm, the BS search stopped after 200 replicates. In BI algorithm, after ten million

generations with an average standard deviation of split frequencies of 0.004707, all chains

converged, which was indicated by the effective sample sizes (ESSs) of all parameters above 3800

and the potential scale reduction factors (PSRFs) close to 1.000. Because BI and ML algorithms

generated nearly congruent topologies, the topology from ML algorithm is presented along with BS

values and BPPs greater than 50% and 0.8, respectively, at the nodes (Fig. 1). Regarding the

molecular clock analysis, chain convergence was indicated by the ESSs above 1200. The

maximum-clade-credibility chronogram is presented along with estimated divergence times of 95%

highest posterior density for all clades as node bars and the crown ages and BPPs above 0.8,

respectively, above and below the branches at the nodes (Fig. 2). While the monophyly of all

sampled orders was well supported, this multilocus-based phylogeny strongly supported the

independence of Sertulicium and Sistotremastrum (BS = 100%, BPP = 1) from other genera within

Trechisporales (BS = 100%, BPP = 1), although these two genera did have a closer relationship

with other genera within Trechisporales than with other orders (Fig. 1). Moreover, the 95% highest

posterior density ages for the clade of Sertulicium plus Sistotremastrum compared to the clade of

other genera within Trechisporales were, respectively, 111.09–167.24 Mya and 118.13–158.89

Mya with mean crown ages of 139.15 Mya and 137.95 Mya, respectively (Fig. 2). These

divergence times fell in the range of mean crown ages of other orders within Agaricomycetes from

38.72 Mya (Phallales) to 134.83 Mya (Russulales) except for Sebacinales having mean crown ages

of 183.61 Mya (Fig. 2). The option of combining Sertulicium, Sistotremastrum and other genera

within Trechisporales as one order produced a clade with a mean crown age of 175.94 Mya

considerably earlier than the option of two separate clades and most additional sampled orders

(Fig. 2). In association with distinct morphological characters, one new family within one new

order are described to accommodate Sertulicium and Sistotremastrum below.

The combined dataset of ITS, nrLSU, tef1-α, rpb2 and mtSSU regions (2) comprised 67

samples and resulted in an alignment of 4102 characters with GTR + I + G as the best-fit

evolutionary model. In ML algorithm, the BS search stopped after 150 replicates. In BI algorithm,

after ten million generations with an average standard deviation of split frequencies of 0.003781, all

chains converged, which was indicated by the ESSs above 3200 and the PSRFs close to 1.000.

Because BI and ML algorithms generated nearly congruent topologies, the topology from ML

algorithm is presented along with BS values and BPPs greater than 50% and 0.8, respectively, at

the nodes (Fig. 3). This phylogeny also supported, at least did not reject, the segregation of

Sertulicium and Sistotremastrum from Trechisporales as an independent lineage at the order level.

Each sampled genus within Trechisporales was well differentiated from others. Moreover, within

Trechisporales one new lineage at the genus level was revealed being composed of two recently

described species in Trechispora and a new lineage at the species level represented by two samples.

In association with morphological characters, one new genus within Hydnodontaceae,

Trechisporales and one new species and two new combinations accommodated in the new genus

are described below.

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Figure 1 – Phylogenetic position of Sistotremastrales and Trechisporales within the

Agaricomycetes inferred from the combined dataset of ITS, nrLSU, tef1-α and rpb2 regions. The

topology is generated by the maximum likelihood algorithm. Bootstrap values and Bayesian

881

posterior probabilities, when simultaneously above 50% and 0.8, respectively, are labelled at the

nodes.

Figure 2 – Maximum-clade-credibility chronogram and estimated divergence times of orders

within the Agaricomycetes inferred from the combined dataset of ITS, nrLSU, tef1-α and rpb2

regions. The estimated divergence times of 95% highest posterior density for all clades were

indicated as node bars, and for Sistotremastrales and Trechisporales were also provided below the

order names as exact numbers. The mean divergence times of clades (crown ages) and Bayesian

882

posterior probabilities above 0.8 were labeled above and below the branches, respectively, at the

nodes.

Figure 3 – Phylogenetic relationship among genera within Sistotremastrales and Trechisporales

inferred from the combined dataset of ITS, nrLSU, tef1-α, rpb2 and mtSSU regions. The topology

883

is generated by the maximum likelihood algorithm. Bootstrap values and Bayesian posterior

probabilities, when simultaneously above 50% and 0.8, respectively, are labelled at the nodes.

Figure 4 – Phylogenetic relationship among species of Fibrodontia inferred from the combined

dataset of ITS and nrLSU regions. The topology is generated by the maximum likelihood

algorithm. Bootstrap values and Bayesian posterior probabilities, when simultaneously above 50%

and 0.8, respectively, are labelled at the nodes. The newly described species are in boldface.

The combined dataset of ITS and nrLSU regions (3) comprised 32 samples and resulted in an

alignment of 1511 characters with GTR + I + G as the best-fit evolutionary model. In ML

algorithm, the BS search stopped after 350 replicates. In BI algorithm, after two million generations

with an average standard deviation of split frequencies of 0. 006526, all chains converged, which

was indicated by the ESSs above 1100 and the PSRFs close to 1.000. Because BI and ML

algorithms generated nearly congruent topologies, the topology from ML algorithm is presented

along with BS values and BPPs greater than 50% and 0.8, respectively, at the nodes (Fig. 4). The

monophyly of Fibrodontia was strongly supported and one new lineage represented by two samples

emerged within Fibrodontia. In association of morphological characters, this new lineage is

described as one new species of Fibrodontia below.

The combined dataset of ITS and nrLSU regions (4) comprised 29 samples and resulted in an

alignment of 1521 characters with GTR + I + G as the best-fit evolutionary model. In ML

884

algorithm, the BS search stopped after 350 replicates. In BI algorithm, after one million generations

with an average standard deviation of split frequencies of 0. 006978, all chains converged, which

was indicated by the ESSs above 700 and the PSRFs close to 1.000. Because BI and ML algorithms

generated nearly congruent topologies, the topology from ML algorithm is presented along with BS

values and BPPs greater than 50% and 0.8, respectively, at the nodes (Fig. 5). The monophyly of

Subulicystidium was strongly supported, and two new lineages, each represented by two samples,

emerged within Subulicystidium. In association of morphological characters and conditions of

specimens, one of these two new lineages is described as one new species of Subulicystidium

below.

Figure 5 – Phylogenetic relationship among species of Subulicystidium inferred from the combined

dataset of ITS and nrLSU regions. The topology is generated by the maximum likelihood

algorithm. Bootstrap values and Bayesian posterior probabilities, when simultaneously above 50%

and 0.8, respectively, are labelled at the nodes. The newly described species are in boldface.

The combined dataset of ITS and nrLSU regions (5) comprised 249 samples and resulted in

an alignment of 1796 characters with GTR + I + G as the best-fit evolutionary model. In ML

algorithm, the BS search stopped after 250 replicates. In BI algorithm, after ten million generations

with an average standard deviation of split frequencies of 0.007705, all chains converged, which

885

was indicated by the ESSs above 2800 and the PSRFs close to 1.000. Because BI and ML

algorithms generated nearly congruent topologies, the topology from ML algorithm is presented

along with BS values and BPPs greater than 50% and 0.8, respectively, at the nodes (Fig. 6). The

monophyly of Trechispora was strongly supported and 16 new lineages, each represented by two or

more samples, emerged within Trechispora. In association of morphological characters, these 16

new lineages are described as 16 new species of Trechispora below. In addition, five species of

Scytinopogon with clear taxonomic background, viz. S. caulocystidiatus, S. dealbatus, S. foetidus,

S. robustus and S. scaber nested within Trechispora (Fig. 6) and thus are transferred to Trechispora

below.

Figure 6 – Phylogenetic relationship among species of Trechispora inferred from the combined

dataset of ITS and nrLSU regions. The topology is generated by the maximum likelihood

886

algorithm. Bootstrap values and Bayesian posterior probabilities, when simultaneously above 50%

and 0.8, respectively, are labelled at the nodes. The newly described species are in boldface.

Figure 6 – Continued.

887

Figure 6 – Continued.

Besides these phylogenetic analyses, molecular sequences were also employed to explore the

taxonomic position of Litschauerella, Sphaerobasidium and Trechispora yunnanense via BLAST

search. Their taxonomic positions at the higher rank are changed below. In addition, a brief

summary of each genus accepted in Trechisporales is provided.

Taxonomy

Sistotremastrales L.W. Zhou & S.L. Liu, ord. nov.

Index Fungorum number: IF559875; Facesoffungi number: FoF12855

Etymology – Sistotremastrales (Latin), refers to the type family Sistotremastraceae.

Type family – Sistotremastraceae L.W. Zhou & S.L. Liu (described below).

Type genus – Sistotremastrum J. Erikss., Symb. bot. upsal. 16(no. 1): 62 (1958).

888

Type species – Sistotremastrum suecicum Litsch. ex J. Erikss., Symb. bot. upsal. 16(no. 1):

62 (1958).

Type specimen – SWEDEN, Upland, Uppsala, Bondkyrka parish, Malma skog, Northwest of

Hälltorpet, on the underside of coniferous fencing material lying in a wet spot, 4 Jan. 1933, Seth

Lundell, F204406 (S, holotype).

Description – Basidiomes resupinate, effused, thin. Hymenophore smooth, grandinioid or

odontioid. Hyphal system monomitic, all septa with clamp connections. Cystidia mostly absent.

Basidia cylindrical to tubular with four to eight sterigmata. Basidiospores smooth, ellipsoid to

cylindrical, inamyloid, acyanophilous. On wood.

Notes – Sertulicium and Sistotremastrum formerly belonging to Trechisporales are

segregated as the new order. Sistotremastrales is characterized by corticioid basidiomes on wood,

basidia with four to eight sterigmata and smooth basidiospores. Comparing with the

circumscription of Trechisporales sensu He et al. (2019) and Spirin et al. (2021), some species in

the reduced concept of Trechisporales after the exclusion of Boidinella (below), Litschauerella

(below), Sertulicium, Sistotremastrum and Sphaerobasidium (below) also have smooth

basidiospores, and thus are similar to species bearing basidia with four sterigmata in

Sistotremastrales; however, species of Trechisporales differ in soft basidiomes, subicular hyphae

with ampullate septa and presence of cystidia with various shapes (Spirin et al. 2021).

Sistotremastraceae L.W. Zhou & S.L. Liu, fam. nov.

Index Fungorum number: IF559876; Facesoffungi number: FoF12856

Etymology – Sistotremastraceae (Latin), refers to the type genus Sistotremastrum.

Type genus – Sistotremastrum J. Erikss., Symb. bot. upsal. 16(no. 1): 62 (1958).

Description – Basidiomes resupinate, effused, thin, usually up to 100 μm thick. Hymenophore

smooth, grandinioid or odontioid, white to pale ochraceous. Hyphal system monomitic, all septa

with clamp connections. Cystidia mostly absent, hyphidia rarely present. Basidia cylindrical to

tubular, often with a slight median constriction, with four to eight sterigmata. Basidiospores

smooth, ellipsoid to cylindrical, inamyloid, acyanophilous. On wood.

Notes – Sistotremastraceae is the single family within Sistotremastrales and comprises

Sertulicium and Sistotremastrum, segregated from Trechisporales. Therefore, the unique

morphological characters of Sistotremastraceae are the same as those indicated to Sistotremastrales

above. From the morphological perspective, Larsson (2007) suggested the distinction of

Sistotremastrum from other genera in Trechisporales and used the designation ‘Sistotremastrum

family’. Spirin et al. (2021) segregated Sertulicium from Sistotremastrum and indicated these two

genera are unique in Trechisporales. However, Spirin et al. (2021) did not propose any formal

taxonomic change at the family or higher rank mainly due to lack of reliable phylogenetic support.

Here, according to multilocus-based phylogenetic analyses, we formally propose Sertulicium and

Sistotremastrum in one new family and one new order independent from Hydnodontaceae and

Trechisporales, respectively.

Sertulicium Spirin, Volobuev & K.H. Larss., in Spirin, Volobuev, Viner, Miettinen, Vlasák,

Schoutteten, Motato-Vásquez, Kotiranta & Larsson, Mycol. Progr. 20(4): 460 (2021).

Type species – Sertulicium niveocremeum (Höhn. & Litsch.) Spirin & K.H. Larss., in Spirin,

Volobuev, Viner, Miettinen, Vlasák, Schoutteten, Motato-Vásquez, Kotiranta & Larsson, Mycol.

Progr. 20(4): 466 (2021)

Description – Basidiomes annual, resupinate, effused, very thin (usually up to 0.1 mm thick).

Hymenophore smooth, white, cream to pale ochraceous. Hyphal system monomitic, generative

hyphae with clamp connections, hyaline. Cystidia mostly absent. Hyphidia rarely present. Basidia

clavate, hyaline, thin-walled, with four to six or six to eight sterigmata and a basal clamp

connection. Basidiospores narrowly ellipsoid to cylindrical, hyaline, thin-walled, smooth,

inamyloid, indextrinoid, acyanophilous. On wood.

889

Notes – Sertulicium was recently segregated from Sistotremastrum as a new genus with S.

niveocremeum as the generic type (Spirin et al. 2021). Six combined species from other genera and

a newly described species from Finland bring the species number of Sertulicium to seven (Spirin et

al. 2021, Liu et al. 2022). While their morphological characters are not distinct, phylogenetically

Sertulicium and Sistotremastrum are close but clearly separated from each other (Spirin et al. 2021,

Figs 1–3).

Sistotremastrum J. Erikss., Symb. bot. upsal. 16(no. 1): 62 (1958).

Type species – Sistotremastrum suecicum Litsch. ex J. Erikss., Symb. bot. upsal. 16(no. 1):

62 (1958).

Type specimen – SWEDEN, Upland, Uppsala, Bondkyrka parish, Malma skog, Northwest of

Hälltorpet, on the underside of coniferous fencing material lying in a wet spot, 4 Jan. 1933, Seth

Lundell, F204406 (S, holotype).

Description – Basidiomes annual, resupinate, effused, thin to rather substantial (up to 0.2 mm

thick). Hymenophore smooth, grandinioid, odontioid, white, cream to buff-yellow. Hyphal system

monomitic, generative hyphae with clamp connections, hyaline. Cystidia mostly absent. Hyphidia

present. Basidia clavate, hyaline, thin-walled, with two to four, four to six or six to eight sterigmata

and a basal clamp connection. Basidiospores narrowly ellipsoid to cylindrical, hyaline, thin-walled,

smooth, inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Sistotremastrum was erected for S. niveocremeum and S. suecicum with the latter

species as the generic type (Eriksson 1958). Later, six species, viz. S. aculeocrepitans, S. chilense,

S. fibrillosum, S. guttuliferum, S. lateclavigerum and S. roseum were successively added to this

genus (Boidin & Gilles 1994, Telleria et al. 2013b, 2014, Dhingra et al. 2014, Gruhn et al. 2018).

However, of these eight species, S. chilense, S. lateclavigerum and S. niveocremeum were recently

transferred to Sertulicium, and S. guttuliferum was considered to be a synonym of Sertulicium

granuliferum by Spirin et al. (2021). Besides, Spirin et al. (2021) also newly described eight species

in Sistotremastrum. In addition, Sistotremastrum limonadense was newly described from French

Guiana (Gruhn & Alvarado 2021), but this species was later transferred to Sertulicium (Liu et al.

2022). In summary, a total of 12 species is accepted in Sistotremastrum. Although being placed in

Trechisporales, Sistotremastrum has long been considered to be separated from additional genera in

the single family Hydnodontaceae within this order (Larsson 2007). Unlike the previous phylogeny

lack of reliable support at the family rank (Spirin et al. 2021), the current multilocus-based

phylogenetic analyses support the proposal of Sistotremastrum in one new family

Sistotremastraceae within one new order Sistotremastrales independent from Trechisporales (Figs

1–3).

Trechisporales K.H. Larss., in Hibbett et al., Mycol. Res. 111(5): 541 (2007).

Type genus – Trechispora P. Karst., Hedwigia 29: 147 (1890) = Hydnodon Banker,

Mycologia 5(6): 297 (1913).

Type species – Trechispora onusta P. Karst., Hedwigia 29: 147 (1890) = T. hymenocystis

(Berk. & Broome) K.H. Larss., Mycol. Res. 98(10): 1167 (1994).

Description – Basidiomes resupinate, effused, stipitate or clavarioid. Hymenophore smooth,

grandinioid, hydnoid or poroid to partly irpicoid. Hyphal system monomitic to dimitic, all septa

with clamp connections. Cystidia present or absent. Basidia cylindrical with two to four sterigmata.

Basidiospores smooth or ornamented, subglobose, ellipsoid, subangular or fusiform, inamyloid,

acyanophilous. On wood or ground.

Notes – Comparing with the circumscription of Trechisporales sensu He et al. (2019) and

Spirin et al. (2021), the current concept of Trechisporales is reduced according to the exclusion of

Boidinella (below), Litschauerella (below), Sertulicium, Sistotremastrum and Sphaerobasidium

(below). For now, Trechisporales is composed of one family Hydnodontaceae comprising 12

genera.

890

Hydnodontaceae Jülich, Biblthca Mycol. 85: 372 (1982) [1981].

Type genus – Hydnodon Banker, Mycologia 5(6): 297 (1913) = Trechispora P. Karst.,

Hedwigia 29: 147 (1890).

Type species – Hydnodon thelephorus (Lév.) Banker [as 'thelephorum'], Mycologia 5(6): 297

(1913) = Trechispora thelephora (Lév.) Ryvarden, Syn. Fung. (Oslo) 15: 32 (2002).

= Subulicystidiaceae Jülich, Biblthca Mycol. 85: 391 (1982) [1981].

Description – Basidiomes resupinate, effused, stipitate or clavarioid. Hymenophore smooth,

grandinioid, odontioid, hydnoid or poroid to partly irpicoid, mycelial cords present or absent.

Hyphal system monomitic to dimitic, all septa with clamp connections, subicular hyphae with or

without ampullate septa. Cystidia present in some species, mostly absent. Basidia cylindrical with

two to four sterigmata, often with a slight median constriction. Basidiospores smooth or

ornamented, subglobose, ellipsoid, subangular or fusiform, inamyloid, acyanophilous.

Conidiospores sometimes present. On wood or ground.

Notes – After the exclusion of Boidinella (below), Litschauerella (below), Sertulicium,

Sistotremastrum and Sphaerobasidium (below), all genera formerly accepted in Trechisporales and

a new genus Allotrechispora (described below) are accommodated in Hydnodontaceae.

The family names Hydnodontaceae and Subulicystidiaceae (as well as Litschauerellaceae,

see Litschauerella under excluded genera from Trechisporales) were published simultaneously by

Jülich (1981). Under this circumstance, the choice of name is governed by Art. 11.5 of the

Shenzhen Code (Turland et al. 2018) and the first effectively published choice establishes priority.

Hibbett et al. (2014) listed Subulicystidiaceae as a synonym of Hydnodontaceae, thereby

establishing the priority of the latter name.

Allotrechispora L.W. Zhou & S.L. Liu, gen. nov.

Index Fungorum number: IF559877; Facesoffungi number: FoF12857

Etymology – Allotrechispora (Latin), refers to the segregation from Trechispora.

Diagnosis – Differs from Trechispora in the absence of stipitate or clavarioid basidiomes, and

the absence of ampullate septa on subicular hyphae (Eriksson et al. 1981, de Meiras-Ottoni et al.

2021).

Type species – Allotrechispora gatesiae L.W. Zhou, S.L. Liu & T.W. May (described below).

Type specimen – AUSTRALIA, Tasmania, Tahune Adventures, The Look-in Look-out, on

fallen Atherosperma moschatum, 15 May 2018, L.W. Zhou, LWZ 20180515-18 (holotype in MEL,

isotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft, loosely attached to the

substrates. Hymenophore smooth to tuberculate, white to cinnamon-buff, cracked with age. Hyphal

system monomitic, generative hyphae with clamp connections, hyaline, thin-walled. Cystidia

absent. Basidia subcylindrical to subclavate, hyaline, thin-walled, with four sterigmata and a basal

clamp connection. Basidiospores oblong ellipsoid to ellipsoid, hyaline, thin-walled, smooth,

inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Compared to other genera in Trechisporales, Allotrechispora is characterized by

cream to cinnamon-buff, corticioid basidiomes, a monomitic hyphal system, absence of

isodiametric subhymenial hyphae, absence of cystidia, and smooth, oblong ellipsoid to ellipsoid

basidiospores. Allotrechispora is phylogenetically closest to Fibrodontia (the clade containing both

genera has BS = 74%, BPP = 0.99; Fig. 3), and these two genera are more or less similar by the

absence of cystidia and smooth, ellipsoid basidiospores. However, Fibrodontia differs in the

odontioid hymenophore and a dimitic hyphal system.

Allotrechispora daweishanensis (C.L. Zhao) L.W. Zhou & S.L. Liu, comb. nov.

Index Fungorum number: IF559878; Facesoffungi number: FoF12858

Basionym. Trechispora daweishanensis C.L. Zhao, in Zong, Liu, Wu & Zhao, Phytotaxa

479(2): 153 (2021).

891

≡ Brevicellicium daweishanense (C.L. Zhao) Z.B. Liu & Yuan Yuan, Frontiers in

Microbiology 13(no. 818358): 15 (2022).

Notes – Trechispora daweishanensis was recently described from southwestern China (Zong

et al. 2021). Although the authors stated that their phylogeny indicates this species nested within

Trechispora, this genus was not recovered as a well-supported clade (Fig. 1 in Zong et al. 2021). A

later phylogenetic analysis revealed the separation of T. daweishanensis from Trechispora, but

unfortunately this phylogeny sampling taxa incomprehensively was incorrectly recognized and T.

daweishanensis was accordingly transferred to Brevicellicium (Liu et al. 2022). According to the

current phylogeny (Fig. 3), T. daweishanensis does not belong to Trechispora and Brevicellicium,

but falls within a distinct clade described as a new genus Allotrechispora in Trechisporales.

Therefore, T. daweishanensis is transferred to Allotrechispora.

Allotrechispora gatesiae L.W. Zhou, S.L. Liu & T.W. May, sp. nov. Figs 7, 8

Index Fungorum number: IF559879; Facesoffungi number: FoF12859

Etymology – gatesiae (Latin), refers to the Australian mycologist, Dr. Genevieve Gates, who

kindly arranged the author Li-Wei Zhou’s field trip in Tasmania, Australia.

Diagnosis – Characterized by the largest basidiospores in the genus.

Typus – AUSTRALIA, Tasmania, Tahune Adventures, The Look-in Look-out, on fallen

trunk of Atherosperma moschatum, 15 May 2018, L.W. Zhou, LWZ 20180515-18 (holotype in

MEL, isotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft, loosely attached to the

substrates. Hymenophore tuberculate, straw-yellow when fresh, straw-yellow to cinnamon-buff,

cracked with age. Margin thinning out, fimbriate, slightly paler than hymenophore surface,

becoming indistinct with age.

Hyphal system monomitic; generative hyphae with clamp connections, hyaline, thin-walled,

frequently branched and septate, loosely interwoven, 2–3 µm in diam. Cystidia absent. Basidia

cylindrical, hyaline, thin-walled, with four sterigmata and a basal clamp connection, 20–27 × 6–8

µm; basidioles in shape similar to basidia, but slightly smaller. Basidiospores oblong ellipsoid to

ellipsoid, hyaline, thin-walled, smooth, inamyloid, indextrinoid, acyanophilous, (5–)5.2–6.8(–7) ×

(3.7–)3.8–4.3(–5) µm, L = 6.0 µm, W = 4.1 µm, Q = 1.4–1.5 (n = 60/2).

Other specimen (paratype) examined – AUSTRALIA, Tasmania, Tahune Adventures, The

Look-in Look-out, on fallen trunk of Atherosperma moschatum, 15 May 2018, L.W. Zhou, LWZ

20180515-20 (HMAS).

Notes – As far as macromorphology, Allotrechispora gatesiae is distinct from other species

in this genus by its colored basidiomes. The basidiospores in A. gatesiae are larger than those in A.

daweishanensis (3.8–5 × 2.7–3.5 µm; Zong et al. 2021: as Trechispora). In comparison to

Allotrechispora gatesiae, basidiospores are overlapping in size in A. xantha (4.3–5.7 × 3.2–4 µm;

transferred from Trechispora xantha below), but this species has a lower length to width ratio of

basidiospores (1.32–1.41; Zong et al. 2021: as Trechispora).

Allotrechispora xantha (C.L. Zhao) L.W. Zhou & S.L. Liu, comb. nov.

Index Fungorum number: IF559880; Facesoffungi number: FoF12860

Basionym. Trechispora xantha C.L. Zhao, in Zong, Liu, Wu & Zhao, Phytotaxa 479(2): 155

(2021).

≡ Brevicellicium xanthum (C.L. Zhao) Z.B. Liu & Yuan Yuan, Frontiers in Microbiology

13(no. 818358): 15 (2022).

Notes – Trechispora xantha was described from southwestern China together with

T. daweishanensis, and these two species have a close phylogenetic relationship (Zong et al. 2021).

Together with T. daweishanensis, T. xantha was inappropriately transferred to Brevicellicium (Liu

et al. 2022). The current phylogeny supports that T. xantha and T. daweishanensis fall within the

clade of the new genus Allotrechispora (Fig. 3). Therefore, T. xantha is transferred to

Allotrechispora.

892

Figure 7 – Basidiomes of Allotrechispora gatesiae (LWZ 20180515-18, holotype). Scale bars: a =

1 cm, b = 2 mm.

Brevicellicium K.H. Larss. & Hjortstam, in Hjortstam & Larsson, Mycotaxon 7(1): 117 (1978).

Type species – Brevicellicium exile (H.S. Jacks.) K.H. Larss. & Hjortstam, Mycotaxon 7(1):

118 (1978).

Description – Basidiomes annual, resupinate, effused, thin, membranaceous. Hymenophore

smooth, grandinioid, white, cream to yellowish. Hyphal system monomitic, generative hyphae with

clamp connections, hyaline, thin-walled, distinctly isodiametric. Cystidia absent. Basidia shortly

cylindrical, hyaline, thin-walled, with four sterigmata and a basal clamp connection. Basidiospores

usually subangular with a distinct apiculus, hyaline, thin-walled, smooth, inamyloid, indextrinoid,

acyanophilous. On wood.

Notes – Brevicellicium was erected by Hjortstam & Larsson (1978) with B. exile as the

generic type. Brevicellicium is a cosmopolitan genus and is characterized by isodiametric

subhymenial hyphae, short basidia, and smooth, often subangular basidiospores with a distinct

893

apiculus. Jülich (1981) put Brevicellicium in Hydnodontaceae, and Larsson (2007) accepted this

placement at the family level and further put it in Trechisporales. Later, Telleria et al. (2013a)

confirmed the classification of Brevicellicium within Hydnodontaceae, Trechisporales from the

phylogenetic perspective. For now, ten species are accepted in Brevicellicium, while the taxonomic

position of certain species needs to be further tested by molecular evidence (Telleria et al. 2013a).

Moreover, two unnamed single-specimen lineages, viz. LWZ 20190809-10b and LWZ 20190918-

13, both from Sichuan, China are revealed from the current multilocus-based phylogenetic analyses

(Figs 1–3).

Figure 8 – Microscopic structures of Allotrechispora gatesiae (drawn from the holotype).

a Vertical section of basidiomes. b Basidia. c Basidiospores. Scale bars = 10 μm.

Brevicellopsis Hjortstam & Ryvarden, Syn. Fung. (Oslo) 25: 15 (2008).

Type species – Brevicellopsis allantospora (Hjortstam & Ryvarden) Hjortstam & Ryvarden,

Syn. Fung. (Oslo) 25: 15 (2008).

Description – Basidiomes annual, resupinate, effused, thin, membranaceous, soft, fragile.

Hymenophore odontioid, greyish white to pale ochraceous. Hyphal system monomitic, generative

hyphae with clamp connections, hyaline, slightly isodiametric. Cystidia absent. Basidia shortly

cylindrical, hyaline, thin-walled, with four sterigmata and a basal clamp connection. Basidiospores

allantoid, hyaline, thin-walled, smooth, inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Brevicellopsis was erected to accommodate B. allantospora (Hjortstam & Ryvarden

2008) that is a species originally described from Africa as a member of Brevicellicium (Hjortstam

& Ryvarden 1980). This monotypic genus differs from Brevicellicium mainly by odontioid

hymenophore and allantoid basidiospores (Hjortstam and Ryvarden 2008). Presumably due to the

segregation from Brevicellicium, Brevicellopsis was also considered to potentially belong to

Trechisporales (Hibbett et al. 2014). Phylogenetic analyses need to be employed to exactly delimit

the taxonomic position of Brevicellopsis.

894

Dextrinocystis Gilb. & M. Blackw., Mycotaxon 33: 376 (1988).

Type species – Dextrinocystis capitata (D.P. Rogers & Boquiren) Gilb. & M. Blackw.,

Mycotaxon 33: 378 (1988).

Description – Basidiomes annual, resupinate, effused, thin, soft, easily separated from the

substrate. Hymenophore smooth, cream to buff, not cracked. Hyphal system monomitic, generative

hyphae with clamp connections, hyaline. Cystidia-like branches present on subicular hyphae,

embedded, hyaline, thick-walled, encrusted at apex. Cystidia subulate, projecting beyond

hymenium, bi- or multi-rooted, hyaline, distinctly thick-walled with a narrow lumen, slightly

encrusted at apex, distinctly dextrinoid. Basidia subcylindrical to subclavate, hyaline, thin-walled,

with four sterigmata and a basal clamp connection. Basidiospores oblong-ellipsoid to short

cylindrical, hyaline, thin-walled, smooth, inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Dextrinocystis was segregated from Epithele to accommodate E. capitata that is

strikingly distinguished by strongly dextrinoid cystidia (Gilbertson & Blackwell 1988). This

monotypic genus was doubtfully put in Trechisporales (Larsson 2007). Later, another two species

also with dextrinoid cystidia, viz. D. calamicola and D. macrospora were added to this genus

(Nakasone 2013, Liu et al. 2019). Of the three species accepted in Dextrinocystis, D. calamicola

was included in a phylogenetic analysis that unambiguously confirmed that within Trechisporales

this genus has a closer relationship with Tubulicium, as indicated by morphological characters (Liu

et al. 2019). This topology is also recovered by the current phylogeny (Fig. 3).

Fibrodontia Parmasto, Consp. System. Corticiac. (Tartu): 174 (1968).

Type species – Fibrodontia gossypina Parmasto, Consp. System. Corticiac. (Tartu): 207

(1968).

Description – Basidiomes annual, resupinate, effused, thin, soft-membranaceous, easily

detached. Hymenophore grandinioid, odontioid, white to pale cinnamon-buff. Hyphal system

dimitic, generative hyphae with clamp connections, hyaline; skeletal hyphae hyaline to yellowish,

thick-walled with a wide lumen. Cystidia absent. Basidia subcylindrical to subclavate, hyaline,

thin-walled, with four sterigmata and a basal clamp connection. Basidiospores broadly ellipsoid to

ovoid, hyaline, thin-walled, smooth, inamyloid, indextrinoid, acyanophilous or slightly

cyanophilous. On wood.

Notes – Fibrodontia gossypina was newly described as the generic type of a new genus

Fibrodontia (Parmasto 1968). Although Fibrodontia is morphologically close to Hyphodontia

(Eriksson et al. 1981) and was formerly treated as a synonym of Hyphodontia (Langer 1994), the

molecular evidence indicated that this genus belongs to Hydnodontaceae (Binder et al. 2005) and

thus the genus was placed in Trechisporales (Larsson 2007). Six species were accepted in

Fibrodontia (Liu et al. 2021), while one additional species is described below, bringing the total

species number in this genus to seven. It is noteworthy that Baltazar et al. (2016) proposed

Cystidiodendron fimbriatum (the generic type of Cystidiodendron) as conspecific with

F. gossypina. To solve the taxonomic delimitation of these two genera, the species affinity of

C. fimbriatum and F. gossypina has to be tested via molecular evidence (Liu et al. 2021). Ideally,

the lectotype of C. fimbriatum collected in 1933 or potentially a later collection designated as an

epitype can be sequenced.

Fibrodontia subaustrosinensis S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 9, 10

Index Fungorum number: IF559881; Facesoffungi number: FoF12861

Etymology – subaustrosinensis (Latin), refers to the new species resembling Fibrodontia

austrosinensis in morphology.

Diagnosis – Differs from Fibrodontia austrosinensis by longer basidia and slightly larger

basidiospores (Liu et al. 2021).

Typus – CHINA, Yunnan, Xichou County, Xiaoqiaogou Forest Park, on fallen angiosperm

trunk, 16 Nov. 2019, S.H. He, He 6279 (holotype in BJFC 033223, isotype in HMAS).

895

Description – Basidiomes annual, resupinate, easily detached, without odor or taste, soft

corky and brittle when dry, up to 13 cm long, 3 cm wide and 0.2 mm thick. Hymenophore

grandinioid, white to cream when fresh, usually with curry-yellow to olive tinge, olivaceous buff to

honey-yellow when dry. Margin white, cottony, up to 0.5 mm wide.

Hyphal system dimitic; generative hyphae with clamp connections. Subiculum composed of a

loose layer of distinct hyphae; generative hyphae, hyaline, thin- to slightly thick-walled,

occasionally branched, smooth, 2–3.5 μm in diam; skeletal hyphae rare, hyaline to yellowish, thick-

walled with a wide to narrow lumen, unbranched, smooth, slightly flexuous, 2–3 μm in diam.

Aculei composed of a central core of compact hyphae and subhymenial and hymenial layers, at

apex terminal hyphae slightly tapered; generative hyphae distinct, hyaline, thin- to slightly thick-

walled, occasionally branched, smooth, subparallel, 2–3 μm in diam; skeletal hyphae rare, hyaline

to yellowish, thick-walled with a wide to narrow lumen, unbranched, smooth, slightly flexuous,

more or less parallel along the aculei, 2–3 μm in diam. Basidia suburniform to clavate, thin-walled,

with four sterigmata and a basal clamp connection, 20–27 × 3.8–5 μm; basidioles similar in shape

to basidia, but smaller. Basidiospores ellipsoid to ovoid, hyaline, thin-walled, smooth, inamyloid,

indextrinoid, acyanophilous, (4.5–)4.8–6 × (3.9–)4–5(–5.5) μm, L = 5.2 μm, W = 4.4 μm, Q = 1.2

(n = 60/2).

Other specimen (paratype) examined – CHINA, Hainan, Changjiang County, Bawangling

National Forest Park, on dead branch of living tree of Arenga pinnata, 16 Nov. 2019, S.H. He, He

6033 (BJFC).

Notes – Fibrodontia subaustrosinensis closely resembles F. austrosinensis, but the latter

species has shorter basidia (13–16 µm in length) and slightly smaller basidiospores (4.2–5.2 × 3.5–

4.5 µm; Liu et al. 2021). Fibrodontia brevidens is also similar to F. subaustrosinensis by the

yellowish hymenophore with an olive tinge, but differs in the presence of moderately encrusted

skeletal hyphae and smaller basidiospores (4–4.5 × 3.5–4.5 μm; Yurchenko & Wu 2014).

Luellia K.H. Larss. & Hjortstam, Svensk bot. Tidskr. 68(1): 59 (1974).

Type species – Luellia recondita (H.S. Jacks.) K.H. Larss. & Hjortstam, Svensk bot. Tidskr.

68(1): 60 (1974).

Description – Basidiomes annual, resupinate, effused, thin, soft, closely attached to the

substrates. Hymenophore smooth, brown. Hyphal system monomitic, generative hyphae clamped or

not, yellowish brown. Cystidia absent. Basidia clavate to pyriform, hyaline, thin-walled, with two

to four sterigmata. Basidiospores fusiform to navicular, with distinct apiculus, thin-walled, smooth,

inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Luellia was erected by Larsson & Hjortstam (1974), who at that time accommodated

two species in the genus, viz. L. furcata and L. recondita. Later, Luellia cystidiata was added to this

genus (Hauerslev 1979). Morphologically, Luellia is characterized by brown hymenophore with

mostly encrusted hyphae and thin-walled, inamyloid basidiospores, and was placed in Atheliaceae,

Atheliales (Jlich 1981). However, even though morphological characters did not suggest affinity

to other genera in Trechisporales, Larsson (2007) considered that Luellia is a member of

Hydnodontaceae, Trechisporales on the basis of unpublished molecular evidence, which is also

confirmed by Spirin et al. (2021) and the current phylogeny (Fig. 3).

Porpomyces Jülich, Persoonia 11(4): 425 (1982).

Type species – Porpomyces mucidus (Pers.) Jülich, Persoonia 11(4): 425 (1982).

Description – Basidiomes annual, resupinate, soft. Hymenophore poroid, white to pale

ochraceous, pores round to angular. Hyphal system monomitic, generative hyphae with clamp

connections, hyaline, with ampullate septa in hyphal cords. Cystidia absent. Basidia barrel-shaped,

hyaline, thin-walled, with four sterigmata and a basal clamp connection. Basidiospores subglobose

to ellipsoid, hyaline, thin-walled, smooth, inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Porpomyces was erected as a monotypic genus typified by P. mucidus (Jlich 1982).

This species was formerly considered to be close to Ceriporiopsis and thus placed in Polyporales

896

(Gilbertson & Ryvarden 1985), but later molecular evidence indicated that it is close to Trechispora

instead of Ceriporiopsis (Larsson 2001). Therefore, Porpomyces was listed as one of the exemplar

genera of Trechisporales when this order was newly proposed (Hibbett et al. 2007). Three species

are accepted in Porpomyces and its poroid hymenophore and ampullate hyphal septa make it

morphologically similar to Trechispora (Spirin et al. 2021). However, Porpomyces bears smooth

basidiospores, while all known poroid species in Trechispora have ornamented basidiospores.

Figure 9 – Basidiomes of Fibrodontia subaustrosinensis (He 6279, holotype). Scale bars: a = 1 cm,

b = 1 mm.

Pteridomyces Jülich, Persoonia 10(3): 331 (1979).

Type species – Pteridomyces galzinii (Bres.) Jülich, Persoonia 10(3): 331 (1979).

Description – Basidiomes annual, resupinate, effused, thin. Hymenophore grandinioid,

odontioid. Hyphal system monomitic to dimitic, generative hyphae with clamp connections,

hyaline. Hyphal pegs consisting of parallelly arranged, thin-walled, hyaline hyphae. Cystidia

897

present or absent, fusoid. Basidia narrowly clavate, hyaline, thin-walled, with two to four

sterigmata and a basal clamp connection. Basidiospores cylindrical, allantoid or navicular, thin-

walled, smooth, inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Pteridomyces was erected as a monotypic genus to accommodate Epithele galzinii

(Jülich 1979). Later, Boidin and his colleagues broadened the morphological delimitation of

Pteridomyces from a monomitic to dimitic hyphal system by adding seven species to this genus

(Boidin & Lanquetin 1983, Boidin & Gilles 1986a, 1988, Boidin et al. 1989). Hjortstam (1991)

redelimited the taxonomic status of the eight species of Pteridomyces, and treated Pteridomyces as

a synonym of Athelopsis, a disposition which is not widely recognized. However, due to the

morphological affinity to Athelopsis, Pteridomyces was placed in Atheliaceae, Atheliales, when

Larsson (2007) dealt with the taxonomic position of corticioid fungi. Subsequently, a new species

of Pteridomyces was described from Chilean Patagonia based solely on morphological characters

(Gorjón & Hallenberg 2013). Recently, Spirin et al. (2021) and Sulistyo et al. (2021) with the aid of

phylogenetic analyses suggested that Pteridomyces is an independent genus in Hydnodontaceae,

Trechisporales. The current phylogenies also recovered this topology (Figs 1–3).

Figure 10 – Microscopic structures of Fibrodontia subaustrosinensis (drawn from the holotype).

a Vertical section of basidiomes. b Basidia. c Basidiospores. Scale bars = 10 μm.

Subulicystidium Parmasto, Consp. System. Corticiac. (Tartu): 120 (1968).

Type species – Subulicystidium longisporum (Pat.) Parmasto, Consp. System. Corticiac.

(Tartu): 121 (1968).

= Aegeritina Jülich, Int. J. Mycol. Lichenol. 1(3): 282 (1984). Type species – Aegeritina

tortuosa (Bourdot & Galzin) Jülich, Int. J. Mycol. Lichenol. 1(3): 282 (1984).

Description – Basidiomes annual, resupinate, effused, thin, soft. Hymenophore smooth, more

or less arachnoid, white, cream to olivaceous buff. Margin not differentiated. Hyphal system

monomitic, generative hyphae with clamp connections, hyaline, thin-walled. Cystidia subulate,

projecting beyond hymenium, hyaline, thick-walled and regularly covered with rectangular crystals

except at the apex. Basidia suburniform, hyaline, thin-walled, with four sterigmata and a basal

898

clamp connection. Basidiospores cylindrical, fusiform to sigmoid, hyaline, thin-walled, smooth,

inamyloid, indextrinoid, acyanophilous. On wood.

Notes – Subulicystidium was erected as a monotypic genus with S. longisporum as the generic

type (Parmasto 1968). Molecular evidence unambiguously supported this genus as a member of

Hydnodontaceae, Trechisporales, although there was a lack of clear morphological affinity

(Larsson 2007). Recently, 13 species were newly described in Subulicystidium on the basis of

morphological and phylogenetic evidence (Ordynets et al. 2018, Liu et al. 2019). One more species

of Subulicystidium is described below, which bring the species number of this genus to 24.

Subulicystidium is morphologically distinct from other genera in Trechisporales in the presence of

a crystalline sheath on the cystidia.

The monotypic genus Aegeritina comprises A. tortuosa, accepted as the asexual stage of

Subulicystidium longisporum (Eriksson et al. 1984). With the transition to one fungus-one name,

Stalpers et al. (2021) recommended retention of the earlier generic name Subulicystidium over

Aegeritina.

Subulicystidium daii S.L. Liu & L.W. Zhou, sp. nov. Figs 11, 12

Index Fungorum number: IF559882; Facesoffungi number: FoF12862

Etymology – daii (Latin), refers to the Chinese mycologist, Prof. Dr. Yu-Cheng Dai, who

opened the door of fungal taxonomy for the author Li-Wei Zhou.

Diagnosis – Differs from Subulicystidium acerosum by the absence of needle-like crystals

and wider basidiospores (1.8–2.2 µm in width in S. acerosum; Liu et al. 2019).

Typus – CHINA, Hubei, Wudangshan Town, Wudangshan National Forest Park, on fallen

angiosperm branch, 20 Aug. 2017, L.W. Zhou, LWZ 20170820-35 (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, very thin, loosely attached to the

substrates, up to 9 cm long, 4 cm wide. Hymenophore smooth, cream to straw-yellow when fresh,

cream to ash-grey with age, not cracked. Margin undifferentiated.

Hyphal system monomitic; generative hyphae with clamp connections, hyaline, slightly thick-

walled, frequently branched and septate, loosely subparallel, 2–3.5 µm in diam. Cystidia abundant,

subulate, projecting beyond hymenium, hyaline, thick-walled, regularly covered with rectangular

crystals except at the apex, 50–80 × 3–5 µm. Basidia subclavate to suburniform, hyaline, thin-

walled, with four sterigmata and a basal clamp connection, 16–22 × 5–7 µm; basidioles in shape

similar to basidia, but slightly smaller. Basidiospores fusiform to slightly vermicular, hyaline, thin-

walled, smooth, inamyloid, indextrinoid, acyanophilous, (15–)15.5–17.5(–18.5) × 2.3–3 µm, L =

16.5 µm, W = 2.6 µm, Q = 6.5–6.9 (n = 60/2).

Other specimen (paratype) examined – CHINA, Guangxi, Longzhou County, Nonggang

National Nature Reserve, on fallen angiosperm branch, 3 July 2007, H.X. Xiong, Xiong 221 (IFP

009160).

Notes – Besides Subulicystidium acerosum, S. daii also resembles S. cochleum and S.

perlongisporum by the long (above 15 µm in length) and straight or slightly curved basidiospores;

however, S. cochleum differs in the presence of a bundle of needle-like crystals at the cystidial

crystalline sheath ends, while S. perlongisporum differs in narrower basidiospores (1.5–2.5 µm in

width; Ordynets et al. 2018).

Suillosporium Pouzar, Česká Mykol. 12(1): 31 (1958).

Type species – Suillosporium cystidiatum (D.P. Rogers) Pouzar, Česká Mykol. 12(1): 31

(1958).

Description – Basidiomes annual, resupinate, effused, thin. Hymenophore smooth,

grandinioid, odontioid, white to cream. Margin not differentiated. Hyphal system monomitic,

generative hyphae with clamp connections, hyaline. Cystidia (septocystidia) thin-walled, more or

less encrusted. Basidia shortly clavate, hyaline, thin-walled, with two to four sterigmata.

Basidiospores fusiform to navicular, thin- to slightly thick-walled, smooth, inamyloid, indextrinoid,

weakly cyanophilous. On wood.

899

Notes – Suillosporium was erected as a monotypic genus to accommodate Pellicularia

cystidiata (Pouzar 1958). Suillosporium was put in Botryobasidiaceae (Jülich 1981) and

morphologically related to Botryobasidium (Eriksson et al. 1984). Later, three additional species

were added to Suillosporium (Boidin & Gilles 1986b, Langer & Langer 2004, Kotiranta &

Saarenoksa 2006). Although Larsson (2007) doubted the placement of Suillosporium at higher

ranks and considered its taxonomic position as uncertain, He et al. (2019) still treated this genus in

Botryobasidiaceae. This issue was for the first time clarified on the basis of phylogenetic evidence

by Spirin et al. (2021), whose analyses indicated it as a member of Hydnodontaceae,

Trechisporales. The current phylogenies also recovered this topology (Figs 1–3).

Figure 11 – Basidiomes of Subulicystidium daii (LWZ 20170820-35, holotype). Scale bars: a = 1

cm, b = 1 mm.

Trechispora P. Karst., Hedwigia 29: 147 (1890).

Type species – Trechispora onusta P. Karst., Hedwigia 29: 147 (1890) = T. hymenocystis

(Berk. & Broome) K.H. Larss., Mycol. Res. 98(10): 1167 (1994).

= Pseudohydnum Rick, Annls mycol. 2(5): 409 (1904) Nom. illegit. non Pseudohydnum P.

Karst., Not. Sällsk. Fauna et Fl. Fenn. Förh. 9: 374 (1868).

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= Hydnodon Banker, Mycologia 5(6): 297 (1913).

= Fibuloporia Bondartsev & Singer, in Singer, Mycologia 36(1): 67 (1944).

= Scytinopogon Singer, Lloydia 8(3): 139 (1945).

= Echinotrema Park.-Rhodes, Trans. Br. mycol. Soc. 38(4): 367 (1955).

= Fibriciellum J. Erikss. & Ryvarden, Cortic. N. Eur., 3 Coronicium-Hyphoderma (Oslo):

373 (1975).

= Osteomorpha G. Arnaud ex Watling & W.B. Kendr., Naturalist (Hull), ser. 3 104(no. 948):

1 (1979).

= Cristelloporia I. Johans. & Ryvarden, Trans. Br. mycol. Soc. 72(2): 189 (1979).

= Dextrinodontia Hjortstam & Ryvarden, Mycotaxon 12(1): 172 (1980).

Figure 12 – Microscopic structures of Subulicystidium daii (drawn from the holotype). a Vertical

section of basidiomes. b Basidiospores. Scale bars = 10 μm.

Description – Basidiomes annual, resupinate, effused, stipitate, or clavarioid, soft, loosely

attached to the substrates. Hymenophore smooth, grandinioid, odontioid, hydnoid or poroid, white,

cream to cinnamon-buff, mycelial cords often present. Hyphal system monomitic to dimitic,

generative hyphae with clamp connections, hyaline, with typical ampullate septa, crystals common

on subicular hyphae. Cystidia present or absent. Basidia cylindrical, often with a slight median

constriction, hyaline, thin-walled, with two to four sterigmata and a basal clamp connection.

Basidiospores usually subglobose to ellipsoid, rarely allantoid or subangular, hyaline, thin-walled,

smooth, verrucose or aculeate, inamyloid, indextrinoid, acyanophilous. Conidiospores sometimes

present. On soil or wood.

901

Notes – Trechispora was erected as a monotypic genus for the corticioid species T. onusta

(Karsten 1890). Later, its species diversity was extremely enriched (e.g. Liberta 1966, 1973,

Larsson 1994, 1996). The concept was further enlarged, especially by absorbing Hydnodon with

stipitate basidiomes (Ryvarden 2002) and Scytinopogon with clavarioid basidiomes (de Meiras-

Ottoni et al. 2021). When the family Hydnodontaceae and the order Hydnodontales were erected,

Trechispora was included as one of exemplar genera (Jülich 1981). Furthermore, Trechispora is the

automatic type genus of Trechisporales (equivalent to Hydnodontales; Hibbett et al. 2007). The

shared morphological characters of species in Trechispora are remarkable ampullate hyphal septa,

and in addition most species in this genus have ellipsoid, ornamented basidiospores, but there is

considerable diversity in basidiome form.

It is noteworthy that of the currently accepted members of Trechispora, T. clancularis, T.

molliuscula, T. mollusca and T. silvae-ryae were, respectively, placed at one time in four

monotypic genera Echinotrema, Dextrinodontia, Fibuloporia and Fibriciellum based on

morphological characters (Larsson 1992, 1994).

Echinotrema was introduced for E. clanculare, a resupinate species from the United

Kingdom, described as having a hymenium of “parallel sinuous plates” and echinulate

basidiospores, with these characters compared by the original author to those of Lindtneria and

Sistotrema (Parker-Rhodes 1955). Larsson (1994) examined the type collection, which he described

as having a poroid to irpicoid hymenium, and found frequently ampullate septa and aculeate

basidiospores. Although Larsson (1994) found some unusual features in E. clanculare, especially

the basidia with an oblique medial widening, such structures were present in some other species of

Trechispora, and consequently he did not consider that there were sufficient grounds to recognize

Echinotrema as distinct from Trechispora. Dextrinodontia was erected to accommodate an African

new species D. molliuscula (Hjortstam & Ryvarden 1980). This genus was listed as a member of

Hydnodontaceae, Trechisporales with doubt by Larsson (2007) and as a potential member of

Trechisporales by Hibbett et al. (2014). Fibriciellum was described to accommodate a new species

F. silvae-ryae from Sweden (Eriksson & Ryvarden 1975), and later was placed in Hydnodontaceae,

Trechisporales (Larsson 2007). Recently, an ITS sequence (MZ159622) of F. silvae-ryae was

submitted to GenBank. Although this ITS sequence was generated from a voucher collected from

England instead of type locality, as the single molecular sequence of this species, its BLAST search

revealed an affinity to members of Trechispora. For now, we accept Echinotrema, Dextrinodontia

and Fibriciellum as synonyms of Trechispora but the placement of species used to typify the three

genera should be further clarified ideally with the aid of multilocus-based phylogenetic analyses

from more samples.

Fibuloporia was introduced for F. mollusca (based on Boletus molluscus Pers.) by Singer

(1944) with minimal discussion and no indication that the fungus has ornamented basidiospores,

from which it can be gathered that the erroneous interpretation of B. molluscus as a smooth-spored

species was being followed. For discussion of the complex situation around the typification of B.

molluscus see Larsson (1994, 2001). With acceptance of material collected by Persoon and

designated by Donk as the neotype of B. molluscus, the name must be interpreted as based on a

species with rough basidiospores, thus the placement in Trechispora by Liberta (1973), which is

followed by subsequent authors including Larsson (1994). Given that sequenced material is

available for Trechispora mollusca, there is no doubt about the synonymy of Fibuloporia and

Trechispora.

Scytinopogon is another noteworthy synonym of Trechispora. This genus, typified by

Scytinopogon pallescens, was erected for clavarioid species (Singer 1945). Although

macromorphology initially indicated placement of Scytinopogon in the family Clavariaceae,

Scytinopogon microscopically deviates by rather short basidia and ellipsoid basidiospores with

ornamentations and thus was put in its own new family Scytinopogonaceae as the type genus

(Jülich 1981). Jülich (1981) also suggested the morphological affinity of Scytinopogon to

Hydnodon (a synonym of Trechispora; Ryvarden 2002) and Trechispora, and placed

Scytinopogonaceae in Hydnodontales (equivalent to Trechisporales; Hibbett et al. 2007).

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Molecular evidence not only supported the close relationship between Scytinopogon and

Trechispora (Larsson et al. 2011), but also grouped these two genera in a clade with strong support

(Birkebak et al. 2013, de Meiras-Ottoni et al. 2021) or weak support (Desjardin & Perry 2015). de

Meiras-Ottoni et al. (2021) stated that the weak support in the phylogeny of Desjardin & Perry

(2015) resulted from one problematic sequence in their dataset. Moreover, de Meiras-Ottoni et al.

(2021) carefully compared the morphological characters of types of Scytinopogon and Trechispora,

and indicated that both share the remarkable ampullate hyphal septa and ellipsoid, ornamented

basidiospores. Therefore, Scytinopogon was formally proposed as a later synonym of Trechispora

(de Meiras-Ottoni et al. 2021). The current phylogeny sampling more taxa of Trechispora clearly

supports the monophyly of this genus with inclusion of species formerly belonging to Scytinopogon

(Fig. 6). After transferring five species from Scytinopogon to Trechispora (combined below), only

two species, viz. S. echinosporus and S. parvus are still left in Scytinopogon due to unavailable of

their molecular sequences.

The illegitimate genus name Pseudohydnum J. Rick is included under Trechispora following

Donk (1956), who treated the type species P. guepinioides as a synonym of Hydnum thelephorum

(Trechispora thelephora).

Osteomorpha was introduced for an asexual species O. fragilis that was associated with a

species of Trechispora. Stalpers et al. (2021) discussed the connection between Osteomorpha and

Trechispora and concluded that the two genera are synonymous.

Cristelloporia was described by Johansen & Ryvarden (1979) for Cristelloporia dimitica I.

Johans. & Ryvarden, a resupinate, poroid fungus with asperulate, ellipsoid to irregularly lobed

basidiospores and needle-like crystals among dimitic hyphae, found in Ghana, Africa. The authors

noted that “When first examined under the microscope, C. dimitica was considered to be a new

species of Trechispora Karst. and there are several characters pointing towards this genus,

including the aculeate spores, the needle-like crystals and the many pleurobasidia”. However, they

concluded that “it would be better placed in the Polyporaceae [i.e. as a new genus] because of the

very distinct dimitic hyphal system giving the fruitbodies a cottony and coriaceous consistency so

typical for many resupinate polypores”. Cristelloporia is listed as a synonym of Trechispora by

Index Fungorum, following the entry in the 10th edition of Dictionary of the Fungi (Kirk et al.

2008) where the source of the synonym is noted as “Larsson in litt.” which would refer to Larsson

(1992) and this placement is followed by Gorjón (2020) and He et al. (2019). Larsson (1992)

considered that the type, Cristelloporia dimitica, was synonymous with C. brasiliensis Corner and

Heterobasidion pahangense Corner. Because Trechispora dimitica Hallenb. blocked transfer of

C. dimitica to Trechispora, Larsson (1992) took up the next available epithet, brasiliensis, but

without making a valid transfer to Trechispora. Later, C. brasiliensis was validly transferred to

Trechispora by Chikowski et al. (2020) on the basis of morphological characters. Even though

sequences are not available for T. brasiliensis, we accept the placement in Trechispora taken up by

Larsson (1992) and Chikowski et al. (2020), which renders Cristelloporia a synonym of

Trechispora. Note that Hattori (2003) treated Heterobasidion pahangense as an independent

species (combined in Cristelloporia). Of the three other species placed in Cristelloporia:

C. trimitica was found by Hattori (2003) to represent a trimitic species of Trametes, for which the

type collection was contaminated by another fungus with echinulate spores; the position of

C. asperispora has not been re-examined since it was originally described from Kenya; and for C.

rutilantiformis see under Murilloporus under Genus of uncertain position (below).

Trechispora caulocystidiata (A.N.M. Furtado & M.A. Neves) L.W. Zhou & S.L. Liu, comb. nov.

Index Fungorum number: IF559883; Facesoffungi number: FoF12863

Basionym. Scytinopogon caulocystidiatus A.N.M. Furtado & M.A. Neves, in Furtado,

Daniëls, Reck & Neves, Mycotaxon 136(1): 113 (2021).

Notes – Trechispora caulocystidiata was recently described as a new species in Scytinopogon

on the basis of morphological and molecular data (Furtado et al. 2021). Although Furtado et al.

(2021) noticed that Trechispora has been formally treated as having priority over Scytinopogon (de

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Meiras-Ottoni et al. 2021), they did not put this new species in Trechispora due to lack of a

comprehensive phylogeny. The current phylogeny (Fig. 6) undoubtedly supports that S.

caulocystidiatus nests within Trechispora, and thus we make this combination. The detailed

description of T. caulocystidiata can be found in Furtado et al. (2021).

Trechispora chaibuxiensis S.L. Liu, L.W. Zhou & S.H. He, sp. nov. Figs 13–15

Index Fungorum number: IF559885; Facesoffungi number: FoF12864

Etymology – chaibuxiensis (Latin), refers to Chaibuxi Grand Canyon Scenic Spot.

Diagnosis – Differs from Trechispora subsinensis (described below) in the presence of

hyphoid cystidia.

Typus – CHINA, Hubei, Wufeng County, Chaibuxi Grand Canyon Scenic Spot, on fallen

angiosperm branch, 14 Aug. 2017, L.W. Zhou, LWZ 20170814-34 (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft and fragile, easily separated

from substrates, up to 5 cm long, 2 cm wide. Hymenophore odontioid with numerous small aculei,

sometimes fertile at the apex of the aculei, cream to straw-yellow when fresh, straw-yellow when

dry, up to 0.3 mm long. Margin white, fimbriate, up to 0.5 mm wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae hyaline,

slightly thick-walled, moderately branched and septate, interwoven, 2.5–4.5 µm diam, ampullate

septa up to 6 μm wide. Aculei composed of a central core of compact hyphae and subhymenial and

hymenial layers; generative hyphae distinct, hyaline, thin to slightly thick-walled, occasionally

branched, smooth, subparallel, 2–3.5 μm in diam, apical ends in aculei with basidia and rare

hyphoid cystidia. Crystals usually present, bipyramidic, aggregated. Hyphoid cystidia rare, smooth,

thin-walled, fusoid, 20–35 × 3–5 µm. Basidia cylindrical with a slight median constriction, hyaline,

thin-walled, with four sterigmata and a basal clamp connection, 12–20 × 3.5–4.5 µm; basidioles in

shape similar to basidia, but slightly smaller. Basidiospores broadly ellipsoid, hyaline, thin-walled,

aculeate, inamyloid, indextrinoid, acyanophilous, (2.6–)2.8–3.3 × (2–)2.2–2.8(–2.9) µm, L = 3 µm,

W = 2.6 µm, Q = 1.1–1.2 (n = 90/3).

Other specimens (paratypes) examined – CHINA, Hubei, Wufeng County, Chaibuxi Grand

Canyon Scenic Spot, on fallen angiosperm branch, 14 Aug. 2017, L.W. Zhou, LWZ 20170814-35

(HMAS), L.W. Zhou, LWZ 20170814-36 (HMAS), L.W. Zhou, LWZ 20170814-42 (HMAS); on

fallen angiosperm branch, 15 Aug. 2017, S.H. He, He 5072 (BJFC 024590).

Notes – Trechispora chaibuxiensis is characterized by the odontioid hymenophore with

numerous small aculei, a monomitic hyphal system, the presence of hyphoid cystidia and basidia in

apical ends of aculei, and broadly ellipsoid, aculeate basidiospores. Besides Trechispora

subsinensis, T. chaibuxiensis could also be confused with T. nivea, but the latter species has a

hymenophore with longer aculei (up to 1 mm in length) and lacks cystidia (Larsson 1995). Similar

to Trechispora chaibuxiensis, other species such as T. caulocystidiata, T. gelatinosa and T.

minispora also have cystidial structures, but they differ in the clavarioid basidiomes (de Meiras-

Ottoni et al. 2021).

Trechispora constricta S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 16–18

Index Fungorum number: IF559886; Facesoffungi number: FoF12865

Etymology – constricta (Latin), refers to constriction on the spines of basidiospores.

Diagnosis – Characterized by the presence of a slight constriction on the spines of

basidiospores.

Typus – CHINA, Jiangxi, Fenyi County, Dagangshan Nature Reserve, on rotten angiosperm

wood, 19 Sept. 2008, Y.C. Dai, Dai 10534 (holotype in BJFC 004783).

Description – Basidiomes annual, resupinate, effused, thin, soft, easily separated from

substrates, up to 3 cm long, 2 cm wide. Hymenophore odontioid, white to cream when fresh, cream

to buff-yellow with age, not cracked. Aculei 5–8 per mm, up to 0.5 mm long. Margin thinning out

as byssoid, white.

904

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin-walled, moderately branched, septate, interwoven, 3–5 µm in diam, ampullate septa

up to 7 µm wide. Tramal generative hyphae distinct, hyaline, thin-walled, moderately branched,

smooth, subparallel, 3–6 μm in diam. Crystals usually present, rhomboidal. Cystidia absent.

Basidia cylindrical with a slight median constriction, hyaline, thin-walled, with four sterigmata and

a basal clamp connection, 11–15 × 4–5.5 µm; basidioles in shape similar to basidia, but slightly

smaller. Basidiospores ellipsoid, hyaline to yellowish, thin-walled, aculeate, with a slight

constriction in the middle-upper part of spines, inamyloid, indextrinoid, acyanophilous, 3–4 × 2.3–

2.9(–3) µm, L = 3.4 µm, W = 2.6 µm, Q = 1.3 (n = 60/2).

Other specimens (paratypes) examined – CHINA, Jiangxi, Fenyi County, Dagangshan Nature

Reserve, on rotten angiosperm wood, 18 Sept. 2008, Y.C. Dai, Dai 10488 (BJFC 004737);

Guangdong, Shixing County, Chebaling National Nature Reserve, on angiosperm stump, 14 June

2019, S.H. He, He 5899 (BJFC 030774).

Figure 13 – Basidiomes of Trechispora chaibuxiensis (LWZ 20170814-34, holotype). Scale bars:

a = 1 cm, b = 1 mm.

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Figure 14 – Microscopic structures of Trechispora chaibuxiensis (drawn from the holotype).

a Vertical section of basidiomes. b Basidia. c Cystidia. d Basidiospores. Scale bars = 10 μm.

Figure 15 – Scanning electron micrograph of basidiospores of Trechispora chaibuxiensis (scanned

from the holotype). Scale bar = 3 μm.

Notes – Trechispora constricta is characterized by wide generative hyphae (3–6 µm in diam)

and aculeate basidiospores with a slight constriction in the middle-upper part of spines (Fig. 18).

Trechispora constricta resembles T. tropica (described below), but the latter species differs also in

906

smaller basidiospores (2.5–3 × 2.2–2.5 µm) besides lack of a constriction on the spines of

basidiospores.

Figure 16 – Basidiomes of Trechispora constricta (Dai 10534, holotype). Scale bars: a = 1 cm, b =

1 mm.

Trechispora crystallina S.L. Liu & L.W. Zhou, sp. nov. Figs 19–21

Index Fungorum number: IF559887; Facesoffungi number: FoF12866

Etymology – crystallina (Latin), refers to crystals.

Diagnosis – Characterized by crystals abundant in subiculum and trama, and verrucose

basidiospores.

Typus – VIETNAM, Ho Chi Minh City, Le Thi Rieng Park, on living angiosperm tree, 13

Oct. 2017, L.W. Zhou, LWZ 20171013-7 (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft and fragile, easily separated

from substrates, up to 10 cm long, 2.5 cm wide. Hymenophore grandinioid with numerous small

aculei, white to cream when fresh, cream to straw-yellow when dry. Margin white, slightly

fimbriate, up to 0.2 mm wide.

907

Figure 17 – Microscopic structures of Trechispora constricta (drawn from the holotype). a Vertical

section of basidiomes. b Basidiospores. Scale bars = 10 μm.

Figure 18 – Scanning electron micrograph of basidiospores of Trechispora constricta (scanned

from the holotype). Scale bar = 2 μm.

908

Figure 19 – Basidiomes of Trechispora crystallina (LWZ 20171013-7, holotype). Scale bars: a = 1

cm, b = 0.5 mm.

Hyphal system monomitic; generative hyphae with clamp connections. Subiculum composed

of indistinct generative hyphae; subicular hyphae hyaline, thin-walled, frequently branched and

septate, interwoven, 2–3.5 µm in diam, ampullate septa up to 6 µm wide. Tramal generative hyphae

distinct, hyaline, thin-walled, frequently branched, smooth, interwoven, 3–5 μm in diam. Crystals

occurring in both subiculum and trama, as small, aggregated rhomboidal flakes. Cystidia absent.

Basidia cylindrical with a slight median constriction, hyaline, thin-walled, with four sterigmata and

a basal clamp connection, 17–22 × 4–6 µm; basidioles in shape similar to basidia, but slightly

smaller. Basidiospores ellipsoid, hyaline, thin to slightly thick-walled, verrucose, inamyloid,

indextrinoid, acyanophilous, (3.2–)3.5–4.2(–5) × (2.8–)3–3.6(–3.8) µm, L = 4 µm, W = 3.2 µm, Q

= 1.2–1.3 (n = 60/2).

Other specimen (paratype) examined – CHINA, Inner Mongolia, Tongliao, Daqinggou

National Nature Reserve, on fallen angiosperm twig, 29 July 2017, L.W. Zhou, LWZ 20170729-2

(HMAS).

909

Figure 20 – Microscopic structures of Trechispora crystallina (drawn from the holotype).

a Vertical section of basidiomes b Basidia. c Basidiospores. Scale bars = 10 μm.

Figure 21 – Scanning electron micrograph of basidiospores of Trechispora crystallina (scanned

from the holotype). Scale bar = 2 μm.

Notes – Trechispora crystallina resembles T. cyatheae and T. torrendii by whitish to

yellowish, grandinioid hymenophore, thin subiculum and a monomitic hyphal system with thin-

910

walled hyphae (Ordynets et al. 2015, Chikowski et al. 2020). However, T. cyatheae differs in

aculeate, smaller basidiospores (3–3.5 × 2–3 µm including spines) and growth exclusively on

Cyathea glauca, an endemic species of tree fern to La Réunion, France (Ordynets et al. 2015); and

T. torrendii differs in the absence of crystals and aculeate, smaller basidiospores (3.2–3.5 × 2.8–3.2

µm including spines; Chikowski et al. 2020).

Trechispora damansaraensis S.L. Liu, L.W. Zhou & S.H. He, sp. nov. Figs 22–23

Index Fungorum number: IF559888; Facesoffungi number: FoF12867

Etymology – damansaraensis (Latin), refers to Kota Damansara Community Forest Reserve.

Diagnosis – Characterized by the combination of cream, smooth hymenophore and the

occasional presence of crystals.

Typus – MALAYSIA, Selangor, Kota Damansara Community Forest Reserve, on fallen

angiosperm branch, 17 Apr. 2018, L.W. Zhou, LWZ 20180417-26 (holotype in HMAS).

Figure 22 – Basidiomes of Trechispora damansaraensis (LWZ 20180417-26, holotype). Scale

bars: a = 1 cm, b = 1 mm.

911

Figure 23 – Microscopic structures of Trechispora damansaraensis (drawn from the holotype).

a Vertical section of basidiomes. b Hyphae in subiculum. c Basidia. d Basidiospores. Scale bars =

10 μm.

Description – Basidiomes annual, resupinate, effused, thin, soft and fragile, loosely attached

to the substrates. Hymenophore smooth, farinaceous, white to cream when fresh, cream,

occasionally cracked with age. Margin thinning out, fimbriate, slightly paler than hymenophore,

becoming indistinct with age. Mycelial cords present, white, 0.5 mm wide. Hyphal system

monomitic; generative hyphae with clamp connections. Subicular hyphae hyaline, thin to slightly

thick-walled, frequently branched and septate, interwoven, 1–4 µm in diam, ampullate septa up to 5

μm wide, thin, flexuous. Subhymenium composed of indistinct generative hyphae, 2–4 µm in diam,

much branched, flexuous. Crystals occasionally present, as aggregated rhomboidal flakes. Cystidia

absent. Basidia cylindrical with a slight median constriction, thin-walled, with four sterigmata and

a basal clamp connection, 9–12 × 5–6.5 µm; basidioles in shape similar to basidia, but slightly

smaller. Basidiospores ellipsoid, hyaline, thin to slightly thick-walled, aculeate, inamyloid,

indextrinoid, acyanophilous, (2.8–)3–3.8 × (2.1–)2.3–3(–3.2) µm, L = 3.2 µm, W = 2.7 µm, Q = 1.2

(n = 60/2).

Other specimen (paratype) examined – MALAYSIA, Selangor, Kota Damansara Community

Forest Reserve, on fallen angiosperm branch, 7 Dec. 2019, S.H. He, He 6415 (BJFC 033359).

Notes – Trechispora minima mostly resembles T. damansaraensis by smooth hymenophore,

a monomitic hyphal system and occasional presence of crystals in subiculum, but differs in parallel

generative hyphae in subiculum, narrower basidia (4.5–5 µm in width) and subglobose to broadly

ellipsoid basidiospores (Larsson 1996).

Trechispora dealbata (Berk.) L.W. Zhou & S.L. Liu, comb. nov.

Index Fungorum number: IF559889; Facesoffungi number: FoF12868

Basionym. Clavaria dealbata Berk., Hooker's J. Bot. Kew Gard. Misc. 8: 275 (1856).

912

≡ Scytinopogon dealbatus (Berk.) Corner, Beih. Nova Hedwigia 33: 89 (1970).

Notes – de Meiras-Ottoni et al. (2021) showed that Scytinopogon dealbatus nested within

Trechispora on the basis of molecular data, but they did not propose any taxonomic change because

of lack of morphological information on relevant specimens. Later but almost simultaneously,

Furtado et al. (2021) provided a detailed morphological description of S. dealbatus, noting that

‘additional DNA regions and species are needed before concluding that Scytinopogon and

Trechispora are fully synonymous’. The current phylogenies sampling more gene regions (Fig. 3)

and sequence-available species of Scytinopogon and Trechispora (Fig. 6) undoubtedly support

Scytinopogon and Trechispora as congeneric. Therefore, we transfer S. dealbatus to Trechispora as

T. dealbata.

Trechispora foetida (A.N.M. Furtado & M.A. Neves) L.W. Zhou & S.L. Liu, comb. nov.

Index Fungorum number: IF559890; Facesoffungi number: FoF12869

Basionym. Scytinopogon foetidus A.N.M. Furtado & M.A. Neves, in Furtado, Daniëls, Reck

& Neves, Mycotaxon 136(1): 119 (2021).

Notes – Scytinopogon foetidus was newly described together with S. caulocystidiatus

(Furtado et al. 2021). Similar to the treatment of S. caulocystidiatus, we transfer S. foetidus to

Trechispora as T. foetida. The detailed description of T. foetida can be found in Furtado et al.

(2021).

Trechispora gracilis S.L. Liu & L.W. Zhou, sp. nov. Figs 24–26

Index Fungorum number: IF559891; Facesoffungi number: FoF12870

Etymology – gracilis (Latin), refers to thin basidiomes.

Diagnosis – Characterized by the combination of thin, corticioid basidiomes with ash-grey,

smooth hymenophore and the absence of crystals.

Typus – CHINA, Jiangxi, Jiujiang, Bailudong Academy, on fallen gymnosperm branch, 26

June 2021, L.W. Zhou, LWZ 20210626-5b (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft, easily separated from

substrates, up to 10 cm long, 2 cm wide, 50 µm thick. Hymenophore smooth, arachnoid, cream to

light ash-grey when fresh, ash-grey when dry, not cracked. Margin white, thinning out as byssoid,

0.5 mm wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

long-celled, hyaline, slightly thick-walled, frequently branched and septate, interwoven, 2.5–3.5

µm in diam, ampullate septa up to 5 μm wide. Subhymenium composed of indistinct generative

hyphae, much branched. Crystals absent. Cystidia absent. Basidia cylindrical with a slight median

constriction, hyaline, thin-walled, with four sterigmata and a basal clamp connection, 10–13 × 4–5

µm; basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid, hyaline,

thin-walled, aculeate, inamyloid, indextrinoid, acyanophilous, (2.5–)2.8–3.2 × (2–)2.3–2.8(–2.9)

µm, L = 3.0 µm, W = 2.5 µm, Q = 1.2 (n = 60/2).

Other specimens (paratypes) examined – CHINA, Hubei, Huanggang, Dabieshan National

Nature Reserve, on fallen gymnosperm branch, 19 Sept. 2021, L.W. Zhou, LWZ 20210919-9a

(HMAS); Macheng, Shizifeng Nature Reserve, on fallen twig of Pinus, 22 Sept. 2021, L.W. Zhou,

LWZ 20210922-7b (HMAS); Wufeng County, Chaibuxi Grand Canyon Scenic Spot, on fallen

angiosperm twig, 17 Aug. 2017, L.W. Zhou, LWZ 20170814-17 (HMAS).

Notes – Trechispora gracilis resembles T. damansaraensis by thin basidiomes and smooth

hymenophore; however, T. damansaraensis differs in white to cream hymenophore, the occasional

presence of crystals in subiculum and slightly longer basidiospores (3–3.8 µm in length).

Trechispora larssonii S.L. Liu, L.W. Zhou & S.H. He, sp. nov. Figs 27–29

Index Fungorum number: IF559892; Facesoffungi number: FoF12872

Etymology – larssonii (Latin), refers to the Swedish mycologist, Prof. Dr. Karl-Henrik

Larsson, who has made most significant contributions to the taxonomy of Trechispora.

913

Diagnosis – Differs from Trechispora minima by the common presence of crystals and

verrucose basidiospores (Larsson 1996).

Typus – CHINA, Sichuan, Muchuan County, Qincaiping Nature Reserve, on fallen

gymnosperm twig, 17 Aug. 2019, L.W. Zhou, LWZ 20190817-11a (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft, fragile, easily separated from

substrates, up to 15 cm long, 2 cm wide. Hymenophore smooth, farinaceous, white to cream when

fresh, cream to buff-yellow with age, finely cracked. Margin thinning out as byssoid, narrow, white

to cream.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin-walled, moderately branched and septate, subparallel, 2.5–5 µm in diam, ampullate

septa up to 7 µm wide. Subhymenial hyphae short-celled and wide, 3–5 µm in diam, much

branched. Crystals common, as aggregated rhomboidal flakes. Cystidia absent. Basidia cylindrical

with a slight median constriction, thin-walled, with four sterigmata and a basal clamp connection,

7–13 × 4–5 µm; basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid,

hyaline, thin-walled, verrucose, inamyloid, indextrinoid, acyanophilous, (2.5–)2.8–3.3(–3.5) ×

(2.5–)2.1–2.8(–3.3) µm, L = 3 µm, W = 2.5 µm, Q = 1.2 (n = 60/2).

Figure 24 – Basidiomes of Trechispora gracilis (LWZ 20210626-5b, holotype). Scale bars: a = 1

cm, b = 1 mm.

914

Figure 25 – Microscopic structures of Trechispora gracilis (drawn from the holotype). a Vertical

section of basidiomes. b Basidia. c Basidiospores. Scale bars = 10 μm.

Figure 26 – Scanning electron micrograph of basidiospores of Trechispora gracilis (scanned from

the holotype). Scale bar = 3 μm.

Other specimens (paratypes) examined – CHINA, Sichuan, Meigu County, Dafengding

National Nature Reserve, on fallen gymnosperm branch, 18 Aug. 2020, L.W. Zhou, LWZ

20200818-10b (HMAS); Guizhou, Chishui County, Alsophila National Nature Reserve, on dead

branch of living Osmunda vachellii, 7 July 2018, S.H. He, He 5450 (BJFC 026511).

915

Notes – Besides Trechispora minima, which differs by occasional occurrence of crystals and

aculeate basidiospores (Larsson 1996), T. larssonii is also similar to T. damansaraensis by smooth

and farinaceous hymenophore, but T. damansaraensis differs in slightly narrower generative

hyphae in subiculum and subhymenium (up to 4 µm), the occasional presence of crystals, and

aculeate, slightly longer basidiospores (3–3.8 µm in length).

Figure 27 – Basidiomes of Trechispora larssonii (LWZ 20190817-11a, holotype). Scale bars: a = 1

cm, b = 1 mm.

Trechispora latehypha S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 30–32

Index Fungorum number: IF559893; Facesoffungi number: FoF12873

Etymology – latehypha (Latin), refers to wide subhymenial hyphae.

Diagnosis – Characterized by the rows of short-celled and wide hyphae in subhymenia, thick-

walled generative hyphae in subiculum, and aculeate basidiospores.

Typus – CHINA, Guangdong, Renhua County, Danxiashan National Nature Reserve, on

fallen angiosperm trunk, 4 June 2019, S.H. He, He 5848 (holotype in BJFC 030723).

916

Figure 28 – Microscopic structures of Trechispora larssonii (drawn from the holotype). a Vertical

section of basidiomes. b Basidia. c Basidiospores. Scale bars = 10 μm.

Figure 29 – Scanning electron micrograph of basidiospores of Trechispora larssonii (scanned from

the holotype). Scale bar = 2 μm.

Description – Basidiomes annual, resupinate, effused, thin, soft, fragile, inseparable from

substrate, up to 15 cm long, 4 cm wide. Hymenophore smooth, arachnoid, white to cream when

fresh, cream to buff-yellow with age, finely cracked with age. Margin white, thinning out as

byssoid, 1 mm wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae long-

celled, hyaline, slightly thick-walled to thick-walled, moderately branched and septate, 3.5–6 µm in

diam. Subhymenial hyphae short-celled and wide, hyaline, thin-walled, 4.5–7 µm in diam, much

917

branched. Cystidia absent. Basidia cylindrical with a slight median constriction, hyaline, thin-

walled, with four sterigmata and a basal clamp connection, 13–16 × 4–5.5 µm; basidioles in shape

similar to basidia, but slightly smaller. Basidiospores ellipsoid, hyaline, thin-walled, aculeate,

inamyloid, indextrinoid, acyanophilous, (2.9–)3–3.5(–3.8) × (2.2–)2.4–2.9(–3) µm, L = 3.2 µm, W

= 2.6 µm, Q = 1.2 (n = 60/2).

Other specimens (paratypes) examined – CHINA, Fujian, Wuyishan, Wuyishan National

Nature Reserve, on fallen branch of Tsuga, 17 Aug. 2016, S.H. He, He 4472 (BJFC 023913).

Guangdong, Renhua County, Danxiashan National Nature Reserve, on fallen angiosperm trunk, 4

June 2019, S.H. He, He 5838 (BJFC 030705). Hainan, Qiongzhong County, Limushan National

Forest Park, on fallen angiosperm trunk, 16 June 2017, L.W. Zhou, LWZ 20170611-16 (HMAS);

Wuzhishan, Wuzhishan National Forest Park, on dead branch of living angiosperm, 10 June 2016,

S.H. He, He 3924 (BJFC 022426).

Figure 30 – Basidiomes of Trechispora latehypha (He 5848, holotype). Scale bars: a = 1 cm, b = 2

mm.

918

Figure 31 – Microscopic structures of Trechispora latehypha (drawn from the holotype). a Vertical

section of basidiomes; b. Basidiospores. Scale bars = 10 μm.

Figure 32 – Scanning electron micrograph of basidiospores of Trechispora latehypha (scanned

from the holotype). Scale bar = 2 μm.

Notes – Trechispora latehypha is similar to T. larssonii by more or less smooth

hymenophoral configuration and rows of short-celled and wide subhymenial hyphae. However, T.

919

larssonii has thin-walled subicular hyphae and verrucose basidiospores. Trechispora farinacea also

resembles T. latehypha by short-celled and wide subhymenial hyphae, but differs in more or less

grandinioid to nearly hydnoid hymenophore, and subglobose to broadly ellipsoid basidiospores

(Larsson 1995). In addition, T. farinacea is a common species in the north temperate zone,

especially in Europe (Bernicchia & Gorjón 2010), whereas T. latehypha is only known in southern

China.

Trechispora longiramosa S.L. Liu, G. He, Shuang L. Chen & L.W. Zhou, sp. nov. Figs 33–35

Index Fungorum number: IF559894; Facesoffungi number: FoF12874

Etymology – longiramosa (Latin), refers to long terminal branches of basidiomes.

Diagnosis – Characterized by U-shaped, dense, long, thin terminal branches with acute tips

and a fishy odor.

Typus – CHINA, Guizhou, Libo County, Maolan National Nature Reserve, on ground, 17

July 2019, G. He, CH 19233 (holotype in HMAS).

Figure 33 – Basidiomes of Trechispora longiramosa (CH 19233, holotype). a Fresh specimen.

b Hymenophore on branches of dried specimens. Scale bars: a = 1 cm, b = 1 mm.

920

Description – Basidiomes annual, clavarioid, solitary or in small groups, densely branched,

moderately open, fleshy consistency, cream to buff turning yellowish brown towards the apex when

fresh, olivaceous buff turning dark brown towards the apex when dry, with fishy odor, 7 cm high.

Branches polychotomous, axils U-shaped, tips acute, white to honey-yellow. Stipe white to cream,

5–15 × 1–2 mm.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin-walled, cylindrical, moderately branched and septate, subparallel, 3–8 μm in diam;

ampullate septa usually present in the hyphae at the base of the stipe, up to 10 μm wide.

Subhymenial hyphae short-celled and wide, 4–8 µm in diam, much branched. Cystidia absent.

Basidia suburniform to subclavate, hyaline, thin-walled, with four sterigmata and a basal clamp

connection, agglutinated, 15–23 × 8–11 µm; basidioles in shape similar to basidia, but slightly

smaller. Basidiospores ellipsoid, hyaline to yellowish, slightly thick-walled, aculeate with spines

slightly swelled in 5% potassium hydroxide, inamyloid, indextrinoid, acyanophilous, (4–)4.8–6 ×

(3–)3.3–4(–4.5) µm, L = 5.2 µm, W = 3.8 µm, Q = 1.4 (n = 60/2).

Figure 34 – Microscopic structures of Trechispora longiramosa (drawn from the holotype).

a Hyphae in subiculum. b Vertical section of basidiomes. c Basidia. d Basidiospores. Scale bars =

10 μm.

921

Figure 35 – Scanning electron micrograph of basidiospores of Trechispora longiramosa (scanned

from the holotype). Scale bar = 3 μm.

Other specimen (paratype) examined – CHINA, Guizhou, Libo County, Maolan National

Nature Reserve, on ground, 13 July 2014, G. He, HG 140168 (MCCNNU 00968).

Notes – Micromorphologically, T. longiramosa is similar to T. copiosa by similar shape and

size of basidiospores; however, T. copiosa differs in V-shaped branches and pale greyish yellow to

beige basidiomes (de Meiras-Ottoni et al. 2021).

Trechispora malayana S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 36–38

Index Fungorum number: IF559895; Facesoffungi number: FoF12875

Etymology – malayana (Latin), refers to the Malay Peninsula.

Diagnosis – Characterized by the combination of hydnoid hymenophore, the presence of

mycelial cords, and long spines on basidiospores (0.6–1 µm in length).

Typus – SINGAPORE, Bukit Timah Nature Reserve, on rotten angiosperm wood, 20 July

2017, Y.C. Dai, Dai 17876 (holotype in BJFC 025408).

Description – Basidiomes annual, resupinate, effused, thin, soft, fragile, easily separated from

substrates, up to 10 cm long, 5 cm wide. Hymenophore hydnoid, cream to buff-yellow with age.

Aculei white when fresh, 5–6 per mm, up to 0.8 mm long. Margin thinning out as byssoid, white,

up to 1.5 mm wide. Mycelial cords present, white, 0.5 mm wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin-walled, frequently branched and septate, interwoven, 1.5–4 µm in diam, ampullate

septa up to 6 µm wide. Tramal generative hyphae distinct, hyaline, thin-walled, frequently

branched, smooth, subparallel to interwoven, 2–4 μm in diam. Crystals usually present,

bipyramidic, aggregated. Cystidia absent. Basidia cylindrical with a slight median constriction,

hyaline, thin-walled, with four sterigmata and a basal clamp connection, 20–28 × 4–5 µm;

basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid, hyaline, thin-

walled, aculeate, inamyloid, indextrinoid, acyanophilous, 3.3–4.1(–4.5) × (2.3–)2.5–3(–3.5) µm, L

= 3.8 µm, W = 2.9 µm, Q = 1.3–1.4 (n = 60/2).

Other specimen (paratype) examined – THAILAND, Krabi, Khao Phanom Bencha National

Park, on rotten angiosperm trunk, 28 July 2016, S.H. He, He 4156 (BJFC 023598).

922

Notes – Trechispora malayana may be confused with T. nivea, but T. nivea differs in slightly

thick-walled tramal hyphae and shorter spines on basidiospores (up to 0.3 µm in length; Larsson

1995).

Trechispora robusta (Rick) L.W. Zhou & S.L. Liu, comb. nov.

Index Fungorum number: IF559896; Facesoffungi number: FoF12876

Basionym. Clavaria robusta Rick, Egatea 16: 120 (1931).

≡ Scytinopogon robustus (Rick) Corner, Beih. Nova Hedwigia 33: 91 (1970).

Notes – The proposal of this combination is on the basis of the same reason as that for

Trechispora dealbata indicated above. The detailed description of T. robusta can be found in

Furtado et al. (2021).

Figure 36 – Basidiomes of Trechispora malayana (Dai 17876, holotype). Scale bars: a = 1 cm, b =

1 mm.

923

Figure 37 – Microscopic structures of Trechispora malayana (drawn from the holotype). a Vertical

section of basidiomes; b. Basidiospores. Scale bars = 10 μm.

Figure 38 – Scanning electron micrograph of basidiospores of Trechispora malayana (scanned

from the holotype). Scale bar = 3 μm.

Trechispora scabra (Berk. & M.A. Curtis) L.W. Zhou & S.L. Liu, comb. nov.

Index Fungorum number: IF559897; Facesoffungi number: FoF12877

924

Basionym. Thelephora scabra Berk. & M.A. Curtis, Amer. J. Sci. Arts, Ser. 2 11: 94 (1851).

≡ Scytinopogon scaber (Berk. & M.A. Curtis) D.A. Reid, Persoonia 2(2): 161 (1962).

Notes – The proposal of this combination is on the basis of the same reason as that for

Trechispora dealbata indicated above. The detailed description of T. scabra can be found in

Furtado et al. (2021).

Trechispora sinensis S.L. Liu, L.W. Zhou & S.H. He, sp. nov. Figs 39–41

Index Fungorum number: IF559898; Facesoffungi number: FoF12878

Etymology – sinensis (Latin), refers to China.

Diagnosis – Differs from Trechispora subsinensis (described below) in verrucose

basidiospores.

Figure 39 – Basidiomes of Trechispora sinensis (LWZ 20180804-19, holotype). Scale bars: a = 1

cm; b = 2 mm.

925

Typus – CHINA, Beijing, Mentougou, Donglingshan Scenic Spot, on fallen angiosperm

branch, 4 Aug. 2018, L.W. Zhou, LWZ 20180804-19 (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft, fragile, easily separated from

substrates, up to 12 cm long, 5 cm wide. Hymenophore odontioid with numerous small aculei,

cream to straw-yellow when fresh, cinnamon-buff when dry. Margin white, fimbriate, up to 3 mm

wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin or thick-walled, moderately branched and septate, interwoven, 2–3.5 µm in diam,

ampullate septa up to 5 μm wide. Aculei composed of a central core of compact hyphae and

subhymenial and hymenial layers; generative hyphae distinct, hyaline, thin or thick-walled,

moderately branched, smooth, subparallel to interwoven, 2–3.5 μm in diam. Crystals abundant,

bipyramidic, aggregated. Cystidia absent. Basidia cylindrical with a slight median constriction,

thin-walled, with four sterigmata and a basal clamp connection, 11–15 × 3.8–5 μm; basidioles

similar in shape to basidia, but smaller. Basidiospores broadly ellipsoid, hyaline, thin-walled,

verrucose, inamyloid, indextrinoid, acyanophilous, 2.8–3.3 × (2.2–)2.5–2.9 μm, L = 3 μm, W = 2.7

μm, Q = 1.1–1.2 (n = 90/3).

Other specimens (paratypes) examined – CHINA, Beijing, Mentougou, Donglingshan Scenic

Spot, on fallen angiosperm branch, 4 Aug. 2018, L.W. Zhou, LWZ 20180804-20 (HMAS); on

fallen angiosperm twig, 9 Sept. 2017, L.W. Zhou, LWZ 20170909-11 (HMAS). Chongqing,

Simianshan National Scenic Spot, on fallen angiosperm branch, 8 July 2018, S.H. He, He 5491

(BJFC 026552). Hubei, Wufeng County, Houhe National Nature Reserve, on fallen angiosperm

branch, 16 Aug. 2017, L.W. Zhou, LWZ 20170816-35 (HMAS). Hunan, Sangzhi County,

Badagongshan National Nature Reserve, on fallen angiosperm branch, 17 Aug. 2017, L.W. Zhou,

LWZ 20170817-5 (HMAS).

Figure 40 – Microscopic structures of Trechispora sinensis (drawn from the holotype). a Vertical

section of basidiomes. b Basidiospores. c Basidia. Scale bars = 10 μm.

926

Figure 41 – Scanning electron micrograph of basidiospores of Trechispora sinensis (scanned from

the holotype). Scale bar = 3 μm.

Notes – Trechispora sinensis has a wide distribution throughout China, including in multiple

provincial regions, like Beijing, Chongqing, Guangdong, Guangxi, Guizhou, Hubei, Hunan,

Jiangsu, Jiangxi, Jilin and Liaoning. Trechispora sinensis may be confused with T. chaibuxiensis

and T. subsinensis; however, T. chaibuxiensis differs in the presence of hyphoid cystidia, while T.

subsinensis differs in predominantly thick-walled tramal hyphae and aculeate basidiospores.

Trechispora subfissurata S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 42–44

Index Fungorum number: IF559899; Facesoffungi number: FoF12879

Etymology – subfissurata (Latin), refers to the similarity to Trechispora fissurata.

Diagnosis – Differs from Trechispora fissurata by its thinner basidiomes and not cracked

hymenophore with shorter aculei (Zhao & Zhao 2021).

Typus – CHINA, Hainan, Baisha County, Yinggeling National Nature Reserve, on dead

branch of living angiosperm, 9 June 2016, S.H. He, He 3907 (holotype in BJFC 022409).

Description – Basidiomes annual, resupinate, effused, thin, soft, fragile, easily separated from

substrates, up to 8 cm long, 3 cm wide, 0.5 mm thick. Hymenophore odontioid to hydnoid with

numerous aculei up to 0.4 mm long, white to cream when fresh, cream to buff-yellow when dry.

Margin white to cream, fimbriate, up to 2 mm wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular generative

hyphae hyaline, thick-walled, moderately branched and septate, subparallel to interwoven, 2.5–4.5

µm in diam, ampullate septa up to 6 µm wide. Aculei composed of a central core of compact

hyphae and subhymenial and hymenial layers; tramal generative hyphae distinct, hyaline, thick-

walled, moderately branched, smooth, subparallel, 2.5–4 μm in diam. Crystals usually present,

bipyramidic, aggregated. Cystidia absent. Basidia cylindrical with a slight median constriction,

hyaline, thin-walled, with four sterigmata and a basal clamp connection, 11–15 × 4–5.5 µm;

basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid, hyaline, thin- to

slightly thick-walled, aculeate with spines that have a sharp apex, inamyloid, indextrinoid,

acyanophilous, 2.8–3.5 × 2.5–3 µm, L = 3.1 µm, W = 2.8 µm, Q = 1.1 (n = 60/2).

Other specimen (paratype) examined – CHINA, Guangdong, Ruyuan County, Nanling

National Forest Park, on the base of living angiosperm, 13 June 2019, L.W. Zhou, LWZ 20190613-

48 (HMAS).

927

Notes – Trechispora subfissurata is characterized by aculeate basidiospores with spines that

have a sharp apex. This character makes T. subfissurata similar to T. echinospora and T. fissurata

(Phookamsak et al. 2019, Zhao & Zhao 2021). However, T. echinospora differs in the absence of

crystals in subiculum and trama, and globose basidiospores (Phookamsak et al. 2019), while T.

fissurata differs in thicker basidiomes (up to 0.8 mm in thickness) with longer aculei (up to 0.9 mm

in length; Zhao & Zhao 2021).

Figure 42 – Basidiomes of Trechispora subfissurata (He 3907, holotype). Scale bars: a = 1 cm, b =

1 mm.

Trechispora subhymenocystis S.L. Liu, H.S. Yuan & L.W. Zhou, sp. nov. Figs 45–47

Index Fungorum number: IF559900; Facesoffungi number: FoF12880

Etymology – subhymenocystis (Latin), refers to the similarity to Trechispora hymenocystis.

Diagnosis – Differs from Trechispora hymenocystis in lack of sphaerocysts in subiculum

(Larsson 1994).

928

Figure 43 – Microscopic structures of Trechispora subfissurata (drawn from the holotype).

a Vertical section of basidiomes. b Basidia. c Basidiospores. Scale bars = 10 μm.

Figure 44 – Scanning electron micrograph of basidiospores of Trechispora subfissurata (scanned

from the holotype). Scale bar = 5 μm.

929

Figure 45 – Basidiomes of Trechispora subhymenocystis (LWZ 20190818-32b, holotype). Scale

bars: a = 1 cm, b = 1 mm.

Typus – CHINA, Sichuan, Meigu County, Dafengding National Nature Reserve, on fallen

trunk of Cryptomeria fortunei, 18 Aug. 2019, L.W. Zhou, LWZ 20190818-32b (holotype in

HMAS).

Description – Basidiomes annual, resupinate, effused, soft, fragile, easily separated from

substrates, soft when fresh, becoming soft corky when dry, up to 6 cm long, 4 cm wide. Pore

surface white when fresh, buff, cinnamon-buff to yellowish brown when dry; pores round to

angular, 3–4 per mm; dissepiments thin, entire. Subiculum white, soft corky, thin, about 0.1 mm

thick. Tubes cinnamon-buff to yellowish brown, soft, up to 1.5 mm long. Margin white, fimbriate,

rhizomorphic.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular generative

hyphae hyaline, thin-walled, moderately branched and septate, subparallel to interwoven, 2.5–5 µm

in diam, ampullate septa up to 7 µm wide; crystals abundant, irregular. Generative hyphae in tubes

hyaline, thin-walled, moderately branched and septate, more or less parallel, 2.5–4 µm in diam;

930

crystals rare, irregular. Cystidia absent. Basidia cylindrical with a slight median constriction,

hyaline, thin-walled, with four sterigmata and a basal clamp connection, 13–22 × 5–7 µm;

basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid, hyaline to

yellowish, slightly thick-walled, aculeate with tubercles on spines, inamyloid, indextrinoid,

acyanophilous, (3.5–)3.8–4.5(–4.8) × (2.8–)3–3.5(–3.9) µm, L = 4.1 µm, W = 3.1 µm, Q = 1.3 (n =

60/2).

Other specimens (paratypes) examined – CHINA, Sichuan, Meigu County, Dafengding

National Nature Reserve, on fallen trunk of Cryptomeria fortunei, 18 Aug. 2019, L.W. Zhou, LWZ

20190818-29b (HMAS). Yunnan, Xianggelila, Shuoduhu, on fallen trunk of Abies, 31 Aug. 2006,

H.S. Yuan, Yuan 2027 (IFP 007168); Xianggelila, Qianhushan, on fallen trunk of Abies, 1 Sept.

2006, H.S. Yuan, Yuan 2089 (IFP 007171).

Figure 46 – Microscopic structures of Trechispora subhymenocystis (drawn from the holotype).

a Vertical section through basidiomes showing position of b and c. b Section through a dissepiment

edge. c Section at the apex of a dissepiment. d Basidia and basidioles. e Basidiospores. Scale bars =

10 μm.

Notes – The tuberculate ornamentations on spines of basidiospores, which were first reported

in T. hymenocystis (Larsson 1994), make Trechispora subhymenocystis similar to T. araneosa, T.

931

hondurensis, T. hymenocystis and T. minima. However, T. araneosa and T. minima differ in non-

poroid hymenophores (Larsson 1995, 1996), T. hondurensis differs in smaller basidiospores (3.67–

3.84 × 2.76–2.89 µm; Haelewaters et al. 2020), and T. hymenocystis differs in the presence of

sphaerocysts in cords and the adjacent part of subiculum and larger basidiospores (4.5–5.5 × 3.5–

4.5 µm; Larsson 1994). Trechispora subhymenocystis also resembles T. mollusca by poroid

hymenophore, a monomitic hyphal system, and ellipsoid, aculeate basidiospores; however, T.

mollusca differs in white to light ochraceous hymenophore, slightly thick-walled subicular hyphae

and shorter basidiospores (2.5–4 µm in length; Liberta 1973, Larsson 1994, Bernicchia & Gorjón

2010).

Figure 47 – Scanning electron micrograph of basidiospores of Trechispora subhymenocystis

(scanned from the holotype). Scale bar = 2 μm.

Trechispora subsinensis S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 48–50

Index Fungorum number: IF559901; Facesoffungi number: FoF12881

Etymology – subsinensis (Latin), refers to the similarity to Trechispora sinensis.

Diagnosis – Differs from Trechispora sinensis in aculeate basidiospores.

Typus – CHINA, Guangdong, Guangzhou, Baiyunshan National Scenic Spot, on fallen

angiosperm branch, 11 June 2019, L.W. Zhou, LWZ 20190611-9 (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft, fragile, easily separated from

substrates, up to 9 cm long, 3 cm wide. Hymenophore odontioid with numerous small aculei, cream

when fresh, straw-yellow to bluish grey when dry. Margin white, fimbriate, up to 0.5 mm wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin to thick-walled, moderately branched and septate, interwoven, 2–3.5 µm in diam,

ampullate septa up to 5 μm wide. Aculei composed of a central core of compact hyphae and

subhymenial and hymenial layers; tramal generative hyphae distinct, hyaline, thick-walled,

moderately branched, smooth, subparallel to interwoven, 2–3.5 μm in diam. Crystals usually

present, bipyramidic, aggregated. Cystidia absent. Basidia cylindrical with a slight median

constriction, thin-walled, with four sterigmata and a basal clamp connection, 11–15 × 3.8–5 μm;

basidioles similar in shape to basidia, but smaller. Basidiospores ellipsoid, hyaline, thin-walled,

aculeate, inamyloid, indextrinoid, acyanophilous, (2.5–)2.7–3.5(–4) × (2–)2.3–2.8(–3) μm, L = 3

μm, W = 2.5 μm, Q = 1.2 (n = 90/3).

932

Other specimens (paratypes) examined – CHINA, Guangdong, Guangzhou, Baiyunshan

National Scenic Spot, on dead branch of living angiosperm, 11 June 2019, L.W. Zhou, LWZ

20190611-19 (HMAS); Zhaoqing, Dinghushan National Nature Reserve, on fallen angiosperm

trunk, 10 June 2019, S.H. He, He 5894 (BJFC 030763). THAILAND, Chiang Mai, Mork Fa

Waterfall, on dead branch of living angiosperm, 25 July 2016, S.H. He, He 4122 (BJFC 023564),

on rotten bamboo, 25 July 2016, S.H. He, He 4125 (BJFC 023567).

Notes – Trechispora subsinensis morphologically resembles T. chaibuxiensis, T. fimbriata, T.

nivea and T. sinensis (Larsson 1995, Zhao & Zhao 2021), and phylogenetically these five species

also nested within a strongly supported clade (BS = 97%, BPP = 1; Fig. 6). Trechispora

chaibuxiensis differs in the presence of hyphoid cystidia, T. fimbriata in cracked hymenial surface

and longer aculei (0.5–0.9 mm; Zhao & Zhao 2021), T. nivea in slightly thick-walled tramal hyphae

and longer aculei (up to 1 mm; Larsson 1995), and T. sinensis in verrucose basidiospores.

Figure 48 – Basidiomes of Trechispora subsinensis (LWZ 20190611-9, holotype). Scale bars: a =

1 cm; b = 1 mm.

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Figure 49 – Microscopic structures of Trechispora subsinensis (drawn from the holotype).

a Vertical section of basidiomes. b Hyphae in subiculum. c Basidiospores. Scale bars = 10 μm.

Figure 50 – Scanning electron micrograph of basidiospores of Trechispora subsinensis (scanned

from the holotype). Scale bar = 2 μm.

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Trechispora taiwanensis S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 51–53

Index Fungorum number: IF559902; Facesoffungi number: FoF12882

Etymology – taiwanensis (Latin), refers to Taiwan, China.

Diagnosis – Differs from Trechispora thailandica (described below) in narrower

basidiospores.

Typus – CHINA, Taiwan, Nantou County, Lianhuachi Nature Reserve, on dead bamboo, 6

Dec. 2016, S.H. He, He 4571 (holotype in BJFC 024012).

Description – Basidiomes annual, resupinate, effused, thin, soft, fragile, loosely attached to

the substrates, up to 10 cm long, 5 cm wide. Hymenophore smooth to grandinioid with numerous

small aculei, farinaceous, cream when fresh, cream to straw-yellow with age, finely cracked with

age. Margin thinning out as byssoid, 2–3 mm wide.

Figure 51 – Basidiomes of Trechispora taiwanensis (He 4571, holotype). Scale bars: a = 1 cm, b =

1 mm.

935

Figure 52 – Microscopic structures of Trechispora taiwanensis (drawn from the holotype). a

Vertical section through basidiomes showing position of b and c. b Section through base of a spine.

c Section through apex of a spine. d Basidia. e Basidiospores. Scale bars = 10 μm.

Figure 53 – Scanning electron micrograph of basidiospores of Trechispora taiwanensis (scanned

from the holotype). Scale bar = 2 μm.

936

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

long-celled, hyaline, thin-walled, frequently branched and septate, subparallel to interwoven, 2.5–

4.5 µm in diam, ampullate septa up to 6 µm wide. Aculei composed of a central core of compact

hyphae and subhymenial and hymenial layers; generative hyphae distinct, hyaline, thin or thick-

walled, moderately branched, smooth, subparallel, 2.5–5.5 μm in diam. Crystals present,

bipyramidic, aggregated. Cystidia absent. Basidia cylindrical with a slight median constriction,

hyaline, thin-walled, with four sterigmata and a basal clamp connection, 13–16 × 4–5.5 µm;

basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid, hyaline, thin-

walled, aculeate, inamyloid, indextrinoid, acyanophilous, sometimes with one oil drop in the

protoplasm, 3–4(–4.2) × 2–2.8(–3.2) µm, L = 3.3 µm, W = 2.5 µm, Q = 1.2–1.5 (n = 60/2).

Other specimen (paratype) examined – CHINA, Taiwan, Nantou County, Lianhuachi Nature

Reserve, on dead bamboo, 6 Dec. 2016, S.H. He, He 4574 (BJFC 024015).

Notes – Macromorphologically, T. taiwanensis resembles T. laevis, but T. laevis differs in

straight or concave basidiospores at the ventral side (Larsson 1996). In addition, T. taiwanensis is

so far only known on bamboo in subtropical Asia, whereas T. laevis grows on coniferous wood in

North Europe (Larsson 1996).

Trechispora thailandica S.L. Liu, S.H. He & L.W. Zhou, sp. nov. Figs 54–56

Index Fungorum number: IF559903; Facesoffungi number: FoF12883

Etymology – thailandica (Latin), refers to Thailand.

Diagnosis – Differs from Trechispora taiwanensis in wider basidiospores.

Typus – THAILAND, Chiang Mai, Doi Saket, on rotten bamboo, 24 July 2016, S.H. He, He

4101 (holotype in BJFC 023542).

Description – Basidiomes annual, resupinate, effused, thin, soft, easily separated from

substrates, up to 9 cm long, 3 cm wide. Hymenophore grandinioid with round and obtuse aculei,

white to cream when fresh, cream to buff-yellow with age, finely cracked with age. Margin

thinning out as byssoid.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin-walled, frequently branched and septate, subparallel, 2.5–5 µm in diam. Tramal

generative hyphae distinct, hyaline, thin-walled, moderately branched, smooth, interwoven, 3–5 μm

in diam. Crystals usually present, bipyramidic, aggregated. Cystidia absent. Basidia cylindrical

with a slight median constriction, hyaline, thin-walled, with four sterigmata and a basal clamp

connection, 11–15 × 4–5.5 µm; basidioles in shape similar to basidia, but slightly smaller.

Basidiospores ellipsoid, hyaline to yellowish, thin-walled, aculeate, inamyloid, indextrinoid,

acyanophilous, (3.5–)3.8–4.3(–4.5) × (2.5–)2.8–3.5 µm, L = 4 µm, W = 3 µm, Q = 1.3 (n = 60/2).

Other specimens (paratypes) examined – THAILAND, Chiang Mai, Doi Saket, on rotten

bamboo, 24 July 2016, S.H. He, He 4112 (BJFC 023554), He 4114 (BJFC 023556).

Notes – Trechispora thailandica resembles T. cyatheae by the white to cream, grandinioid

hymenophore and ellipsoid basidiospores; however, T. cyatheae has smaller basidiospores (3–3.5 ×

2–3 µm including spines) and grows exclusively on Cyathea glauca, an endemic species of tree

fern to La Réunion, France (Ordynets et al. 2015).

Trechispora tropica S.L. Liu & L.W. Zhou, sp. nov. Figs 57–59

Index Fungorum number: IF559904; Facesoffungi number: FoF12884

Etymology – tropica (Latin), refers to tropics.

Diagnosis – Characterized by the absence of crystals in trama and subhymenium, and

cystidium-like hyphal ends at the apex of aculei.

Typus – CHINA, Hainan, Ledong County, Jianfengling National Forest Park, on fallen

angiosperm branch, 13 June 2017, L.W. Zhou, LWZ 20170613-14 (holotype in HMAS).

Description – Basidiomes annual, resupinate, effused, thin, soft and fragile, easily separated

from substrates, up to 11 cm long, 5 cm wide. Hymenophore grandinioid to odontioid with

937

numerous, small aculei, white to cream when fresh, cream to straw-yellow when dry. Margin white,

fimbriate, up to 2 mm wide.

Hyphal system monomitic; generative hyphae with clamp connections. Subicular hyphae

hyaline, thin-walled, frequently branched and septate, subparallel, 2.5–6 µm in diam, ampullate

septa up to 8 μm wide. Aculei composed of a central core of compact hyphae and subhymenial and

hymenial layers; generative hyphae distinct, hyaline, thin or thick-walled, frequently branched,

smooth, subparallel, 2.5–5.5 μm in diam; hyphal ends present at apex, cystidium-like, long-celled,

2.5–4 μm in diam. Crystals only present in subiculum, usually flat and basically rhomboidal.

Cystidia absent. Basidia cylindrical with a slight median constriction, hyaline, thin-walled, with

four sterigmata and a basal clamp connection, 7–10 × 4–5 µm; basidioles in shape similar to

basidia, but slightly smaller. Basidiospores ellipsoid, hyaline, thin-walled, aculeate, inamyloid,

indextrinoid, acyanophilous, 2.5–3 × 2.2–2.5(–2.6) µm, L = 2.8 µm, W = 2.3 µm, Q = 1.2 (n =

60/2).

Figure 54 – Basidiomes of Trechispora thailandica (He 4101, holotype). Scale bars: a = 1 cm, b =

0.5 mm.

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Figure 55 – Microscopic structures of Trechispora thailandica (drawn from the holotype).

a Vertical section through basidiomes showing position of b. b Section through apex of a spine.

c Basidia. d Basidiospores. Scale bars = 10 μm.

Figure 56 – Scanning electron micrograph of basidiospores of Trechispora thailandica (scanned

from the holotype). Scale bar = 2 μm.

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Figure 57 – Basidiomes of Trechispora tropica (LWZ 20170613-14, holotype). Scale bars: a = 1

cm; b = 1 mm.

Other specimens (paratypes) examined – CHINA, Hainan, Ledong County, Jianfengling

National Forest Park, on fallen angiosperm branch, 13 June 2017, L.W. Zhou, LWZ 20170613-16

(HMAS). VIETNAM, Da Lat, Bidoup Nui Ba National Park, on fallen angiosperm branch, 15 Oct.

2017, L.W. Zhou, LWZ 20171015-22 (HMAS).

Notes – Trechispora tropica is phylogenetically close to T. laevis, T. stevensonii and T.

taiwanensis (Fig. 6). However, T. laevis differs in smooth hymenophore (Larsson 1996), T.

stevensonii in the presence of arthroconidia (Larsson 1995), and T. taiwanensis in longer

basidiospores (3–4 µm in length) sometimes with one oil drop in the protoplasm and the growth on

bamboo.

Tubulicium Oberw., Sydowia 19(1-3): 53 (1966) [published June 1965].

Type species – Tubulicium vermiferum (Bourdot) Oberw. ex Jülich, Persoonia 10(3): 335

(1979).

= Tubulixenasma Parmasto, Izv. Akad. Nauk Estonsk. SSR, Ser. Biol. 14: 231 (1965)

[published July 1965]. Type species – Tubulixenasma vermiferum (Bourdot) Parmasto, Eesti NSV

Tead. Akad. Toim., Biol. seer 14(2): 231 (1965).

940

Description – Basidiomes annual, resupinate, effused, thin, closely adnate. Hymenophore

smooth, more or less arachnoid, white, cream to buff. Hyphal system monomitic, generative hyphae

with clamp connections, hyaline, thin- to slightly thick-walled. Cystidia (lyocystidia) conical,

subulate, projecting beyond hymenium, multi-rooted, hyaline, distinctly thick-walled, slightly

amyloid, covered with dendroid branching hyphae. Basidia suburniform, hyaline, thin-walled, with

four sterigmata and a basal clamp connection. Basidiospores navicular to sigmoid, hyaline, thin-

walled, smooth, inamyloid, indextrinoid, acyanophilous. On wood.

Figure 58 Microscopic structures of Trechispora tropica (drawn from the holotype). a Vertical

section of basidiomes b Basidia. c Basidiospores. Scale bars = 10 μm.

Notes – Tubulicium was erected for two species, viz. Hypochnus dussii and Peniophora

vermifera (Oberwinkler 1965). However, the combinations of these two species to Tubulicium by

Oberwinkler (1965) were invalid according to the International Code of Botanical Nomenclature

(Montreal Code published in 1961). Later, Jülich (1979) validated these two combinations.

Tubulicium was formerly put in the family Tubulicrinaceae (Jülich 1981), but later molecular

evidence indicated that it belonged in the trechisporoid clade (Larsson et al. 2004). Therefore,

Larsson (2007) formally accepted it as a member of Hydnodontaceae, Trechisporales, although

there was lack of clear affinity to other genera within Trechisporales in morphology. Recently, two

new species of Tubulicium were almost simultaneously described on the basis of morphological and

phylogenetic evidence (Liu et al. 2019, Ushijima et al. 2019). Besides bringing the species number

of Tubulicium to 11, these two studies also confirmed the taxonomic placement of Tubulicium

proposed by Larsson (2007), which is also recovered by the current phylogenies (Figs 1, 2, 3).

Morphologically, Tubulicium is characterized by the presence of multirooted, subulate and thick-

walled cystidia covered by dendroid-branching hyphae (Liu et al. 2019, Ushijima et al. 2019).

941

Tubulixenasma Parmasto is an obligate synonym of Tubulicium, because it is based on the same

type species, Peniophora vermifera. Tubulicium and Tubulixenasma were published in the same

year, but according to the title page of Sydowia volume 19, the paper on “Primitive

Basidiomyceten” by Oberwinkler “wurde als Sonderdruck im Juni 1965 ausgegeben [was issued as

an offprint in June 1965]” while Parmasto (1965) appeared in July of that year, according to

Hjortstam et al. (1988).

Figure 59 Scanning electron micrograph of basidiospores of Trechispora tropica (scanned from the

holotype). Scale bar = 4 μm.

Genera excluded from Trechisporales

Boidinella Nakasone, Cryptog. Mycol. 32(2): 192 (2011).

Type species – Boidinella globulispora (Boidin & Lanq.) Nakasone, Cryptog. Mycol. 32(2):

193 (2011).

Notes – Boidinella was introduced for B. cystidiolophora (originally described in

Sistotremella) and the type species B. globulispora (originally described in Dendrothele) by

Nakasone (2011). The genus was characterized by the combination of “effuse, soft, densely

farinaceous or membranous basidioma, urniform basidia with 4-sterigmata, obclavate lcptocystidia,

dendrohyphidia, and basidiospores with smooth, slightly thickened. cyanophilous walls”. There are

no sequences for either species currently in GenBank. Nevertheless, Boidinella was placed in

Hydnodontaceae, Trechisporales by Kirk (2019) who does not indicate the source of this

placement. Until molecular data are available, the genus is best treated as a member of the

Cantharellales as indicated by the original author according to morphological characters (Nakasone

2011), which are distinct from members in Trechisporales.

Litschauerella Oberw., Sydowia 19(1-3): 43 (1966) [1965].

Type species – Litschauerella abietis (Bourdot & Galzin) Oberw. ex Jülich, Persoonia 10(3):

335 (1979).

Notes – Litschauerella is a small and not well-known genus. Larsson (2007) place this genus

in Trechisporales with doubt. Two authors of the current paper, viz. S.L. Liu and S.H. He, and their

colleagues for the first time generated molecular sequences from a species in this genus, but they

did not mention its taxonomic position (Liu et al. 2019). Here, via BLAST search of ITS

942

(MK204555) and nrLSU (MK204556) regions from Litschauerella gladiola represented by the

specimen He 3171 (BJFC 021566) from Yunnan, China, we confirmed that this species has an

affinity to Hymenochaetales rather than Trechisporales. Although L. gladiola is not the generic

type, its placement based on molecular evidence does provide more doubts about the position of

Litschauerella within Trechisporales. Given above, no evidence supports the taxonomic position of

Litschauerella in Trechisporales and thus we tentatively exclude it from Trechisporales. We note

that the family Litschauerellaceae (Jülich 1981) is based on Litschauerella, but even if

Litschauerella should prove to belong in Hydnodontaceae, this latter family name could stand as

the two families were introduced in the same publication, and Hydnodontaceae has already been

established as the preferred choice over Subulicystidiaceae and, if necessary, a similar choice could

be made against Litschauerellaceae.

Sphaerobasidium Oberw., Sydowia 19(1-3): 57 (1966) [1965].

Type species – Sphaerobasidium minutum (J. Erikss.) Oberw. ex Jülich, Persoonia 10(3): 335

(1979).

Notes – For unexplained reasons, He et al. (2019) and Kirk (2019) listed Sphaerobasidium in

Hydnodontaceae despite the fact that molecular data on the type species of this genus indicate a

placement in the vicinity of Tubulicrinis in the Hymenochaetales (Larsson et al. 2006, Larsson

2007). BLAST search with all three available sequences of Sphaerobasidium minutum (GenBank

accession numbers: AJ406446, DQ873652 and DQ873653) also suggests its affinity to

Hymenochaetales rather than Trechisporales.

Tomentella P. Karst., Bidr. Känn. Finl. Nat. Folk 48: 419 (1889) non Tomentella Pers. ex Pat.,

Hyménomyc. Eur. (Paris): 154 (1887).

Type species – Tomentella sulphurea (Pers.) P. Karst., Bidr. Känn. Finl. Nat. Folk 48: 419

(1889).

Notes – The type of the illegitimate name Tomentella P. Karst. was placed at one time in

Trechispora as Trechispora sulphurea, but was subsequently accepted as a species of Phlebiella as

P. sulphurea and considered synonymous with Trechispora vaga, such as by Ginns & Lefebvre

(1993). While Tomentella sulphurea (based on Corticium sulphureum) and Trechispora vaga

(based on Phlebia vaga) are both sanctioned, the former has the earlier basionym as is the name

that should be taken up when the two names are treated as referring to the same species. Larsson

(2007) placed Phlebiella outside of Hydnodontaceae, in the “Phlebiella family”. Piątek (2005)

pointed out that Phlebiella was not validly published and should be known as Xenasmatella. While

the name Xenasmatella vaga exists for another species, no combination has been made yet in

Xenasmatella for T. sulphurea. Despite the unresolved issues around the correct name for

Tomentella sulphurea, it is clear that Tomentella P. Karst. does not belong in Hydnodontaceae,

despite the placement in that family by Index Fungorum (http://www.indexfungorum.org/).

Genus of uncertain position

Murrilloporus Ryvarden, Mycotaxon 23: 192 (1985).

Type species – Murrilloporus rutilantiformis (Murrill) Ryvarden, Mycotaxon 23: 192 (1985).

Notes – Trametes rutilantiformis, type of the monotypic genus Murrilloporus, was placed by

Stalpers (1996) in Heterobasidion, at that time making Murrilloporus a synonym of

Heterobasidion. Later, Hattori (2003) synonymized Loweporus corticicola with Trametes

rutilantiformis and placed the latter in Cristelloporia, making Murrilloporus a synonym of

Cristelloporia. In this case, given that Cristelloporia is now a synonym of Trechispora,

Murrilloporus would also be placed under Trechispora. However, T. rutilantiformis differs from

Trechispora in the combination of coriaceous basidiomes with context up to 1 cm thick and

dextrinoid skeletal hyphae. Given this morphological divergence from Trechispora, sequencing is

required to confirm the appropriate placement of Trametes rutilantiformis.

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Species excluded from Trechispora

Trechispora yunnanensis C.L. Zhao, in Xu, Chen & Zhao, Phytotaxa 424(4): 256. (2019).

Notes – Trechispora yunnanensis was recently described from Yunnan, China (Xu et al.

2019). Chikowski et al. (2020) noted that the deposited molecular sequences of this species are

questionable according to BLAST search. Although the morphological characters of T. yunnanensis

partially fit the concept of Trechispora, we confirmed that the ITS and nrLSU regions of this

species (put in reverse order in the columns in Table 1 of the original publication) actually

represent two taxa, respectively, from the orders Trechisporales and Hymenochaetales instead of

being a single taxon. Therefore, the identity of type specimens of T. yunnanensis needs to be further

clarified. For now, we tentatively exclude this species from Trechispora, Trechisporales.

Key to 12 genera in Trechisporales

1. Basidiomes clavarioid or pileate-stipitate, on ground or termite mounds..................Trechispora A

1. Basidiomes resupinate, on wood.....................................................................................................2

2. Hymenophore poroid.......................................................................................................................3

2. Hymenophore non-poroid................................................................................................................4

3. Basidiospores smooth.....................................................................................................Porpomyces

3. Basidiospores ornamented..........................................................................................Trechispora B

4. Hymenophore brown..............................................................................................................Luellia

4. Hymenophore usually light colored.................................................................................................5

5. Cystidia present, distinct..................................................................................................................6

5. Cystidia absent or indistinct.............................................................................................................9

6. Cystidia distinctly dextrinoid......................................................................................Dextrinocystis

6. Cystidia amyloid or negative in Melzer’s reagent...........................................................................7

7. Cystidia neither bi- nor multi-rooted, multiseptated...................................................Suillosporium

7. Cystidia bi- or multi-rooted, not multiseptated................................................................................8

8. Cystidia regularly encrusted with rectangular crystals.............................................Subulicystidium

8. Cystidia usually covered with dendroid hyphae..............................................................Tubulicium

9. Generative hyphae with ampullate septa....................................................................Trechispora C

9. Generative hyphae without ampullate septa..................................................................................10

10. Subhymenial hyphae isodiametric...............................................................................................11

10. Subhymenial hyphae not isodiametric.........................................................................................12

11. Hymenophore firmly granular or almost smooth; basidiospores rhomboid or short ellipsoid........

...................................................................................................................................Brevicellopsis

11. Hymenophore distinctly odontioid; basidiospores narrowly allantoid ....................Brevicellicium

12. Hyphal system dimitic..................................................................................................Fibrodontia

12. Hyphal system monomitic...........................................................................................................13

13. Sterile hyphal pegs present.........................................................................................Pteridomyces

13. Sterile hyphal pegs absent.......................................................................................Allotrechispora

Key to 87 species in Trechispora (separated into three parts following the key to 12 genera in

Trechisporales)

Trechispora A (Basidiomes clavarioid or pileate-stipitate, on ground or termite mounds)

1. Hymenophore pileate-stipitate.........................................................................................................2

1. Hymenophore clavarioid..................................................................................................................4

2. Aculei up to 0.4 mm long; basidiospores < 3.5 μm wide.............................................T. hypogeton

2. Aculei up to 1 mm long; basidiospores > 3.5 µm wide...................................................................3

3. The stipe and abhymenial surface white to sordid cream...................................................T. gillesii

3. The stipe and abhymenial surface light yellow brown.................................................T. thelephora

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4. Hymenophore on termite mounds..............................................................................T. termitophila

4. Hymenophore on ground.................................................................................................................5

5. Hymenophore more or less minutely papillate or hydnoid..............................................................6

5. Hymenophore smooth .....................................................................................................................7

6. Hymenophore white............................................................................................................T. scabra

6. Hymenophore pale orange to reddish.............................................................................T. papillosa

7. Hymenophore light brown to reddish brown when fresh................................................................8

7. Hymenophore pure white, greyish yellow to beige when fresh....................................................10

8. Inflated hyphae absent in subiculum, basidia with two sterigmata...........................T. havencampii

8. Inflated hyphae present in subiculum, basidia with four sterigmata...............................................9

9. Subicular hyphae 3.5–8 µm in diam.................................................................................T. foetidus

9. Subicular hyphae inflated to 6–23 µm in diam.................................................................T. robusta

10. Hymenia with cystidial structures................................................................................................11

10. Hymenia without cystidial structures...........................................................................................13

11. Cystidia clavate...........................................................................................................T. minispora

11. Cystidia lanceolate, narrowly utriform or capitate......................................................................12

12. Basidiospores subglobose, Q = 1.06.................................................................T. caulocystidiatus

12. Basidiospores ellipsoid, Q = 1.29...............................................................................T. gelatinosa

13. Inflated hyphae present in subiculum..........................................................................................14

13. Inflated hyphae absent in subiculum............................................................................................16

14. Branches U-shaped, cream to buff turning yellowish brown towards the apex......T. longiramosa

14. Branches V-shaped, pale greyish yellow to beige.......................................................................15

15. Basidiomes pale yellow when dry..................................................................................T. copiosa

15. Basidiomes reddish brown when dry............................................................................T. dealbata

16. Hymenia unilateral......................................................................................................T. chartacea

16. Hymenia amphigenous................................................................................................T. pallescens

Trechispora B (Basidiomes resupinate, on wood, hymenophore poroid)

1. Hyphal system dimitic.....................................................................................................................2

1. Hyphal system monomitic...............................................................................................................3

2. Basidiospores ventrally concave.................................................................................T. brasiliensis

2. Basidiospores ventrally convex......................................................................................T. dimitiella

3. Basidiospores subglobose to subangular, sparsely verrucose................................T. polygonospora

3. Basidiospores subglobose to broadly ellipsoidal, densely aculeate.................................................4

4. Cystidial structures present.............................................................................................T. regularis

4. Cystidial structures absent...............................................................................................................5

5. Basidiospores including spines > 6 µm long...............................................................T. clancularis

5. Basidiospores including spines < 6 µm long...................................................................................6

6. Subicular hyphae slightly thick-walled, up to 3 µm wide...............................................................7

6. Subicular hyphae thin-walled, up to 6 µm wide..............................................................................8

7. Ampullate septa present on subicular hyphae.................................................................T. mollusca

7. Ampullate septa absent on subicular hyphae..................................................................T. suberosa

8. Crystals in subiculum as numerous rodlets..............................................................T. candidissima

8. Crystals in subiculum as rhomboidal plates or various shapes........................................................9

9. Sphaerocysts present in cords and the adjacent part of subiculum..........................T. hymenocystis

9. Sphaerocysts absent.......................................................................................................................10

10. Basidiospores < 3.8 µm long, < 3 µm wide.............................................................T. hondurensis

10. Basidiospores > 3.8 µm long, > 3 µm wide......................................................T. subhymenocystis

Trechispora C (Basidiomes resupinate, on wood, hymenophore non-poroid)

1. Basidiospores smooth......................................................................................................................2

1. Basidiospores ornamented.............................................................................................................12

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2. Hyphal system dimitic.....................................................................................................................3

2. Hyphal system monomitic...............................................................................................................4

3. Hymenophore hydnoid................................................................................................T. molliuscula

3. Hymenophore smooth..................................................................................................T. silvae-ryae

4. Hymenophore hydnoid................................................................................................T. kavinioides

4. Hymenophore smooth to farinaceous..............................................................................................5

5. Basidiospores subglobose, angular to turbinate...............................................................................6

5. Basidiospores ellipsoid to lacrymoid...............................................................................................8

6. Hymenophore pellicular, smooth.......................................................................................T. mellina

6. Hymenophore more or less adnate, farinaceous..............................................................................7

7. Basidiospores turbinate.....................................................................................T. subsphaerospora

7. Hymenophore subglobose................................................................................................T. confinis

8. Crystals bipyramid, aggregate.......................................................................................T. byssinella

8. Crystals differently shaped..............................................................................................................9

9. Basidiospores > 4 µm long..........................................................................................T. amianthina

9. Basidiospores < 4 µm long............................................................................................................10

10. Crystals as rodlets with incised ends..........................................................................T. cohaerens

10. Crystals otherwise........................................................................................................................11

11. Basidiospores thick-walled.............................................................................................T. confinis

11. Basidiospores thin-walled...........................................................................................T. laevispora

12. Hyphal system dimitic.................................................................................................................13

12. Hyphal system monomitic...........................................................................................................17

13. Thick-walled conidia present.......................................................................................................14

13. Thick-walled conidia absent........................................................................................................15

14. Conidia rugose..............................................................................................................T. invisitata

14. Conidia smooth.............................................................................................................T. tenuicula

15. Crystalline sphere covered by numerous needle-like crystals present.............T. echinocristallina

15. Crystalline sphere covered by numerous needle-like crystals absent..........................................16

16. Subicular crystals acicular..............................................................................................T. dimitica

16. Subicular crystals differently shaped...............................................................................T. minuta

17. Basidiomes reflexed.....................................................................................................T. fastidiosa

17. Basidiomes not reflexed...............................................................................................................18

18. Basidiomes < 50 µm thick..............................................................................................T. gracilis

18. Basidiomes > 50 µm thick...........................................................................................................19

19. Hymenophore smooth..................................................................................................................20

19. Hymenophore non-smooth...........................................................................................................35

20. Basidia with two sterigmata.........................................................................................................21

20. Basidia with four sterigmata........................................................................................................22

21. Basidiospores > 4.5 µm long, > 3.5 µm wide.................................................................T. antipus

21. Basidiospores < 4.5 µm long, < 3.5 µm wide.................................................................T. bispora

22. Basidiospores ventrally concave..................................................................................................23

22. Basidiospores ventrally straight or convex..................................................................................25

23. Arthroconidia absent, basidiospores < 4 µm long..............................................................T. laevis

23. Arthroconidia present, basidiospores > 4 µm long......................................................................24

24. Basidiospores including spines mostly > 5 µm long...................................................T. caucasica

24. Basidiospores including spines mostly < 5 µm long.....................................................T. elongata

25. Crystals acicular in subiculum.....................................................................................................26

25. Crystals differently shaped in subiculum.....................................................................................27

26. Basidiospores including spines < 5 µm long, < 4 µm wide, subglobose to.....................................

lacrymoid...................................................................................................................T. microspora

26. Basidiospores including spines > 5 µm long, > 4 µm wide, ellipsoid.......................T. praefocata

27. Basidiospores in shape irregular...............................................................................T. canariensis

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27. Basidiospores in shape regular.....................................................................................................28

28. Basidiospores sparsely ornamented.............................................................................................29

28. Basidiospores densely ornamented..............................................................................................30

29. Basidiospores turbinate, verrucose...................................................................T. subsphaerospora

29. Basidiospores ellipsoid, aculeate..................................................................................T. stellulata

30. Subicular generative hyphae thick-walled...................................................................T. latehypha

30. Subicular generative hyphae thin-walled.....................................................................................31

31. Basidiospores verrucose................................................................................................T. larssonii

31. Basidiospores aculeate.................................................................................................................32

32. Crystals abundant in subhymenium...............................................................................T. cyatheae

32. Crystals occasionally present or absent in subhymenium............................................................33

33. Crystals as single or aggregated rodlets with incised ends..................................................T. incia

33. Crystals as aggregate rhomboidal flakes......................................................................................34

34. Basidia < 5 µm wide, basidiospores subglobose to broadly ellipsoid.............................T. minima

34. Basidia > 5 µm wide, basidiospores ellipsoid...................................................T. damansaraensis

35. Hymenophore colliculose or grandinioid.....................................................................................36

35. Hymenophore odontioid to hydnoid............................................................................................44

36. Hymenophore yellowish to ochraceous; conidia present.............................................................37

36. Hymenophore white to cream; conidia absent.............................................................................38

37. Basidiospores verrucose, ventrally straight or convex...................................................T. alnicola

37. Basidiospores aculeate, ventrally concave..................................................................T. caucasica

38. On bamboo...................................................................................................................................39

38. Not on bamboo.............................................................................................................................40

39. Basidiospores mostly < 2.8 µm wide........................................................................T. taiwanensis

39. Basidiospores mostly > 2.8 µm wide........................................................................T. thailandica

40. Basidiospores ventrally concave........................................................................................T. rigida

40. Basidiospores ventrally straight or convex..................................................................................41

41. Basidiospores globose to subglobose............................................................................T. torrendii

41. Basidiospores ellipsoid to broadly ellipsoid................................................................................42

42. Basidiospores verrucose.............................................................................................T. crystallina

42. Basidiospores aculeate.................................................................................................................43

43. Crystals butterfly-shaped in subiculum........................................................................T. araneosa

43. Crystals as prisms to rhomboidal flakes in subiculum.................................................T. farinacea

44. Tramal hyphae distinctly thick-walled.........................................................................................45

44. Tramal hyphae thin-walled or slightly thick-walled....................................................................52

45. Hymenophore aculei > 0.4 mm long............................................................................................46

45. Hymenophore aculei < 0.4 mm long............................................................................................49

46. Margin undifferentiated.................................................................................................T. fissurata

46. Margin thinning out, fimbriate.....................................................................................................47

47. Basidiomes irpicoid............................................................................................................T. denta

47. Basidiomes typically odontioid or hydnoid.................................................................................48

48. Hymenophore aculei sparse, cream to buff-yellow when fresh....................................T. fimbriata

48. Hymenophore aculei dense, white when fresh.................................................................T. fragilis

49. Basidiospores with sharp spines...............................................................................T. subfissurata

49. Basidiospores without sharp spines.............................................................................................50

50. Basidiospores verrucose.................................................................................................T. sinensis

50. Basidiospores aculeate.................................................................................................................51

51. Basidiospores thick-walled.....................................................................................T. bambusicola

51. Basidiospores thin-walled.........................................................................................T. subsinensis

52. Crystals absent, basidiospores globose....................................................................T. echinospora

52. Crystals present, basidiospores ellipsoid to broadly ellipsoid.....................................................53

53. Arthroconidia present.................................................................................................T. stevensonii

947

53. Arthroconidia absent....................................................................................................................54

54. Crystals absent in trama...............................................................................................................55

54. Crystals present in trama..............................................................................................................56

55. Basidiospores excluding spines < 3.5 µm long, < 3 µm wide.........................................T. tropica

55. Basidiospores excluding spines > 3.5 µm long, > 3 µm wide..................................T. verruculosa

56. Tramal hyphae 3–6 µm wide, spines of basidiospores constricted.............................T. constricta

56. Tramal hyphae 2–4 µm wide, spines of basidiospores not constricted.......................................57

57. Basidia present at the apical ends of aculei, hyphoid cystidia present..................T. chaibuxiensis

57. Basidia absent at the apical ends of aculei, hyphoid cystidia absent...........................................58

58. Tramal hyphae thin-walled, spines on basidiospores > 0.6 µm long...........................T. malayana

58. Tramal hyphae slightly thick-walled, spines on basidiospores < 0.3 µm long...................T. nivea

Discussion

In this study, the relationships among members of Trechisporales within Agaricomycetes are

for the first time explored with the help of the most comprehensive multilocus-based phylogenetic

analyses. In association with morphological characters, a new family Sistotremastraceae within a

new order Sistotremastrales are introduced for Agaricomycetes to accommodate Sertulicium and

Sistotremastrum (type genus) segregated from Trechisporales; a new genus Allotrechispora is

proposed within Hydnodontaceae, Trechisporales; one new species is introduced for each of

Allotrechispora, Fibrodontia and Subulicystidium, and 16 new species for Trechispora are

described; seven new combinations are proposed for Allotrechispora and Trechispora; and

Boidinella, Litschauerella and Sphaerobasidium are excluded from Hydnodontaceae,

Trechisporales, while Trechispora yunnanensis from Trechispora with uncertain taxonomic

position at these ranks. Moreover, a brief introduction to two accepted genera within

Sistotremastrales and all 12 accepted genera within Trechisporales, along with keys to these 12

genera of Trechisporales and to all 87 species of Trechispora are provided. In addition, Tomentella

P. Karst. non-Pers. ex Pat. and Murrilloporus, potential synonyms of Trechispora, are excluded

from Trechisporales and of uncertain position, respectively.

Sistotremastrum, as the type genus of the new family Sistotremastraceae and the new order

Sistotremastrales, was formerly known as the ‘Sistotremastrum family’ within Trechisporales, and

distantly related to other genera belonging to Hydnodontaceae, the single formally named family of

this order (Larsson 2007). Recently, Spirin et al. (2021) segregated an additional genus Sertulicium

from Sistotremastrum, and also suggested that Sertulicium and Sistotremastrum should be

independent at the family level from Hydnodontaceae. However, even if the morphological

characters of Sertulicium and Sistotremastrum are distinct, the dataset of nrLSU region failed to

generate a strongly supported phylogeny (Spirin et al. 2021). The current phylogenies, for the first

time both sampling taxa comprehensively and employing multiloci, recover the clade of

Sertulicium and Sistotremastrum as highly distinct from Hydnodontaceae at the family level and

also from Trechisporales at the order level within Agaricomycetes (Figs 1–3). Although the close

relationship between Sistotremastrales and Trechisporales cannot be rejected, it is assumed that all

nodes in phylogenetic trees will receive strong supports when sampling enough gene regions, like

the phenomena in recent phylogenomic analyses (Nagy et al. 2014, Kiss et al. 2019, Miyauchi et al.

2020, Jiang et al. 2021, Li et al. 2021). In addition, in the current phylogeny (Fig. 1), the clade

being composed of five well-accepted independent orders, viz. Geastrales, Gomphales,

Hysterangiales, Phallales and Stereopsidales, receives almost full statistical support (BS = 98%,

BPP = 1), while Gloeophyllales, Jaapiales, Polyporales and Thelephorales also group together

with strong support (BS = 86%, BPP = 1). Therefore, the phylogenetic affinity between these two

orders is not the obstacle to separating them. Moreover, the divergence time of this clade also fits

well within the range of all known orders in Agaricomycetes (Fig. 2). A recent whole-scale

phylogenomic analysis has proved that the current fungal classification at higher ranks is basically

consistent with the evolutionary divergence in Basidiomycota (Li et al. 2021). That is to say that the

divergence time can be considered to be one of important supports for circumscribing fungal orders

948

as used here. Given above, the segregation of Sistotremastrales from Trechisporales is justified

following the practice of integrated taxonomy.

Within the small family Sistotremastraceae and order Sistotremastrales, only two genera

accommodating 19 species are known to date (Dhingra et al. 2014, Gruhn & Alvarado 2021, Spirin

et al. 2021). Although this study does not describe new species of Sertulicium and Sistotremastrum,

three unnamed single-specimen lineages are recovered in Sistotremastrum, viz. LWZ 20171015-32

collected from Vietnam, LWZ 20191107-25 from Yunnan, China and LWZ 20191207-26 from

Malaysia (Fig. 3). This indicates that more species in Sistotremastraceae, Sistotremastrales await

description. Moreover, the phylogenetic relationships among species of Sistotremastrum indicate

that additional new genera may need to be segregated as was Sertulicium (Fig. 3).

Besides one new species of Subulicystidium, one unnamed single-specimen (LWZ 20180804-

5) lineage and one unnamed double-specimen (LWZ 20170816-7 and LWZ 20190816-24a) lineage

were newly revealed in this genus (Fig. 5). Following a desired taxonomic practice (Aime et al.

2021), the unnamed single-specimen new lineage is better not to be described as a new species until

more related samples can be examined. Regarding the unnamed double-specimen new lineage, it

was phylogenetically related to S. perlongisporum (Fig. 5). However, the phylogenetic identity of

S. perlongisporum is not clear (Fig. 5; Volobuev 2016, Ordynets et al. 2018). More importantly, the

condition of both specimens in this new lineage is not good enough to justly determine the

morphological characters. Therefore, we leave this unnamed double-specimen new lineage open

until more related samples in a good condition can be secured.

With the inclusion of 16 new species, a total of 87 species are accepted in Trechispora. In

addition, 29 unnamed single-specimen new lineages were revealed in this genus (Fig. 6) and their

taxonomic identities await to be further determined with more related samples (Aime et al. 2021).

Consequently, more than one hundred species could be present in Trechispora. Although

Trechispora is strongly supported as a monophyletic genus (Figs 3, 6), it is noticed that nearly one

quarter of known species (21 out of 87) in Trechispora are not subjected to phylogenetic analyses

with molecular data. Taking the segregation of Allotrechispora into consideration, the monophyly

of Trechispora still needs to be tested by sampling the remaining one quarter of known species in

phylogenetic analyses.

As the most species-rich genus in Trechisporales, Trechispora produces highly diverse

morphological characters. The most striking character is the clavarioid basidiomes adopted by

synonymizing Scytinopogon (de Meiras-Ottoni et al. 2021), which was originally put in

Clavariaceae (Corner 1950, García-Sandoval et al. 2005) but later moved to Trechisporales (Jülich

1981, Birkebak et al. 2013). In addition, even for the species of Trechispora with resupinate

basidiomes, their hymenophoral configurations can range from smooth, grandinioid, hydnoid to

poroid. Previous study on Hymenochaetales indicated that grandinioid hymenophoral configuration

represents an adaptive advantage in balancing protection and dispersal of basidiospores (Wang et

al. 2021). Even though of interest, the trait evolution of basidiomes in Trechispora is not yet

explored. Microscopically, as sexual reproductive cells of species in Basidiomycota (Wallen &

Perlin 2018), the morphology of basidiospores is considered to be one of the most important and

distinguishable taxonomic characters. In species of Trechispora, the surface of basidiospores is

morphologically diverse, including contrasts between ornamented vs. not ornamented and

ornamentation-verrucose vs. ornamentation-aculeate. However, the biological and ecological

functions of these diverse characters in the process of sexual reproduction are unknown. Besides

variation in morphology, the nutritional modes of Trechispora may be saprotrophic or biotrophic

(Vanegas-León et al. 2019). The answer to how and why the nutritional modes shift from one mode

to another will help to understand the evolutionary history of Trechispora. In the current phylogeny

of Trechispora (Fig. 6), the interspecific relationships are not well resolved, and moreover the

phylogenetic identity of certain species, like T. nivea and T. pallescens, is not clear. To solve the

above-mentioned series of evolutionary issues in macromorphology, micromorphology and

nutrition mode, a much higher resolution of phylogenetic relationships among species of

Trechispora is needed. Undoubtedly, gene regions additional to ITS and nrLSU should be used to

949

construct an intrageneric phylogeny with reliable statistical supports. Moreover, omics analyses of

certain representative species could help to elucidate the evolutionary dynamics of corresponding

traits, like fungal multicellularity (Kiss et al. 2019) and symbiosis (Miyauchi et al. 2020).

In conclusion, via a wider sampling especially from Asia Pacific, careful morphological

examinations and comprehensive multilocus-based phylogenetic analyses, an emended

classification of Trechisporales within Agaricomycetes is constructed. This classification will help

to further clarify species diversity and explore trait evolution within Trechisporales.

Supplementary material

File S1 Alignment resulted from the dataset (1).

File S2 Alignment resulted from the dataset (2).

File S3 Alignment resulted from the dataset (3).

File S4 Alignment resulted from the dataset (4).

File S5 Alignment resulted from the dataset (5).

Acknowledgements

Prof. Karl-Henrik Larsson (University of Oslo, Norway) is thanked for helpful comments on

an early version of the manuscript. Profs. Yu-Cheng Dai (Beijing Forestry University, China), Ping

Zhang (Hunan Normal University, China) and Yu-Lian Wei (Institute of Applied Ecology, Chinese

Academy of Sciences, China) are thanked for sending specimens on loan. Prof. Yu-Cheng Dai is

also thanked for providing photos of basidiomes in situ. The research was financed by the National

Key Research and Development Program of China (No. 2022YFC2601203), the National Natural

Science Foundation of China (Nos. 31770008, 31870011, 31860007, 31970012 & 32100004), the

Biodiversity Survey and Assessment Project of the Ministry of Ecology and Environment, China

(No. 2019HJ2096001006) and National Science & Technology Fundamental Resources

Investigation Program of China (No. 2019FY101800).

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Taxonomy of Hyphodermella: a case study to show that simple phylogenies cannot always accurately place species in appropriate genera

Article

Full-text available

Jun 2023

The genus is a special and crucial taxonomic rank compared with others above the species level, because a species has to be placed in a certain genus instead of any other higher ranks. With more and more new species being described, the placements of their generic position are sometimes incorrect due to the simple phylogenies resulting from inappropriate sampling. Here, we focus on the taxonomy of a small wood-inhabiting fungal genus Hyphodermella . With the most comprehensive sampling to date, the phylogenetic position of Hyphodermella within Phanerochaetaceae is rearranged by employing the same ITS and nLSU regions as in previous studies and also the ITS, nLSU, rpb1 , rpb2 and tef1α regions. Three species are excluded from Hyphodermella : H. poroides is placed in a newly introduced monotypic genus Pseudohyphodermella , while H. aurantiaca and H. zixishanensis are transferred to Roseograndinia . Hyphodermella suiae is described as a new species from South China and Vietnam. Keys to eight species in Hyphodermella and five in Roseograndinia are provided. Beyond solving the taxonomic issue of Hyphodermella itself, the current study also aims to suggest that all fungal taxonomists especially beginners should keep in mind to sample as many comprehensive taxa as possible in phylogenetic analyses.

Preliminary species diversity and community phylogenetics of wood-inhabiting basidiomycetous fungi in the Dabie Mountains, Central China reveal unexpected richness

Article

Full-text available

Nov 2023

Wood-inhabiting fungi have important economic values as well as playing a major ecological role in forest ecosystem cycles. The Dabie Mountains, at the junction of Henan, Hubei, and Anhui Provinces, Central China, provide an ideal climate and favorable niches for the speciation and diversification of various forms of life including fungi. We studied the species diversity and community phylogenetics of wood-inhabiting basidiomycetous fungi that revealed 175 wood-inhabiting basidiomycetous species, of which 20 represented unidentified species, based on morphological and phylogenetic analyses of 575 specimens collected from ten sampling sites. These species belonged to two classes, 11 orders, 42 families, and 106 genera of Basidiomycota, and included 12 edible species, 28 medicinal species, four poisonous species, and seven forest pathogens. Four types of fungal distribution pattern at the genus level were recognized for 65 genera, while another 41 genera could not be placed in any known distribution pattern. The five sampling sites in the eastern part of the Dabie Mountains had significantly higher species diversity and phylogenetic diversity of wood-inhabiting basidiomycetous fungi than those in the western part, and thus deserve priority in terms of conservation. The community of wood-inhabiting basidiomycetous fungi in the Dabie Mountains is generally affected by a combination of habitat filtering and competitive exclusion. This study provides a basis on which to build actions for the comprehensive recognition, utilization, and conservation of wood-inhabiting basidiomycetous fungi in the region. Supplementary Information The online version contains supplementary material available at 10.1186/s43008-023-00130-9.

A new species, new combination, and a nomen novum in the genus Trechispora from India

Preprint

Full-text available

May 2023

The genus Trechispora accommodate species with corticioid, clavarioid, and polyporoid basidiomata, smooth to verrucose, angular basidiospores, monomitic or dimitic hyphal systems with clamp-connections, and ampullaceous septa. During a study of clavarioid fungi of Kerala, seven Trechispora species with clavarioid basidiomata were collected and studied. Morphological and molecular analyses revealed one species, T. cystidiata , characterized by cystidia, to be new to science. In addition, a new combination ( T. angulispora ) and a nomen novum ( T. corneri ) are also proposed. Descriptions of all the species collected and studied from Kerala, along with their phylogenetic placement in the order Trechisporales is presented. A taxonomic key to the clavarioid species of Trechisporales from India is also provided.

Morphological and molecular identification for four new wood-inhabiting species of Trechispora (Basidiomycota) from China

Article

Full-text available

May 2024

Four new wood-inhabiting fungi, Trechispora albofarinosa, T. bisterigmata, T. pileata and T. wenshanensisspp. nov., are proposed based on a combination of morphological features and molecular evidence. Trechispora albofarinosa is characterized by the farinose basidiomata with flocculence hymenial surface, a monomitic hyphal system with clamped generative hyphae, and ellipsoid, warted basidiospores. Trechispora bisterigmata is characterized by the membranous basidiomata with odontioid hymenial surface, rhizomorphic sterile margin, barrelled basidia and subglobose to broad ellipsoid, smooth basidiospores. Trechispora pileata is characterized by the laterally contracted base, solitary or imbricate basidiomata, fan shaped pileus, radially striate-covered surface with appressed scales, odontioid hymenophore surface, and subglobose to broad ellipsoid, thin-walled, smooth basidiospores. Trechispora wenshanensis is characterized by a cottony basidiomata with a smooth hymenial surface, and ellipsoid, thin-walled, warted basidiospores. Sequences of ITS and LSU marker of the studied samples were generated, and phylogenetic analyses were performed with the maximum likelihood, maximum parsimony, and Bayesian inference methods. The phylogenetic tree inferred from the ITS+nLSU sequences highlighted that four new species were grouped into the genus Trechispora.

Catalogue of fungi in China 1. New taxa of plant-inhabiting fungi

Article

Full-text available

Mar 2024

China has a huge area of diverse landscapes and is believed to conceive incredibly high fungal diversity. To systematically and promptly report Chinese fungal species, we initiate the series of Catalogue of fungi in China here. In the first paper of this series, we focus on plant-inhabiting fungi. A total of 33 new taxa are described all over China. These taxa include two new genera, viz., and N. yoshinagae. The morphological characteristics and phylogenetic evidence are used to support the establishment of these new taxa and the accuracy of their taxonomic placements. We hope that the series of Catalogue of fungi in China will contribute to Chinese fungal diversity and promote the significance of recording new fungal taxa from China. ARTICLE HISTORY

Fungal diversity notes 1717–1817: taxonomic and phylogenetic contributions on genera and species of fungal taxa

Article

Full-text available

Feb 2024
FUNGAL DIVERS

As the continuation of Fungal Diversity Notes series, the current paper is the 16th contribution to this series. A total of 103 taxa from seven classes in Ascomycota and Basidiomycota are included here. Of these 101 taxa, four new genera, 89 new species, one new combination, one new name and six new records are described in detail along with information of hosts and geographic distributions. The four genera newly introduced are Ascoglobospora, Atheliella, Rufoboletus and Tenuimyces. Newly described species are Akanthomyces xixiuensis, Agaricus agharkarii, A. albostipitatus, Amphisphaeria guttulata, Ascoglobospora marina, Astrothelium peudostraminicolor, Athelia naviculispora, Atheliella conifericola, Athelopsis subglaucina, Aureoboletus minimus, A. nanlingensis, Autophagomyces incertus, Beltrania liliiferae, Beltraniella jiangxiensis, Botryobasidium coniferarum, Calocybella sribuabanensis, Calonarius caesiofulvus, C. nobilis, C. pacificus, C. pulcher, C. subcorrosus, Cortinarius flaureifolius, C. floridaensis, C. subiodes, Crustomyces juniperi, C. scytinostromoides, Cystostereum subsirmaurense, Dimorphomyces seemanii, Fulvoderma microporum, Ginnsia laricicola, Gomphus zamorinorum, Halobyssothecium sichuanense, Hemileccinum duriusculum, Henningsomyces hengduanensis, Hygronarius californicus, Kneiffiella pseudoabdita, K. pseudoalutacea, Laboulbenia bifida, L. tschirnhausii, L. tuberculata, Lambertella dipterocarpacearum, Laxitextum subrubrum, Lyomyces austro-occidentalis, L. crystallina, L. guttulatus, L. niveus, L. tasmanicus, Marasmius centrocinnamomeus, M. ferrugineodiscus, Megasporoporia tamilnaduensis, Meruliopsis crystallina, Metuloidea imbricata, Moniliophthora atlantica, Mystinarius ochrobrunneus, Neomycoleptodiscus alishanense, Nigrograna kunmingensis, Paracremonium aquaticum, Parahelicomyces dictyosporus, Peniophorella sidera, P. subreticulata, Phlegmacium fennicum, P. pallidocaeruleum, Pholiota betulicola, P. subcaespitosa, Pleurotheciella hyalospora, Pleurothecium aseptatum, Resupinatus porrigens, Russula chlorina, R. chrysea, R. cruenta, R. haematina, R. luteocarpa, R. sanguinolenta, Synnemellisia punensis, Tenuimyces bambusicola, Thaxterogaster americanoporphyropus, T. obscurovibratilis, Thermoascus endophyticus, Trechispora alba, T. perminispora, T. subfarinacea, T. tuberculata, Tremella sairandhriana, Tropicoporus natarajaniae, T. subramaniae, Usnea kriegeriana, Wolfiporiella macrospora and Xylodon muchuanensis. Rufoboletus hainanensis is newly transferred from Butyriboletus, while a new name Russula albocarpa is proposed for Russula leucocarpa G.J. Li & Chun Y. Deng an illegitimate later homonym of Russula leucocarpa (T. Lebel) T. Lebel. The new geographic distribution regions are recorded for Agaricus bambusetorum, Bipolaris heliconiae, Crinipellis trichialis, Leucocoprinus cretaceus, Halobyssothecium cangshanense and Parasola setulosa. Corresponding to morphological characters, phylogenetic evidence is also utilized to place the above-mentioned taxa in appropriate taxonomic positions. The current morphological and phylogenetic data is helpful for further clarification of species diversity and exploration of evolutionary relationships in the related fungal groups.

Three coralloid species of the genus Trechispora (Trechisporales, Basidiomycota) in China: two newly discovered taxa and one reported for the first time

Article

Full-text available

Sep 2023

Two new species of Trechispora indigenous to southern China, T. laxa and T. tongdaoensis , are described and illustrated, and the first record of T. khokpasiensis in China is reported. Molecular phylogenetic analyses of the concatenated nuclear rDNA ITS1–5.8S–ITS2 and nuclear large subunit sequences supported the inclusion of the three species within the Trechispora clade, together with species formerly classified in Scytinopogon . The new species are similar in micromorphology to species of Trechispora (as traditionally circumscribed) but are distinguished by having coralloid basidiomata. A key to the known coralloid Trechispora species in China is provided.

Xenasmatellales ord. nov. and Xenasmatellaceae fam. nov. for Xenasmatella ( Agaricomycetes, Basidiomycota )

Article

Full-text available

Jun 2023

In the era of molecular phylogeny as dominant evidence in fungal taxonomy, the taxonomic framework of fungi adopted from morphological characteristics has been largely updated. Compared with other fungal groups, macrofungi underwent fewer updates at the order and higher level. In this study, the taxonomic placement of a poorly known macro-basidiomycetous genus Xenasmatella is studied. Phylogenetic and molecular clock analyses inferred from a seven-locus dataset support that the genus represents an order rank lineage. Accordingly, a monotypic order Xenasmatellales and a monotypic family Xenasmatellaceae are newly introduced for Xenasmatella within Agaricomycetes. The species diversity and relationships of Xenasmatella are further clarified with the aid of the phylogenetic analysis inferred from a four-locus dataset. In association with morphological characteristics, a new species Xenasmatella hjortstamii is described. Moreover, the distribution of Xenasmatella ailaoshanensis, X. gossypina, and X. wuliangshanensis previously known only from type localities in Yunnan Province, China are expanded. In addition, two unnamed single-specimen lineages of Xenasmatella from Victoria State, Australia and Sichuan, China are revealed, likely representing two potential new species of this genus. In summary, the current study updates the taxonomic framework of Agaricomycetes and provides a crucial supplement for comprehensively understanding the evolutionary history of this fungal class.

An updated taxonomic framework of Hymenochaetales (Agaricomycetes, Basidiomycota)

Article

Apr 2023

Three new species of Trechispora from Northern and Northeastern Thailand

Article

May 2023

During the rainy season in Thailand, specimens of coral fungi were collected from community forests in Chiang Mai and Kalasin Provinces. The species presented here, Trechispora khokpasiensis, T. saluangensis, and T. sanpapaoensis are characterized by coralloid basidiomata and described as new taxa in Hydnodontaceae (Trechisporales) based on analyses of their morphological characters and phylogenetic relationships from the nuclear ribosomal large subunit (LSU) as well as the internal transcribed spacer regions’ (ITS) sequence data. Morphological descriptions and illustrations, including color photographs, line drawings and phylogenetic trees, are presented. Comparisons with similar species are also provided.

Outline, divergence times and phylogenetic analyses of Trechisporales (Agaricomycetes, Basidiomycota)

Article

Full-text available

Apr 2022

Phylogenetic analyses inferred from the nuc rDNA ITS1-5.8S-ITS2 (ITS) data set and the combined 2-locus data set [5.8S + nuc 28S rDNA (nLSU)] of taxa of Trechisporales around the world show that Sistotremastrum family forms a monophyletic lineage within Trechisporales. Bayesian evolutionary and divergence time analyses on two data sets of 5.8S and nLSU sequences indicate an ancient divergence of Sistotremastrum family from Hydnodontaceae during the Triassic period (224.25 Mya). Sistotremastrum family is characterized by resupinate and thin basidiomata, smooth, verruculose, or odontoid- semiporoid hymenophore, a monomitic hyphal structure, and generative hyphae bearing clamp connections, the presence of cystidia and hyphidia in some species, thin-walled, smooth, inamyloid, and acyanophilous basidiospores. In addition, four new species, namely, Trechispora dentata, Trechispora dimitiella, Trechispora fragilis, and Trechispora laevispora, are described and illustrated. In addition, three new combinations, namely, Brevicellicium daweishanense, Brevicellicium xanthum, and Sertulicium limonadense, are also proposed.

Two New Species of Fibrodontia (Trechisporales, Basidiomycota) with a Key to Worldwide Species

Article

Full-text available

Nov 2021
J. Fungi

Fibrodontia is a genus of wood-inhabiting fungi consisting of four species so far, including F. gossypina as generic type. Two new species, Fibrodontia austrosinensis and F. subalba, are described and illustrated from China. Fibrodontia austrosinensis from southwestern China is characterized by a grandinioid to odontioid hymenophore with numerous small aculei, a dimitic hyphal system with scattered, smooth skeletal hyphae and ellipsoid basidiospores measuring 4.2–5.2 × 3.5–4.5 μm. Fibrodontia subalba from the West Tianshan Mountain in northwestern China is distinguished by an odontioid to hydnoid hymenophore, a dimitic hyphal system, and ellipsoid basidiospores measuring 3.7–4.4 × 2.8–3.4 μm. The phylogenies inferred from the data set of nuc rDNA ITS1-5.8S-ITS2 (ITS) and D1–D2 domains of nuc 28S rDNA (28S), and that of ITS, 28S, translation elongation factor (tef1α), and RNA polymerase II second largest subunit (rpb2) supported Fibrodontia as a monophyletic genus in the Trechisporales, and F. austrosinensis and F. subalba as separate lineages within Fibrodontia. Multi-rate Poisson Tree Processes, Automatic Barcode Gap Discovery and genetic distance methods based on ITS sequences of Fibrodontia also supported F. austrosinensis and F. subalba as distinct species. The taxonomic status of F. fimbriata that was recently transferred from Cystidiodendron, is briefly discussed. A key to all six known species of Fibrodontia is provided.

The First Whole Genome Sequencing of Sanghuangporus sanghuang Provides Insights into Its Medicinal Application and Evolution

Article

Full-text available

Sep 2021
J. Fungi

Sanghuangporus is a medicinal macrofungal genus typified by S. sanghuang, the very species utilized in traditional Chinese medicines by Chinese ancient people. To facilitate the medicinal application of S. sanghuang, we, for the first time, perform its genome sequencing and analyses from a monokaryon strain. A 33.34 Mb genome sequence was assembled to 26 contigs, which lead to the prediction of 8278 protein-coding genes. From these genes, the potential biosynthesis pathway of sesquiterpenoids was, for the first time, identified from Sanghuangporus, besides that of triterpenoids. While polysaccharides are the main medicinal metabolites in S. sanghuang, flavonoids are especially abundant medicinal metabolites comparing with other medicinal macrofungal groups. From the genomic perspective, S. sanghuang has a tetrapolar heterothallic mating system, and has its special nutritional strategy and advantageous medicinal properties compared with S. baumii and S. vaninii. A phylogenomics analysis indicates that Sanghuangporus emerged 15.39 million years ago and S. sanghuang has a closer phylogenetic relationship with S. baumii than S. vaninii. However, S. sanghuang shares a higher region of synteny and more orthologous genes, including carbohydrate-active enzymes with S. vaninii than S. baumii. A comparative genomics analysis with S. baumii and S. vaninii indicates that species diversification within Sanghuangporus may be driven by the translocation and translocation plus inversion of genome sequences, while the expansion and contraction of gene families may contribute to the host specificity of Sanghuangporus species. In general, the genome sequence of S. sanghuang provides insights into its medicinal application and evolution.

Competing sexual-asexual generic names in Agaricomycotina (Basidiomycota) with recommendations for use

Article

Full-text available

Dec 2021

With the change to one scientific name for fungal taxa, generic names typified by species with sexual or asexual morph types are being evaluated to determine which names represent the same genus and thus compete for use. In this paper generic names of the Agaricomycotina ( Basidiomycota ) were evaluated to determine synonymy based on their type. Forty-seven sets of sexually and asexually typified names were determined to be congeneric and recommendations are made for which generic name to use. In most cases the principle of priority is followed. However, 16 generic names are recommended for use that do not have priority and thus need to be protected: Aleurocystis over Matula; Armillaria over Acurtis and Rhizomorpha; Asterophora over Ugola; Botryobasidium over Acladium , Allescheriella, Alysidium, Haplotrichum , Physospora, and Sporocephalium; Coprinellus over Ozonium; Coprinopsis over Rhacophyllus; Dendrocollybia over Sclerostilbum and Tilachlidiopsis; Diacanthodes over Bornetina; Echinoporia over Echinodia; Neolentinus over Digitellus; Postia over Ptychogaster; Riopa over Sporotrichum; Scytinostroma over Artocreas, Michenera , and Stereofomes; Tulasnella over Hormomyces; Typhula over Sclerotium; and Wolfiporia over Gemmularia and Pachyma. Nine species names are proposed for protection: Botryobasidium aureum, B. conspersum , B. croceum , B. simile, Pellicularia lembosporum (syn. B. lembosporum ), Phanerochaete chrysosporium , Polyporus metamorphosus (syn. Riopa metamorphosa ), Polyporus mylittae (syn. Laccocephalum mylittae ), and Polyporus ptychogaster (syn . Postia ptychogaster ). Two families are proposed for protection: Psathyrellaceae and Typhulaceae . Three new species names and 30 new combinations are established, and one lectotype is designated.

Towards a Natural Classification of Hyphodontia Sensu Lato and the Trait Evolution of Basidiocarps within Hymenochaetales (Basidiomycota)

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Full-text available

Jun 2021
J. Fungi

Hyphodontia sensu lato, belonging to Hymenochaetales, accommodates corticioid wood-inhabiting basidiomycetous fungi with resupinate basidiocarps and diverse hymenophoral characters. Species diversity of Hyphodontia sensu lato has been extensively explored worldwide, but in previous studies the six accepted genera in Hyphodontia sensu lato, viz. Fasciodontia, Hastodontia, Hyphodontia, Kneiffiella, Lyomyces and Xylodon were not all strongly supported from a phylogenetic perspective. Moreover, the relationships among these six genera in Hyphodontia sensu lato and other lineages within Hymenochaetales are not clear. In this study, we performed comprehensive phylogenetic analyses on the basis of multiple loci. For the first time, the independence of each of the six genera receives strong phylogenetic support. The six genera are separated in four clades within Hymenochaetales: Fasciodontia, Lyomyces and Xylodon are accepted as members of a previously known family Schizoporaceae, Kneiffiella and Hyphodontia are, respectively, placed in two monotypic families, viz. a previous name Chaetoporellaceae and a newly introduced name Hyphodontiaceae, and Hastodontia is considered to be a genus with an uncertain taxonomic position at the family rank within Hymenochaetales. The three families emerged between 61.51 and 195.87 million years ago. Compared to other families in the Hymenochaetales, these ages are more or less similar to those of Coltriciaceae, Hymenochaetaceae and Oxyporaceae, but much older than those of the two families Neoantrodiellaceae and Nigrofomitaceae. In regard to species, two, one, three and 10 species are newly described from Hyphodontia, Kneiffiella, Lyomyces and Xylodon, respectively. The taxonomic status of additional 30 species names from these four genera is briefly discussed; an epitype is designated for X. australis. The resupinate habit and poroid hymenophoral configuration were evaluated as the ancestral state of basidiocarps within Hymenochaetales. The resupinate habit mainly remains, while the hymenophoral configuration mainly evolves to the grandinioid-odontioid state and also back to the poroid state at the family level. Generally, a taxonomic framework for Hymenochaetales with an emphasis on members belonging to Hyphodontia sensu lato is constructed, and trait evolution of basidiocarps within Hymenochaetales is revealed accordingly.

Notes on Taxonomy and Nomenclature of the Polypores

Article

Sep 2018

Rolf Singer

New Genera of Fungi

Article

Sep 2018

Rolf Singer

Forecasting the number of species of asexually reproducing fungi (Ascomycota and Basidiomycota)

Article

Mar 2022

Asexually reproducing fungi play a significant role in essential processes in managed and wild ecosystems such as nutrients cycling and multitrophic interactions. A large number of such taxa are among the most notorious plant and animal pathogens. In addition, they have a key role in food production, biotechnology and medicine. Taxa without or rare sexual reproduction are distinguished based on their sporulating structures and conidiomata in traditional morphology-based taxonomy. The number, variation and diversity of asexually reproducing taxa are insufficiently known, even though fungi capable of asexual reproduction may provide an untapped, rich biological resource for future exploitation. Currently, ca. 30,000 asexual species belonging to ca. 3800 genera have been reported (including 1388 coelomycetous and 2265 hyphomycetous genera). Recent reports (2017–2020) reiterate that the number of asexually producing fungi is higher than the number of frequently sexually-reproducing fungi. With the advent of molecular tools and the abandonment of the dual nomenclature system for pleomorphic fungi, priority criteria were established and revisited in the latest outline of fungi and fungus-like taxa. However, species numbers and taxonomic boundaries of pleomorphic taxa and their synanamorphs or synasexual morphs have yet to be addressed. The number of species of speciose genera (e.g. Alternaria, Aspergillus, Cercospora, Fusarium, Phoma and Pseudocercospora), cryptic species, species of pleomorphic genera, less studied life modes (such as lichenicolous taxa, taxa from extreme environments) and species from biodiversity-rich areas still need evaluation to achieve more reliable estimates of their diversity. This paper discusses the current knowledge on the matter, with diversity estimates, and potential obstacles in several chapters on (1) speciose genera; (2) pleomorphic genera; (3) cryptic species; (4) well-studied but insufficiently resolved taxa, e.g. leaf inhabiting species, marine fungi, (5) less studied life modes, e.g. lichenicolous, rock-inhabiting fungi, insect-associated and yeast-forming taxa and (6) species from biodiversity-rich areas.

Sistotremastrum limonadense sp. nov. from French Guiana

Article

Jun 2021

A new species of Sistotremastrum J. Erikss. (Basidiomycota, Trechisporales) with (6-8)-spored basidia is described and supported by morphology and phylogenetic analyses based on ITS rDNA sequences.

Scytinopogon caulocystidiatus and S. foetidus spp. nov., and five other species recorded from Brazil

Article

Apr 2021

Seven species of Scytinopogon in the Atlantic Forest of Brazil are described. Scytinopogon caulocystidiatus and S. foetidus are proposed as new species based on morphological and molecular data. Five other species are presented: S. chartaceus, S. dealbatus and S. robustus, originally described from Brazil; and S. pallescens and S. scaber, previously reported for Brazil. Illustrations of the basidiomata and microstructures, including SEM images of ornamented basidiospores, are provided for all taxa. Comparisons with morphologically similar taxa and a key to the seven species of Scytinopogon known to occur in Brazil are also presented.

Trechisporales emended with a segregation of Sistotremastrales ord. nov. (Basidiomycota)

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