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Two new species of Infundibulicybe (Omphalinaceae, Agaricales) from northern China

May 2024
Phytotaxa 649(2):193-210

May 2024
649(2):193-210

DOI:10.11646/phytotaxa.649.2.4

Authors:

Ning Mao

Capital Normal University

Li Fan

Capital Normal University

Infundibulicybe confusa sp. nov. and I. phaeocentralis sp. nov. were described based on specimens from Shanxi Province in northern China. Morphologically, I. phaeocentralis is similar to I. rufa but the latter is differentiated by its isolated phylogenetic position and varied colors of the basidiomata. Infundibulicybe confusa is similar to I. alkaliviolascens, but I. alkaliviolascens has a pileus that turns violet to purplish pink in KOH and long basidiospores. Our morphological and phylogenetic analyses supported the taxonomic position of the two new species in Infundibulicybe. Additionally, a key to the species of Infundibulicybe in Shanxi Province is provided.

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Phytotaxa 649 (2): 193–210

https://www.mapress.com/pt/

ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)

Accepted by Genevieve Gates: 4 May 2024; published: 20 May 2024

https://doi.org/10.11646/phytotaxa.649.2.4

193

Two new species of Infundibulicybe (Omphalinaceae, Agaricales) from northern

China

YU-XIN ZHANG1,3, NING MAO1,4, TING LI2,5 & LI FAN1,6*

1College of Life Sciences, Capital Normal University, Xisanhuanbeilu 105, Haidian, Beijing 100048, China

2Department of Life Sciences, National Natural History Museum of China, Tianqiaonandajie126, Dongcheng, Beijing, 100050, China

�

zhangyuxin7237@163.com; https://orcid.org/0000-0002-6757-1903

�

373898825@qq.com; https://orcid.org/0000-0003-1564-9446

�

1377090363@qq.com; https://orcid.org/0000-0001-7809-3217

�

fanli@mail.cnu.edu.cn; https://orcid.org/0000-0001-9887-7086

*Corresponding author:

�

fanli@mail.cnu.edu.cn

Abstract

Infundibulicybe confusa sp. nov. and I. phaeocentralis sp. nov. were described based on specimens from Shanxi Province

in northern China. Morphologically, I. phaeocentralis is similar to I. rufa but the latter is differentiated by its isolated

phylogenetic position and varied colors of the basidiomata. Infundibulicybe confusa is similar to I. alkaliviolascens, but

I. alkaliviolascens has a pileus that turns violet to purplish pink in KOH and long basidiospores. Our morphological and

phylogenetic analyses supported the taxonomic position of the two new species in Infundibulicybe. Additionally, a key to the

species of Infundibulicybe in Shanxi Province is provided.

Key words: clitocyboid, multigene, novel taxa, taxonomy

Introduction

The genus Infundibulicybe Harmaja (2003:215) (Omphalinaceae, Agaricales) was proposed to accommodate those

species with lacrymoid basidiospores that traditionally were placed in Clitocybe (Fr.) Staude (1857:122). However, the

classification status of the genus was not widely accepted until the advent of molecular phylogenetics. Matheny et al.

(2006) recognized the genus in the Agaricales and as a sister group of the Tricholomatoid clade, based on a multilocus

analysis of a six-gene region. Binder et al. (2010) conducted a phylogenetic study of the Agaricomycetidae using a

six-gene combination, which further confirmed that Infundibulicybe was a distinct genus independent of Clitocybe, if

Clitocybe is not typified by Clitocybe infundibuliformis (Schaeff.) Quél (1872:88) (He et al. 2023, He & Yang 2023).

The species of Infundibulicybe are mainly characterized by a slightly depressed to infundibuliform pileus, decurrent

lamellae, and smooth, cyanophobic and lacrymoid basidiospores with confluent base. Furthermore, the basidiospores

do not adhere in tetrads and the mycelium is incapable of reducing nitrate (Harmaja 2003, Vizzini et al. 2011, Xu

et al. 2022, Vizzini et al. 2024). Except for I. geotropa (Bull.) Harmaja (2003:216) (Harmaja 2003, Akata et al.

2012, Ali et al. 2020), species of Infundibulicybe are saprobes, mainly occurring in leaf or needle litter, sometimes

in remnants of herbaceous plants (Harmaja 2003), and mainly occur in temperate, boreal and alpine regions of the

Northern Hemisphere (Harmaja 2003, Kirk et al. 2008, Vizzini et al. 2011, Zhao et al. 2016, Ishaq et al. 2019, Ali et

al. 2020, Vizzini et al. 2024). Currently, a total of 31 binomials are listed in Index Fungorum (www. indexfungorum.

org/Names/Names.asp), but only 25 species are currently classified in this genus. In China, eight Infundibulicybe

species have been reported and confirmed (Wei et al. 2015, Zhao et al. 2016, Liu et al. 2019, Xu et al. 2022, He &

Yang 2023), including I. alkaliviolascens (Bellù) Bellù (2012:6), I. bispora Z.M. He & Zhu L. Yang (2023:702), I.

ellipsospora Z.M. He & Zhu L. Yang (2023:703), I. gibba (Pers.) Harmaja (2003:207), I. hongyinpan L. Fan & H. Liu

(2018:277), I. rufa Q. Zhao, K.D. Hyde, J.K. Liu & Y.J. Hao (2016:136), I. squamulosa (Pers.) Harmaja (2003:217)

and I. trachyspora J.Z. Xu, J.C. Qin & Yu Li (2022:2).

During the study of the macrofungal flora of Shanxi Province, northern China, specimens of Infundibulicybe have

been collected since 2017. Based on morphological and molecular phylogenetic analyses, we found some specimens

ZHANG ET AL.

representing two taxa new to science. The aim of this paper is to describe and illustrate the two new species occurring

in Shanxi Province.

Materials and methods

Site description

The specimens we studied were collected in Shanxi Province, northern China. The Guancen Mountains are situated

in the northwest of Shanxi Province which has a northern temperate continental climate being cold and dry, with an

annual average temperature of 7.9 °C–11.7 °C, and an annual rainfall of 700–800 mm. Heyeping is the highest peak

with an elevation of 2,784 m. The vegetation mainly includes Betula spp., Larix gmelinii var. principis-rupprechtii

(Mayr) Pilg. (1926:327), Picea asperata Mast. (1906:419), Picea meyeri Rehder & E.H.Wilson (1914:28) and Picea

wilsonii Mast. (1903:133). The soil types mainly include mountain brown soil, mountain cinnamon soil, subalpine

meadow soil and brown forest soil.

The Guandi Mountain range is in the west of Shanxi Province, and experiences a temperate continental climate.

The average annual temperature is 3.5 °C, the annual precipitation is 830 mm, and the highest point has an elevation

of 2,831 m. The vegetation mainly includes Betula spp., Larix gmelinii var. principis-rupprechtii, Picea spp., Pinus

tabuliformis Carrière (1867:510) and Populus spp.

Morphological studies

The fresh basidiomata were photographed in the field and macroscopic characteristics were recorded. ColorHexa (https://

www.colorhexa.com/) was used to match colors. The specimens were dried in a fruit drier at 40–45 °C and deposited

in BJTC (Herbarium Biology Department, Capital Normal University, Beijing, China). Microscopic characteristics

were examined from sections of dried material, rehydrated in distilled water, and then mounted in either 3 % KOH,

Congo Red, Melzer’s reagent or Cotton Blue. All measurements were made in Congo Red. For each specimen at

least 30 spores and 15 basidia were measured. Spore measurements exclude apiculus. Basidia measurements exclude

sterigmata, which are given separately. The dimensions of basidiospores are given by (a–)b–c(–d). The range ‘b–c’

covers a minimum of 90 % of the measured values. Extreme values ‘a’ and ‘d’ are given in parentheses. Q refers to the

length/width ratio in the profile view of basidiospores, and Qm refers to the average Q of all basidiospores ± sample

standard deviation.

To observe any reaction to alkali, a drop of 5 % KOH and a drop of distilled water (as the control) were applied

separately to the dried pileus. White filter paper was placed beneath the specimen as a background to better observe

any color change. Each species was subjected to three replications of the test.

DNA extraction, PCR amplification and DNA sequencing

Dried basidiomata were crushed by shaking for 45 s at 30 Hz 2−4 times (Mixer Mill MM301, Retsch, Haan, Germany)

in a 1.5 mL tube together with 3 mm diam tungsten carbide balls; total genomic DNA was extracted using the modified

CTAB method (Gardes & Bruns 1993). The ITS region was amplified using the primers ITS1F/ITS4 (White et al.

1990, Gardes & Bruns 1993), primers LR0R and LR5 were employed for nrLSU (Vilgalys & Hester 1990), primers

EF1-983F and EF1-1953R were used to amplify the TEF1(Rehner & Buckley 2005), primers RPB1-Af and fRPB1-Cr

were used to amplify the RPB1(Matheny et al. 2002), primers bRPB2-6F and bRPB2-7.1R were used to amplify the

RPB2 (Matheny 2005), and primers ATP6-3 and ATP6-6r were used to amplify ATP6 (Kretzer & Bruns 1999, Binder

& Hibbett 2003).

Polymerase Chain Reactions (PCR) for all regions were performed in 25 µL reaction containing 2 µL DNA

template, 1 µL primer (10 µM) each, 12.5 µL of 2 × Master Mix [Tiangen Biotech (Beijing) Co.], 8.5 µL ddH2O.

PCR reactions were implemented as follows: for the ITS region: an initial denaturation at 94 ℃ for 3 min, followed

by 35 cycles at 94 ℃ for 45 s, 55 ℃ for 45 s, 72 ℃ for 1 min, and a final extension at 72 ℃ for 10 min. For the other

five regions the protocols described by He & Yang (2023) were followed. The PCR products were sent to Beijing

Zhongkexilin Biotechnology Co. Ltd. for purification, sequencing, and editing. Validated sequences were deposited

in GenBank (http://www.ncbi.nlm.nih.gov/). Other sequences of Infundibulicybe and related species were selected

mainly from those used by Walther et al. (2005), Matheny et al. (2006), Maeta et al. (2008), Bjorbaekmo et al. (2010),

Vizzini et al. (2011), Geml et al. (2012), Osmundson et al. (2013), Tian et al. (2015), Tomáš et al. (2016), Zhao et al.

(2016), Vu et al. (2018), Ishaq et al. (2019), Liu et al. (2019), Ali et al. (2020), Giusti et al. (2021), He & Yang (2022),

Xu et al. (2022), and He & Yang (2023). The accession numbers of new and downloaded sequences stored in the NCBI

database (http://www.ncbi.nlm.nih.gov/) are provided in Table 1 and Supplementary Table 2.

TABLE 1. Infundibulicybe specimens used in molecular phylogenetic analyses and their GenBank accession numbers (ITS).

Newly generated sequences are in black bold.

Species Voucher Locality ITS References

Clitopaxillus fibulatus KUN-HKAS 115964 China: Gansu OP964650 He & Yang (2023)

I. alkaliviolascens KUN-HKAS 115938 China: Xinjiang OP983898 He & Yang (2023)

I. alkaliviolascens KUN-HKAS 115942 China: Xinjiang OP983901 He & Yang (2023)

I. alkaliviolascens LUGO:ECC17121604 Spain MW376672 Unpublished

I. alkaliviolascens 13243 Italy JF907807 Osmundson et al. (2013)

I. alkaliviolascens BJTC FM1383 China: Shanxi PP492517 This study

I. alkaliviolascens BJTC FM4108 China: Shanxi PP492512 This study

I. alkaliviolascens BJTC FM3164 China: Shanxi PP492508 This study

I. alkaliviolascens BJTC FM2324 China: Shanxi PP492515 This study

I. alkaliviolascens BJTC FM2106 China: Shanxi PP492511 This study

I. alkaliviolascens BJTC FM2363 China: Shanxi PP492507 This study

I. alkaliviolascens BJTC FM2965 China: Shanxi PP492504 This study

I. alkaliviolascens HSA123 China: Shanxi PP492516 This study

I. alkaliviolascens BJTC FM3396 China: Shanxi PP492506 This study

I. alkaliviolascens BJTC FM3298 China: Shanxi PP492514 This study

I. alkaliviolascens BJTC FM2910 China: Shanxi PP492509 This study

I. alkaliviolascens BJTC FM1458 China: Shanxi PP492503 This study

I. alkaliviolascens BJTC FM3081 China: Shanxi PP492505 This study

I. alkaliviolascens BJTC FM1653 China: Shanxi PP492497 This study

I. alkaliviolascens BJTC FM910 China: Shanxi PP492496 This study

I. alkaliviolascens HSA120 China: Shanxi PP492501 This study

I. alkaliviolascens BJTC FM1658 China: Shanxi PP492500 This study

I. alkaliviolascens BJTC FM1890 China: Shanxi PP492499 This study

I. alkaliviolascens BJTC FM1708 China: Shanxi PP492498 This study

I. alkaliviolascens BJTC FM2690 China: Shanxi PP492510 This study

I. alkaliviolascens BJTC FM604 China: Shanxi PP492502 This study

I. alkaliviolascens BJTC FM1540 China: Shanxi PP492513 This study

I. bispora KUN-HKAS 115961 China: Jilin OP983927 He & Yang (2023)

I. bispora KUN-HKAS 115962 China: Jilin OP983928 He & Yang (2023)

I. bispora KUN-HKAS 115963 China: Gansu OP983929 He & Yang (2023)

I. bispora Z7 China: Hei Longjiang GU134502 Unpublished

I. bresadolana Mushroom Observer 470945 USA OM980624 Unpublished

I. bresadolana 16016 Italy JF907810 Osmundson et al. (2013)

I. catinus I France HM631720 Vizzini et al. (2011)

I. confusa (holotype) BJTC FM1281 China: Shanxi PP492518 This study

I. confusa BJTC FM1285 China: Shanxi PP492519 This study

I. ellipsospora KUN-HKAS 114671 China: Yunnan OP983917 He & Yang (2023)

I. ellipsospora KUN-HKAS 70065 China: Yunnan OP983918 He & Yang (2023)

I. ellipsospora KUN-HKAS 105353 China: Yunnan OP983919 He & Yang (2023)

I. ellipsospora KUN-HKAS 115959 China: Tibet OP983920 He & Yang (2023)

I. ellipsospora KUN-HKAS 54633 China: Yunnan OP983921 He & Yang (2023)

......continued on the next page

ZHANG ET AL.

TABLE 1. (Continued)

Species Voucher Locality ITS References

I. ellipsospora KUN-HKAS 56928 China: Yunnan OP983922 He & Yang (2023)

I. ellipsospora KUN-HKAS 56965 China: Yunnan OP983923 He & Yang (2023)

I. ellipsospora KUN-HKAS 116251 China: Tibet OP983924 He & Yang (2023)

I. ellipsospora KUN-HKAS 57667 China: Yunnan OP983925 He & Yang (2023)

I. ellipsospora KUN-HKAS 115960 China: Yunnan OP983926 He & Yang (2023)

I. geotropa ALV4344 — KT122792 Unpublished

I. geotropa IZS189 Italy MZ005561 Giusti et al. (2021)

I. geotropa AODJ.161.15 — MZ374498 Unpublished

I. gibba KUN-HKAS 81200 China: Yunnan MZ718993 Unpublished

I. gibba KUN-HKAS 115948 China: Yunnan OP983907 He & Yang (2023)

I. gibba KUN-HKAS 115950 China: Jilin OP983909 He & Yang (2023)

I. gibba KUN-HKAS 115951 China: Liaoning OP983910 He & Yang (2023)

I. gibba KUN-HKAS 53323 China: Yunnan MZ718994 Unpublished

I. gibba KUN-HKAS 57657 China: Yunnan MZ718995 Unpublished

I. gibba KUN-HKAS 105606 China: Gansu MZ718996 Unpublished

I. gibba KUN-HKAS 73334 China: Yunnan MZ718997 Unpublished

I. gibba KUN-HKAS 115952 China: Yunnan OP983911 He & Yang (2023)

I. gibba AFTOL-ID 1508 USA DQ490635 Matheny et al. (2006)

I. gibba KUN-HKAS 92032 China: Liaoning MZ675563 He & Yang (2022)

I. gibba BJTC FM99 China: Shanxi MH581237 Liu et al. (2019)

I. gibba TZ16 Czech Republic LN714556 Tomáš et al. (2016)

I. gibba PNGS/304 Poland KX882674 Unpublished

I. gibba CBS 128.42 Sweden MH856103 Vu et al. (2018)

I. gibba H01IK17 Germany HF675204 Unpublished

I. gibba CNF 1/7258 Croatia MK169232 Unpublished

I. gibba LUGO:ECC18071802 Spain MW376674 Unpublished

I. gibba AB 17-08-16 France MN661006 Unpublished

I. gibba M Italy HM631722 Vizzini et al. (2011)

I. gibba CM052 Algeria KP826742 Unpublished

I. gibba AR09751 Mexico KT875020 Unpublished

I. gibba ASIS22552 — KF668316 Unpublished

I. gibba BJTC FM3133 China: Shanxi PP492455 This study

I. gibba BJTC FM1455 China: Shanxi PP492460 This study

I. gibba BJTC FM3476 China: Shanxi PP492464 This study

I. gibba BJTC FM723 China: Shanxi PP492461 This study

I. gibba BJTC FM2287 China: Shanxi PP492463 This study

I. gibba BJTC FM682 China: Shanxi PP492456 This study

I. gibba BJTC FM3121 China: Shanxi PP492454 This study

I. gibba BJTC FM1706 China: Shanxi PP492459 This study

I. gibba BJTC FM761 China: Shanxi PP492458 This study

I. gibba BJTC FM757 China: Shanxi PP492457 This study

I. gibba BJTC FM2104 China: Shanxi PP492462 This study

I. gibba BJTC FM2123 China: Shanxi PP492453 This study

I. gibba BJTC FM3367 China: Shanxi PP492468 This study

I. gibba HSA438 China: Shanxi PP492465 This study

I. gibba BJTC FM2052 China: Shanxi PP492466 This study

I. gibba HSA331 China: Shanxi PP492467 This study

I. gibba BJTC FM1879 China: Shanxi PP492469 This study

......continued on the next page

TABLE 1. (Continued)

Species Voucher Locality ITS References

I. gibba BJTC FM2279 China: Shanxi PP492470 This study

I. gibba BJTC FM2283 China: Shanxi PP492451 This study

I. gibba BJTC FM1086 China: Shanxi PP492450 This study

I. gibba BJTC FM2535 China: Shanxi PP492452 This study

I. gibba BJTC FM1713 China: Shanxi PP492449 This study

I. gibba BJTC FM174 China: Shanxi PP492448 This study

I. gibba BJTC FM1492 China: Shanxi PP492471 This study

I. gibba BJTC FM2048 China: Shanxi PP492472 This study

I. gibba BJTC FM2471 China: Shanxi PP492475 This study

I. gibba BJTC FM1851 China: Shanxi PP492473 This study

I. gibba BJTC FM3016 China: Shanxi PP492474 This study

I. gibba BJTC FM3293 China: Shanxi PP492476 This study

I. gibba BJTC FM934 China: Shanxi PP492477 This study

I. hongyinpan KUN-HKAS 115955 China: Tibet OP983914 He & Yang (2023)

I. hongyinpan KUN-HKAS 115956 China: Yunnan OP983915 He & Yang (2023)

I. hongyinpan KUN-HKAS 105573 China: Gansu MZ718999 Unpublished

I. hongyinpan BJTC FM187 China: Shanxi MH581239 Liu et al. (2019)

I. hongyinpan BJTC FM394 China: Shanxi MH581238 Liu et al. (2019)

I. hongyinpan BJTC FM216 China: Shanxi MH581240 Liu et al. (2019)

I. hongyinpan iNAT:17303634 USA MZ146330 Unpublished

I. hongyinpan NBRC 30524 Japan AB301607 Maeta et al. (2008)

I. hongyinpan BJTC FM2504 China: Shanxi PP492482 This study

I. hongyinpan BJTC FM181 China: Shanxi PP492478 This study

I. hongyinpan BJTC FM294 China: Shanxi PP492479 This study

I. hongyinpan BJTC FM3202 China: Shanxi PP492483 This study

I. hongyinpan HSA391 China: Shanxi PP492485 This study

I. hongyinpan BJTC FM1469 China: Shanxi PP492481 This study

I. hongyinpan HSA171 China: Shanxi PP492484 This study

I. hongyinpan HSA244 China: Shanxi PP492480 This study

I. kotanensis LAH35902 Pakistan MN017278 Ishaq et al. (2019)

I. kotanensis LAH35902 Pakistan MN017274 Ishaq et al. (2019)

I. lateritia 19_N1F29_F35 Norway HQ445603 Bjorbaekmo et al. (2010)

I. lateritia 0731-39 Norway MW748452 Unpublished

I. lateritia KH74 Norway GU234109 Geml et al. (2012)

I. macrospora LAH35863 (holotype) Pakistan MN066565 Ali et al. (2020)

I. macrospora LAH35864 Pakistan MN066566 Ali et al. (2020)

I. macrospora SWAT0901356 Pakistan MT156434 Unpublished

I. macrospora SWAT0901357 Pakistan MT156435 Unpublished

I. mediterranea IZS118 Italy MZ005521 Giusti et al. (2021)

I. mediterranea O Italy HM631724 Vizzini et al. (2011)

I. phaeocentralis (holotype) BJTC FM2899 China: Shanxi PP492487 This study

I. phaeocentralis BJTC FM3215 China: Shanxi PP492486 This study

I. rufa KUN-HKAS 77865 China: Hubei MZ719000 Unpublished

I. rufa KUN-HKAS 57811 China: Tibet MZ719001 Unpublished

I. rufa KUN-HKAS 87741 China: Sichuan KX239806 Zhao et al. (2016)

I. rufa BJTC FM562 China: Shanxi PP492492 This study

I. rufa HSA134 China: Shanxi PP492493 This study

I. rufa BJTC FM2056 China: Shanxi PP492494 This study

......continued on the next page

ZHANG ET AL.

TABLE 1. (Continued)

Species Voucher Locality ITS References

I. rufa BJTC FM1271 China: Shanxi PP492495 This study

I. squamulosa CBS:128.63 France MH858237 Vu et al. (2018)

I. squamulosa 10423(9) — KP226187 Tian et al. (2015)

I. squamulosa BJTC FM2353 China: Shanxi PP492488 This study

I. squamulosa BJTC FM2483 China: Shanxi PP492490 This study

I. squamulosa BJTC FM2452 China: Shanxi PP492491 This study

I. squamulosa BJTC FM2451 China: Shanxi PP492489 This study

I. trachyspora HMJU 744 China: Jilin MW736885 Xu et al. (2022)

I. trachyspora KUN-HKAS 78801 China: Hubei MZ719002 Unpublished

Omphalina pyxidata AMB:19294 Italy OR863467 Vizzini et al. (2024)

Omphalina pyxidata AMB:19295 Italy OR863468 Vizzini et al. (2024)

Pseudoclitocybe cyathiformis KUN-HKAS 73390 China: Yunnan MZ718991 Unpublished

Phylogenetic analyses

The ITS dataset and the combined LSU-TEF1-RPB1-RPB2-ATP6 dataset were used to investigate the phylogenetic

positions of the Infundibulicybe species of Shanxi. Clitopaxillus fibulatus P.-A. Moreau, Dima, Consiglio & Vizzini

(2018a:116) and Pseudoclitocybe cyathiformis (Bull.) Singer (1956:725) were chosen as outgroups (Alvarado et al.

2018b, He & Yang 2022). The sequences of the markers were aligned in MAFFT v.7.110 (Katoh & Standley 2013)

under default parameters. Ambiguously aligned sites were identified by Gblocks v.0.91b (Castresana 2000) with

default parameters.

Phylogenetic analyses were conducted using Maximum Likelihood (ML) and Bayesian Inference (BI). Maximum

Likelihood (ML) was performed using RAxML 8.0.14 (Stamatakis 2014) by running 1000 bootstrap replicates under

the GTRGAMMAI model. Bayesian Inference (BI) was conducted using MrBayes v3.1.2 (Ronquist & Huelsenbeck

2003) as an additional method of evaluating branch support. In the BI analysis, for the ITS dataset, we selected the

best substitution model (GTR+I+G) determined by MrModeltest v2.3 (Nylander 2004); for the combined dataset, a

partitioned mixed model was used by defining the sequences of LSU, TEF1, RPB1, RPB2 and ATP6 as five independent

partitions, and each gene was separately modeled with different parameters. The best-fitted substitution model for

each gene marker was determined through MrModeltest v2.3 (Nylander 2004) using Akaike Information Criterion

(AIC). GTR+I was chosen as the best model for LSU, GTR+I+G was chosen as the best model for TEF1, K80+I+G

was chosen as the best model for RPB1 and SYM+I+G was chosen as the best model for RPB2 whereas HKY+I was

selected as the best model for ATP6. We used two independent runs with four Markov chains Monte Carlo (MCMC)

for 1,215, 000 generations under the default settings. The average standard deviation of split frequencies (ASDSF)

values was far lower than 0.01 at the end of the runs. Trees were sampled every 100 generations after burn-in (25 %

of trees were discarded as the burn-in phase of the analyses, set up well after convergence), and 50 % majority-rule

consensus trees were constructed.

Clades with bootstrap support (BS) ≥ 70 % and Bayesian posterior probability (PP) ≥ 0.99 were considered

significantly supported (Hillis & Bull 1993, Alfaro et al. 2003). All phylogenetic trees were viewed with TreeView

(Page 2001).

Results

Molecular phylogenetic analyses

In this study, 71 ITS sequences were newly generated from our specimens. The ITS dataset contained 16 taxa,

comprising 569 characters. ML and BI analyses yielded similar tree topologies and therefore only the tree inferred

from the ML analysis is shown (Fig. 1). According to the ITS phylogenetic tree, our specimens were resolved into

seven strongly supported branches, and five of them were identified as known species, i.e., I. alkaliviolascens, I. gibba,

I. hongyinpan, I. rufa and I. squamulosa, while the remaining two clade were independent, supporting they are new to

science, and these are described in this paper as I. confusa and I. phaeocentralis.

FIGURE 1. The ML phylogenetic tree based on the ITS sequences of Infundibulicybe. Clitopaxillus fibulatus and Pseudoclitocybe

cyathiformis are the outgroups. Values at the node represent ML bootstrap (≥ 70 % BS, left) and Bayesian posterior probability (≥ 0.99 PP,

right) support. Our specimen sequences are printed in bold and the new species sequences are highlighted in blue in the tree.

ZHANG ET AL.

FIGURE 1. The ML phylogenetic tree based on the ITS sequences of Infundibulicybe. (Continued)

The combined LSU-TEF1-RPB1-RPB2-ATP6 dataset included 12 taxa and 4,102 characters (886 from LSU, 935

from TEF1, 1116 from RPB1, 728 from RPB2 and 437 from ATP6, gaps included). ML and BI analyses yielded similar

tree topologies and therefore only the tree inferred from the ML analysis is shown (Fig. 2). The multilocus phylogenetic

tree further determined that the two new species recognized in the ITS phylogeny were distinct species. Infundibulicybe

phaeocentralis sp. nov. was phylogenetically related to I. bispora and I. gibba, whereas, Infundibulicybe confusa sp.

nov, was related to I. alkaliviolascens.

FIGURE 2. The ML phylogenetic tree based on the combined LSU-TEF1-RPB1-RPB2-ATP6 sequences of Infundibulicybe. Clitopaxillus

fibulatus and Pseudoclitocybe cyathiformis are outgroups. Values at the node represent ML bootstrap (≥ 70 % BS, left) and Bayesian

posterior probability (≥ 0.99 PP, right) support. Our specimen sequences are printed in bold and the new species sequences are highlighted

in blue in the tree.

Taxonomy

Infundibulicybe confusa L. Fan & Y.X. Zhang sp. nov. Figs. 3, 4

MycoBank:—MB852897

Diagnosis:—Basidiomata small to medium; pileus 1.7–3.9 cm diam, infundibuliform; surface dry, smooth to finely

felty, brownish flesh to pale brown, center and margin always much lighter, pale brown to cream; stipe cylindrical,

coarser towards the base, hollow, surface with longitudinally fibrils, yellowish brown, base inflated with abundant

white tomentum, and shorter basidiospores (5–)5.5–6.5(–7) × 3.5–4.5 μm.

ZHANG ET AL.

Etymology:—confusa, refers to this species being confused with Infundibulicybe alkaliviolascens due to the

appearance of the basidioma.

Holotype:—CHINA. Shanxi Province, Jiaocheng County, Guandi Mountain, Chailugou Village, on soil, in mixed

forest of Larix gmelinii var. principis-rupprechtii and Picea asperata, elev. 1,600 m, 111°24’2’’E, 37°47’25’’N, 29

July 2020, H. Liu, J. Yang LH970 (BJTC FM1281).

Description:—Basidiomata small to medium, clitocyboid. Pileus 1.7–3.9 cm diam, infundibuliform; dry, smooth

to finely felty, brownish flesh (#e0b89a) to pale brown (#c19472), center and margin always much lighter, pale brown

(#edceba) to cream (#d8d0bb), margin straight, sometimes slightly involute. Lamellae long-decurrent, white (#eef0e8)

to light cream (#f1f0de), moderately crowded, with lamellulae of 1 or 3 tiers, edges smooth and concolorous with faces.

Stipe 2.5–3.9 × 0.6–1 cm, central, cylindrical, coarser towards the base, hollow, longitudinally fibrillose, concolorous

with the pileus, base inflated with abundant white tomentum. 5 % KOH on the surface of the dried pileus produced no

color change.

FIGURE 3. Basidiomata of Infundibulicybe confusa. a, b. BJTC FM1281 (holotype). Scale bars: a, b= 10 mm. Photos by: Hong Liu.

FIGURE 4. Microscopic features of Infundibulicybe confusa. a. Basidiospores. b. Pileipellis. c. Basidia. Scale bars: a = 5 μm. b, c = 10

μm. Drawings by: Yuxin Zhang.

Basidiospores (5–)5.5–6.5(–7) × 3.5–4.5 μm, Q=1.4–1.6, Qm=1.5±0.1, ellipsoidal in face view, lacrymoid in

lateral view, hyaline, smooth, inamyloid, cyanophilous, color not changed in KOH. Basidia 21–37.5 × 4–7 μm,

clavate, hyaline, 4-spored or 2-spored, sterigmata 2–5 μm long. Cystidia absent. Lamellar trama regular; hyphae 3–8

µm wide, colorless, hyaline, cylindrical. Pileipellis a cutis composed of repent, cylindrical, subparallel, septate, thin-

walled hyphae, with intracellular yellowish brown pigment 3–11 μm wide. Stipitipellis a cutis composed of subparallel

cylindrical hyphae 5–8 μm wide. Clamp connections present in all parts of the basidiomata.

Ecology and distribution:—associated with the mixed forest of Larix gmelinii var. principis-rupprechtii and Picea

asperata, at elevation 1,600 m, only known from Shanxi Province, northern China.

Additional specimen examined:—CHINA. Shanxi Province, Jiaocheng County, Guandi Mountain, Chailugou

Village, on soil, in mixed forest of Larix gmelinii var. principis-rupprechtii and Picea asperata, elev. 1,600 m,

111°24’2’’E, 37°47’25’’N, 29 July 2020, H. Liu, J. Yang LH974 (BJTC FM1285).

Notes:—In the ITS phylogenetic tree (Fig. 1), Infundibulicybe confusa and I. bresadolana are phylogenetically

closely related to each other, but I. bresadolana has smooth basidiomata, more reddish colors, brownish gills in mature

basidiomata, longer and narrower basidiospores (5.5–8.0 × 3.1–4 μm), which are usually lacrymoid, or ellipsoid to

oblong, and usually grows on grasslands associated with Dryas sp. (Harmaja 1969). Morphologically, I. alkaliviolascens

shares similarities with I. confusa in the yellowish brown basidiomata, but it can be differentiated from the latter by the

violet to purplish pink reaction to KOH on the dried pileus, and long basidiospores (6–8.5(9) × (3)3.5–4.5 μm) (He &

Yang 2023).

Infundibulicybe phaeocentralis L. Fan & Y.X. Zhang sp. nov. Fig. 5

MycoBank:—MB852896

Diagnosis:—Basidiomata small to medium; pileus 2.6–5.4 cm diam, infundibuliform, dry, smooth, brick red to reddish

brown, stipe hollow, yellowish brown, surface with inconspicuous longitudinal fibrils; shorter basidiospores (6–)6.5–

8(–8.5) × 4–6 μm and bigger basidia 25–40 × 6–7.5 μm.

Etymology:—phaeocentralis, refers to the darker center of the pileus.

Holotype:—CHINA. Shanxi Province, Guancen Mountain, on soil, in mixed forest of Larix gmelinii var. principis-

rupprechtii and Picea asperata, 2 September 2022, J.Z. Cao CF1197 (BJTC FM2899).

Description:—Basidiomata small to medium, clitocyboid. Pileus 2.6–5.4 cm diam, infundibuliform; dry, smooth,

glabrous, brick red (#c2847a) to reddish brown (#8c564a), center and margin always darker, deep reddish brown

(#67393b), margin straight. Lamellae long-decurrent, yellowish white (#f2f6b6) to deep cream (#d2b86b), moderately

crowded, with 1 to 4 tiers of lamellulae, edges smooth and concolorous with faces. Stipe 3.7–5.7 × 0.6–0.9 cm,

centrally attached, cylindrical, coarser towards the base, hollow, surface with inconspicuous longitudinal fibrils,

yellowish brown (#a97e3d), base inflated with abundant white tomentum. 5 % KOH on the surface of the dried pileus

produced no color change.

FIGURE 5. Infundibulicybe phaeocentralis. a. Basidiomata (BJTC FM2899, holotype). b. Basidia. c. Pileipellis. d. Basidiospores. Scale

bars: a = 10 mm, b, c, d = 5 μm. Photos by: Jinzhong Cao, drawings by: Yuxin Zhang.

ZHANG ET AL.

Basidiospores (6–)6.5–8(–8.5) × 4–6 μm, Q=(1.2)1.3–1.6(1.8), Qm=1.4±0.1, ellipsoidal to broadly ellipsoidal in

face view, lacrymoid in lateral view, hyaline, smooth, inamyloid, cyanophilous, color not changing in KOH. Basidia

25–40 × 6–7.5 μm, clavate, hyaline, usually 4-spored, sometimes 2-spored, sterigmata 2.5–5 μm long. Cystidia

absent. Lamellar trama regular; hyphae 4–10 µm wide, colorless, hyaline, cylindrical. Pileipellis a cutis composed of

repent, cylindrical, subparallel, septate, thin-walled hyphae containing yellowish brown pigment 4.5–11.5 μm diam.

Stipitipellis a cutis composed of subparallel cylindrical hyphae 4–10 μm diam. Clamp connections present in all parts

of the basidiomata.

Ecology and distribution:—on soil of a mixed forest of Larix gmelinii var. principis-rupprechtii and Picea

asperata, only known from Shanxi Province, northern China.

Additional specimens examined:—CHINA. Shanxi Province, Ningwu County, Guancen Mountain, on soil, in

mixed forest of Larix gmelinii var. principis-rupprechtii and Picea asperata, 14 August 2017, H. Liu 1-LH170731

(BJTC FM402); CHINA. Shanxi Province, Ningwu County, on soil, in mixed forest of Larix gmelinii var. principis-

rupprechtii and Picea asperata, 5 July 2022, H. Liu LH1563 (BJTC FM2067); CHINA. Shanxi Province, Ningwu

County, Luya Mountain, elev. 2,293m, 111°58’17’’E, 38°47’58’’N, 16 August 2023, H.Y. Fu, H.M. Ji MS270 (BJTC

FM3215).

FIGURE 6. Basidiomata of Infundibulicybe species. a–c. I. alkaliviolascens (BJTC FM1653, BJTC FM1658, BJTC FM3396). d–f. I.

gibba (BJTC FM682, BJTC FM757, BJTC FM1706). g–j. I. hongyinpan (BJTC FM187, BJTC FM216, HSA391, BJTC FM394). k. I. rufa

(BJTC FM562). l–m. I. squamulosa (BJTC FM2451, BJTC FM2452). Scale bars: a–m = 1cm. Photos by: Hong Liu, Jinzhong Cao.

Notes:—Infundibulicybe phaeocentralis formed an isolated position in our phylogenetic trees (Figs. 1, 2), which

strongly supported its uniqueness. The new species is very similar to the samples of I. rufa that are also characterised

by a red-brown cap (Zhao et al. 2016). We also recognized the occurrence of I. rufa in the northern region of Shanxi

Province in this study. However, most of the Shanxi samples of I. rufa supported by our molecular analysis clearly

lack the red-brown in the cap, and are, instead, brown or yellow-brown, and sometimes a shade of purple-brown (Fig.

6, k). Thus, both of the two species are easily separated from each other by the color of the basidiomata in the area of

Shanxi Province. Infundibulicybe squamulosa, originally described from Europe, and confirmed from China before

this study (He & Yang 2023), is another species similar to Infundibulicybe phaeocentralis in morphology, and this

species is also distributed in Shanxi Province (Fig. 6, l–m). Actually, the two species are impossible to separate from

each other by morphology alone based on the examination of the Shanxi samples in this study. However, I. squamulosa

has been recorded only in the forest of pure Pinus tabuliformis in the lowland area in Shanxi Province, whereas I.

phaeocentralis is exclusively found in the mixed forest of Picea sp. and Larix sp. in alpine or subalpine regions in

this province. Infundibulicybe bispora and I. gibba are phylogenetically related to I. phaeocentralis according to the

multilocus phylogeny (Fig. 2). Morphologically, Infundibulicybe bispora can be distinguished from I. phaeocentralis

by its white lamellae, mostly 2-spored basidia, longer sterigmata (4–10 μm) and longer basidiospores (6–11.5 × 4–6

μm) (He & Yang 2023). I. gibba is differentiated by its tan, pinkish tan, or flesh-colored pileus, whitish, off-white, or

very pale stipe and smaller basidiospores (5–9 × 3.5–6 μm) (Harmaja 2003).

Discussion

In this study, seven species of Infundibulicybe have been recognized from Shanxi Province, northern China (Fig. 1).

Besides the two new species described in this paper, the other species are Infundibulicybe alkaliviolascens, I. gibba, I.

hongyinpan, I. rufa. and I. squamulosa. Both I. alkaliviolascens and I. gibba are the most frequently encountered and

widely distributed species in Shanxi Province. They occur from the lowland of the south to the high mountainous area

of the north in this province, and from early summer to late fall. They can grow under both conifer and broad-leaved

trees. Both I. hongyinpa and I. rufa are also often found, and grow under a mixed forest of Larix and Picea but only in

the mountainous area in the northern region where the climate is normally colder. I. squamulosa is a rarely collected

species, currently observed from the central area of this province and only grows in the forest of Pinus tabuliformis.

The color of the basidiomata of Infundibulicybe species varies greatly according to our observations, which leads

to misidentification of species in the field. For example, the surface of the pileus and stipe of I. alkaliviolascens

are usually yellow-brown, but white basidioma and basidioma with a white stipe are not uncommon in this area.

Another example is that the stipe of I. gibba is usually pale, and it is an important feature to differentiate it from I.

alkaliviolascens in the field, but basidiomata with a yellow-brown stipe are also observed. We also noted that white I.

gibba was recorded by He & Yang (2023). Infundibulicybe rufa, a species originally described from Sichuan Province

of southwestern China, is characterised by the red-brown color of both the pileus and the stipe (Zhao et al. 2016).

However, this species showed a wide range of colors in Shanxi Province, ranging from red-brown to pale yellow-

brown and even presents with purple tints. In general, in Shanxi Province, I. hongyinpan is easily recognized as it has

extremely large basidiomata, while the other six species of Infunbulicybe are nearly the same in their small to medium

basidiomata, and actually they are often confused with each other in the field although the color combination of the

pileus and stipe can be used to tell them apart. Consequently, an accurate examination of the species of Infudibulicybe

has to be accompanied with DNA analysis.

Species of Infundibulicybe have been traditionally harvested for food in the northern region of Shanxi Province by

locals for many hundreds of years, particularly among the people living around Wutai and Guancen Mountain, which

is another reason for investigating the taxonomy of this genus.

Key to the species of Infundibulicybe from Shanxi Province of northern China

1. Pileipellis with intracellular pigment only .........................................................................................................................................2

1. Pileipellis with incrusting pigment .....................................................................................................................................................3

2. Pileus brick red to reddish brown ..............................................................................................................................I. phaeocentralis

2. Pileus brownish flesh to pale brown ..................................................................................................................................... I. confusa

3. Pileipellis with faint incrusting pigment .................................................................................................................................. I. gibba

3. Pileipellis with conspicuous incrusting pigment ................................................................................................................................4

4. Pileus finely scaly .......................................................................................................................................................... I. squamulosa

ZHANG ET AL.

4. Pileus not scaly ...................................................................................................................................................................................5

5. Pileal surface produces a positive reaction to KOH ................................................................................................I. alkaliviolascens

5. Pileal surface produces a negative reaction to KOH ..........................................................................................................................6

6. Pileipellis an ixocutis ..................................................................................................................................................................I. rufa

6. Pileipellis a cutis ............................................................................................................................................................ I. hongyinpan

Acknowledgements

The study was supported by the National Natural Science Foundation of China (No. 32370010). We thank the two

anonymous reviewers for their suggestions and corrections to improve this paper.

References

Akata, I., Kalyoncu, F., Solak, M.H. & Kalmış, E. (2012) Growth of mycelium of three ectomycorrhizal macrofungi, Infundibulicybe

geotropa, Tricholoma anatolicum and Lactarius deliciosus in culture media containing various carbon sources. African Journal of

Microbiology Research 6: 3042–3046.

https://doi.org/10.5897/AJMR12.196

Alfaro, M.E., Zoller, S. & Lutzoni, F. (2003) Bayes or bootstrap? A simulation study comparing the performance of Bayesian Markov

chain Monte Carlo sampling and bootstrapping in assessing phylogenetic confidence. Molecular Biology & evolution 20: 255–266.

https://doi.org/10.1093/molbev/msg028

Ali, M., Khan, J., Bashir, H., Niazi, A.R., Sher, H. & Khalid, A.N. (2020) Infundibulicybe macrospora sp. nov. a new and noteworthy

species from Himalayan moist temperate forests of Pakistan. Phytotaxa 452: 268–277.

https://doi.org/10.11646/phytotaxa.452.4.2

Alvarado, P., Moreau, P.A., Dima, B., Vizzini, A., Consiglio, G., Moreno, G., Setti, L., Kekki, T., Huhtinen, S., Liimatainen, K. &

Niskanen, T. (2018a) Pseudoclitocybaceae fam. nov. (Agaricales, Tricholomatineae), a new arrangement at family, genus and species

level. Fungal Diversity 90: 109–133.

https://doi.org/10.1007/s13225-018-0400-1

Alvarado, P., Moreau, P.A., Sesli, E., Khodja, L.Y., Contu, M. & Vizzini, A. (2018b) Phylogenetic studies on Bonomyces (Tricholomatineae,

Agaricales) and two new combinations from Clitocybe. Cryptogamie Mycologie 39: 149–168.

https://doi.org/10.7872/crym/v39.iss2.2018.149

Bellù, F. (2012) Clitocybe alkaliviolascens is now an Infundibulicybe: its recombination with additional distribution data. Bresadoliana

1: 3–7.

Binder, M. & Hibbett, D.S. (2003) Oligonucleotides. Worcester (MA): The Education Network of Clark University. Available from: http://

www2.clarku.edu/faculty/dhibbett/Protocols_Folder/Primers/Primers.htm (accessed: 18 Sep 2003).

Binder, M., Larsson, K.H., Matheny, P.B. & Hibbett, D.S. (2010) Amylocorticiales ord. nov. and Jaapiales ord. nov.: Early diverging clades

of Agaricomycetidae were dominated by corticioid forms. Mycologia 102: 865–880.

https://doi.org/10.3852/09-288

Bjorbækmo, M.F.M., Carlsen, T., Brysting, A., Vrålstad, T., Høiland, K., Ugland, K.I., Geml, J., Schumacher, T. & Kauserud, H. (2010)

High diversity of root associated fungi in both alpine and arctic Dryas octopetala. BMC Plant Biology 10: 1–12.

https://doi.org/10.1186/1471-2229-10-244

Carrière, E.A. (1867) Traite General des Coniferes, ed. 2: 510.

Castresana, J. (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology

and Evolution 17: 540–552.

https://doi.org/10.1093/oxfordjournals.molbev.a026334

Engler, H.G.A. (1926) Natürlichen Pflanzenfamilien, ed. 2. Vol. 13. pp. 327.

Gardes, M. & Bruns, T.D. (1993) ITS primers with enhanced specificity for Basidiomycetes – application to the identification of

mycorrhizae and rusts. Molecular Ecology 2: 113–118.

https://doi.org/10.1111/j.1365-294X.1993.tb00005.x

Geml, J., Timling, I., Robinson, C.H., Lennon, N., Nusbaum, H.C., Brochmann, C., Noordeloos, M.E. & Taylor, D.L. (2012) An arctic

community of symbiotic fungi assembled by long-distance dispersers: phylogenetic diversity of ectomycorrhizal basidiomycetes in

Svalbard based on soil and sporocarp DNA. Journal of Biogeography 30: 74–88.

https://doi.org/10.1111/J.1365-2699.2011.02588.X

Giusti, A., Ricci, E., Gasperetti, L., Galgani, M., Polidori, L., Verdigi, F., Narducci, R. & Armani, A. (2021) Building of an Internal

Transcribed Spacer (ITS) Gene Dataset to Support the Italian Health Service in Mushroom Identification. Foods 10: 1–20.

https://doi.org/10.3390/foods10061193

Harmaja, H. (2003) Notes on Clitocybe s. lato (Agaricales). Annales Botanici Fennici 40: 213–218.

Harmaja, H. (1969) The genus Clitocybe (Agaricales) in Fennoscandia. Karstenia 10: 5–168.

https://doi.org/10.29203/ka.1969.62

He, Z.M. & Yang, Z.L. (2022) The genera Bonomyces, Harmajaea and Notholepista from Northwestern China: two new species and a

new record. Mycological progress 21: 1–16.

https://doi.org/10.1007/s11557-022-01786-0

He, Z.M. & Yang, Z.L. (2023) A contribution to the knowledge of the genus Infundibulicybe (Tricholomatineae, Agaricales) in China: Two

new species and five redescribed taxa. Mycologia 115: 693–713.

https://doi.org/10.1080/00275514.2023.2226893

He, Z.M., Chen, Z.H., Bau, T., Wang, G.S. & Yang, Z.L. (2023) Systematic arrangement within the family Clitocybaceae (Tricholomatineae,

Agaricales): phylogenetic and phylogenomic evidence, morphological data and muscarine-producing innovation. Fungal Diversity

123: 1–47.

https://doi.org/10.1007/s13225-023-00527-2

Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic

Biology 42: 182–192.

https://doi.org/10.1093/sysbio/42.2.182

Ishaq, M., Khan, M.B., Ullah, S., Fiaz, M. & Khalid, A.N. (2019) Infundibulicybe kotanensis sp. nov. (Tricholomataceae), a new species

from Buner, Pakistan. Phytotaxa 418: 195–202.

https://doi.org/10.11646/phytotaxa.418.2.4

Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability.

Molecular Biology and Evolution 30: 772–780.

https://doi.org/10.1093/molbev/mst010

Kirk, P.M., Cannon, P.F., Minter, D.W. & Stalpers, J.A. (2008) Dictionary of the fungi. Oxford University Press, Wallingford, UK, 341

pp.

Kretzer, A.M. & Bruns, T.D. (1999) Use of atp6 in fungal phylogenetics: an example from the Boletales. Molecular Phylogenetics &

Evolution 13: 483–492.

https://doi.org/10.1006/mpev.1999.0680

Liu, H., Guo, S. & Fan, L. (2018) Infundibulicybe hongyinpan sp. nov., a well-known edible fungus from Shanxi based on both nrDNA-

ITS and morphological analysis. Journal of Shanxi University 42: 275–280.

https://doi.org/10.13451/j.cnki.shanxi.univ (nat.sci.).2018.03.16.007

Maeta, K., Ochi, T., Tokimoto, K., Shimomura, N., Maekawa, N., Kawaguchi, N., Nakaya, M., Kitamoto, Y. & Aimi, T. (2008) Rapid

Species Identification of Cooked Poisonous Mushrooms by Using Real-Time PCR. Applied and Environmental Microbiology 74:

3306–3309.

https://doi.org/10.1128/AEM.02082-07

Mast. (1903) The Gardeners’ chronicle. 33: 133.

Mast. (1906) The Journal of the Linnean Society. Botany 37: 419.

Matheny, P.B. (2005) Improving phylogenetic inference of mushrooms with RPB1 and RPB2 nucleotide sequences (Inocybe; Agaricales).

Molecular Phylogenetics & Evolution 35: 1–20.

https://doi.org/10.1088/1126-6708/2004/10/014

Matheny, P.B., Curtis, J.M., Hofstetter, V., Aime, M.C., Moncalvo, J.M., Ge, Z.W., Yang, Z.L., Slot, J.C., Ammirati, J.F., Baroni, T.J.,

Bougher, N.L., Hughes, K.W., Lodge, D.J., Kerrigan, R.W., Seidl, M.T., Aanen, D.K., DeNitis, M., Daniele, G.M., Desjardin, D.E.,

Kropp, B.R., Norvell, L.L., Parker, A., Vellinga, E.C., Vilgalys, R. & Hibbett, D.S. (2006) Major clades of Agaricales: a multilocus

phylogenetic overview. Mycologia 98: 982–995.

https://doi.org/10.3852/mycologia.98.6.982

Matheny, P.B., Liu, Y.J., Ammirati, J.F. & Hall, B.D. (2002) Using RPB1 sequences to improve phylogenetic inference among mushrooms

(Inocybe, Agaricales). American Journal of Botany 89: 688–698.

https://doi.org/10.3732/ajb.89.4.688

Murrill, W.A. (1915) The genus Clitocybe in North America. Mycologia 7: 256–283.

https://doi.org/10.1080/00275514.1915.12021718

Nylander, J.A.A. (2004) MrModeltest 2.2. Computer software distributed by the University of Uppsala. Sweden: Evolutionary Biology

Centre.

ZHANG ET AL.

Osmundson, T.W., Robert, V.A., Schoch, C.L., Baker, L.J., Smith, A., Robich, G., Mizzan, L. & Garbelotto, M.M. (2013) Filling Gaps

in Biodiversity Knowledge for Macrofungi: Contributions and Assessment of an Herbarium Collection DNA Barcode Sequencing

Project. PLOS ONE 8: 1–8.

https://doi.org/10.1371/journal.pone.0062419

Page, R.D. (2001) TreeView. Glasgow University, Glasgow.

Quélet, L. (1872) Les Champignons du Jura et des Vosges. Mémoires de la Société d'Émulation de Montbéliard 5: 43–332.

Rehner, S.A. & Buckley, E. (2005) A Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: evidence for cryptic

diversification and links to Cordyceps teleomorphs. Mycologia 97: 84–98.

https://doi.org/10.3852/mycologia.97.1.84

Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–

1574.

https://doi.org/10.1093/bioinformatics/btg180

Sargent, C.S. (1914) Plantae Wilsonianae 2: 28.

Singer, R. (1956) New genera of fungi. VII. Mycologia 48: 719–727.

https://doi.org/10.1080/00275514.1956.12024585

Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30:

1312–1313.

https://doi.org/10.1093/bioinformatics/btu033

Staude, F. (1857) Die Schwämme Mitteldeutschlands, in besondere des Herzogthums. pp. 1–150.

Tian, H.M, Liu, T.Z., Li, G.L. & Bateer. (2015) Morphology and ITS sequencing analysis of Russulaceae in Inner Mongolia. Journal of

Inner Mongolia University (Natural Science Edition) 46: 520–526.

https://doi.org/10.13484/j.nmgdxxbzk.20150512

Tomáš, V., Miroslav, K., Lucia, Ž., Tomáš, Z. & Petr, B. (2016) The rpb2 gene represents a viable alternative molecular marker for the

analysis of environmental fungal communities. Molecular Ecology Resources 16: 388–401.

https://doi.org/10.1111/1755-0998.12456

Vilgalys, R. & Hester, M. (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several

Cryptococcus species. Journal of Bacteriology 172: 4238–4246.

https://doi.org/10.1128/jb.172.8.4238-4246.1990

Vizzini, A., Contu, M., Musumeci, E. & Ercole, E. (2011) A new taxon in the Infundibulicybe gibba complex (Basidiomycota, Agaricales,

Tricholomataceae) from Sardinia (Italy). Mycologia 103: 203–208.

https://doi.org/10.3852/10-137

Vizzini, A., Alvarado, P., Consiglio, G., Marchetti, M. & Xu, J. (2024) Family matters inside the order Agaricales: systematic reorganization

and classification of incertae sedis clitocyboid, pleurotoid and tricholomatoid taxa based on an updated 6-gene phylogeny. Studies

in Mycology 107: 67–148.

https://doi.org/10.3114/sim.2024.107.02

Vu, D., Groenewald, M., de Vries, M., Gehrmann, T., Stielow, B., Eberhardt, U., Al-Hatmi, A., Groenewald, J.Z., Cardinali, G., Houbraken,

J., Boekhout, T., Crous, P.W., Robert, V. & Verkley, G.J.M. (2018) Large-scale generation and analysis of filamentous fungal DNA

barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal species and higher taxon delimitation. Studies in

Mycology 92: 135–154.

https://doi.org/10.1016/j.simyco.2018.05.001

Walther, G., Garnica, S. & Weiß, M. (2005) The systematic relevance of conidiogenesis modes in the gilled Agaricales. Mycological

Research 109: 525–544.

https://doi.org/10.1017/S0953756205002868

Wei, T.Z., Li, B.B., Wang, W.J. & Yao, Y.J. (2015) Infundibulicybe alkaliviolascens, a new agaric record of China. Journal of Fungal

Research 13: 284–288.

https://doi.org/10.13341/j.jfr.2014.2063

White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics.

PCR Protocols: A guide to Methods and Applications 18: 315–322.

https://doi.org/10.1016/B978-0-12-372180-8.50042-1

Xu, J.Z., Zhao, W., Yu, X.D., Idrees, M., Li, Y. & Qin, J.C. (2022) Infundibulicybe trachyspora, a new species from northeastern China

based on morphology and molecular phylogeny. Current Microbiology 79: 1–9.

https://doi.org/10.1007/s00284-022-02808-6

Zhao, Q.I., Hao, Y.J., Liu, J.K., Hyde, K.D., Cui, Y.Y., Brooks, S. & Zhao, Y.C. (2016) Infundibulicybe rufa sp. nov. (Tricholomataceae),

a reddish-brown species from southwestern China. Phytotaxa 266: 134–140.

https://doi.org/10.11646/phytotaxa.266.2.7

Supplementary TABLE 2. Infundibulicybe specimens used in molecular phylogenetic analyses and their GenBank accession

numbers (other five gene). Newly generated sequences are in black bold.

Species Voucher Locality LSU TEF1 RPB1 RPB2 ATP6

Clitopaxillus fibulatus KUN-HKAS 115964 China: Gansu OP962073 OP966740 OP966741 OP966742 OP966743

I. alkaliviolascens KUN-HKAS 115938 China: Yunnan OP972477 OQ030273 OQ030303 OQ030335 OQ030366

I. alkaliviolascens KUN-HKAS 115939 China: Xinjiang OP972478 OQ030274 OQ030304 OQ030336 OQ030367

I. alkaliviolascens KUN-HKAS 115940 China: Xinjiang OP972479 OQ030275 OQ030305 OQ030337 OQ030368

I. alkaliviolascens KUN-HKAS 115942 China: Xinjiang OP972480 OQ030276 OQ030306 OQ030338 OQ030369

I. alkaliviolascens KUN-HKAS 115943 China: Yunnan OP972481 OQ030277 OQ030307 OQ030339 OQ030370

I. alkaliviolascens KUN-HKAS 115944 China: Yunnan OP972482 OQ030278 OQ030308 OQ030340 OQ030371

I. bispora KUN-HKAS 115961 China: Jilin OP972506 OQ030300 OQ030332 OQ030363 OQ030395

I. bispora KUN-HKAS 115962 China: Jilin OP972507 OQ030301 OQ030333 OQ030364 OQ030396

I. bispora KUN-HKAS 115963 China: Gansu OP972508 OQ030302 OQ030334 OQ030365 OQ030397

I. confusa (holotype) BJTC FM1281 China: Shanxi PP492520 PP496079 PP496067 PP496073 PP496061

I. confusa BJTC FM1285 China: Shanxi PP492521 PP496080 PP496068 PP496074 PP496062

I. ellipsospora KUN-HKAS 114671 China: Yunnan OP972496 OQ030291 OQ030322 OQ030353 OQ030385

I. ellipsospora KUN-HKAS 70065 China: Yunnan OP972497 OQ030292 OQ030324 OQ030354 OQ030386

I. ellipsospora KUN-HKAS 105353 China: Yunnan OP972498 OQ030293 OQ030329 OQ030355 OQ030387

I. ellipsospora KUN-HKAS 115959 China: Tibet OP972499 OQ030294 OQ030325 OQ030360 OQ030388

I. ellipsospora KUN-HKAS 54633 China: Yunnan OP972500 OQ030289 OQ030330 OQ030356 OQ030389

I. ellipsospora KUN-HKAS 56928 China: Yunnan OP972501 OQ030295 OQ030331 OQ030357 OQ030390

I. ellipsospora KUN-HKAS 56965 China: Yunnan OP972502 OQ030296 OQ030323 OQ030358 OQ030391

I. ellipsospora KUN-HKAS 116251 China: Tibet OP972503 OQ030297 OQ030326 OQ030361 OQ030392

I. ellipsospora KUN-HKAS 57667 China: Yunnan OP972504 OQ030290 OQ030327 OQ030362 OQ030393

I. ellipsospora KUN-HKAS 115960 China: Yunnan OP972505 OQ030298 OQ030328 OQ030359 OQ030394

I. geotropa GLM 45881 Germany AY207154 — DQ067938 — —

I. geotropa ALV4344 — KT122793 — — — —

I. gibba KUN-HKAS 115945 China: Yunnan OP972483 OQ030279 OQ030309 OQ030341 OQ030372

I. gibba KUN-HKAS 80468 China: Yunnan MZ719004 MZ753646 — — —

I. gibba KUN-HKAS 81200 China: Yunnan MZ719005 MZ753647 — — —

I. gibba KUN-HKAS 115946 China: Yunnan OP972484 OQ030280 OQ030312 OQ030343 OQ030373

I. gibba KUN-HKAS 115947 China: Yunnan OP972485 OQ030281 OQ030313 OQ030344 OQ030374

I. gibba KUN-HKAS 115948 China: Yunnan OP972486 OQ030282 OQ030310 OQ030342 OQ030375

I. gibba KUN-HKAS 115949 China: Yunnan OP972487 OQ030283 OQ030311 OQ030345 OQ030376

I. gibba KUN-HKAS 115950 China: Jilin OP972488 OQ030284 OQ030314 OQ030346 OQ030377

I. gibba KUN-HKAS 115951 China: Liaoning OP972489 OQ030285 OQ030315 OQ030347 OQ030378

I. gibba KUN-HKAS 53323 China: Yunnan MZ719006 MZ753648 MZ753621 MZ753630 MZ753638

I. gibba KUN-HKAS 57657 China: Yunnan MZ719007 MZ753649 MZ753622 MZ753631 MZ753639

I. gibba KUN-HKAS 105606 China: Gansu MZ719008 MZ753650 MZ753623 — MZ753640

I. gibba KUN-HKAS 73334 China: Yunnan MZ719009 MZ753651 MZ753624 MZ753632 MZ753641

I. gibba KUN-HKAS 73336 China: Yunnan MZ719010 MZ753652 MZ753625 MZ753633 MZ753642

I. gibba KUN-HKAS 115952 China: Yunnan OP972490 OQ030286 OQ030316 OQ030348 OQ030379

I. gibba KUN-HKAS 115953 China: Yunnan OP972491 OQ030287 OQ030317 OQ030349 OQ030380

I. gibba AFTOL-ID 1508 USA DQ457682 GU187759 DQ447913 DQ472727 —

I. gibba KUN-HKAS 92032 China: Liaoning MZ675574 MZ681874 MZ681885 MZ681895 MZ681904

I. hongyinpan KUN-HKAS 115954 China: Gansu OP972492 OQ067523 OQ030318 — OQ030381

I. hongyinpan KUN-HKAS 115955 China: Tibet OP972493 OQ067524 OQ030319 OQ030350 OQ030382

I. hongyinpan KUN-HKAS 115956 China: Yunnan OP972494 OQ030288 OQ030320 OQ030351 OQ030383

I. hongyinpan KUN-HKAS 105573 China: Gansu MZ719011 MZ753653 MZ753626 MZ753634 MZ753643

I. hongyinpan KUN-HKAS 115957 China: Gansu OP972495 OQ067525 OQ030321 OQ030352 OQ030384

......continued on the next page

ZHANG ET AL.

Supplementary TABLE 2. (Continued)

Species Voucher Locality LSU TEF1 RPB1 RPB2 ATP6

I. phaeocentralis

(holotype) BJTC FM2899 China: Shanxi PP492523 PP496082 PP496070 PP496076 PP496064

I. phaeocentralis BJTC FM3215 China: Shanxi PP492524 PP496084 PP496072 PP496078 PP496066

I. phaeocentralis BJTC FM402 China: Shanxi PP492525 PP496083 PP496071 PP496077 PP496065

I. phaeocentralis BJTC FM2067 China: Shanxi PP492522 PP496081 PP496069 PP496075 PP496063

I. rufa KUN-HKAS 77865 China: Hubei MZ719012 MZ753654 — — —

I. rufa KUN-HKAS 57811 China: Tibet MZ719013 MZ753655 MZ753627 MZ753635 MZ753644

I. squamulosa CBS:128.63 France MH869837 MH858237 MH869837

I. trachyspora HMJU 744 China: Jilin MW880692 — OL677427 OL677428 —

I. trachyspora KUN-HKAS 78801 China: Hubei MZ719014 MZ753656 MZ753628 MZ753636 MZ753645

Omphalina pyxidata AMB:19294 Italy OR863534 OR828282

Omphalina pyxidata AMB:19295 Italy OR863535 OR828283

Pseudoclitocybe

cyathiformis KUN-HKAS 73390 China: Yunnan MZ719003 MZ681907 MZ753620 MZ753629 MZ753637

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