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Research Article
Iugisporipsathyra reticulopilea gen. et sp. nov. (Agaricales, Psathyrellaceae) from tropical China produces unique ridge-ornamented spores with an obvious suprahilar plage
expand article infoSheng-Nan Wang§, Yu-Guang Fan|, Jun-Qing Yan§
‡ Jiangxi Agricultural University, Nanchang, China
§ Fujian Academy of Agricultural Sciences; National and Local Joint Engineering Research Center for Breeding & Cultivation of Features Edible Mushrooms, Fuzhou, China
| Hainan Medical University, Haikou, China
Open Access

Abstract

Iugisporipsathyra, a new psathyrelloid genus from tropical red soil of China, is established with I. reticulopilea as the type species. The new genus is characterised by basidiomata psathyrelloid, pileus rugose to appearing reticulate ridged, covered by persistent, but inconspicuous villus, pleurocystidia absent and ridge-ornamented spores with an obvious suprahilar plage. The genus is unique amongst Psathyrellaceae in producing ridge-ornamented spores with an obvious suprahilar plage and forms a distinct lineage within Psathyrellaceae, based on the Maximum Likelihood and Bayesian Inference analyses of a combined three-gene sequence dataset (ITS, LSU and β-tub). Full descriptions and photographs of the new genus and species are presented.

Keywords

Basidiomycete, fungal phylogeny, taxonomy

Introduction

The Psathyrellaceae Vilgalys, Moncalvo & Redhead was established in 2001, based on the type genus Psathyrella (Fr.) Quél. by Vilgalys and Redhead (Redhead et al. 2001). More than 1300 names within the family, including synonyms and subspecies, are listed in Index Fungorum (http://www.indexfungorum.org). Species of Psathyrellaceae are cosmopolitan and often grow on decaying logs, woody debris, humus or soil, in woodlands, lawns or bogs and can have either broad or specific substrate relationships (Kirk et al. 2008).

Traditionally, the family included two types of species: psathyrelloid species and coprinoid species. During the classic period of morphological research, Fries (1838) classified the psathyrelloid species to Agaricus L. trib. Psathyrella Fr. Quélet (1872) promoted this group to the rank of genus. Psathyrella was finally accepted after the transfer of Drosophila Quél. species and emendations by Singer (1951,1975). Subsequently, Kits van Waveren (1985) removed the species with warty spores from Psathyrella and treated these as the genus Lacrymaria Pat. Although the boundaries of the genus were disputed, most researchers agreed that the psathyrelloid species should be classified in Coprinaceae R.Heim ex Pouzar subfamily Psathyrelloideae (Kuhner) Singer (Hawksworth et al. 1983; Kirk et al. 2001). During this same period, the coprinoid species were classified in Coprinus Pers. (Coprinaceae subfamily Coprinoideae Henn.) (Hawksworth et al. 1995; Kirk et al. 2001). Coprinus was circumscribed by Persoon (1797). However, Fries (1821) did not recognise the genus in his monograph Systema Mycologicum and classified the species in Agaricus. However, in his subsequent monograph Epierisis systematis Mycologici, Fries discarded his previous classification and again placed the coprinoid species in the independent genus Coprinus (Fries, 1838).

Although morphological studies provide abundant support for recognition of Psathyrellaceae, morphological data are inadequate to conclusively resolve the systematic relationships amongst the constituent genera and species. When the works of Hopple and Vilgalys (1999) and Redhead et al. (2001) were published, it became apparent that molecular biology techniques would profoundly alter the classical systematics of psathyrelloid species and coprinoid species. Based on these studies, Coprinus was split into four genera (Coprinellus P.Karst., Coprinopsis P.Karst., Coprinus and Parasola Redhead, Vilgalys & Hopple) (Redhead et al. 2001), restraining the generic name Coprinus to a small group centred on the type species Coprinu comatus (O.F.Müll.) Pers., which is now classified in the Agaricaceae Chevall. The other three genera, together with Psathyrella and Lacrymaria, were incorporated into the newly-established Psathyrellaceae. In 2015, Psathyrella, as a paraphyletic group, was also split, with the establishment of the segregate genera Cystoagaricus Singer emend. Örstadius & E.Larss., Homophron (Britzelm.) Örstadius & E.Larss., Kauffmania Örstadius & E.Larss. and Typhrasa Örstadius & E.Larss. (Örstadius et al. 2015). In 2020, Candolleomyces D.Wächt. & A.Melzer, Britzelmayria D.Wächt. & A.Melzer and Olotia D.Wächt. & A.Melzer were separated from Psathyrella, Punjabia D.Wächt. & A.Melzer and Tulosesus D.Wächt. & A.Melzer were separated from Coprinellus, Narcissea D.Wächt. & A.Melzer was segregated from Coprinopsis and Hausknechtia D.Wächt. & A.Melzer was erected for Galerella floriformis Hauskn. (Wächter and Melzer 2020). Heteropsathyrella T.Bau & J.Q.Yan was established in 2021, based on the new species He. macrocystidia T.Bau & J.Q.Yan (Bau and Yan 2021a). Thus, the main systematic framework of Psathyrellaceae has been confirmed. In addition, Ozonium Link and Hormographiella Guarro & Gené, formerly members of the Psathyrellaceae, were established to accommodate the conidial anamorphs of certain species, now classified in Coprinellus (Nagy et al. 2013). Gasteroagaricoides D.A.Reid and Macrometrula Donk & Singer, two genera that, to date, have not been included in phylogenetic analyses, are retained in the Psathyrellaceae. There were 19 genera, in total, in the Psathyrellaceae before the new taxon we discovered was added.

From 2015, we initiated a study of Chinese psathyrelloid species and described 15 new taxa (Yan and Bau 2017, 2018a,b; Yan et al. 2019; Bau and Yan 2021a,b; Wang et al. 2021). By chance, we collected a psathyrelloid species with a reticulate-ridged pileus, that was reminiscent of Pluteus thomsonii (Berk. & Broome) Dennis, on the roadside in tropical China. After examining the micromorphology of the specimens, we observed that it produced ridge-ornamented spores with an obvious suprahilar plage. Surprisingly, phylogenetic analysis of molecular data revealed that it belonged to the Psathyrellaceae. Although abundant genera and species are recognised in the Psathyrellaceae, the majority of species have smooth spores. Verrucous spores have been observed only in Lacrymaria. Rough spores have been observed in Coprinopsis, Coprinellus and Psathyrella, but are extremely rare. Thus, the specimens are unique amongst Psathyrellaceae in producing ridge-ornamented spores with an obvious suprahilar plage. On the basis of our morphological and phylogenetic analyses, the specimens are described herein as a new species and a new genus is erected to accommodate the new species.

Materials and methods

Morphological studies

Macroscopic descriptions and habitat details were based on detailed field notes of fresh basidiomata and photos. The location of the collection point is marked on the map (Fig. 1). Colour codes follow the Methuen Handbook of Colour (Kornerup and Wanscher 1978). Microscopic structures were observed and measured from dried specimens mounted in water, 5% potassium hydroxide (KOH), 10% ammonium hydroxide (NH4OH) or Melzer’s Reagent. Congo red was used as a stain when necessary (Horak 2005). A minimum of 100 basidiospores, basidia and cystidia from seven basidiomata (three collections) were randomly measured using an Olympus BX53 microscope. Detailed observations of spores were made by SEM. The measurements and Q values are recorded as (a)b–c(d), in which “a” is the lowest value, “b–c” covers a minimum of 90% of the values and “d” is the highest value. “Q” stands for the ratio of length and width of a spore (Bas 1969; Yu et al. 2020). Specimens were deposited in the Herbarium of Fungi, Jiangxi Agricultural University (HFJAU).

Figure 1. 

Map showing the location of the collection site of the specimens (red triangle).

DNA extraction and sequencing

DNA was extracted from dried specimens with the NuClean Plant Genomic DNA kit (CWBIO, China) (Ge et al. 2021; Na et al. 2022). Three regions (ITS, LSU and β-tub) were selected for the study and were amplified using the primer pairs by ITS1/ITS4 (White et al. 1990), LR0R/LR7 (Hopple and Vilgalys 1999) and B36f/B12r (Nagy et al. 2011), respectively. PCR was performed using a touchdown programme for all regions: 5 min at 95 °C; 1 min at 95 °C; 30 s at 65 °C (add -1 °C per cycle); 1 min at 72 °C; cycle 15 times; 1 min at 95 °C; 30 s at 50 °C; 1 min at 72 °C; cycle 20 times; 10 min at 72 °C (Bau and Yan 2021a). The sequencing was performed by Qing Ke Biotechnology Co. Ltd. (Wuhan City, China).

Data analyses

A total of 221 nucleotide DNA (ITS, LSU and β-tub) sequences representing 93 taxa were used in subsequent analyses. Details are presented in Table 1. Some species of Agaricaceae, Mythicomycetaceae Vizzini, Consiglio & M. Marchetti and Nidulariaceae Dumort. were chosen as outgroup taxa according to the results of Zhao et al. (2017) and Vizzini et al. (2019). ITS, LSU and β-tub sequence datasets were separately aligned on the MAFFT online server (Katoh et al. 2019). Bayesian Inference (BI) and Maximum Likelihood (ML) phylogenetic analyses of the aligned concatenated dataset were respectively carried out in MrBayes v.3.2.7a and IQTREE v.2.1.2 (Nguyen et al. 2014) via the CIPRES web portal. For the BI analyses, optimal evolutionary models were selected using PartitionFinder2 (Lanfear et al. 2017) with the greedy algorithm and the AICc criterion. Four Monte Carlo Markov chains were run for 2 million generations, sampling every 100th generation, with the first 25% of trees discarded as burn-in (Ronquist et al. 2012). For the ML analysis, models of sequence evolution were assessed in IQ-Tree prior to the analysis. The ML analysis was conducted using the ultrafast bootstrap option with 1,000 replicates and allowing partitions to have different seeds (--p). A nexus file contains alignment sequence and original tree of ML and Bayes is deposited in Suppl. material 1.

Table 1.

Sequences used in this study.

Taxon Voucher ITS LSU β-tub
Britzelmayria multipedata LÖ237-04 KC992888 KC992888 KJ664867
B. supernula LÖ250-04 KC992867 KC992867 KJ664849
Candolleomyces eurysporus GLM-F126263 Type MT651560 MT651560 MW369460
C. subcacao HMJAU37807 Type MW301064 MW301092 MW314063
C. subminutisporus HMJAU37801 Type MW301066 MW301094 MW314065
C. subsingeri HMJAU37913 Type MG734725 MW301098 MW314068
Coprinellus andreorum CS1247 Type MW621497 MW621007
C. aureogranulatus CBS973.95 GQ249274 GQ249283 GQ249258
C. aureogranulatus CBS753.96 Isotype MH862611
C. curtus NL-2339 FM878016 FM876273 FN396281
C. deminutus NL-0761 JN159572 JN159592 JN159636
C. disseminatus NL-2337 FM878017 FM876274 FN396282
C. domesticus NL-1292 FN396102 HQ847132 FN396330
C. silvaticus LÖ172-08 KC992943 KC992943 KJ664911
Coprinopsis babosiae NL-4139 Type FN396128 FN396177 FN396352
C. calospora CBS612.91 Type GQ249275 GQ249284 GQ249259
C. cortinatus NL-1621 FN396121 FN396171 FN396346
C. musae JV06-179 Type KC992965 KC992965 KJ664920
C. musae JV06-180 KC992966 KC992966 KJ664921
C. semitalis CBS291.77 Type GQ249278 GQ249287 GQ249262
C. udicola AM1240 Type KC992967 KC992967 KJ664922
C. villosa NL-1758 Type JN943128 JQ045877 HQ847173
Cystoagaricus hirtosquamulosa Ramsholm800927 KC992945 KC992945
C. olivaceogrisea WK8/15/63-5 Type KC992948 KC992948
C. silvestris LÖ191-92 KC992949 KC992949
C. squarrosiceps Laessoe44835 KC992950
C. strobilomyces E.Nagasawa9740 AY176347 AY176348
Hausknechtia floriformis WU22833 Type JX968254 JX968371
Heteropsathyrellamacrocystidia HMJAU37803 MW405101 MW413358
H. macrocystidia HMJAU37802 Type MW405102 MW413359 MW410997
Homophron camptopodum 1997/956 KC992956 KC992956
H. cernuum LÖ134-98 DQ389726 DQ389726 KJ664915
H. crenulata W-K8/10/64-5 Type KC992957
H. spadiceum Enderle Epitype DQ389729 DQ389729
Iugisporipsathyra reticulopilea HFJAU1352 Type ON207138 ON207137 ON210974
I. reticulopilea HFJAU3181 ON207139 ON210975
I. reticulopilea HFJAU3182 ON207140 ON210976
Kauffmania larga LÖ223-90 DQ389694 DQ389694 KJ664912
K. larga LAS97-054 DQ389695 DQ389695
Lacrymaria glareosa LAS06-019 KC992954 KC992954 KJ664914
L. hypertropicalis Guzman29585 Type KC992958 KC992958 KJ664916
L. lacrymabunda EL70-03 DQ389724 DQ389724
L. pyrotricha CBS573 GQ249280 GQ249289 GQ249264
L. rigidipes LAS00-081 KC992953 KC992953 KJ664913
L. subcinnamomea Smith16957 Type KC992951 KC992951
Narcissea cordispora SFSUDEH2073 AY461827
N. cordispora LÖ41-01 DQ389723 KJ664910
N. patouillardi NL-1687 FM878009 FM876265 FN396257
Olotia codinae GLM-F112430 Type MG696611 MG674714
Parasola auricoma NL-0087 JN943107 JQ045871 FN396252
P. conopilea LÖ186-02 Neotype DQ389725 DQ389725
P. kuehneri Ulje31-V-1987 Type KY928608 KY928633
P. lactea NL-0466 FM163192 FM160717 FN396254
P. misera NL-0280 Neotype FM163210 FM160699
P. ochracea NL-3621 Type JN943134 JQ045875
P. parvula CAL1667 Type NR_160509 NG064556
P. plicatilis NL-0295 FM163216 FM160693 FN396253
P. plicatilis NL-0075a Epitype NR_171786 NG075167
P. psathyrelloides CAL1753 Type MK682756 MK682754
Psathyrella amygdalinospora HMJAU37952 Type MW405104 MW413361 MW410991
P. amygdalinospora HMJAU57044 MW405105
P. fagetophila LÖ210-85 (M) Type KC992902 KC992902 KJ664879
P. fennoscandica HMJAU37918 MG734723 MW413365 MW410993
P. fennoscandica LÖ484-05 Type KC992903 KC992903 KJ664881
P. noli-tangere LÖ83-03 Neotype DQ389713 DQ389713 KJ664890
P. seminuda Smith34091 (MICH) Type KC992907 KC992907
P. warrenensis Smith70162 (MICH) Type KC992906 KC992906
Punjabia pakistanica MEL2382843 KP012718 KP012718
P. pakistanica LAH35323 Type MH366736
Tulosesus canistri Walleyn877 Isotype HQ846985 HQ847142
T. cinereopallidus NL-0177 Type HQ847001 HQ847090 HQ847149
T. fuscocystidiatus NL-2720 Type HQ846977 HQ847064 HQ847152
T. hiascens NL-2536 FM878018 FM876275 FN396284
T. pseudoamphithallus Ulje1288 Type HQ846973 HQ847059
T. radicellus NL-3168 Type GU227719 HQ847077 GU227737
T. sassii NL-1495 FN396101 FN396155 FN396329
Typhrasa gossypina Schumacher024 KC992946 KC992946
T. nanispora Barta980706 Type KC992947 KC992947
T. polycystis HFJAU1454 Type MW466538 MW466544
T. rugocephala HFJAU1467 Type MW466541 MW466546
Outgroup
Coprinus comatus AFTOL_ID_626 AY854066 AY635772
Crucibulum laeve REGCrul1/DSH96-02 DQ486696 AF336246
Cyathus striatus DSH96-028/Cyst1/DSH96-001 DQ486697 AF336247
Lepiota cristata ZRL20151133 LT716026 KY418841
Leucocoprinus fragilissimus ZRL20151466 LT716029 KY418844
Lycoperdon ericaeum ZRL20151498 LT716030 KY418845
Macrolepiota dolichaula xml2013058 LT716021 KY418836
Mycocalia denudata AFTOL2018/CBS494.85 DQ911596 DQ911597
Mythicomyces corneipes AFTOL-ID972 DQ404393 AY745707
M. corneipes KB51 KY648897
Nidula niveotomentosa AFTOL1945/CBS250.84 DQ917654 DQ986295
Stagnicola perplexa AH25260 Holotype MK351609 MK353793
S. perplexa AH25282 Paratype MK351610 MK353794

Results

Phylogenetic analysis

Based on the BLAST results, the new species were found sharing less than 90.82% (ITS), 97.66% (LSU) and 87.03% (β-tub) similarity with the known species. The aligned concatenated dataset comprised 2,591 characters (ITS 835 bp, LSU 1338 bp and β-tub 418 bp), of which 983 sites were variable and 757 were parsimony informative. The best-fit evolutionary models used for the phylogenetic analyses were as follows: for the BI analysis, GTR + I + G for ITS and LSU and TIM + I + G for β-tub; and for the ML analysis, TIM2 + F + I + G4 for ITS, GTR + F + R4 for LSU and HKY + F + I + G4 for β-tub. The log-likelihood of the ML consensus tree was –27426.323 and the average standard deviation of split frequencies was less than 0.01 after 1,115,000 generations in the BI analysis. In the resulting trees, clades with a Bayesian posterior probability (BI-PP) ≥ 0.95 and ML bootstrap support (ML-BP) ≥ 75% were considered to be well supported.

As shown in the BI tree in Fig. 2, all taxa of Psathyrellaceae formed a well-supported monophyletic lineage (BI-PP = 1; ML-BP = 100%). Within Psathyrellaceae, 18 major supported clades with a high statistical support value (BI-PP ≥ 0.95, ML-BP ≥ 75%) represented a total of 17 (out of 19) known genera and a new genus. Iugisporipsathyra formed a distinct lineage (BI-PP = 1; ML-BP = 100%) clearly separated from currently recognised genera.

Figure 2. 

Phylogeny generated by Bayesian Inference, based on a concatenated sequence dataset for three nuclear DNA regions (ITS, LSU and β-tub). The tree was rooted with Agaricaceae spp., Mythicomycetaceae spp. and Nidulariaceae spp. Bayesian Inference posterior probabilities (BI-PP) ≥ 0.95 and Maximum Likelihood bootstrap percentages (ML-BP) ≥ 75% are shown as PP/BP at relevant nodes. (black circle) indicates newly-described taxa.

Taxonomy

Iugisporipsathyra J.Q. Yan, Y.G. Fan & S.N. Wang, gen. nov.

MycoBank No: 843734

Etymology

Iugi-, iugis (Latin), ridge; -spori-, sporis (Latin), spores; Iugispori-, refers to its spore ornamentation; -psathyra, one of the synonyms of Psathyrella, refers to its similarity to Psathyrella.

Description

Basidiomata psathyrelloid, fragile, non-deliquescent. Pileus hygrophanous, rugose to appearing reticulate ridged, covered by persistent and inconspicuous villus. Lamellae adnexed, brown. Stipe white, central, hollow. Spores amygdaliform in profile view, ovoid to elongate in face view, inamyloid, brown, fades in concentrated sulphuric acid, ridged and rarely verrucose ornamentation, suprahilar plage obvious. Basidia monomorphic. Pseudoparaphyses abundant. Pleurocystidia absent. Cheilocystidia present. Pileipellis hymeniderm, pyriform cell mixed with simple hairs.

Type species

Iugisporipsathyra reticulopilea J.Q. Yan, Y.G. Fan & S.N. Wang

Notes

The combination of veil absent, pleurocystidia absent and spores ornamented with ridges or rarely verrucose, with an obvious suprahilar plage is unique in Psathyrellaceae.

Iugisporipsathyra reticulopilea J.Q. Yan, Y.G. Fan & S.N. Wang, sp. nov.

MycoBank No: 843801
Fig. 3

Etymology

reticulo-, reticular; reticulopilea, referring to the surface characteristic of the pileus.

Description

Pileus 30–90 mm broad, oblate when young, expanding to plane, surface dry, rugose to appearing reticulate ridged, hygrophanous, pale yellow to greyish-yellow (4A3–4B2), becoming yellowish-white (4A2) as pileus dries, centre and ridged area darker, brown to dark brown (7D6–7F6), becoming greyish-yellow (4B2) as pileus dries. Pileus surface covered by inconspicuous villus. Villus very short, white (4A2), persistent. Veil absent. Context 3.0–4.0 mm broad, fragile, dirty white (7A1–7B2). Lamellae 3.5–10 mm broad, crowded, adnexed, 2–3 tiers of lamellulae, dirty white (7A1–7B2), becoming brown (7E6–7E8) as spores mature, edge white (7A1–7B1) and saw-toothed under 20× magnification. Stipe 50–80 mm long, 3.0–10 mm thick, fragile to fibrous, white to dirty white (7A1–7B1), cylindrical, hollow, gradually thickening towards base, 8.0–17 mm thick at base. Stipe surface covered with small, white, evanescent fibrils.

Figure 3. 

Macroscopic and microscopic structures of Iugisporipsathyra reticulopilea a–d Basidiomata e, f spores viewed by scanning electron microscopy g spores in Melzer’s Reagent h spores in water i hymenophore j, k cheilocystidia l, m pileipellis and hairs hyphae n, o caulocystidia. Scale bars: 20 mm (a–d); 20 μm (g–o). Structures of i–o were observed in 5% KOH solution and Congo red was used as the stain.

Spores (7.5–)8.0–9.7(–10.5) × (4.0–)4.5–6.0 μm, Q = 1.5–2.0, amygdaliform in profile view, (4.5–)4.8–6.0(–6.3) μm broad, ovoid to elongate in face view, inamyloid, red-brown in water, brown in alkaline solution, fades in concentrated sulphuric acid, ornamentation up to 1.0 μm high, composed of irregular ridges and rarely verrucose, variable in length, partly connected, sometimes forming a zebroid pattern or closed meshes, suprahilar plage obvious, germ pore absent. Basidia (19–)22–29 × 9.5–12.0 μm, clavate, hyaline, 4- or 2-spored. Pseudoparaphyses abundant. Pleurocystidia absent. Cheilocystidia (37–)40–61(–68) × (9.5–)12–18(–22) μm, hyaline, utriform with obtuse to broadly obtuse apex, base tapering to a short or long stipe. Caulocystidia 50–90 × 6.0–14 μm, scattered or caespitose, various, mostly narrow clavate, hyaline. Trama of gills subparallel. Pileipellis hymeniderm, composed of a 1-cell-deep layer of pyriform cells, mixed with sparsely simple hairs, pyriform cells (35–)38–60 (–62) × (12–)14–23 μm, hairs hyphae, separate, 7.0–10 μm broad. Clamps present.

Known distribution

Tropical China (Hainan Province).

Habit and habitat

Scattered or 2–3 caespitose on red soil of roadside under broadleaf tree.

Specimens examined

China. Hainan Province, Ding’an County, Longhu Town, 2 Jan 2019, Yu-Guang Fan, Jun-Qing Yan HFJAU 1352 (holotype); 4 Jan 2019, Jun-Qing Yan, Sheng-Nan Wang, HFJAU 3181, HFJAU 3182.

DNA sequence of type

ON207138 (ITS), ON207137 (LSU), ON210974 (β-tub).

Notes

Differs from other species in Psathyrellaceae by having ridge-ornamented spores with an obvious suprahilar plage.

Discussion

The discovery of I. reticulopilea has transformed our traditional understanding of Psathyrellaceae. The species is unique amongst Psathyrellaceae in producing ridge-ornamented spores with an obvious suprahilar plage. This feature is so unusual that it seems difficult to associate it with Psathyrellaceae. However, the characteristic of the spores of fading in concentrated sulphuric acid is in common with other species in this family (Singer 1986; Kirk et al. 2008; Padamsee et al. 2008; Nagy et al. 2013; Örstadius et al. 2015; Wächter and Melzer 2020).

Macroscopically, the psathyrelloid basidiomata of I. reticulopilea enables ready discrimination from the coprinoid taxa of Psathyrellaceae. Gasteroagaricoides spp. have a densely granular-warty pileus and Macrometrula spp. have a volva (Singer 1948; Reid 1986). Iugisporipsathyra reticulopilea can be distinguished from these species by the smooth pileus and absence of a volva. Amongst the abundant psathyrelloid taxa of Psathyrellaceae, only the species of Typhrasa have slight to distinct ridge-like folds on the pileus. However, no species has a reticulate-ridged pileus similar to that of I. reticulopilea. In addition, the pileus surface of I. reticulopilea is covered by a white, inconspicuous, but persistent villus. This feature also readily distinguishes I. reticulopilea from known species of Typhrasa (Örstadius et al. 2015; Wang et al. 2021).

Microscopically, almost all species of Psathyrellaceae have smooth spores. Granulose spores are observed only in Coprinopsis, Coprinellus and Psathyrella, but are extremely rare. Verrucose spores are known only in Lacrymaria. No species has an obvious suprahilar plage as in I. reticulopilea (Guzmán et al. 1990; Örstadius and Knudsen 2012; Örstadius et al. 2015). In the classification system of Smith (Smith 1972), some species with ornamented spores were classified in Psathyrella subg. Panaeolina (Maire) A.H. Smith. Those species are now excluded from the Psathyrellaceae and are classified in Panaeolina Maire, based on phylogenetic relationships and spores that do not fade in concentrated sulphuric acid (Kirk et al. 2013; Zhao et al. 2017). Detailed morphological comparison of Iugisporipsathyra and psathyrelloid genera of Psathyrellaceae is presented in Table 2.

Table 2.

Summary of morphological characteristics used to discriminate psathyrelloid genera in the Psathyrellaceae.

Britzelmayria Candolleomyces Cystoagaricus Heterospathyrella Homophron Iugisporipsathyra Kauffmania Lacrymaria Olotia Psathyrella Typhrasa
Pileus surface smooth smooth fibrillose, squamulose, spiny, or squarrose; hyphae smooth smooth non-obvious villus; hyphae smooth tomentose; hyphae smooth smooth slight to distinct ridge-like folds
Veil wipeable; hyphae wipeable; hyphae absent wipeable; hyphae absent absent wipeable; hyphae absent wipeable; hyphae wipeable; hyphae, rarely subglobose cells wipeable; hyphae
Cap or lamellae non-deliquescent non-deliquescent non-deliquescent non-deliquescent non-deliquescent non-deliquescent non-deliquescent non-deliquescent non-deliquescent non-deliquescent non-deliquescent
Spore surface smooth smooth smooth smooth smooth ridges ornamentation with obvious suprahilar plage smooth often warty smooth smooth, rarely granulose or with myxosporium smooth
Basidia monomorphic monomorphic monomorphic monomorphic monomorphic monomorphic monomorphic mono- to dimorphic monomorphic monomorphic monomorphic
Pseudoparaphyses absent absent absent present absent present absent absent absent rarely present absent
Pileipellis paraderm hymeniderm to paraderm paraderm hymeniderm to paraderm, covered by a 1 cell deep layer of periclinal hyphae hymeniderm to paraderm. simple hairs sometimes present Hymeniderm, mixes with sparsely simple hairs hymeniderm to paraderm hymeniderm hymeniderm to paraderm hymeniderm, paraderm, rarely cutis hymeniderm to paraderm
Pleurocystidia thin-walled absent thin-walled thin-walled thick-walled absent thin-walled thin-walled predominantly spatula-shaped and strongly pediculated thin-walled or rarely slight thick-walled thin-walled, with intracellular oily drops or globules
Cheilocystidia present present present present present present present present present present present
Pileocystidia present absent absent absent absent absent absent absent absent very rarely present absent

Acknowledgements

This work was financed by the National Natural Science Foundation of China (31960008, 31860009), The Project of FAAS (XTCXGC2021007) and Jiangxi Provincial Natural Science Foundation (20202BABL213041). Sincere thanks to the anonymous reviewers of the manuscript.

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Supplementary material

Supplementary material 1 

Iugisporipsathyra reticulopilea gen. et sp. nov. (Agaricales, Psathyrellaceae) from Tropical China Produces Unique Ridge-ornamented Spores with an Obvious Suprahilar Plage

Jun-Qing Yan, Yu-Guang Fan, Sheng-Nan Wang

Data type: phylogenetic

Explanation note: A nexus file contains alignment sequence and original tree of ML and Bayes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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