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 AgaricusL.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 Coprinucomatus (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 Galerellafloriformis 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 Pluteusthomsonii (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 |
|---|---|---|---|---|
| Britzelmayriamultipedata | LÖ237-04 | KC992888 | KC992888 | KJ664867 |
| B.supernula | LÖ250-04 | KC992867 | KC992867 | KJ664849 |
| Candolleomyceseurysporus | 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 |
| Coprinellusandreorum | 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 |
| Coprinopsisbabosiae | 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 |
| Cystoagaricushirtosquamulosa | 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 | – |
| Hausknechtiafloriformis | WU22833 Type | JX968254 | JX968371 | – |
| Heteropsathyrellamacrocystidia | HMJAU37803 | MW405101 | MW413358 | – |
| H.macrocystidia | HMJAU37802 Type | MW405102 | MW413359 | MW410997 |
| Homophroncamptopodum | 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 | – |
| Iugisporipsathyrareticulopilea | HFJAU1352 Type | ON207138 | ON207137 | ON210974 |
| I.reticulopilea | HFJAU3181 | ON207139 | – | ON210975 |
| I.reticulopilea | HFJAU3182 | ON207140 | – | ON210976 |
| Kauffmanialarga | LÖ223-90 | DQ389694 | DQ389694 | KJ664912 |
| K.larga | LAS97-054 | DQ389695 | DQ389695 | – |
| Lacrymariaglareosa | 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 | – |
| Narcisseacordispora | SFSUDEH2073 | AY461827 | – | – |
| N.cordispora | LÖ41-01 | DQ389723 | – | KJ664910 |
| N.patouillardi | NL-1687 | FM878009 | FM876265 | FN396257 |
| Olotiacodinae | GLM-F112430 Type | MG696611 | MG674714 | – |
| Parasolaauricoma | 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 | – |
| Psathyrellaamygdalinospora | 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 | – |
| Punjabiapakistanica | MEL2382843 | KP012718 | KP012718 | – |
| P.pakistanica | LAH35323 Type | MH366736 | – | – |
| Tulosesuscanistri | 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 |
| Typhrasagossypina | Schumacher024 | KC992946 | KC992946 | – |
| T.nanispora | Barta980706 Type | KC992947 | KC992947 | – |
| T.polycystis | HFJAU1454 Type | MW466538 | MW466544 | – |
| T.rugocephala | HFJAU1467 Type | MW466541 | MW466546 | – |
| Outgroup | ||||
| Coprinuscomatus | AFTOL_ID_626 | AY854066 | AY635772 | – |
| Crucibulumlaeve | REGCrul1/DSH96-02 | DQ486696 | AF336246 | – |
| Cyathusstriatus | DSH96-028/Cyst1/DSH96-001 | DQ486697 | AF336247 | – |
| Lepiotacristata | ZRL20151133 | LT716026 | KY418841 | – |
| Leucocoprinusfragilissimus | ZRL20151466 | LT716029 | KY418844 | – |
| Lycoperdonericaeum | ZRL20151498 | LT716030 | KY418845 | – |
| Macrolepiotadolichaula | xml2013058 | LT716021 | KY418836 | – |
| Mycocaliadenudata | AFTOL2018/CBS494.85 | DQ911596 | DQ911597 | – |
| Mythicomycescorneipes | AFTOL-ID972 | DQ404393 | AY745707 | – |
| M.corneipes | KB51 | KY648897 | – | – |
| Nidulaniveotomentosa | AFTOL1945/CBS250.84 | DQ917654 | DQ986295 | – |
| Stagnicolaperplexa | 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.
3F06F9DF-4D54-56D4-94DA-8EB9ED2BF448
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.
Iugisporipsathyrareticulopilea 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.
0E3EF4D4-73C8-54DA-99EA-4CAA3C9FDC86
843801
Figure 3.
Macroscopic and microscopic structures of Iugisporipsathyrareticulopileaa–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.
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.
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.
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). Iugisporipsathyrareticulopilea 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 Psathyrellasubg.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 |
Supplementary Material
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.
Citation
Wang S-N, Fan Y-G, Yan J-Q (2022) Iugisporipsathyra reticulopilea gen. et sp. nov. (Agaricales, Psathyrellaceae) from tropical China produces unique ridge-ornamented spores with an obvious suprahilar plage. MycoKeys 90: 147–162. https://doi.org/10.3897/mycokeys.90.85690
Funding Statement
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.
Supplementary materials
Iugisporipsathyrareticulopilea gen. et sp. nov. (Agaricales, Psathyrellaceae) from Tropical China Produces Unique Ridge-ornamented Spores with an Obvious Suprahilar Plage
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.
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.
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Iugisporipsathyrareticulopilea gen. et sp. nov. (Agaricales, Psathyrellaceae) from Tropical China Produces Unique Ridge-ornamented Spores with an Obvious Suprahilar Plage
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.
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.