Research Article |
Corresponding author: Yupeng Ge ( gaiyupeng@126.com ) Academic editor: Maria-Alice Neves
© 2024 Qin Na, Hui Zeng, Yaping Hu, Hui Ding, Binrong Ke, Zhiheng Zeng, Changjing Liu, Xianhao Cheng, Yupeng Ge.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Na Q, Zeng H, Hu Y, Ding H, Ke B, Zeng Z, Liu C, Cheng X, Ge Y (2024) Morphological and phylogenetic analyses reveal five new species of Porotheleaceae (Agaricales, Basidiomycota) from China. MycoKeys 105: 49-95. https://doi.org/10.3897/mycokeys.105.118826
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The first occurrence of Marasmiellomycena and Pulverulina in the Chinese mycobiota are reported, M. tomentosa and P. flavoalba, two new species and M. albodescendens, a new combination, revealed by phylogenetic analyses and morphological study. These newly-recorded genera, Marasmiellomycena, which can be distinguished by their agaricoid basidiomata, dark-coloured stipe, sarcodimitic tramal structure, stipitipellis with yellow to yellowish-brown pigments and yellow-pigmented thick-walled caulocystidia and Pulverulina, which differs from other genera of Porotheleaceae by its pruinose stipe, decurrent lamellae, inamyloid basidiospores and absence of hymenial cystidia. We also formally describe three other new species of Porotheleaceae collected from Chinese temperate to subtropical zones of Fujian and Zhejiang Provinces: Clitocybula fuscostriata, Gerronema brunneosquamulosum and Leucoinocybe subglobispora. Furthermore, we include the results of a phylogenetic analysis of Porotheleaceae, based on a multi-locus (ITS, nrLSU and rpb2) dataset. According to this analysis, Chrysomycena, Clitocybula, Delicatula, Hydropodia, Hydropus, Leucoinocybe, Marasmiellomycena, Megacollybia, Pulverulina, Trogia and Vizzinia are monophyletic. However, Gerronema is identified as polyphyletic and, additionally, Porotheleum does not form a monophyletic group either because Porotheleum parvulum and Porotheleum albidum are “unassigned” in phylogenetic analysis. The results of our phylogenetic analyses, coupled with morphological observations, confirm recognition of these new taxa. Morphological descriptions, photographs, line drawings and comparisons with closely-related taxa are presented for the new species. A key to the 22 species belonging to nine genera of Porotheleaceae in China is also provided.
cyphelloid polypores, new taxon, Porotheleum, systematics, white-spored omphalinoid fungi
The family Porotheleaceae (order Agaricales), formally proposed by
A comprehensive phylogenetic analysis of Porotheleaceae has not been performed because few sequences are available. Prior to 2012, the family was informally cited in literature as the ‘hydropoid’ clade within the ‘marasmioid’ clade (
Seventeen species belonging to seven genera of Porotheleaceae, namely, one species of Clitocybula (Singer) Singer ex Métrod, one species of Delicatula, seven species of Gerronema, four species of Hydropus, one species of Leucoinocybe, two species of Megacollybia and one species of Trogia, have been recognised in China as of 2023 (
Macroscopic descriptions were based on the study of fresh specimens, whereas micromorphological descriptions relied on dried materials. In our descriptions, colour abbreviations follow the colour standards and colour nomenclature of
Genomic DNAs of the putative new species were extracted from dried materials using a NuClean PlantGen DNA kit (Kangwei Century Biotechnology Co., Beijing, China). Gene regions were amplified using the following primer pairs: ITS1/ITS4 (
For phylogenetic analysis, we constructed a concatenated dataset of 168 ITS, 87 nrLSU and 14 rpb2 sequences from 58 taxa of 14 genera of Porotheleaceae. In addition, six sequences (three ITS and three nrLSU) of Mycena purpureofusca (Peck) Sacc. were included as outgroups according to the results of
Specimens used in phylogenetic analysis, with geographic origin and GenBank accession numbers.
No. | Taxa | Voucher | Locality | ITS Sequence ID | LSU Sequence ID | rpb2 Sequence ID | Reference |
---|---|---|---|---|---|---|---|
1 | Chrysomycena perplexa | MCVE:30184 TYPE | Italy | NR172974 | NG071251 | – |
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2 | Clitocybula albida | CUH AM064 | India | MG250188 | – | – |
|
3 | Clitocybula albida | CUH AM065 | India | MG250189 | – | – |
|
4 | Clitocybula abundans | STU:SMNS-B-FU-2017/00898 | Germany | MF627833 | – | – | Unpublished |
5 | Clitocybula familia | 2319-QFB-25741 | Canada | KM406970 | – | – | Unpublished |
6 | Clitocybula familia | PRM 921866 | Czech Republic | JF730327 | JF730320 | – |
|
7 | Clitocybula familia | BRNM 736053 | Slovakia | JF730328 | JF730323 | – |
|
8 | Clitocybula familia | STU:SMNS-B-FU-2017/00926 | Germany | MF627834 | – | – | Unpublished |
9 | Clitocybula familia | NAMA 2017-349 | USA | MH979253 | – | – | Unpublished |
10 | Clitocybula fuscostriata | FFAAS1029 | China | OR238881 | OR238893 | OR258374 | This study |
11 | Clitocybula fuscostriata | FFAAS1030 Holotype | China | OR238882 | OR238894 | OR258375 | This study |
12 | Clitocybula fuscostriata | FFAAS1031 | China | OR238883 | OR238895 | OR258376 | This study |
13 | Clitocybula lacerata | LE 6639 | Russia | HM191746 | – | – |
|
14 | Clitocybula lacerata | LE 262744 | Russia | HM191747 | – | – |
|
15 | Clitocybula lacerata | LE 262743 | Russia | HM191748 | – | – |
|
16 | Clitocybula lacerata | PRM 915404 | Czech Republic | LT854054 | LT854030 | – |
|
17 | Clitocybula lacerata | WU 19575 | Austria | LT854053 | LT854031 | – |
|
18 | Clitocybula oculus | 3512 | Canada | KM406971 | – | – | Unpublished |
19 | Clitocybula oculus | WU 20008 | Canada | LT854017 | LT854017 | – |
|
20 | Clitocybula oculus | S.D. Russell iNaturalist # 8606755 | India | MN906165 | – | – | Unpublished |
21 | Clitocybula oculus | S.D. Russell iNaturalist # 8591258 | India | MN906164 | – | – | Unpublished |
22 | Clitocybula oculus | BIOUG24046-B03 | Canada | KT695321 | – | – |
|
23 | Clitocybula oculus | AFTOL-ID 1554 | USA | DQ192178 | DQ192178 | – |
|
24 | Delicatula integrella | KA12-1305 | Korea | KR673538 | – | – |
|
25 | Delicatula integrella | S.D. Russell MycoMap # 6067 | USA | MN906231 | – | – | Unpublished |
26 | Delicatula integrella | G0060 | USA | – | MK277924 | – |
|
27 | Gerronema baishanzuense | FFAAS0359 Holotype | China | OL985962 | OL985984 | – |
|
28 | Gerronema baishanzuense | FFAAS0360 | China | OL985963 | – | – |
|
29 | Gerronema baishanzuense | FFAAS0361 | China | OL985964 | OL985985 | – |
|
30 | Gerronema baishanzuense | FFAAS0362 | China | OL985965 | OL985986 | – |
|
31 | Gerronema baishanzuense | FFAAS0363 | China | OL985966 | OL985987 | – |
|
32 | Gerronema baishanzuense | FFAAS0366 | China | OL985967 | OL985988 | – |
|
33 | Gerronema brunneosquamulosum | FFAAS1032 Holotype | China | OR238884 | OR238896 | OR258377 | This study |
34 | Gerronema brunneosquamulosum | FFAAS1033 | China | OR238885 | OR238897 | OR258378 | This study |
35 | Gerronema indigoticum | HMJAU 47636 | China | MK693727 | MK693732 | – |
|
36 | Gerronema indigoticum | HMJAU 47942 | China | MK693728 | MK693733 | – |
|
37 | Gerronema indigoticum | HMJAU 47943 | China | MK693729 | MK693734 | – |
|
38 | Gerronema keralense | 2 | India | MH156555 | NG_064531 | – |
|
39 | Gerronema keralense | BKF10263 | Thailand | MZ452107 | MZ452144 | Direct Submission | |
40 | Gerronema kuruvense | CAL 1665 | India | NG_159831 | NG_064530 | – |
|
41 | Gerronema kuruvense | BKF10266 | Thailand | MZ452090 | MZ452669 | – | Direct Submission |
42 | Gerronema kuruvense | DCY3362(HGASMF01-15010) | Chian | MZ951144 | – | – | Direct Submission |
43 | Gerronema microcarpum | FFAAS0365 | China | – | OL985989 | – |
|
44 | Gerronema microcarpum | FFAAS0371 | China | OL985968 | OL985990 | – |
|
45 | Gerronema microcarpum | FFAAS0372 | China | OL985969 | OL985991 | – |
|
46 | Gerronema microcarpum | FFAAS0373 Holotype | China | OL985970 | OL985992 | – |
|
47 | Gerronema microcarpum | FFAAS0374 | China | OL985971 | – | – |
|
48 | Gerronema microcarpum | FFAAS0375 | China | OL985972 | OL985993 | – |
|
49 | Gerronema nemorale | KACC 43599 | Korea | EU883592 | – | – | Unpublished |
50 | Gerronema nemorale | KACC 43600 | Korea | EU883593 | – | – | Unpublished |
51 | Gerronema nemorale | not indicated | Korea | EU883594 | – | – | Unpublished |
52 | Gerronema nemorale | FA249 | Pakistan | MN744686 | – | – |
|
53 | Gerronema nemorale | FA236 | Pakistan | MN744687 | – | – |
|
54 | Gerronema nemorale | FA239 | Pakistan | MN744688 | – | – |
|
55 | Gerronema strombodes | DJL05NC72 | USA | EU623639 | – | – |
|
56 | Gerronema strombodes | TFB12519/TENN60718 | USA | EU623640 | – | – |
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57 | Gerronema strombodes | TFB12783/TENN61350 | USA | EU623641 | – | – |
|
58 | Gerronema strombodes | TFB11947 clone C2 | USA | KY242503 | – | – |
|
59 | Gerronema strombodes | TFB11947 clone C3 | USA | KY242504 | – | – |
|
60 | Gerronema strombodes | TFB11947 clone C5 | USA | KY242506 | – | – |
|
61 | Gerronema strombodes | TFB14234 | USA | KY242507 | – | – |
|
62 | Gerronema strombodes | TFB14514 | USA | KY242509 | – | – |
|
63 | Gerronema strombodes | TFB11947 | USA | KY271083 | – | – | from GenBank |
64 | Gerronema subclavatum | Redhead 5175, DAOM | not indicated | U66434 | – | – |
|
65 | Gerronema subclavatum | FLAS-F-60986 | USA | MH016932 | – | – | from GenBank |
66 | Gerronema subclavatum | FLAS-F-61518 | USA | MH211945 | – | – | from GenBank |
67 | Gerronema subclavatum | Smith-2018 | USA | MK573888 | – | – | Direct Submission |
68 | Gerronema subclavatum | Mushroom Observer # 243440 | USA | MK607510 | – | – | Direct Submission |
69 | Gerronema subclavatum | iNaturalist # 8545787 | India | MN906021 | – | – | from GenBank |
70 | Gerronema subclavatum | S.D. Russell MycoMap # 6854 | India | MN906138 | – | – | from GenBank |
71 | Gerronema waikanaense | PDD:87667 | New Zealand | JQ694117 | – | – | from GenBank |
72 | Gerronema wildpretii | BRNM 788347 | Madeira | LT854045 | LT854043 | – | Antonin et al. (2019) |
73 | Gerronema xanthophyllum | PRM 924657 | Czech Republic | LT854023 | LT854023 | – | Antonin et al. (2019) |
74 | Gerronema zhujian | FFAAS0364 | China | OL985973 | OL985994 | – |
|
75 | Gerronema zhujian | FFAAS0370 | China | OL985974 | OL985995 | – |
|
76 | Gerronema zhujian | FFAAS0376 Holotype | China | OL985975 | OL985996 | – |
|
77 | Hydropodia subalpina (≡Hydropus subalpinus) | STU:SMNS-STU-F-0900123 | Germany | MF039248 | – | – |
|
78 | Hydropodia subalpina (≡Hydropus subalpinus) | Montri-291 | not indicated | MK028414 | – | – | Unpublished |
79 | Hydropodia subalpina (≡Hydropus subalpinus) | Montri-312 | not indicated | MK028415 | – | – | Unpublished |
80 | Hydropodia subalpina (≡Hydropus subalpinus) | Montri-323 | not indicated | MK028416 | – | – | Unpublished |
81 | Hydropodia subalpina (≡Hydropus subalpinus) | OKA-TR-K364 | Turkey | MN701620 | MN700170 | – | Unpublished |
82 | Hydropodia subalpina (≡Hydropus subalpinus) | OKA-TR-K380 | Turkey | MN701621 | MN700171 | – | Unpublished |
83 | Hydropodia subalpina (≡Hydropus subalpinus) | OKA-TR-B400 | Turkey | MN701622 | MN700172 | – | Unpublished |
84 | Hydropus atramentosus | 918 | Italy | JF908050 | – | – |
|
85 | Hydropus fuliginarius | S.D. Russell ONT iNaturalist # 130794969 | USA | OP643427 | – | – | Unpublished |
86 | Hydropus fuliginarius | DAOM196062 | USA | – | AF261368 | – |
|
87 | Hydropus marginellus | AFTOL-ID 1720 | not indicated | DQ490627 | DQ457674 | DQ472722 |
|
88 | Hydropus marginellus | OSC 112834 | USA | EU669314 | EU852808 | – | Unpublished |
89 | Hydropus rugosodiscus | MGW1257 | USA | KY777386 | – | – | Unpublished |
90 | Hydropus rugosodiscus | PBM4022 | USA | KY777390 | – | – | Unpublished |
91 | Hydropus rugosodiscus | Taxon 10 | not indicated | MW399385 | – | – | Unpublished |
92 | Leucoinocybe auricoma (≡Mycena auricoma) | HKAS126433 | China | OQ025169 | – | – | Direct Submission |
93 | Leucoinocybe auricoma (≡Mycena auricoma) | AFTOL-ID 1341 (specimen_voucher HKAS41510) | China | DQ490647 | – | – |
|
94 | Leucoinocybe danxiashanensis | GDGM79543 | China | MZ667475 | MZ667479 | – | Unpublished |
95 | Leucoinocybe danxiashanensis | GDGM80113 | China | MZ667476 | MZ667480 | – | Unpublished |
96 | Leucoinocybe danxiashanensis | GDGM80114 | China | MZ667477 | MZ667481 | – | Unpublished |
97 | Leucoinocybe danxiashanensis | GDGM80184 | China | MZ667478 | MZ667482 | – | Unpublished |
98 | Leucoinocybe flavoaurantia | D | Italy | HM191743 | – | – |
|
99 | Leucoinocybe flavoaurantia | GDOR | Italy | HM191744 | – | – |
|
100 | Leucoinocybe flavoaurantia | LE 262757 | Russia | HM191745 | – | – |
|
101 | Leucoinocybe lenta | BOZ (EPITYPE) | Italy | – | LT854032 | – |
|
102 | Leucoinocybe lishuiensis | FFAAS 0111 (HOLOTYPE) | China | MW424488 | MW424492 | – |
|
103 | Leucoinocybe lishuiensis | FFAAS 0112 | China | MW424489 | MW424493 | – |
|
104 | Leucoinocybe lishuiensis | FFAAS 0113 | China | MW424490 | MW424494 | – |
|
105 | Leucoinocybe lishuiensis | FFAAS 0115 | China | MW424491 | MW424495 | – |
|
106 | Leucoinocybe sp. | KA12-0435 | South Korea | KR673482 | – | – |
|
107 | Leucoinocybe subglobispora | FFAAS1034 Holotype | China | OR238886 | OR238898 | OR258379 | This study |
108 | Leucoinocybe subglobispora | FFAAS1035 | China | OR238887 | OR238899 | OR258380 | This study |
109 | Leucoinocybe sulcata | CAL 1246 (HOLOTYPE) | India | KR029720 | KR029721 | – |
|
110 | Leucoinocybe taniae | BCN-SCM B-4064 | Italy | LT854057 | LT854028 | – |
|
111 | Marasmiellomycena albodescendens | PDD 96142 | New Zealand | OL998341 | OL998380 | – |
|
112 | Marasmiellomycena albodescendens | PDD 96321 | New Zealand | OL998343 | OL998382 | – |
|
113 | Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) | WU 16775 | Italy | OM422777 | OM423654 | – | Direct Submission |
114 | Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) | LIP 0401689 | France | OM422780 | OM423655 | – | Direct Submission |
115 | Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) | AMB 18850 | France | OM422781 | OM423656 | – | Direct Submission |
116 | Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) | AMB 18845 | France | OM422782 | – | – | Direct Submission |
117 | Marasmiellomycena pseudoomphaliiformis | BRNM:552721 | USA | OR913562 | OR913566 | – |
|
118 | Marasmiellomycena pseudoomphaliiformis | BRNM:552654 | USA | OR913560 | OR913564 | – |
|
119 | Marasmiellomycena pseudoomphaliiformis | BRNM:552658 | USA | OR913561 | OR913565 | – |
|
120 | Marasmiellomycena tomentosa | FFAAS1036 Holotype | China | OR238888 | OR238900 | OR258381 | This study |
121 | Marasmiellomycena tomentosa | FFAAS1037 | China | OR238889 | OR238901 | OR258382 | This study |
122 | Marasmiellomycena tomentosa | FFAAS1038 | China | OR238890 | OR238902 | OR258383 | This study |
123 | Megacollybia clitocyboidea | TFB11884/TENN60766 | USA | EU623658 | – | – |
|
124 | Megacollybia clitocyboidea | TENN62231 | USA | EU623664 | – | – |
|
125 | Megacollybia clitocyboidea | TENN62230 clone c4 | USA | EU623673 | – | – |
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126 | Megacollybia clitocyboidea | TENN62230 clone c5 | USA | EU623674 | – | – |
|
127 | Megacollybia fallax | MICH 45002 | USA | EU623714 | – | – |
|
128 | Megacollybia fallax | TFB11561/TENN59447 | USA | EU623723 | – | – |
|
129 | Megacollybia fallax | DAOM208710 | USA | EU623724 | – | – |
|
130 | Megacollybia fallax | Mushroom Observer 291302 | USA | MN176984 | – | – | Direct Submission |
131 | Megacollybia fallax | Mushroom Observer 286893 | USA | MT437075 | – | – | Direct Submission |
132 | Megacollybia marginata | PRM 860926 | Czech Republic | LT854022 | – | – |
|
133 | Megacollybia marginata | PRM 859785 | Czech Republic | LT854046 | LT854042 | – |
|
134 | Megacollybia marginata | HR 91642 | Czech Republic | LT854050 | – | – |
|
135 | Megacollybia marginata | HR 91607 | Czech Republic | LT854051 | – | – |
|
136 | Megacollybia platyphylla | AFTOL-ID 560 | USA | DQ249275 | AY635778 | DQ385887 | Unpublished |
137 | Megacollybia platyphylla | BRNM 737654 | Czech Republic | LT854048 | LT854036 | – |
|
138 | Megacollybia platyphylla | BRNM 766972 | Czech Republic | LT854049 | LT854037 | – |
|
141 | Megacollybia rodmani | BHS2009-06 | USA | GQ397989 | – | – | from GenBank |
149 | Megacollybia rodmani | PUL F27039 | USA | MW448576 | – | – | from GenBank |
150 | Megacollybia subfurfuracea | TFB11075/TENN59558 clone c3 | USA | EU623744 | – | – |
|
151 | Megacollybia subfurfuracea | TFB11075/TENN59558 clone c8 | USA | EU623745 | – | – |
|
152 | Megacollybia texensis | DPL7405/TENN62058 clone c1 | USA | EU623725 | – | – |
|
153 | Megacollybia texensis | DPL7405/TENN62058 clone c2 | USA | EU623726 | – | – |
|
154 | Megacollybia texensis | FLAS-F-61511 | USA | MH211940 | – | – | from GenBank |
155 | Mycena purpureofusca | HMJAU 43554 | China | MG654740 | MK629356 | – |
|
156 | Mycena purpureofusca | HMJAU 43624 | China | MG654741 | MK629357 | – |
|
157 | Mycena purpureofusca | HMJAU 43640 | China | MG654742 | MK629358 | – |
|
158 | Porotheleum fimbriatum | Dai 12276 | China | KX081137 | KX161656 | – | from GenBank |
159 | Porotheleum fimbriatum | Dai 12289 | China | KX081138 | KX161654 | – | from GenBank |
160 | Porotheleum fimbriatum | CLZhao 1120 | China | MH114870 | – | – | from GenBank |
161 | Porotheleum fimbriatum | CLZhao 2368 | China | MH114871 | – | – | from GenBank |
162 | Porotheleum fimbriatum | SWFC 006350 | China | MK894078 | – | – | from GenBank |
163 | Porotheleum fimbriatum | SWFC 006399 | China | MK894079 | – | – | from GenBank |
164 | Porotheleum parvulum | JBSD131802 Type | Dominican Republic | NR_182714 | OM423657 | – |
|
165 | Pseudohydropus floccipes | AMB 18768 | Spain | – | OM423637 | – |
|
166 | Pseudohydropus floccipes | BRNM 825631 | Spain | OM422760 | OM423636 | – |
|
167 | Pseudohydropus floccipes | BRNM 751633 | Spain | OM422759 | OM423635 | – |
|
168 | Pseudohydropus globosporus | BAP 661 (Holotype, SFSU) | USA | OM422758 | OM423634 | – |
|
169 | Pseudohydropus sp | MushroomObserver490861 | Jamaica | OR879917 | – | – | Direct Submission |
170 | Pulverulina flavoalba | FFAAS1039 Holotype | China | OR238891 | OR238903 | OR258384 | This study |
171 | Pulverulina flavoalba | FFAAS1040 | China | OR238892 | OR238904 | OR258385 | This study |
172 | Pulverulina ulmicola | TENN 029208 Holotype | USA | NR_119887 | HQ179668 | – |
|
173 | Pulverulina ulmicola | TFB13871 | USA | MT237476 | MT237446 | – |
|
174 | Pulverulina ulmicola | KUBOT-KRMK-2020-13 | India | MW425325 | MW425344 | – | Unpublished |
175 | Trogia benghalensis | CUH AM031 | India | KU647630 | – | – |
|
176 | Trogia benghalensis | CUH AM122 | India | MF967246 | – | – |
|
177 | Trogia infundibuliformis | KUN_HKAS63661 | China | JQ031775 | JQ031780 | – |
|
178 | Trogia infundibuliformis | KUN_HKAS56709 | China | JQ031776 | JQ031781 | – |
|
179 | Trogia venenata | KUN_HKAS54710 | China | JQ031772 | JQ031778 | – |
|
180 | Trogia venenata | KUN_HKAS56679 | China | JQ031773 | JQ031779 | – |
|
181 | Trogia venenata | TC2-28 | China | KT968080 | – | – |
|
182 | Trogia venenata | MHHNU 8750 | China | KX268227 | – | – | Unpublished |
183 | Vizzinia domingensis (≡Porotheleum domingense) | JBSD131801a | Dominican Republic | OM422768 | OM423646 | – |
|
184 | Vizzinia nigripes (≡Porotheleum nigripes) | JBSD131803 | Dominican Republic | OM422771 | OM423648 | – |
|
A data matrix was created for 59 taxa, including 58 taxa of Porotheleaceae and, as an outgroup, one taxon of Mycena. Including gaps, the aligned dataset comprised 2,274 nucleotide sites: 974 for ITS, 610 for nrLSU and 690 for rpb2 exons (all sites without introns). For the ML analysis, the best-fit substitution models selected for ITS, nrLSU and rpb2-exon partitions in the concatenated dataset were TPM2uf+I+G4, GTR +I+G4 and TIM2+I+G4, respectively. For the BI analysis, the best-fit substitution model selected for each of the three DNA regions (ITS, nrLSU and rbp2 exons) was GTR+I+G. Phylogenetic reconstructions, based on BI and ML methods, yielded similar topologies. The BI topology was, therefore, selected as a representative phylogeny (Fig.
In the tree shown in Fig.
In the phylogenetic tree, samples of the new species and new combination are placed in Marasmiellomycena, where they constitute monophyletic lineages, each with high statistical support (M. albodescendens: MLB = 93%, BPP = 1.00; M. tomentosa: MLB = 100%, BPP = 1.00). The four other new species are strongly supported as members of Gerronema 3, Pulverulina, Leucoinocybe and Clitocybula clades (C. fuscostriata: MLB = 92%, BPP = 1.00; G. brunneosquamulosum: MLB = 100%, BPP = 1.00; L. subglobispora: MLB = 100%, BPP = 1.00; and Pulverulina flavoalba: MLB = 100%, BPP = 1.00). Marasmiellomycena tomentosa is closely related to a clade containing two species and a new combination, M. albodescendens, M. omphaliiforme and M. pseudoomphaliiformis. Pulverulina flavoalba sp. nov. is grouped with high statistical support (MLB = 100%; BPP = 1.00) with three sequences of Pulverulina ulmicola (H.E. Bigelow) Matheny & K.W. Hughes from India and the USA (including holotype voucher no. TENN 029208). Within the Leucoinocybe clade, L. subglobispora constitutes a monophyletic lineage that is most closely related to Leucoinocybe lishuiensis, a new species recently described from China (
Clitocybula albida A.K. Dutta, K. Acharya & Antonín, reported from India as a new species, was transferred to Porotheleum [as Porotheleum albidum (A.K. Dutta, K. Acharya & Antonín) Vizzini & Consiglio] and Porotheleum parvulum Angelini, Vizzini, Consiglio & M. Marchetti as a new species from the Dominican Republic (
China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu, Yaping Hu, Changjing Liu and Hui Ding, FFAAS1030 (collection number MY0460).
Name refers to the pileus with radially fuscous striae.
Microscopic features of Clitocybula fuscostriata (FFAAS1030, holotype) A–E basidiospores F basidia G margin of lamellae H lamellar trama I pileipellis and pileocystidia J caulocystidia. Scale bars: 5 μm (A–E); 10 μm (F–J). Structures were stained with 1% Congo Red aqueous solution before photographing.
Pileus 3.0–28.5 mm in diameter, hemispherical at first, then convex with depressed centre, expanded with age, infundibuliform with deeply umbilicate at the centre when old, thin-fleshed, dry, surface innately radially Fuscous (XLVI13′′′′k) to Fuscous-Black (XLVI13′′′′m) striate, surface somewhat fibrillose, becoming glabrous, radially cracked at margin when old, Benzo Brown (XLVI13′′′′i), Hair Brown (XLVI17′′′′i), Fuscous (XLVI13′′′′k) to Fuscous-Black (XLVI13′′′′m) at the centre, Pale Smoke Grey (XLVI21′′′′f) in the margin when young, Pale Smoke Grey (XLVI21′′′′f) to Smoke Grey (XLVI21′′′′d) with Bone Brown (XL13′′′m) at the centre when old. Context thin, white, fragile. Lamellae subdecurrent, white, with 1–3 tiers of lamellulae, irregularly intervenose, edges concolorous with the face. Stipe 17.0–52.0 × 1.0–2.5 mm, hollow, cylindrical, strongly and coarsely grooved, slightly bulbous at the base, fragile, finely whitish fibrillose, white in the upper part, Citrine Drab (XL21′′′i) in the base, base covered with a few white fibrils. Odour and taste inconspicuous.
Basidiospores (80/4/3) (5.2) 5.4–5.8–6.2 (6.5) × (4.2) 4.3–4.7–5.0 (5.1) μm [Q = 1.13–1.34, Q = 1.25 ± 0.050] [holotype (40/2/1) (5.3) 5.5–5.8–6.2 (6.5) × (4.2) 4.4–4.6–5.0 (5.1) μm, Q = 1.17–1.32, Q = 1.26 ± 0.040], broadly ellipsoid, hyaline in 5% KOH, smooth, thin-walled, guttulate, amyloid. Basidia 22–32 × 5–9 μm, 2- or 4-spored, clavate, sterigmata 2.5–4.7 × 0.6–1.6 μm. Hymenial cystidia absent. Lamellae edge cells scattered, cylindrical, narrowly clavate, thin-walled. Lamellar trama subregular; hyphae 3–7 μm wide, thin-walled, hyaline, non-dextrinoid. Pileipellis hyphae 4–9 μm wide, smooth; pileocystidia 70–162 × 7–19 μm, cylindrical or narrowly clavate, apically obtuse, thin-walled, hyaline, smooth. Stipitipellis a cutis made up of 3–8 μm wide hyphae, smooth, thin-walled; caulocystidia 27–63 × 5–8 μm, cylindrical, clavate, fusoid, apically obtuse, thin-walled base, smooth, transparent. Clamps present in all tissues.
Scattered on rotten branches or twigs in Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea mixed forests.
Zhejiang Province, China.
China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Hui Zeng and Yulan Sun, FFAAS1029 (collection number MY0459); Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu, Yaping Hu, Changjing Liu and Hui Ding, FFAAS1031 (collection number MY0466).
Clitocybula fuscostriata is considered to be a distinct species in the genus on account of its pileus with dark-brown striae, broadly ellipsoid basidiospores, absence of cheilocystidia and pleurocystidia and thin-walled pileipellis and stipitipellis hyphae. Five recorded species morphologically resemble this new species: C. familia (Peck) Singer, C. lacerata (Scop.) Métrod, C. oculata (Murrill) H.E. Bigelow, C. striata Dähncke, Contu & Vizzini and C. tilieti (Singer) Singer (
Differs from G. zhujian in having a fuscous stipe densely covered with deep-brown pubescence or scales and by the presence of large basidiospores.
China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, and Hui Zeng, FFAAS1032 (collection number MY0481).
Name refers to the pileus and stipe covered with dark-brown scales.
Microscopic features of Gerronema brunneosquamulosum (FFAAS1032, holotype) A–E basidiospores F basidia G–J cheilocystidia K lamellar trama L pileipellis and pileocystidia M caulocystidia. Scale bars: 5 μm (A–E); 10 μm (structures A–K, M were stained with 1% Congo Red aqueous solution and L in 5% KOH aqueous solution before photographing).
Pileus 4.5–42.0 mm in diam., applanate and centrally depressed, subumbilicate to umbilicate when young, concave to deeply infundibulate with age, pellucid-striate or sulcate, always ± distinctly radially striped with darkened lines, Buffy Brown (XL17′′′k) at the centre, Olive Buff (XL21′′′d) in margin when young, Olive Brown (XL17′′′k), Clove Brown (XL17′′′m), Light Greyish-Olive (XLVI21′′′′b) in margin with age, densely covered with tiny, Warm Blackish-Brown (XXXIX1′′′m) granules, pubescence or scales, slightly sparse with age, dry, lustreless, with a slightly involuted margin. Context white, thin, tough. Lamellae narrowly adnexed to subdecurrent, moderately broad, pure white, edges concolorous with the sides. Stipe 6.0–32.0 × 1.5–2.0 mm, central, cylindrical, almost equal above, white, densely covered with Warm Blackish-Brown (XXXIX1′′′m) scales, hollow, base Light Seal Brown (XXXIX9′′′m), slightly swollen with tiny, inconspicuous fine white hairs. Odourless. Taste mild.
Basidiospores [60/3/2] (9.0) 9.2–10.0–11.2 (12.9) × (4.9) 5.2–5.8–6.6 (7.2) μm [Q = 1.54–1.91, Q = 1.73 ± 0.097] [holotype [40/2/1] (9.0) 9.2–10.2–11.2 (12.9) × (5.3) 5.5–5.9–6.5 (7.2) μm, Q = 1.54–1.90, Q = 1.71 ± 0.086], ellipsoid to narrowly ellipsoid, hyaline, guttulate, thin-walled, inamyloid. Basidia 22–39 × 7–9 μm, hyaline, clavate, 2- or 4-spored, sterigmata 2.3–6.0 × 0.8–2.2 μm. Cheilocystidia 23–59 × 6–9 μm, subfusiform, clavate, apex usually swollen, hyaline. Pleurocystidia absent. Lamellar trama subregular; hyphae 2–7 μm wide, thin-walled, hyaline, inamyloid. Pileus trama subregular, sarcodimitic. Pileipellis hyphae 3–7 μm wide, a cutis, light yellow (2B2); terminal elements clavate or utriform with rounded apex, 53–95 × 7–16 μm, Dark Citrine (IV21m), Olive Brown (XL17′′′k) to Clove Brown (XL17′′′k) pigmented; true pileocystidia absent. Hyphae of the stipitipellis 5–11 μm wide, hyaline, smooth; caulocystidia long cylindrical, sometimes with rounded apex, 40–76 × 5–12 μm, hyaline, thin-walled. All tissues non-reactive in iodine. Clamps present in all tissues.
Solitary to scattered on rotten wood, branches and twigs in Acer, Ginkgo, Liriodendron, Picea and Tsuga.
Fujian Province, Zhejiang Province, China.
China. Fujian Province: Wuyi Mountain, Nanping City, 13 Aug 2021, Qin Na, Yupeng Ge, Junqing Yan, Hui Zeng, and Zewei Liu, FFAAS1033 (collection number MY0571).
Gerronema brunneosquamulosum is unique amongst members of Gerronema on account of its fuscous pileus and stipe with dark-brown to blackish-brown pubescence or scales, larger basidiospores and a dark-pigmented pileipellis. Gerronema zhujian, reported from Anhui and Fujian Provinces in China, is the most closely allied congener of G. zhujian on the basis of the brown colouration of the umbilicus of its pileus, its whitish stipe and similarly-shaped cheilocystidia and terminal elements of the pileipellis (
Pileus dark brown. Basidiospores subglobose to broadly ellipsoid. Pileocystidia and caulocystidia thick-walled. Differs from L. lishuiensis in having broader basidiospores.
China. Zhejiang Province: Tianmu Mountain, Hangzhou City, 1 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu and Yulan Sun, FFAAS1034 (collection number MY0444).
Name refers to the subglobose to broadly ellipsoid basidiospores.
Microscopic features of Leucoinocybe subglobispora (FFAAS1034, holotype) A–E basidiospores F basidia G–J cheilocystidia K lamellar trama L pileipellis and pileocystidia M caulocystidia. Scale bars: 5 μm (A–E); 10 μm (F–M). Structures were stained with 1% Congo Red aqueous solution before photographing.
Pileus 2.5–8.0 mm in diameter, hemispherical or campanulate when young, becoming campanulate with age, umbilicate at the centre, sulcate, finely granulose all over, Dark Livid Brown (XXXIX1′′′k), Benzo Brown (XLVI13′′′′i) to Fuscous (XLVI13′′′′k) at the centre, Pale Smoke Grey (XLVI21′′′′f) in the margin, uplifted or recurved at the margin and sometimes rimose in age, dry. Context white, thin, fragile. Lamellae adnexed to slightly subdecurrent, white, with 1–2 tiers of lamellulae, edges concolorous with the face. Stipe 9.5–14.0 × 1.0–1.5 mm, equal or slightly broadened at the base, hollow, fragile, white, sometimes inconspicuous Pale Olive-Buff (XL21′′′d) at the base, densely pruinose, but sparsely with age, base covered with small white fibrils. Odour and taste indistinctive.
Basidiospores (60/3/2) (5.6) 5.8–6.4–7.1 (7.5) × (4.8) 5.0–5.6–6.5 (6.8) μm [Q = 1.06–1.27, Q = 1.16 ± 0.054] [holotype (40/2/1) (5.7) 5.9–6.5–7.2 (7.5) × (4.9) 5.0–5.5–6.5 (6.8) μm, Q = 1.07–1.27, Q = 1.18 ± 0.052], subglobose to broadly ellipsoid, hyaline in 5% KOH, smooth, thin-walled, guttulate, amyloid. Basidia 28–37 × 7–9 μm, 4-spored, clavate, sterigmata 1.4–2.7 × 0.8–1.7 μm. Cheilocystidia 28–62 × 9–15 μm, distinct, flexuose, narrowly utriform, fusoid or lageniform, subcapitate, thin-walled, hyaline. Pleurocystidia absent. Lamellae trama subregular; hyphae 2–6 μm wide, thin-walled, hyaline, amyloid. Pileipellis hyphae 2–8 μm wide, smooth; pileocystidia 62–116 × 10–19 μm, lageniform, subulate, apically obtuse, distinctly 0.8–1.8 μm thick-walled, with a thin-walled base, hyaline, smooth. Stipitipellis a cutis made up of 3–9 μm wide hyphae, smooth, thin-walled; caulocystidia 34–62 × 5–10 μm, subulate, fusoid, lageniform, sometimes clavate, always thick-walled in the middle part and with a thin-walled base, smooth, transparent. Clamps present in all tissues.
Solitary or scattered on rotten wood or branches in Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea mixed forests.
Zhejiang Province, China.
China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu, Yaping Hu and Hui Ding, FFAAS1035 (collection number MY0475).
Leucoinocybe subglobispora is considered to be a distinct species of Leucoinocybe on account of its subdecurrent lamellae, subglobose to broadly ellipsoid basidiospores, thick-walled pileocystidia and caulocystidia and saprophytic habitat. Leucoinocybe lenta, the type species of Leucoinocybe, also has a white stipe and lamellae, similarly-shaped cheilocystidia and thick-walled pileocystidia, but differs from the new species by the presence of a reddish-brown pileus with pinkish shades or pale pinkish-beige at the centre that fades to white towards the margin, larger basidiomata and ellipsoid basidiospores [(5.3)6.0–7.3(7.9) × (3.8)4.0–4.5(5.1) μm] (
Porotheleum albodescendens J.A. Cooper, in Consiglio, Vizzini, Cooper, Marchetti, Angelini, Brugaletta & Setti, Riv. Micol. 64(2): 117, 2022.
Holotype : New Zealand: North Island, Taupo, Tauhara Centre, 15 May 2011, PDD 96321.
New Zealand.
Marasmiellomycena albodescendens has marasmielloid basidiomes, a pure-white pileus, relatively large spores, no hymenial cystidia and abundant, thick-walled pileocystidia and caulocystidia with yellowish contents. Unlike other species of Marasmiellomycena possessing a yellow, reddish-brown or yellowish-brown pileus, M. albodescendens can be easily recognised by its white pileus. The pileus of Marasmiellomycena albodescendens is macromorphologically more similar to some species of Marasmiellus Murrill (
Pileus and stipe distinctly tomentose. Pileus dark brown, subsquamulose. Basidiospores narrowly ellipsoid, slightly amyloid. Hymenial cystidia absent. Pileipellis and stipitipellis sarcodimitic, hyphae thick-walled with yellowish-brown pigments. Pileocystidia and caulocystidia thick-walled with yellow contents. Differs from M. pseudoomphaliiformis by possessing a distinctly tomentose, dark-brown subsquamulose pileus, narrowly ellipsoid basidiospores and absence of cheilocystidia.
China. Zhejiang Province: Tianmu Mountain, Hangzhou City, 30 Jul 2021, Qin Na, Zewei Liu, Yulan Sun and Yupeng Ge, FFAAS1036 (collection number MY0421).
Pileus 0.5–18.5 mm in diameter, at first convex or campanulate, soon expanding to plano-convex, always depressed to umbilicate at the centre, surface dry, densely covered with minute white (LIII) pubescence, tomentose all over, subsquamulose, ground colour Verona Brown (XXIX13′′k) to Warm Sepia (XXIX13′′m), Mustard Yellow (XVI19′b), Old Gold (XVI19′i) to Buffy Citrine (XVI19′k), Saccardo’s Olive (XVI19′m) at the centre, fading to Wax Yellow (XLVI21′′′′f) when old, margin slightly sulcate, uplifted or recurved in age. Context thin, Primrose Yellow (XXX23′′d). Lamellae decurrent to subdecurrent, Wax Yellow (XLVI21′′′′f), Mustard Yellow (XVI19′b), with 1–2 tiers of lamellulae, edges concolorous with the face, slightly fimbriate edge. Stipe 7.5–21.0 × 1.0–1.6 mm, central, terete, curved, equal or slightly broadened at the base, hollow or stuffed, dry, Mustard Yellow (XVI19′b) in the upper part, Saccardo’s Olive (XVI19′m), Benzo Brown (XLVI13′′′′i), Fuscous (XLVI13′′′′m), Deep Greyish-Olive (XLVI21′′′′b) towards the base, densely and minutely silky-fibrillose and white (LIII) pruinose-floccose to tomentose throughout, base covered with white mycelium. Odour indistinct to fungoid, taste mild.
Basidiospores (80/4/3) (6.8) 7.2–7.6–8.2 (8.4) × (3.7) 3.9–4.1–4.5 (4.6) μm [Q = 1.75–1.98, Q = 1.83 ± 0.052] [holotype (40/2/1) (6.8) 7.2–7.7–8.4 × 3.9–4.2–4.6 μm, Q = 1.75–1.98, Q = 1.82 ± 0.050], narrowly ellipsoid, hyaline in 5% KOH, smooth, thin-walled, multiguttulate, slightly amyloid. Basidia 20–35 × 5–8 μm, 2- or 4-spored, clavate, sterigmata 2.2–4.8 × 0.6–1.6 μm. Hymenial cystidia absent. Lamellar trama subregular; hyphae 3–10 μm wide, with 0.5–1.0 µm thick-walled, light yellow, dextrinoid. Pileipellis hyphae 3–8 μm wide, sarcodimitic, cutis, smooth, 0.4–1.0 μm thick-walled, with intracellular yellowish-brown pigment; pileocystidia 38–223 × 5–12 μm, in clusters, narrowly subulate or narrowly lageniform to fusiform with very long and tapering neck, distinctly 0.6–1.5 μm thick-walled, yellow, smooth. Stipitipellis made up of cylindrical, 4–9 µm wide hyphae, sarcodimitic, smooth, 0.5–1.0 μm thick-walled, with intracellular brownish-orange pigment; caulocystidia 45–327 × 5–9 μm, similar to the pileocystidia, but usually longer, 0.5–1.3 μm thick-walled, smooth, with intracellular yellowish pigment. Clamps present in all tissues.
Solitary or scattered on rotten branches, twigs and wood debris in Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea mixed forests.
Zhejiang Province, China.
China. Zhejiang Province: Tianmu Mountain, Hangzhou City, 30 Jul 2021, Qin Na, Zewei Liu, Yulan Sun and Yupeng Ge, FFAAS1037 (collection number MY0422); Zhejiang Province: Tianmu Mountain, Hangzhou City, 1 Aug 2021, Qin Na, Zewei Liu, Yulan Sun and Yupeng Ge, FFAAS1038 (collection number MY0443).
Marasmiellomycena tomentosa is a rare thermophilous species reported from south-eastern areas of China from July to August on rotten branches, twigs and woody debris of deciduous and coniferous trees (Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea). The most distinctive characteristics of this species are a tomentose, brown subsquamulose pileus, a tomentose stipe, narrowly ellipsoid and slightly amyloid basidiospores, the absence of hymenial cystidia and thick-walled pileipellis, stipitipellis, pileocystidia and caulocystidia with yellow or brownish-orange contents. Species morphologically most closely allied to Marasmiellomycena tomentosa include M. omphaliiforme, M. pseudoomphaliiformis and M. albodescendens. Marasmiellomycena pseudoomphaliiformis resembles M. tomentosa by the presence of a pale beige to brown pileus with finely tomentose to pubescent pileus, but differs in having white to cream-white or beige lamellae rather than yellow, ellipsoid to ellipsoid-fusiform basidiospores [(6.5–)7.0–9.0(–9.5) × 4.0–5.5 µm] and clavate, fusiform to lageniform cheilocystidia (
Pileus white to light orange yellow. Basidiospores cylindrical. Hymenial cystidia absent. Lamellar trama, pileipellis and stipitipellis hyphae thin-walled. Differs from Pu. ulmicola in having larger and longer basidiospores and possessing thin-walled lamellar trama, pileipellis and stipitipellis hyphae.
China. Guangxi Zhuang Autonomous Region: Liangfengjiang National Forest Park, Nanning City, 13 Jul 2022, Yupeng Ge and Renxiu Wei, FFAAS1039 (collection number MY0863).
Name refers to the white to light-yellow pileus and stipe.
Microscopic features of Pulverulina flavoalba (FFAAS1039, holotype) A–E basidiospores F basidia G lamellae margin H lamellar trama I pileipellis J caulocystidia. Scale bars: 5 μm (A–E); 10 μm (F–J). Structures A–F were stained in 5% KOH aqueous solution and G–J with 1% Congo Red aqueous solution before photographing.
Pileus 1.2–5.8 mm in diameter, arched or plano-convex with a slight depression at the centre when young, becoming more depressed with age; translucent striate, floccose or granulose, glabrescent when old, surface dull, dry; white (LIII) when young, aniline yellow (IV19i) or light orange-yellow (III17d) at the centre and in the margin with age, margin decurved. Context white, thin, not fragile. Lamellae decurrent, white, orange citrine (IV19k) tinged when old, with 1–2 tiers of lamellulae, edges even, medium-broad. Stipe 1.6–14.4 × 0.5–1.0 mm, terete or slightly broadened at the base, curved, dry, white, with a pruinose, pubescent or fibrillose surface, sparser with age, hollow, not fragile, white, sometimes aniline yellow (IV19i), light orange-yellow (III17d) in the middle and at the base; base covered with white mycelium. Odour absent, taste mild.
Basidiospores (60/3/2) (6.8) 7.0–7.9–8.8 (9.1) × (3.3) 3.7–4.1–4.4 (4.7) μm [Q = 1.81–2.19, Q = 1.93 ± 0.099] [holotype (40/2/1) (6.8) 7.0–7.8–8.9 (9.1) × (3.3) 3.7–4.1–4.4 (4.7) μm, Q = 1.77–2.19, Q = 1.92 ± 0.084], cylindrical, hyaline in 5% KOH, smooth, thin-walled, guttulate, inamyloid, with a small, but discernible apiculus. Basidia 21–30 × 4–6 μm, 2- or 4-spored, clavate, sterigmata 1.9–5.6 × 0.6–1.6 μm. Hymenial cystidia absent. Lamellar trama subregular to interwoven; hyphae 5–15 µm wide, hyaline, thin-walled. Pileipellis a cutis of cylindrical hyphae 3–7 µm wide, smooth; end cells often protruding, 35–105 × 3–12 μm, cylindrical, subfusiform, apically obtuse, thin-walled, hyaline, smooth. Stipitipellis hyphae 3–8 μm wide, smooth, thin-walled; caulocystidia 19–50 × 4–9 μm, clavate, subfusiform, thin-walled, smooth, transparent. Clamps present in all tissues.
Scattered to gregarious on rotten wood, branches or fruits in mixed forests of Acacia, Ficus, Ilex, Parashorea, Picea and Trachycarpus etc.
Guangxi Zhuang Autonomous Region, China.
China. Guangxi Zhuang Autonomous Region: Liangfengjiang National Forest Park, Nanning City, 13 Jul 2022, Yupeng Ge and Renxiu Wei, FFAAS1040 (collection number MY0865).
Clitocybe ulmicola H.E. Bigelow was established by Bigelow in 1982 and published as a new combination, Pulverulina ulmicola (H.E. Bigelow) Matheny & K.W. Hughes (
1 | Lamellae not well developed | Delicatula integrella |
– | Lamellae well developed | 2 |
2 | Pileocystidia present | 3 |
– | Pileocystidia absent | 9 |
3 | Cheilocystidia not seen | Marasmiellomycena tomentosa |
– | Cheilocystidia abundant | 4 |
4 | Basidiospores inamyloid Megacollybia | 5 |
– | Basidiospores amyloid | 6 |
5 | Cheilocystidia digitate, narrowly or broadly clavate or sphaeropedunculate, rarely with short apical outgrowths | Me. clitocyboidea |
– | Cheilocystidia clavate, without outgrowths | Me. platyphylla |
6 | Cheilocystidia distinctly thick-walled overall Leucoinocybe | 7 |
– | Cheilocystidia thin-walled or slightly thick-walled in the base Clitocybula | 8 |
7 | Basidiospores narrowly ellipsoid | L. lishuiensis |
– | Basidiospores subglobose to broadly ellipsoid | L. subglobispora |
8 | Basidiospores (5.2) 5.4–5.8–6.2 (6.5) × (4.2) 4.3–4.7–5.0 (5.1) μm, broadly ellipsoid | C. fuscostriata |
– | Basidiospores 3.5–5.3(–5.5) × 3.5–5.0 μm, globose, subglobose to broadly elliptic | C. familia |
9 | Pileus trama sarcodimitic | 10 |
– | Pileus trama not sarcodimitic | 18 |
10 | Basidiospores inamyloid | Trogia venenata |
– | Basidiospores amyloid Gerronema 11 | |
11 | Basidiomata distinctly small (Pileus < 9 mm in diam.) | G. microcarpum |
– | Basidiomata moderately small (Pileus > 9 mm in diam.) | 12 |
12 | Pleurocystidia present | G. chrysocarpum |
– | Pleurocystidia absent | 13 |
13 | Pileus blue | G. indigoticum |
– | Pileus not blue | 14 |
14 | Pileus and stipe pure white | G. albidum |
– | Pileus yellow to brown, stipe white to yellowish-brown | 15 |
15 | Pileus without pubescence or scales | 16 |
– | Pileus densely covered with deep brown pubescence or scales | 17 |
16 | Cheilocystidia up to 48 μm long | G. baishanzuense |
– | Cheilocystidia less than 35 μm long | G. nemorale |
17 | Stipe without fuscous pubescence or scales, basidiospores (6.3) 6.7–7.4–8.0 (8.5) × (3.2) 3.7–4.1–4.6 (4.8) μm | G. zhujian |
– | Stipe with deep brown fuscous pubescence or scales, basidiospores (9.0) 9.2–10.0–11.2 (12.9) × (4.9) 5.2–5.8–6.6 (7.2) μm | G. brunneosquamulosum |
18 | Cheilocystidia absent | Pulverulina flavoalba |
– | Cheilocystidia present | 19 |
19 | Dermatocystidia inconspicuous and rare | Pseudohydropus floccipes |
– | Dermatocystidia abundant Hydropus | 20 |
20 | Carpophore blackening when touched or bruised | H. nigrita |
– | Carpophore not blackening in any part when touched or bruised | 21 |
21 | Basidiospores ellipsoid | H. marginellus |
– | Basidiospores broadly ellipsoid | H. atriceps |
Previous molecular phylogenetic analyses of the so-called hydropoid clade and the Porotheleaceae have been conducted, based on various combinations of ITS, 28S, 18S, 5.8S, 25S, rpb1 and rpb2 loci (
Morphologically, Marasmiellomycena is easily recognisable as an omphalinoid mushroom in the field owing to its pileus that is depressed to umbilicate at the centre, decurrent to subdecurrent lamellae, dark-coloured stipe, sarcodimitic structure and thick-walled caulocystidia with contents. Marasmiellomycena is most similar to Vizzinia, but Vizzinia differs in basidiomata turning brownish on handling, distinctly squamulose pileus, weakly amyloid spores and absence of cheilocystidia. Pulverulina resembles Clitocybula in being an omphalinoid basidiocarps with decurrent lamellae, but can be distinguished by pruinose stipes, inamyloid basidiospores and absence of hymenial cystidia. Gerronema, Megacollybia and Trogia are more similar to Marasmiellomycena on the basis of their sarcodimitic structure. Marasmiellomycena can be readily discriminated in possessing dark-coloured stipe, inamyloid basidiospores and thick-walled caulocystidia with yellow to yellowish-brown pigments. Pulverulina species are characterised by their inamyloid basidiospores, non-sarcodimitic structure, thin-walled caulocystidia and non-pigmented pileocystidia and caulocystidia.
Our multi-gene phylogenetic analysis divided Gerronema into several highly-supported clades. This finding is consistent with the analyses of
Several species of Porotheleaceae have been reported to be edible or have toxic or ecological effects. Megacollybia platyphylla (Pers.) Kotl. & Pouzar (
We thank Dr Junqing Yan (Jiangxi Agriculture University), Ms. Zewei Liu (Ludong University), Ms. Yulan Sun (Ludong University) and Ms. Renxiu Wei (Ludong University) for their kind help during fieldwork. We sincerely thank the reviewers for their corrections and suggestions to improve our work.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was supported by the National Natural Science Foundation of China (grant no. 32200008), the Natural Science Foundation of Shandong Province (grant no. ZR2020QC001), the 5511 Collaborative innovation project of Fujian Province (grant no. XTCXGC2021007), the Central Public-Interest Scientific Institution Basal Research Fund (grant no. GYZX200203), the East-west cooperation project, FAAS (grant no. DKBF-2022-12), the Natural Science Foundation of Fujian Province (grant no. 2023J01379), the Project of Biological Resources Survey in Wuyishan National Park (grant no. HXQT2020120701), the Project of Biodiversity Conservation in Lishui, Zhejiang Province (grant no. HXYJCP2021110648), the Biodiversity investigation, observation and assessment program of Ministry of Ecology and Environment of China (grant no. 2110404 and 2019-2023), the Shandong Agricultural Industry Technology System (2021 grant no. 26, SDAIT-07-03) and the Soft Science project of Ministry of Public Security (grant no. 2022LL75).
Qin Na, Xianhao Cheng, and Yupeng Ge were responsible for designing the research and contributed significantly to data analysis and interpretation. Hui Zeng, Yaping Hu, Zhiheng Zeng, Bingrong Ke, Changjing Liu, and Hui Ding actively participated in the field investigation. All authors have contributed to the manuscript and approved the version submitted for publication.
Qin Na https://orcid.org/0000-0001-8406-6389
Hui Zeng https://orcid.org/0000-0003-2025-844X
Yaping Hu https://orcid.org/0000-0003-1242-1139
Hui Ding https://orcid.org/0000-0003-4490-2105
Binrong Ke https://orcid.org/0009-0008-7209-7362
Zhiheng Zeng https://orcid.org/0009-0008-4208-2629
Xianhao Cheng https://orcid.org/0000-0002-5922-9913
Yupeng Ge https://orcid.org/0000-0001-5754-201X
All of the data that support the findings of this study are available in the main text.