MycoKeys 88: 79_ | 08 (2022) : ye “." A peer-reviewed open-access journal doi: 10.3897/mycokeys.88.79266 RESEARCH ARTICLE . 03 MycoKkeys https://mycokeys.pensoft. net Launched to accelerate biodiversity research Taxonomic study of Collybiopsis (Omphalotaceae, Agaricales) in the Republic of Korea with seven new species Ji Seon Kim', Yoonhee Cho', Ki Hyeong Park', Ji Hyun Park’, Minkyeong Kim}, Chang Sun Kim‘, Young Woon Lim! I School of Biological Sciences and Institute of Microbiology, Seoul National University, Seoul 08826, Republic of Korea 2. Water Supply and Sewerage Research Division, National Institute of Environmental Research, Incheon 22689, Republic of Korea 3 Microorganism Resources Division, National Institute of Biological Resources, Incheon, Republic of Korea 4 Forest Biodiversity Division, Korea National Arboretum, Pocheon-si 11186, Republic of Korea Corresponding author: Young Woon Lim (ywlim@snu.ac.kr) Academic editor: Bao-Kai Cui | Received 13 December 2021 | Accepted 4 March 2022 | Published 30 March 2022 Citation: Kim JS, Cho Y, Park KH, Park JH, Kim M, Kim CS, Lim YW (2022) Taxonomic study of Collybiopsis (Omphalotaceae, Agaricales) in the Republic of Korea with seven new species. MycoKeys 88: 79-108. https:// doi.org/10.3897/mycokeys.88.79266 Abstract Collybiopsis is a genus of the gymnopoid/marasmioid complex of the family Omphalotaceae. The clas- sification system of Collybiopsis has recently undergone large changes through molecular approaches. The new classification system has not been applied for Collybiopsis in the Republic of Korea, and general research on this genus was also lacking. In this study, we analyzed the Collybiopsis species in the Republic of Korea by assessing all gymnopoid/marasmioid specimens collected nationwide for ten years by combin- ing morphological approaches and multilocus (ITS + nrLSU) phylogenetic analysis. We thus confirmed that 16 species of Collybiopsis are present in the Republic of Korea, including two previously unreported species (Co. nonnulla and Co. dichroa) and seven new species (Co. albicantipes sp. nov., Co. clavicystidiata sp. nov., Co. fulva sp. nov., Co. orientisubnuda sp. nov., Co. subumbilicata sp. nov., Co. undulata sp. nov., and Co. vellerea sp. nov.). A thorough examination of the Collybiopsis suggested that it is difficult to distinguish or identify the species based on morphological characteristics only; a combined molecular ap- proach is needed for accurate identification. The Co/lybiopsis database of the Republic of Korea is updated, and information on the new species is provided. Five new combinations from Marasmiellus to Collybiopsis are also proposed (Co. istanbulensis comb. nov., Co. koreana comb. nov., Co. omphalodes comb. nov., Co. pseudomphalodes comb. nov., and Co. ramuliciola comb. nov.). Keywords Collybia, gymnopoid, Gymnopus, ITS, Marasmiellus, marasmioid, nrLSU Copyright Ji Seon Kim et al. 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. 80 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Introduction Collybiopsis Earle (1909) is a genus of gymnopoid/marasmioid mushrooms belong- ing to the family Omphalotaceae Bresinsky (Earle 1909; Petersen and Hughes 2021). Species of Collybiopsis are characterized by collybioid, gymnopoid, marasmielloid, om- phalioid, and pleurotoid basidiomata; free to decurrent lamellae; a central to eccentric, insititious to subinsititious stipe; ellipsoid to oblong, inamyloid, and hyaline basidi- ospores with white sporeprints; presence of caulocystidia; and coralloid or diverticulate terminal elements of pileipellis (Murrill 1915; Singer 1973; Antonin and Noordeloos 1993; Retnowati 2018; Oliveira et al. 2019). Owing to its relatively uncharacteristic basidiocarp and little variation in morphological characteristics, most gymnopoid/mar- asmioid species were previously placed within the genus Collybia Staude (1857) and Marasmius Fr. (1835) before molecular identification was introduced actively to tax- onomy. However, recent molecular studies have clarified the phylogenetic relationship of gymnopoid/marasmioid species belonging to the family Omphalotaceae and family Marasmiaceae Roze ex Kiihner (Wilson and Desjardin 2005; Oliveira et al. 2019). Initial molecular studies have segregated Collybia and Marasmius and some species of both genera transferred into several genera such as Gymnopus Roussel, Marasmiellus Murril, Rhodocollybia Singer, etc. (Moncalvo et al. 2002; Mata et al. 2004b; Mata et al. 2004c; Wilson and Desjardin 2005; Hughes et al. 2010; Oliveira et al. 2019; Petersen and Hughes 2017, 2021). Five Collybia sections (locephalae Halling, Levipedes Quel, Striipedes Quél, Subfumosae Singer, and Véestipedes Quél) were subsumed into Gymnopus sensu lato (s.l.) (Mata et. al., 2004c). However, Gymnopus s. |. is polyphy- letic, and there has been much debate on the delimitation of this genus (Mata et al, 2004c; Wilson and Desjardin 2005; Mata et al. 2006; Oliveira et al. 2019; Petersen and Hughes 2016). Prior to this debate, a monophyletic genus, Marasmiellus sensu stricto (s. str.), was proposed (Wilson and Desjardin 2005), with Marasmiellus juni- perinus Murrill as the monotype species (Wilson and Desjardin 2005; Sandoval-Leiva et al. 2016; Oliveira et al. 2019). A recent study showed that if judged congeneric, Collybiopsis Earle (1909) has priority over Marasmiellus Murrill (1915) based on the nomenclature rule (Petersen and Hughes 2021). Hereupon, Collybiopsis has been re- defined based on the type species, Collybiopsis ramealis Earle, with at least 44 closely related species (Petersen and Hughes 2021). All species of Collybiopsis and some species of Gymnopus sect. Vestipedes, as well as some species of Marasmiellus, are included in the genus Collybiopsis (Petersen and Hughes 2021). Collybiopsis is morphologically similar and phylogenetically close to Gymnopus (Desjardin et al. 1999; Mata 2002; Dutta et al. 2015). Both genera are reported to be distinguishable through like types of the terminal element of pileipellis, attachment of lamellae, the character of stipe, basidiospores, and cheilocystidia. However, as the characteristics of each genus cannot be seen as absolute because exceptions exist, and some characteristics overlap, it is difficult to distinguish Collybiopsis from Gymnopus solely on morphology. Furthermore, the morphological characteristics of their basidi- omata vary greatly depending on the environment and developmental stage. ‘Therefore, Seven new Collybiopsis species 81 molecular data play an important role in distinguishing these genera (Antonin and Herink 1999; Hughes et al. 2014; Hughes and Petersen 2015). Although there have been many taxonomic changes for gymnopoid/marasmioid species, these changes have not been reflected in the gymnopoid/marasmioid species in the Republic of Korea. Since the first report of Collybiopsis confluens (Pers.) R.H. Peters- en, as its previous name Collybia confluens Fr. (Kaburagi 1940), nine current Collybi- opsis species have been reported until recently (National list of species of Korea 2020). However, they were identified and classified as Collybia, Gymnopus, and Marasmiellus based on their macroscopic morphological features. Owing to the uncertain placement of previous morphologically identified collybioid collections, it was necessary to re-ex- amine Korean collections of collybioids and marasmioids based on molecular data. In this study, we investigated gymnopoid/marasmioid specimens collected over 10 years and deposited in three Korean herbaria based on their molecular analysis. As a result, we provide a list of Collybiopsis species in the Republic of Korea with seven new species. Methods Collections of specimens A total of 372 specimens deposited in three Korean fungal herbarium — Seoul Na- tional University Fungus Collection (SFC), Korea National Arboretum (KA), and the National Institute of Biological Resources (NIBR) — were used in this study. The specimens were collected from 2012 to 2021 and stored in dried condition. All speci- mens were identified based on their morphological characteristics by each herbarium. The collection information (e.g. collection date, collection site, collector, etc.) and the notes of fresh basidiomata of each specimen were provided from each herbarium. Molecular analysis Genomic DNA was extracted from each specimen using a modified CTAB DNA ex- traction protocol (Rogers and Bendich 1994). The primer set ITS1F/ITS4B (Gardes and Bruns 1993) was used to amplify the internal transcribed spacer (ITS) region for all specimens, and the primer set LROR/LRS5 (Vilgalys and Hester 1990; Rehner and Samuels 1994) was used to amplify the nuclear large subunit ribosomal RNA (nrLSU) region. PCR was conducted by a C1000 thermal cycler (Bio-Rad, Richmond, CA, USA) using AccuPower PCR master premix (Bioneer Co., Daejeon, the Republic of Korea). PCR conditions for ITS and nrLSU region were: 5 min initial denaturation at 95 °C followed by 35 cycles of 40 s at 95 °C, 40 s at 55 °C and 60 s at 72 °C with a final extension step for 7 min at 72 °C. The amplifications of the PCR products were verified by visualization using 1% agarose gels with EcoDye DNA staining solution (SolGent Co., Daejeon, the Republic of Korea). The PCR products were purified us- ing the ExpinTM PCR Purification Kit (GeneAll Biotechnology, Seoul, the Republic 82 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) of Korea) following the manufacturer’s instructions. The purified PCR amplicons were sequenced using an ABI Prism 3700 Genetic Analyzer (Life Technologies, Gaithers- burg, MD, USA) at Macrogen (Seoul, the Republic of Korea). All sequences generated in this study were proofread using MEGA version 7 (Ku- mar et al. 2016). The sequences used for analyses were deposited in GenBank (Table 1). We then selected the closely related sequences from NCBI databases mainly referred to Oliveira et al. (2019) and Petersen and Hughes (2021). After retrieving all published ITS and nrLSU sequences of all Collybiopsis species in GenBank, phylogenetic analyses were performed together with new sequences generated from specimens. ‘The sequenc- es were respectively aligned for each loci using Multiple Alignment Fast Fourier Trans- form (MAFFT ver. 7) with the L-NSI-I option algorithm (Katoh and Standley 2013). The aligned sequence data were manually checked and edited. The final sequence of each specimen was created as a concatemer by manually attaching the aligned sequenc- es of the two loci. Maximum likelihood (ML) phylogenetic tree was constructed on the CIPRES Science Gateway (Miller et al. 2012) using the GTR+GAMMA model with 1000 bootstrap replicates. Rhodocollybia butyraceae Lennox (TFB14382), Rho- docollybia dotae JL Mata and Halling (REH7007), and Rhodocollybia maculate Singer (TFB13989) were used as outgroups (Oliveira et al. 2019). Bootstraps higher than 70% were considered to support a clade and are shown in the tree (Figure 1). Morphological observation All specimens were preliminarily observed and macro/micro-structures of two to four representative specimens, which were in the best condition among the specimens, were presented in figures. Photographs and notes of fresh basidiomata taken at the time of collection were used for macro-morphological description. For micro-morphological ob- servations, tissues of dried specimens were rehydrated in 5% (w/v) KOH and mounted in Congo red solution (Clémengon 1973) and Melzer’s reagent. The observation was per- formed by using a Nikon Eclipse 80i optical microscope (Nikon, Tokyo, Japan) at 20 x to 1000 x magnification. More than thirty basidiospores and more than twenty other micro- structures (e.g., basidia, cheilocystidia, etc.) were measured to analyze the microstructures based on the microscopic pictures of specimens stained with Congo red. The Methuen Handbook of Colour (Kornerup and Wanscher 1978) was used for color indications. The following abbreviations and acronyms were used: Co = Collybiopsis; G = Gymnopus; Ma = Marasmiellus; L = the number of complete lamellae; 1 = the number of lamellulae tiers between neighboring complete lamellae; and Q = the values of the length divided by the width of basidiospores (Petersen and Hughes 2021; Ryoo et al. 2020). Results Through ITS sequence analysis of 372 gymnopoid/marasmioid specimens, 201 speci- mens were confirmed to belong to Collybiopsis. The remaining 160 specimens were 83 Seven new Collybiopsis species Table |. Information about the Collybiopsis specimens and published Collybiopsis sequences used in phy- logenetic analysis. Species with an asterisk are those proposed as new species. Sequences newly produced in this study are presented in bold. Organisms Specimen Collection Location GenBank Accession Number Date ITS nrLSU Collybiopsis SFC20170725-35 25.7.2017 Yeosu-si, Jeollanam-do, the Republic of Korea OL467272. +=OL462811 albicantipes* SFC20180704-86 4.7.2018 Jindo-gun, Jeollanam-do, the Republic of Korea OL467273 OL462812 Co. biformis TFB14251 USA: Tennessee, GSMNP KJ416245 KJ189567 TFB13890 USA: North Carolina KJ416248 KJ189570 TFB13814 USA: Tennessee KJ416249 KJ189569 KA14-0526 15.7.2014 Suncheon, Jeollanam-do, the Republic of Korea OL467227 OL462784 KA16-0526 13.7.2016 Sinan-gun, Jeollanam-do, the Republic of Korea OL467228 OL462785 SFC20180704-36 4.7.2018 Wando-gun, Jeollanam-do, the Republic of O1467229 OL462789 Korea SFC20180831-16 31.8.2018 Jindo-gun, Jeollanam-do, the Republic of Korea OL467230 OL462790 Co. AWWO01 Java/Bali AY263434 AY639412 brunneigracilis Co. californica TFB5787 Canada: British Columbia MN413338 Co. SFC20180705-26 5.7.2018 Haenam-gun, Jeollanam-do, the Republic of OL467250 OL462816 clavicystidiata* Korea SFC20180705-84 5.7.2018 Jindo-gun, Jeollanam-do, the Republic of Korea OL467252 OL462817 SFC20180705-92 5.7.2018 Jindo-gun, Jeollanam-do, the Republic of Korea OL467253 OL462818 SFC20180713-09 13.7.2018 Gwanak-gu, Seoul, the Republic of Korea OL467251 O1L462819 Co. confluens SFC20190731-06 31.7.2019 ‘Taebaek-si, Gangwon-do, the Republic of Korea OL467237 OL462797 SFC20190731-48 31.7.2019 ‘Taebaek-si, Gangwon-do, the Republic of Korea OL467238 OL462798 TFB14115 Germany, Thuringia KP710292 ~—-KJ 189578 110116MFBPL0425 China MW554401 HMAS 290186 China MK966541 Co. confluens ssp. | TFB14409 Canada: New Brunswick KP710278 ~—-KJ189585 americana TFB14075 USA: North Carolina KP710281 KJ189581 Co. dichroa KA14-0969 19.8.2014 Hwasun-gun, Jeollanam-do, the Republic of OL467254 OL462799 Korea KA18-0389 10.7.2018 Cheongdo-gun, Gyeongsangbuk-do, the O1L467255 = O1L546541 Republic of Korea SFC20180712-16 12.7.2018 Gwangju, Gyeonggi-do, the Republic of Korea O1467256 OL462800 TFB9623 USA: North Carolina MW396865 MW396865 TENNG0014c2 USA: Tennessee, GSMNP JF313671 TFB7920 USA DQ450007 TENNG1624cla USA: Tennessee, GSMNP JF313678 TFB2028 USA DQ450008 TENNG61624c9 USA: Tennessee, GS MNP JF313692 Co. disjuncta TFB14339 USA: Connecticut NR_137865 TFB14281 USA: Mississippi KJ416253 KY019643 Co. eneficola 09-09-26AV13 Canada: Newfoundland NR_137613 NG_059502 MICH:PK6975 Alaska KP710270 KP7 10304 Co. fibrosipes FB9699 Costa Rica AF505763 Co. filamentipes TFB13962 USA: Tennessee MN897832 MN897832 Co. foliiphila CUH AM090 India NR_154176 NG_060320 CUM AM101 India KP317638 —_KP317636 Co. fulva* KA13-0216 19.6.2013 Geochang-gun, Gyeongsangnam-do, the O1467257. = OL462793 Republic of Korea KA13-0333 10.7.2013 Pocheon-si, Gyeonggi-do, the Republic of Korea OL467258 OL462794 KA15-0210 21.7.2015 Pocheon-si, Gyeonggi-do, the Republic of Korea OL467259 OL462795 Co. furtiva TFB4796 USA: Georgia MN413343, MW396879 Co. gibbosa MEL:2382838 Australia KP012713, = KP012713 URM 90012 Brazil KY061202 KY061202 Co. hasanskyensis. TFB11846 Russia: Kedrovaya MN897829 TFB11847 Russia MN897830 84 Organisms Co. indoctus Co. istanbulensis Co. juniperina Co. koreana Co. luxurians Co. melanopus Co. menehune Co. mesoamericana Co. micromphaleoides Co. minor Co. neotropica Co. nonnulla Co. nonnulla var. attenuatus Co. obscuroides Co. omphalodes Co. orientisubnuda* (as Gymnopus subnudus) Co. parvula Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Specimen AWW04 KATO Fungi 3596 BRNM 781163 TFB9889 TFB10782 SFC20120821-84 SFC20130711-05 SFC20150721-10 BRNM 714972 BRNM 718782 NIBRFG0000502888 SFC20190731-18 TFB10350 ZD16102301 TFB9121 AWW54 CUH AM093 SFC20150811-29 SFC20180905-33 SFSU: DED5866 CUH:AM074 SFSU-AWW15 TFB11005 REH7379 TENN 68165 TFB14282 TFB11930 TFB5434 TFB10416 KA13-0254 KA13-0741 KA15-0129 TFB14492 TFB14278 AWW05 AWW55 RAK369.2 RAK372.2 GB-0150514 FB11021 TFB 10427 TFB 10022 NIBRFG0000500990 SFC20170823-39 SFC20180830-29 KUC20150911-19 TFB10419 TFB10422 Collection Date 21.8.2012 11.7.2013 21.7.2015 4.9,.2018 31.7.2019 11.8.2015 5.9.2018 20.6.2013 21.8.2013 14.7.2015 19.7.2016 23.8.2017 30.8.2018 Location Unknown Turkey Turkey USA: Louisiana Argentina: Missiones Boryeong-si, Chungcheongnam-do, the Republic of Korea Pyeongchang-gun, Gangwon-do, the Republic of Korea Inje-gun, Gangwon-do, the Republic of Korea Korea Korea Ongjin-gun, Incheon, the Republic of Korea Taebaek-si, Gangwon-do, the Republic of Korea USA: North Carolina China USA: Louisiana Java/Bali India Guri-si, Gyeonggi-do, the Republic of Korea Anyang City, Gyeonggi Province, the Republic of Korea Hawaii India Java/Bali Costa Rica Costa Rica USA: Tennessee, GSMNP USA: South Carolina Costa Rica Geochang-gun, Gyeongsangnam-do, the Republic of Korea Geochang-gun, Gyeongsangnam-do, the Republic of Korea Gangneung-si, Gangwon-do, the Republic of Korea USA: Mississippi USA: Mississippi Java/Bali Java/Bali Cameroon Cameroon Norway: Svalbard Costa Rica Costa Rica Costa Rica Ulleung-gun, Gyeongsangbuk-do, the Republic of Korea Hapcheon-gun, Gyeongsangnam-do, the Republic of Korea Hapcheon-gun, Gyeongsangnam-do, the Republic of Korea Korea Costa Rica Costa Rica GenBank Accession Number ITS nrLSU AY263439 KX184795 —-KX184796 KY250435 AY256708 KY019637 KY026661 KY026661 O1467269 O1L546545 O1467270 = OL462801 OL467271 OL462802 GU319113, GU319117 GU319114 GU319118 OL467248 OL462803 O1467249 O1L462804 AY256709 AY256709 MN523270 KY026649 KY026649 NR_137539 NG_060624 KM896875 — KP100305 O1467235 OL462805 O1467236 OL462806 AY263426 KJ778753 KP100302 AY263443 AY639424 DQ450035 KY019632 AF505768 MN413334 MW396880 MW396872 MWW396872 AF505769 O1L467242 OL462820 O1467243 OL462807 O1L467244 OL462808 KY026699 KY026699 KY026701 KY026701 AY263445 AY639426 AY263446 MN930621 MN930622 KX958399 = KX958399 AF505761 DQ450011 AY256700 O1467262 O1L546546 O1467263 OL546547 OL467264 OL462796 KX513748 DQ450060 AF505774 Organisms Co. peronata Co. polygramma (as Gymnopus iocephalus) Co. pseudoluxurians Co. pseudomphalodes Co. quercophila Co. ramealis Co. ramulicola Co. readiae Co. stenophylla Co. subcyathiformis Co. subnuda Co. subpruinosus Co. subumbilicata* Co. trogivides Co. undulata* Specimen TFB13743 LE-Bin1364 CBS 223.37 SFC20170807-35 SFC20180905-63 SFC20210629-01 PR2542TN CUH:AM082 URM 90015 MHHNU 30912 TFB9628 SFC20120821-64 HEJAU 0425 KUC20140804-02 TFB14290 REH7348 PR24TN TFB14570 TFB14615 Seven new Collybiopsis species Collection Date 7.8.2017 5.9.2018 29.6.2021 Location Belgium Russia unknown Hapcheon-gun, Gyeongsangnam-do, the Republic of Korea Gwanak-gu, Seoul, the Republic of Korea Gwanak-gu, Seoul, the Republic of Korea Puerto Rico India Brazil: Amapa China Puerto Rico Korea China: Jiangxi Korea USA: Mississippi Costa Rica Puerto Rico Slovakia USA: California NIBRFG0000508888 29.7.2020 Jeongseon-gun, Gangwon-do, the Republic of TFB13769 TFB13770 DED4425 TFB14555 BR72_41 GDGM 43884 GDGM 44256 GDGM 50060 TFB7571 PDD:95844 TFB13998 TFB4798 TFB9629 URM 90023 URM 90022 TFB12577 WRW 08-462 TFB14043 BRNM781138 TFB11063 SFC20120802-03 SFC20140701-03 SFC20150902-50 SFC20170822-14 AWW51 SFC20120821-04 SFC20130808-08 SFC20150813-04 2.8.2012 1.7.2014 2.9.2015 22.8.2017 21.8.2012 8.8.2013 13.8.2015 Korea Belgium: Couvin Belgium: Couvin USA: North Carolina Slovakia Belgium China China China New Zealand New Zealand USA: Tennessee, USA: Georgia Brazil: Para Brazil: Para USA: Tennessee, GSMNP USA: West Virginia USA: North Carolina Portugal: Madeira USA Goseong-gun, Gangwon-do, the Republic of Korea Inje-gun, Gangwon-do, the Republic of Korea Ulleung-gun, Gyeongsangbuk-do, the Republic of Korea Ulleung-gun, Gyeongsangbuk-do, the Republic of Korea Indonesia Boryeong-si, Chungcheongnam-do, the Republic of Korea Sangju-si, Gyeongsangbuk-do, the Republic of Korea Goyang-si, Gyeonggi-do, the Republic of Korea 85 GenBank Accession Number ITS nrLSU KY026677. ~=KY026677 KY026755 KY026755 MH855896 MH867405 O1467245 O1L546542 O1467246 = OL546544 O1467247 =©01L546543 DQ450028 KJ778752 KP100303 KY074640 KY088275 MK214392 DQ450028 KJ609162 MN258643 KX513745 NR_137863 AF505762 AY842957 KY026728 KY026728 KY026736 KY026736 O1467260 O1L546549 MN413345 MN413345 MN413346 + MW396882 DQ450031 AF042650 MW405779 MW396884 MW396875 MW396875 KU057798 KU321529 KU321530 DQ450034 HQ533036 MN413331 MW396886 MN413330 MW396887 KY404982 KY404982 KY404983 = KY404983 KY026667 FJ750262 KY026765 KY026765 MW396876 MW396876 MK646034 DQ450025 OL467231 OL462786 O1L467232 OL462787 OL467234 OL546540 O1467233 OL462788 AY263428 AY639431 O1467239 =O1462813 O1467240 O1462814 OL467241 OL462815 86 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Organisms Specimen Collection Location GenBank Accession Number Date ITS nrLSU Co. utriformis TFB14334h1 USA: Connecticut KY026708 KY026708 WRW05-1170 USA: West Virginia KY026764 KY026764 Co. vellerea* NIBRFG0000502858 4.9.2018 Ongjin-gun, Incheon, the Republic of Korea 01467265 OL462791 SFC20120708-02 8.7.2012 Seosan-si, Chungcheongnam-do, the Republic = OL467266 OL462809 of Korea SFC20140821-29 21.8.2014 Gwanak-gu, Seoul, the Republic of Korea OL467267 = OL462810 SFC20180705-90 5.7.2018 Jindo-gun, Jeollanam-do, the Republic of Korea OL467268 OL462792 Co. vallianti TFB13739 USA: Tennessee, GSMNP KY026676 KY026676 Co. villosipes TFB9539 USA DQ450058 TFB12836 New Zealand: Fiordland KJ416255 FJ750264 Collybiopsis cf. SFC20180829-20 29.8.2018 Shinan-gun, Jeollanam-do, the Republic of OL467261 OL546548 ramealis Korea Rhodocollybia TFB 14382 Canada: New Brunswick KY026716 KY026716 butyracea Rhodocollybia dotae REH7007 Costa Rica AF505758 Rhodocollybia TEB 13989 USA: Mississippi KY026688 KY026688 maculata identified as members of the following genera: Gymnopus, Marasmius, or Rhodocollybia and were excluded from this study. A total of 201 specimens were segregated into 16 putative taxa based on ITS phylogenetic analyses (Table 2). To confirm the species’ identity and to infer the phylogenetic relationships within Collybiopsis, the nrLSU re- gion was amplified and sequenced from 47 representative specimens of 16 taxa (Table 1). The final phylogenetic analyses were conducted with datasets of two loci from 16 Collybiopsis species (Table 1). In ML analysis, 178 multigene sequences (110 for ITS and 68 for nrLSU) were retrieved from GenBank and used. The adjusted alignments comprised 535 to 794 bases for ITS and 324 to 904 bases for nrLSU. The phylogenetic analysis results of the two combined loci revealed that Collybiopsis specimens from the Republic of Korea were identified as 16 taxa (Fig. 1). Of the 16 putative taxa, nine matched with previously described species — Co. biformis (Peck) R.H. Petersen, Co. confluens, Co. dichroa (Berk. & M.A. Curtis) Earle, Co. luxurians (Peck) R.H. Petersen, Co. menehune (Desjardin, Halling & Hemmes) R.H. Petersen, Co. nonnulla (Corner) R.H. Petersen, Co. polygramma (Mont.) R.H. Petersen, Co. ramealis (Bull.) Earle, and Marasmiellus koreanus An- tonin, Ryoo & H.D. Pictures of basidiomata are shown in Fig. 2. The other seven taxa formed distinct clades and did not correspond to any known Collybiopsis spe- cies. Furthermore, based on the comparison with other Collybiopsis species, these seven species have distinct morphological characteristics, confirming that they were new to science — Co. albicantipes sp. nov., Co. clavicystidiata sp. nov., Co. fulva sp. nov., Co. orientisubnuda sp. nov., Co. subumbilicata sp. nov., Co. undulata sp. nov., and Co. vellerea sp. nov. Illustrations of basidiomata and micro-morphological fea- tures are shown in Figs 3 and 4. Five species (G. omphalodes Halling & J.L. Mata, G. pseudomphalodes J.L. Mata, G. ramulicola T.H. Li & S.E Deng, Ma. istanbulensis E. Sesli, Antonin and E.Aytag, and Ma. koreanus), previously placed in Gymnopus section Vestipedes, were confirmed to be- Table 2. Identification information of Korean Collybiopsis specimens confirmed in the names in bold indicate new species. Species Co. albicantipes Co. biformis Co. clavicystidiata Co. confluens Co. dichroa Co. fulva Co. koreana Co. luxurians Co. menehune Co. nonnulla Co. orientisubnuda Co. polygramma Co. ramealis Co. subumbilicata Co. undulata Co. vellerea Collybiopsis cf. ramealis SFC20170725-35 NIBRFG0000502789 KA16-0307 SFC20140724-41 SFC20180831-16 KA14-0667 KA18-0282 SFC20180713-09 Seven new Collybiopsis species Specimen Number SFC20180704-86 KA14-0259 KA16-0371 SFC20160719-42 KA14-0724 KA18-0353 NIBRFG0000508913 NIBRFG0000508991 SFC2019073 1-06 KA14-0969 KA13-0215 KA14-0386 KA16-0428 SFC20120821-84 NIBRFG0000502888 SFC20180905-86 NIBRFG0000502876 SFC20160719-15 KA13-0254 SFC20170823-39 SFC20150811-48 KA17-0600 KA16-0724 F20200701-11 KA13-0506 KA14-1089 NIBRFG0000508098 NIBRFG0000508059 NIBRFG0000508089 SFC20170822-66 SFC20130711-05 KA13-1214 SFC20140701-03 KA17-0335 SFC20150715-24 KA14-0132 KA14-0412 KA14-0734 KA14-1061 KA14-1555 KA15-0485 KA16-0252 KA16-0986 KA17-1074 KA18-0348 KA19-0125 SFC20150714-01 F20200729-14 SFC20190731-32 KA17-0344 KA13-0216 KA14-0666 KA17-0388 SFC20150702-25 SFC20190731-18 SFC20180905-43 KA13-0887 SFC20180905-33 KA13-0741 SFC20170708-14 SFC20150701-100 KA16-1154 KA15-0179 F20200630-30 KA13-0956 KA14-1092 SFC20180905-49 KA15-0173 SFC20150902-50 KA18-0651 SFC20150813-04 KA14-0163 KA14-0446 KA14-0735 KA14-1147 KA14-1558 KA15-0502 KA16-0485 KA16-0992 KA18-0089 KA18-0795 SFC20120708-02 SFC20170705-06 KA14-0526 KA16-0526 SFC20180704-36 KA15-0211 SFC20180705-84 KA16-0696 SFC20190731-34 SFC20180706-60 KA13-0333 KA14-0691 KA17-0596 SFC20170713-06 SFC20190731-08 KA18-0321 KA14-0494 KA15-0129 SFC20150902-01 QM 2020091 1-57 KA16-0925 KA14-0985 F20180904KCM21 KA13-1101 KA18-0115 SFC20180905-63 KA15-0185 SFC20170822-14 KA18-0651 KA14-0196 KA14-0447 KA14-0774 KA14-1349 KA15-0213 KA15-0527 KA16-0783 KA17-0368 KA18-0139 KA18-0836 SFC20120820-02 SFC20180705-90 KA14-0917 KA18-0657 SFC20180706-05 KA17-0287 SFC20180705-92 KA18-0338 SFC20190731-48 SFC20180712-16 KA13-0357 KA15-0210 KA18-0233 SFC20180704-17 SFC20190730-36 KA14-0579 KA14-0510 SFC20150820-59 QM20200911-52 KA16-0902 KA13-1225 F20160719-12 KA13-1333 KA18-0724 SFC20170712-08 KA15-0787 SFC20210623-03 SFC20120821-04 KA14-0245 KA14-0474 KA14-0787 KA14-1426 KA15-0215 KA15-0568 KA16-0982 KA17-0586 KA18-0151 KA18-0987 SFC20140821-29 SFC20180829-30 87 study. Scientific KA14-0924 KA18-0673 SFC20180831-13 KA17-0369 SFC20180706-04 SFC20150626-26 KA14-0168 KA16-0425 KA18-0241 SFC20180907-105 SFC20150811-29 SFC20150820-01 KA17-0787 KA16-0780 F20200730-24 KA14-0904 QM20200721-07 SFC20170807-35 SFC20120802-03 SFC20130808-08 KA14-0397 KA14-0725 KA14-1005 KA14-1475 KA15-0473 KA16-0191 KA16-0985 KA17-0742 KA18-0152 KA18-1027 SFC20150630-38 SFC20180901-01 long to the genus Collybiopsis, and we thus propose to reclassify them as Co. omphalodes comb. nov., Co. pseudomphalodes comb. nov., Co. ramulicola comb. nov., Co. istan- bulensis comb. nov., and Co. koreana comb. nov. respectively. 88 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Gymnopus menehune CUH AMO74 (India) Gymnopus menehune DEDS5866 (Hawali) 2 100} Collybiopsis menehune SFC20150811-29 Collybiopsis menehune 100 Collybiopsis menehune SFC20180905-33 Gymnopus menehune AWW 15 (Java) ‘ Gymnopus subcyathiformis TF B9629 (Puerto Rico) 100] Gymnopus subeyathiformis URM 90023 (Brazil) Gymnopus subcyathiformis URM 90022 (Brazil) Gymnopus neotropicus FB10416 (Costa Rica) 31 Gymnopus indoctus ZD16061307 (China) 91 Gymmnopus indoctus AWW 04 (Unknown) Gymnopus mesoamericanus REH7379(Costa Rica) 100'Gymnopus mesoamericanus TFB 11005 (Costa Rica) Gymnopus confluens TFB14075(USA) 100’ Gymnopus confluens TFB14409 (Canada) i Ie PEEL oasien mnopus confluens 425 (China) ji A 77 Coliybiopsis confluens SEC20190731-48 Collybiopsis confluens Gymmnopus confluens HMAS 290186 (China) Collybiopsis confluens SFC20190731-06 Gymncnus readiae PDD 95844 (New Zealand) 100" Gymnopus readiae TFB757 1 (New Zealand) Gymnopus disjunctus TFB14281 (USA) 76 94 Gymnopus disjunctus TFB14339(USA) 99) Collybiopsis subumbilicata SFC20140701-03 | Collybiopsis subumbilicata SFC20120802-03 700] Collybiopsis subumbilicata SFC20150902-50 Collybiopsis subumbilicata SFC20170822-14 85) Gymnopus biformis TFB13890(USA) 85 ymmnopus biformis TFB13814 (USA) z "eoipbogl Bs Kaos lybiopsis biformi B ‘ollybiopsis biformis | 291A collybiopsis biformis KA16-0526 Collybiopsis biformis 91 Collybiopsis biformis SFC20180831-16 100 Collybiopsis biformis SFC20180704-36 100) Collybiopsis undulata SFC20150813-04 . ; * Collybiopsis undulata SFG20130808-08 | Collybiopsis undulata * 4100 Collybiopsis undulata SFC20120821-04 G4| —100- Gymnopus villosipes TFB9539(USA) Gymnopus villosipes TFB12836 (New Zealand) Gymnopus parvulus TFB10419 (Costa Rica) 700'Gymnopus parvulus FB10422 (Costa Rica) Collybiopsis hasanskyensis TFB11846 (Russia) 100" Collybiopsis hasanskyensis TFB1 1847 (Russia) 92 100 Collybiopsis subumbilicata * 100 100 Marasmieilus vaillantii TFB13739 (USA) as Gymnopus abscuroides GB-0150514 (Norway) 700 Gymnopus eneficala MICH PK6975 (Alaska) 78'— Gymnopus eneficola 09-09-26AV13 (Canada) 100) Gymnopus nonnullus var attenuatus AWWOS5 (Java) 99 Gymnopus nonnullus var attenuatus AWW55 (Java) Gymnopus nonnullus var attenuatus RAK 369 2 (Cameroon) 100" Gymnopus nonnullus var attenuatus RAK 372 2(Cameraon) . . 99 Marasmiellus nonnullus TFB14492(USA) Col lybi opsis nonn ulla 98+ Gymnopus nonnullus TFB14278 (USA) Bo colliopsie nonnulla KA13-0741 700 Col nonnulia KA15-0126 Collybiopsis nonnulla KA13-0254 Gymnopus juniperinus TFB107682 (Argentina) 100' Marasmiellus juniperinus TFB9889 (USA) Collybiopsis polygramma SFC20170807-35 400] Collybiopsis polygramma SFC20210629-01 Collybiopsis polygramma SFC20180905-63 Gymnopus iocephalus KUC20140804-02 (Korea) Gymnopus polygrammus SFC20120821-64 (Korea) j, A Gyinnopus polygrammus HEJAUO425 (china) Collybiopsis polygramma Gymnopus polygrammus MHHNU 30912 (China) 100} = 74] -— Gymnopus polygrammus CUH AMQ82 (india) gg Gymnopus polygrammus TFB9628 (Puerto Rico) Gymnopus polygrammus URM 90015 (Brazil) Gymnopus subpruinosus TFB11063 (USA) 100 ' Gymnopus subpruinosus BRNM 781138 (Portugal) Collybiopsis clavicystidiata SFC20160705 92 ‘ollybiopsis clavicystidiata -92 | i ji ii id: * 100 100| Coilybiopsis clavicystidiata SFC20180713-09 | Collybiopsis clavicystidiata Collybiopsis clavicystidiata SFC20180705-84 1007 Gymnopus pséudoomphalodes PR24TN (Puerto Rico) Gymnopus fibrosipes FB9699 (Costa Rica) 78 Gymnopus psaudoomohelodes REH7348 (Costa Rica) 100] 738fGymnopus luxurians ZD16102301 (China) Collybiopsis luxurians SFC20190731-18 I, a é _|' Collybiopsis luxurians NIBRFGO000502888 Collybiopsis luxurians 98} _)Gymnopus luxurians TFB10350 (USA) ymnopus luxurians TFB9121(USA) Gymnopus pseudoluxurians TFB14290 (USA) Gymnopus trogicides AWW 51 (indonesia) Gymnopus brunneigracilis AW W01 (Java) Gymnopus gibbosus URM 90012 (Brazil) 100'- Gymnopus gibbosus MEL 2382838 (Australia) 100 Gymnopus melanopus SFSU AWW54 (Java) Gymnopus melanopus CUH AMO93 (India) Collybiopsis utriformis TFB14334 (USA) 100° Collybiopsis utriformis WRW 05-1170(USA) ; 95;- Gymnopus ramulicola GDGM 44256 (China) 82 Gymnopus ramulicola GDGM 50060 (China) 100! Gymnopus ramulicola GDGM 43884 (China) 98] Collybiopsis fulva KA13-0333 i 4 . 98lf Collybiopsis fulva KA13-0216 | Collybiopsis fulva * 717 Collybiopsis fulva KA15-0210 96) Gymnopus dichrous TENN60014c2(USA) Gymnopus dichrous TFB7920 (USA). Gymnopus dichrous TENN61624c1a (USA) 95 Collybiopsis dichroa KA14-0969 Collybiopsis dichroa SFC20180712-16 j ji i Collybiopsis dichroa KA18-0389 Collybiopsis dichroa 89) Gymnopus dichrous TENN60014c5 (USA) 97| Gymnopus dichrous TFB2028 (USA) Gymmnopus dichrous TENN61624c9 (USA) a0] 8) _ | Marasmiellus dichrous TFB9623 (USA) Marasmielius istanbulensis KATO fungi 3596 (Turkey) 90 + Marasmiellus istanbulensis BRNM 781163 (Turkey) Gymnopus micromphalecides TENN 68165 (USA) 100° Gymnopus micromphalecides TFB 14282 (USA) Collybiopsis quercophila 1FB14615 (USA) 00 Collybiopsis quercophila TFB14570 (Slovakia) Collybiopsis minor TFB5434 (USA) 100'— Collybiopsis minor TFB11930 (USA) Collybiopsis koreana SFC20150721-10 96 Coulyblopsis Korsana eit ee ; Cc II bi :. k jarasmiellus koreanus orea Too) | Collybiopsis koreana SFC20120821- 84 OnYBIOpSIs KOfeana a7 700 Marasmiellus koreanus BRNM 718782 (Korea) Collybiopsis albicantipes SFC20170725-35 A H A . ES 700! Collybiopsis albicantipes SFC20180704-86 | Collybiopsis albicantipes 7Qy Collybiopsis vellerea SFC20120708-02 Collybiopsis vellerea NIBRFG0000502858 00! Collybiopsis vellerea SFC20140821-29 100) 100! Collybiopsis vellerea SFC20180705-90 100} Gymnopus omphalodes FB11021 (Costa Rica) 400 Gymnopus omphalodes TENN56734 (Costa Rica) 100- Gymnopus omphalodes TFB10427 (Costa Rica) 99) Marasmiellus subnudus TFB14043 100 96)" Marasmiellus subnudus TFB14043 Gymnopus subnudus W RW 08-462 CL. Gymnepus subnudus TFB12577 Gymnopus peronatus LE-BIN1364 100] 79q| Gymnopus peronatus TFB13743 Gymnopus peronatus CBS 223 37 91] [ Collybiopsis orientisubnuda NIBRFG0000500990 Collybiopsis orientisubnuda SFC20170823-39 Collybiopsis orientisubnuda SFC20180830-29 - Gymnopus subnudus KUC20150911-19 Marasmiellus stenophylius TFB13998 100! Marasmiellus Stenophyllus TFB4798 Marasmiellus folliiphilus CUH AM101 a8 100! Marasmiellus folliphilus CUH AMOSO 100 —_ Collybiopsis californica TENN-F-052617 100 Collybiopsis filamentipes TFB13962 99 Collybiopsis cf. ramealis SFC20180829-20 Collybiopsis furtiva TFB4796 me iy Mare ens faineale vie NIBREGOOOOSOBaas ollybiopsis ramealis i, i i 99 Marasmietas ramealis TFB 13770 Colly biop: sis ramealis 95|_| Marasmiellus ramealis TFB13769 99 Collybiopsis ramealis TFB14555 Collybiopsis ramealis BR72-41 Rhodocollybia maculata TFB13989 F o7 Rhadocollybia dotae REH7007 70 Rhodocollybia butyracea TFB14382 100 Collybiopsis vellerea * * Collybiopsis orientisubnuda 0.05 Figure |. Phylogenetic tree based on maximum likelihood analysis using combined sequence data of ITS and nrLSU. ML bootstrap values greater than 70% are indicated at the nodes. Collybiopsis species that were newly sequenced in this study are represented in bold. Species with an asterisk are those proposed as new species. Seven new Collybiopsis species 89 PN Figure 2. Basidiomata of the described Collybiopsis species in the Republic of Korea A Co. biformis (SFC20180706-05) B Co. confluens (SFC20190731-06) C Co. dichroa (KA18-0389) D Co. koreana (SFC20180704-17) E Co. luxurians (SFC20190731-18) F Co. menehune (SFC20150811-29) G Ca. nonnulla (KA13—0254) H Co. polygramma (SFC20170712-08) I Co. ramealis (SFC20180829-—20). Scale bar: 1 cm (A=). Taxonomy Collybiopsis albicantipes J.S. Kim & Y.W. Lim, sp. nov. MycoBank No: 842053 Fig. 3A—B, Suppl. material 1: Fig. SIA Etymology. Epithet “a/bicantipes” refers to having a whitish base of the stipe. Holotype. The Republic of Korea, Jeollanam-do: Yeosu-si, Dolsan-eup, Hyangi- ram, 34°35'27"N, 127°47'55"E, alt. 183 m, 25 July 2017, Jae Young Park, Komsit Wisitrassameewong, SFC20170725-35 (GenBank accession no. ITS: OL467272; nrLSU: OL462811). Diagnosis. ‘This species notably has hemispherical to convex, 4-23 mm pileus, distant lamellae, central to eccentric, tomentose, 5-15 x 0.5-1.5 mm stipe with a white base; ellipsoid to ovoid, 5.8—7.4 x 2.84 um basidiospores, clavate (often con- stricted), 25.5—34.8 x 4.8-6.7 um basidia, broadly clavate, irregular, sometimes lobed, 26-49 x 5.4—10.6 um cheilocystidia, and a habit of fruiting on branches. Description. Pileus: 4-23 mm, eccentric, convex to hemispherical when young, becoming depressed and undulating with age; Surface smooth, brownish orange (5C3 to 6D4) at the center, becoming paler to the margin (4A3 to 3A2). Lamellae: distant, 90 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Figure 3. Basidiomata and microscopic characters of the four new Collybiopsis species A, B Co. albican- tipes (SFC20170725-35) C, D Co. clavicystidiata (SFC20180705—84) E, F Co. fulva (KA15—0210) G, H Co. orientisubnuda (NIBRFG0000502862). Scale bars: lcm (A, C, E, G); 20 4m (B, D, F, H). Abbrevia- tions: s basidiospores; b basidia; ch cheilocystidia; p pleurocystidia; ca caulocystidia. Seven new Collybiopsis species a! L= 10-16, | = 3-7, adnate, whitish to yellowish white (3A2). Stipe: 5-15 x 0.5-1.5 mm, central to eccentric, cylindrical, tomentose, apex brownish orange (5C3) to light brown (6D4), gradually becoming paler downwards (5B2 to 6C2), with whitish basal tomen- tum. Basidiospores: 5.8—7.4 x 2.8—4 um (average 5.5 x 3.2 um), Q= 1.6—2.1 (mean = 1.97), ellipsoid to ovoid, amygdaliform, smooth, hyaline, non-dextrinoid, with drops. Basidia: (23) 25.5—34.8 x 4.86.7 (7) um, 4-spored, clavate, often constricted. Cheilo- cystidia: 26-49 x 5.4—10.6 (14) um, broadly clavate, irregular, sometimes lobed. Pleu- rocystidia: 25.8-56.4 (62) x 6.2—12.5 um, clavate, subulate, sometimes lobed. Trama hyphae: cylindrical, often sub-inflated, smooth, non-dextrinoid 1.7—9 (12) um wide. Pileipellis: a cutis made up of cylindrical, often sub-inflated, with weak annular orna- mentation, 2.0—-7.5 um wide hyphae; terminal elements adpressed, cylindrical, clavate, sometimes constricted or curved, 2.0-5 um wide. Stipitipellis: a cutis of cylindrical, smooth, 2.7-9.7 (11) um wide hyphae. Caulocystidia: 21.7—90 x 3.9-11.7 um, cylin- drical, flexuose, sometimes curved. Clamp connections: present in all tissues. Other specimens examined. The Republic of Korea, Jeollanam-do: Jindo-gun, Maenggoldo island, 34°12'21"N, 125°51'41"E, alt. 24 m, 4 July 2018, Jae Young Park, SFC20180704—86. Habit and habitat. Scattered to gregarious on the branch in mixed forest domi- nated by Camellia japonica Linne, in summer. Distribution. The Republic of Korea. Remark. Collybiopsis albicantipes is similar to Co. ramulicola and Co. koreana when comparing macro-morphological characteristics. Collybiopsis ramulicola is dis- tinguishable from Co. albicantipes by a reddish pileus, fewer and buff lamellulae (1-4), a shorter and thinner stipe (12-23 x 2-3 mm), shorter and slightly elongated basidi- ospores (6.6—-8.4 x 3.5—-4.5 um), shorter basidia (23-27 x 3.8-5.5 um), and shorter cheilocystidia (23—27 x 3-6 um) (Deng et al. 2016). Collybiopsis koreana differs from Co. albicantipes by having a larger pileus (27-60 mm), more lamellae (15-20) and lamellulae (2—3), longer and thicker stipe (14-70 x 2-3.5 um), bigger and elongated basidiospores (7.5-10 x 4—5 um), cheilocystidia with different shapes and sizes (25— 55 x 4-10 um), and incrustation dark brown in KOH (Antonin et al. 2010). Collybiopsis clavicystidiata J.S. Kim & Y.W. Lim, sp. nov. MycoBank No: 842054 Fig. 3C—D, Suppl. material 1: Fig. S1B Etymology. Epithet “clavicystidiata’ indicates that the new species has clavate cheilocystidia. Holotype. The Republic of Korea, Jeollanam-do: Jindo-gun, Jodo-myeon, Donggeocha island, 34°23'34"N, 125°93'84"E, alt. 70 m, 05 July 2018, Jae Young Park, Tae Heon Kim, SFC20180705-—84, (GenBank accession no. ITS: OL467252; nrLSU: OL462817). Diagnosis. The prominent features of this species include a greyish orange to brownish, 6-45 mm pileus, whitish lamellae, a subinstitious, tomentose, whitish, 15— 92 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) 26 x 1.2-1.6 mm stipe, oblong to subcylindrical, 6.7—9.4 x 3.1-4.6 um basidiospores, utriform, clavate, 20.1-37.5 x 6.8-12.2 um cheilocystidia, and cylindrical, flexuose, irregular, 17-50 x 3.5—7 um caulocystidia. Description. Pileus: 6-45 mm, convex to hemispherical, becoming plano-convex to flat with an uplifted margin with age; Surface smooth, dull, hygrophanous, greyish orange (6B3) to brownish (7D8 to E8) at the center, being whitish at the margin (4A2 to 6C8), being paler with age. Lamellae: subdistant, L = 20-32, | = 1-7, adnexed, white. Stipe: 15-26 x 1.2-1.6 mm, cylindrical, tomentose, subinsititious, whitish to reddish grey (9B2). Basidiospores: 6.7-9.4 x 3.1-4.6 um, average 8.13 x 3.62 um, Q = 22.4 (mean = 2.26), oblong to cylindrical, smooth, hyaline, non-dextrinoid, with drops. Basidia: 18.3—30 x 4.1-8.8 um, 4-spored, narrowly clavate, narrowly utri- form, often curved. Cheilocystidia: 20.1-37.5 x 6.8—12.2 um, utriform, clavate, some- times with mucronate apex. Pleurocystidia: absent. Trama hyphae: cylindrical, often subinflated, smooth, branched, non-dextrinoid, 2-12 um wide. Pileipellis: transition between cutis and trichoderm, composed of cylindrical, with heavy annular ornamen- tation, 4-12 um wide hyphae; terminal elements adpressed to suberect, cylindrical, clavate, often incrusted (often incrusted), thin-walled, 3-6 um wide. Stipitipellis: a cutis of cylindrical, smooth, 2-7 um wide hyphae. Caulocystidia: 17-50 x 3.5—7 um, cylindrical, flexuose, irregular or curved. Clamp connections: present in all tissues. Other specimens examined. The Republic of Korea, Jeollanam-do: Haenam-gun, Mt. Duryun, 34°29'6"N, 126°38'54"E, alt. 169 m, 5 July 2018, Young Woon Lim, Abel Severin Lupala, Jun Won Lee, SFC20180705—26. The Republic of Korea, Seoul: Gwanak-gu, Gwanak-ro 1, Seoul National University, 37° 27' 37"N, 126° 56' 59"E, alt. 80m, 13 July 2018, Jae Young Park, SFC20180713-09. Habit and habitat. Solitary to scattered on dead wood debris of conifers, in summer. Distribution. The Republic of Korea Remark. Collybiopsis clavicystidiata is morphologically similar to G. omphalodes and Co. menehune. Collybiopsis omphalodes differs in their larger pileus (2-30 mm), a darker colored stipe, smaller basidiospores (5-6 x 2.5—3 um), and thinner hyphae in the pileipellis (S—8 um wide). Collybiopsis menehune can be distinguished from Co. clavicystidiata by its larger pileus (8-30 mm), buff lamellae, longer stipe (15-60 mm), longer basidiospores (7.5—9.5 x 3.5—4.2 um, Q = 2.2), and longer caulocystidia (16— 67 x 3-5 um) (Desjardin et al. 1999). Co. clavicystidiata is phylogenetically close to Co. pseudomphalodes. Collybiopsis pseudomphalodes has relatively few references for com- parison, but differences can be found in the lengths of the stipe (3—4 mm) and cheilo- cystidia (40 x 3 um) when compared with Co. clavicystidiata (Dennis 1961). Collybiopsis fulva J.S. Kim & Y.W. Lim, sp. nov. MycoBank No: 842055 Fig. 3E-E Suppl. material 1: Fig. S1C Diagnosis. This species has a pale orange to brownish-colored, 4-20 mm pileus, an or- ange white colored to light brownish colored, 7-30 x 0.7—1 mm stipe with pubescence, Seven new Collybiopsis species 93 spheropedunculate, pleurocystidia, oblong to subcylindrical, 6.8—9.2 x 3.1-4.9 um ba- sidiospore, lobed, clavate with rostrate apex, 24.8—38.4 x 6.5-11.8 um cheilocystidia. Etymology. Epithet “fulva” referring to fox-colored pileus. Holotype. The Republic of Korea, Gyeonggi-do: Pocheon-si, Soheul-eup, Gwangneungsumogwon-ro 415, 37°45'17"N, 127°9'59"E, alt. 101 m, Sang Kook Han, 21 July 2015, KA15—0210 (GenBank accession no. ITS: OL467259; nrLSU: OL462795). Description. Pileus: 4-20mm, hemispherical, convex to plane, sometimes concave with slightly reflexed, wavy margin, hygrophanous, pale orange (6A3) to greyish orange, becoming more brownish to the center (5B4 to 7C4). Lamellae: distant, L = 16-28, / = 1-5, sinuate, broad, whitish to yellowish white (4A2) to brownish orange (6C4 to 7C4). Stipe: 7-30 x 0.7—-1 mm, cylindrical, gradually widened towards the base, tomentose, apex orange white (5A2) to brownish orange (6C6), becoming dense downwards (6D8), covered with pubescence. Basidiospores: 6.8-9.2 x 3.1-4.9 um (average 7.47 x 3.69 um), Q = 2.05, oblong to cylindrical, smooth, colorless, non-dextrinoid, with drops. Basidia: 20.4-29.4 x 4.7—7.8 um, 4-spored, narrowly clavate, sometimes constricted or curved. Cheilocystidia: (20.5) 24.8—38.4 x 6.5-11.8 um, lobed, clavate, sometimes with rostrate apex. Pleuro- cystidia: 31.5—46.9 x 12—20.6 um, spheropedunculate, obovoid, sometimes with mucronate apex. Trama hyphae: cylindrical to subinflated, irregular, thin-walled, smooth, branched, non-dextrinoid, 2.0-15 um wide. Pileipellis: a cutis of cylindri- cal, thin-walled, 4-15 um wide hyphae; terminal elements adpressed to suberect, narrowly clavate, thin-walled, with heavy annular ornamentation, 3-8 um wide. Stipitipellis: a cutis of cylindrical, thin-walled, smooth, 5-15 um wide hyphae. Cau- locystidia: 45.6-108.3 (131) x 6.8-14.8 um, cylindrical, irregular, curved. Clamp connections: present in all tissues. Other specimens examined. The Republic of Korea, Gyeonggi-do: Pocheon-si, Soheul-eup, Gwangneung forest exhibition hall, 37°45'19"N, 127°9'58"E, alt. 99 m, 8 July 2016, Sang Kook Han, KA16—0428. The Republic of Korea, Gyeongsangnam- do: Geochang-gun, Mt. Gibaek, 35°43'6"N, 127°45'49"E, alt. 1095 m, 19 June 2013, Sang Kook Han, KA13—0216. Habit and habitat. Scattered or gregarious on the bark of deciduous trees or on the rotting branch of both broadleaf trees and conifers, in summer. Distribution. The Republic of Korea. Remark. Collybiopsis fulva morphologically resembles Co. menehune and Co. ramealis. They can be distinguished based on several morphological differences. Collybiopsis menehune has a longer stipe (15-60 mm length), denser lamellae, and larg- er basidiospores (7.5—9.5 x 3.5-4.2 um) (Desjardin et al. 1999). Collybiopsis ramealis has a smaller basidiocarp (2-20 mm), shorter basidiospores (7.8—11 x 2.5-4 mm) and different type of pileipellis (Rameales-structure) (Noordeloos 1983; Desjardin et al. 1997). Phylogenetically, Co. fulva is closely related to Co. ramulicola. Collybiopsis ramu- licola differs in having a more yellowish pileus, fewer lamellae (9-12) that are brighter in color, a more reddish and thicker stipe (2-3 mm), and smaller sized cheilocystidia (23-27 x 3-6 mm) (Deng et al. 2016). 94 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Collybiopsis orientisubnuda J.S. Kim & Y.W. Lim, sp. nov. MycoBank No: 842056 Fig. 3G—H, Suppl. material 1: Fig. S1D Etymology. Epithet “orientisubnuda” meaning the new species has originated from the East and is morphologically similar to Co. subnuda. Holotype. The Republic of Korea, Gyeongsangbuk-do: Ulleung-gun, 37°31'21"N, 130°53'14"E, alt. 757 m, 19 July 2016, Changmu Kim, Jinsung Lee, Jae Young Park, NIBRFG0000500990 (GenBank accession no. ITS: OL467262; nrLSU: OL546546). Diagnosis. It features a brownish, 15-50 mm pileus, orangish cream-colored la- mellae, greyish to brownish orange, tomentose, 20-80 x 2.5-6 mm stipe, subcylindri- cal to fusoid, 6.7—8.6 x 1.8—3.2 um basidiospores, and cylindrical, flexuose, sometimes irregular or curved, 26.3—52 (63) x 3.5—6.5 um caulocystidia. This species is morpho- logically similar to Co. subnuda. Description. Pileus: 15-50 mm, convex to plano-convex, sometimes subum- bonate; Surface smooth, brownish orange (6C5 to 7C4), becoming paler to the margin (5A2). Lamellae: distant, L = 16-28, | = 3-7, adnexed, pale yellow (4A3) to orange white (5A2). Stipe: 20-80 (100) x 2.5-6 mm, central to eccentric, cylin- drical, tomentose, often twisted, greyish orange (6B4) to brownish orange(7C4), becoming paler and thinner to the base. Basidiospores: 6.7—8.6 x 1.8—3.2 um (av- erage 7.5 x 2.5 um), Q = 2.5-3.2 (mean = 2.92), cylindrical to fusoid, smooth, hyaline, non-dextrinoid, with drops. Basidia: (17) 19.8-28.7 (29) x 3.7-7.3 um, 4-spored, narrowly clavate, often constricted. Cheilocystidia: variable in shape and size, 21-33.3 x 4.7-8.2 um, lobed, clavate, slightly sphaeropendunculate, some- times constricted or with rostrate apex. Pleurocystidia: 24.7-52.3 x 5.1-9.1 um, narrowly utriform, clavate, sometimes clavate with rostrate apex. Trama hyphae: cylindrical, often subinflated, smooth, branched, non-dextrinoid, 2.0—7.0 um wide. Pileipellis: a cutis made up of cylindrical, 2-8 um wide hyphae; terminal elements adpressed, cylindrical, often subinflated, with weak annular ornamentation, 3-6 um wide. Stipitipellis: a cutis of cylindrical, smooth, 2.5-7 um wide hyphae. Caulo- cystidia: 26.3-52 (63) x 3.5-6.5 um, cylindrical, flexuose, sometimes irregular or curved. Clamp connections: present in all tissues. Other specimens examined. The Republic of Korea, Chungcheongnam-do: Yesan-gun, Mt. Gaya, 35°48'14"N, 128°5'49"E, alt. 863 m, 23 August 2017, Hae Jin Cho, Ki Hyeong Park, SFC20170823-—39. The Republic of Korea, Gangwon-do: Pyeongchang-gun, Mt. Odae, 37°43'54"N, 128°35'42"E, alt. 683 m, 8 July 2017, Nam Kyu Kim, SFC20170708-14. The Republic of Korea, Gyeongsangbuk-do: Ulle- ung-gun, 37°31'30"N, 130°52'21"E, alt. 718 m, 2 September 2015, Jae Young Park, SFC20150902-01. Habit and habitat. Scattered to gregarious on the ground covered with dead and decaying leaves of broadleaf forest, from summer to autumn. Distribution. The Republic of Korea. Seven new Collybiopsis species 95 Remark. Collybiopsis orientisubnuda is morphologically similar to Co. peronata (Bolton) R.H. Petersen and Co. subnuda (Ellis ex Peck) R.H. Petersen. Collybiopsis peronata can be distinguished from Co. orientisubnuda by fewer and buff lamellulae (1-3), a thicker stipe (3-8 mm), smaller Q value (2.3), longer basidia (20-40 um), and longer cheilocystidia (25-90 x 5-10 um) (Noordeloos et al. 1999). Collybiopsis subn- uda differs from Co. orientisubnuda with thinner stipe (~3 mm), larger basidiospores (8-11 x 3—4.5 um) and the absence of pleurocystidia (Tekpinar and Acar 2020). Collybiopsis subumbilicata J.S. Kim & Y.W. Lim, sp. nov. MycoBank No: 842057 Fig. 4A, B, Suppl. material 1: Fig. SIE Etymology. Epithet “swbumbilicata” referring to having a small depressed center in pileus. Holotype. The Republic of Korea, Seoul, Gwanak-gu, Mt. Gwanak, 37°12'39"N, 128°19"E, alt. 877 m, 01 July 2014, Young Woon Lim, SFC20140701-03 (GenBank accession no. ITS: OL467232; nrLSU: OL462787). Diagnosis. The distinctive features include a brownish, 10-35 mm pileus, white colored lamellae, a brownish, 25-60 x 1-3 mm stipe covered with pubescence, ellipsoid to oblong basidiospores, narrowly clavate and cylindrical, 17—24.3 x 3.5—5.1 um basid- ia, and cylindrical, flexuose, sometimes curved, 12.6—38.2 x 2.4—6.6 um caulocystidia. Description. Pileus: 10-35 mm, plano-convex to plano-concave, subumbili- cate, becoming undulate and uplifted in age; Surface smooth, greyish orange (5B3) to brown (GE5). Lamellae: subdistant, L = 22—38, | = 3-7, free to adnexed, white. Stipe: 25-60 x 1-3 mm, cylindrical, tomentose, hollow, light brown (7D4) to dark brown (9F8), becoming paler to the apex, covered with pubescence. Basidiospores: 5.5-7.5 x 2.5-3.6 pm (average 6.47 x 3.0 um), Q = 1.8-2.2 (mean = 2), oblong to fusiform, smooth, hyaline, non-dextrinoid, with drops. Basidia: (15.6) 17—24.3 (27.6) x 3.5-5.1 (5.9) um, 4-spored, narrowly clavate, cylindrical. Cheilocystidia: 17.6— 38.4 x 5—7.8 pm, various in shape, lobed. Pleurocystidia: 20.3-30.7 x 6.8-9.5 um, clavate, fusiform, slightly sphaeropedunculate. Trama hyphae: cylindrical, subinflated, branched, smooth, non-dextrinoid, 1.5-8 um wide. Pileipellis: a cutis made up of cy- lindrical, often incrusted, with heavy annular ornamentation, 5.0-15 um wide hyphae; terminal elements adpressed to suberect, fusoid, clavate, 6.0—16 um wide. Stipitipellis: a cutis of cylindrical, smooth, thin-walled, 2.0-6.0 um wide hyphae. Caulocystidia: 12.6-38.2 x 2.4—6.6 pm, cylindrical, flexuose, sometimes irregular or curved. Clamp connections: present in all tissues. Other specimens examined. The Republic of Korea, Gangwon-do: Goseong-gun, Hwajinpo, Hwajinpo Condominium, 38°28'24"N, 128°26'30"E, alt. 7 m, 2 August 2012, Young Woon Lim, SFC20120802-—03. The Republic of Korea, Gyeongsangbuk- do: Ulleung-gun, Ulleung island, 37°30'38"N, 130°51'44"E, alt. 429 m, 22 August 2017, Jae Young Park, Nam Kyu Kim, SFC20170822-14. 96 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Figure 4. Basidiomata and microscopic characters of the three new Collybiopsis species A, B Co. subum- bilicata (SFC20120802-03) C, D Co. undulata (SFC20150813—04) E, F Co. vellerea (SFC20140821- 29). Scale bars: 1cm (A, C, E); 20 ym (B, D, F). Abbreviations: s basidiospores; b basidia; ch cheilocyst- idia; p pleurocystidia; ca caulocystidia. Habit and habitat. Scattered to gregarious on the ground covered with dead leaves in temperate mixed forests, from summer to autumn. Distribution. The Republic of Korea. Remark. Collybiopsis subumbilicata appears similar to Co. villosipes (Cleland) R.H. Petersen. Collybiopsis villosipes is distinguished from Co. subumbilicata by fewer and brownish lamellae (also lamellulae), a noninsititious, light-colored stipe, larger basidi- ospores (6.5-10.5 x 3.5-4.5 um) and basidia (25-34 x 6.5-7.5 um) (Desjardin et al. Seven new Collybiopsis species 97 1997). Furthermore, Co. subumbilicata is phylogenetically close to Co. biformis and Co. disjuncta (R.H. Petersen & K.W. Hughes) R.H. Petersen & K.W. Hughes. Col lybiopsis biformis is morphologically similar to Co. subumbilicata but can be distin- guished by elongated basidiospores (6.4-9.2 x 2.4-4.8 um), thicker basidia (6-7 um thick) and cheilocystidia (6-12 ym thick) (Morgan 1905; Mata 2002). Collybiopsis disjuncta can be distinguished from Co. subumbilicata by a smaller pileus (7-12 mm) with olivaceous tint, pinkish lamellae, slender stipe (0.5—1 mm thick), bigger basidi- ospores (6—7.5 x 3—3.5 um), bigger basidia (22—34 x 5—7 um), and a seldom incrusted pileipellis (Petersen and Hughes 2014). Collybiopsis undulata J.S. Kim & Y.W. Lim, sp. nov. MycoBank No: 842058 Fig. 4C—D, Suppl. material 1: Fig. S1F Etymology. Epithet “undulata” referring to having an undulate margin of pileus. Holotype. The Republic of Korea, Chungcheongnam-do, Boryeong-si, recrea- tion forest of Mt Sungju, 36°20'4"N, 126°39'50"E, alt. 241 m, 21 August 2012, Jae Young Park, SFC20120821-—04 (GenBank accession no. ITS: OL467239; nrLSU: OL462813). Diagnosis. It is characterized by having 10-23 mm sized pileus that is particularly brown in the middle with a wavy margin, subdistant and creamy lamellae, a dark brown, 35— 55 x 0.8-2 mmstipe that becomes lighter to the apex, subcylindrical, broadly clavate or irregu- lar, sometimes lobed, 16.7—28 x 4.8—8 um cheilocystidia, and 27—60 x 3.5—6 pm sized cau- locysitida which has a morphology similar to cheilocystidia and sometimes grows in bundles. Description. Pileus: 10-23 mm, convex to concave, margin becoming undulate with age; Surface smooth, hygrophanous, brown (7D2 to 7E6) in the center, becom- ing paler to the margin (5A2—5B3 to 7B2). Lamellae: subdistant, L = 15-30, | = 3-9, adnexed, cream. Stipe: 35-55 x 0.8-2 mm, cylindrical, tomentose, dark brown (7F5 to 8F8), gradually becoming paler to apex (7B2 to 7C2). Basidiospores: 5.6-9.5 x 2—3.4 um (average 7.3 x 2.8 um), Q = 2—3.1 (mean = 2.58), cylindrical, smooth, hya- line, non-dextrinoid, with drops. Basidia: 15—22.3 x 3.6-6.8 um, 4-spored, cylindri- cal, narrowly clavate to utriform, often curved. Cheilocystidia: 16.7—28 x 4.8—-8 um, subcylindrical, broadly clavate or irregular, sometimes lobed. Pleurocystidia: absent. Trama hyphae: cylindrical, sometimes subinflated, smooth, branched, non-dextri- noid, 2—8 um wide. Pileipellis: a cutis made up of cylindrical, often incrusted, slightly brownish, with heavy annular ornamentation, 2.4—7 um wide hyphae; terminal ele- ments adpressed to suberect, cylindrical to clavate, 3-6 um wide. Stipitipellis: a cutis of cylindrical, smooth, 2.0-3.5 um wide hyphae. Caulocystidia: 27-60 x 3.5—6 um, irregularly cylindrical, narrowly utriform, seldom apically lobed, sometimes gathered in a bunch. Clamp connections: present in all tissues. Other specimens examined. The Republic of Korea, Gyeonggi-do: Goyang-si, Deog- yang-gu, Seooreung, 37°37'26"N, 126°54'4"E, alt. 35 m, 13 August 2015, Jae Young Park, 98 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) SFC20150813-04. The Republic of Korea, Gyeongsangbuk-do, Sangju-si, Mt Noheum, 36°26'20"N, 128°5'48"E, alt. 695 m, 8 August 2013, Jae Young Park, SFC20130808-08. Habit and habitat. Scattered to gregarious on leaf litter in mixed forest dominated with broadleaf trees, in summer. Distribution. The Republic of Korea. Remark. Collybiopsis undulata is morphologically similar to Co. subpruinosa (Mur- rill) R.H. Petersen. Collybiopsis subpruinosa has differences in having small central pa- pilla on pileus, fewer lamellulae (3-4 series), vivid colored lamellae, thicker basidi- ospores (4.5—5.2 um wide), larger basidia (30-36 x 7.5—8.5 um) and cheilocystidia (25-80 x 5-16 um), thick-walled trama hyphae (0.5—1 um), caulocystidia with a wid- er size range, and a habit of growing solitary on rotten twigs or logs (Desjardin et al. 1999). Collybiopsis undulata is phylogenetically close to Co. villosipes but Co. villosipes can be differentiated by having fewer lamelluale (2—3 series), vivid colored lamellae, thicker stipe (1.5—4.0 mm), slightly thicker basidiospores (3.5—4.5 um wide), and ba- sidia (25-34 x 6.5-7.5 um) (Desjardin et al. 1997). Collybiopsis vellerea J.S. Kim & Y.W. Lim, sp. nov. MycoBank No: 842059 Fig. 4E—-E Suppl. material 1: Fig. S1G Etymology. Epithet “vellerea” refers to having a velvety stipe. Holotype. The Republic of Korea, Seoul: Gwanak-gu, Mt. Gwanak, 37°27'32"N, 126°56'49"E, alt. 90 m, 21 August 2014, Young Woon Lim, SFC20140821—29 (Gen- Bank accession no. ITS: OL467267; nrLSU: OL462810). Diagnosis. It has a dull, greyish orange, 18-45 mm pileus with darker center, a tomen- tose (like velvet), insititious, orangish, 15—55 x 3-5 mm stipe, sphaeropendunculate, subo- void, 23.449 x 7.5—13.4 um pleurocystidia, oblong to subcylindrical basidiospores, nar- rowly clavate with rostrate apex, sometimes lobed, 7.7-49.7 x 3.8-14.6 um cheilocystidia. Description. Pileus: 18-45 mm, hemispherical, appendiculate to convex, subum- bonate with an uplifted margin when old; Surface smooth, dull, hygrophanous, orange white (5A2) to greyish orange (6E8 to 7F8) on the center, gradually becoming paler to, the,edge.(5A) to 5B2)? Lamellae: crowded to-close, £:= 38-52, 7 = 3-7, hurcate, white. Stipe: 15-55 x 3-5 mm, cylindrical, finely tomentose, insititious, pale orange (5A3) to reddish grey (7B2), becoming darker to the base (GA2 to 7C2). Basidiospores: 5.2-7 x 2.5-3.8 um (average 6.17 x 3.06 um), Q = 1.8—2.4 (mean = 2.03), oblong to subcylindrical, smooth, hyaline, non-dextrinoid, with drops. Basidia: 16.2—24.8 x 3.3— 5.3 um, 4-spored, (narrowly) clavate, often curved or constricted. Cheilocystidia: 7.7-49.7 x 3.8-14.6 um, narrowly clavate with rostrate apex, sometimes lobed. Pleu- rocystidia: 23.4—49 x 7.5-13.4 um, sphaeropendunculate, subovoid. Trama hyphae: cylindrical, often subinflated, thin-walled, smooth, branched, non-dextrinoid, 2-5 um wide. Pileipellis: a cutis made up of cylindrical, thin-walled, with weak annular orna- mentation, 3-10 um wide hyphae; terminal elements adpressed to suberect, cylindrical, fusoid, clavate, 5-11 um wide. Stipitipellis: a cutis of cylindrical, thin-walled, smooth, Seven new Collybiopsis species 99 2.0-6.0 um wide hyphae. Caulocystidia: 12-38 x 2.4-6.6 um, cylindrical, narrowly utriform, sometimes irregular, or curved. Clamp connections: present in all tissues. Other specimens examined. The Republic of Korea, Chungcheongnam-do: Seosan- si, Mt. Gaya, 36°41'0"N, 126°35'19"E, alt. 260 m, 20 August 2012, Jae Young Park, SFC20120820-02. The Republicof Korea, Incheon: Ongjin-gun, 37°13'10"N, 126°10'4"E, alt. 6 m, 4 September 2018, Changmu Kim, Jin Sung Lee, NIBRFG0000502858. The Re- public of Korea, Jeollanam-do: Jindo-gun, Seogeocha island, 34°15'22"N, 125°55'11"E, alt. 38 m, 5 July 2018, Jae Young Park, Tae Heon Kim, SFC20180705-90. Habit and habitat. Scattered to gregarious on the ground covered with dead and decaying conifer needles, from summer to autumn. Distribution. The Republic of Korea. Remark. Collybiopsis vellerea is morphologically similar to Co. menehune and G. spon- giosus Halling. Collybiopsis menehune has a paler stipe, a smaller pileus (3-30 mm), and fewer lamellulae (4-6 series) (Desjardin et al. 1999). Gymnopus spongiosus has a smaller pi- leus (8-20 mm) and longer stipe (20-55 mm). Micromorphologically, Co. menehune has larger basidiospores, basidia, and caulocystidia (Desjardin et al. 1999). Gymnopus spon- giosus differs from Co. vellerea in that its pileipellis is a Dryophila-type cutis and its color changes in alkalies. Furthermore, its basidia (18—25 x 6—9 um) and trama hyphae (3.5-17 um) are thicker and its caulocystidia (3.5-10.5 um broad) are smaller (Halling 1996). Collybiopsis vellerea is phylogenetically close to Co. omphalodes. Collybiopsis omphalodes differs in having smaller basidiomata (20-30 mm) and its habit on logs (Dennis 1951). Proposal for Collybiopsis recombination In this study, many epithets were found that required an additional transfer of species from Marasmiellus to Collybiopsis apart from the study done by Petersen and Hughes (2021). Oliveira et al. (2019) had previously suggested to replace these species from Gymnopus to Marasmiellus s. str., but this study suggests that these species should be further transferred from Marasmiellus s. str. to Collybiopsis. Collybiopsis istanbulensis (E.Sesli, Antonin & E.Aytag) J.S. Kim & Y.W. Lim, comb. nov. MycoBank No: 842060 Basionym. Marasmiellus istanbulensis E. Sesli, Antonin & E.Aytag. Pl. Biosystems 152(4): 669. 2018. Collybiopsis koreana (Antonin, Ryoo & H.D.Shin) J.S. Kim & Y.W. Lim, comb. nov. MycoBank No: 842061 Basionym. Marasmiellus koreanus Antonin, Ryoo and H.D.Shin. Mycotaxon 112: 190. 2010. 100 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Collybiopsis omphalodes (Berk.) J.S. Kim & Y.W. Lim, comb. nov. MycoBank No: 842062 Chamaeceras omphalodes (Berk.) Kuntze, Revis. gen. pl. (Leipzig) 3(3): 456. 1898. Collybia omphalodes (Berk.) Dennis, Trans. Br. mycol. Soc. 34(4): 443. 1951. Marasmiellus omphalodes (Berk.) Singer. Sydowia 9(1—G): 385. 1955. Gymnopus omphalodes (Berk.) Halling & J.L. Mata, in Mata, Halling, and Petersen, Fungal Diversity 16: 122. 2004. Basionym. Marasmius omphalodes Berk., Hooker's J. Bot. Kew Gard. Misc. 8: 138. 1856. Collybiopsis pseudomphalodes (Dennis) J.S. Kim & Y.W. Lim, comb. nov. MycoBank No: 842063 Gymnopus pseudomphalodes (Dennis) J.L. Mata, in Mata, Hughes, and Petersen, Sydowia 58(2): 289. 2006, as “pseudo-omphalodes”. Marasmiellus pseudomphalodes (Dennis) J.S. Oliveira, in Oliveira, Vargas-Isla, Cabral, Rodrigues and Ishikawa, Mycol. Progr. 18(5): 735. 2019, as “pseudomphalioides”. Basionym. Collybia pseudomphalodes Dennis, Kew Bull. 15(1): 74 (1961). Collybiopsis ramulicola (T.H. Li & S.F. Deng) J.S. Kim & Y.W. Lim, comb. nov. MycoBank No: 842064 Basionym. Gymnopus ramulicola T.H. Li & S.E Deng, in Deng, Li, Jiang and Song, Mycotaxon 131(3): 665. 2016. Taxonomic key to Collybiopsis in Korea 1 Pileus:< 25 sr diata. cea one eee a 2 - Ee rele ees oat hae CL: 18g ORC. Me ay RRS rs. in, REST ie Fy cet Se REY S AN ee 11 2 Lamellae-subdistantao- distant Cl OS 40 i... sors nesslecectdoncyeteticeenebeceetrbercete tes 3 = Lamellae close to crowded (> 30) ..ccccccccccesseccceccccccccccccccesesssessssssssssssssssscesecs 9 3 Basidiomes-on bark, branch,-or woody debris 5.,.:2s:.+1+eeqnssescseseeedereseecegeeeetse “. — Basidiomes: ondti-o wor Soil wen ccoer ener eens araeeuneretha sprmmevresban coevreaarshetl 8 4 PleWe CyStI dia PTE Se MIE 22h. cic ack ack P a anndeeroctst hott aaendl tcioaec rial ce hd waste TontRnetince 5 - Plewieeysticiidabsctit. Metin ce rteee sts rtm eereserda eccrine Zz D Stipe base covered with dense whitish basal tomentum. ....... Co. albicantipes - Stipe base not covered with whitish basal tomentum.............eeeeeeeeeseeeeeeeee 6 6 Pileipellis composed of a coarse Rameales-structure hyphae.......Co. ramealis = Pileipellis composed of a cylindrical, often sub-inflated hyphae, not a RA WTE RICE SIEGAL cieek esa nehdtevs rebated ek Se denbeide deeb pecs tbe acsbendsketainebgh nee Co. fulva Seven new Collybiopsis species 101 7 Pileus distinctly sulcate. Stipe base covered with dense whitish basal tomen- LG 5 ¢ By ett Wer vr eit oe Oe ork ry i tye gE ae re POL dO Oni e RAEN ne OTA. Co. koreana — Pileus slightly sulcate. Stipe base covered with weak whitish basal tomentum besatuc suet eM ath tbat ue Buk tos lees chd ena sbitue Mache alae ahY ebb: Co. nonulla 8 Stipe < 2 cm long. Q value of basidiospores 1.6-2.2 oe Co. dichroa - Stipe > 2 cm long. Q value of basidiospores 2.0-3.1 ....... ce Co. undulata | armel he-croweledntss TOO) niu. Sases.ctsl on Abe biden, tees Co. confluens = Lamellae*closeto. crowded. oP OO) 5 sc.:2 oc cass cea loetovencectincnd adh us ionantletenes 10 10 Basidiates 22) uit LO 8 isda ccteeencessanpeaeavouestaeh cera eaarresecy arsccct Co. menehune Basidiar< 22 wit lone Se. ccgesurehnsetsaten tae cbesnceceeeacaue tegen Co. biformis 11 Lamellaesubdistantstosdisrant-O— 38), <5s. fest es. asedeedetbeoeeSeedesthenocetbastoetboe’ 12 = Team ae: ClOSEcE G8) ric caeccente encondicne.ddvrseventtevedixeebcstepeadisamasinnedbsavesotivese 15 iJ Pléurocystidia present-co 0s... 2 eee ee 13 — Pleurocysticlia absent tre. 89.8 Meus. aS-cbetuser eer Fait tsa Mette naela eal eal 14 1} Oxvalue ot: Dasichosporesty) 2 ile Neve e etl Aaa Co. orientisubnuda — (value*or basidiospores!4) 2.2 cco ieccceneceeostis, eevoosees Co. subumbilicata 14 Pileus convex, hemispherical, plano-convex to flat. Cheilocystidia utriform ANG CAVA geveeusay ees vcaeem tad Minune lesa bon stat abn putes Co. clavicystidiata — Pileus convex to broad-convex. Cheilocystidia narrowly clavate ...........:000 BT RG Fe Ba CAN YAN re tae ina, Seay oy Co. polygramma 15 JEALSI Terex cla (at Fey o}n on) 01 reeiartne, ronan. Rie ee an en, AULA a lear one Co. vellerea _ RleutoeystidiasabSent.. ,...ch.c.ecereivccay eovonesPiee sor eenestiesens totes Co. luxurians Discussion Of the 372 gymnopoid/marasmioid specimens, we confirmed 201 specimens (54%) to belong to Collybiopsis. These results indicate that the species of Collybiopsis can be confused with those of similar genera as well as with other Collybiopsis members when identification is based solely on morphological information. ‘This is because some char- acteristics are overlapped between species (Suppl. material 2: Fig. S2) and the charac- teristics can be different depending on developmental stage or environmental condi- tions. Further, the high misidentification ratio may be caused by the slow rate of adop- tion of the current names. Sequence-based taxonomy has introduced rapid changes in the classification of gymnopoid/marasmioid species (Mata 2002; Mata et al. 2004a; Mata et al. 2004c; Hughes et al. 2010; Oliveira et al. 2019; Petersen and Hughes 2017, 2021). As such, taxonomic confusion has been resolved in taxa that have been well re- searched based on molecular data (Desjardin et al. 1999; Mata 2002; Lee et al. 2019). Nine of the sixteen Collybiopsis species were identified as already known species. Of the nine described species, seven species were identified as the species previously recorded in the Republic of Korea: Collybiopsis biformis, Co. confluens, Co. koreana, Co. luxurians, Co. menehune, Co. polygramma, and Co. ramealis. Two species, Co. dichroa and Co. non- nulla, were reported for the first time in the Republic of Korea. Most of the nine described species formed a monophyletic clade with each corresponding species. However, sequence 102 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) variations by continent were detected in Co. biformis, Co. confluens, Co. dichroa, and Co. nonnulla. Asian samples, including our specimens, were clearly separated from those of Europe, North America, and Africa. These results have also been reported in previous studies on Collybiopsis biformis (Mata 2002; Petersen and Hughes 2014; Razaq et al. 2020) and Co. confluens (Hughes and Petersen 2015). Especially, Co. confluens is known as a rep- resentative example of intra-specific variation between continents. Percent ITS sequence divergence of this species was reported to be 3.25% when comparing the sequences of the North America and Europe (Hughes and Petersen 2015). We confirmed that percent ITS sequence divergence of Asian Co. confluens (our Korean samples and Chinese sequences) were each about 3% when compared to American and European sequences. Similarly, Co. dichroa showed sequence variations that were previously reported in association with intraspecific hybridization and dramatic sequence variations including frequent nucleotide substitutions of Adenine and Guanine (Hughes et al. 2015). The Korean Co. dichroa was closely related to Co. dichroa taxa 2 mentioned in Hughes et al. 2015. Similarly, the intraspecific genetic variation depending on environmental condi- tions or geographical distribution has been reported in many other fungal species (Manian et al. 2001; Kauserud et al. 2007; Seierstad et al. 2013). For the last, Korean Co. nonnulla showed high intra-specific divergence when matching with sequences of Co. nonnulla of America and Cameroon. According to the phylogenetic analysis results, there is a slight sequence variation, but it forms a clade supported by a high bootstrap and morphologi- cally almost coincides with the reference. Therefore, we view this sequence variation as due to different environments by continent and identify the specimens as Co. nonnulla. Nev- ertheless, compared to the fact that it was reported as a new species a long time ago, only seven sequences were deposited in the NCBI, so further study on this species is necessary. Morphologically, the morphological characteristics of the seven described species were also in agreement with the previous descriptions (Suppl. material 2: Fig. $2). However, Co. luxurians and Co. polygramma found in the Republic of Korea showed few differences compared to the Western descriptions in the previous literature (Mata 2003; Noordeloos et al. 1999). In the case of the Co. /uxurians, Korean sequences formed a slightly distinct clade in the phylogenetic tree, along with the Chinese se- quence (ZD 16102301), from European sequences. In this study, direct morphologi- cal comparison studies with European and Chinese samples were difficult and there was no significant morphological difference from the references. For these reasons, we identified Korean specimens as Co. /uxurians, but further studies are needed with more samples from other countries for this species. Seven new species have common characteristics of Colybiopsis such as insititious to subinsititious stipe, ellipsoid to oblong, inamyloid basidiospores, and presence of caulocystidia. However, it is difficult to distinguish them from other Collybiopsis spe- cies based on morphological characteristics alone. Upon molecular phylogenetic analy- ses, each of them clearly formed a distinct clade clearly in the ML phylogenetic tree (Fig. 1). Their morphological features may or may not be distinguished from their phylogenetically close relatives. The morphological differences between new species and morphologically similar or phylogenetically close species are discussed in the re- marks for each species. Seven new Collybiopsis species 103 Two species previously reported in the Republic of Korea, Co. peronata (Cho & Lee, 1979) and Co. subnuda (National list of species of Korea 2020), were not confirmed in this study. Co. peronata and Co. subnuda, which are typical collybioid mushrooms, have been reported in Asia based on their morphological characteristics (Cho and Lee 1979; Kim et al. 1991; Park and Cho 1992; Yoshida and Muramatsu 1998; Tolgor and Yu 2000). Mo- lecular analyses showed that none of the Korean specimens examined in this study could be identified as Co. peronata nor Co. subnuda. Instead, the specimens labelled as Co. pero- nata or Co. subnuda were identified as different species — Gymnopus similis Antonin, Ryoo & Ka and Co. orientisubnuda. Collybiopsis peronata were originally mostly reported from Europe and America and Co. subnuda were originally reported from America (Desjardin et al. 1999; Mata et al. 2006). Furthermore, there have been no recent sequence uploads to GenBank or reports of Co. peronata and Co. subnuda from Asia, making it difficult to confirm whether they exist in the Republic of Korea. Although Co. orientisubnuda is close- ly related to Co. peronata and Co. subnuda, there are clear differences in the ITS regions of these three species (Suppl. material 3: Fig. $3). Morphologically, Co. orientisubnuda is highly similar to Co. subnuda and considerably different from Co. peronata. The detailed comparisons of the morphological features are provided in the remarks for each species. In conclusion, we identified 16 Collybiopsis species in the Republic of Korea through morphological and molecular analyses and we update the Korean inventory of Collybiopsis. Our study showed that the identification of Collybiopsis species requires both morphological and molecular analyses. Further, this study has the following sig- nificance as in the previous study conducted by Petersen and Hughes (2021): addi- tional combinations of Marasmiellus species under Collybiopsis, detailed morphological characterization of Collybiopsis species in the Republic of Korea along with photo- graphs and drawings, and specific approaches to species differentiation and identifica- tion through morphological and molecular analyses. 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Journal of Environmental Radioactivity 41(2): 183-205. https://doi.org/10.1016/S0265- 93 1X(97)00098-2 Supplementary material | Figure S1 Authors: Ji Seon Kim, Yoonhee Cho, Ki Hyeong Park, Ji Hyun Park, Minkyeong Kim, Chang Sun Kim, Young Woon Lim Data type: Jpg file. Explanation note: Pileipellis elements of seven new species. A Collybiopsis albicantipes B Co. clavicystidiata C Co. fulva D Co. orientisubnuda E Collybiopsis subumbilicata F Co. undulata G Co. vellerea. Scale bars: 10 wm. Copyright notice: 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. Link: https://doi.org/10.3897/mycokeys.88.79266.suppl1 Supplementary material 2 Figure S2 Authors: Ji Seon Kim, Yoonhee Cho, Ki Hyeong Park, Ji Hyun Park, Minkyeong Kim, Chang Sun Kim, Young Woon Lim Data type: Jpg file. Explanation note: Comparison of the morphological characters of 16 Korean Collybi- opsis species. Copyright notice: 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. Link: https://doi.org/10.3897/mycokeys.88.79266.suppl2 108 Ji Seon Kim et al. / MycoKeys 88: 79-108 (2022) Supplementary material 3 Figure S3 Authors: Ji Seon Kim, Yoonhee Cho, Ki Hyeong Park, Ji Hyun Park, Minkyeong Kim, Chang Sun Kim, Young Woon Lim Data type: Jpg file. Explanation note: Sequence difference between the three species in the ITS region. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODDbL) 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. Link: https://doi.org/10.3897/mycokeys.88.79266.suppl3