Research Article |
Corresponding author: Young Woon Lim ( ywlim@snu.ac.kr ) Academic editor: Bao-Kai Cui
© 2022 Ji Seon Kim, Yoonhee Cho, Ki Hyeong Park, Ji Hyun Park, Minkyeong Kim, Chang Sun Kim, Young Woon Lim.
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:
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
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Collybiopsis is a genus of the gymnopoid/marasmioid complex of the family Omphalotaceae. The classification 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 combining 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 approach is needed for accurate identification. The Collybiopsis 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.).
Collybia, gymnopoid, Gymnopus, ITS, Marasmiellus, marasmioid, nrLSU
Collybiopsis
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. (
Collybiopsis is morphologically similar and phylogenetically close to Gymnopus (
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. Petersen, as its previous name Collybia confluens Fr. (
A total of 372 specimens deposited in three Korean fungal herbarium – Seoul National University Fungus Collection (
Genomic DNA was extracted from each specimen using a modified CTAB DNA extraction protocol (
All sequences generated in this study were proofread using MEGA version 7 (
Information about the Collybiopsis specimens and published Collybiopsis sequences used in phylogenetic analysis. Species with an asterisk are those proposed as new species. Sequences newly produced in this study are presented in bold.
Organisms | Specimen | Collection Date | Location | GenBank Accession Number | |
---|---|---|---|---|---|
ITS | nrLSU | ||||
Collybiopsis albicantipes * | SFC20170725-35 | 25.7.2017 | Yeosu-si, Jeollanam-do, the Republic of Korea | OL467272 | OL462811 |
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 Korea | OL467229 | OL462789 | |
SFC20180831-16 | 31.8.2018 | Jindo-gun, Jeollanam-do, the Republic of Korea | OL467230 | OL462790 | |
Co. brunneigracilis | AWW01 | Java/Bali | AY263434 | AY639412 | |
Co. californica | TFB5787 | Canada: British Columbia | MN413338 | ||
Co. clavicystidiata * | SFC20180705-26 | 5.7.2018 | Haenam-gun, Jeollanam-do, the Republic of Korea | OL467250 | OL462816 |
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 | OL462819 | |
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 | KJ189578 | ||
110116MFBPL0425 | China | MW554401 | |||
HMAS 290186 | China | MK966541 | |||
Co. confluens ssp. americana | TFB14409 | Canada: New Brunswick | KP710278 | KJ189585 | |
TFB14075 | USA: North Carolina | KP710281 | KJ189581 | ||
Co. dichroa | KA14-0969 | 19.8.2014 | Hwasun-gun, Jeollanam-do, the Republic of Korea | OL467254 | OL462799 |
KA18-0389 | 10.7.2018 | Cheongdo-gun, Gyeongsangbuk-do, the Republic of Korea | OL467255 | OL546541 | |
SFC20180712-16 | 12.7.2018 | Gwangju, Gyeonggi-do, the Republic of Korea | OL467256 | OL462800 | |
TFB9623 | USA: North Carolina | MW396865 | MW396865 | ||
TENN60014c2 | USA: Tennessee, GSMNP | JF313671 | |||
TFB7920 | USA | DQ450007 | |||
TENN61624c1a | USA: Tennessee, GSMNP | JF313678 | |||
TFB2028 | USA | DQ450008 | |||
TENN61624c9 | USA: Tennessee, GSMNP | 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 | KP710304 | ||
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 Republic of Korea | OL467257 | OL462793 |
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 | |||
Co. indoctus | AWW04 | Unknown | AY263439 | ||
Co. istanbulensis | KATO Fungi 3596 | Turkey | KX184795 | KX184796 | |
BRNM 781163 | Turkey | KY250435 | |||
Co. juniperina | TFB9889 | USA: Louisiana | AY256708 | KY019637 | |
TFB10782 | Argentina: Missiones | KY026661 | KY026661 | ||
Co. koreana | SFC20120821-84 | 21.8.2012 | Boryeong-si, Chungcheongnam-do, the Republic of Korea | OL467269 | OL546545 |
SFC20130711-05 | 11.7.2013 | Pyeongchang-gun, Gangwon-do, the Republic of Korea | OL467270 | OL462801 | |
SFC20150721-10 | 21.7.2015 | Inje-gun, Gangwon-do, the Republic of Korea | OL467271 | OL462802 | |
BRNM 714972 | Korea | GU319113 | GU319117 | ||
BRNM 718782 | Korea | GU319114 | GU319118 | ||
Co. luxurians | NIBRFG0000502888 | 4.9.2018 | Ongjin-gun, Incheon, the Republic of Korea | OL467248 | OL462803 |
SFC20190731-18 | 31.7.2019 | Taebaek-si, Gangwon-do, the Republic of Korea | OL467249 | OL462804 | |
TFB10350 | USA: North Carolina | AY256709 | AY256709 | ||
ZD16102301 | China | MN523270 | |||
TFB9121 | USA: Louisiana | KY026649 | KY026649 | ||
Co. melanopus | AWW54 | Java/Bali | NR_137539 | NG_060624 | |
CUH AM093 | India | KM896875 | KP100305 | ||
Co. menehune | SFC20150811-29 | 11.8.2015 | Guri-si, Gyeonggi-do, the Republic of Korea | OL467235 | OL462805 |
SFC20180905-33 | 5.9.2018 | Anyang City, Gyeonggi Province, the Republic of Korea | OL467236 | OL462806 | |
SFSU: DED5866 | Hawaii | AY263426 | |||
CUH:AM074 | India | KJ778753 | KP100302 | ||
SFSU-AWW15 | Java/Bali | AY263443 | AY639424 | ||
Co. mesoamericana | TFB11005 | Costa Rica | DQ450035 | KY019632 | |
REH7379 | Costa Rica | AF505768 | |||
Co. micromphaleoides | TENN 68165 | ||||
TFB14282 | |||||
Co. minor | TFB11930 | USA: Tennessee, GSMNP | MN413334 | MW396880 | |
TFB5434 | USA: South Carolina | MW396872 | MW396872 | ||
Co. neotropica | TFB10416 | Costa Rica | AF505769 | ||
Co. nonnulla | KA13-0254 | 20.6.2013 | Geochang-gun, Gyeongsangnam-do, the Republic of Korea | OL467242 | OL462820 |
KA13-0741 | 21.8.2013 | Geochang-gun, Gyeongsangnam-do, the Republic of Korea | OL467243 | OL462807 | |
KA15-0129 | 14.7.2015 | Gangneung-si, Gangwon-do, the Republic of Korea | OL467244 | OL462808 | |
TFB14492 | USA: Mississippi | KY026699 | KY026699 | ||
TFB14278 | USA: Mississippi | KY026701 | KY026701 | ||
Co. nonnulla var. attenuatus | AWW05 | Java/Bali | AY263445 | AY639426 | |
AWW55 | Java/Bali | AY263446 | |||
RAK369.2 | Cameroon | MN930621 | |||
RAK372.2 | Cameroon | MN930622 | |||
Co. obscuroides | GB-0150514 | Norway: Svalbard | KX958399 | KX958399 | |
Co. omphalodes | FB11021 | Costa Rica | AF505761 | ||
TFB 10427 | Costa Rica | DQ450011 | |||
TFB 10022 | Costa Rica | AY256700 | |||
Co. orientisubnuda * | NIBRFG0000500990 | 19.7.2016 | Ulleung-gun, Gyeongsangbuk-do, the Republic of Korea | OL467262 | OL546546 |
SFC20170823-39 | 23.8.2017 | Hapcheon-gun, Gyeongsangnam-do, the Republic of Korea | OL467263 | OL546547 | |
SFC20180830-29 | 30.8.2018 | Hapcheon-gun, Gyeongsangnam-do, the Republic of Korea | OL467264 | OL462796 | |
(as Gymnopus subnudus) | KUC20150911-19 | Korea | KX513748 | ||
Co. parvula | TFB10419 | Costa Rica | DQ450060 | ||
TFB10422 | Costa Rica | AF505774 | |||
Co. peronata | TFB13743 | Belgium | KY026677 | KY026677 | |
LE-Bin1364 | Russia | KY026755 | KY026755 | ||
CBS 223.37 | unknown | MH855896 | MH867405 | ||
Co. polygramma | SFC20170807-35 | 7.8.2017 | Hapcheon-gun, Gyeongsangnam-do, the Republic of Korea | OL467245 | OL546542 |
SFC20180905-63 | 5.9.2018 | Gwanak-gu, Seoul, the Republic of Korea | OL467246 | OL546544 | |
SFC20210629-01 | 29.6.2021 | Gwanak-gu, Seoul, the Republic of Korea | OL467247 | OL546543 | |
PR2542TN | Puerto Rico | DQ450028 | |||
CUH:AM082 | India | KJ778752 | KP100303 | ||
URM 90015 | Brazil: Amapa | KY074640 | KY088275 | ||
MHHNU 30912 | China | MK214392 | |||
TFB9628 | Puerto Rico | DQ450028 | |||
SFC20120821-64 | Korea | KJ609162 | |||
HFJAU 0425 | China: Jiangxi | MN258643 | |||
(as Gymnopus iocephalus) | KUC20140804-02 | Korea | KX513745 | ||
Co. pseudoluxurians | TFB14290 | USA: Mississippi | NR_137863 | ||
Co. pseudomphalodes | REH7348 | Costa Rica | AF505762 | ||
PR24TN | Puerto Rico | AY842957 | |||
Co. quercophila | TFB14570 | Slovakia | KY026728 | KY026728 | |
TFB14615 | USA: California | KY026736 | KY026736 | ||
Co. ramealis | NIBRFG0000508888 | 29.7.2020 | Jeongseon-gun, Gangwon-do, the Republic of Korea | OL467260 | OL546549 |
TFB13769 | Belgium: Couvin | MN413345 | MN413345 | ||
TFB13770 | Belgium: Couvin | MN413346 | MW396882 | ||
DED4425 | USA: North Carolina | DQ450031 | AF042650 | ||
TFB14555 | Slovakia | MW405779 | MW396884 | ||
BR 72_41 | Belgium | MW396875 | MW396875 | ||
Co. ramulicola | GDGM 43884 | China | KU057798 | ||
GDGM 44256 | China | KU321529 | |||
GDGM 50060 | China | KU321530 | |||
Co. readiae | TFB7571 | New Zealand | DQ450034 | ||
PDD:95844 | New Zealand | HQ533036 | |||
Co. stenophylla | TFB13998 | USA: Tennessee, | MN413331 | MW396886 | |
TFB4798 | USA: Georgia | MN413330 | MW396887 | ||
Co. subcyathiformis | TFB9629 | ||||
URM 90023 | Brazil: Para | KY404982 | KY404982 | ||
URM 90022 | Brazil: Para | KY404983 | KY404983 | ||
Co. subnuda | TFB12577 | USA: Tennessee, GSMNP | KY026667 | FJ750262 | |
WRW 08-462 | USA: West Virginia | KY026765 | KY026765 | ||
TFB14043 | USA: North Carolina | MW396876 | MW396876 | ||
Co. subpruinosus | BRNM781138 | Portugal: Madeira | MK646034 | ||
TFB11063 | USA | DQ450025 | |||
Co. subumbilicata * | SFC20120802-03 | 2.8.2012 | Goseong-gun, Gangwon-do, the Republic of Korea | OL467231 | OL462786 |
SFC20140701-03 | 1.7.2014 | Inje-gun, Gangwon-do, the Republic of Korea | OL467232 | OL462787 | |
SFC20150902-50 | 2.9.2015 | Ulleung-gun, Gyeongsangbuk-do, the Republic of Korea | OL467234 | OL546540 | |
SFC20170822-14 | 22.8.2017 | Ulleung-gun, Gyeongsangbuk-do, the Republic of Korea | OL467233 | OL462788 | |
Co. trogioides | AWW51 | Indonesia | AY263428 | AY639431 | |
Co. undulata * | SFC20120821-04 | 21.8.2012 | Boryeong-si, Chungcheongnam-do, the Republic of Korea | OL467239 | OL462813 |
SFC20130808-08 | 8.8.2013 | Sangju-si, Gyeongsangbuk-do, the Republic of Korea | OL467240 | OL462814 | |
SFC20150813-04 | 13.8.2015 | Goyang-si, Gyeonggi-do, the Republic of Korea | OL467241 | OL462815 | |
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 | OL467265 | OL462791 |
SFC20120708-02 | 8.7.2012 | Seosan-si, Chungcheongnam-do, the Republic of Korea | OL467266 | OL462809 | |
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. ramealis | SFC20180829-20 | 29.8.2018 | Shinan-gun, Jeollanam-do, the Republic of Korea | OL467261 | OL546548 |
Rhodocollybia butyracea | TFB 14382 | Canada: New Brunswick | KY026716 | KY026716 | |
Rhodocollybia dotae | REH7007 | Costa Rica | AF505758 | ||
Rhodocollybia maculata | TFB 13989 | USA: Mississippi | KY026688 | KY026688 |
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.
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 observations, tissues of dried specimens were rehydrated in 5% (w/v) KOH and mounted in Congo red solution (
Through ITS sequence analysis of 372 gymnopoid/marasmioid specimens, 201 specimens were confirmed to belong to Collybiopsis. The remaining 160 specimens were 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
Identification information of Korean Collybiopsis specimens confirmed in the study. Scientific names in bold indicate new species.
Species | Specimen Number | ||||
---|---|---|---|---|---|
Co. albicantipes | SFC20170725-35 | SFC20180704-86 | |||
Co. biformis | NIBRFG0000502789 | KA14-0259 | KA14-0526 | KA14-0917 | KA14-0924 |
KA16-0307 | KA16-0371 | KA16-0526 | KA18-0657 | KA18-0673 | |
SFC20140724-41 | SFC20160719-42 | SFC20180704-36 | SFC20180706-05 | SFC20180831-13 | |
SFC20180831-16 | |||||
Co. clavicystidiata | KA14-0667 | KA14-0724 | KA15-0211 | KA17-0287 | KA17-0369 |
KA18-0282 | KA18-0353 | SFC20180705-84 | SFC20180705-92 | SFC20180706-04 | |
SFC20180713-09 | |||||
Co. confluens | NIBRFG0000508913 | NIBRFG0000508991 | KA16-0696 | KA18-0338 | SFC20150626-26 |
SFC20190731-06 | SFC20190731-32 | SFC20190731-34 | SFC20190731-48 | ||
Co. dichroa | KA14-0969 | KA17-0344 | SFC20180706-60 | SFC20180712-16 | |
Co. fulva | KA13-0215 | KA13-0216 | KA13-0333 | KA13-0357 | KA14-0168 |
KA14-0386 | KA14-0666 | KA14-0691 | KA15-0210 | KA16-0425 | |
KA16-0428 | KA17-0388 | KA17-0596 | KA18-0233 | KA18-0241 | |
Co. koreana | SFC20120821-84 | SFC20150702-25 | SFC20170713-06 | SFC20180704-17 | |
Co. luxurians | NIBRFG0000502888 | SFC20190731-18 | SFC20190731-08 | SFC20190730-36 | SFC20180907-105 |
SFC20180905-86 | SFC20180905-43 | KA18-0321 | KA14-0579 | ||
Co. menehune | NIBRFG0000502876 | KA13-0887 | KA14-0494 | KA14-0510 | SFC20150811-29 |
SFC20160719-15 | SFC20180905-33 | ||||
Co. nonnulla | KA13-0254 | KA13-0741 | KA15-0129 | ||
Co. orientisubnuda | SFC20170823-39 | SFC20170708-14 | SFC20150902-01 | SFC20150820-59 | SFC20150820-01 |
SFC20150811-48 | SFC20150701-100 | QM20200911-57 | QM20200911-52 | KA17-0787 | |
KA17-0600 | KA16-1154 | KA16-0925 | KA16-0902 | KA16-0780 | |
KA16-0724 | KA15-0179 | KA14-0985 | KA13-1225 | F20200730-24 | |
F20200701-11 | F20200630-30 | F20180904KCM21 | F20160719-12 | ||
Co. polygramma | KA13-0506 | KA13-0956 | KA13-1101 | KA13-1333 | KA14-0904 |
KA14-1089 | KA14-1092 | KA18-0115 | KA18-0724 | QM20200721-07 | |
NIBRFG0000508098 | NIBRFG0000508059 | NIBRFG0000508089 | SFC20170712-08 | SFC20170807-35 | |
SFC20170822-66 | SFC20180905-49 | SFC20180905-63 | |||
Co. ramealis | SFC20130711-05 | ||||
Co. subumbilicata | KA13-1214 | KA15-0173 | KA15-0185 | KA15-0787 | SFC20120802-03 |
SFC20140701-03 | SFC20150902-50 | SFC20170822-14 | SFC20210623-03 | ||
Co. undulata | KA17-0335 | KA18-0651 | KA18-0651 | SFC20120821-04 | SFC20130808-08 |
SFC20150715-24 | SFC20150813-04 | ||||
Co. vellerea | KA14-0132 | KA14-0163 | KA14-0196 | KA14-0245 | KA14-0397 |
KA14-0412 | KA14-0446 | KA14-0447 | KA14-0474 | KA14-0725 | |
KA14-0734 | KA14-0735 | KA14-0774 | KA14-0787 | KA14-1005 | |
KA14-1061 | KA14-1147 | KA14-1349 | KA14-1426 | KA14-1475 | |
KA14-1555 | KA14-1558 | KA15-0213 | KA15-0215 | KA15-0473 | |
KA15-0485 | KA15-0502 | KA15-0527 | KA15-0568 | KA16-0191 | |
KA16-0252 | KA16-0485 | KA16-0783 | KA16-0982 | KA16-0985 | |
KA16-0986 | KA16-0992 | KA17-0368 | KA17-0586 | KA17-0742 | |
KA17-1074 | KA18-0089 | KA18-0139 | KA18-0151 | KA18-0152 | |
KA18-0348 | KA18-0795 | KA18-0836 | KA18-0987 | KA18-1027 | |
KA19-0125 | SFC20120708-02 | SFC20120820-02 | SFC20140821-29 | SFC20150630-38 | |
SFC20150714-01 | SFC20170705-06 | SFC20180705-90 | SFC20180829-30 | SFC20180901-01 | |
Collybiopsis cf. ramealis | F20200729-14 |
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 Antonín, Ryoo & H.D. Pictures of basidiomata are shown in Fig.
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 Co. nonnulla (KA13–0254) H Co. polygramma (SFC20170712–08) I Co. ramealis (SFC20180829–20). Scale bar: 1 cm (A–I).
Basidiomata and microscopic characters of the four new Collybiopsis species A, B Co. albicantipes (SFC20170725–35) C, D Co. clavicystidiata (SFC20180705–84) E, F Co. fulva (KA15–0210) G, H Co. orientisubnuda (NIBRFG0000502862). Scale bars: 1cm (A, C, E, G); 20 µm (B, D, F, H). Abbreviations: s basidiospores; b basidia; ch cheilocystidia; p pleurocystidia; ca caulocystidia.
Basidiomata and microscopic characters of the three new Collybiopsis species A, B Co. subumbilicata (SFC20120802–03) C, D Co. undulata (SFC20150813–04) E, F Co. vellerea (SFC20140821–29). Scale bars: 1cm (A, C, E); 20 µm (B, D, F). Abbreviations: s basidiospores; b basidia; ch cheilocystidia; p pleurocystidia; ca caulocystidia.
Five species (G. omphalodes Halling & J.L. Mata, G. pseudomphalodes J.L. Mata, G. ramulicola T.H. Li & S.F. Deng, Ma. istanbulensis E. Sesli, Antonín and E.Aytaç, and Ma. koreanus), previously placed in Gymnopus section Vestipedes, were confirmed to belong 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. istanbulensis comb. nov., and Co. koreana comb. nov. respectively.
Epithet “albicantipes” refers to having a whitish base of the stipe.
This species notably has hemispherical to convex, 4–23 mm pileus, distant lamellae, central to eccentric, tomentose, 5–15 × 0.5‒1.5 mm stipe with a white base; ellipsoid to ovoid, 5.8–7.4 × 2.8–4 μm basidiospores, clavate (often constricted), 25.5–34.8 × 4.8–6.7 μm basidia, broadly clavate, irregular, sometimes lobed, 26–49 × 5.4–10.6 μm cheilocystidia, and a habit of fruiting on branches.
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, L = 10–16, l = 3–7, adnate, whitish to yellowish white (3A2). Stipe: 5–15 × 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 tomentum. Basidiospores: 5.8–7.4 × 2.8–4 μm (average 5.5 × 3.2 μm), Q = 1.6–2.1 (mean = 1.97), ellipsoid to ovoid, amygdaliform, smooth, hyaline, non-dextrinoid, with drops. Basidia: (23) 25.5–34.8 × 4.8–6.7 (7) μm, 4-spored, clavate, often constricted. Cheilocystidia: 26–49 × 5.4–10.6 (14) μm, broadly clavate, irregular, sometimes lobed. Pleurocystidia: 25.8–56.4 (62) × 6.2–12.5 μm, clavate, subulate, sometimes lobed. Trama hyphae: cylindrical, often sub-inflated, smooth, non-dextrinoid 1.7–9 (12) μm wide. Pileipellis: a cutis made up of cylindrical, often sub-inflated, with weak annular ornamentation, 2.0‒7.5 μm wide hyphae; terminal elements adpressed, cylindrical, clavate, sometimes constricted or curved, 2.0‒5 μm wide. Stipitipellis: a cutis of cylindrical, smooth, 2.7‒9.7 (11) μm wide hyphae. Caulocystidia: 21.7–90 × 3.9–11.7 μm, cylindrical, flexuose, sometimes curved. Clamp connections: present in all tissues.
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.
Scattered to gregarious on the branch in mixed forest dominated by Camellia japonica Linne, in summer.
The Republic of Korea.
Collybiopsis albicantipes is similar to Co. ramulicola and Co. koreana when comparing macro-morphological characteristics. Collybiopsis ramulicola is distinguishable from Co. albicantipes by a reddish pileus, fewer and buff lamellulae (1–4), a shorter and thinner stipe (12–23 × 2–3 mm), shorter and slightly elongated basidiospores (6.6–8.4 × 3.5–4.5 μm), shorter basidia (23–27 × 3.8–5.5 μm), and shorter cheilocystidia (23–27 × 3–6 μm) (
Epithet “clavicystidiata” indicates that the new species has clavate cheilocystidia.
The prominent features of this species include a greyish orange to brownish, 6–45 mm pileus, whitish lamellae, a subinstitious, tomentose, whitish, 15–26 × 1.2‒1.6 mm stipe, oblong to subcylindrical, 6.7‒9.4 × 3.1‒4.6 μm basidiospores, utriform, clavate, 20.1–37.5 × 6.8–12.2 μm cheilocystidia, and cylindrical, flexuose, irregular, 17–50 × 3.5–7 μm caulocystidia.
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, l = 1–7, adnexed, white. Stipe: 15–26 × 1.2‒1.6 mm, cylindrical, tomentose, subinsititious, whitish to reddish grey (9B2). Basidiospores: 6.7‒9.4 × 3.1‒4.6 μm, average 8.13 × 3.62 μm, Q = 2–2.4 (mean = 2.26), oblong to cylindrical, smooth, hyaline, non-dextrinoid, with drops. Basidia: 18.3–30 × 4.1–8.8 μm, 4-spored, narrowly clavate, narrowly utriform, often curved. Cheilocystidia: 20.1–37.5 × 6.8–12.2 μm, utriform, clavate, sometimes with mucronate apex. Pleurocystidia: absent. Trama hyphae: cylindrical, often subinflated, smooth, branched, non-dextrinoid, 2‒12 μm wide. Pileipellis: transition between cutis and trichoderm, composed of cylindrical, with heavy annular ornamentation, 4‒12 μm wide hyphae; terminal elements adpressed to suberect, cylindrical, clavate, often incrusted (often incrusted), thin-walled, 3‒6 μm wide. Stipitipellis: a cutis of cylindrical, smooth, 2‒7 μm wide hyphae. Caulocystidia: 17–50 × 3.5–7 μm, cylindrical, flexuose, irregular or curved. Clamp connections: present in all tissues.
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.
Solitary to scattered on dead wood debris of conifers, in summer.
The Republic of Korea
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 × 2.5–3 μm), and thinner hyphae in the pileipellis (5–8 μm 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 × 3.5–4.2 μm, Q = 2.2), and longer caulocystidia (16–67 × 3–5 μm) (
This species has a pale orange to brownish-colored, 4–20 mm pileus, an orange white colored to light brownish colored, 7–30 × 0.7–1 mm stipe with pubescence, spheropedunculate, pleurocystidia, oblong to subcylindrical, 6.8–9.2 × 3.1–4.9 μm basidiospore, lobed, clavate with rostrate apex, 24.8–38.4 × 6.5–11.8 μm cheilocystidia.
Epithet “fulva” referring to fox-colored pileus.
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, l = 1–5, sinuate, broad, whitish to yellowish white (4A2) to brownish orange (6C4 to 7C4). Stipe: 7–30 × 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 × 3.1–4.9 μm (average 7.47 × 3.69 μm), Q = 2.05, oblong to cylindrical, smooth, colorless, non-dextrinoid, with drops. Basidia: 20.4–29.4 × 4.7–7.8 μm, 4-spored, narrowly clavate, sometimes constricted or curved. Cheilocystidia: (20.5) 24.8–38.4 × 6.5–11.8 μm, lobed, clavate, sometimes with rostrate apex. Pleurocystidia: 31.5–46.9 × 12–20.6 μm, spheropedunculate, obovoid, sometimes with mucronate apex. Trama hyphae: cylindrical to subinflated, irregular, thin-walled, smooth, branched, non-dextrinoid, 2.0‒15 μm wide. Pileipellis: a cutis of cylindrical, thin-walled, 4–15 μm wide hyphae; terminal elements adpressed to suberect, narrowly clavate, thin-walled, with heavy annular ornamentation, 3–8 μm wide. Stipitipellis: a cutis of cylindrical, thin-walled, smooth, 5‒15 μm wide hyphae. Caulocystidia: 45.6–108.3 (131) × 6.8–14.8 μm, cylindrical, irregular, curved. Clamp connections: present in all tissues.
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.
Scattered or gregarious on the bark of deciduous trees or on the rotting branch of both broadleaf trees and conifers, in summer.
The Republic of Korea.
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 larger basidiospores (7.5–9.5 × 3.5–4.2 μm) (
Epithet “orientisubnuda” meaning the new species has originated from the East and is morphologically similar to Co. subnuda.
It features a brownish, 15‒50 mm pileus, orangish cream-colored lamellae, greyish to brownish orange, tomentose, 20–80 × 2.5‒6 mm stipe, subcylindrical to fusoid, 6.7–8.6 × 1.8–3.2 μm basidiospores, and cylindrical, flexuose, sometimes irregular or curved, 26.3–52 (63) × 3.5–6.5 μm caulocystidia. This species is morphologically similar to Co. subnuda.
Pileus: 15‒50 mm, convex to plano-convex, sometimes subumbonate; Surface smooth, brownish orange (6C5 to 7C4), becoming paler to the margin (5A2). Lamellae: distant, L = 16–28, l = 3–7, adnexed, pale yellow (4A3) to orange white (5A2). Stipe: 20–80 (100) × 2.5‒6 mm, central to eccentric, cylindrical, tomentose, often twisted, greyish orange (6B4) to brownish orange(7C4), becoming paler and thinner to the base. Basidiospores: 6.7–8.6 × 1.8–3.2 μm (average 7.5 × 2.5 μm), Q = 2.5–3.2 (mean = 2.92), cylindrical to fusoid, smooth, hyaline, non-dextrinoid, with drops. Basidia: (17) 19.8–28.7 (29) × 3.7–7.3 μm, 4-spored, narrowly clavate, often constricted. Cheilocystidia: variable in shape and size, 21–33.3 × 4.7–8.2 μm, lobed, clavate, slightly sphaeropendunculate, sometimes constricted or with rostrate apex. Pleurocystidia: 24.7–52.3 × 5.1–9.1 μm, narrowly utriform, clavate, sometimes clavate with rostrate apex. Trama hyphae: cylindrical, often subinflated, smooth, branched, non-dextrinoid, 2.0–7.0 μm wide. Pileipellis: a cutis made up of cylindrical, 2‒8 μm wide hyphae; terminal elements adpressed, cylindrical, often subinflated, with weak annular ornamentation, 3‒6 μm wide. Stipitipellis: a cutis of cylindrical, smooth, 2.5–7 μm wide hyphae. Caulocystidia: 26.3–52 (63) × 3.5–6.5 μm, cylindrical, flexuose, sometimes irregular or curved. Clamp connections: present in all tissues.
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: Ulleung-gun, 37°31'30"N, 130°52'21"E, alt. 718 m, 2 September 2015, Jae Young Park, SFC20150902-01.
Scattered to gregarious on the ground covered with dead and decaying leaves of broadleaf forest, from summer to autumn.
The Republic of Korea.
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 μm), and longer cheilocystidia (25–90 × 5–10 μm) (
Epithet “subumbilicata” referring to having a small depressed center in pileus.
The distinctive features include a brownish, 10–35 mm pileus, white colored lamellae, a brownish, 25–60 × 1‒3 mm stipe covered with pubescence, ellipsoid to oblong basidiospores, narrowly clavate and cylindrical, 17–24.3 × 3.5–5.1 μm basidia, and cylindrical, flexuose, sometimes curved, 12.6–38.2 × 2.4–6.6 μm caulocystidia.
Pileus: 10–35 mm, plano-convex to plano-concave, subumbilicate, becoming undulate and uplifted in age; Surface smooth, greyish orange (5B3) to brown (6E5). Lamellae: subdistant, L = 22–38, l = 3–7, free to adnexed, white. Stipe: 25–60 × 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 × 2.5‒3.6 μm (average 6.47 × 3.0 μm), Q = 1.8–2.2 (mean = 2), oblong to fusiform, smooth, hyaline, non-dextrinoid, with drops. Basidia: (15.6) 17–24.3 (27.6) × 3.5–5.1 (5.9) μm, 4-spored, narrowly clavate, cylindrical. Cheilocystidia: 17.6–38.4 × 5–7.8 μm, various in shape, lobed. Pleurocystidia: 20.3‒30.7 × 6.8‒9.5 μm, clavate, fusiform, slightly sphaeropedunculate. Trama hyphae: cylindrical, subinflated, branched, smooth, non-dextrinoid, 1.5‒8 μm wide. Pileipellis: a cutis made up of cylindrical, often incrusted, with heavy annular ornamentation, 5.0‒15 μm wide hyphae; terminal elements adpressed to suberect, fusoid, clavate, 6.0‒16 μm wide. Stipitipellis: a cutis of cylindrical, smooth, thin-walled, 2.0‒6.0 μm wide hyphae. Caulocystidia: 12.6–38.2 × 2.4–6.6 μm, cylindrical, flexuose, sometimes irregular or curved. Clamp connections: present in all tissues.
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.
Scattered to gregarious on the ground covered with dead leaves in temperate mixed forests, from summer to autumn.
The Republic of Korea.
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 basidiospores (6.5‒10.5 × 3.5‒4.5 μm) and basidia (25‒34 × 6.5‒7.5 μm) (
Epithet “undulata” referring to having an undulate margin of pileus.
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 × 0.8‒2 mm stipe that becomes lighter to the apex, subcylindrical, broadly clavate or irregular, sometimes lobed, 16.7–28 × 4.8–8 μm cheilocystidia, and 27–60 × 3.5–6 μm sized caulocysitida which has a morphology similar to cheilocystidia and sometimes grows in bundles.
Pileus: 10‒23 mm, convex to concave, margin becoming undulate with age; Surface smooth, hygrophanous, brown (7D2 to 7E6) in the center, becoming paler to the margin (5A2–5B3 to 7B2). Lamellae: subdistant, L = 15–30, l = 3–9, adnexed, cream. Stipe: 35–55 × 0.8‒2 mm, cylindrical, tomentose, dark brown (7F5 to 8F8), gradually becoming paler to apex (7B2 to 7C2). Basidiospores: 5.6–9.5 × 2–3.4 μm (average 7.3 × 2.8 μm), Q = 2–3.1 (mean = 2.58), cylindrical, smooth, hyaline, non-dextrinoid, with drops. Basidia: 15–22.3 × 3.6–6.8 μm, 4-spored, cylindrical, narrowly clavate to utriform, often curved. Cheilocystidia: 16.7–28 × 4.8–8 μm, subcylindrical, broadly clavate or irregular, sometimes lobed. Pleurocystidia: absent. Trama hyphae: cylindrical, sometimes subinflated, smooth, branched, non-dextrinoid, 2–8 μm wide. Pileipellis: a cutis made up of cylindrical, often incrusted, slightly brownish, with heavy annular ornamentation, 2.4–7 μm wide hyphae; terminal elements adpressed to suberect, cylindrical to clavate, 3–6 μm wide. Stipitipellis: a cutis of cylindrical, smooth, 2.0‒3.5 μm wide hyphae. Caulocystidia: 27–60 × 3.5–6 μm, irregularly cylindrical, narrowly utriform, seldom apically lobed, sometimes gathered in a bunch. Clamp connections: present in all tissues.
The Republic of Korea, Gyeonggi-do: Goyang-si, Deogyang-gu, Seooreung, 37°37'26"N, 126°54'4"E, alt. 35 m, 13 August 2015, Jae Young Park, 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.
Scattered to gregarious on leaf litter in mixed forest dominated with broadleaf trees, in summer.
The Republic of Korea.
Collybiopsis undulata is morphologically similar to Co. subpruinosa (Murrill) R.H. Petersen. Collybiopsis subpruinosa has differences in having small central papilla on pileus, fewer lamellulae (3–4 series), vivid colored lamellae, thicker basidiospores (4.5–5.2 μm wide), larger basidia (30–36 × 7.5–8.5 μm) and cheilocystidia (25–80 × 5–16 μm), thick-walled trama hyphae (0.5–1 μm), caulocystidia with a wider size range, and a habit of growing solitary on rotten twigs or logs (
Epithet “vellerea” refers to having a velvety stipe.
It has a dull, greyish orange, 18‒45 mm pileus with darker center, a tomentose (like velvet), insititious, orangish, 15–55 × 3‒5 mm stipe, sphaeropendunculate, subovoid, 23.4–49 × 7.5–13.4 μm pleurocystidia, oblong to subcylindrical basidiospores, narrowly clavate with rostrate apex, sometimes lobed, 7.7–49.7 × 3.8–14.6 μm cheilocystidia.
Pileus: 18‒45 mm, hemispherical, appendiculate to convex, subumbonate 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 (5A1 to 5B2). Lamellae: crowded to close, L = 38‒52, l = 3‒7, furcate, white. Stipe: 15–55 × 3‒5 mm, cylindrical, finely tomentose, insititious, pale orange (5A3) to reddish grey (7B2), becoming darker to the base (6A2 to 7C2). Basidiospores: 5.2‒7 × 2.5‒3.8 μm (average 6.17 × 3.06 μm), Q = 1.8–2.4 (mean = 2.03), oblong to subcylindrical, smooth, hyaline, non-dextrinoid, with drops. Basidia: 16.2–24.8 × 3.3–5.3 μm, 4-spored, (narrowly) clavate, often curved or constricted. Cheilocystidia: 7.7–49.7 × 3.8–14.6 μm, narrowly clavate with rostrate apex, sometimes lobed. Pleurocystidia: 23.4–49 × 7.5–13.4 μm, sphaeropendunculate, subovoid. Trama hyphae: cylindrical, often subinflated, thin-walled, smooth, branched, non-dextrinoid, 2‒5 μm wide. Pileipellis: a cutis made up of cylindrical, thin-walled, with weak annular ornamentation, 3‒10 μm wide hyphae; terminal elements adpressed to suberect, cylindrical, fusoid, clavate, 5‒11 μm wide. Stipitipellis: a cutis of cylindrical, thin-walled, smooth, 2.0‒6.0 μm wide hyphae. Caulocystidia: 12–38 × 2.4–6.6 μm, cylindrical, narrowly utriform, sometimes irregular, or curved. Clamp connections: present in all tissues.
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 Republic of 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 Republic 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.
Scattered to gregarious on the ground covered with dead and decaying conifer needles, from summer to autumn.
The Republic of Korea.
Collybiopsis vellerea is morphologically similar to Co. menehune and G. spongiosus Halling. Collybiopsis menehune has a paler stipe, a smaller pileus (8–30 mm), and fewer lamellulae (4–6 series) (
In this study, many epithets were found that required an additional transfer of species from Marasmiellus to Collybiopsis apart from the study done by
Marasmiellus istanbulensis E. Sesli, Antonín & E.Aytaç. Pl. Biosystems 152(4): 669. 2018.
Marasmiellus koreanus Antonín, Ryoo and H.D.Shin. Mycotaxon 112: 190. 2010.
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(1–6): 385. 1955.
Gymnopus omphalodes (Berk.) Halling & J.L. Mata, in Mata, Halling, and Petersen, Fungal Diversity 16: 122. 2004.
Marasmius omphalodes Berk., Hooker’s J. Bot. Kew Gard. Misc. 8: 138. 1856.
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”.
Collybia pseudomphalodes Dennis, Kew Bull. 15(1): 74 (1961).
Gymnopus ramulicola T.H. Li & S.F. Deng, in Deng, Li, Jiang and Song, Mycotaxon 131(3): 665. 2016.
1 | Pileus < 25 mm diam | 2 |
– | Pileus > 25 mm diam | 11 |
2 | Lamellae subdistant to distant (10–30) | 3 |
– | Lamellae close to crowded (> 30) | 9 |
3 | Basidiomes on bark, branch, or woody debris | 4 |
– | Basidiomes on duff or on soil | 8 |
4 | Pleurocystidia present | 5 |
– | Pleurocystidia absent | 7 |
5 | Stipe base covered with dense whitish basal tomentum | Co. albicantipes |
– | Stipe base not covered with whitish basal tomentum | 6 |
6 | Pileipellis composed of a coarse Rameales-structure hyphae | Co. ramealis |
– | Pileipellis composed of a cylindrical, often sub-inflated hyphae, not a Rameales-structure | Co. fulva |
7 | Pileus distinctly sulcate. Stipe base covered with dense whitish basal tomentum | Co. koreana |
– | Pileus slightly sulcate. Stipe base covered with weak whitish basal tomentum | Co. nonulla |
8 | Stipe < 2 cm long. Q value of basidiospores 1.6–2.2 | Co. dichroa |
– | Stipe > 2 cm long. Q value of basidiospores 2.0–3.1 | Co. undulata |
9 | Lamellae crowded (> 100) | Co. confluens |
– | Lamellae close to crowded (< 100) | 10 |
10 | Basidia > 22 µm long | Co. menehune |
– | Basidia < 22 µm long | Co. biformis |
11 | Lamellae subdistant to distant (10–38) | 12 |
– | Lamellae close (> 38) | 15 |
12 | Pleurocystidia present | 13 |
– | Pleurocystidia absent | 14 |
13 | Q value of basidiospores > 2.2 | Co. orientisubnuda |
– | Q value of basidiospores < 2.2 | Co. subumbilicata |
14 | Pileus convex, hemispherical, plano-convex to flat. Cheilocystidia utriform and clavate | Co. clavicystidiata |
– | Pileus convex to broad-convex. Cheilocystidia narrowly clavate | Co. polygramma |
15 | Pleurocystidia present | Co. vellerea |
– | Pleurocystidia absent | Co. luxurians |
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 characteristics are overlapped between species (Suppl. material
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. nonnulla, 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 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 (
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 (
Morphologically, the morphological characteristics of the seven described species were also in agreement with the previous descriptions (Suppl. material
Seven new species have common characteristics of Collybiopsis 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 species based on morphological characteristics alone. Upon molecular phylogenetic analyses, each of them clearly formed a distinct clade clearly in the ML phylogenetic tree (Fig.
Two species previously reported in the Republic of Korea, Co. peronata (Cho & Lee, 1979) and Co. subnuda (
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 significance as in the previous study conducted by
We greatly appreciate Prof. R.H. Petersen for his valuable comments on this manuscript. This study was supported by the National Institute of Biological Resources (NIBR202203112) and the Korea National Arboretum (KNA1-1-25, 19-2).
Figure S1
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 µm.
Figure S2
Data type: Jpg file.
Explanation note: Comparison of the morphological characters of 16 Korean Collybiopsis species.
Figure S3
Data type: Jpg file.
Explanation note: Sequence difference between the three species in the ITS region.