菌物学报

Species	Voucher specimen	Geographical origin	GenBank accession number
R. badia	587/BB 07.324 (PC0124709)	Slovakia	PX726806	KU237571	KU237715	KU237857	KU237999
R. carminipes	531/BB 07.192 (PC0124681)	Slovakia	PX726805	KU237523	KU237673	KU237809	KU237953
R. claroflava	WXH10952 (HKAS 146581)	Sweden	PV974715	PV973112	PV999097	PV999168	PV999017
R. cremeoavellanea	WXH10943 (HKAS 149035)	Sweden	PV974716	PV973113	PV999098	PV999169	PV999018
R. decolorans	WXH11050 (HKAS 146587)	Sweden	PV974717	PV973114	PV999099	PV999170	PV999019
R. decolorans	WXH11069 (HKAS 146590)	Sweden	PV974718	PV973115	PV999100	PV999171	PV999020
R. dhakuriana	CXH02 (HKAS 86727)	Shanghang, Fujian	PX622224 PX622225	PX558766	PX753901	PX580453 PX580454	PX694204
R. dhakuriana	WXH4518 (HKAS 104867)	Xinping, Yunnan	PX622226 PX622227	PX558767	PX580446 PX580447	-	PX580449
R. dhakuriana	YZL7713 (HKAS 150857)	Tengchong, Yunnan	PX622230	PX558768	PX694206	PX580455 PX580456	PX580450
R. griseocarnosa cladeⅠ	CXH03 (HKAS 86728)	Dapu, Guangdong	PV974585	PV973066	PV999049	PV999126	PV998967
R. griseocarnosa cladeⅠ	CXH04 (HKAS 86729)	Tengxian, Guangxi	PV974586	PV973067	PV999050	PV999127	PV998968
R. griseocarnosa cladeⅠ	CXH06 (HKAS 86731)	Pubei, Guangxi	PV974587	PV973068	PV999051	PV999128	PV998969
R. griseocarnosa cladeⅠ	CXH07 (HKAS 86732)	Yanling, Hunan	PV974588	PV973069	PV999052	PV999129	PV998970
R. griseocarnosa cladeⅠ	CXH10 (HKAS 86735)	Pubei, Guangxi	PV974589	PV973070	PV999053	PV999130	PV998971
R. griseocarnosa cladeⅠ	RTJ05 (HKAS 149431)	Sanming, Fujian	PV974578 PV974579	PV973065	PV999048	PV999125	PV998966
R. griseocarnosa cladeⅠ	WXH2125b (HKAS 51839b) holotype p.p.	Jinghong, Yunnan	PV974584*	PV973140*	PV999118*	PV999191*	PV999040*
R. griseocarnosa cladeⅡ	CXH01 (HKAS 86726)	Shanghang, Fujian	PV974639	PV973077	PV999060	PV999137	PV998980
R. griseocarnosa cladeⅡ	CXH05 (HKAS 86730)	Rongxian, Guangxi	PV974640	PV973078	PV999061	PV999138	PV998981
R. griseocarnosa cladeⅡ	HJW02 (HKAS 149378)	Changting, Fujian	PV974605	PV973073	PV999056	PV999133	PV998975
R. griseocarnosa cladeⅡ	HJW07 (HKAS 149383)	Changting, Fujian	PV974610	PV973074	PV999057	PV999134	PV998976
R. griseocarnosa cladeⅡ	LJM12 (HKAS 149404)	Pucheng, Fujian	PV974599	PV973072	PV999055	PV999132	PV998973 PV998974
R. griseocarnosa cladeⅡ	RTJ02 (HKAS 149428)	Nanping, Fujian	PV974625 PV974626	PV973075	PV999058	PV999135	PV998977 PV998978
R. griseocarnosa cladeⅡ	WXH3163 (HKAS 73659)	Shixing, Guangdong	PV974630	PV973076	PV999059	PV999136	PV998979
R. griseocarnosa cladeⅡ	YYB04 (HKAS 149449)	Gutian, Fujian	PV974641	PV973079	PV999062 PV999063	PV999139	PV998982
R. griseocarnosa cladeⅡ	YYB10 (HKAS 149455)	Gutian, Fujian	PV974642 PV974643	PV973080	PV999064	PV999140	PV998983
R. griseocarnosa cladeⅡ	ZGY02 (HKAS 149353)	Rongxian, Guangxi	PV974591	PV973071	PV999054	PV999131	PV998972
R. griseocarnosa clade Ⅲ	WXH4426 (HKAS 104790)	Maguan, Yunnan	PV974644	PV973089	PV999072	PV999148	PV998991
R. griseocarnosa clade Ⅲ	WXH4506 (HKAS 104808)	Xinping, Yunnan	PV974645	PV973090	PV999073	PV999149	PV998992
R. griseocarnosa clade Ⅳ	CXH12 (HKAS 86737)	Pu'er, Yunnan	PV974647	PV973091	PV999075	PV999150	PV998993
R. griseocarnosa clade Ⅳ	LSH28 (HKAS 149482)	Jingdong, Yunnan	PV974646	PV973141*	PV999074	PV999192*	PV999041
R. griseocarnosa clade Ⅴ	WXH3527 (HKAS 76064)	Lüchun, Yunnan	PV974649	PV973092	PV999076	PV999151	PV998994
R. griseocarnosa clade Ⅴ	ZP-2152 (MHHNU 8652)	Sangzhi, Hunan	PV974648*	PV973142*	PV999120*	PV999193*	PV999043*
R. griseocarnosa clade Ⅵ	LYC4252 (HKAS 149392)	Pingbian, Yunnan	PV974652	PV973094	PV999078	PV999153	PV998996
R. griseocarnosa clade Ⅵ	WXH2125c (HKAS 51839c) holotype p.p.	Jinghong, Yunnan	PV974650*	PV973143*	PV999119*	PV999194*	PV999042*
R. griseocarnosa clade Ⅵ	YYB07 (HKAS 149452)	Gutian, Fujian	PV974651	PV973093	PV999077	PV999152	PV998995
R. griseocarnosa clade Ⅵ	YZL6649 (HKAS 126726)	Ruili, Yunnan	PV974653	PV973095	PV999079	PV999154	PV998997
R. griseocarnosa clade Ⅶ	CXH09 (HKAS 86734)	Jiaoling, Guangdong	PV974657	PV973100	PV999085	PV999158	PV999003
R. griseocarnosa clade Ⅶ	ZGY08 (HKAS 149357)	Maoming, Guangdong	PV974654	PV973096 PV973097	PV999080	PV999155	PV998998 PV998999
R. griseocarnosa clade Ⅶ	ZGY09 (HKAS 149358)	Maoming, Guangdong	PV974655	PV973098	PV999081 PV999082	PV999156	PV999000 PV999001
R. griseocarnosa clade Ⅶ	ZGY19 (HKAS 149364)	Bobai, Guangxi	PV974656	PV973099	PV999083 PV999084	PV999157	PV999002
R. griseocarnosa clade Ⅷ	MJM01 (HKAS 149417)	Nanping, Fujian	PV974677 PV974678	PV973101 PV973102	PV999086	PV999159	PV999004
R. griseocarnosa clade Ⅷ	MJM08 (HKAS 149423)	Nanping, Fujian	PV974682	PV973103	PV999087	PV999160	PV999005 PV999006
R. griseocarnosa clade Ⅷ	RTJ04 (HKAS 149430)	Nanping, Fujian	PV974687 PV974688	PV973104	PV999088	PV999161	PV999007
R. griseocarnosa clade Ⅷ	WXH1989a (HKAS 51713) paratype	Jinghong, Yunnan	PV974692*	PV973144*	PV999121*	PV999195*	PV999044*
R. griseocarnosa clade Ⅷ	WXH1989b (HKAS 51713) paratype	Jinghong, Yunnan	PV974693*	PV973145*	PV999122*	PV999196*	PV999045*
R. griseocarnosa clade Ⅷ	WXH2125a (HKAS 51839a) lectotype	Jinghong, Yunnan	PV974694*	PV973146*	PV999123*	PV999197*	PV999046*
R. griseocarnosa clade Ⅷ	WXH3509 (HKAS 76051)	Pu'er, Yunnan	PV974695	OP794377	OP805334	OP804711	PV999008
R. griseocarnosa clade Ⅷ	YYB02 (HKAS 149447)	Gutian, Fujian	PV974698 PV974699	PV973105	PV999089	PV999162	PV999009 PV999010
R. griseocarnosa var. ailaoshanensis	LSH43 (HKAS 149483)	Jingdong, Yunnan	PV974624	PV973086	PV999069	PV999145	PV998988
R. griseocarnosa var. ailaoshanensis	SKM2 (HKAS 149437) holotype	Jingdong, Yunnan	PV974615	PV973081	PV999065	PV999141	PV998984
R. griseocarnosa var. ailaoshanensis	SKM5 (HKAS 149439)	Jingdong, Yunnan	PV974618	PV973082	PV999066	PV999142	PV998985
R. griseocarnosa var. ailaoshanensis	SKM6 (HKAS 149440)	Jingdong, Yunnan	PV974619	PV973083	PV999067	PV999143	PV998986
R. griseocarnosa var. ailaoshanensis	SKM7 (HKAS 149441)	Jingdong, Yunnan	PV974620	PV973084	PV999068	PV999144	PV998987
R. griseocarnosa var. ailaoshanensis	WXH4646 (HKAS 109819)	Nanhua, Yunnan	PV974636	PV973087	PV999070	PV999146	PV998989
R. griseocarnosa var. ailaoshanensis	WXH4649 (HKAS 104735)	Nanhua, Yunnan	PV974638	PV973088	PV999071	PV999147	PV998990
R. griseocarnosa var. ailaoshanensis	YZL7597 (HKAS 150851)	Baoshan, Yunnan	PX622248	PX558769	PX580448	PX580457	PX580451
R. integra	518/BB 07.198 (PC0124682)	Slovakia	PX726804	KU237513	KU237663	KU237799	KU237943
R. laeta	519/BB 07.267 (PC0124692)	Slovakia	PX726803	KU237514	KU237664	KU237800	KU237944
R. laricina	575/BB 08.681		JN944008	KU237560	KU237704	KU237846	KU237991
R. lepida	437/BB 07.189 (PC0124680)	Slovakia	PX726807 PX726808	KU237500	KU237649	KU237786	KU237930
R. aff. lepida	QWQ-R064 (HKAS 110974)	Zhangjiajie, Hunan	PV974708	PV973107	PV999091	PV999164	PV999012 PV999013
R. aff. lepida	WJ193 (HKAS 104840)	Jingdong, Yunnan	PV974707	PV973106	PV999090	PV999163	PV999011
R. aff. lepida	WXH4489 (HKAS 104801)	Zhenyuan, Yunnan	PV974710 PV974711	PV973108 PV973109	PV999092	PV999165	PV999014
R. cf. lepida	WJ211 (HKAS 104857)	Jingdong, Yunnan	PV974713	PV973110	PV999093 PV999094	PV999166	PV999015
R. cf. lepida	WXH4488 (HKAS 104869)	Zhenyuan, Yunnan	PV974714	PV973111	PV999095 PV999096	PV999167	PV999016
R. occulta	WJ192 (HKAS 104858) holotype	Jingdong, Yunnan	PV974729	PV973122 PV973123	PV999107	PV999178 PV999179	PV999027
R. occulta	CXH11 (HKAS 86736)	Pu'er, Yunnan	PV974728	PV973121	PV999106	PV999177	PV999026
R. occulta	WJ194 (HKAS 110373)	Jingdong, Yunnan	PV974730	PV973124	PV999108	PV999180	PV999028
R. occulta	WXH4492 (HKAS 104866)	Zhenyuan, Yunnan	PV974735	PV973126	PV999110	PV999183	PV999029
R. occulta	WXH4656 (HKAS 104854)	Nanhua, Yunnan	PV974737 PV974738	PV973127	PV999111	PV999184	PV999030
R. occulta	WXH4657 (HKAS 104853)	Nanhua, Yunnan	PV974739	PV973128	PV999112	PV999185	PV999031
R. occulta	WXH4662 (HKAS 104852)	Nanhua, Yunnan	PV974741	PV973130	PV999113	PV999186	PV999034
R. occulta	WXH4665 (HKAS 104851)	Nanhua, Yunnan	PV974742	PV973131	PV999114	PV999187	PV999035
R. occulta	ZP01 (HKAS 149414)	Sanming, Fujian	PV974743	PV973132	PV999115	PV999188	PV999036
R. paludosa	442/BB 07.330 (PC0142538)	Slovakia	PX754887	KU237505	KU237654	KU237791	KU237935
R. purpureozonata	WXH5155 (HKAS 111615)	Deqin, Yunnan	PV974723	PV973118	PV999103	PV999174	PV999023
R. purpureozonata	WXH8294 (HKAS 117918)	Deqin, Yunnan	PV974720	PV973116	PV999101	PV999172	PV999021
R. purpureozonata	WXH8707 (HKAS 118314)	Deqin, Yunnan	PV974722	PV973117	PV999102	PV999173	PV999022
R. quercina	WP-1 (HKAS 149469)	Kunming, Yunnan	PV974725	PV973119	PV999104	PV999175	PV999024
R. quercina	WP-3 (HKAS 149471)	Kunming, Yunnan	PV974727	PV973120	PV999105	PV999176	PV999025
R. subdensifolia	552/BB 05.158 (PC0124674)	USA	PX754886	KU237544	-	KU237830	KU237974
R. vinosa	WXH11053 (HKAS 146588)	Sweden	PV974746	PV973133	-	PV999189	PV999037
R. yanheensis	WXH4661 (HKAS 104879)	Nanhua, Yunnan	PV974748 PV974748	PV973136	PV999117	PV999190	PV999039
R. yanheensis	WXH7157 (HKAS 120113)	Jianchuan, Yunnan	PV974747	PV973134 PV973135	PV999116	-	PV999038
Russula sp.1	WXH4493 (HKAS 110297)	Zhenyuan, Yunnan	PV974736	PV973147*	PV999124*	PV999198*	PV999047*
Russula sp.2	WXH4658 (HKAS 104855)	Nanhua, Yunnan	PV974740	PV973129	PX694205	PV999182	PV999032
Russula sp.2	WJ50 (HKAS 104863)	Maguan, Yunnan	PV974731	PV973125	PV999109	PV999181	PV999033

Species	Voucher specimen	Geographical origin	GenBank accession number
R. badia	587/BB 07.324 (PC0124709)	Slovakia	PX726806	KU237571	KU237715	KU237857	KU237999
R. carminipes	531/BB 07.192 (PC0124681)	Slovakia	PX726805	KU237523	KU237673	KU237809	KU237953
R. claroflava	WXH10952 (HKAS 146581)	Sweden	PV974715	PV973112	PV999097	PV999168	PV999017
R. cremeoavellanea	WXH10943 (HKAS 149035)	Sweden	PV974716	PV973113	PV999098	PV999169	PV999018
R. decolorans	WXH11050 (HKAS 146587)	Sweden	PV974717	PV973114	PV999099	PV999170	PV999019
R. decolorans	WXH11069 (HKAS 146590)	Sweden	PV974718	PV973115	PV999100	PV999171	PV999020
R. dhakuriana	CXH02 (HKAS 86727)	Shanghang, Fujian	PX622224 PX622225	PX558766	PX753901	PX580453 PX580454	PX694204
R. dhakuriana	WXH4518 (HKAS 104867)	Xinping, Yunnan	PX622226 PX622227	PX558767	PX580446 PX580447	-	PX580449
R. dhakuriana	YZL7713 (HKAS 150857)	Tengchong, Yunnan	PX622230	PX558768	PX694206	PX580455 PX580456	PX580450
R. griseocarnosa cladeⅠ	CXH03 (HKAS 86728)	Dapu, Guangdong	PV974585	PV973066	PV999049	PV999126	PV998967
R. griseocarnosa cladeⅠ	CXH04 (HKAS 86729)	Tengxian, Guangxi	PV974586	PV973067	PV999050	PV999127	PV998968
R. griseocarnosa cladeⅠ	CXH06 (HKAS 86731)	Pubei, Guangxi	PV974587	PV973068	PV999051	PV999128	PV998969
R. griseocarnosa cladeⅠ	CXH07 (HKAS 86732)	Yanling, Hunan	PV974588	PV973069	PV999052	PV999129	PV998970
R. griseocarnosa cladeⅠ	CXH10 (HKAS 86735)	Pubei, Guangxi	PV974589	PV973070	PV999053	PV999130	PV998971
R. griseocarnosa cladeⅠ	RTJ05 (HKAS 149431)	Sanming, Fujian	PV974578 PV974579	PV973065	PV999048	PV999125	PV998966
R. griseocarnosa cladeⅠ	WXH2125b (HKAS 51839b) holotype p.p.	Jinghong, Yunnan	PV974584*	PV973140*	PV999118*	PV999191*	PV999040*
R. griseocarnosa cladeⅡ	CXH01 (HKAS 86726)	Shanghang, Fujian	PV974639	PV973077	PV999060	PV999137	PV998980
R. griseocarnosa cladeⅡ	CXH05 (HKAS 86730)	Rongxian, Guangxi	PV974640	PV973078	PV999061	PV999138	PV998981
R. griseocarnosa cladeⅡ	HJW02 (HKAS 149378)	Changting, Fujian	PV974605	PV973073	PV999056	PV999133	PV998975
R. griseocarnosa cladeⅡ	HJW07 (HKAS 149383)	Changting, Fujian	PV974610	PV973074	PV999057	PV999134	PV998976
R. griseocarnosa cladeⅡ	LJM12 (HKAS 149404)	Pucheng, Fujian	PV974599	PV973072	PV999055	PV999132	PV998973 PV998974
R. griseocarnosa cladeⅡ	RTJ02 (HKAS 149428)	Nanping, Fujian	PV974625 PV974626	PV973075	PV999058	PV999135	PV998977 PV998978
R. griseocarnosa cladeⅡ	WXH3163 (HKAS 73659)	Shixing, Guangdong	PV974630	PV973076	PV999059	PV999136	PV998979
R. griseocarnosa cladeⅡ	YYB04 (HKAS 149449)	Gutian, Fujian	PV974641	PV973079	PV999062 PV999063	PV999139	PV998982
R. griseocarnosa cladeⅡ	YYB10 (HKAS 149455)	Gutian, Fujian	PV974642 PV974643	PV973080	PV999064	PV999140	PV998983
R. griseocarnosa cladeⅡ	ZGY02 (HKAS 149353)	Rongxian, Guangxi	PV974591	PV973071	PV999054	PV999131	PV998972
R. griseocarnosa clade Ⅲ	WXH4426 (HKAS 104790)	Maguan, Yunnan	PV974644	PV973089	PV999072	PV999148	PV998991
R. griseocarnosa clade Ⅲ	WXH4506 (HKAS 104808)	Xinping, Yunnan	PV974645	PV973090	PV999073	PV999149	PV998992
R. griseocarnosa clade Ⅳ	CXH12 (HKAS 86737)	Pu'er, Yunnan	PV974647	PV973091	PV999075	PV999150	PV998993
R. griseocarnosa clade Ⅳ	LSH28 (HKAS 149482)	Jingdong, Yunnan	PV974646	PV973141*	PV999074	PV999192*	PV999041
R. griseocarnosa clade Ⅴ	WXH3527 (HKAS 76064)	Lüchun, Yunnan	PV974649	PV973092	PV999076	PV999151	PV998994
R. griseocarnosa clade Ⅴ	ZP-2152 (MHHNU 8652)	Sangzhi, Hunan	PV974648*	PV973142*	PV999120*	PV999193*	PV999043*
R. griseocarnosa clade Ⅵ	LYC4252 (HKAS 149392)	Pingbian, Yunnan	PV974652	PV973094	PV999078	PV999153	PV998996
R. griseocarnosa clade Ⅵ	WXH2125c (HKAS 51839c) holotype p.p.	Jinghong, Yunnan	PV974650*	PV973143*	PV999119*	PV999194*	PV999042*
R. griseocarnosa clade Ⅵ	YYB07 (HKAS 149452)	Gutian, Fujian	PV974651	PV973093	PV999077	PV999152	PV998995
R. griseocarnosa clade Ⅵ	YZL6649 (HKAS 126726)	Ruili, Yunnan	PV974653	PV973095	PV999079	PV999154	PV998997
R. griseocarnosa clade Ⅶ	CXH09 (HKAS 86734)	Jiaoling, Guangdong	PV974657	PV973100	PV999085	PV999158	PV999003
R. griseocarnosa clade Ⅶ	ZGY08 (HKAS 149357)	Maoming, Guangdong	PV974654	PV973096 PV973097	PV999080	PV999155	PV998998 PV998999
R. griseocarnosa clade Ⅶ	ZGY09 (HKAS 149358)	Maoming, Guangdong	PV974655	PV973098	PV999081 PV999082	PV999156	PV999000 PV999001
R. griseocarnosa clade Ⅶ	ZGY19 (HKAS 149364)	Bobai, Guangxi	PV974656	PV973099	PV999083 PV999084	PV999157	PV999002
R. griseocarnosa clade Ⅷ	MJM01 (HKAS 149417)	Nanping, Fujian	PV974677 PV974678	PV973101 PV973102	PV999086	PV999159	PV999004
R. griseocarnosa clade Ⅷ	MJM08 (HKAS 149423)	Nanping, Fujian	PV974682	PV973103	PV999087	PV999160	PV999005 PV999006
R. griseocarnosa clade Ⅷ	RTJ04 (HKAS 149430)	Nanping, Fujian	PV974687 PV974688	PV973104	PV999088	PV999161	PV999007
R. griseocarnosa clade Ⅷ	WXH1989a (HKAS 51713) paratype	Jinghong, Yunnan	PV974692*	PV973144*	PV999121*	PV999195*	PV999044*
R. griseocarnosa clade Ⅷ	WXH1989b (HKAS 51713) paratype	Jinghong, Yunnan	PV974693*	PV973145*	PV999122*	PV999196*	PV999045*
R. griseocarnosa clade Ⅷ	WXH2125a (HKAS 51839a) lectotype	Jinghong, Yunnan	PV974694*	PV973146*	PV999123*	PV999197*	PV999046*
R. griseocarnosa clade Ⅷ	WXH3509 (HKAS 76051)	Pu'er, Yunnan	PV974695	OP794377	OP805334	OP804711	PV999008
R. griseocarnosa clade Ⅷ	YYB02 (HKAS 149447)	Gutian, Fujian	PV974698 PV974699	PV973105	PV999089	PV999162	PV999009 PV999010
R. griseocarnosa var. ailaoshanensis	LSH43 (HKAS 149483)	Jingdong, Yunnan	PV974624	PV973086	PV999069	PV999145	PV998988
R. griseocarnosa var. ailaoshanensis	SKM2 (HKAS 149437) holotype	Jingdong, Yunnan	PV974615	PV973081	PV999065	PV999141	PV998984
R. griseocarnosa var. ailaoshanensis	SKM5 (HKAS 149439)	Jingdong, Yunnan	PV974618	PV973082	PV999066	PV999142	PV998985
R. griseocarnosa var. ailaoshanensis	SKM6 (HKAS 149440)	Jingdong, Yunnan	PV974619	PV973083	PV999067	PV999143	PV998986
R. griseocarnosa var. ailaoshanensis	SKM7 (HKAS 149441)	Jingdong, Yunnan	PV974620	PV973084	PV999068	PV999144	PV998987
R. griseocarnosa var. ailaoshanensis	WXH4646 (HKAS 109819)	Nanhua, Yunnan	PV974636	PV973087	PV999070	PV999146	PV998989
R. griseocarnosa var. ailaoshanensis	WXH4649 (HKAS 104735)	Nanhua, Yunnan	PV974638	PV973088	PV999071	PV999147	PV998990
R. griseocarnosa var. ailaoshanensis	YZL7597 (HKAS 150851)	Baoshan, Yunnan	PX622248	PX558769	PX580448	PX580457	PX580451
R. integra	518/BB 07.198 (PC0124682)	Slovakia	PX726804	KU237513	KU237663	KU237799	KU237943
R. laeta	519/BB 07.267 (PC0124692)	Slovakia	PX726803	KU237514	KU237664	KU237800	KU237944
R. laricina	575/BB 08.681		JN944008	KU237560	KU237704	KU237846	KU237991
R. lepida	437/BB 07.189 (PC0124680)	Slovakia	PX726807 PX726808	KU237500	KU237649	KU237786	KU237930
R. aff. lepida	QWQ-R064 (HKAS 110974)	Zhangjiajie, Hunan	PV974708	PV973107	PV999091	PV999164	PV999012 PV999013
R. aff. lepida	WJ193 (HKAS 104840)	Jingdong, Yunnan	PV974707	PV973106	PV999090	PV999163	PV999011
R. aff. lepida	WXH4489 (HKAS 104801)	Zhenyuan, Yunnan	PV974710 PV974711	PV973108 PV973109	PV999092	PV999165	PV999014
R. cf. lepida	WJ211 (HKAS 104857)	Jingdong, Yunnan	PV974713	PV973110	PV999093 PV999094	PV999166	PV999015
R. cf. lepida	WXH4488 (HKAS 104869)	Zhenyuan, Yunnan	PV974714	PV973111	PV999095 PV999096	PV999167	PV999016
R. occulta	WJ192 (HKAS 104858) holotype	Jingdong, Yunnan	PV974729	PV973122 PV973123	PV999107	PV999178 PV999179	PV999027
R. occulta	CXH11 (HKAS 86736)	Pu'er, Yunnan	PV974728	PV973121	PV999106	PV999177	PV999026
R. occulta	WJ194 (HKAS 110373)	Jingdong, Yunnan	PV974730	PV973124	PV999108	PV999180	PV999028
R. occulta	WXH4492 (HKAS 104866)	Zhenyuan, Yunnan	PV974735	PV973126	PV999110	PV999183	PV999029
R. occulta	WXH4656 (HKAS 104854)	Nanhua, Yunnan	PV974737 PV974738	PV973127	PV999111	PV999184	PV999030
R. occulta	WXH4657 (HKAS 104853)	Nanhua, Yunnan	PV974739	PV973128	PV999112	PV999185	PV999031
R. occulta	WXH4662 (HKAS 104852)	Nanhua, Yunnan	PV974741	PV973130	PV999113	PV999186	PV999034
R. occulta	WXH4665 (HKAS 104851)	Nanhua, Yunnan	PV974742	PV973131	PV999114	PV999187	PV999035
R. occulta	ZP01 (HKAS 149414)	Sanming, Fujian	PV974743	PV973132	PV999115	PV999188	PV999036
R. paludosa	442/BB 07.330 (PC0142538)	Slovakia	PX754887	KU237505	KU237654	KU237791	KU237935
R. purpureozonata	WXH5155 (HKAS 111615)	Deqin, Yunnan	PV974723	PV973118	PV999103	PV999174	PV999023
R. purpureozonata	WXH8294 (HKAS 117918)	Deqin, Yunnan	PV974720	PV973116	PV999101	PV999172	PV999021
R. purpureozonata	WXH8707 (HKAS 118314)	Deqin, Yunnan	PV974722	PV973117	PV999102	PV999173	PV999022
R. quercina	WP-1 (HKAS 149469)	Kunming, Yunnan	PV974725	PV973119	PV999104	PV999175	PV999024
R. quercina	WP-3 (HKAS 149471)	Kunming, Yunnan	PV974727	PV973120	PV999105	PV999176	PV999025
R. subdensifolia	552/BB 05.158 (PC0124674)	USA	PX754886	KU237544	-	KU237830	KU237974
R. vinosa	WXH11053 (HKAS 146588)	Sweden	PV974746	PV973133	-	PV999189	PV999037
R. yanheensis	WXH4661 (HKAS 104879)	Nanhua, Yunnan	PV974748 PV974748	PV973136	PV999117	PV999190	PV999039
R. yanheensis	WXH7157 (HKAS 120113)	Jianchuan, Yunnan	PV974747	PV973134 PV973135	PV999116	-	PV999038
Russula sp.1	WXH4493 (HKAS 110297)	Zhenyuan, Yunnan	PV974736	PV973147*	PV999124*	PV999198*	PV999047*
Russula sp.2	WXH4658 (HKAS 104855)	Nanhua, Yunnan	PV974740	PV973129	PX694205	PV999182	PV999032
Russula sp.2	WJ50 (HKAS 104863)	Maguan, Yunnan	PV974731	PV973125	PV999109	PV999181	PV999033

Clade	ITS	LSU	RPB2	TEF1-α	RPB1	Five-locus combined	Criterion satisfied
Ⅰ	98%	52%	-	55%	94%	100%	a & b
Ⅱ	100%	74%	96%	81%	-	100%	a & b
Ⅲ	96%	99%	100%	-	100%	100%	a & b
Ⅳ	84%	98%	-	-	-	90%	b
Ⅴ	90%	90%	-	-	-	100%	b
Ⅵ	Supported paraphyly	-	63%	-	76%	84%	Neither
Ⅶ	Supported paraphyly	-	96%	-	-	69%	Neither
Ⅵ+Ⅶ	Supported paraphyly	-	86%	23%	93%	98%	Neither
Ⅷ	-	56%	90%	51%	100%	100%	a & b
B	69%	80%	89%	-	100%	100%	a & b

Clade	ITS	LSU	RPB2	TEF1-α	RPB1	Five-locus combined	Criterion satisfied
Ⅰ	98%	52%	-	55%	94%	100%	a & b
Ⅱ	100%	74%	96%	81%	-	100%	a & b
Ⅲ	96%	99%	100%	-	100%	100%	a & b
Ⅳ	84%	98%	-	-	-	90%	b
Ⅴ	90%	90%	-	-	-	100%	b
Ⅵ	Supported paraphyly	-	63%	-	76%	84%	Neither
Ⅶ	Supported paraphyly	-	96%	-	-	69%	Neither
Ⅵ+Ⅶ	Supported paraphyly	-	86%	23%	93%	98%	Neither
Ⅷ	-	56%	90%	51%	100%	100%	a & b
B	69%	80%	89%	-	100%	100%	a & b

商品野生食用蘑菇“大红菌”物种多样性的重新评估

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张佩 ¹^,² , 李育武 ³ , 曹书琴 ² , 王晶 ⁴ , 苏开美 ⁵^,^* , 李树红 ⁵ , BUYCK Bart ⁶ , 王向华 ²^,^*

菌物学报 | 研究论文 2026,45(2): 250280

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菌物学报 | 研究论文 2026, 45(2): 250280

商品野生食用蘑菇“大红菌”物种多样性的重新评估

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张佩¹^,², 李育武³, 曹书琴², 王晶⁴, 苏开美⁵^,^*, 李树红⁵, BUYCK Bart⁶, 王向华²^,^*

作者信息

1 中国科学院大学生命科学学院，北京 101408

2 中国科学院昆明植物研究所云南省真菌多样性与绿色发展重点实验室，云南昆明 650201

3 哀牢山国家级自然保护区楚雄管护局，云南楚雄 675000

4 贵州科学院贵州省生物研究所，贵州贵阳 550009

5 云南省农业科学院，云南昆明 650221

6 系统进化与生物多样性研究所法国国家自然历史博物馆，巴黎 F-75005，法国

Critical re-assessment of species diversity of marketed wild edible mushroom ‘dahongjun'

Pei ZHANG¹^,², Yuwu LI³, Shuqin CAO², Jing WANG⁴, Kaimei SU⁵^,^*, Shuhong LI⁵, Bart BUYCK⁶, Xianghua WANG²^,^*

Affiliations

1 College of Life Sciences, University of Chinese Academy of Sciences, Beijing 101408, China

2 Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China

3 Chuxiong Management Bureau of Yunnan Ailao Mountains National Nature Reserve, Chuxiong 675000, Yunnan, China

4 Guizhou Institute of Biological Research, Guizhou Academy of Sciences, Guiyang 550009, Guizhou, China

5 Yunnan Academy of Agricultural Sciences, Kunming 650221, Yunnan, China

6 Institut pour la Systématique, Evolution, Biodiversité (ISYEB), Muséum national d'histoire naturelle, Sorbonne Université, CNRS, EPHE, Université des Antilles, F-75005 Paris, France

出版时间: 2026-02-22 doi: 10.13346/j.mycosystema.250280

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摘要

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“大红菌”是一类在我国南部广泛采食的具有红色菌盖的红菇。最初我国的大红菌被误定为欧洲的酒红菇。自2009年灰肉红菇Russula griseocarnosa被描述后，这一种名成为大红菌唯一可用的名称。然而，前人研究发现该种下存在多个系统发育分支，而这些分支是否代表了真正的物种尚待研究。通过在我国南方大红菌的主产区进行广泛取样，采用多基因片段进行系统发育物种识别，我们鉴定出除灰肉红菇外的9个物种，包括已知种达哈库尔红菇R. dhakuriana、栎生红菇R. quercina、紫环红菇R. purpureozonata和沿河红菇R. yanheensis。研究发现了灰肉红菇的姊妹种隐匿红菇R. occulta (新种)以及4个尚待描述的种，它们分别与鳞盖红菇R. lepida、落叶松红菇R. laricina和紫环红菇近缘。基于5个DNA位点，我们通过多基因谱系一致法进行系统发育物种识别，在灰肉红菇内界定了6个系统发育分支。基因谱系间存在的冲突提示将灰肉红菇处理为一个系统发育种较为合适，其下的6个分支可处理为种下分类单元。对灰肉红菇主模式标本进行的浅层基因组测序发现它的3个个体分属于3个不同的分支，故使用其中的一个个体为其指定了后选模式。我们描述了一个主产区在云南哀牢山区的一个新变种：灰肉红菇哀牢山变种R. griseocarnosa var. ailaoshanensis，它与原变种的区别在于其近白色的菌肉、较大的孢子和较膨大的盖表皮菌丝。本研究提供的形态和分子数据将有助于鉴定红菇亚属内的野生食用菌。

关键词

中国 / 后选模式 / 新种 / 系统发育物种识别 / 红菇属 / 野生食用菌

Abstract

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The informal term ‘dahongjun' or ‘big red mushroom' is a group of red-coloured russulas widely collected and consumed in southern China, originally mistakenly recognized as the European Russula vinosa. Russula griseocarnosa was described from China in 2009, and since then, it has become the only available scientific name for ‘dahongjun'. Nevertheless, several lineages exist within this complex, and whether these lineages represent real species is still an open question. Sampling in major producing areas of ‘dahongjun' and phylogenetic species recognition using multi-locus data were conducted. Besides R. griseocarnosa nine additional species were identified under the commercial name ‘dahongjun', including R. dhakuriana, R. quercina, R. purpureozonata, R. yanheensis and a new species sister to R. griseocarnosa here described as R. occulta. The others were undescribed species close to R. laricina, R. lepida, and R. purpureozonata respectively. Six evolutionary lineages within R. griseocarnosa were defined by genealogical concordance phylogenetic species recognition based on five-locus DNA data. Conflicts among different genealogies suggested that R. griseocarnosa is best considered as a single phylogenetic species, comprising several infraspecific taxa. Lower coverage whole genome sequencing of R. griseocarnosa holotype discovered that its three individuals belong to three different clades. Lectotypification was made using one of the three individuals. A new variety R. griseocarnosa var. ailaoshanensis is described to name one of the terminal clades that is mainly distributed in Ailao Mountains. This variety differs morphologically from the type variety in its nearly white context, bigger spores and more inflated hyphae in the pileipellis. The morphological and molecular data provided in this study are helpful to identify wild edible mushrooms of Russula subgen. Russula.

Key words

China / lectotype / new species / phylogenetic species recognition / Russula / wild edible fungi

引用本文

张佩, 李育武, 曹书琴, 王晶, 苏开美, 李树红, BUYCK Bart, 王向华. 商品野生食用蘑菇“大红菌”物种多样性的重新评估. 菌物学报, 2026 , 45 (2) : 250280 - . DOI: 10.13346/j.mycosystema.250280

Pei ZHANG, Yuwu LI, Shuqin CAO, Jing WANG, Kaimei SU, Shuhong LI, Bart BUYCK, Xianghua WANG. Critical re-assessment of species diversity of marketed wild edible mushroom ‘dahongjun'[J]. Mycosystema, 2026 , 45 (2) : 250280 - . DOI: 10.13346/j.mycosystema.250280

正文

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INTRODUCTION

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In southern China, Russula (Russulaceae, Russulales) species are frequently sold in local markets (Wang et al. 2004; Wu et al. 2019; Deng et al. 2020; Wang et al. 2022; Huang et al. 2023). Among them ‘dahongjun' (big red mushroom) is the most prized and sold at high price (RMB ¥70-100/kg or US $17.00 when fresh). The price of dried products (RMB ¥700-1 500/kg or US $100-200) is even higher than that of Tricholoma matsutake (S. Ito & S. Imai) Singer (matsutake mushroom). As an important non-timber forest product, ‘dahongjun' represents the main source of income for people living in the mountains in eastern (Jiangxi), central (Hunan), southern (Guangdong, Guangxi), southeastern (Fujian) and southwestern (Yunnan) China.

In field and markets, ‘dahongjun' can be recognized by a set of characteristics: basidiocarps robust, thick-fleshed; pileus vivid to dark red, dry or subgreasy, with a frequently much darker center and non-striate margin; context often graying; stipe surface often reddish; taste mostly mild (Wang et al. 2004; Chen 2010; Li & Hu 2010). Most of these characters are similar to those of European R. vinosa Lindblad, a name that was adopted for ‘dahongjun' until about 15 years ago. Wang et al. (2009) compared the ITS sequences and morphology of Chinese samples with those of R. vinosa and found that ‘dahongjun' was a different species, thus a distinct species R. griseocarnosa X.H. Wang et al. was described.

Soon after the publication of R. griseocarnosa, Li et al. (2010) sequenced a large number of samples of ‘dahongjun' from southern China and identified three divergent lineages. Similar genetic variation was simultaneously presented by Chen (2010) based on eight samples from Fujian, Guangdong and Guangxi. Furthermore, Yu (2013) found that two ‘dahongjun' samples from Fujian have ITS sequences only 91% similar to that of R. griseocarnosa. A large number of samples with such ITS type were recently discovered from central and southern Yunnan by Feng et al. (2020). These samples formed an additional major lineage to those discovered by Li et al. (2010). Yu et al. (2020) sequenced the genome of a sample of R. griseocarnosa collected from Guangxi. The ITS sequence of this sample is 97.7% similar to EF627042 (ITS of R. griseocarnosa holotype). In GenBank there are more sequences (e.g. MT211446 and MT252594) similar to those of R. griseocarnosa but are not covered by the lineages of Li et al. (2010) and Feng et al. (2020). Whether the samples outside the core R. griseocarnosa lineage belong to different species is still an open question.

R. griseocarnosa previously was regarded as a species with dark red pileus and mainly distributed in the tropics (Wang et al. 2009). Das et al. (2010) reported a specimen of R. griseocarnosa with less reddish but more ochraceus pileus collected from mid-montane forest in Sikkim Himalaya, India. Specimens with such colour (HKAS 104790 and HKAS 104808, KUN) were also collected in Ailao Mountains (mid-south Yunnan) at altitude > 2 000 m. Moreover, we obtained some specimens with nearly white context in the pileus and the sequences of these specimens did not fall into the same lineage of typical R. griseocarnosa. It was suggested that there might hide multiple species under the local name ‘dahongjun'.

This study aims at re-evaluating the divergent lineages of R. griseocarnosa in former studies and detecting whether they represent species different from R. griseocarnosa; determining the identity of the specimens morphologically different from typical R. griseocarnosa; locating these species in the most up-to-date multi-gene phylogeny of Russula (Buyck et al. 2018). Broader sampling, phylogenetic species recognition (PSR) and morphological comparison were taken to achieve these aims.

MATERIALS AND METHODS

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1.1 Sampling

Materials of ‘dahongjun' were obtained from the main producing areas in southern China (Fujian, Guangdong, Guangxi, Hunan, Sichuan and Yunnan). Ailao Mountains National Nature Reserve in central-southern Yunnan, one of the most famous localities of ‘dahongjun', was intensively visited. In total, 188 specimens were obtained: 82 from Fujian, 6 from Guangdong, 22 from Guangxi, 3 from Hunan and 75 from Yunnan. Among them, 63 specimens were collected in the field and 125 were purchased from markets or mushroom hunters. In addition, five specimens of four European edible species, i.e. R. cremeoavellanea Singer, R. decolorans (Fr). Fr., R. claroflava Grove and R. vinosa, were used for comparison purpose. Specimens are deposited in the Cryptogamic section (HKAS) of Herbarium of Kunming Institute of Botany, Chinese Academy of Sciences (KUN).

1.2 Morphological analysis

Macroscopical descriptions are based on field notes of fresh materials and microscopical ones on dried specimens. Colour codes follow Kornerup and Wanscher (1961). Methods of morphological observation followed Wang et al. (2009). Microscopic photos were taken with a charge- coupled device SC 2000C connected to a Nikon ECLIPSE Ni-U compound microscope (Nikon Corporation).

1.3 DNA extraction and sequencing

DNA extractions and PCR amplification followed Wang et al. (2009). Five nuclear markers commonly used for PSR and phylogenetic inference in Russulaceae (e.g. Buyck et al. 2018; de Lange et al. 2021; Huang et al. 2023), i.e. ITS, nrLSU, RPB1, RPB2 and TEF1-α, were amplified. Universal primers published by White et al. (1990), Moncalvo et al. (2000), Liu et al. (1999), Matheny et al. (2002) and Morehouse et al. (2003) and specific primers by Cao et al. (2023) were used. For sequences having INDELs among copies or ambiguous sites, PCR products were sequenced bidirectionally to make contigs of the amplified regions or verify the ambiguous sites. The method of Flot et al. (2006) was used to determine the INDEL heterozygosity and phase the sequences into two copies. Raw sequences generated with Sanger sequencing were edited and assembled in Sequencher v4.1.4 (Gene Codes Corporation). In rare cases where the amount of PCR products is not sufficient for sequencing or direct sequencing failed, they were cloned following the method of Wang et al. (2015). PCR of the holotype and one paratype (HKAS 51713) of R. griseocarnosa and several problematic samples failed. We thus conducted lower-coverage whole genome sequencing (“genome skimming”) by using the next generation sequencing on an Illumina NovoSeq-5500 platform to recover the target loci. Library preparation followed the protocol described by Zeng et al. (2018). The raw data were de novo assembled using GetOrganelle toolkit (Jin et al. 2020).

1.4 Multi-gene phylogenetic constructions

Representative species were selected from the Russula phylogeny of Buyck et al. (2018, 2020, 2024). Two ITS-nrLSU-RPB1-RPB2-TEF1-α combined datasets were used: dataset A to construct the phylogeny of R. subgen. Russula and B to resolve the phylogeny of R. griseocarnosa complex. In dataset A, ITS sequences were only included for the target species and their closely related species in the same section or subsection. We followed Buyck et al. (2018) to exclude the ambiguous sites of RPB2 and introns and to set partitions. In dataset B, introns could be well aligned and were thus kept. The partitioning was the same as in dataset A, with introns treated as additional partitions. Alignments were made by MAFFT v7 (Katoh & Toh 2008) using the L-INS-I strategy and manually adjusted in BioEdit v.7.1.3.0 (Hall 1999). All best tree searches and bootstrap analyses of maximum likelihood (ML) were conducted in RAxML v. 7.7.7 (Stamatakis 2014) using the rapid bootstrap algorithm. Other settings of ML analysis followed Buyck et al. (2024). Trees were viewed and exported in FigTree v1.3.1 and further processed with Adobe Illustrator (Fig. 1).

1.5 Phylogenetic species recognition

We followed the genealogical concordance phylogenetic species recognition (GCPSR) method of Dettman et al. (2003) to determine species limits in the R. griseocarnosa complex. Five loci, i.e., ITS, nrLSU, RPB1, RPB2 and TEF1-α, were used. Duplicate sequences were filtered, leaving those covering genetic and morphological variation and geographical origins. Single genealogies were constructed in RAxML v. 7.7.7 (Stamatakis 2014). All introns were kept and partitioning was similar to that in the multi-gene phylogenetic analysis. To identify independent evolutionary lineages under the GCPSR, two criteria were adopted: (i) genealogical concordance, the clade was present in the majority (4/5) of the single-locus genealogies; (ii) genealogical non-discordance, the clade was well supported in at least one single-locus genealogy, as judged by ML bootstrap (ML-bs) ≥ 70% and was not contradicted in any other single-locus genealogy at the same level of support. A conflict was assumed to be significant when two different relationships (one monophyletic and the other non-monophyletic) for the same set of taxa were both supported with ML-bs ≥ 70%. To decide which independent evolutionary lineages represented phylogenetic species, we conducted “exhaustive subdivision”, i.e., all individuals had to be placed within a phylogenetic species in the combined phylogeny.

2 RESULTS

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2.1 Multi-gene phylogenetic inference

In total, 534 sequences were generated and deposited in GenBank (Table 1, plus PV974553- PV974750 and PX622224-PX622256 for ITS). Analysis of these sequences indicated that all the samples of ‘dahongjun' collected from local markets belonged to the crown clade of clade Ⅷ in the phylogeny of Russula (Buyck et al. 2018) representing the majority of R. subgen. Russula. Ten species were identified. Among them, R. griseocarnosa, R. occulta (HKAS 104858, holotype), R. purpureozonata K. Das et al., R. quercina J.J. Zhou & R.Q. Ji and R. sp.1 (HKAS 110297) were grouped with European R. decolorans, R. cremeoavellanea, R. integra, R. vinosa and R. claroflava, forming a clade with moderate support (ML-bs 70%). Taxonomically this clade corresponds to R. subsect. Integrinae Sarnari. The other five species are R. yanheensis T.C. Wen et al. of R. subsect. Paludosinae Jul. Schäff., Russula sp.2 (HKAS 104863 and HKAS 104855) of R. sect. Laricinae (Romagn.) Bon., R. dhakuriana K. Das et al. of R. subsect. Roseinae Singer ex Sarnari and two species (R. aff. lepida and R. cf. lepida) of Russula sect. Flavisiccantes Buyck & V. Hofst. in Das et al. (2017).

2.2 Phylogenetic species recognition of the Russula griseocarnosa complex

Five loci of 57 representative samples were successfully sequenced. In the five-locus phylogeny, these specimens formed two major clades. Clade B (ML-bs 100%) consisted of nine specimens collected from Fujian (Sanming) and Yunnan (Nanhua, Jingdong and Xinping). This clade was present in all single genealogies with significant supports except for in TEF1-α, but the paraphyly in TEF1-α had no bootstrap support (Table 2). It met both genealogical concordance and genealogical non-discordance and was defined as an independent evolutionary lineage. Clade A (ML-bs 100%) was composed of 48 samples (Fig. 2) from Fujian, Guangdong, Guangxi, Hunan and Yunnan. These samples formed seven well-supported terminal clades (Ⅰ-Ⅵ and Ⅷ) and one clade with low support (Ⅶ, ML-bs 69%) (Fig. 2). All clades except for Ⅵ and Ⅶ met one or both criteria of evolutionary lineage (Table 2). Three of the four samples of clade Ⅵ (HKAS 51839c, 126726, 149452) formed a well-supported clade (ML-bs 75%) with HKAS 149364 of clade Ⅶ in the ITS genealogy (tree not shown). This contradicted the supported monophyly of clade Ⅵ in the RPB1 genealogy (ML-bs 76%) and the monophyly of clade Ⅶ in the RPB2 genealogy (ML-bs 96%). In the ITS genealogy, sample HKAS 86734 of clade Ⅶ and the two samples of clade Ⅳ(HKAS 86737 and HKAS 149482) formed a monophyly with significant support (ML-bs 81%, tree not shown). This also contradicted the monophyly of clade Ⅶ in the RPB2 genealogy and invalidated the recognition of evolutionary lineages of the clade Ⅵ+Ⅶ, despite the significant support (ML-bs 98%) in the combined phylogeny. To achieve exhaustive subdivision, all the samples of clade A were lumped into one phylogenetic species. Only two phylogenetic species were recognized: R. griseocarnosa and R. occulta, a new species to be described below. Within cladeⅡ, seven specimens from Ailao Mts. formed a terminal clade with long branch.

3 Nomenclature and taxonomy

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3.1 Lectotypification of Russula griseocarnosa

The holotype of R. griseocarnosa is composed of three fruiting bodies. By next-generation sequencing, we obtained multi-genes from all of them. These individuals fell into three different clades of R. griseocarnosa by GCPSR: HKAS 51839a in Ⅷ, HKAS 51839b in Ⅰ and HKAS 51839c in Ⅵ (Fig. 2). The ITS sequence of HKAS 51839c is identical with the published holotype sequence EF627042. The spore measurements of the holotype in the original description are from HKAS 51839a and HKAS 51839b and the line drawings are from HKAS 51839a.

To avoid potential confusion of the identity of R. griseocarnosa, we need to select a fruiting body (specimen) to determine the application of this species name. HKAS 51839b does not have any priority, since neither the published ITS sequence (EF627042) nor the line drawings was from it. In GCPSR, clade Ⅷ, where HKAS 51839a was nested, met both genealogical concordance and non-disconcordance, whereas clade Ⅵ, where HKAS 51839c was clustered, met neither. Moreover, clade Ⅷ includes the paratype HKAS 51713, and is the clade from which the morphological recognition of R. griseocarnosa is de facto made. Thus, HKAS 51839a has qualification to be a type and we here make a lectotypification for R. griseocarnosa using it: Lectotype (hic designatus): Yunnan Province, Jinghong, Xiaomengyang Township, Xishuangbanna National Nature Reserve, Guanping Station, 28 Jul. 2006, X.H. Wang 2125a (HKAS 51839a, KUN) (MycoBank registration number: MBT 10030945). The GenBank accession numbers of this lectotype are: PV974694-ITS, PV973146-nrLSU, PV999123- RPB1, PV999197-RPB2 and PV999046-TEF1-α.

3.2 Taxonomy

Russula griseocarnosa var. ailaoshanensis K.M. Su, S.H Li, Pei Zhang, Yu-Wu Li & X.H. Wang, var. nov. (灰肉红菇哀牢山变种) Fig. 3

MycoBank: MB 861588

Etymology: named after the type locality, Ailao Mts.

Holotype: China, Yunnan Province, Jingdong County, Taizhong Township, Ailao Mts. National Nature Reserve, 5 Aug. 2019, K.M. Su, no. SKM2 (HKAS 149437, KUN!).

Diagnosis: differing from R. griseocarnosa var. griseocarnosa by the white or less gray pileus context, bigger spores and more inflated hyphae in the subpellis of pileipellis.

Basidiocarps medium-sized to big, thick- fleshed, fragile. Pileus 70-90 mm diam., convex with a broadly depressed center; surface smooth, greasy when wet, soon dry, often with concentrical wrinkles, cuticle cracked when mature, dark red to vivid red, locally paler or with orange tinge, cuticle peeling around 70% towards the center; context white, rarely gray. Lamellae 3−6 mm broad, adnate, sub-crowded to crowded, brittle and fragile, nearly equal, whitish cream when young, then turning warm ochre, with rose red edge near the pileus margin; context 3-6 mm thick, white, rarely gray. Stipe 75-115 mm × 15-20 mm, cylindrical or slightly expanding downwards; surface smooth, white or red when young, pale to dark rose red when mature; context pale gray. Taste mild, rarely acrid then hurting.

Spores (180/8/8)(8-)8.5-9.3-10(-10.5) × (7-)7.5-7.9-8.5(-9) μm, Q= (1.04-)1.11-1.17- 1.25(-1.29), subglobose to broadly ellipsoid; ornamentation 1-1.8(-2) μm, composed of conical warts, warts mostly isolated, rarely fused; suprahilar spot totally amyloid. Basidia 40-65 × 11-15 μm, clavate, 4-spored. Hymenophoral gloeocystidia common to abundant, emergent, 85-130 × 12-18 μm, projecting 20-50(-85) μm beyond the basidia layer, base originating deeply from subhymenium or trama, fusiform, with flocculent, granular or crystalline contents, middle part often thick-walled (wall to 1 μm thick), apex rarely moniliform or mamillate. Lamellar edge fertile or sterile; cystidia rare to abundant, 67-126 × 8-16 μm, fusiform, some cylindrical-clavate, with flocculent or granular contents. Pileipellis orthochromatic in cresyl blue, an (ixo) trichoderm, 120-220 μm thick, hyphae compactly ascending, branching, locally with long repent hyphae, with disperse rose pink pigmentation; hyphae in subpellis mostly 3-7 μm wide, some inflated to 13-15 µm wide; hyphae in suprapellis 3-7 µm wide, terminal cells 19-80 × 4-8 (-10) μm, cylindrical, fusiform, with scattered zebroid ornamentation; pileocystidia absent. Stipitipellis a cutis, 50-150 μm; hyphae 2-5 μm wide, hyaline; cystidia absent. Rosettes abundant in gill, pileus and stipe trama. Clamp connections absent.

Ecology: in group, in fagaceous forests, subtropical, mid-montane.

Distribution: Yunnan (Baoshan City, Dali City, Jingdong, Jinggu, Lincang, Nanhua).

Additional specimens examined: China, Yunnan Province, Chuxiong City, Xishelu Township, Ailao Mts. National Nature Reserve, Gamo Village, collected from local mushroom hunters, 7 Sept. 2024, X.H. Wang & P. Zhang, no. XHW12556 (HKAS 149442), XHW12557 (HKAS 149443), XHW12558 (HKAS 149444), XHW12559 (HKAS 149445); Jingdong County, Taizhong Township, Ailao Mts. National Natural Reserve, Xujiaba, 5 Aug. 2019, K.M. Su, no. SKM1-1 (HKAS 149435), SKM1-2 (HKAS 149436), SKM2 (HKAS 149437), SKM4 (HKAS 149438), SKM4-2 (HKAS 149468), SKM5 (HKAS 149439), SKM6 (HKAS 149440), SKM7 (HKAS 149441), SKM8 (HKAS 149464), SKM9 (HKAS 149465), SKM10 (HKAS 149466), SKM11 (HKAS 149467); Jingdong County, Ailao Mts. National Nature Reserve, 10 Aug. 2019, J. He, no. LSH03 (HKAS 149481), LSH43 (HKAS 149483); Nanhua County, Majie Township, Ailao Mts. National Nature Reserve, Dazhongshan, 16 Aug. 2017, X.H. Wang, B. Buyck & J. Wang, no. XHW4646 (HKAS 109819), XHW4648 (HKAS 111683), XHW4649 (HKAS 104735); Nanhua County, Majie Township, Ailao Mts. National Nature Reserve, Dazhongshan, Boluoshan, in fagaceous forest, 24°48ʹ11.10ʺN 100°51ʹ34.59ʺE, alt. 2 551 m, 14 Aug. 2024, X.H. Wang & P. Zhang, no. XHW11935 (HKAS 145185).

Note: This variety corresponds to lineage 1-1 in Li et al. (2010) and part of cladeⅠin Feng et al. (2020). All the specimens of this variety in Li et al. (2010) and in the present study are from Ailao Mt, except for HKAS 150851 from Gaoligong Mts. Some specimens of Feng et al. (2020) are from Jinggu, Lincang and Dali, surrounding areas of Ailao Mts.

Different from R. griseocarnosa var. griseocarnosa as represented by HKAS 51839a and HKAS 51713a and 51713b (clade Ⅷ in Fig. 2), this variety has white or less gray context in the pileus. Among the lineages of R. griseocarnosa, this variety and the other samples in clade Ⅱ, clades Ⅵ and Ⅶ have bigger spores. The taste of context is often mild, but we did find specimen with acrid taste (HKAS 104735). Hyphae in the subpellis are often more inflated than those of R. occulta and R. griseocarnosa var. griseocarnosa.

Russula occulta X.H. Wang, Pei Zhang & J. Wang, sp. nov. (隐匿红菇) Fig. 4

MycoBank: MB 861589

Holotype: China. Yunnan Province, Jingdong County, Yinsheng market, 15 Aug. 2017, Jing Wang 192 (HKAS 104858).

Etymology: occulta = hidden, naming a species that has been hidden in its sibling species R. griseocarnosa.

Diagnosis: A species characterized by thick- fleshed basidiocarps, smooth dark red pileus hardly with striates, white context in the pileus, relatively small spores (6.5-8 × 5.5-7 μm) with blunt warts and short ridges partly connected and pileipellis an (ixo)trichoderm. Differing from R. griseocarnosa in the whitish context in pileus and smaller spores with more coalescent ornamentation.

Basidiocarps medium-sized to big, thick- fleshed, fragile. Pileus 40-120 mm diam., broadly depressed in the center, margin applanate, not or with very short striates; surface slightly greasy when wet, soon dry, smooth, locally cracked when mature, particularly so closer to the margin, margin sometimes rugose, cuticle peeling 10% to 50% towards the center, greyish red (10B4), red (10B5), brownish red (10D7) to violet brown (11F7) in the center, sometimes with cream-ochre spots, paler (11B7) towards margin; context 3-7 mm thick, white. Lamellae 3−9 mm broad, equal, adnate, sub-crowded to crowded, 6-11 L/cm, brittle, not or exceptionally forking away from stipe, cream-coloured, warm ochre when mature. Stipe 60-80 mm × 20-25 mm, cylindrical or slightly narrowing downwards, neither extremely fragile nor very hard, white (10A1) or pink (11A3, 11A4), sometimes with grayish tinge, spongy inside, white or pale gray (paler than 10B1). Taste mild or slightly acrid, rarely strongly acrid.

Spores (260/13/7)(6-)6.5-7.2-8(-8.5) × (5-)5.5-6.2-7(-7.5) μm, Q= (1.03-)1.11-1.16- 1.20(-1.23), subglobose, broadly ellipsoid, rarely globose; ornamentation 0.3-0.7(-1) μm high, composed of blunt warts and short ridges fused or connected by fine lines, rarely forming a broken reticulum, suprahilar spot totally amyloid. Basidia (27-)30-55 × 8-13(-16) μm, clavate, 4-spored. Hymenophoral gloeocystidia scattered to common, 56-112 × 7-17 μm, originating deeply from the subhymenium or trama, emergent, projecting 10-35(-74) μm beyond the basidia layer, fusiform, cylindrical-clavate, rarely thick-walled (wall 0.5-1 μm thick), with dense pale yellow granular, crystalline or flocculent contents in the middle and upper parts. Lamellar edge fertile or sterile; cheilocystidia rare to common, 23-97 × 8-13 μm, subfusiform, cylindrical, rarely thick-walled (wall up to 1 µm thick), with flocculent or granular contents. Pileipellis orthochromatic in cresyl blue, an (ixo)trichoderm, 100-180 μm thick, sometimes with a mucous layer 30-50 µm thick, hyphae compactly ascending, locally with long repent hyphae, with disperse rose pink pigmentation; hyphae in subpellis 3-5 μm wide, in suprapellis 3-7 µm wide, terminal cells 30-45 × 3-7 μm, cylindrical, subfusiform, surface rarely with zebroid ornamentation; pileocystidia absent. Stipitipellis a cutis, 65-165 μm, hyphae 2-5 μm wide; cystidia absent. Rosettes abundant in gill, pileus and stipe trama. Clamp connections absent.

Ecology: habitat not fully known, growing in Quercus forests in subtropical mid-montane areas.

Distribution: southeast and southwest China (Fujian and Yunnan).

Additional materials examined: China. Fujian Province, Sanming, Dec. 2024, collected from local markets by Y.J. Ma, no. ZP01 (HKAS 149414), ZP02 (HKAS 149415), ZP03 (HKAS 149416); Nanping City, Wuyi Mt., collected from local market by J.M. Miao, no. MJM04 (HKAS 149419); Yunnan Province, Dali City, Cangshan, west slope, Xueshanhe reserve, 12 Aug. 2020, J. He, Z.Q. Zhang & J. Gao, no. sj136 (HKAS 128490); Jingdong County, Yinsheng market, 15 Aug. 2017, J. Wang 194 (HKAS 110373); Nanhua County, wild mushroom market, 17 Aug. 2017, X.H. Wang, B. Buyck & J. Wang, no. XHW4656 (HKAS 104854), XHW4657 (HKAS 104853), XHW4662 (HKAS 104852), XHW4665 (HKAS 104851); Simao, Nanping Township, Manxie Village, 7 Sept. 2013, Bo Xiao, no. CXH11 (HKAS 86736); Yunlong County, Caojian Township, market, X.H. Wang 7887 (HKAS 117515); Zhenyuan County, Heping Township, market, X.H. Wang, B. Buyck & J. Wang, no. XHW4492 (HKAS 104866).

Note: This is the real sibling of R. griseocarnosa. It is not as common as R. griseocarnosa and seems to prefer less tropical areas. In the field and markets, it can be separated from typical R. griseocarnosa by the white pileus context. Microscopically it can be easily separated from R. griseocarnosa by the much smaller spores with more coalescent ornamentation. Taste of flesh varies among individuals, being mild (HKAS 104852 and HKAS 104854), weakly acrid (HKAS 105851), acrid but not very strong (HKAS 104853), to strongly acrid (HKAS 104858). The acrid taste and the white context suggest R. chichuensis W.F. Chiu, described by Chiu (1945) from central Yunnan. Russula chichuensis has paler pileus and bigger spores. Russula griseocarnosa var. ailaoshanensis has nearly white pileus context, but it has much bigger spores and more inflated hyphae in the pileipellis.

Russula purpureozonata K. Das, A. Ghosh & Buyck, European Journal of Taxonomy 782: 164 (2021) (紫环红菇，新拟) Fig. 5F

Basidiocarps big, stout, thick-fleshed, compact. Pileus 65-135 mm diam., convex with a broadly depressed center; surface smooth, subgreasy to greasy, first nearly black, cuticle cracked into patchy scales, these forming concentrical zones, center reddish gray (12C2), pale yellowish brown or olivaceous brown with purplish tinge towards margin, locally paler or with orange tinge, cuticle peeling around 1/5-2/3 towards the center, margin not striate; context 5-7 mm thick, smoky gray to grayish black, then ±discoloring reddish brown (similar to the discoloration in Russula sect. Nigricantinae Bataille). Lamellae 5−11 mm broad, adnate, sub-crowded (5-7 L/cm), equal, ±brittle, slightly buttery when touched, edge even or toothed, anastomosing towards context, forking near the stipe apex, yellowish white (2A2, 3A2) to pale yellow (2A3, 3A3), sometimes with grayish tinge, then turning pale brownish, with rose red edge near the pileus margin. Stipe 50-90 mm × 16-25 mm, expanding downwards, hard, spongy inside; surface smooth, yellowish white with pinkish flush or nearly purplish red; context smoky gray. Taste mild.

Spores (40/2/2) (7.5-)8-8.4-9(-9.5) × 6.5- 7.3-8 μm, Q= 1.09-1.14-1.20(-1.23), broadly ellipsoid; ornamentation 0.1-0.5 μm high, composed of blunt warts and short irregular ridges, those mostly isolated, rarely fused; suprahilar spot totally amyloid. Basidia 45-60 × 11-14 μm, clavate, 4-spored, some with dark brownish contents. Hymenophoral gloeocystidia abundant, emergent, 68-100 × 9-13 μm, projecting 10-30 μm beyond the basidia, base originating deeply from subhymenium or trama, slightly thick-walled, fusiform with a long narrow base and wide and blunt upper part, tips rarely constricted, with needle-like contents. Lamellar edge sterile, marginal cells 20-35 × 4-7 μm; cheilocystidia common, 55-70 × 8-16 μm, cylindrical, subfusiform, with granular or crystalline contents. Pileipellis orthochromatic in cresyl blue, a thick subtrichoderm, 170-260 μm thick, hyphae compactly packed and interwoven, those in suprapellis mostly ascending, in subpellis more horizontally oriented, branching, 3-5 μm wide; pileocystidia common, distributed at different depths, cylindrical, 60-140 × 5-8 µm, 2-3 celled, with pale yellowish brown granular or crystalline contents. Stipitipellis a cutis, 70-100 μm; hyphae compactly packed, 2.5-4 μm wide, hyaline, hyphal ends projecting, clavate, 4-7 µm wide; caulocystidia common, long, 4-7 µm wide. Rosettes abundant in gill, pileus and stipe trama, locally with dark brown contents. Clamp connections absent.

Ecology: scattered in Abies forests, subalpine.

Distribution: China (northwestern Yunnan) and India (Sikkim).

Specimens examined: China, Yunnan Province, Deqin County, Benzilan Township, Baima Snow Mts., road from Benzilan to Deqin, 28°18ʹ40ʺN 99°07ʹ40ʺE, alt. 3 600 m, 11 Aug. 2018, X.H Wang 5155 (HKAS 111615); Deqin County, Yanmen Township, Yugong Village, west slope of Baima Snow Mts., 28°30ʹ13.83ʺN 98°54ʹ47.41ʺE, alt. 3 406 m, 23 Aug. 2020, X.H. Wang & Z. Wang, no. XHW8294 (HKAS 117918); Yunling Township, road from Deqin to Gongshan, Kongqueshan mountain pass, 26 Aug. 2018, X.H. Wang 5846 (HKAS 116519); Deqin County, Yunling Township, road from Nanzhong mountain pass to Xidang Village, 28°23ʹ39.49ʺN 98°49ʹ15.17ʺE, alt. 3 712 m, 26 Aug. 2020, X.H. Wang & Z. Wang, no. XHW8634 (HKAS 118247); ibid., 27 Aug. 2020, X.H. Wang & Z. Wang, no. XHW8707 (HKAS 118314).

Note: The morphology of the Chinese specimens cited above meets the original description of this species given by Das et al. (2021), except that the flesh of the Chinese species is initially smoky gray (i.e. not changing to gray) and the spore ornamentation is much lower (0.3-1.2 µm in Indian specimens). This is a typical subalpine species, specifically growing with Abies trees. In the field, it looks like a “dingy olivaceus- purple” R. griseocarnosa. Microscopically the spores with blunt ornamentation and presence of pileocystidia are good characters separating it from R. griseocarnosa.

4 DISCUSSION AND CONCLUSION

收起

4.1 Russula griseocarnosa: a phylogenetic species with high genetic and morphological variations

Although Li et al. (2010) and Feng et al. (2020) defined several lineages in R. griseocarnosa, they did not answer whether these lineages represent real phylogenetic species. By broader sampling, adding two loci (RPB1 and TEF1-α) and conducting GCPSR, the present study concludes that these lineages are not real phylogenetic species. There exists disconcordance among the ITS, RPB1 and RPB2 genealogies. On the context of current sampling, if we take each of the evolutionary lineages as different species, the samples of clade Ⅵ and Ⅶ will be left as unclassified, since these clades do not meet the criterion of being an evolutionary lineage and thus independent species. We recognized only two phylogenetic species: R. griseocarnosa and R. occulta.

The lineages of R. griseocarnosa vary considerably in size and geographical range. By combining all the ITS sequences of Li et al. (2010), Feng et al. (2020), Yu et al. (2020) with our own data, we could compare the genetic variations among the lineages based on ITS sequences. CladeⅠhas the biggest within-group genetic distance. It was found from 11 counties of five province-level administrative regions (Fujian, Guangdong, Guangxi, Hunan, Yunnan). Clade Ⅶ is the most common lineage, with 205 documented samples from 15 counties of four province-level administrative regions (Fujian, Guangdong, Guangxi, Yunnan). In this clade, 150 specimens belong to R. griseocarnosa var. ailaoshanensis and they are only found in subtropical Yunnan, mainly from Ailao Mts. The three exclusive lineages discovered in this study (Ⅲ, Ⅳ and Ⅴ) have smallest population size and genetic variation. The lowest within-species ITS similarity is 92% after all these clades were lumped in R. griseocarnosa. We have to admit that such cutoff is too tolerant and even much lower than the popularly acknowledged cutoff of OTU in environmental studies. Russula griseocarnosa may eventually fall into several species if broader sampling and/or more DNA loci, e.g. genome data are used. For the time being, this taxonomic treatment does not seem to be unacceptable, as at least the trade of ‘dahongjun' will benefit more from recognizing only one species than splitting into many small species.

Russula griseocarnosa as defined in this study holds high morphological variations. The two specimens of clade Ⅲ (HKAS 104790 and HKAS 104808) have pileus with mixed dingy ochraceous and dull reddish color (Fig. 5B). Spores of this clade are smallest among all the lineages (av. 7.8 × 6.8 µm). This lineage merits an infraspecific taxon if the variations are confirmed by more specimens. The Indian “Russula griseocarnosa” with a less reddish and more ochraceous pileus documented by Das et al. (2010) might belong here, except that their spores are slightly bigger (av. 8.5 × 7.3 µm). Spore of clades Ⅱ, Ⅵ, and Ⅶ are much bigger (av. 9.3-9.7 × 8-8.2 µm), whereas those of clades Ⅰ, Ⅳ, Ⅴ, and Ⅷ are intermediate (av. 8.3-8.7 × 7.2-7.5 µm). All the clades have greyish context, except that the populations of Ailao and Gaoligong Mts. of cladeⅡhave white or nearly white context. Microscopically they all have spores ornamented with isolated warts and lack pileocystidia in the pileipellis. If R. griseocarnosa is split into more species, some of them would be cryptic. Given that the geographical ranges of these lineages strongly overlap, we speculated a prevalent sympatric speciation scenario.

4.2 ‘dahongjun': an assemblage of lookalikes consisting of several phylogenetically distinct species

After R. griseocarnosa was described to replace the name R. vinosa for Chinese specimens (Wang et al. 2009), for a long time it was nearly the only available scientific name for marketed ‘dahongjun' in China. This study discovered a high species diversity under ‘dahongjun'. These species are placed in five sections/subsections (Fig. 1). The most acknowledged and prized species is still R. griseocarnosa. It is widely distributed in subtropical-tropical areas, including Fujian, Guangdong, Hunan, Guangxi, Yunnan and Sichuan (voucher specimen SAAS 3706). The less common species is R. occulta, living in untypically tropical places (Ailao Mts. and northwards and Fujian). This species has whitish context and much smaller spores (Fig. 3). In the subalpine coniferous forests of Yunnan, R. purpureozonata, a species originally described from subalpine Himalayan India (Ghosh et al. 2021), is firstly discovered. This thick-fleshed species has dulled purple pileus and typically greyish context (Fig. 5F). In the field it is strongly reminiscent of R. griseocarnosa. However, multi-gene phylogenetic analysis showed that it was not sister to R. griseocarnosa, but to an undescribed species (HKAS 110297) which was also sold in Ailao Mts. under ‘dahongjun'. These two species formed a clade sister to European R. cremeoavellanea. We have not seen R. purpureozonata in local market, but it has potential commercial value. In a very small local market near Kunming, fresh fruiting bodies of R. quercina, a species originally described from Heilongjiang, northeast China (Zhou et al. 2022), were occasionally seen (Fig. 5H). Apparently, they were collected from the forest nearby and taken as edible mushrooms by the local people. It has less dark pileus, white flesh, yellower lamellae and less stout basidiocarps. The five species above all belong to R. subsect. Integrae or to the “Decolorantes” group.

The other four species regarded as ‘dahongjun' in Yunnan markets are dispersed in three section/subsections. In sect. Flavisiccantes, R. aff. lepida (Figs. 1 and 5D) formed a sister of European R. lepida. This species is very common in subtropical-tropical markets in Yunnan. In Yunnan market, there is another less common species of this section (R. cf. lepida in Figs. 1 and 5E, represented by HKAS 104857 and HKAS 104869). In GenBank an ITS sequence from China (PP930946) and two from India (OQ653085 and PP582965), which are misidentified as R. shigatseensis S.Hui Wang et al. belong to this species. We did not find reliable difference among this species, R. aff. lepida and European R. lepida and here left it as a cryptic species. Russula shigatseensis has much smaller spores (Wang et al. 2024). Microscopically the more or less reticulate spores and the presence of pilocystidia of the species above can help to separate them from R. griseocarnosa and R. occulta.

Russula yanheensis, a species originally described from Guizhou (Tibpromma et al. 2017), was seen in the markets of Nanhua and Jianchuan, Yunnan. It was placed in R. subsect. Paludosinae in our phylogeny. This species has white context and quite stout basidiocarps, hardly separated from R. occulta based simply on macromorphology. In the original description, the spores of R. yanheensis were described as having “moderately large and distant amyloid warts”. The spore photos were not taken in Melzer's reagent, therefore it is not possible to see the details of ornamentation. Our two specimens have spores with very fine and low ornamentation (0.1-0.3 µm high), composed of ridges and warts partly connected into a broken reticulum. Wang et al. (2022) reported “Russula vinosa” from Pu'er market in tropical Yunnan. Their voucher specimen (HKAS 122380) is actually R. yanheensis. “Russula paludosa” reported by Zhang et al. (2021, ITS GenBank no. MW874552) from Yunnan is also R. yanheensis. Further analysis of ITS sequences shows that R. yanheensis has wide geographical distribution. We collected this species in Quercus forest from Bomi, Xizang (HKAS 116446; ITS GenBank no. PV974750). The Japanese sample OMS-113 (ITS GenBank no. LC667117) reported by Shirakawa et al. (2022) and UDB0777983 from Laos and UDB013215 from Papua New Guinea in the UNITE database also belong to this species.

The last known species, R. dhakuriana, was originally described from India (Das et al. 2006). It belongs to R. subsect. Roseinae. We did not see it in markets, but local people in Gaoligong Mts. picked and ate it. When fresh, the white flesh, vivid red pileus and the relatively slender fruiting bodies are helpful to recognition (Fig. 5C). Microscopically the subpellis of pileus with regular layers of globose cells and the short erect hyphae in the suprapellis, together with the medium-sized spores ornamented with isolated warts are characteristic. Russula dhakuriana reported by Zhao & Ji (2021) does not fit this species. The photos represent a species of R. subgen. Archaeae, most likely R. butyroindica K. Das & Buyck. Their spores (5.2-6.8 × 3.2-4.8 µm) are too small for R. dhakuriana. This species is common in Ailao Mts. and also detected from environmental samples in central Japan (GenBank accessions LC096755 and LC096756, Uesugi et al. 2016).

To separate the species of ‘dahongjun', a key is provided here:

1. Context of pileus grayish or grayish black 2

1. Context of pileus white to cream-colored, at most pale gray 6

2. Pileus predominantly red to dark red 3

2. Pileus dark purple or dingy ochraceous or olivaceous with reddish tinge 5

3. Pileocystidia present; spores av. 8.1 × 7.1 µm, ornamentation 0.2-0.5 µm high, of low and blunt warts partly connected; subtropical Yunnan Russula sp.1 (HKAS 110297)

3. Pileocystidia absent; ornamentation 1-1.5(2) µm high, composed of high and conical isolated warts 4

4. Spores av. 9.3-9.5 × 8-8.1 µm; Fujian, Guangdong, Guangxi, Sichuan, Yunnan R. griseocarnosa clades Ⅱ, Ⅵ and Ⅶ

4. Spores av. 8.3-8.7 × 7.2-7.5 µm; Fujian, Guangdong, Guangxi, Hunan, Yunnan R. griseocarnosa cladesⅠ, Ⅳ,Ⅴand Ⅷ

5. Pileus dingy ochraceous with dull reddish tinge at centre; spores 7-8 × 6-7 µm (av. 7.8 × 6.8 µm); pileocystidia absent; in subtropical mid-montane broad-leaved forest in Yunnan R. griseocarnosa clade Ⅲ

5. Pileus dingy olivaceous brown, often with purplish or ochraceous; spores 8-9 × 6.5-8 µm (av. 8.3 × 7.3 µm); pileocystidia present; in subalpine coniferous forest in Yunnan R. purpureozonata

6. Pileocystidia present 7

6. Pileocystidia absent 10

7. Lamellae pale yellow; spore ornamentation composed of isolated warts rarely fused; in northeast and mid-montane southwest China R. quercina

7. Lamellae white; spore ornamentation more or less reticulate 8

8. Stipe with a hard cortex, not fragile; pileus rose-red, pruinose; pileipellis an (ixo)trichoderm; spore ornamentation composed of low warts and short ridges, partly connected by fine lines, forming a broken or nearly complete reticulum; Hunan and Yunnan

R. cf. lepida (HKAS 104869), R. aff. lepida (HKAS 104869, HKAS 105857)

8. Stipe soft, fragile; pileus not pruinose, often subgreasy 9

9. Spores ornamentation up to 0.3 µm high, of blunt warts and ridges partly connected forming a very broken reticulum; pleurocystidia 74-120(150) × 10-16 μm, strongly emergent; Guizhou, Xizang and Yunnan R. yanheensis

9. Spores ornamentation up to 0.7 µm high, of acute warts and ridges connected and forming a more complete reticulum; pleurocystidia smaller, 53-93 × 8-11 μm, not strongly emergent; Yunnan Russula sp.2 (HKAS 104855, HKAS 104863)

10. Pileus vivid red, margin with short striates; pileipellis of a typical “virescens-type” structure; spore ornamentation of fine and dense conical warts; subtropical Yunnan R. dhakuriana

10. Pileus dark red, margin without striates; pileipellis a trichoderm, hyphae of subpellis at most inflated 11

11. Spores 6.5-8 × 5.5-7 μm (av. 7.2 × 6.2 µm), ornamentation 0.3-0.8(1) μm high, composed of blunt warts connected by fine lines; pileus dry to subgreasy; subtropical Yunnan R. occulta

11. Spores 8.5-10 × 7.5-8.5 μm (av. 9.3 × 7.9 µm), ornamentation 1-1.5(2) μm, composed of isolated conical warts rarely fused; pileus often greasy; subtropical Yunnan R. griseocarnosa var. ailaoshanensis

Acknowledgments

收起

The earliest plan of this study was made by Dr. CHEN Xinhua (1977-2015), Department of Pharmacy, Guilin Medicinal College. He collected specimens from Guangdong, Guangxi and Yunnan and shared some of the specimens and data with XHW. This paper is dedicated to him in memory of his contribution to the study of ‘dahongjun'. LI Weibiao, HUANG Lianhui, SHI Shufeng and WANG Zhen, former members of XHW's research group participated in lab and field work. Dr. Karen Hansen, Swedish Museum of Natural History arranged the field trip in Sweden. Prof. ZENG Guangyu, Guangxi Academy of Forestry and Mr. LIU Liangjun, Xiamen Evening News Agency helped to collect dry specimens from mushroom hunters and dealers in Fujian, Guangdong and Guangxi. Prof. QIN Weiqiang, Zhangjiajie Campus, Jishou University provided specimen of R. aff. lepida, Mr. HE Jun, Dali University collected specimen of R. occulta and Dr. HE Xiaolan, Sichuan Institute of Edible Fungi arranged the loan of R. griseocarnosa. Amateur from Yunnan, Mr. WANG Peng collected specimens of R. quercina from local market. Prof. LI Taihui and Prof. DENG Wangqiu, Institute of Microbiology, Guangdong Academy of Sciences arranged the field trip in Chebaling National Nature Reserve, Guangdong. Prof. CHEN Zuohong, Hunan Normal University kindly arranged the loan of R. griseocarnosa specimen in MHHNU. Dr. ZENG Chunxia and Dr. ZHANG Zhirong, Germplasm Bank of Wild Species, KIB helped with the genome skimming sequencing and data analysis.

Author contributions

收起

ZHANG Pei: Field work, data analysis, microscopic observation, manuscript writing. LI Yuwu: Field investigation. CAO Shuqin: Field work, data analysis, microscopic observation. WANG Jing: Field work, molecular lab work, microscopic observation. SU Kaimei: Field investigation. LI Shuhong: Field investigation, DNA data producing. BUYCK Bart: Data analysis, manuscript writing. WANG Xianghua: Project design, data analysis and manuscript writing.

Declaration of competing interests

收起

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

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2026年第45卷第2期

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doi: 10.13346/j.mycosystema.250280

接收时间：2025-09-26
首发时间：2026-04-29
出版时间：2026-02-22

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收稿日期：2025-09-26
录用日期：2025-12-10

基金

National Natural Science Foundation of China(32170022)

2023 Biodiversity Monitoring-special Investigation of Wild Fungi Issued by Chuxiong Management Bureau of Yunnan Ailao Mountains National Nature Reserve, the Biodiversity Survey and Assessment Project(2019HJ2096001006)

The “Investigation of Macrofungi of Maguan County” Issued by Ministry of Ecology and Environment of the People's Republic of China

作者信息

1 中国科学院大学生命科学学院，北京 101408

2 中国科学院昆明植物研究所云南省真菌多样性与绿色发展重点实验室，云南昆明 650201

3 哀牢山国家级自然保护区楚雄管护局，云南楚雄 675000

4 贵州科学院贵州省生物研究所，贵州贵阳 550009

5 云南省农业科学院，云南昆明 650221

6 系统进化与生物多样性研究所法国国家自然历史博物馆，巴黎 F-75005，法国

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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Species	Voucher specimen	Geographical origin	GenBank accession number
R. badia	587/BB 07.324 (PC0124709)	Slovakia	PX726806	KU237571	KU237715	KU237857	KU237999
R. carminipes	531/BB 07.192 (PC0124681)	Slovakia	PX726805	KU237523	KU237673	KU237809	KU237953
R. claroflava	WXH10952 (HKAS 146581)	Sweden	PV974715	PV973112	PV999097	PV999168	PV999017
R. cremeoavellanea	WXH10943 (HKAS 149035)	Sweden	PV974716	PV973113	PV999098	PV999169	PV999018
R. decolorans	WXH11050 (HKAS 146587)	Sweden	PV974717	PV973114	PV999099	PV999170	PV999019
R. decolorans	WXH11069 (HKAS 146590)	Sweden	PV974718	PV973115	PV999100	PV999171	PV999020
R. dhakuriana	CXH02 (HKAS 86727)	Shanghang, Fujian	PX622224 PX622225	PX558766	PX753901	PX580453 PX580454	PX694204
R. dhakuriana	WXH4518 (HKAS 104867)	Xinping, Yunnan	PX622226 PX622227	PX558767	PX580446 PX580447	-	PX580449
R. dhakuriana	YZL7713 (HKAS 150857)	Tengchong, Yunnan	PX622230	PX558768	PX694206	PX580455 PX580456	PX580450
R. griseocarnosa cladeⅠ	CXH03 (HKAS 86728)	Dapu, Guangdong	PV974585	PV973066	PV999049	PV999126	PV998967
R. griseocarnosa cladeⅠ	CXH04 (HKAS 86729)	Tengxian, Guangxi	PV974586	PV973067	PV999050	PV999127	PV998968
R. griseocarnosa cladeⅠ	CXH06 (HKAS 86731)	Pubei, Guangxi	PV974587	PV973068	PV999051	PV999128	PV998969
R. griseocarnosa cladeⅠ	CXH07 (HKAS 86732)	Yanling, Hunan	PV974588	PV973069	PV999052	PV999129	PV998970
R. griseocarnosa cladeⅠ	CXH10 (HKAS 86735)	Pubei, Guangxi	PV974589	PV973070	PV999053	PV999130	PV998971
R. griseocarnosa cladeⅠ	RTJ05 (HKAS 149431)	Sanming, Fujian	PV974578 PV974579	PV973065	PV999048	PV999125	PV998966
R. griseocarnosa cladeⅠ	WXH2125b (HKAS 51839b) holotype p.p.	Jinghong, Yunnan	PV974584*	PV973140*	PV999118*	PV999191*	PV999040*
R. griseocarnosa cladeⅡ	CXH01 (HKAS 86726)	Shanghang, Fujian	PV974639	PV973077	PV999060	PV999137	PV998980
R. griseocarnosa cladeⅡ	CXH05 (HKAS 86730)	Rongxian, Guangxi	PV974640	PV973078	PV999061	PV999138	PV998981
R. griseocarnosa cladeⅡ	HJW02 (HKAS 149378)	Changting, Fujian	PV974605	PV973073	PV999056	PV999133	PV998975
R. griseocarnosa cladeⅡ	HJW07 (HKAS 149383)	Changting, Fujian	PV974610	PV973074	PV999057	PV999134	PV998976
R. griseocarnosa cladeⅡ	LJM12 (HKAS 149404)	Pucheng, Fujian	PV974599	PV973072	PV999055	PV999132	PV998973 PV998974
R. griseocarnosa cladeⅡ	RTJ02 (HKAS 149428)	Nanping, Fujian	PV974625 PV974626	PV973075	PV999058	PV999135	PV998977 PV998978
R. griseocarnosa cladeⅡ	WXH3163 (HKAS 73659)	Shixing, Guangdong	PV974630	PV973076	PV999059	PV999136	PV998979
R. griseocarnosa cladeⅡ	YYB04 (HKAS 149449)	Gutian, Fujian	PV974641	PV973079	PV999062 PV999063	PV999139	PV998982
R. griseocarnosa cladeⅡ	YYB10 (HKAS 149455)	Gutian, Fujian	PV974642 PV974643	PV973080	PV999064	PV999140	PV998983
R. griseocarnosa cladeⅡ	ZGY02 (HKAS 149353)	Rongxian, Guangxi	PV974591	PV973071	PV999054	PV999131	PV998972
R. griseocarnosa clade Ⅲ	WXH4426 (HKAS 104790)	Maguan, Yunnan	PV974644	PV973089	PV999072	PV999148	PV998991
R. griseocarnosa clade Ⅲ	WXH4506 (HKAS 104808)	Xinping, Yunnan	PV974645	PV973090	PV999073	PV999149	PV998992
R. griseocarnosa clade Ⅳ	CXH12 (HKAS 86737)	Pu'er, Yunnan	PV974647	PV973091	PV999075	PV999150	PV998993
R. griseocarnosa clade Ⅳ	LSH28 (HKAS 149482)	Jingdong, Yunnan	PV974646	PV973141*	PV999074	PV999192*	PV999041
R. griseocarnosa clade Ⅴ	WXH3527 (HKAS 76064)	Lüchun, Yunnan	PV974649	PV973092	PV999076	PV999151	PV998994
R. griseocarnosa clade Ⅴ	ZP-2152 (MHHNU 8652)	Sangzhi, Hunan	PV974648*	PV973142*	PV999120*	PV999193*	PV999043*
R. griseocarnosa clade Ⅵ	LYC4252 (HKAS 149392)	Pingbian, Yunnan	PV974652	PV973094	PV999078	PV999153	PV998996
R. griseocarnosa clade Ⅵ	WXH2125c (HKAS 51839c) holotype p.p.	Jinghong, Yunnan	PV974650*	PV973143*	PV999119*	PV999194*	PV999042*
R. griseocarnosa clade Ⅵ	YYB07 (HKAS 149452)	Gutian, Fujian	PV974651	PV973093	PV999077	PV999152	PV998995
R. griseocarnosa clade Ⅵ	YZL6649 (HKAS 126726)	Ruili, Yunnan	PV974653	PV973095	PV999079	PV999154	PV998997
R. griseocarnosa clade Ⅶ	CXH09 (HKAS 86734)	Jiaoling, Guangdong	PV974657	PV973100	PV999085	PV999158	PV999003
R. griseocarnosa clade Ⅶ	ZGY08 (HKAS 149357)	Maoming, Guangdong	PV974654	PV973096 PV973097	PV999080	PV999155	PV998998 PV998999
R. griseocarnosa clade Ⅶ	ZGY09 (HKAS 149358)	Maoming, Guangdong	PV974655	PV973098	PV999081 PV999082	PV999156	PV999000 PV999001
R. griseocarnosa clade Ⅶ	ZGY19 (HKAS 149364)	Bobai, Guangxi	PV974656	PV973099	PV999083 PV999084	PV999157	PV999002
R. griseocarnosa clade Ⅷ	MJM01 (HKAS 149417)	Nanping, Fujian	PV974677 PV974678	PV973101 PV973102	PV999086	PV999159	PV999004
R. griseocarnosa clade Ⅷ	MJM08 (HKAS 149423)	Nanping, Fujian	PV974682	PV973103	PV999087	PV999160	PV999005 PV999006
R. griseocarnosa clade Ⅷ	RTJ04 (HKAS 149430)	Nanping, Fujian	PV974687 PV974688	PV973104	PV999088	PV999161	PV999007
R. griseocarnosa clade Ⅷ	WXH1989a (HKAS 51713) paratype	Jinghong, Yunnan	PV974692*	PV973144*	PV999121*	PV999195*	PV999044*
R. griseocarnosa clade Ⅷ	WXH1989b (HKAS 51713) paratype	Jinghong, Yunnan	PV974693*	PV973145*	PV999122*	PV999196*	PV999045*
R. griseocarnosa clade Ⅷ	WXH2125a (HKAS 51839a) lectotype	Jinghong, Yunnan	PV974694*	PV973146*	PV999123*	PV999197*	PV999046*
R. griseocarnosa clade Ⅷ	WXH3509 (HKAS 76051)	Pu'er, Yunnan	PV974695	OP794377	OP805334	OP804711	PV999008
R. griseocarnosa clade Ⅷ	YYB02 (HKAS 149447)	Gutian, Fujian	PV974698 PV974699	PV973105	PV999089	PV999162	PV999009 PV999010
R. griseocarnosa var. ailaoshanensis	LSH43 (HKAS 149483)	Jingdong, Yunnan	PV974624	PV973086	PV999069	PV999145	PV998988
R. griseocarnosa var. ailaoshanensis	SKM2 (HKAS 149437) holotype	Jingdong, Yunnan	PV974615	PV973081	PV999065	PV999141	PV998984
R. griseocarnosa var. ailaoshanensis	SKM5 (HKAS 149439)	Jingdong, Yunnan	PV974618	PV973082	PV999066	PV999142	PV998985
R. griseocarnosa var. ailaoshanensis	SKM6 (HKAS 149440)	Jingdong, Yunnan	PV974619	PV973083	PV999067	PV999143	PV998986
R. griseocarnosa var. ailaoshanensis	SKM7 (HKAS 149441)	Jingdong, Yunnan	PV974620	PV973084	PV999068	PV999144	PV998987
R. griseocarnosa var. ailaoshanensis	WXH4646 (HKAS 109819)	Nanhua, Yunnan	PV974636	PV973087	PV999070	PV999146	PV998989
R. griseocarnosa var. ailaoshanensis	WXH4649 (HKAS 104735)	Nanhua, Yunnan	PV974638	PV973088	PV999071	PV999147	PV998990
R. griseocarnosa var. ailaoshanensis	YZL7597 (HKAS 150851)	Baoshan, Yunnan	PX622248	PX558769	PX580448	PX580457	PX580451
R. integra	518/BB 07.198 (PC0124682)	Slovakia	PX726804	KU237513	KU237663	KU237799	KU237943
R. laeta	519/BB 07.267 (PC0124692)	Slovakia	PX726803	KU237514	KU237664	KU237800	KU237944
R. laricina	575/BB 08.681		JN944008	KU237560	KU237704	KU237846	KU237991
R. lepida	437/BB 07.189 (PC0124680)	Slovakia	PX726807 PX726808	KU237500	KU237649	KU237786	KU237930
R. aff. lepida	QWQ-R064 (HKAS 110974)	Zhangjiajie, Hunan	PV974708	PV973107	PV999091	PV999164	PV999012 PV999013
R. aff. lepida	WJ193 (HKAS 104840)	Jingdong, Yunnan	PV974707	PV973106	PV999090	PV999163	PV999011
R. aff. lepida	WXH4489 (HKAS 104801)	Zhenyuan, Yunnan	PV974710 PV974711	PV973108 PV973109	PV999092	PV999165	PV999014
R. cf. lepida	WJ211 (HKAS 104857)	Jingdong, Yunnan	PV974713	PV973110	PV999093 PV999094	PV999166	PV999015
R. cf. lepida	WXH4488 (HKAS 104869)	Zhenyuan, Yunnan	PV974714	PV973111	PV999095 PV999096	PV999167	PV999016
R. occulta	WJ192 (HKAS 104858) holotype	Jingdong, Yunnan	PV974729	PV973122 PV973123	PV999107	PV999178 PV999179	PV999027
R. occulta	CXH11 (HKAS 86736)	Pu'er, Yunnan	PV974728	PV973121	PV999106	PV999177	PV999026
R. occulta	WJ194 (HKAS 110373)	Jingdong, Yunnan	PV974730	PV973124	PV999108	PV999180	PV999028
R. occulta	WXH4492 (HKAS 104866)	Zhenyuan, Yunnan	PV974735	PV973126	PV999110	PV999183	PV999029
R. occulta	WXH4656 (HKAS 104854)	Nanhua, Yunnan	PV974737 PV974738	PV973127	PV999111	PV999184	PV999030
R. occulta	WXH4657 (HKAS 104853)	Nanhua, Yunnan	PV974739	PV973128	PV999112	PV999185	PV999031
R. occulta	WXH4662 (HKAS 104852)	Nanhua, Yunnan	PV974741	PV973130	PV999113	PV999186	PV999034
R. occulta	WXH4665 (HKAS 104851)	Nanhua, Yunnan	PV974742	PV973131	PV999114	PV999187	PV999035
R. occulta	ZP01 (HKAS 149414)	Sanming, Fujian	PV974743	PV973132	PV999115	PV999188	PV999036
R. paludosa	442/BB 07.330 (PC0142538)	Slovakia	PX754887	KU237505	KU237654	KU237791	KU237935
R. purpureozonata	WXH5155 (HKAS 111615)	Deqin, Yunnan	PV974723	PV973118	PV999103	PV999174	PV999023
R. purpureozonata	WXH8294 (HKAS 117918)	Deqin, Yunnan	PV974720	PV973116	PV999101	PV999172	PV999021
R. purpureozonata	WXH8707 (HKAS 118314)	Deqin, Yunnan	PV974722	PV973117	PV999102	PV999173	PV999022
R. quercina	WP-1 (HKAS 149469)	Kunming, Yunnan	PV974725	PV973119	PV999104	PV999175	PV999024
R. quercina	WP-3 (HKAS 149471)	Kunming, Yunnan	PV974727	PV973120	PV999105	PV999176	PV999025
R. subdensifolia	552/BB 05.158 (PC0124674)	USA	PX754886	KU237544	-	KU237830	KU237974
R. vinosa	WXH11053 (HKAS 146588)	Sweden	PV974746	PV973133	-	PV999189	PV999037
R. yanheensis	WXH4661 (HKAS 104879)	Nanhua, Yunnan	PV974748 PV974748	PV973136	PV999117	PV999190	PV999039
R. yanheensis	WXH7157 (HKAS 120113)	Jianchuan, Yunnan	PV974747	PV973134 PV973135	PV999116	-	PV999038
Russula sp.1	WXH4493 (HKAS 110297)	Zhenyuan, Yunnan	PV974736	PV973147*	PV999124*	PV999198*	PV999047*
Russula sp.2	WXH4658 (HKAS 104855)	Nanhua, Yunnan	PV974740	PV973129	PX694205	PV999182	PV999032
Russula sp.2	WJ50 (HKAS 104863)	Maguan, Yunnan	PV974731	PV973125	PV999109	PV999181	PV999033

Species

Voucher
specimen

Geographical
origin

GenBank accession number

ITS

LSU

RPB1

RPB2

TEF1-α

R. badia

587/BB 07.324
(PC0124709)

Slovakia

PX726806

KU237571

KU237715

KU237857

KU237999

R. carminipes

531/BB 07.192
(PC0124681)

Slovakia

PX726805

KU237523

KU237673

KU237809

KU237953

R. claroflava

WXH10952
(HKAS 146581)

Sweden

PV974715

PV973112

PV999097

PV999168

PV999017

R. cremeoavellanea

WXH10943
(HKAS 149035)

Sweden

PV974716

PV973113

PV999098

PV999169

PV999018

R. decolorans

WXH11050
(HKAS 146587)

Sweden

PV974717

PV973114

PV999099

PV999170

PV999019

R. decolorans

WXH11069
(HKAS 146590)

Sweden

PV974718

PV973115

PV999100

PV999171

PV999020

R. dhakuriana

CXH02
(HKAS 86727)

Shanghang,
Fujian

PX622224
PX622225

PX558766

PX753901

PX580453
PX580454

PX694204

R. dhakuriana

WXH4518
(HKAS 104867)

Xinping,
Yunnan

PX622226
PX622227

PX558767

PX580446
PX580447

PX580449

R. dhakuriana

YZL7713
(HKAS 150857)

Tengchong,
Yunnan

PX622230

PX558768

PX694206

PX580455
PX580456

PX580450

R. griseocarnosa
cladeⅠ

CXH03
(HKAS 86728)

Dapu,
Guangdong

PV974585

PV973066

PV999049

PV999126

PV998967

R. griseocarnosa
cladeⅠ

CXH04
(HKAS 86729)

Tengxian,
Guangxi

PV974586

PV973067

PV999050

PV999127

PV998968

R. griseocarnosa
cladeⅠ

CXH06
(HKAS 86731)

Pubei,
Guangxi

PV974587

PV973068

PV999051

PV999128

PV998969

R. griseocarnosa
cladeⅠ

CXH07
(HKAS 86732)

Yanling,
Hunan

PV974588

PV973069

PV999052

PV999129

PV998970

R. griseocarnosa
cladeⅠ

CXH10
(HKAS 86735)

Pubei,
Guangxi

PV974589

PV973070

PV999053

PV999130

PV998971

R. griseocarnosa
cladeⅠ

RTJ05
(HKAS 149431)

Sanming,
Fujian

PV974578
PV974579

PV973065

PV999048

PV999125

PV998966

R. griseocarnosa
cladeⅠ

WXH2125b
(HKAS 51839b)
holotype p.p.

Jinghong,
Yunnan

PV974584*

PV973140*

PV999118*

PV999191*

PV999040*

R. griseocarnosa
cladeⅡ

CXH01
(HKAS 86726)

Shanghang,
Fujian

PV974639

PV973077

PV999060

PV999137

PV998980

R. griseocarnosa
cladeⅡ

CXH05
(HKAS 86730)

Rongxian,
Guangxi

PV974640

PV973078

PV999061

PV999138

PV998981

R. griseocarnosa
cladeⅡ

HJW02
(HKAS 149378)

Changting,
Fujian

PV974605

PV973073

PV999056

PV999133

PV998975

R. griseocarnosa
cladeⅡ

HJW07
(HKAS 149383)

Changting,
Fujian

PV974610

PV973074

PV999057

PV999134

PV998976

R. griseocarnosa
cladeⅡ

LJM12
(HKAS 149404)

Pucheng,
Fujian

PV974599

PV973072

PV999055

PV999132

PV998973
PV998974

R. griseocarnosa
cladeⅡ

RTJ02
(HKAS 149428)

Nanping,
Fujian

PV974625
PV974626

PV973075

PV999058

PV999135

PV998977
PV998978

R. griseocarnosa
cladeⅡ

WXH3163
(HKAS 73659)

Shixing,
Guangdong

PV974630

PV973076

PV999059

PV999136

PV998979

R. griseocarnosa
cladeⅡ

YYB04
(HKAS 149449)

Gutian,
Fujian

PV974641

PV973079

PV999062
PV999063

PV999139

PV998982

R. griseocarnosa
cladeⅡ

YYB10
(HKAS 149455)

Gutian,
Fujian

PV974642
PV974643

PV973080

PV999064

PV999140

PV998983

R. griseocarnosa
cladeⅡ

ZGY02
(HKAS 149353)

Rongxian,
Guangxi

PV974591

PV973071

PV999054

PV999131

PV998972

R. griseocarnosa
clade Ⅲ

WXH4426
(HKAS 104790)

Maguan,
Yunnan

PV974644

PV973089

PV999072

PV999148

PV998991

R. griseocarnosa
clade Ⅲ

WXH4506
(HKAS 104808)

Xinping,
Yunnan

PV974645

PV973090

PV999073

PV999149

PV998992

R. griseocarnosa
clade Ⅳ

CXH12
(HKAS 86737)

Pu'er,
Yunnan

PV974647

PV973091

PV999075

PV999150

PV998993

R. griseocarnosa
clade Ⅳ

LSH28
(HKAS 149482)

Jingdong,
Yunnan

PV974646

PV973141*

PV999074

PV999192*

PV999041

R. griseocarnosa
clade Ⅴ

WXH3527
(HKAS 76064)

Lüchun,
Yunnan

PV974649

PV973092

PV999076

PV999151

PV998994

R. griseocarnosa
clade Ⅴ

ZP-2152
(MHHNU 8652)

Sangzhi,
Hunan

PV974648*

PV973142*

PV999120*

PV999193*

PV999043*

R. griseocarnosa
clade Ⅵ

LYC4252
(HKAS 149392)

Pingbian,
Yunnan

PV974652

PV973094

PV999078

PV999153

PV998996

R. griseocarnosa
clade Ⅵ

WXH2125c (HKAS
51839c) holotype p.p.

Jinghong,
Yunnan

PV974650*

PV973143*

PV999119*

PV999194*

PV999042*

R. griseocarnosa
clade Ⅵ

YYB07
(HKAS 149452)

Gutian,
Fujian

PV974651

PV973093

PV999077

PV999152

PV998995

R. griseocarnosa
clade Ⅵ

YZL6649
(HKAS 126726)

Ruili,
Yunnan

PV974653

PV973095

PV999079

PV999154

PV998997

R. griseocarnosa
clade Ⅶ

CXH09
(HKAS 86734)

Jiaoling,
Guangdong

PV974657

PV973100

PV999085

PV999158

PV999003

R. griseocarnosa
clade Ⅶ

ZGY08
(HKAS 149357)

Maoming,
Guangdong

PV974654

PV973096
PV973097

PV999080

PV999155

PV998998
PV998999

R. griseocarnosa
clade Ⅶ

ZGY09
(HKAS 149358)

Maoming,
Guangdong

PV974655

PV973098

PV999081
PV999082

PV999156

PV999000
PV999001

R. griseocarnosa
clade Ⅶ

ZGY19
(HKAS 149364)

Bobai,
Guangxi

PV974656

PV973099

PV999083
PV999084

PV999157

PV999002

R. griseocarnosa
clade Ⅷ

MJM01
(HKAS 149417)

Nanping,
Fujian

PV974677
PV974678

PV973101
PV973102

PV999086

PV999159

PV999004

R. griseocarnosa
clade Ⅷ

MJM08
(HKAS 149423)

Nanping,
Fujian

PV974682

PV973103

PV999087

PV999160

PV999005
PV999006

R. griseocarnosa
clade Ⅷ

RTJ04
(HKAS 149430)

Nanping,
Fujian

PV974687
PV974688

PV973104

PV999088

PV999161

PV999007

R. griseocarnosa
clade Ⅷ

WXH1989a
(HKAS 51713)
paratype

Jinghong,
Yunnan

PV974692*

PV973144*

PV999121*

PV999195*

PV999044*

R. griseocarnosa
clade Ⅷ

WXH1989b
(HKAS 51713)
paratype

Jinghong,
Yunnan

PV974693*

PV973145*

PV999122*

PV999196*

PV999045*

R. griseocarnosa
clade Ⅷ

WXH2125a
(HKAS 51839a)
lectotype

Jinghong,
Yunnan

PV974694*

PV973146*

PV999123*

PV999197*

PV999046*

R. griseocarnosa
clade Ⅷ

WXH3509
(HKAS 76051)

Pu'er,
Yunnan

PV974695

OP794377

OP805334

OP804711

PV999008

R. griseocarnosa
clade Ⅷ

YYB02
(HKAS 149447)

Gutian,
Fujian

PV974698
PV974699

PV973105

PV999089

PV999162

PV999009
PV999010

R. griseocarnosa
var. ailaoshanensis

LSH43
(HKAS 149483)

Jingdong,
Yunnan

PV974624

PV973086

PV999069

PV999145

PV998988

R. griseocarnosa
var. ailaoshanensis

SKM2 (HKAS
149437) holotype

Jingdong,
Yunnan

PV974615

PV973081

PV999065

PV999141

PV998984

R. griseocarnosa
var. ailaoshanensis

SKM5
(HKAS 149439)

Jingdong,
Yunnan

PV974618

PV973082

PV999066

PV999142

PV998985

R. griseocarnosa
var. ailaoshanensis

SKM6
(HKAS 149440)

Jingdong,
Yunnan

PV974619

PV973083

PV999067

PV999143

PV998986

R. griseocarnosa
var. ailaoshanensis

SKM7
(HKAS 149441)

Jingdong,
Yunnan

PV974620

PV973084

PV999068

PV999144

PV998987

R. griseocarnosa
var. ailaoshanensis

WXH4646
(HKAS 109819)

Nanhua,
Yunnan

PV974636

PV973087

PV999070

PV999146

PV998989

R. griseocarnosa
var. ailaoshanensis

WXH4649
(HKAS 104735)

Nanhua,
Yunnan

PV974638

PV973088

PV999071

PV999147

PV998990

R. griseocarnosa
var. ailaoshanensis

YZL7597
(HKAS 150851)

Baoshan,
Yunnan

PX622248

PX558769

PX580448

PX580457

PX580451

R. integra

518/BB 07.198
(PC0124682)

Slovakia

PX726804

KU237513

KU237663

KU237799

KU237943

R. laeta

519/BB 07.267
(PC0124692)

Slovakia

PX726803

KU237514

KU237664

KU237800

KU237944

R. laricina

575/BB 08.681

JN944008

KU237560

KU237704

KU237846

KU237991

R. lepida

437/BB 07.189
(PC0124680)

Slovakia

PX726807
PX726808

KU237500

KU237649

KU237786

KU237930

R. aff. lepida

QWQ-R064
(HKAS 110974)

Zhangjiajie,
Hunan

PV974708

PV973107

PV999091

PV999164

PV999012
PV999013

R. aff. lepida

WJ193
(HKAS 104840)

Jingdong,
Yunnan

PV974707

PV973106

PV999090

PV999163

PV999011

R. aff. lepida

WXH4489
(HKAS 104801)

Zhenyuan,
Yunnan

PV974710
PV974711

PV973108
PV973109

PV999092

PV999165

PV999014

R. cf. lepida

WJ211
(HKAS 104857)

Jingdong,
Yunnan

PV974713

PV973110

PV999093
PV999094

PV999166

PV999015

R. cf. lepida

WXH4488
(HKAS 104869)

Zhenyuan,
Yunnan

PV974714

PV973111

PV999095
PV999096

PV999167

PV999016

R. occulta

WJ192 (HKAS
104858) holotype

Jingdong,
Yunnan

PV974729

PV973122
PV973123

PV999107

PV999178
PV999179

PV999027

R. occulta

CXH11
(HKAS 86736)

Pu'er,
Yunnan

PV974728

PV973121

PV999106

PV999177

PV999026

R. occulta

WJ194
(HKAS 110373)

Jingdong,
Yunnan

PV974730

PV973124

PV999108

PV999180

PV999028

R. occulta

WXH4492
(HKAS 104866)

Zhenyuan,
Yunnan

PV974735

PV973126

PV999110

PV999183

PV999029

R. occulta

WXH4656
(HKAS 104854)

Nanhua,
Yunnan

PV974737
PV974738

PV973127

PV999111

PV999184

PV999030

R. occulta

WXH4657
(HKAS 104853)

Nanhua,
Yunnan

PV974739

PV973128

PV999112

PV999185

PV999031

R. occulta

WXH4662
(HKAS 104852)

Nanhua,
Yunnan

PV974741

PV973130

PV999113

PV999186

PV999034

R. occulta

WXH4665
(HKAS 104851)

Nanhua,
Yunnan

PV974742

PV973131

PV999114

PV999187

PV999035

R. occulta

ZP01
(HKAS 149414)

Sanming,
Fujian

PV974743

PV973132

PV999115

PV999188

PV999036

R. paludosa

442/BB 07.330
(PC0142538)

Slovakia

PX754887

KU237505

KU237654

KU237791

KU237935

R. purpureozonata

WXH5155
(HKAS 111615)

Deqin,
Yunnan

PV974723

PV973118

PV999103

PV999174

PV999023

R. purpureozonata

WXH8294
(HKAS 117918)

Deqin,
Yunnan

PV974720

PV973116

PV999101

PV999172

PV999021

R. purpureozonata

WXH8707
(HKAS 118314)

Deqin,
Yunnan

PV974722

PV973117

PV999102

PV999173

PV999022

R. quercina

WP-1
(HKAS 149469)

Kunming,
Yunnan

PV974725

PV973119

PV999104

PV999175

PV999024

R. quercina

WP-3
(HKAS 149471)

Kunming,
Yunnan

PV974727

PV973120

PV999105

PV999176

PV999025

R. subdensifolia

552/BB 05.158
(PC0124674)

USA

PX754886

KU237544

KU237830

KU237974

R. vinosa

WXH11053
(HKAS 146588)

Sweden

PV974746

PV973133

PV999189

PV999037

R. yanheensis

WXH4661
(HKAS 104879)

Nanhua,
Yunnan

PV974748
PV974748

PV973136

PV999117

PV999190

PV999039

R. yanheensis

WXH7157
(HKAS 120113)

Jianchuan,
Yunnan

PV974747

PV973134
PV973135

PV999116

PV999038

Russula sp.1

WXH4493
(HKAS 110297)

Zhenyuan,
Yunnan

PV974736

PV973147*

PV999124*

PV999198*

PV999047*

Russula sp.2

WXH4658
(HKAS 104855)

Nanhua,
Yunnan

PV974740

PV973129

PX694205

PV999182

PV999032

Russula sp.2

WJ50
(HKAS 104863)

Maguan,
Yunnan

PV974731

PV973125

PV999109

PV999181

PV999033

Clade	ITS	LSU	RPB2	TEF1-α	RPB1	Five-locus combined	Criterion satisfied
Ⅰ	98%	52%	-	55%	94%	100%	a & b
Ⅱ	100%	74%	96%	81%	-	100%	a & b
Ⅲ	96%	99%	100%	-	100%	100%	a & b
Ⅳ	84%	98%	-	-	-	90%	b
Ⅴ	90%	90%	-	-	-	100%	b
Ⅵ	Supported paraphyly	-	63%	-	76%	84%	Neither
Ⅶ	Supported paraphyly	-	96%	-	-	69%	Neither
Ⅵ+Ⅶ	Supported paraphyly	-	86%	23%	93%	98%	Neither
Ⅷ	-	56%	90%	51%	100%	100%	a & b
B	69%	80%	89%	-	100%	100%	a & b

Clade

ITS

LSU

RPB2

TEF1-α

RPB1

Five-locus combined

Criterion satisfied

Ⅰ

98%

52%

55%

94%

100%

a & b

Ⅱ

100%

74%

96%

81%

100%

a & b

Ⅲ

96%

99%

100%

a & b

Ⅳ

84%

98%

90%

Ⅴ

90%

100%

Ⅵ

Supported paraphyly

63%

76%

84%

Neither

Ⅶ

Supported paraphyly

96%

69%

Neither

Ⅵ+Ⅶ

Supported paraphyly

86%

23%

93%

98%

Neither

Ⅷ

56%

90%

51%

100%

a & b

69%

80%

89%

100%

a & b