灰褶鹅膏

Amanita griseofolia Zhu L . Yang , sp . nov . Fig1-7
Amanita ceciliae sensu Zhu L . Yang , Biblioth . Mycol .170:116,1997, p . p .
Pileus 3-7(10.5) cm diametro , convexus vel applanatus , obscure umbonatus vel non - umbonatus , griseo - brun - neus vel brunneo - griseus , non luteus , non flavus , non ochraceus , margine sulcata (0.2-0.4R), non appendiculata , eliuiis volvae coactis , flocculosis vel verrucosis , griseis vel atrogriseis ornatus . Lamellae liberae , griseolae vel griseae , confertae ; lamellulae truncatae . Stipes (6)8-17x0.5-1.5 cm , subcylindricus vel sursum attenuatus , haud bulbosus , albidus , cavus , squamellis griseis , fibrilloseis vel farinoseis , exannulatus , volvatus . Volva grisea vel atrogrisea , verrucolosa . Caro albida . Basidia 4- sporigera . Basidiosporae (9.5)10.0-13.5(16.5) x (8.5)9.5-13.0(15.0) um , globosae vel subglobosae , non amyloideae . Fibulae absentes .
Holotype : China , Yunnan , Kunming , Heilongtan ,1980 m asl ., on soil in forest dominated by Quercus sp ., Pinus armandi , and P . yunnanensis ,27.6.2001, Z . L . Yang 3081( HKAS 38159).
Etymology : griseofolia - referring to the greyish to grey lamellae.
Basidiocarps ( Figs .1,5-7) small to medium - sized , rarely large . Pileus 3-7(10.5) cm diam , at first nearly campanulate to hemispherical , then convex to plano - convex , without an umbo or slightly um - bonate , brownish grey to grey - brown [ Natal Brown , Buffy Brown , Snuff Brown , Medal Bronze ,4B1-2,5C2-4,5D2-4,5E3-5], darker over disc , becoming somewhat paler towards the margin , lacking any yellow or ochreous tint at all stages of development ; margin tuberculate - striate (0.20-0.43R), non - appendiculate ; volval remnants grey to dark grey [ Iron Gray , somewhat darker than Smoke Gray ,4C1-2,4D1-2,4E1-2], verrucous to felty or farinose , sometimes irregularly formed ,0.5-1.5 mm thick , easily washed away by rain ; trama thin ,2-5 mm thick , white to whitish , unchan - ging . Lamellae free , crowded , whitish [ Cream Color , IAI -2J, but soon becoming greyish to grey [ Pale Smoke Gray to Light Drab , paler than 5D2], often becoming somewhat darker when dried ; lamellar edges grey to dark grey [ Iron Gray , darker than Smoke Gray ,4E1-2,5E1-3]; lamellulae truncate to subtruncate , plentiful , evenly distributed . Stipe (6)8-17 x 0.5-1.5 cm , subcylindric or slightly tapering upward , with apex slightly expanded , white to dirty white , lower half covered with grey to greyish [ Pale Smoke Gray ,4D2] fibrillose squamules , upper half densely covered with grey farinose squamules , fistulous ; context white to dirty white , unchanging ; basal bulb lacking ; volval remnants felty to granular or verrucous , grey to dark grey [ Iron Gray , Smoky Gray , Drab ,4E1-2,SE1-3], arranged irregularly or sometimes in incomplete belts or rings at stipe base . Annulus lacking . Smell indistinct .
Lamellar trama ± bilateral. Mediostratum 30–40 μm wide, composed of fairly abundant, long ellipsoid to subfusiform inflated cells (70–90 × 15–20 μm) and abundant interwoven, often anastomosing, 3–7 μm wide filamentous hyphae; vascular hyphae rare. Lateral stratum composed of fairly abundant, long ellipsoid to subfusiform inflated cells (40–60 × 10–15 μm), diverging at an angle of ca. 30° to mediostratum; filamentous hyphae fairly abundant to abundant, 3–7 μm wide, frequently branching, interwoven, sometimes anastomosing; vascular hyphae rare, 3–9 μm wide. Subhymenium (Fig. 2) 30–50 μm thick, with 2–3(4) layers of subglobose to ovoid or shortly ellipsoid cells, 12–25 × 10–20 μm, occasionally mixed with hardly inflated cells. Basidia (Fig. 2) 40–70 × 15–20 μm, clavate, 4-spored; sterigmata 5–7 μm long; basal septa without clamps. Basidiospores (Fig. 3) [456/28/22] (9.5)10.0–13.5(16.5) × (8.5)9.5–13.0(15.0) μm [Q = 1.0–1.10(1.18), Q = 1.04 ± 0.03], globose to subglobose, rarely broadly ellipsoid, inamyloid, colorless, hyaline, thin-walled, smooth; apiculus ca. 1 μm long. Lamellar edge as sterile, gelatinized, yellow-brown strip up to 250 μm wide in side view, predominantly composed of ovoid to subglobose, sometimes sphaeropedunculate inflated cells (20–50 × 15–35 μm), single and terminal or in chains of 2–3, colorless and hyaline, or with yellowish brown vacuolar pigments; filamentous hyphae fairly abundant to scattered, 2–7 μm wide, colorless and hyaline, sometimes with brownish vacuolar pigments, irregularly arranged or running parallel to the lamellar edge, elements embedded in yellow-brown amorphous matrix gradually disappearing in KOH. Pileipellis 50–100 μm thick; upper layer (20–60 μm thick) strongly gelatinized, composed of ± radially and moderately compactly arranged, 1–4 μm wide, thin-walled, colorless or brownish vacuolar pigmented filamentous hyphae; lower layer (30–50 μm thick) composed of radially and compactly arranged, 3–8(15) μm wide filamentous hyphae with brownish yellow, vacuolar pigments; terminal cells present, 5–10 μm wide, hardly inflated; vascular hyphae rare. Volval remnants on pileus (Fig. 4) composed of ± vertically (over disc) to irregularly (on other parts of the pileus) arranged elements. Inflated cells very abundant to dominant, subglobose, ovoid to shortly ellipsoid (20–70 × 15–50 μm), sometimes broadly clavate (40–60 × 20–30 μm) or sphaeropedunculate (50–80 × 25–45 μm), often in chains of 2–3 and then terminal, thin- to slightly thick-walled (up to 0.5 μm thick), often with brownish yellow vacuolar pigments; filamentous hyphae scattered to fairly abundant, 2–7 μm wide, frequently branching , often anastomosing , thin - walled , colorless and hyaline , sometimes with brownish yellow vacuolar pigments ; vascular hyphae rare ,2-8 μm wide . Volval remnants on stipe base similar to those on the pileus , but inflated cells and filamentous hyphae irregularly arranged . Stipe trama composed primarily of longitudinally arranged , long clavate , terminal cells ,250-350x 25-40μm ; filamentous hyphae 2-7μm wide , scattered ( in interior ) or abundant ( on stipe surface ); vascular hyphae rare ,2-10 um wide . Clamps absent in fruitbody .
Specimens examined : CHINA , Beijing , Tanzhesi , on soil ,25.7.1958, S . C . Teng 6063( HMAS 22610); Beijing , Tanzhesi , on soil ,29.7.1959, W . X . Wang and X . L . Kong 74( HMAS 26499); Hainan , Changjiang , Bawangling ,1350 m asl ., on soil in broad - leaved forest ,24.8.1990, Q . Chen ( GDGM 16689); Hainan , Ledong , Jianfengling ,900 m asl ., on soil in broad - leaved forest ,25.9.1987, Z . S . Bi and T . H . Li ( GDGM 12414); Henan , Shiziping , on soil ,26.8.1968, H . Z . Li et al .162b( HMAS 35975b); Jilin , Antu , Changbaishan ,1750 m asl ., on soil ,5.8.1960, Y . C . Yang et al .564( HMAS 29135); Jilin , Antu , Changbaishan , no date , Z . X . Xie 820373( IFP ); Xizang ( Tibet ), Bomi , Galongla ,3000 m asl ., on soil in forest ,24.8.1982, X . L . Mao 227( HMAS 51439). Yunnan , Chuxiong , Zixishan ,2400 m asl ., on soil under trees of Lithocarpus sp . and Pinus armandi ,2.8.2001, Z . L . Yang 3144( HKAS 38321); Yunnan , Guangnan , Maojie ,1500 m asl .,27.6.1959, Q . Z . Wang 688( HMAS 26498); Yunnan , Kunming ,17.6.1973, Y . C . Zong et al .155( HMAS 36284); Yunnan , Kunming , on soil ,30.6.1973, Q . M . Ma et al .285 and Y . C . Zong et al .285( HMAS 36036 and 37441 respectively ); Yunnan , Kunming , Dapiji , on soil under trees of Quercus sp .,8.7.1942, W . F . Chiu 8216( HMAS 4216); Yunnan , Kunming , Heilongtan ,21.8.1987, Y . Xuan 1385( HKAS 18318); Yunnan , Kunming , Heilongtan ,1980 m asl ., on soil in foret dominated by Quercus sp ., Pinus armandi , and P . yunnanensis ,27.6.2001, Z . L . Yang 3081( HKAS 38159, HOLOTYPE ); Yunnan , Kunming , Heilongtan , on soil in forest with species of Quercus and Pinus ,30.6.1991, Z . L . Yang 1074( HKAS 24219); Yunnan , Kunming , Heilongtan ,1980 m asl ., on soil under trees of Picea sp .,4.9.1999, Z . L . Yang 2628( HKAS 34081); Yunnan , Kunming , Jindian ,1700 m asl ., on soil ,29.9.1995, Z . L . Yang 2199( HKAS 29630); Yunnan , Kunming , Jindian , on soil under Pinus sp .,4.8.1995, M . Wang 3( HKAS 32507); Yunnan , Songming , Liangyushan , on soil in forest ,17.9.1979, G . H . Feng 13( HKAS 4646); Yunnan , Yingjiang , Xima ,1700 m asl ., on soil under trees of Lithocarpus sp .,16.7.2003, Z . L . Yang 3708( HKAS 42894).
Notes : Amanita griseofolia , a member of Amanita subgenus Amanita section Vaginatae ( Fr .) Qu é l . in the sense of YANG (1997), is characterized by its small to medium - sized , slender fruitbody with grey to dark grey , felty to verrucose or farinose , easily removed volval remnants on a grey , dark grey to brownish grey pileus , greyish to grey lamellae , a slender greyish stipe , felty to granular or verru - cose , grey to dark grey volval remnants in incomplete belts around the non - bulbous stipe base , and globose to subglobose , inamyloid basidiospores .
HKAS 4646,18318,24219, and 32507, as well as HMAS 4216 and 36284 were regarded as A . ceciliae by YANG (1997). However , A . ceciliae ( Berk . et Broome ) Bas , originally described from Europe , differs from A . griseofolia by its much more robust fruitbody with a yellow - brown , reddish brown to grey - brown or olive - brown pileus covered with lighter colored ( greyish to brownish ) volval remnants , white lamellae with white edges , and a relatively thicker stipe . Furthermore , the volval rem - nants at the base of the stipe of A . ceciliae often form a ring - zone above a strangulate region and a floccose , nearly cupulate structure at the very base of the stipe ( PHILLIPS 1990, Breitenbach & Kr ä nzlin 1995), and there are more filamentous hyphae in the volval remnants of European A . ce - ciliae [ France , Jura , near Noga ,2.10.1992, C . Bas 9341( L )] than in A . griseofolia . The basidio - spores of A . ceciliae are slightly larger than those of A . griseofolia ( TULLOSS , OVREBO & HALLING 1992, BREITENBACH & KR ä nzlin 1995).
Amanita griseofolia is also similar to A . beckeri Huijsman ex Huijsman , A . cinctipes Corner et Bas , A . sororcula Tulloss , Ovrebo et Halling , and A . liquii Zhu L . Yang , M . Wei ß et Oberw . However , Amanita beckeri , originally described from Europe , differs from A . griseofolia by its differently colored pileus , white volval remnants browning in age , but never becoming grey or darker , white to crcam lamellae with brownish spots in age , and smaller basidiospores ( HUIJSMAN 1962a,1962b, TuLLoss 1994).
Amanita cinctipes , originally described from Singapore , is distinguished from A . griseofolia by its abundant volval remnants on the base of the stipe forming 2-4 rings , much thinner pileipellis , smaller basidia , and smaller basidiospores ( CORNER & BAS 1962).
Amanita sororcula , originally described from Colombia , South America , differs from A . griseo - folia by its smaller fruitbody , thinner pileipellis with common vascular hyphae , and somewhat smaller and less perfectly globose basidiospores ( TULLOSS , OvREBO & HALLING 1992).
Amanita liquii , originally described from southwestern China , differs from A . griseofolia by its larger , much more robust and fleshier fruitbody with a dark brown to blackish pileus covered with thicker and darker volval remnants , thicker pileipellis , somewhat larger basidia , and larger basidio - spores . Furthermore , the pigment in the cells of volval remnants of A . liquii is much darker than the similarly located pigment in A . griseofolia , and the volval remnants at the stipe base of A . liquii are more abundant than those of A . griseofolia ( YANG , WEISS & OBERWINKLER 2004).

菌盖直径3~7(9)cm，扁半球形至平展，中央无凸起或稍凸起;菌盖表面灰色、暗灰色至褐灰色，具灰色至深灰色易脱落的粉质颗粒状至毡状菌幕残余;菌盖边缘具沟纹(0.2~0.5R)。菌褶幼时污白色，成熟时淡灰色;褶缘色较深，干后变为灰色至深灰色;短菌褶近菌柄端多平截。菌柄长(6)8~16cm，直径0.5~1.5cm，细长，白色至污白色，下半部被灰色纤丝状鳞片，上半部被灰色粉末状鳞片，内部空心，基部不膨大，无球状体;菌环阙如;菌幕残余(菌托)灰色至深灰色粉质，排列不规则或有时呈带状。

灰褶鹅膏 图44
Amanita griseofolia Zhu L. Yang, Frontiers in Basidiomycote Mycology: 315, Figs. 1~7, 2004.
Amanita ceciliae sensu Zhu L. Yang, Biblioth. Mycol. 170: 116, 1997, p. p.; non A. ceciliae (Berk. & Broome) Bas, Persoonia 12: 192, 1984.
担子果小型至中等。菌盖直径3~7（9）cm，初期钟形至半球形，后期平展，中央无凸起或稍凸起，灰色、暗灰色至褐灰色，无黄色色调，具灰色至深灰色易脱落的粉质颗粒状至毡状菌幕残余，湿时稍黏；边缘具棱纹（0.2~0.4R）。菌肉白色，在菌盖中央厚2~5 mm。菌褶离生，成熟时浅灰色，褶缘色较深，较密，不等长，干后变为灰色至深灰色；短菌褶近菌柄端多平截。菌柄长（6）8~16 cm，粗0.5~1.5 cm，细长，近圆柱形，白色至污白色，下半部被灰色、纤丝状鳞片，上半部被灰色、粉状鳞片，内部空心，基部不膨大，无球状体。菌柄基部菌幕残余（菌托）灰色至深灰色粉质，不规则排列，有时呈带状。菌环缺如。
菌褶多少两侧型：中央菌髓宽约30（40）μm，由菌丝（直径3~7 μm）和椭圆形至近梭形的膨大细胞（70~90×15~20 μm）构成；侧生菌髓不明显。亚子实层厚30~50 μm，由2~3（4）层近球形、卵状至短椭圆形的膨大细胞（12~25×10~20 μm）组成。担子45~70×15~20 μm，棒状，具4小梗；小梗长5~7 μm；担子基部横隔上无锁状联合。担孢子（9.5）10.0~13.5（16.5）×（8.5）9.5~13.0（15.0）μm [Q = 1.0~1.10（1.18），Q = 1.04±0.03]，球形至近球形，非淀粉质，无色，光滑，薄壁；侧生小尖长约1 μm。褶缘（侧面观）有宽达250 μm的不育带，由菌丝和膨大细胞构成：膨大细胞非常丰富，卵状至近球形，有时球头短柄状，20~50×15~35 μm，无色，壁薄，或有时有黄褐色胞内色素，单个顶生或2~3个连成念珠状；菌丝较丰富，直径2~7 μm，无色，不规则排列，有时其走向与褶缘大致平行。菌盖表皮厚50~100 μm，其中上层厚约20~60 μm，强烈胶化，由近辐射状排列较为疏松的菌丝组成，菌丝直径1~4 μm，无色或有浅褐色胞内色素；下层厚30~50 μm，由紧密多少近辐射状排列的菌丝组成，菌丝直径3~8（15）μm，常有褐黄色胞内色素。菌盖表面菌幕残余由近纵向排列（菌盖中央）至不规则排列（菌盖其他部分）的菌丝和膨大细胞构成：膨大细胞非常丰富，近球形、卵形至短椭圆形（20~70×15~50 μm），有时宽棒状（40~60×20~30 μm），有时球头短柄状（50~80×25~45 μm），壁厚达0.5 μm，常有黄褐色至褐色胞内色素，常2~3个连成念珠状；菌丝稀疏至较丰富，直径2~7 μm，无色或有时有褐黄色胞内色素。菌柄基部的菌幕残余结构与菌盖表面的相似，但菌丝和膨大细胞不规则排列。菌柄菌髓主要由纵向排列的长棒状的膨大细胞（250~350×25~40 μm）构成，其间夹杂的菌丝稀少（菌柄内部）至较多（菌柄表面），直径2~7 μm。子实体无锁状联合。
模式产地：中国（云南）。
生境：生壳斗科、松科植物组成的林中地上。
研究标本：北京：潭柘寺，1958年7月25日，邓叔群6063（HMAS 22610）；潭柘寺，1959年7月29日，王维兴、孔显良74（HMAS 26499）。吉林：安图，长白山，时间不详，谢支锡820373（IFP）；同上，海拔1750 m，1960年8月5日，杨玉川等564（HMAS 29135）。河南：狮子坪，1968年8月26日，李惠中等162b（HMAS 35975b）。湖南：永顺，1975年6月4日，无名氏（MHHNU 1235）；长沙，1984年5月，无名氏（MHHNU 4450）；长沙，1984年7月，无名氏（MHHNU 4188）。海南：乐东，尖峰岭，1987年9月25日，毕志树、李泰辉（HMIGD 12414）；昌江，霸王岭，1990年8月24日，陈庆（HMIGD 16689）。云南：楚雄，紫溪山，2001年8月2日，杨祝良3144（HKAS 38321）；广南，猫街，海拔1500 m，1959年6月27日，王庆之688（HMAS 26498）；昆明，1973年6月30日，马启明等285（HMAS 36036）、宗毓臣等285（HMAS 37441）；昆明，1973年6月17日，宗毓臣等155（HMAS 36284）；昆明，大普吉，1942年7月8日，W. F. Chiu 8216 [HMAS 4216（清华8216）]；昆明，金殿，1995年8月4日，王明3（HKAS 32507）；同上，1995年9月29日，杨祝良2199（HKAS 29630）；昆明，黑龙潭，1991年6月30日，杨祝良1074（HKAS 24219）；同上，1999年9月4日，杨祝良2628（HKAS 34081）；同上，2001年6月27日，杨祝良3081（HKAS 38159，主模式）；同上，1987年8月21日，宣宇1385（HKAS 18318）；嵩明，梁玉山，1979年9月17日，冯桂华13（HKAS 4646）；盈江，昔马，海拔1700 m，2003年7月16日，杨祝良3708（HKAS 42894）。西藏：波密，嘎隆拉，海拔3000 m，1982年8月24日，卯晓岚227（HMAS 51439）。
世界分布：中国。
讨论：灰褶鹅膏 A. griseofolia 的主要特征是：担子果小型至中等；菌盖灰色、暗灰色至褐灰色，无黄色色调，具灰色至褐灰色易脱落的粉质颗粒状至毡状菌幕残余，菌肉较薄；菌褶浅灰色，干后变为灰色至深灰色；菌柄细长，被灰色纤丝状至粉状鳞片，基部不膨大；菌幕残余（菌托）灰色至深灰色，粉质，不规则排列或呈带状；菌环缺如。
在我国，过去一般将菌柄基部不膨大、菌托多少呈圈状的鹅膏，定为“圈托鹅膏”，其拉丁学名常用 Amanita inaurata Secr.、Amanita strangulata (Fr.) Quel. 或 Amanitopsis strangulata Fr.，近来也见有用 Amanita ceciliae (Berk. & Broome) Bas 的（Teng 1932a, 1936；邓叔群1963；应建浙和臧穆1994；Teng 1996；Yang 1997；卯晓岚1998）。值得指出的是，Secretan（1833）发表的名称属不合格发表（Greuter et al. 2000），所以 Amanita inaurata Secr. 这一名称是不能使用的。Amanita inaurata Secr. ex Gillet 虽然是合格的，但它是 Agaricus ceciliae Berk. & Broome [= Amanita ceciliae (Berk. & Broome) Bas] 的晚出异名（Bas 1984）。根据 Bas（1984）的意见，Amanita strangulata (Fr.) Quel. [= Amanitopsis strangulata Fr.] 是个有菌环的种，具体是什么尚不清楚。因此，欧洲的“圈托鹅膏”目前一般使用“Amanita ceciliae (Berk. & Broome) Bas”这一拉丁名。
圈托鹅膏 A. ceciliae 原初描述于欧洲，它的菌柄粗壮，菌盖常带黄色色调，菌盖菌肉较厚，菌褶白色，干后不变为灰色，菌幕残余（菌托）常呈环带状；菌盖表面的菌幕残余中菌丝较多（Phillips 1990; Breitenbach & Kränzlin 1995）。近来的研究表明，我国过去文献中记载的“圈托鹅膏”可能包囊了至少两个种，即李逵鹅膏（A. liquii Zhu L. Yang et al.）和灰褶鹅膏，这二者的区别可见本志鞘托鹅膏组 sect. Vaginatae 分种检索表。
李惠中等162（HMAS 35975）包括了两个种的标本，李惠中等162a（HMAS 35975a）是褐烟色鹅膏 A. brunneofuliginea Zhu L. Yang。