在过去很长时期,蜈蚣衣科曾置于茶渍目Lecanorales。第10版《真菌辞典》(Kirk et al. 2008)将蜈蚣衣科置于黄枝衣目Teloschistales。近期研究(Gaya et al. 2012;Miadlikowska et al. 2014)已将蜈蚣衣科置于粉衣目Caliciales。
本研究基于形态学与地衣化学确认以下5个种为中国新记录：美洲哑铃孢Heterodermia galactophylla,红髓外蜈蚣叶Hyperphyscia crocata,狭叶红髓黑蜈蚣叶Phaeophyscia fumosa,毛裂芽黑蜈蚣叶P. kairamoi和变白蜈蚣衣Physcia albata。
1 材料与方法
1.1 研究标本
研究标本均保存在中国科学院微生物研究所菌物标本馆地衣标本室(HMSA-L)。
1.2 形态和解剖
形态观察使用Zeiss Stemi SV6或Motic体视显微镜,并拍摄物种形态特征图。子囊盘和分生孢子器分别徒手切片或使用冷冻切片机切片,置于光学显微镜(Zeiss Axioscop 40,Zeiss Axioscop 2或Olympus BH-2)下观察子囊盘结构、子囊孢子以及分生孢子等。
1.3 地衣化学
显色反应采用常规试剂10%氢氧化钾(K);5%-10%对苯二胺乙醇溶液(PD);漂白粉饱和水溶液或巴士液(C);Lugols碘液(I)。地衣物质的检测采用薄层色谱(TLC)方法(Culberson 1972),通常使用C溶剂系统。
2 结果与分析
2.1 美洲哑铃孢 图1A
图1
图1
中国蜈蚣衣科5种地衣 A：美洲哑铃孢的粉芽堆(陈健斌等9395);B：红髓外蜈蚣叶(赵继鼎、徐连旺8567);C：狭叶红髓黑蜈蚣叶(陈健斌等9564-1);D：毛裂芽黑蜈蚣叶的裂芽和小裂片上有毛(王先业1261);E：变白蜈蚣衣(王先业等8860). 标尺：A-D=2 mm;E=5 mm
Fig. 1
Five species of Physciaceae. A: Soralia of Heterodermia galactophylla (Chen et al. 9395); B: Hyperphyscia crocata (Zhao & Xu 8567); C: Phaeophyscia fumosa (Chen et al. 9564-1); D: Hairs on isidia and lobules of Phaeophyscia kairamoi (Wang 1261); E: Physcia albata (Wang 8860). Bars: A-D=2 mm; E=5 mm.
Heterodermia galactophylla (Tuck.) W.L. Culb. Bryologist 69: 482, 1967.
≡ Parmelia ciliaris var. galactophylla Tuck., Proc. Amer. Acad. Arts Sci. 1: 224, 1848.
地衣体叶状,连座状至不规则扩展,2-3.5 cm宽,较疏松的常常仅以中央部位贴于基物;裂片二叉式至不规则分裂,亚覆瓦状,裂片顶部抬升,基部0.7-1.5 mm宽,顶部加宽,常呈匙形,顶端略圆,可达3-4 mm宽;上表面淡灰白色至淡灰绿色,平坦或微凸(边缘稍向下卷);髄层白色;下表面无皮层,似蛛网和脉纹状,白色,粉芽顶生,粉芽堆略唇形,无黄色色素;边缘生假根与地衣体同色,顶部常变暗色,单一不分枝至稀分枝,长0.5-1.5(-2.0) mm。子囊盘未见。
化学：上皮层K+黄色;髓层K+黄色,C-,KC-,PD-;含有atranorin,zeorin。
研究标本：福建,武夷山,海拔2 150 m,张恩然wy364-1,wy372。江西,庐山植物园,赵继鼎等399。湖南,桑植县天平山,1 300- 1 400 m,陈健斌等 9395,9525,9873;衡山,海拔750-1 200 m,赵继鼎、徐连旺9992,10134。广西,花坪,海拔900 m,赵继鼎、徐连旺9728。
世界分布：热带至暖温带(Kurokawa 1973)。主要分布于美洲(北美,中美,南美)。
讨论：本种主要特征是具有抬升和匙形裂片,无下皮层,无黄色色素,下表面顶部有粉芽,缺乏norstictic acid,salazinic acid。本种十分相似于丛毛哑铃孢H. comosa和翘哑铃孢H. subascendens,主要区别在于丛毛哑铃孢具有表面生的缘毛型白色毛,不同于翘哑铃孢在于本种下表面缺乏黄色色素。本种还与透明哑铃孢H. pellucida的外形和化学相似,但后者无粉芽。
2.2 红髓外蜈蚣叶 图1B
Hyperphyscia crocata Kashiw., Bull. Natn. Mus., Tokyo, Ser. B, 11(3): 92, 1985.
地衣体叶状,略圆形,直径1-3 cm,紧密贴生基物;裂片二叉至不规则分裂,狭窄,0.5- 1.5 mm宽;上表面淡灰绿色至橄榄色,粉芽堆主要表面生,小圆形和半球形,粉芽颗粒状;髓层红色,至少髓层下半部位红色;下皮层不明显,下表面近黑色,顶部颜色稍浅,假根缺乏或非常稀少。子囊盘表面生,直径0.5-1.5 mm,盘面近黑色,盘缘完整至轻微缺刻,盘缘有时粉芽化,孢子Pachysporaria型,17-25×8-11 µm。
化学：地衣体K-;髓层含有skyrin,zeorin。
基物：树皮。
研究标本：广西,临桂县宛田,赵继鼎、徐连旺8567。云南,思茅附近,陈健斌6587。
世界分布：中国,日本(Kashiwadani 1985)。
讨论：本种主要特征是具有表面生粉芽堆,髓层红色,含有zerion,skyrin。本研究引证的2份标本中有一份标本被Moberg来华期间鉴定为H. pandani (H. Magn.) Moberg。但H. pandani的裂片十分狭窄,0.2-0.5 mm宽(Rogers 2011),或最大达1 mm宽(Moberg 1987),缺乏zeorin。本研究引证的2份标本的裂片0.5-1.5 mm宽,用TLC方法测定的结果含有zeorin。在外蜈蚣叶属地衣中含有zerion是少见的。中国的这2份标本因裂片0.5-1.5 mm宽,具粉芽,红色髓层及含有zeorin等特征,符合产于日本的H. crocata Kashiw. (Kashiwadani 1985)。与原描述相比,中国标本的下表面的颜色较暗近黑色,顶部颜色稍浅,而日本的标本下表面淡褐色至褐色。此外,中国标本的孢子17-25×8-11 µm,稍大于日本标本的孢子16-20×7-8 µm (Kashiwadani 1985)。
2.3 狭叶红髓黑蜈蚣叶 图1C
Phaeophyscia fumosa Moberg, Nord. J. Bot. 3: 514, 1983.
地衣体叶状,近圆形,约2-4 cm宽,裂片近放射状排列,分离至中央部位紧密相连或稍微重叠,十分狭窄,0.3-0.6 mm宽,最大者不超过1 mm宽,平坦至稍稍下凹,上表面淡灰褐色至褐色,无粉芽和裂芽,髓层红色(色素skyrin),下表面黑色,顶部淡色;假根黑色。子囊盘直径0.5-1.5 mm,盘缘完整或缺刻,基部常常有假根,子囊棍棒状,内含8孢;子囊孢子褐色,双胞,厚壁,Pachysporaria型或主要是Pachysporaria型,(20-)23-27(-31)×8.5-12 µm。
化学：地衣体K-;髓层含有色素skyrin。
基物：树皮。
研究标本：福建,武夷山自然保护区专家楼附近,海拔800 m,黄满荣924。湖南,桑植天平山,海拔1 350-1 400 m,陈健斌等9490,9564-1,9636;湖南宜章县莽山,海拔1 300 m,陈健斌等21013。四川,峨眉山万年寺,海拔1 000 m,赵继鼎、徐连旺8320。云南,贡山县,海拔2 300 m,苏京军2323;独龙江,海拔2 000 m,苏京军3923-1;保山市高黎贡山白华岭林场,海拔1 700 m,姜玉梅697;泸水县片马,海拔2 400 m,王先业等2147。
世界分布：中国,东非(Moberg 1983),澳大利亚(McCarthy 2018)。
讨论：本种的主要特征是裂片十分狭窄,通常0.3-0.6 mm宽,最宽不超过1 mm,无粉芽和裂芽,髓层橙红色,不含zeorin,树生。与P. pyrrophora的主要区别在于裂片狭窄,P. pyrrophora的裂片宽1-2.5 mm,日本的标本可达4 mm宽。本种与P. endococcina的区别在于后者是一个石生种,而且髓层含有zeorin。Moberg(1983)记载本种孢子为24-32×12-14 µm,稍大于中国标本的孢子(20-) 23-27(-31)×8.5-12 µm。
2.4 毛裂芽黑蜈蚣叶 图1D
Phaeophyscia kairamoi (Vain.) Moberg, Symb. Bot. Upsal. 22(1): 40, 1977.
≡ Physcia kairamoi Vain., Acta Soc. Fl. Fenn 7: 3, 1921.
地衣体叶状,不规则状至略圆形,疏松着生于基物;裂片不规则分裂,1-2(-3.5) mm宽;上表面灰褐色至暗褐色,平坦或稍凹,末端多向上翘起;上表面末端时有稀疏无色皮层毛,裂芽多边缘生,偶有表面生,常发展成背腹分明的小裂片,这些裂芽和小裂片有时密集呈簇,裂芽或小裂片生有淡色或暗色的毛;髓层白色;下表面黑色,末端常为灰白色;假根通常稠密,黑色,单一不分枝或有简单枝,常伸出裂片边缘。子囊盘少见,圆盘状,直径可达2 mm,盘面黑色,无粉霜,盘缘完整或缺刻,基部常常具黑色假根。子囊孢子双胞,褐色,Physcia型,22-27×9-11 µm。分生孢子器埋生于地衣体中,孔口开于地衣体上表面,暗褐色至黑色,圆点状。分生孢子椭圆形、卵形,2-3×1-1.5 µm。
化学：地衣体K-;髓层K-;未检测出地衣物质。
基物：岩石,石表藓层,树皮。
研究标本：湖北,神农架千家坪,海拔2 000 m,陈健斌11719-1a。四川,若尔盖县铁布,海拔2 800 m,王先业、肖勰11091;卧龙,海拔2 750 m,陈健斌6339-1。云南,中甸小中甸林场,海拔3 600 m,王先业等5529;西畴法斗,海拔1 400 m,陈健斌5313。西藏,林芝县,海拔3 200 m,胡光荣h217,h247。新疆,天山夏塔,海拔2 300-2 900 m,王先业920;北木扎尔台河谷,海拔2 500-2 600 m,王先业1068,1167,1218,1261;天山昭苏县林场和巩留林场,海拔1 950-2 000 m,赵从福2555,2594-1。
世界分布：中国,欧洲,北美,格陵兰,东亚(Moberg 1994,1995)。
讨论：本种由于具有边缘生裂芽和小裂片而相似于P. exornatula,但本种的裂芽和小裂片上有淡色或暗色的毛,而且裂片上表面顶部区域常存在无色皮层毛。在本研究的标本中,新疆、四川的标本比较容易识别。有的标本无皮层毛,或这些皮层毛比较微小,需仔细观察。Moberg(1995)曾经报道过中国和俄罗斯远东地区的黑蜈蚣叶属Phaeophyscia地衣,但其中P. kairamoi并没有来自中国的标本,引证的标本全部来自俄罗斯和蒙古国的有关地区。
2.5 变白蜈蚣衣 图1E
Physcia albata (F. Wilson) Hale, Bryologist 66: 73, 1963.
≡ Parmelia albata F.Wilson, Pap. Proc. Roy. Soc. Tasmania: 173, 1893
地衣体叶状,近圆形或不规则,直径可达9 cm,疏松着生于基物;裂片不规则分裂,相对较宽,1.5-4.0 mm宽,顶端钝圆,时有缺刻;上表面灰白色至淡灰绿色,顶端部位常为暗灰色,无白斑,有时具微弱粉霜,粉芽堆表面生,开始呈斑点形(maculiform),多呈白色,然后粉芽堆发育成大型或连成一片,粉芽颗粒状,有时粉芽消失后,上表面留有疤痕,露出髓层;髓层白色;下表面灰白色至淡褐色,下皮层为假薄壁组织;假根与下表面同色或有时变暗色,中等稠密,单一不分枝。子囊盘及分生孢子器未见。文献记载本种子囊盘稀少,孢子双胞,Pachysporaria-type,17-25×8-12 μm (Moberg 1986);分生孢子杆状,4-6×1 μm (Elix 2011)。
化学：地衣体K+黄色;髓层K+黄色;含有atranorin,zeorin。
基物：树皮(文献记载也生于岩石)。
研究标本：四川,西昌市泸山,海拔1 500 m,王先业等8069;贡嘎山,海拔1 900-3 450 m,王先业等8860,9023。云南,昆明西山,陈健斌6470-1。
世界分布：中国,东非、南非(Moberg 1986,2004),南美(Moberg 1990),澳大利亚,新西兰(Moberg 2001;Galloway & Moberg 2005)。
讨论：本种的主要特征为裂片较宽阔(宽1.5-4.0 mm),粉芽堆表面生,含有atranorin和zeorin。P. albata和P. dilatata是中国蜈蚣衣属地衣中裂片最宽阔的两个种。但P. dilatata无粉芽,二者互为种对(Moberg 1986,1990)。
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