一些树栖斑痣盘菌的分类及生物学特性研究
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摘要
本文采用现代菌物学研究技术，对我国斑痣盘菌科(Rhytismataceae Chev.)的部分属进行了较系统的研究。主要内容包括：各菌种形态解剖学、发育学特性的研究；安徽一些种的生态习性、培养性状、子囊孢子萌发特性的探讨及酯酶同工酶酶谱的测定与分析；各属的多性状综合分类。
赞同Cannon & Minter (1986)提出的分类观点，将散斑壳属、环绵盘菌属、皮下盘菌属、小皮下盘菌属、丝环盘菌属、黑皮盘菌属、小沟盘菌属置于盘菌纲(Discomycetes)斑痣盘菌目(Rhytismatales)斑痣盘菌种(Rhytismataceae)下。对来自全国16个省份的近200号标本进行鉴定与分类，共查明7属18种。其中包括1个新种：Hypoderma sabinae；1个中国新记录属、种：Naemacyclus fimbriatus；1个新组合：Lirula filiformis；国内13个仅见报道或描述不祥的已知种。编制了该7属分种的二叉式检索表，对各种的形态解剖学特征、寄主和地理分布分别作了记载，并加以讨论。在参考前人研究资料的基础上，综述并评价了斑痣盘菌科下7个属的分类历史、形态学、生态与地理分布以及生物学特性。新种的模式标本保存在安徽农业大学森保教研室(AAUFP)。
本次研究的18种斑痣盘菌来自安徽、江苏、海南、西藏、黑龙江等16省，总体上该类真菌由于喜温暖、潮湿的环境条件而呈南多北少的地理分布格局，并且寄主范围广泛。多数种仅知在单寄主属发生，也有少数种能在多寄主上产果。它们共涉及寄主植物2科(松科、柏科)7属24种。此类真菌侵染林木活的叶片、枝干等部位而引起不同程度的病害，或生于衰老或正腐败的部位，一般在枯死部位产生子实体。
测定了白皮松散斑壳(Lophodermium pini-bungeanae)的子囊孢子萌发同温度、pH值、营养、光照及紫外线的关系。结果表明：白皮松散斑壳子囊孢子萌发的最适温度是20～25℃，并在30℃下也有较大的萌发率。在pH3-9的酸度范围内子囊孢子均能萌发，最适pH值为4～5。不同营养条件对子囊孢子的萌发率影响较大：其中以2％葡萄糖中的萌发率最高，25％的松针煎汁次之，再次为2％的蔗糖溶液及20％马铃薯煎汁，在无菌水中的萌发率极低。本次研究还发现全黑暗条件最适宜于白皮松散斑壳子囊孢子的萌发。紫外光对子囊孢子的萌发率有一定的影响，随着紫外光处理时间的延长，子囊孢子的萌发率随之降低，紫外光照射时间超过10min时，子囊孢子完全不萌发。
以组织分离法对采自安徽黄山及合肥市苗圃的13种斑痣盘菌进行了分离培养。研究了营养、光照、温度、pH值及3种药剂与白皮松散斑壳菌落生长量的关系。结果如下：对在不同培养条件下的菌落生长情况的观察，其菌落的生长与时间呈线性相关。由方差分析可知，各菌落在不同培养基上生长速度为OA＞PA＞MA、PDA。白皮松散斑壳营养菌丝在2～32℃的范围内均可生长，最适生长温度为24℃。光照处理对白皮松散斑壳的菌丝生长有一定的影响，其生长量在室内漫射光与黑暗自然交替、全光照之
间差异不显著，而在全黑暗条件下生长最快。菌落生长的酸碱度范围为pH3－8，最适
pH值为5．5巧．5。在农药处理中，甲基托布津、百菌清、多菌灵3种药剂对白皮松散
斑壳菌落的生长均有一定的抑制作用，其抑制中浓度（EC印）分别为0．13PPm八53PPm、
0刀gppm，故多菌灵的抑菌作用最强。
采用聚丙烯酚胺凝胶电泳方法获得12个斑病盘菌菌株的酯酶同工酶酶谱，根据系
统聚类，可将其归为5种类型。结果表明不同菌株间酯酶谱存在差异，而亲缘关系愈
近的菌株其酶谱的相似性愈大，属间差异明显大于种间，同菌种的酶谱可因寄主、环
境因素的不同而存在一定的差异性。认为酯酶同工酶酶谱的测定与分析可作为区分近
似种的一种辅助手段。
In this present thesis, some genera of Rhytismataceae Chev. were studied in the taxonomic method of synthesizing several characters. The main content included researches on morphology and developmental biology, discussion on the ecology, cultural features, the regulation of ascospora gemination in different condition and enzymology of several species in Anhui, and seven genera taxology.
Cyclaneusma, Hypoderma, Hypodermella, Lirula, Lophodermium, Meloderma and Naemacyclus were placed in Discomycetes, Rhytismatales, Rhytiamataceae, according to Cannon & Minter (1986). About 200 samples collected from 16 provinces in China were examined and 7 genera and 18 species were identified. Among them there were 1 new species: Hypodema sabinae; 1 new recording genus and species to China: Naemacyclus fimbriatus; 1 new recombination: Lirula filiformis; and still included 13 species which were nor systematically described or only mentioned simply known from the country. Keys were provided for identification of all recognised species. The anatomical configuration characteristics were depicted and illustrated, also the hosts and geographical distribution were recorded here. Taxological history, anatomical configuration fetures, ecological properties and geographical distribution of these seven genera were summarized and estimated on the basis of referring to former scholars' data. The new species type specie
smen was deposited in the Forest Protection Laboratory of Anhui Agricultural University (AAUFP), Hefei, China.
18 species of Rhytismataceae were collected from 16 proviences such as Anhui Provience, Jiangsu Provience, Hainan Provience, Xizhang Provience, Heilongjiang Provience, ect. As a whole this group of fungi presented the main distribution of more species in South and fewer in North because of used to the warm and humid circumstances. The range of host plants were abroad. Most species occur only on a single host gunus, but there were a few species had a wide range of host plants. Two families (Pinaceae, Cupressaceae), 7 genera and 24 speceis of host were involved in the study. This type of fungi infected arboreous live place such as leaves and limbs leading to deseases in different degrees. Sometimes they also infected senile or decaying parts. Commonly the ascomata were produced in the blasted places.
The Rhytismatacese fungi collected from the Huangshan Mountains and the Hefei Nersery were isolated by tissue isolation techniques. The biological characters of Lophodermium pini-bungeanae were studied. The results were reported as follows: the optimum temperature ranged from 20℃ to 25℃, and the rate of ascospora germination was
relatively high at 30℃. Ascospores could germinate in the acidic scope from pH3~9, but the optimum range of acidity alkalinity was pH4-5. The different nutrient substance had great effects on the rate of ascospora germination which was the highest in the 25% glucose and lowered in the solution of 25% decoctde pine-needle, the solution of 2% sucrose , the solution of 20% potato in order. The rate of germination was special low in sterile water. And in the condition of entire darkness the rate of ascospora germination was higher than in the condition of natural alteration of indoor diffused light and darkness ,or the condition of entire light. Ultraviolet radiation restrain ascospores to germinate. The rate of germination was failed along with the time prolonged when the ascospores were treated with UV.
The effect of the nutrition, light, temperature, acidity alkalinity and three fungicides on the fugal hyphal growth rate were also studied. The results were reported as follows: the diameter of the fungal colony was linear correlated with time. And the hyphal growth rate in different culture medium, OA>PA>MA PDA, analyzed in the method of regression analysis. The hyphae could grow from 2℃ to 32℃, and the optimum temperature for the hyphal growth was from 24℃ to 25℃. The treatment of light had effects on the hyphal growth. The condition of entire darkness speeded the growth of the co
赞同Cannon & Minter (1986)提出的分类观点，将散斑壳属、环绵盘菌属、皮下盘菌属、小皮下盘菌属、丝环盘菌属、黑皮盘菌属、小沟盘菌属置于盘菌纲(Discomycetes)斑痣盘菌目(Rhytismatales)斑痣盘菌种(Rhytismataceae)下。对来自全国16个省份的近200号标本进行鉴定与分类，共查明7属18种。其中包括1个新种：Hypoderma sabinae；1个中国新记录属、种：Naemacyclus fimbriatus；1个新组合：Lirula filiformis；国内13个仅见报道或描述不祥的已知种。编制了该7属分种的二叉式检索表，对各种的形态解剖学特征、寄主和地理分布分别作了记载，并加以讨论。在参考前人研究资料的基础上，综述并评价了斑痣盘菌科下7个属的分类历史、形态学、生态与地理分布以及生物学特性。新种的模式标本保存在安徽农业大学森保教研室(AAUFP)。
本次研究的18种斑痣盘菌来自安徽、江苏、海南、西藏、黑龙江等16省，总体上该类真菌由于喜温暖、潮湿的环境条件而呈南多北少的地理分布格局，并且寄主范围广泛。多数种仅知在单寄主属发生，也有少数种能在多寄主上产果。它们共涉及寄主植物2科(松科、柏科)7属24种。此类真菌侵染林木活的叶片、枝干等部位而引起不同程度的病害，或生于衰老或正腐败的部位，一般在枯死部位产生子实体。
测定了白皮松散斑壳(Lophodermium pini-bungeanae)的子囊孢子萌发同温度、pH值、营养、光照及紫外线的关系。结果表明：白皮松散斑壳子囊孢子萌发的最适温度是20～25℃，并在30℃下也有较大的萌发率。在pH3-9的酸度范围内子囊孢子均能萌发，最适pH值为4～5。不同营养条件对子囊孢子的萌发率影响较大：其中以2％葡萄糖中的萌发率最高，25％的松针煎汁次之，再次为2％的蔗糖溶液及20％马铃薯煎汁，在无菌水中的萌发率极低。本次研究还发现全黑暗条件最适宜于白皮松散斑壳子囊孢子的萌发。紫外光对子囊孢子的萌发率有一定的影响，随着紫外光处理时间的延长，子囊孢子的萌发率随之降低，紫外光照射时间超过10min时，子囊孢子完全不萌发。
以组织分离法对采自安徽黄山及合肥市苗圃的13种斑痣盘菌进行了分离培养。研究了营养、光照、温度、pH值及3种药剂与白皮松散斑壳菌落生长量的关系。结果如下：对在不同培养条件下的菌落生长情况的观察，其菌落的生长与时间呈线性相关。由方差分析可知，各菌落在不同培养基上生长速度为OA＞PA＞MA、PDA。白皮松散斑壳营养菌丝在2～32℃的范围内均可生长，最适生长温度为24℃。光照处理对白皮松散斑壳的菌丝生长有一定的影响，其生长量在室内漫射光与黑暗自然交替、全光照之
间差异不显著，而在全黑暗条件下生长最快。菌落生长的酸碱度范围为pH3－8，最适
pH值为5．5巧．5。在农药处理中，甲基托布津、百菌清、多菌灵3种药剂对白皮松散
斑壳菌落的生长均有一定的抑制作用，其抑制中浓度（EC印）分别为0．13PPm八53PPm、
0刀gppm，故多菌灵的抑菌作用最强。
采用聚丙烯酚胺凝胶电泳方法获得12个斑病盘菌菌株的酯酶同工酶酶谱，根据系
统聚类，可将其归为5种类型。结果表明不同菌株间酯酶谱存在差异，而亲缘关系愈
近的菌株其酶谱的相似性愈大，属间差异明显大于种间，同菌种的酶谱可因寄主、环
境因素的不同而存在一定的差异性。认为酯酶同工酶酶谱的测定与分析可作为区分近
似种的一种辅助手段。
In this present thesis, some genera of Rhytismataceae Chev. were studied in the taxonomic method of synthesizing several characters. The main content included researches on morphology and developmental biology, discussion on the ecology, cultural features, the regulation of ascospora gemination in different condition and enzymology of several species in Anhui, and seven genera taxology.
Cyclaneusma, Hypoderma, Hypodermella, Lirula, Lophodermium, Meloderma and Naemacyclus were placed in Discomycetes, Rhytismatales, Rhytiamataceae, according to Cannon & Minter (1986). About 200 samples collected from 16 provinces in China were examined and 7 genera and 18 species were identified. Among them there were 1 new species: Hypodema sabinae; 1 new recording genus and species to China: Naemacyclus fimbriatus; 1 new recombination: Lirula filiformis; and still included 13 species which were nor systematically described or only mentioned simply known from the country. Keys were provided for identification of all recognised species. The anatomical configuration characteristics were depicted and illustrated, also the hosts and geographical distribution were recorded here. Taxological history, anatomical configuration fetures, ecological properties and geographical distribution of these seven genera were summarized and estimated on the basis of referring to former scholars' data. The new species type specie
smen was deposited in the Forest Protection Laboratory of Anhui Agricultural University (AAUFP), Hefei, China.
18 species of Rhytismataceae were collected from 16 proviences such as Anhui Provience, Jiangsu Provience, Hainan Provience, Xizhang Provience, Heilongjiang Provience, ect. As a whole this group of fungi presented the main distribution of more species in South and fewer in North because of used to the warm and humid circumstances. The range of host plants were abroad. Most species occur only on a single host gunus, but there were a few species had a wide range of host plants. Two families (Pinaceae, Cupressaceae), 7 genera and 24 speceis of host were involved in the study. This type of fungi infected arboreous live place such as leaves and limbs leading to deseases in different degrees. Sometimes they also infected senile or decaying parts. Commonly the ascomata were produced in the blasted places.
The Rhytismatacese fungi collected from the Huangshan Mountains and the Hefei Nersery were isolated by tissue isolation techniques. The biological characters of Lophodermium pini-bungeanae were studied. The results were reported as follows: the optimum temperature ranged from 20℃ to 25℃, and the rate of ascospora germination was
relatively high at 30℃. Ascospores could germinate in the acidic scope from pH3~9, but the optimum range of acidity alkalinity was pH4-5. The different nutrient substance had great effects on the rate of ascospora germination which was the highest in the 25% glucose and lowered in the solution of 25% decoctde pine-needle, the solution of 2% sucrose , the solution of 20% potato in order. The rate of germination was special low in sterile water. And in the condition of entire darkness the rate of ascospora germination was higher than in the condition of natural alteration of indoor diffused light and darkness ,or the condition of entire light. Ultraviolet radiation restrain ascospores to germinate. The rate of germination was failed along with the time prolonged when the ascospores were treated with UV.
The effect of the nutrition, light, temperature, acidity alkalinity and three fungicides on the fugal hyphal growth rate were also studied. The results were reported as follows: the diameter of the fungal colony was linear correlated with time. And the hyphal growth rate in different culture medium, OA>PA>MA PDA, analyzed in the method of regression analysis. The hyphae could grow from 2℃ to 32℃, and the optimum temperature for the hyphal growth was from 24℃ to 25℃. The treatment of light had effects on the hyphal growth. The condition of entire darkness speeded the growth of the co
引文
1. Hawksworth, D. L., Sutton, B. C., Ainsworth, G. C. Ainsworth & Bisby's Dictionary of the Fungi. 8th edn. CMI, Kew, 1995, 680pp.
2．林英任，侯成林，刘和云．中国北部地区松生斑痣盘菌科分类研究．真菌学报，1989，14(2)：92-100．
3. Candolle, A. F. de. Folre Francaise edn 3, 1805, vol. 2. 600pp. Paris: H. Agasse.
4. Chevallier, F. F. Essai sur les Hypoxylons Lichénoides. Journ. De Physique, 1822, 94:28-61.
5. Chevallierv, F. F. Flore générale de Paris vol. 1. 1826, 676pp. Paris: Ferra Jeunev.
6. Notaris, Giuseppe De. Prime linee di una nuova disposizione de Pirenomiceti Isterini. Giorn. Bot. Ital. Firenze 2: 5-52,1847.
7. Rehm, H. Revision der Hyterineen im herb. Duby. Hedwigia. 1886, 25:173-202.
8. Rehm, H. Hysteriaceae. In Rabenhorst's Kryptogamen-Flora yon Deutschland, Oesterreich und der Schweiz. 2nd Ed. 1887, 3: 1-56.
9. Lindau, Gustav. Hysteriineae. In Engler & Prantl, Die Natürlichen Pflánzentamilien. 1896, 1(1): 265-278.
10. Hhnel, F. von. System der Phacidiales von H. Bet. Deutscher Bot. Ges, 1917a, 35: 416～422.
11. Hhnel, F. von. Mycologische. CXXXVI. ber Rhytisma lineare Peck. Ann. Mycol, 1917b, 15: 318～319.
12. Hhnel, F. von. Mycologische Pragments CCLXXII. Ueber die Hysteriaceen. Ann. Mycol, 1918, 16: 145-154.
13. Nannfeldt, J. A. Studisn ber die morphologie und systematic der nicht-lichenisierten inoperculaten Discomyceten. Nova Acta Regiae Soc. Sci. Upsal. Ⅳ, 1932, 8(2): 1～368.
14. Darker, G. D. A Revision of the genera of the Hypodermaceae. Canadian Journal Botany. 1967, 45: 1400～1404.
15. Korf, R. P. Discomycetes and Tuberales. In Ainsworth, G. C., Sparrow, F. K., Sussman, A. S., The Fungi. 1973, 4A: 249-319.
16. Cannon, P. F. & Minter, D. W. The Rhytismataceae of the Indian Subcontinent. Mycological Papers, 1986, 155: 1～123.
17．林英任，刘和云，唐燕平．中国斑痣盘菌目分类研究，安徽农业大学学报，1995，22(2)：95-102．
18. Powell, P. E. Taxonomic Studies in the Genus Hypoderma (Rhytismataceae). Ph. D. Thesis, Cornell University. 1974,117pp.
19. Sherwood, M. A. Taxonomic studieis in the Phacidiales:The genus Coccomyces (Rhytismataceae). Occ. pap. Farlow herb. Crypt. bot. 1980, 15: 1～120.
20. Chevallier, F. F. Flore générale des environs de Paris. Paris. 1826,1:432-438.
21. Schrader, H. A. Plantae cryptogamicae novae, rariores aut minus vognitae. Schrad. Journ. Bot. 1799, 2: 55-70.
22. Darker, G. D. The Hypodermaceae of Conifers. Contributions from the Arnold Arboretum. 1932; 1: 6～15 etc.: Academic Press.
23. Korf, R. P. Discomycetes and Tuberales. In Ainsworth, G. C., Sparrow, F. K. & Sussman, A. S., The Fungi, 1973, 4A:249-319. New York etc.: Academic Press.
24. Dennis, R. W. G. British Ascomycetes. 1978, 585. Vaduz: J. Cramer.
25. Hunt, R. S., Ziller, W. G. Host-genus keys to the Hypodermataceae of conider leaves. Mycotaxon, 1978, 6:481～496.
26. Spooner, B. M. Lophodermium and Hypoderma (Rhytismatales) from Mt Kinabalu, Sabah. Kew Bull, 1991, 46 (1): 73～100.
27. Minter, D.W. Lophodermium on pines. Mycol, 1981, 147: 1～54.
28. Johnston, P. R. Rhytismataceae in New Zealand 1. Some foliicolous species of Coccomyces de Notaris and Propolis (Fries) Corda. New Zealand J. Bort, 1986, 24: 89～124.
29．邓叔群．中国的真菌．科学出版社，1963，808页．
30. Korf, R. P., Zhuang, W. Y. Some new species and new records of Discomycetes in China. Mycotaxon, 1985, 22(2): 483-514.
31．林英任，唐燕子．松树上的七种散斑壳．真菌学报，1988，7(3)：129-137．
32. F. DiCosmo, H. Peredo and D. W. Minter. Cyclaneusma gen.nov. Naemacyclus and Lasiostictis, a nomenclatural problem resolved. Eur. J. Path, 1983, 13: 206-212.
33. Butin, H. Morphologische und taxonomische Untersuchungen an Naemacyclus niveus (Pers. ex Fr.) Fuck. ex Sacc. Und verwandten Arten. Eur. J. For. Path, 1973, 3: 146-163.
34. Millar, C. S.; Minter, D. W. Naemacyclus minor. C. M. I. Descriptions of Pathoogenic Fungi and Bacteria 659. Commonwealth Mycological Institute, Kew. 1980.
35. Minter, D. W., Millar. C.S. Naemacyclus niveus. C. M. I. Descripions of Pathogenic Fungi and Bacteria 660. Commonwealth Mycological Institue, Kew. 1980.
36. DiCosmo, F. A Revision of the Phacidiaceae and Related Anamorphs. Ph. D. Thesis. Univerisity of Waterloo, Waterloo, Canada. 1981, 444.
37. DiCosmo, F.; Nag Raj, T. R.; Kendrick, W. B. A prodromus for a revision of the Phacidiaceae, Can. J. Bot, 1983.
38. Korf, R. P., Discomycetes and Tuberales. In: The Fungi Vol. Iva, Ed. by Ainsworth, G. C.; Sparrow, F. K.; Sussman, A. S. New York, San Francisco, London: Academic Press. 1973.
39. Fuckel, L., Symbolae mycologicae. Beitrage zur Kenntnis der rhein. Pilze. Nachtr. Ⅱ. Jahrb. d. nassauischen Ver. f. Naturkde, 1873, 27/28, 49-50.
40. Saccardo, P. A., Conspectus generum Discomycctum hucusque cognitorum. Bot. Centralbl, 1884, 18,
251.
41. Rehm, H. Discomyceten. In: Rabenhorst, L.: Kryptogamenflora von Deutschland, Osterreich und der Schweiz. Bd. 1, Teibd. 3. Leipzig: E. Kummer, 1896.
42. Petrak, F. Uber die Gattung Naemacyclus Fuck. und Lasiostictis Sacc. Sydowia, 1974, 1: 89-93.
43. Duby, J. E. Mémoire sur la tribu des Hystérinees de la famille des Hypoxylees (Pyrénomycètes). Mem. Soc. Phys. Et Hist. Nat. Geneve, 1861, 16:15-70.
44. Saccardo. P. A. Sylloge Fungorum. 1883, 2: 813pp.
45. Tubeuf, Karl. von. Beitrage zur Kentniss der Baumkrankheiten. Berlin, J. Springer, 1888, 61pp.
46. Tubeuf, Karl. von. Die Nadelschüitte der Weymouthskiefer. Naturw. Zeitschr. Forst. Landw, 1908, 6: 326-330.
47. Johnston, P. R. An undescribed pattern of ascocarp development in some non-coniferous Lophodermium species. Mycotaxon, 1988, 31 (2): 383～394.
48．林英任．长江以南地区松赤落叶病病原菌研究．森林病虫通讯，1991，4：4-6．
49. Persoon, C. H. Einige Bermerkungen ber die Flechten, nebst Beschreibungen einiger neuen Arten aus dieser Familie der Aftermoose. Usteri's Annal. d. Botan, 1794, 7: 1-32; 155-158.
50. Chevallier, F. F. Essai sur les Hypoxylons Lichénoides. Journ. De Physique, 1822, 94: 28-61.
51. Chevallier, F. F. Flore générale des environs de Paris. Paris, 1826, 1: 432-438.
52．戴芳澜．中国真菌总汇．科学出版社，1979，1527页．
53. Saccardo, P. A. Sylloge Fungorum. 1889, vol. 8.; 1143pp. Patavii: Saccardo.
54. Johston, P. R. Rhytismataceae in New Zealang 2. The genus Lophodermiurn of indigenous Plants. New Zealand J. Bot, 1989a, 27:243-274.
55. Johnston, P. R. Lophodermium (Rhytismataceae) on Clusia. Sydowia, 1989b, 41:170-179.
56．林英任．我国松树上的散斑壳菌—十二个种的检索、记述和图解．安徽农学院学报，1990，17(1)：26-31．
57. Spooner, B. M. Coccomyces and Propolis (Rhytismatales) from Mt Kinabalu, Borneo. Kew Bull, 1990,45(3): 451-484.
58. Morgan-Jones, J. F., Hulton, R. L. Ascocarp development in Lophodermium pinastri. Mycologia, 1977, 55: 2605～2612.
59. Reid, D. A. & Minter, D. W. Pseudostypella translucens (Gordon) Reid & Minter comb. nov., a hyperparasite on Lophodermium conigenum. Trans. Br. Mycol. Soc., 1979, 72: 345-347.
60. Osorio, M. & Stephan, B. R. Morphological studies of Lophodermium species (Fuckel) v. Hhnel on Norway spruce needles. Eur. J. F. Patbol., 1991, 21: 389-403.
61. Sakuyama, T. Morphology of Lophodermium species causing needle cast of pines in Japan. Trans. Mycol. Soc. Japan, 1993, 34, 433-437.
62．郭锡华．红松落针病及其防治技术的研究．林业科技通讯，1987，5：10～12．
63．何秉章．樟子松落针病的病原菌和防治技术的研究．东北林学院学报，1985；13(2)：75～81．
64．胡炳福．马尾松赤落叶病初步研究．植物病理学报，1983，13(3)：20～24．
65．林英任．松落针病的主要病原—松针散斑壳的研究．安徽农学院学报，1988，16(6)：14-25．
66．原戈．红松落针病流行规律及最佳防治期的确定．东北林业大学学报，1988，16(6)：14-25．
67．胡能书，万贤国．同工酶技术及应用．湖南科学技术出版社，1985，P1-199．
68．卢怀乡．裂殖酵母属的酶及可溶性蛋白聚丙烯酰胺凝胶电泳．真菌学报，1990，9(1)：50～55．
69. Zervakis G. et al., Taxonomial relationships within the fungal genus Pearotus as determined by isoelectric focusing analysis of enzyme patterns. J Gen Microbiol, 1992, 138: 635—645.
70. Matsuyama N, Moromizoto Z, Ogoshi A, Wakimoto S. Grouping Rhizoctonia solani with nonspecific esterase zymogram. Ann Phytopath Soc Japan, 1978,44: 652～658.
71. Spooner, B. M. Lophodermium and Hypoderma (Rhytismataceae) from Mt Kinabalu, Sabah. Kew Bull, 1991,46 (1): 73-100.
72．方中达．植病研究方法．农业出版社，1979．
73. Durrieu, Guy. Deux Hypodermataceae nouvelles sur Coniferes. Sydowia Beihefte, 1957, 1: 355-358.
74. Yasuo Suto. A new species of Hypoderma causing dieback in Chamaecyparis obtusa and Thuja orientalis. Trans. Mycol. Japan, 1983, 24: 419-424.
75. CMl Description of Pathogenic Fungi and Bacteria No. 943. Mycopathologia, 1988 102: 55-56.
76. Tubuef, Karl von. Demonstration von Lrchenzweigen. Bot. Centralb, 1895, 61: 48-49.
77. Lagerberg. Uber Naevia piniperda Rehm. Svensk Botanisk Tidskr, 1928, 22: 282-288.
78. Hartig, Robert. Wichtige Krankheiten der waldbume. Berlin. J. Springer, 1874, 127pp.
79. Minter, D. W. & Millar, C. W. Lirula nervisequia C. M. I. Descripions of Pathogenic Fungi and Bacteria 660. Commonwealth Mycological Institue, Kew, 783.1884.
80．林英任，侯成林，承河元，刘和．中国北部地区松生斑痣盘菌分类研究．真菌学报，1995，14(2)：92-100．
81．林英任，黎志，梁师文，余盛明．中国北部地区一些针叶树生斑痣盘菌．真菌学报，1995，14(3)：179-183．
82．林英任，唐燕平，刘和云．我国南部地区一些针叶树上的斑痣盘菌．真菌学报，1993，12(2)：93-98．
83．何秉章，杨殿清，齐兴武．红松上的散斑壳．真菌学报，1986，5(2)：70-73．
84．林英任，侯成林．中国北部地区松生斑痣盘菌分类研究．真菌学报，1995，14(2)：92-100．
85. Gordon, C. C. Ascocarpic centrum ontogeny of species of Hypodermatacese of conifers. Am. J. Bot, 1966, 53:319-327.
86. Gordon, C. C. Ascocarpic centrum ontogeny of species of Hypodermataceae of conifers Ⅱ. Am. J. Bot, 1968, 55: 45-52.
87. Staley, J. M. The taxonomy of Lophodermium on pines with special reference to problems in North
American christmas tree plantations. Mitt. BwuiForschAnst. Forst-u. Holzw, 1975, 108: 79-85.
88．侯成林，王有智．针叶树斑痣盘菌科真菌病原调查．林业科学研究，1995，8(4)：426-428．
89. Ouellette, G. B., Magasi, L. P. Lophomerum, a new genus of Hypodermataceae. Mycologia, 1966, 58: 275-280.
90. Rostrup, E. Fortsatte Underogolser over Snyltesvampes Angreb paa Skovtraeome. Tidskr. Skogbr, 1883, 6: 199-300.
91. Tubeuf, Karl yon. Demonstration yon Lrchenzweigen. Bot. Centralb, 1895, 61: 48-49.
92. Birula, I., Z. Dabrowski. and L. Kr6likowski. Gr6zny szkodnik Limby (Pinus cerebra) Hypoderma nova. (Ein geidling der Arve (P. cembra) Hypoderrna Namyslowski sp. nov.). Roezn. Mauk. Rolnicz. Lesn, 1928, 20: 35-41.
93．侯成林．松落针病菌的初步研究．一九九0届硕士学位研究生论文，1990，30-38．
94．何秉章，邓兴林，杨殿清等．樟子松落针病的病原菌和防治的研究．东北林学院学报，1985，13(6)：75-81．
95．李清铣，朱华，王彰明．江苏麦类禾谷镰刀菌酯酶同工酶测定．植物病理学报，1985，15(4)：217-223．
96．徐敬友．六种疫霉菌体可溶性蛋白质和酯酶同工酶的聚丙烯酰胺凝胶垂直平板电泳研究．真菌学报，1993，12(1)：54-64．
97．朱婉华．几种真菌酯酶同工酶聚丙烯酰胺凝胶电泳的初步研究．南京林业大学学报，1987，23(1)：186-190．
2．林英任，侯成林，刘和云．中国北部地区松生斑痣盘菌科分类研究．真菌学报，1989，14(2)：92-100．
3. Candolle, A. F. de. Folre Francaise edn 3, 1805, vol. 2. 600pp. Paris: H. Agasse.
4. Chevallier, F. F. Essai sur les Hypoxylons Lichénoides. Journ. De Physique, 1822, 94:28-61.
5. Chevallierv, F. F. Flore générale de Paris vol. 1. 1826, 676pp. Paris: Ferra Jeunev.
6. Notaris, Giuseppe De. Prime linee di una nuova disposizione de Pirenomiceti Isterini. Giorn. Bot. Ital. Firenze 2: 5-52,1847.
7. Rehm, H. Revision der Hyterineen im herb. Duby. Hedwigia. 1886, 25:173-202.
8. Rehm, H. Hysteriaceae. In Rabenhorst's Kryptogamen-Flora yon Deutschland, Oesterreich und der Schweiz. 2nd Ed. 1887, 3: 1-56.
9. Lindau, Gustav. Hysteriineae. In Engler & Prantl, Die Natürlichen Pflánzentamilien. 1896, 1(1): 265-278.
10. Hhnel, F. von. System der Phacidiales von H. Bet. Deutscher Bot. Ges, 1917a, 35: 416～422.
11. Hhnel, F. von. Mycologische. CXXXVI. ber Rhytisma lineare Peck. Ann. Mycol, 1917b, 15: 318～319.
12. Hhnel, F. von. Mycologische Pragments CCLXXII. Ueber die Hysteriaceen. Ann. Mycol, 1918, 16: 145-154.
13. Nannfeldt, J. A. Studisn ber die morphologie und systematic der nicht-lichenisierten inoperculaten Discomyceten. Nova Acta Regiae Soc. Sci. Upsal. Ⅳ, 1932, 8(2): 1～368.
14. Darker, G. D. A Revision of the genera of the Hypodermaceae. Canadian Journal Botany. 1967, 45: 1400～1404.
15. Korf, R. P. Discomycetes and Tuberales. In Ainsworth, G. C., Sparrow, F. K., Sussman, A. S., The Fungi. 1973, 4A: 249-319.
16. Cannon, P. F. & Minter, D. W. The Rhytismataceae of the Indian Subcontinent. Mycological Papers, 1986, 155: 1～123.
17．林英任，刘和云，唐燕平．中国斑痣盘菌目分类研究，安徽农业大学学报，1995，22(2)：95-102．
18. Powell, P. E. Taxonomic Studies in the Genus Hypoderma (Rhytismataceae). Ph. D. Thesis, Cornell University. 1974,117pp.
19. Sherwood, M. A. Taxonomic studieis in the Phacidiales:The genus Coccomyces (Rhytismataceae). Occ. pap. Farlow herb. Crypt. bot. 1980, 15: 1～120.
20. Chevallier, F. F. Flore générale des environs de Paris. Paris. 1826,1:432-438.
21. Schrader, H. A. Plantae cryptogamicae novae, rariores aut minus vognitae. Schrad. Journ. Bot. 1799, 2: 55-70.
22. Darker, G. D. The Hypodermaceae of Conifers. Contributions from the Arnold Arboretum. 1932; 1: 6～15 etc.: Academic Press.
23. Korf, R. P. Discomycetes and Tuberales. In Ainsworth, G. C., Sparrow, F. K. & Sussman, A. S., The Fungi, 1973, 4A:249-319. New York etc.: Academic Press.
24. Dennis, R. W. G. British Ascomycetes. 1978, 585. Vaduz: J. Cramer.
25. Hunt, R. S., Ziller, W. G. Host-genus keys to the Hypodermataceae of conider leaves. Mycotaxon, 1978, 6:481～496.
26. Spooner, B. M. Lophodermium and Hypoderma (Rhytismatales) from Mt Kinabalu, Sabah. Kew Bull, 1991, 46 (1): 73～100.
27. Minter, D.W. Lophodermium on pines. Mycol, 1981, 147: 1～54.
28. Johnston, P. R. Rhytismataceae in New Zealand 1. Some foliicolous species of Coccomyces de Notaris and Propolis (Fries) Corda. New Zealand J. Bort, 1986, 24: 89～124.
29．邓叔群．中国的真菌．科学出版社，1963，808页．
30. Korf, R. P., Zhuang, W. Y. Some new species and new records of Discomycetes in China. Mycotaxon, 1985, 22(2): 483-514.
31．林英任，唐燕子．松树上的七种散斑壳．真菌学报，1988，7(3)：129-137．
32. F. DiCosmo, H. Peredo and D. W. Minter. Cyclaneusma gen.nov. Naemacyclus and Lasiostictis, a nomenclatural problem resolved. Eur. J. Path, 1983, 13: 206-212.
33. Butin, H. Morphologische und taxonomische Untersuchungen an Naemacyclus niveus (Pers. ex Fr.) Fuck. ex Sacc. Und verwandten Arten. Eur. J. For. Path, 1973, 3: 146-163.
34. Millar, C. S.; Minter, D. W. Naemacyclus minor. C. M. I. Descriptions of Pathoogenic Fungi and Bacteria 659. Commonwealth Mycological Institute, Kew. 1980.
35. Minter, D. W., Millar. C.S. Naemacyclus niveus. C. M. I. Descripions of Pathogenic Fungi and Bacteria 660. Commonwealth Mycological Institue, Kew. 1980.
36. DiCosmo, F. A Revision of the Phacidiaceae and Related Anamorphs. Ph. D. Thesis. Univerisity of Waterloo, Waterloo, Canada. 1981, 444.
37. DiCosmo, F.; Nag Raj, T. R.; Kendrick, W. B. A prodromus for a revision of the Phacidiaceae, Can. J. Bot, 1983.
38. Korf, R. P., Discomycetes and Tuberales. In: The Fungi Vol. Iva, Ed. by Ainsworth, G. C.; Sparrow, F. K.; Sussman, A. S. New York, San Francisco, London: Academic Press. 1973.
39. Fuckel, L., Symbolae mycologicae. Beitrage zur Kenntnis der rhein. Pilze. Nachtr. Ⅱ. Jahrb. d. nassauischen Ver. f. Naturkde, 1873, 27/28, 49-50.
40. Saccardo, P. A., Conspectus generum Discomycctum hucusque cognitorum. Bot. Centralbl, 1884, 18,
251.
41. Rehm, H. Discomyceten. In: Rabenhorst, L.: Kryptogamenflora von Deutschland, Osterreich und der Schweiz. Bd. 1, Teibd. 3. Leipzig: E. Kummer, 1896.
42. Petrak, F. Uber die Gattung Naemacyclus Fuck. und Lasiostictis Sacc. Sydowia, 1974, 1: 89-93.
43. Duby, J. E. Mémoire sur la tribu des Hystérinees de la famille des Hypoxylees (Pyrénomycètes). Mem. Soc. Phys. Et Hist. Nat. Geneve, 1861, 16:15-70.
44. Saccardo. P. A. Sylloge Fungorum. 1883, 2: 813pp.
45. Tubeuf, Karl. von. Beitrage zur Kentniss der Baumkrankheiten. Berlin, J. Springer, 1888, 61pp.
46. Tubeuf, Karl. von. Die Nadelschüitte der Weymouthskiefer. Naturw. Zeitschr. Forst. Landw, 1908, 6: 326-330.
47. Johnston, P. R. An undescribed pattern of ascocarp development in some non-coniferous Lophodermium species. Mycotaxon, 1988, 31 (2): 383～394.
48．林英任．长江以南地区松赤落叶病病原菌研究．森林病虫通讯，1991，4：4-6．
49. Persoon, C. H. Einige Bermerkungen ber die Flechten, nebst Beschreibungen einiger neuen Arten aus dieser Familie der Aftermoose. Usteri's Annal. d. Botan, 1794, 7: 1-32; 155-158.
50. Chevallier, F. F. Essai sur les Hypoxylons Lichénoides. Journ. De Physique, 1822, 94: 28-61.
51. Chevallier, F. F. Flore générale des environs de Paris. Paris, 1826, 1: 432-438.
52．戴芳澜．中国真菌总汇．科学出版社，1979，1527页．
53. Saccardo, P. A. Sylloge Fungorum. 1889, vol. 8.; 1143pp. Patavii: Saccardo.
54. Johston, P. R. Rhytismataceae in New Zealang 2. The genus Lophodermiurn of indigenous Plants. New Zealand J. Bot, 1989a, 27:243-274.
55. Johnston, P. R. Lophodermium (Rhytismataceae) on Clusia. Sydowia, 1989b, 41:170-179.
56．林英任．我国松树上的散斑壳菌—十二个种的检索、记述和图解．安徽农学院学报，1990，17(1)：26-31．
57. Spooner, B. M. Coccomyces and Propolis (Rhytismatales) from Mt Kinabalu, Borneo. Kew Bull, 1990,45(3): 451-484.
58. Morgan-Jones, J. F., Hulton, R. L. Ascocarp development in Lophodermium pinastri. Mycologia, 1977, 55: 2605～2612.
59. Reid, D. A. & Minter, D. W. Pseudostypella translucens (Gordon) Reid & Minter comb. nov., a hyperparasite on Lophodermium conigenum. Trans. Br. Mycol. Soc., 1979, 72: 345-347.
60. Osorio, M. & Stephan, B. R. Morphological studies of Lophodermium species (Fuckel) v. Hhnel on Norway spruce needles. Eur. J. F. Patbol., 1991, 21: 389-403.
61. Sakuyama, T. Morphology of Lophodermium species causing needle cast of pines in Japan. Trans. Mycol. Soc. Japan, 1993, 34, 433-437.
62．郭锡华．红松落针病及其防治技术的研究．林业科技通讯，1987，5：10～12．
63．何秉章．樟子松落针病的病原菌和防治技术的研究．东北林学院学报，1985；13(2)：75～81．
64．胡炳福．马尾松赤落叶病初步研究．植物病理学报，1983，13(3)：20～24．
65．林英任．松落针病的主要病原—松针散斑壳的研究．安徽农学院学报，1988，16(6)：14-25．
66．原戈．红松落针病流行规律及最佳防治期的确定．东北林业大学学报，1988，16(6)：14-25．
67．胡能书，万贤国．同工酶技术及应用．湖南科学技术出版社，1985，P1-199．
68．卢怀乡．裂殖酵母属的酶及可溶性蛋白聚丙烯酰胺凝胶电泳．真菌学报，1990，9(1)：50～55．
69. Zervakis G. et al., Taxonomial relationships within the fungal genus Pearotus as determined by isoelectric focusing analysis of enzyme patterns. J Gen Microbiol, 1992, 138: 635—645.
70. Matsuyama N, Moromizoto Z, Ogoshi A, Wakimoto S. Grouping Rhizoctonia solani with nonspecific esterase zymogram. Ann Phytopath Soc Japan, 1978,44: 652～658.
71. Spooner, B. M. Lophodermium and Hypoderma (Rhytismataceae) from Mt Kinabalu, Sabah. Kew Bull, 1991,46 (1): 73-100.
72．方中达．植病研究方法．农业出版社，1979．
73. Durrieu, Guy. Deux Hypodermataceae nouvelles sur Coniferes. Sydowia Beihefte, 1957, 1: 355-358.
74. Yasuo Suto. A new species of Hypoderma causing dieback in Chamaecyparis obtusa and Thuja orientalis. Trans. Mycol. Japan, 1983, 24: 419-424.
75. CMl Description of Pathogenic Fungi and Bacteria No. 943. Mycopathologia, 1988 102: 55-56.
76. Tubuef, Karl von. Demonstration von Lrchenzweigen. Bot. Centralb, 1895, 61: 48-49.
77. Lagerberg. Uber Naevia piniperda Rehm. Svensk Botanisk Tidskr, 1928, 22: 282-288.
78. Hartig, Robert. Wichtige Krankheiten der waldbume. Berlin. J. Springer, 1874, 127pp.
79. Minter, D. W. & Millar, C. W. Lirula nervisequia C. M. I. Descripions of Pathogenic Fungi and Bacteria 660. Commonwealth Mycological Institue, Kew, 783.1884.
80．林英任，侯成林，承河元，刘和．中国北部地区松生斑痣盘菌分类研究．真菌学报，1995，14(2)：92-100．
81．林英任，黎志，梁师文，余盛明．中国北部地区一些针叶树生斑痣盘菌．真菌学报，1995，14(3)：179-183．
82．林英任，唐燕平，刘和云．我国南部地区一些针叶树上的斑痣盘菌．真菌学报，1993，12(2)：93-98．
83．何秉章，杨殿清，齐兴武．红松上的散斑壳．真菌学报，1986，5(2)：70-73．
84．林英任，侯成林．中国北部地区松生斑痣盘菌分类研究．真菌学报，1995，14(2)：92-100．
85. Gordon, C. C. Ascocarpic centrum ontogeny of species of Hypodermatacese of conifers. Am. J. Bot, 1966, 53:319-327.
86. Gordon, C. C. Ascocarpic centrum ontogeny of species of Hypodermataceae of conifers Ⅱ. Am. J. Bot, 1968, 55: 45-52.
87. Staley, J. M. The taxonomy of Lophodermium on pines with special reference to problems in North
American christmas tree plantations. Mitt. BwuiForschAnst. Forst-u. Holzw, 1975, 108: 79-85.
88．侯成林，王有智．针叶树斑痣盘菌科真菌病原调查．林业科学研究，1995，8(4)：426-428．
89. Ouellette, G. B., Magasi, L. P. Lophomerum, a new genus of Hypodermataceae. Mycologia, 1966, 58: 275-280.
90. Rostrup, E. Fortsatte Underogolser over Snyltesvampes Angreb paa Skovtraeome. Tidskr. Skogbr, 1883, 6: 199-300.
91. Tubeuf, Karl yon. Demonstration yon Lrchenzweigen. Bot. Centralb, 1895, 61: 48-49.
92. Birula, I., Z. Dabrowski. and L. Kr6likowski. Gr6zny szkodnik Limby (Pinus cerebra) Hypoderma nova. (Ein geidling der Arve (P. cembra) Hypoderrna Namyslowski sp. nov.). Roezn. Mauk. Rolnicz. Lesn, 1928, 20: 35-41.
93．侯成林．松落针病菌的初步研究．一九九0届硕士学位研究生论文，1990，30-38．
94．何秉章，邓兴林，杨殿清等．樟子松落针病的病原菌和防治的研究．东北林学院学报，1985，13(6)：75-81．
95．李清铣，朱华，王彰明．江苏麦类禾谷镰刀菌酯酶同工酶测定．植物病理学报，1985，15(4)：217-223．
96．徐敬友．六种疫霉菌体可溶性蛋白质和酯酶同工酶的聚丙烯酰胺凝胶垂直平板电泳研究．真菌学报，1993，12(1)：54-64．
97．朱婉华．几种真菌酯酶同工酶聚丙烯酰胺凝胶电泳的初步研究．南京林业大学学报，1987，23(1)：186-190．