中国科技论文在线 http://www.paper.edu.cn A new species of Terriera, a newly recorded genus of () from China Yang Zhongzhou1, Lin Yingren1, Hou Chenglin2 (1. School of Forestry & Landscape Architecture, Anhui Agricultural University, HeFei 230036; 2. College of Life Science, Capital Normal University, Beijing 100048) Abstract: A new member of Terriera, T. huangshanensis on leaves of Eurya muricata var. huiana, is described in this paper. Terriera is also the first record for China. Features of the genus are revised according to relevant documents, The present species is placed in Terriera on the basis of the presence of strongly carbonized extension adjacent to both sides of the ascoma opening and textura angularis-prismatica composed of colorless to lightly brown, somewhat thin-walled cells at the marginal parts of the ascoma, and the absence of lip cells. The new species is similar to T. minor, but their differences are that the latter possesses smaller ascomata with rounded ends, a textura prismatica in the corner between the covering and basal stroma, paraphyses branching 2-3 times in upper 30-40µm, asci ripening sequentially and ascospores tapering towards the ends. The type specimen is deposited in the Herbarium of Forest Fungi of Anhui Agricultural University, China (AAUF). Keywords:taxonomy; ; Theaceae

0 Introduction Terriera B. Erikss. is a member of Rhytismataceae (Rhytismatales, , Ascomycota ) (Kirk et al. 2008). Since Eriksson (1970) established Terriera as a new genus for T. cladophila (Lév.) B. Erikss. (syn. Lophodermium cladophilum Lév.), 17 species and 2 varieties have been reported so far. Species of Terriera are distributed worldwide and occur on angiosperms and gymnosperms. In the past, members of the genus were incorrectly placed in Lophodermium Chevall., Hypoderma De Not., Clithris (Fr.) Bonorden., Dermascia Tehon (Johnston, 2001). Despite the classifications of Höhnel (1917) and Tehon (1935) based heavily on developmental characteristics of the ascomatal primordium were criticized and rejected by later researchers, they are likely to be phylogenetically highly significant for recognizing the genus Terriera as distinct from Lophodermium s. lat. Ortiz-García et al. (2003) noted that ‘Terriera minor differs from Lophodermium in structure of the ascomatal primordium’. Terriera represents a morphologically quite distinct group of species within Lophodermium, and its establishment would be a start to rearranging this heterogeneous genus. Johnston (2001) contributed a monograph on the species of Lophodermium primordially described or reported from monocotyledonous hosts and realized the importance of some characteristics, such as variation in ascus and ascospore size, shape of the ascus apex, shape of the paraphysis apex and features of the conidiomata in the taxonomy of Terriera. Johnston (2001, in Ortiz-García et al. 2003) had transferred many species with typical ascomata structure of Terriera placed in Lophodermium and other genera, such as Clithris arundinacea Penz. & Sacc., Hypoderma fuegianum (Speg.) Kuntze, Lophodermium javanicum Penz. & Sacc. and L. sacchari Lyon etc. to Terriera. Ortiz-García et al. (2003) attempted to explain phylogenetic relationships within Lophodermium by analyzing rDNA-ITS sequences from species of Lophodermium and other rhytismataceous genera. The results showed that Lophodermium from pine hosts formed an independent clade with Meloderma Darker and Elytroderma Darker, not formed a monophyletic sister group to L. piceae (Fuckel) Höhn from spruce plants, and had farther relationships with Terriera.

Foundations: The National Natural Science Foundation of China (No. 30870014 and 30770006); the Specialized Research Foundation for the Doctoral Program of Higher Education of China (No. 20070364002) Brief author introduction:杨中周，（1984-），男，硕士研究生，森林保护学 Correspondance author: 林英任，（1951-），男，教授，微生物学、森林保护学. E-mail: [email protected]

- 1 - 中国科技论文在线 http://www.paper.edu.cn 1 Materials and Methods Mature fruit bodies were selected from the collected specimens. External shapes, size, color and opening ways of ascomata and conidiomata as well as zone line were observed under the dissecting microscope. Than the materials were rehydrated in water for 15min, and 10−15µm thick sections of the fruit bodies were cut using a freezing microtome, and then mounted in lactic acid or cotton blue with pretreatment in water for observing the outlines of ascomata and conidiomata in vertica section. Gelatinous sheaths surrounding ascospores and paraphyses were examined in water or 0.1% (w/v) cotton blue in lactic acid. Color of diversified structures and ascospore contents were observed in water. Measurements and drawings were made using materials mounted in 5% KOH and from 30 asci, ascospores, and paraphyses for each specimen. Line and point integrated illustrations of internal structures of fruit bodies were drawn using the microscopic painting device. 2 Results and Discussions Terriera B. Erikss. Symb. bot. upsal. 19: 58, 1970, a newly recorded genus of China TYPE SPECIES: T. cladophila (Lév.) B. Erikss. Symb. bot. upsal. 19: 59, 1970 ASCOMATA black to gray, elliptical, oblong-elliptical to sublinear or somewhat irregular in surface view, opening by a longitudinal split, variously embedded. LIPS absent. COVERING STROMA composed of dark, thick-walled cells, but along the edge of the ascoma opening is a flattened extension comprising strongly carbonized tissue, adjacent to the covering stroma. Reduced excipulum closely adhering to sides of the extension and the covering stroma. BASAL STROMA composed of dark, thick-walled cells. SUBHYMENIUM with a region of colorless to slightly dark, textura angularis or prismatica at the edge of the ascoma. PARAPHYSES filiform, simple or branched and slightly swollen at the apex, sometimes forming epithecium. ASCI ripening sequentially or seldom synchronously, cylindrical to clavate, with undifferentiated or slightly thickened wall at the apex, J-, usually with a central ostiole, 8-spored. ASCOSPORES hyaline, filiform, often tapered towards the base or both ends, with or without a gelatinous sheath. If present, conidiomata circular to irregular, concolorous with the substrate, but the edge, and the surrounding of the ostiole (s) dark, sometimes the entirety dark in surface view, subcuticular to subepidermal. CONIDIOGENOUS CELLS cylindrical or ampulliform, proliferating sympodially or percurrently. CONIDIA hyaline, cylindrical, elliptical or elongate-elliptical. KNOWN DISTRIBUTION: Brazil, Chile, China, Columbia, India, Indonesia, New Zealand, USA, Venezuela etc. HABITAT: Collected on dead or fallen leaves, and dead twigs or stem (s) of many angiosperms and gymnosperms. NOTES: Eriksson (1970) erected the segregate monotypic genus Terriera, with type species T. cladophila possessing paraphyses swelling at the apex and agglutinating to form a dark epithecium, tissue composed of small, hyaline, ± angular cells at the marginal parts of the ascoma, and lacking labial cells. Terrier (1942) described the paraphyses as ‘simple’, presumably meaning unbranched. The paraphyses type of the same was described by Terrier and Eriksson so different that subsequent taxonomists raised many suspicions about the type species, though in all other respects the descriptions are compatible. Terriera is distinguished from Lophodermium s. str. by the strongly carbonized extension of the covering stroma, the reduced excipulum, the absence of labial cells, the paraphyses often branched or swollen at the apex, and the tissue comprising thin-walled, prismatic or angular cells in the corner between the covering and basal stroma. Hypoderma is similar to Terriera, but differs in the usually well-developed lip cells, the paraphyses swelling rarely at the apex, and the ellipsoidal, cylindrical to clavate ascospores (Cannon & Minter 1986).

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Johnston (2001) pionted out that T. cladophila was collected from Vaccinium in northern temperate regions, but the Terriera species distributed predominantly in tropical areas and spread widely on monocotyledons and dicotyledons, such as Arecaceae, Cyperaceae, Juncaceae, Liliaceae, Pandanaceae, Poaceae etc., and a minority of the genus were conifer-inhabiting species. Terriera huangshanensis Z.Z. Yang, Y.R. Lin & C.L. Hou, sp.nov. FIGURES 1–7 Ascomata amphigena, conspersa, aggregata usque coalescentia, amplitudine admodum variabilia, 550−1900 × 200−550 µm, elliptica, subfusiformia vel lunaria, interdum triloba, nigra, subepidermalia, valde protuberantia, labiis carentia sed structura dehiscenti preformata instructa, e rima longitudinali vel rimis ternis aperientia. Stroma tegens prope apicem 18−22µm crassum necnon extentione maxime carbonacea habens, versus marines ascomatis gradatim spissescens, extensum tenus stromate basali moderate evoluto 10−18µm crasso. Excipulum reductum hymenium cingens. Subhymenium 15−25µm crassum, hyalinum, ex cellulis angulatis et elongatis. Paraphyses filiformes, 120−140µm longae et 1−1.2µm latae, ad apicem plerumque gradatim tumidae vel irregulatim semel aut iterum ramosae, necon epithecio 10−18μm crasso formantes. Asci in simul maturescentes, 100−120 × 5−7µm, anguste cylindrici, stipite 15−28 µm longo, 8-spori. Ascosporae 58−90 × 1.5−2 µm, filiformes, hyalinae, aseptatae, vagina gelatinosa 1−1.5μm crassa indutae. Conidiomata amphigena, conspersa usque coalescentia, 75−130 × 70−90 µm, rotunda vel elliptica, fusca ad atro-brunnea, subcuticularia, apice poro instructa. Cellulae conidiogenae 8−12 × 2−3 μm, cylindricae, gradatim angustatae versus apicem, in sympodialio prolificae. Conidia 4.5−6 × ca.1 μm, cylindrica, hyalina, continua. Lineae zonarum perpaucae, dilute aterae, graciles. ETYMOLOGY: huangshanensis, referring to the place where the specimen was collected. HOLOTYPE: On leaves of Eurya muricata var. huiana (Kobuski) Hu & L.K. Ling (Theaceae), CHINA. ANHUI, Mt Huangshan, Wenquan. alt. ca. 700m, 12 June 2006, Y. R. Lin et al. L2217 (AAUF68325). ASCOMATA developing on both sides of fallen leaves, principally on the upper side of the leaf, scattered to clustered, sometimes two or three confluent, in deadgress or gray-white, 17−25 mm diam., bleached areas. In surface view, ascomata disparate in dimensions, 550−1900 ×200−550 µm, elliptical, fusiform or subfusiform, straight or curved into lunular, sometimes 3-lobed or triangular, ends rounded to subacute, strongly raising above the surface of the substrate at maturity, opening by a single longitudinal split which is branched in the triangular ascomata. Lips absent, split extending almost the whole length of the ascoma. Entirety of ascomata black, no shiny or slightly shiny, the edge defined. In median vertical section, ascomata subepidermal with epidermal cells becoming filled with fungal tissue as ascomata develops, 175−205 µm deep. COVERING STROMA 18−22 μm thick near the centre of the ascoma, slightly thicker towards the edges, extending to the basal stroma, consisting of dark brown to light black, thick-walled textura angularis and globulosa with cells 3.5−5.5 µm diam., along the edge of the ascoma opening is a 12−20 µm thick extension adjacent to the covering stroma which covers the hymenium, and which comprises strongly carbonized tissue with no obvious cellular structure. EXCIPULUM very poorly developed, closely adhering to sides of the extension and the covering stroma. BASAL STROMA 10−18 µm thick, dark brown, consisting of 2−4 rows of 4−7 µm diam., angular, thick-walled cells. SUBHYMENIUM 15−25µm thick, composed of hyaline textura angularis and porrecta, with a region of colorless to grey-brown, 12 −30 µm thick textura angularis-prismatica at the edge of the ascoma. HYMENIUM often going beyond the top of the extension when ripening. PARAPHYSES 120−140 × 1−1.2 µm, filiform, thin-walled, hyaline, branching 1−2 times and slightly swollen at the apex, with a ca.1 μm thick gelatinous matrix, forming distinct epithecium above asci. ASCI ripening synchronously, 100−120 × 5−7µm, narrow-cylindrical, thin-walled and equal, apex rounded or

- 3 - 中国科技论文在线 http://www.paper.edu.cn subacute, not bluing in iodine, discharging spores through a small apical hole, 8-spored, stalk 15−28 µm in length. Ascospores borne in a fascicle, sometimes helically arranged, 58−90 × 1.5−2 µm, filiform, slightly tapered towards the base, hyaline, aseptate, thin-walled, covered by a 1−1.5 µm thick gelatinous sheath. CONIDIOMATA on both sides of leaves, predominantly on the upper side, scattered to crowded, sometimes coalescing. In surface view, conidiomata 75−130 × 70−90 µm, rotund to elliptical, raising the leaf surface, lightly brown but in the regions of the edge and the surrounding of apical ostiole dark brown, in entirety grey-brown or dark brown after discharging spores. In vertical section, conidiomata subcuticular, somewhat lenticular in outline, 50−65 µm deep. UPPER WALL so poorly developed that it is only present in surrounding of the ostiole. BASAL WALL well-developed, 6−8.5 μm, dark brown, consisting of 2−3 rows of 2−3.5 µm diam., angular, slightly thick-walled cells. SUBCONIDIOGENOUS LAYER ca.8µm thick, composed of very light, thin-walled, angular cells. CONIDIOGENOUS CELLS 8−12 × 2−3 μm, cylindrical, slightly tapered towards the apex, hyaline, proliferating sympodially. CONIDIA 4.5−6 × ca.1 μm, cylindrical, hyaline, aseptate. ZONE LINES very infrequent, thin, grey-black or black, rounding or partly rounding the bleached areas. KNOWN DISTRIBUTION: Only from the type locality. HABITAT: T. huangshanensis was collected from dead leaves of Eurya muricata var. huiana in litter. NOTES: T. huangshanensis is miraculous within the genus because of the asci synchronously ripening and the hymenium often overtopping the extension of the covering stroma when mature. This new species is similar in some aspects to T. minor (Tehon) P. R. Johnst. the differences are that the latter possesses ascomata with rounded ends, which don’t curve and aren’t associated with conidiomata and zone lines; a textura prismatica at the marginal parts of the ascoma; paraphyses branching 2−3 times in upper 30−40 µm; asci sequentially ripening; ascospores with 0−1 septate, which slightly taper towards the both ends. Other host plants of the former have not been observed, except Eurya muricata var. huiana in China, while the latter is widely distributed in New Zealand, Chile, Columbia and Venezuela, and could invade 22 genera of monocotyledons and dicotyledons (Johnston 1988, 1989a, 1989b). T. cladophila is distinguished from T. huangshanensis by the ascomata being subcuticular, circular to elliptical and not curved, by the textura prismatica comprising vertically oriented cells in the corner between the covering and basal stroma, by the asci sequentially ripening, by the ascospores not tapering, by the subhymenium consisting of textura angularis and intricata, by the conidiomata up to 100−200µm in diameter, and by the brown and diffuse zone lines (Minter 1996).

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FIGS 1−7. Terriera huangshanense on fallen leaves of Eurya muricata var. huiana. 1. Habit on leaf. 2. Detail of ascomata and conidiomata. 3. Portion of ascoma in median vertical section. 4. Paraphyses and asci. 5. Discharged ascospores. 6. Conidioma in vertical section. 7. Conidiogenous cells and conidia.

3 Acknowledgements We are grateful to Dr P.R. Johnston and Dr D.W. Minter for providing related literatures, to Dr W.Y. Zhuang and Dr Y.C. Dai for giving valuable suggestions, and to Dr. S.J. Wang, Dr. L. Chen and Dr. X.M. Gao for field investigations and micrograph. The study was supported by the National Natural Science Foundation of China (No. 30870014 and 30770006), the Specialized Research Fund for the Doctoral Program of Higher Education of China (No. 20070364002) and the Natural Science Foundation of Anhui Province of China (No. 070411005).

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References

[1] Cannon PF, Minter DW. The Rhytismataceae of the India subcontinent[M]. Mycol. Pap, 1986. 155: 1–123. [2] Eriksson B. On Ascomycetes on Diapensales and Ericales in Fennoscandia[J]. Symb. Bot. Upsal, 1970. 19: 1–71. [3] Höhnel F. von. System der Phacidiales v. H. Berichtr der Deutsche Botanische Gesellschaft[M], 1917. 35: 416–422. [4] Johnston PR. An undescribed pattern of ascocarp development in some non-coniferous Lophodermium species[J]. Mycotaxon, 1988. 31: 383–394. [5] Johnston PR. Lophodermium (Rhytismataceae) on Clusia[J]. Sydowia, 1989a. 41: 170–179. [6] Johnston PR. Rhytismataceae in New Zealand 2. The genus Lophodermium on indigenous plants[J]. New Zealand J. Bot, 1989b. 27: 243–274. [7] Johnston PR. Monograph of the Monocotyledon-inhabiting species of Lophodermium[M]. Mycol. Pap, 2001. 176: 1–239. [8] Kirk PM, Cannon PF, Minter DW, et al. Ainsworth & Bisby’s Dictionary of the Fungi 10th ed[M]. CAB International. Wallingford, 2008. pp771. [9] Minter DW. Terriera cladophila. IMI Descr[J]. Fungi & Bact, 1996. no. 1296. [10] Ortiz-García S, Gernandt DS, Stone JK, et al. Phylogenetics of Lophodermium from pines[J]. Mycologia, 2003. 95: 846–859. [11] Tehon LR. A monographic rearrangement of Lophodermium[M]. Illinois Biological Monographs, 1935. 13: 1–151. [12] Terrier CA. Essai sur la systématique des Phacidiaceae (Fr.) sensu Nannfeldt (1932) [J]. Beitr. Krypt.-Fl. Schweiz, 1942. 9: 1–99.

斑痣盘菌目（子囊菌门）一中国新纪录属 及其一新种 杨中周1，林英任1，侯成林2 （1. 安徽农业大学林学与园林学院，合肥 230036； 2. 首都师范大学生命科学院，北京 100048） 摘要：报道特里尔盘菌属的 1 个新分类单元，即生于毛枝格药柃落叶上的黄山特里尔盘菌 Terriera huangshanensis sp. nov.。这也是该属在中国的首次报道。根据相关文献，该属 的特征被修订，本种被置于 Terriera 属是基于其子囊果开口两边存在一强烈碳化的延伸物， 在子囊果的毗邻处存在着无色至淡褐色的、薄壁的角胞-矩胞组织，且唇细胞缺乏。本种类 似于小特里尔盘菌 T. minor，主要区别在于后者子囊果端部圆形，覆盖层与基部层毗邻处 为角状细胞组织，侧丝在上端 30-40µm 处分枝 2-3 次，子囊陆续成熟，子囊孢子朝着两端渐 细。模式标本存放在安徽农业大学森林菌物标本室（AAUF）。 关键词：分类学；斑痣盘菌科；山茶科 中图分类号：Q939.5

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