Taxonomic Revision of Lepisorus (J. Sm.) Ching Sect. Lepisorus

Journal of Systematics and Evolution 47 (6): 581–598 (2009) doi: 10.1111/j.1759-6831.2009.00056.x

Taxonomic revision of Lepisorus (J. Sm.) Ching sect. Lepisorus (Polypodiaceae) from China 1,2Xin-Ping QI 1Xian-Chun ZHANG∗ 1(State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China) 2(Graduate University of Chinese Academy of Sciences, Beijing 100049, China)

Abstract The taxonomy of Lepisorus (J. Sm.) Ching sect. Lepisorus in China was revised based on herbarium specimen examinations, field observations, and microscopic study of rhizome scales, soral paraphyses, leaf epidermis, and spores. As a result, nine species were recognized: Lepisorus macrosphaerus (Baker) Ching, Lepisorus asterolepis (Baker) Ching, Lepisorus marginatus Ching, Lepisorus kuchenensis (Y.C.Wu) Ching, Lepisorus megasorus (C. Chr.) Ching, Lepisorus kawakamii (Hayata) Tagawa, Lepisorus subsessilis Ching & Y.X. Lin, Lepisorus affinis Ching, and Lepisorus nudus (Hook.) Ching. Lepisorus kawakamii (Hayata) Tagawa was reinstated; Lepisorus gyirongensis Ching & S.K. Wu and Lepisorus longus Ching were reduced to synonyms of L. nudus and L. affinis, respectively. The subdivision of Lepisorus macrosphaerus was not accepted. Rhizome scales and paraphyses are the most useful characters for species delimitation as well as for infrageneric classification. Characteristics of the leaf epidermis and spore ornamentation are usually stable and thus of great significance in understanding the relationships among groups within the genus. Key words China, Lepisorus, Lepisorus sect. Lepisorus, Polypodiaceae, taxonomic revision.

The fern genus Lepisorus (J. Sm.) Ching com- quently, Lepisorus nudus (Hook.) Ching (= Pleopeltis prises approximately 40 species (Hennipman et al., nuda) should be selected as the type of Lepisorus as has 1990; Zink, 1993), with its main distribution in Asia, a been done by Farr et al. (1979) and Zink (1993). When few species in Africa and Madagascar, and one in the Yu and Lin (1997) published their first comphrensive Hawaiian islands. The genus was first subdivided by infrageneric classification of Lepisorus they unfortu- Ching (1933) into two sections based on sorus shape: nately did not accept L. nudus as type, but instead used sect. Pseudovittaria with linear sori (Lepisorus sinen- L. thunbergianus. Therefore, the names of some of their sis and Lepisorus vittarioides), and sect. Eulepisorus sections should be revised. with discrete sori comprising the remaining members Yu and Lin (1997) subdivided Lepisorus into six of the genus. Yu and Lin (1997) published a new in- sections: sect. Lepisorus, sect. Pleioomma S.L. Yu, sect. frageneric classification of Lepisorus with six sections Macrophyllon S.L. Yu, sect. Sclerophyllon S.L. Yu, sect. mainly based on rhizome scales and soral paraphyses. Pachyphyllon S.L. Yu, and sect. Hymenophyton Ching However, later, Lin (2000) divided the genus into only ex S.L. Yu & Y.X. Lin. This system, based on morphol- two sections: sect. Lepisorus and sect. Hymenophy- ogy, seems rather natural and it was partly supported by ton, based on the possession of evergreen or deciduous a phylogenetic analysis based on chloroplast sequence leaves. data (Wang et al., 2009. in press). Lepisorus was first proposed as a section of Dry- Lepisorus sect. Macrophyllon S.L. Yu is one of the naria by Smith (1846). In the protolog only two elements six sections proposed by Yu and Lin (1997). It origi- were included, Drynaria sesquipedalis and Pleopeltis nally contained only three Chinese species, Lepisorus nuda. When Ching raised Lepisorus to a genus he macrosphaerus, Lepisorus asterolepis, and Lepisorus adopted the then well-known species, Polypodium lin- marginatus. All have coriaceous and evergreen leaves, eare Thunb. (= Lepisorus thunbergianus) as the type and deciduous ovate scales, with small and dense lu- which is contrary to the code, because the type should mina. Some species clearly belonging to this section, be chosen from the two elements cited by Smith. Conse- such as Lepisorus affinis Ching and Lepisorus kuchenensis (Y.C. Wu) Ching, were put into sect. Pachy- phyllon S.L. Yu based on their lanceolate scales, with Received: 22 March 2009 Accepted: 7 July 2009 clathrate lumina. However, both of these species with ∗ Author for correspondence. E-mail: [email protected]; Tel: 86-10- evergreen leaves can easily be distinguished from the 62836291; Fax: 86-10-62836291.

C 2009 Institute of Botany, Chinese Academy of Sciences 582 Journal of Systematics and Evolution Vol. 47 No. 6 2009 other species of sect. Pachyphyllon, typified by Lep- sect. Pachyphyllon, and Lepisorus subsessilis Ching & isorus bicolor with deciduous leaves. Recent molec- Y.X. Lin. All of these species are mainly distributed in ular analysis also found L. macrosphaerus, Lepisorus China with L. nudus and L. macrosphaerus extending megasorus, Lepisorus kawakamii and L. nudus form- into the Himalayas. ing one clade with high support (Kreier et al., 2008). The scales of L. macrosphaerus and L. kawakamii are ovate, while those of L. megasorus and L. nudus are 1 Material and methods lanceolate. Although different in shape, these scales are similar in texture and lumina pattern. Based on its char- 1.1 Rhizome scales, leaf scales, and paraphyses acters of scales, paraphyses, leaf epidermis, and spores, For the study of rhizome scales, leaf scales, and L. nudus should be a member of sect. Macrophyllon. paraphyses, herbarium material for examination was Therefore, sect. Macrophyllon is a synonym of sect. taken from the following herbaria: GAUA, GXMI, IBK, Lepisorus which is typified by L. nudus. IBSC, IMC, KUN, PE, PYU, TI, US, and WUK. For The latest taxonomic revision of the Chinese each species, between 20 and 30 samples per charac- species of Lepisorus was published by Lin (2000), and ter were studied and selected from different specimens; the most reliable infrageneric classification was pub- photographs were taken with a Nikon LV-TV camera lished by Yu and Lin (1997). During our revision study, connected to a stereo microscope and computer, and we have studied almost all of the Asian species of Lep- measured using Nikon Act-1. Taxonomical terminol- isorus except some from the Malaysian regions. Our ogy generally followed Lellinger (2002). observations of macromorphology and micromorphol- ogy (such as leaf epidermis and spores) confirmed that 1.2 Leaf epidermis the genus can be naturally subdivided into several sec- Mature leaves were taken from 22 herbarium (PE) tions based mainly on characters of the rhizome scales specimens, representing all nine species of sect. Lep- and soral paraphyses, and in agreement with the classi- isorus (Table 1). Lepisorus scolopendrium (Ching) fication of Yu and Lin (1997) in most part. As a result of Mehra & Bir, belonging to sect. Pachyphyllon,was part of our taxonomic revision of the genus from China, used for comparison because it has similar scales as nine species were recognized in sect. Lepisorus, which L. kuchenensis and always keyed out together in identi- includes all the species of sect. Macrophyllon, part of fication keys in former publications. Material for light

Table 1 Vouchers for leaf epidermis of Lepisorus sect. Lepisorus Taxon Locality Voucher L. macrosphaerus (Baker) Ching Pingwu, Sichuan, China () H.L. Tsiang () 11118 (PE) Lushui, Yunnan, China () X.C. Zhang et. al () 4520 (PE) Medog,ˆ Xizang, China () B.S. Li () 3215 (PE) L. marginatus Ching Wuxi, Chongqing, China () X.P. Qi () 5178 (PE) Ankang, Shaanxi, China ()P.Y.Li() 3199 (WUK) Barkam, Sichuan, China ()X.Li() 72707 (PE) L. asterolepis (Baker) Ching Mt. Emei, Sichuan, China () K.H. Shing () 1643 (PE) Mt. Jinfo, Chongqing, China () X.P. Qi () 4587 (PE) Daguan, Yunnan, China () NE Yunnan Exped. () 1089 (PE) L. kawakamii (Hayata) Tagawa Taihotou, Taiwan, China ()M.Tagawa47(PE) L. megasorus (C. Chr.) Ching Taizhong, Taiwan, China () W. Hancock 31 (PE) L. kuchenensis (Y. C. Wu) Ching Mt. Yaoshan, Guangxi, China ()C.Wang() 40271 (PE) L. subsessilis Ching & Y. X. Lin Fengshan, Guangxi, China ()Z.T.Li() 601709 (PE) Baise, Guangxi, China () Baise Exped. () 01558 (PE) L. nudus (Hook.) Ching Mussoorie, India S.P. Khular 3 (PE) Simla, India X.C. Zhang () 2754 (PE) L. gyirongensis Ching & S. K. Wu Gyirong, Xizang, China () Qinghai-Xizang Complex Exp. (Geobot. Sect.) () 5184 (PE) L. afﬁnis Ching Mt. Wuzhi, Hainan, China () C L. Tso & N.K. Chun (, ) 44067 (PE) Mt. Wuzhi, Hainan, China () X.C. Zhang () 1681 (PE) L. longus Ching Mt. Diaoluo, Hainan, China () Diaoluoshan Exped. ()3302 (PE) Lingshui, Hainan, China () Hainan Exped. ()1925 (PE) Lingshui, Hainan, China () S.Y. Dong () 355 (PE) L. scolopendrium (Ching) Mehra & Bir Binchuan, Yunnan, China () X.P. Qi () Q007 (PE)

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Table 2 Vouchers for spore observations of Lepisorus sect. Lepisorus Taxon Locality Voucher L. macrosphaerus Malipo, Yunnan, China () K.M. Feng () 12714 (PE) Medog,ˆ Xizang, China () B.S. Li et al. () 3215 (PE) L. marginatus Barkam, Sichuan, China ()X.Li() 72707 (PE) Songxian, Henan, China () Henan Exped. () 1898 (PE) L. asterolepis Mt. Emei, Sichuan, China () Ann 815 (PE) Mt. Jinfo, Chongqing, China () X.P. Qi () 4590 (PE) Daguan, Yunnan, China () NE Yunnan Exped. () 491 (PE) L. kawakamii Hualian, Taiwan, China ()T.C.Chen() 9862 (PE) L. megasorus Hualian, Taiwan, China ()T.C.Chen() 12733 (PE) L. kuchenensis Mt. Yaoshan, Guangxi, China ()C.Wang() 40271 (PE) Taizhong, Taiwan, China () Imaseki 70500 (TI) L. subsessilis Fengshan, Guangxi, China ()Z.T.Li() 601733 (PE) L. nudus Milke Danda, Nepal R.L. Fleming 2087 (PE) L. afﬁnis Mt. Wuzhi, Hainan, China () C.L. Tso & N.K. Chun (, ) 44067 (PE) Baisha, Hainan, China () E Hainan Exped. () 512 (PE) L. longus Mt. Diaoluo, Hainan, China () Mt Diaoluo Exped. () 3302 (PE) L. scolopendrium Maguan, Yunnan, China () S.K. Wu () 61–3578 (PE)

microscopy were boiled in water before being macer- L. macrosphaerus (GZ = Guizhou, YN = Yunnan, ated in 35% NaOCl solution. Pieces of epidermis were XZ = Xizang). Vouchers are deposited in PE. prepared then stained in a solution of 1% safranin (in 50% alcohol), and then dehydrated in an ethanol series before being mounted in Canada balsam. The terminol- 2 Results and discussion ogy for epidermis followed Zhang et al. (1999), Kong et al. (2006), and Sun and Zhang (2009). 2.1 Rhizome scales, leaf scales, and paraphyses The rhizome scales of all species of sect. Lepisorus 1.3 Spore morphology are clathrate, translucent with square or rectangular lu- Mature spores of 22 samples were gathered from mina, entire margins, and either ovate or lanceolate out- herbarium specimens in PE and TI (Table 2). Spores line (Fig. 1). Lepisorus macrosphaerus, L. marginatus, were stuck to aluminum stubs with double-sided tape, L. kawakamii, and L. asterolepis have ovate or ovate- sputter-coated with gold, observed and photographed acuminate scales, L. megasorus, L. kuchenensis, L. sub- using a Hitachi S-4800 scanning electron microscope. sessilis, L. nudus, and L. afﬁnis show lanceolate scales. Terms for spore morphology and ornamentation fol- Paraphyses (Fig. 2: A–L) and leaf scales (Fig. 2: M–T) lowed Wang and Yu(2003), Tryon and Lugardon (1991), are clathrate, lumina almost square. The shape of the and Dai et al. (2006). leaf scales varies from round to lanceolate, whereas the paraphyses are rounded in shape, their diameter ranging from 0.3 to 0.8 mm. 1.4 Principal coordinates analysis of the L. macrosphaerus group Table 3 Characters used in principal coordinates analysis for Lepisorus Lepisorus marginatus, L. asterolepis, and macrosphaerus group L. macrosphaerus are three closely related species; L. 1 Laminae shape: (0) linear-lanceolate; (1) lanceolate asterolepis has often been treated as a variety of L. 2 Length of laminae (cm) macrosphaerus. A total of 15 characters were used in 3 Width of laminae (cm) 4 Widest position of leaves: (0) middle; (2) inframedial the analysis, including six binary and nine continuous 5 Venation: (0) hidden; (1) distinct (Table 3). A data matrix used these 15 characters for 6 Length of stipes (cm) each individual operational taxonomic unit (OTU). 7 Arrangement of sori: (0) close; (1) sparse 8 Distribution of sori: (0) extend from upper part to lower part of Ninety-three OTUs out of nine populations from leaf; (1) from upper part to middle part different regions were constructed, and used for 9 Diameter of sori (mm) principal coordinate analysis by MVSP version 3.13b 10 Sori apart from midvein (cm) 11 Sori apart from leaf margin (cm) analysis software. The material used for the statistical 12 Scales shape: (0) ovate-acuminate; (2) ovate analysis includes L. marginatus (BA = Barkam, 13 Length of scales (mm) CK = Chengkou, SNJ = Shennongjia), L. asterolepis 14 Width of scales (mm) (EM = Mt. Emei, HN = Hunan, JF = Mt. Jinfo), and 15 Diameter of paraphyses (mm)

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Fig. 1. Rhizome scales of Lepisorus. A, L. marginatus (L. Xi 72707). B, L. kawakamii (T.C. Chen 12888). C, L. asterolepis (X.P. Qi 4587). D, L. macrosphaerus (B.S. Li et al. 3215). E, L. afﬁnis (X.C. Zhang 1681). F, L. longus (S.Y. Dong 355).G, L. gyirongensis (Qinghai-Xizang Complex Exped. (Geobot. Sect.) 5184). H, L. nudus (R.L. Fleming 2087). I, L. subsessilis (X.C. Zhang 1075). J, L. megasorus (T.C. Chen 12585). K, L. kuchenensis (K.M. Feng 22641). L, L. scolopendrium (X.P. Qi Q007). Scale bars: A–D, 300 μm; E–L, 900 μm.

Plants of sect. Lepisorus share some synapomor- completely clathrate scales with dense, translucent lu- phies. The rhizome scales are normally deciduous when mina different from those in other sections. A tuft of mature (except in L. kuchenensis), and have entire mar- long hairs is always present at the attachment of scales gins and often with opaque or partly transparent or (Fig. 1: C, D, I). The shape and structure of the rhizome

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Fig. 2. Paraphyses and leaf scales of Lepisorus. A–L, Paraphyses. M–T, Leaf scales. A, L. marginatus. B, L. kawakamii. C, L. asterolepis. D, L. macrosphaerus. E, L. longus. F, L. afﬁnis. G, L. gyirongensis. H, L. nudus. I, L. subsessilis. J, L. megasorus. K, L. kuchenensis. L, L. scolopendrium. M, L. marginatus. N, L. asterolepis. O, L. macrosphaerus. P, L. kawakamii. Q, L. longus. R, L. afﬁnis. S, L. subsessilis. T, L. megasorus. Vouchers: A, M, L. Xi 72707; B, P, T.C. Chen 12888; C, N, X.P. Qi 4587; D, O, B.S. Li et al. 3215; E, Q, S.Y. Dong 355; F, R, X.C. Zhang 1681; G, Qinghai-Xizang Complex Exped. (Geobot. Sect.) 5184; H, R.L. Fleming 2087; I, S, X.C. Zhang 1075; J, T, T.C. Chen 12585; K, K.M. Feng 22641; L, X.P. Qi Q007. Scale bar = 100 μm.

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Fig. 3. Leaf epidermis of Lepisorus. Abaxial on the left, adaxial on the right. A, B, L. macrosphaerus (X.C. Zhang 4520). C, D, L. marginatus (X. Li 72707). E, F, L. asterolepis (K.H. Shing 1643). G, H, L. kawakamii (M. Tagawa 47). Scale bar = 100 μm. scales are the main characters for species delimitation. It (Figs. 3–5). Within species the shape of the epider- has been pointed out that the rhizome scales, leaf scales, mal cells was similar on the adaxial and abaxial side. and paraphyses are correlative in evolution (Yu & Lin, The shape of epidermal cells and anticlinal walls was 1996). In sect. Lepisorus the rhizome scales are translu- variable among different taxa, but constant within the cent, and accordingly the lumina of the leaf scales and same taxon; it also showed resemblance among relative paraphyses are translucent too. species. Stomata were only found on the abaxial side in all studied species. Stomatal types were variable: axil- 2.2 Leaf epidermis locytic and coaxillocytic types are the most common, The leaf epidermal cells of sect. Lepisorus were and polocytic, copolocytic, anomocytic, anomotetra- usually irregular in shape, with sinuate anticlinal walls cytic, anisocytic, actinocytic, and amphicycloanisocytic

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Fig. 4. Leaf epidermis of Lepisorus. Abaxial on the left, adaxial on the right. A, B, L. megasorus (W. Hancock 31). C, D, L. kuchenensis (C. Wang 40271). E, F, L. subsessilis (Z.T. Li 601709). G, H, L. nudus (X.C. Zhang 2754). Scale bar = 100 μm. types were also present. The leaf epidermal characters be a function of their maturity and developmental stage of species of sect. Lepisorus are quite different from (van Uffelen, 1985, 1993, 1997). Lepisorus scolopen- those of L. scolopendrium (Fig. 5: G, H). drium can be distinguished from all species of sect. Lepisorus by its verrucate spore surface. In the clas- 2.3 Spore morphology siﬁcation of Yu and Lin (1997), L. scolopendrium, L. All species within sect. Lepisorus have spores with afﬁnis, and L. kuchenensis belong to sect. Pachyphyllon a similar rugate surface ornamentation (Figs. 6, 7). The because of their lanceolate scales. However, the leaves spores of L. marginatus (Fig. 6: C, D) have prominent of L. scolopendrium are deciduous and the latter two are rugae, others are various in rugate degree, which may evergreen. Different leaf types might indicate a more

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Fig. 5. Leaf epidermis of Lepisorus. Abaxial on the left, adaxial on the right. A, B, L. gyirongensis (Qinghai-Xizang Complex Exped. (Geobot. Sect.) 5184). C, D, L. affinis (X.C. Zhang 1681). E, F, L. longus (Diaoluo Shan Exped. 3302). G, H, L. scolopendrium (X.P. Qi Q007). Scale bar = 100 μm. distant relationship, the different spore type of L. affinis JFs representing populations of L. asterolepis form the and L. kuchenensis with that of L. scolopendrium can second group; and GZs, YNs, and XZs representing further sustain that they belong to different groups. populations of L. macrosphaerus form the third group. Group 1 (BAs, CKs, and SNJs), group 2 (EMs, HNs, and 2.4. Principal coordinates analysis for L. JFs), and group 3 (GZs, YNs, and XZs) are clearly sepa- macrosphaerus group rated from each other, but the latter two are closer to each As shown in Fig. 8, the OTUs formed three distinct other. Lepisorus marginatus can be distinguished from groups: BAs, CKs, and SNJs representing populations the latter two by its ovate-acuminate scales, small and of L. marginatus form the first group; EMs, HNs, and subcostular sori, linear-lanceolate leaves, and undulate

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Fig. 6. Spores of Lepisorus.A,B,L. macrosphaerus (K.M. Feng 12714). C, D, L. marginatus (X. Li 72707). E, F, L. asterolepis (X.P. Qi 4590). G, H, L kawakamii (T.C. Chen 9862). I, J, L. megasorus (T.C. Chen 12733). K, L, L. kuchenensis (Imaseki 70500). M, N, L. subsessilis (Z.T. Li 601733). O, P, L. nudus (R.L. Fleming 2087). Scale bars: A, C, E, G, I, K, 20 μm; B, D, F, H, J, L, 5 μm. margin when dry. Lepisorus asterolepis with ovate- leaves. Therefore, it is reasonable to keep L. marginatus, acuminate to ovate, small scales, large medial sori, and L. asterolepis, and L. macrosphaerus as distinct species. lanceolate leaves, L. macrosphaerus with large ovate In our experience, the scales are the most impor- scales, submarginal sori, lanceolate or linear-lanceolate tant character to distinguish the three species. Lepisorus

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Fig. 7. Spores of Lepisorus. A, B, L.afﬁnis (C.L. Tso & N.K. Chun 44067). C, D, L. longus (Diaoluo Shan Exped. 3302). E, F, L. scolopendrium (S.K. Wu 61–3587). Scale bars: A, C, E, 20 μm; B, D, F, 5 μm. macrosphaerus has large ovate scales 2.0–3.0 × 1.0– marginatus shows small, subcostular sori close to each 2.0 mm, L. asterolepis ovate-acuminate to ovate scales other and parallel with the vein. Leaf shape is also a rel- 1.2–2.0 × 0.8–1.5 mm, and L. marginatus ovate- atively useful character as L. macrosphaerus has lance- acuminate scales 0.5–1.5 mm in length and width. How- olate or linear-lanceolate leaves, L. marginatus linear- ever, the sorus position is another important character. lanceolate leaves, and L. asterolepis lanceolate leaves. Lepisorus macropshaerus has submarginal sori, L. as- Because the venation is either visible or hidden, it is terolepis medial, much bigger and sparser sori, and L. not a good character. Consequently, rhizome scales and

Fig. 8. Scatter plot of principal coordinate analysis. BA + CK + SNJ = Lepisorus marginatus;EM+ HN + JF = L. asterolepis;GZ+ YN + XZ = L. macrosphaerus. BA, Barkam, Sichuan; CK, Chengkou, Sichuan; EM, Mt. Emei, Sichuan; GZ, Guizhou Province; HN, Hunan Province; JF, Mt. Jinfo, Sichuan; SNJ, Shennongjia, Hubei; YN, Yunnan Province; XZ, Xizang Province.

C 2009 Institute of Botany, Chinese Academy of Sciences QI & ZHANG: Taxonomic revision of Lepisorus sect. Lepisorus 591 paraphyses are the most reliable characters. It is also interesting to note that these species are vicarious in distribution. diameter (mm)

2.5 General morphology of sect. Lepisorus Table 4 summarizes the morphology of sect. Lep- 1.0–2.00.5–1.50.8–1.5 0.5–0.8 1.0–1.1 0.3–0.5 1.0–1.5 0.4–0.55 1.3–2.0 0.5–0.7 0.8–1.5 0.5–0.8 0.6–1.2 0.4–0.8 1.0–1.5 0.3–0.5 0.4–0.5 0.35–0.5

isorus, based on observations of a large number of spec- × × × × × × × × × imens, particularly the leaf shape and size, the shape and size of the scales, the diameter of the paraphyses, and the position of the sorus. These characters are useful for the identiﬁcation of the species of sect. Lepisorus.

3 Geographical distributions

The species of sect. Lepisorus are distributed mainly in the subtropical to tropical montane forests in southern China from the Yangtze River regions to Hainan and Taiwan. Lepisorus nudus, L. kuchenensis, L. affinis, L. megasorus, and L. kawakamii are rare lo- cal endemic species (Fig. 9), L. affinis is endemic to the tropical rain forest of Hainan, and L. megasorus and L. kawakamii are endemic to Taiwan. Lepisorus marginatus, L. asterolepis, and L. macrosphaerus are the three most common and widely distributed species in China (Fig. 10), L. macrosphaerus extends in the west to northeastern India. These three species are morphologi- cally similar and allopatric. Lepisorus macrosphaerus is mainly distributed in southwest China, L. asterolepis is distributed along the YangtzeRiver, whereas L. marginatus is found mainly to the north of the Yangtze River. It is interesting to note that in the distribution map (Fig. 9), the species with large and thin texture scales are found mainly in the southern areas, whereas the species with small and thick texture scales are found mainly in Lepisorus the northern areas in China. It has been proposed that leaves (cm) leaves (cm) rhizome scales have significant roles such as protection sect. against desiccation, external water storage, and absorp- tion of water and nutrients (Tsutsumi & Kato, 2008). The elevational distribution of sect. Lepisorus Lepisorus ranges from 500 to 3000 m. Lepisorus subsessilis is an endemic species of limestone areas at elevations of 800–1000 m, L. kuchenensis, L. affinis, and L. megasorus usually grow below 2000 m, whereas L. nudus Linear-lanceolateLanceolateLinear-lanceolateLanceolateBroad-lanceolateLanceolateLinear-lanceolateLanceolate 15–22 25–40 1.5–2.0 15–30 20–30 1.0–2.0 Middle 15–40 20–30 1.5–4.0 Middle 4.0–6.0 18–30 1.5–4.0 Inframedial 1.0–2.0 Middle 20–65 1.0–2.5 Near base Middle Medial 1.0–3.0 Middle or inframedial Medial Medial Medial Middle or inframedial Ovate-acuminate Subcostule Submarginal Medial Ovate-acuminate to Ovate-acuminate Subcostule ovate Lanceolate Lanceolate Lanceolate 1.2–2.0 0.5–1.5 Ovate-lanceolate Lanceolate 1.0–1.2 2.5–4.0 2.0–4.0 2.5–4.0 3.0–5.0 2.0–3.0 and L. kawakamii occur at higher elevations from 1500 Lanceolate or linear-lanceolate 20–40 1.5–4.0 Middle or inframedial Submarginal Ovate 2.0–3.0 to 2300 m. The vertical distributions of L. marginatus, L. asterolepis, and L. macrosphaerus are very interest-

ing, and is situated between 700 and 3000 m. Generally Morphological variation within these species are found at lower elevations in the eastern and northern part of their range, but at higher elevations L. marginatus L. asterolepis L. kawakamii L. megasorus L. kuchenensis L. subsessilis L. nudus L. afﬁnis TaxonL. macrosphaerus Leaf shape Length of Width of Widest position Sorus position Scale shape Scale size Paraphysis in the west and southwest. Table 4

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Fig. 9. Distribution of Lepisorus marginatus, L. asterolepis, L. macrosphaerus,andL. kawakamii.

4 Taxonomic treatment Sect. Lepisorus has the closest relationship with sect. Pachyphyllon. Some species in these sections Lepisorus (J. Sm.) Ching sect. Lepisorus. Type: share lanceolate and translucent scales. However, the L. nudus (Hook.) Ching. former has coriaceous and evergreen leaves, and Sect. Macrophyllon S.L. Yu in S.L. Yu & Y.X. Lin the latter shows deciduous and chartaceous leaves. in Acta Phytotax. Sin. 35: 345. 1997. These two sections can also be distinguished by Sect. Pachyphyllon S.L. Yu in S.L. Yu & Y.X. Lin different spore ornamentation and leaf epidermal in Acta Phytotax. Sin. 35: 346. 1997, pro parte. characters. Rhizome thick, long creeping; scales translucent, Nine species in China, lithophytes, or epiphytes. ovate with uniform and dense lumina, or lanceolate Although mainly distributed in southern China, L. with short oblong lumina, more or less darker in the marginatus extends to temperate regions in the Qin- central part and lighter at margin, often falling off ling Mts., and L. macrosphaerus and L. nudus are also when mature; sometimes a tuft of long hairs present present in the Himalayas. at the attachment of scales on rhizome; leaves lance- Key to Lepisorus (J. Sm.) sect. Lepisorus olate or linear-lanceolate, coriaceous, evergreen; leaf scales rounded or lanceolate, with rectangular lumina; 1a. Scales ovate to ovate-acuminate, margin entire, paraphyses rounded, with uniform and clear lumina; lumina square ...... 2 shape of epidermis cells similar on adaxial and abaxial 1b. Scales lanceolate or ovate-lanceolate, margin side with sinuate anticlinal walls; spores have rugate or- entire or denticulate, lumina rectangular, rarely namentation, semicircular or kidney-shaped, zygomor- square ...... 5 phic, 30–50 μm in polar axis length, 60–100 μmin 2a. Sori submarginal; scales ovate, thin, 2.0–3.0 mm equatorial diameter. long...... 1. L. macrosphaerus

C 2009 Institute of Botany, Chinese Academy of Sciences QI & ZHANG: Taxonomic revision of Lepisorus sect. Lepisorus 593

Fig. 10. Distribution of Lepisorus kuchenensis, L. subsessilis, L. nudus, L. afﬁnis,andL. megasorus.

2b. Sori subcostular or medial; scales ovate to ovate- 6a. Leaves ovate-lanceolate, more than 4 cm wide; acuminate, thick, less than 2 mm long ...... 3 scales pale brown...... 6. L. kuchenensis 3a. Sori small, close, medial or subcostular, generally 6b. Leaves lanceolate or linear-lanceolate, 1.0–3.0 cm extending from the upper part to lower part of wide; scales brown ...... 7 leaves; leaves linear-lanceolate, margins undulate 7a. Scale margin yellow-brown, centrally dark brown; whendry...... 2.L. marginatus lumina small and square in central part ...... 3b. Sori large, distant, medial, generally extending ...... 7.L. subsessilis from the upper part to middle part; leaves lance- 7b. Scale margin pale brown, centrally brown; lumina olate to linear-lanceolate, margins not undulate large and rectangular in central part ...... 8 whendry...... 4 8a. Sori costular; leaves less than 30 cm 4a. Leaves 1.5–4.0 cm wide; scales 1.2–2.0 mm long; rhizome 1.5–2.0 mm in diameter long and 1.0–1.3 wide; paraphyses brown ...... 8.L. nudus ...... 3.L. asterolepis 8b. Sori submarginal; leaves up to 65 cm 4b. Leaves 1.0–2.0 cm wide; scales approximately long; rhizome 2.0–4.0 mm in diameter 1.0 mm long and wide; paraphyses pale brown ...... 9.L. afﬁnis ...... 4.L. kawakamii 5a. Scales ovate-lanceolate, margin denticulate; Enumeration of species leaves widest at the base ...... 5. L. megasorus 1. Lepisorus macrosphaerus (Baker) Ching in Bull. 5b. Scales lanceolate, margin entire; leaves widest at Fan Mem. Inst. Biol. 4: 73. 1933; S.L. Yu & Y.X. Lin in the middle or inframedial ...... 6 Bull. Bot. Res. 16 (1): 19. 1996; Y.X. Lin in Fl. Reip.

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Pop. Sin. 6 (2): 64. 2000; Polypodium macrosphaerum ), Qinghai-Xizang Complex Exped. () 13318 (PE); Pingwu ( Baker in Kew Bull. 55. 1895. Type: China. Yunnan ( ), H.L. Tsiang () 11118 (PE); Rongjing (), C.W.Yao ( ): Mengtze (), Hancock 49 (K, PE). ) 2189 (PE); Tianquan ( ), W.Sichuan Exped. ( ) 3257 (PE). Xizang (): Bomi (), T.S. Ying et al. () 91715 (PE); Lepisorus macrosphaerus var. maximus Ching Cona (), Qinghai-Tibet Plateau Group ( ) 2429 (PE); in C.Y. Wu, Fl. Xizang. 1: 303. 1983; Lepisorus Nyingchi (), Qinghai-Xizang Complex Exped. (Suppl. Sect.) ( macrosphaerus f. maximus (Ching) Y.X. Lin in Fl. Reip. ) 2429 (PE); Medogˆ (), B.S. Li et al. () 3034 Pop. Sin. 6 (2): 64. 2000. Type: China. Xizang (): (PE); Zayu(¨ ), T.P. Yi () 79147 (PE). Yunnan (): Cheli Zayu(¨ ), Shang Zayu(¨ ), T. P. Yi () ( ), C.W. Wang ( ) 78259 (PE); Daguan ( ), W.M. Chu et al. () 3318 (PE); Dali (), T.N. Liou () 23106 (PE); 79179 (PE). Deqˆ enˆ (), C.W. Wang () 69184 (PE); Eshan (), S.K. Lepisorus macrosphaerus var. minimus Ching Wu () 292 (PE); Fengqing (), P.I. Mao ( )93(PE); in C.Y. Wu, Fl. Xizang. 1: 304. 1983; Lepisorus Fugong (), H.T. Tsai ( ) 59092 (PE); Gongshan (), C.W. macrosphaerus f. minimus (Ching) Y.X. Lin in Fl. Reip. Wang () 66815 (PE); Hekou (), Z.D. Que ( ) 1231 Pop. Sin. 6 (2): 65. 2000. Type: China. Xizang (): (PE); Heqing ( ), R.C. Ching ( ) 24465 (PE); Huaping ( ), S. Jiang et al. () 6250 (PE); Huize (), S.B. Lan () Zayu(¨ ), Gyigang ( ), T.P. Yi ( ) 79121 135 (PE); Lushui (), Hengduanshan Exped. () 219 (PE); (PE). Jingdong (), T.N. Liou () 52882 (PE); Kunming (), T.N. Fig. 1: D; Fig. 2: D, O; Fig. 3: A, B; Fig. 6: Liou () 19238 (PE); Lanping ( ), Hengduanshan Exped. ( A, B ) 660 (PE); Luquan (), P.I. Mao ( ) 1600 (PE); Maguan Rhizome long-creeping, 2.2–3.0 mm in diameter; ( ), Z.R. Wang ( ) 789 (PE); Malipo ( ), S.Y. Yu ( ) 935 (PE); Mile (), X.P. Qi () Q051 (PE); Pingbian ( scales often deciduous, mainly present at the apex or ), P.I. Mao ( ) 4168 (PE); Shangri-la (), Zhongdian near phyllopodia; scales ovate, 2.0–3.0 mm long, 1.0– Exped. () 1524 (PE); Shuangbai ( ), W.M. Chu et al. ( 2.0 mm wide, basiﬁxed, margin entire, with uniform ) 4078 (PE); Simao (), anonymous 5999 (PE); Songming ( and small thin lumina; leaves 0.5–2.0 cm apart, cori- ), B.Y. Qiu () 54224 (PE); Suijiang (), W.M. Chu ( aceous or soft coriaceous; stipes 3–16 cm long, 1.0– ) 4756 (PE); Weixi ( ), H.T. Tsai ( ) 59864 (PE); Wenshan (), S.L. Yu ( ) 38151 (PE); Wuding (), J.S. Xin ( ) 2.0 mm in diameter; laminae lanceolate, sometimes 44 (PE); Xichou (), Z.R. Wang () 493 (PE); Xinping (), linear-lanceolate, 20–40 cm long, 1.5–4.0 cm wide, H.T. Tsai ( ) 53381 (PE); Yimen (), W.Q. Yin () 213 widest at middle; leaf scales rounded or lanceolate with (PE); Yongping ( ), Yunnan Tropic Pl. Exped. ( clear and hyaline lumina; venation distinct or hidden; ) 976 (PE); Zhenkang (), C.W. Wang () 72621 sori submarginal, 3.0–4.0 mm in diameter; paraphyses (PE). rounded, 0.5–0.8 mm in diameter. 2. Lepisorus marginatus Ching in Fl. Tsinling 2: 184 The types of L. macrosphaerus f. maximus and L. et in Addenda 233. 1974; S.L. Yu & Y.X. Lin in Bull. macrosphaerus f. minimus represent two extreme forms Bot. Res. 16 (1): 20. 1996; Y.X. Lin in Fl. Reip. Pop. Sin. 6 (2): 65. 2000. Type: China. Sichuan (): in leaf size, which do not merit to be named as different taxa. Barkam( ), Songgang ( ), L. Xin ( ) 72707 (PE). China (Gansu, Guangxi, Guizhou, Sichuan, Yun- nan, Xizang), India. Epiphyte on rocks or trunks, alt. Fig. 1: A; Fig. 2: A, M; Fig. 3: C, D; Fig. 1000–3000 m. 6: C, D Additional specimens examined: Rhizome long-creeping, 2.0–2.5 mm in diameter; China. Gansu (): Wenxian (), Q.Z. Li et al. () scales often deciduous, mainly present at the apex or 2311 (PE). Guangxi (): Mt. Damiao (), S.H. Chun () near phyllopodia; scales broadly ovate-acuminate, ap- 14930 (PE); Longlin (), K. Li () 356 (PE); Longsheng (), proximately 0.5–1.5 mm long and wide, base or central Guangfu Forest Exped. () 936 (PE); Xing,an (), Guangxi Exped. () 427 (PE); Mt. Yaoshan (), K.K. Whong ﬁxed, margin entire, with uniform and small lumina, the 443 (PE). Guizhou (): Anlong (), Guizhou Exped. () apex of scales usually dark brown, however, the base 4845 (PE); Anshun (), P.S. Wang () 1074 (PE); Dushan ( brown; leaves 0.2–3.0 cm apart, soft coriaceous, rarely ), Libo Exped. () 1411 (PE); Jiangkou (), C.P. Tsian et al. coriaceous; stipes 1–4 cm long, 1.0–1.5 mm in diam- () 32583 (PE); Kaili ( ), S. Guizhou Exped. () 3972 eter; laminae nearly linear-lanceolate, 15–25 cm long, (PE); Leishan ( ), C.P.Tsian et al. () 51116 (PE); Pu’an ( ), Anshun Exped. () 1382 (PE); Songtao (), Sino-America 1.5–2.0 cm wide, sometimes to 3.0 cm wide, middle Guizhou Pl. Exped. () 2298 (PE); Xingren (), Guizhou widest, undulate margin when dry; leaf scales rounded Exped. () 779 (PE); Yinjiang ( ), N. Guizhou Exped. () or lanceolate, lumina dark brown, hyaline; venation hid- 1136 (PE). Sichuan (): Butuo (), Sichuan Econ. Pl. Exped. ( den; sori medial or subcostular, 2.5–4.0 mm in diameter; ) 1055 (PE); Dujiangyan (), Z.T. Wang et al. ( paraphyses deciduous, rounded in shape, 0.3–0.5 mm in ) 870196 (PE); Ganluo (), anonymous 3996 (PE); Hongxi (), C.T. Kuan () 6970 (PE); Luding (), W.Sichuan Exped. ( diameter. ) 1678 (PE); Mabian (), W.P. Fang () 6479 (PE); Miyi ( China (Chongqing, Gansu, Hebei, Henan, Hubei, ), Qinghai-Xizang Complex Exped. () 11789 (PE); Muli ( Shaanxi, Shanxi, Sichuan). On rocks, alt. 900–2800 m.

C 2009 Institute of Botany, Chinese Academy of Sciences QI & ZHANG: Taxonomic revision of Lepisorus sect. Lepisorus 595

Additional specimens examined: Rhizome long-creeping, 1.8–3.0 mm in diameter; China. Chongqing (): Chengkou (), T.L. Dai () scales often fall off, ovate-acuminate to ovate in shape, 104872 (PE); Wushan (), G.H. Yang ( ) 59636 (PE); Wuxi (), G.H. Yang ( ) 58407 (PE). Gansu (): Ankang (), 1.2–2.0 mm long, 0.8–1.5 mm wide, base ﬁxed, entire P. Y. L i ( ) 7698 (PE); Kangxian (), W.Y. Hsia () 6320 margin, lumina uniform and small; the center of scales (PE); Tianshui (), Z.W. Zhang ( ) 225 (PE); Wenxian (), usually darker colored with thick atro-brown walls, how- Z.Y. Zhang () 11220 (PE); Wudu (), Z.Y. Zhang () ever, margin much lighter; leaves 0.5–2.0 cm apart; 5577 (PE); Wushan (), Yellow River Exped. () 476 (PE); stipes 5–15 cm long, 1.0–2.0 mm in diameter; lami- Zhouqu ( ), X.C. Zhang et al. () 1771 (PE). Hebei (): Fuping ( ), C.H. Wang 900 (PE). Henan (): Luanchuan (), nae lanceolate, coriaceous, sometimes soft coriaceous, M. Liu () H40087 (PE); Lushan (), anonymous 18566 (PE); 15–30 cm long, 1.5–4.0 cm wide, generally inframedial Lushi ( ), anonymous 34589 (PE); Xixia ( ), Henan Exped. ( widest; leaf scales rounded or lanceolate, lumina brown, ) 1161 (PE); Yiyang ( ), anonymous 21660 (PE). Hubei (): hyaline; venation undistinct; sori medial, 3–4 mm in di- Danjiangkou (), J.Q. Xing ( ) 15003 (PE); Shennongjia ( ameter, distributed from upper part to middle part of

), Shennongjia Pl. Exped. ( ) 31074 (PE); Xingshan (), Q.L. Chen ( ) 2075 (PE); Zhuxi (), P.Y. Li () leaves; paraphyses rounded, 0.4–0.55 mm in diameter. 9348 (WUK). Shaanxi (): Chang,an (), Y.T. Hsieh ( ) Lepisorus macrosphaerus, L. margiantus, and L. 001 (WUK); Danfeng (), J.X. Yang et al. ( ) 3027 (WUK); asterolepis are three closely related species. Lepisorus Fengxian (), K.J. Fu () 12800 (WUK); Foping (), K.J. macrosphaerus with thin and large ovate scales, sub- Fu () 4663 (WUK); Huashan ( ), W.Y. Hsia ()s.n. marginal sori, and L. marginatus with thick and small (WUK); Huayin ( ), X.P. Wang ( ) 20681 (WUK); Huxian (), P.C. Kuo ( ) 666 (WUK); Lantian (), G.X. Su ( ovate-acuminate scales, sori small, medial or subcos- ) 430 (WUK); Lueyang ( ), anonymous L12 (WUK); Meixian tular and close to each other, and L. asterolepis with (), T.P. Wang () 19909 (WUK); Ningshan ( ), Y.P. Xu ovate-acuminate to ovate scales, sori large, medial, and ( ) 819 (WUK); Pingli (), G.Y. Bai ( ) 016 (WUK); discrete from each other. Shanyang ( ), Z.Y. Zhang () 15991 (WUK); Shiquan ( ), China (Anhui, Chongqing, Fujian, Guizhou, J.Q. Xing () 8846 (WUK); Weinan ( ), T.P. Wang () 15745 (WUK); Xunyang ( ), Shaanxi Med. Pl. Exped. ( Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Yunnan, Zhe- ) 4054 (WUK); Yangxian ( ), J.X. Yang ( ) 1465 (WUK); jiang). On rocks, alt. 700–2200 m. Zhashui ( ), J.X. Yu () 1075 (WUK); Zhenping (), G.Y. Additional specimens examined: Xu et al. ( ) 5818 (WUK); Zhouzhi ( ), Y.P. Xu ( ) China. Anhui ( ): Mt. Huangshan (), T.W. Liou et al. ( 6729 (WUK). Shanxi (): Yicheng (), Yellow River Exped. ( ) 3082 (PE); Qingyang ( ), Statio Orientali-Sinensis ( ) 148 (PE); Yongji( ), T. W. Liu () 0283 (PE); Yuncheng ) 6014 (PE); Xiuning ( ), anonymous 2333 (PE). Chongqing (), Yellow River Exped. () 477 (WUK). Sichuan (): (): Chongqing (), X.Z. Deng et al. () 0353 (PE); Mt. Baoxing ( ), T.P. Soong ( ) 39131 (PE); Barkam (), X. Jinfo (), M.G. Li ( ) 61818 (PE). Guizhou (): Bijie ( Li () 72707 (Type, PE); Danba (), S. Jiang et al. () 9718 ), P.H. Yu ( ) 721 (PE); Kaili ( ), S. Guizhou Exped. ( (PE); Fengjie ( ), H.F. Zhou et al. ( ) 108739 (PE); Heishui ) 3851 (PE); Guiding (), Y.Tsiang () 5419 (PE); Yinjiang ( (), X. Li () 73324 (PE); Jinchuan (), anonymous 0074 ), T.S. Ying et al. () 32203 (PE). Fujian ( ): Wushan ( (PE); Kangding (), K.C. Kuan et al. () 113 (PE); Lixian ), anonymous s. n. (PE). Hubei (): Hefeng ( ), H.J. Li ( (), X. Li () 46670 (PE); Luding (), H.S. Kung () ) 5880 (PE); Xuan’en (), H.J. Li () 3611 (PE). Hunan ( 6289 (PE); Maoxian (), Z. He et al. ( ) 14234 (PE); Xikang ): Dawei Shan (), J. M. Xi ( ) 7488 (PE); Hongjiangshi (), K.L. Chu ( ) 7011 (PE). ( ), X.L. Zhou ( ) 1767 (PE); Longshan (), L.H. Liu ( 3. Lepisorus asterolepis (Baker) Ching in Fl. Jiangsu ) 16027 (PE); Pingjiang (), S.X. Qi () 47 (PE); Qianyang ( ), anonymous 1348 (PE); Sangzhi (), S.F. Wu ( ) 1056 (Jiangsu Zhi Wu Zhi) 1: 74, plate 112. 1977; S.L. (PE); Shimen (), S.F. Wu ( )s.n.(PE);Wugang(), L.H. Yu & Y.X. Lin in Bull. Bot. Res. 16 (1): 21. 1996; Liu et al. () 16027 (PE). Jiangxi (): Jiujiang (), Y.H. Y.X. Lin in Fl. Reip. Pop. Sin. 6 (2): 65. 2000; Poly- Yan et al. ( ) 3208 (PE). Lushan ( ), Y.G. Xiong ( podium asterolepis Baker in Journ. Bot. 26 (7): 230. ) 9936 (PE); Luxi ( ), L. Wu ( ) 1319 (PE); Yanshan (), 1888; Polypodium excavatum var. asterolepis C. Chr Jiangxi Exped. () 1166 (PE). Sichuan (): Daxiangling ( ), H.S. Kung () 4195 (PE); Dujiangyan (), X.C. Zhang in Index Filic. 511. 1906; Polypodium macrosphaerum () 0557 (PE); Mt. Emei (), G.H. Yang ( )s.n.(PE); var. asterolepis C. Chr in Acta. Hort. Goteberg. 1: Fengjie ( ), H.F. Zhou ( ) 26390 (PE); Hanyuan ( ), T.P. 101. 1924; Lepisorus macrosphaerus var. asterolepis Wang () 8835 (PYU); Leibo ( ), Q.S. Zhao () 1316 (Baker) Ching in Bull. Fan Mem. Inst. Biol. 4: 74. (PE); Luding (), L. Liu et al. () 903 (PE); Pingshan (), 1933. Type: China. Sichuan (): Mt. Emei, E. Faber T.T. Yu ( ) 3523 (PE); Qiuyang ( ), S.Y. Hou ( ) 786 (PE); Tianquan (), H.L. Tsiang () 34850 (PE); Wenchuan ( 1063 (K). ), K.Y. Lang et al. ( ) 1394 (PE); Ya‘an (), S. Jiang et al. Lepisorus longipes Ching & Z.Y. Liu in Bull. Bot. () 10062 (PE). Yunnan (): Daguan (), W.M. Chu ( Res. 4 (4): 25. 1984. Type: China. Chongqing (): ) 5278 (PYU); Yiliang ( ), NE Yunnan Exped. () 831 Nanchuan (), Mt. Jinfo (), Z.Y. Liu ( (PE); Xuanwei (), G.F. Zhang ( ) 22075 (PYU); Zhenxiong ) 4033 (IMC, PE). (), NE Yunnan Exped. () 1098 (PE). Zhejiang ( ): Anji (), X.Y. He () 24224 (PE); Changhua (), X.Y. He Fig. 1: C; Fig. 2: C, N; Fig. 3: E, F; Fig. 6: () 23108 (PE); Suichang (), P.S.Chiu et al. ( ) 5675 E, F (PE); Mt. Tianmu (), X.Y. He () 22304 (PE).

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4. Lepisorus kawakamii (Hayata) Tagawa in Acta Phy- Lepisorus megasorus and L. kawakamii are en- totax. Geobot. 5: 109. 1936. Devol in Fl. Taiwan 1: demic to Taiwan. They were treated as one species by 489. 1994; Polypodium kawakamii Hayata in Bot. Mag. Ching (1933) and Lin (2000). We observed more spec- Tokyo. 23: 77. 1909. Type: China. Taiwan (), imens of these two species and found they can be easily Toroku: Nanshikiaku, Kawakamii & Mori 2346 (TI); distinguished from scales and paraphyses, so we rein- Arizan, Nakahara s. n. (?TI). state L. kawakamii as a distinct species. Lepisorus megasorus (C. Chr.) Ching in Bull. Fan Endemic to Taiwan. On trunks, alt. 500–1800 m. Mem. Inst. Biol. 4: 83. 1933, pro parte; Y.X. Lin in Fl. Additional specimens examined: Reip. Pop. Sin. 6 (2): 72. 2000, pro parte. China. Taiwan (): Hualian (), T.C. Chen () 12339, 12661, 12585, 12733 (PE), C.M. Kuo () 128 (PE); Nantou ( ) Fig. 1: B; Fig. 2: B, P; Fig. (), D.Z. Fu () 96028 (PE); Nantou (), Y.C. Liu () 3: G, H; Fig. 6: G, H 3635 (PE); Sanwenquan ( ), N. Inagaki s. n. (PE); Taizhong ( Rhizome long-creeping, 2.0–3.0 mm in diameter; ), W. Hancock 31 (PE, US), Yamamoto 2853 (PE); Xinzhu (), G. scales often deciduous, mainly present at the apex or Kokubugata 4104 (TI). near phyllopodia; scales ovate-acuminate, 1–1.2 mm 6. Lepisorus kuchenensis (Y.C.Wu) Ching in Bull. Fan long, 1.0–1.1 mm wide, basiﬁxed, margin entire, with Mem. Inst. Biol. 4: 69. 1933; Y.X. Lin in Fl. Reip. Pop. uniform and small lumina; leaves 0.5–1.5 cm apart, Sin. 6 (2): 72. 2000; Polypodium kuchenense Y.C. Wu coriaceous or soft coriaceous; stipes 7–15 cm long, in Bull. Dept. Biol. Sun Yatsen Univ. 3: 276, pl. 129. 1.5–2.0 mm in diameter; laminae linear-lanceolate, 25– 1932. Type: China. Guangxi (): Mt. Yaoshan ( 40 cm long, 1–2 cm wide, middle or inframedial widest; ), Kuchen (), S.S. Sin () 2121 (SYS?), leaf scales rounded or lanceolate; venation hidden; sori 8980 (PE). medial, sometimes near margin, 3.0–4.5 mm in diame- Fig. 1: K; Fig. 2: K; Fig. 4: C, D; Fig. 6: ter, paraphyses 0.5–0.7 mm in diameter. K, L Endemic to Taiwan Island. On trunks, alt. 1800– Rhizome long-creeping, oblate when dry, as wide 2200 m. as 3.0–5.0 mm, with not easily detachable scales; scales Additional specimens examined: pale brown, lanceolate, 3.0–5.0 mm long, 1.3–2.0 mm China. Taiwan (): Hualian (), T.C. Chen () 9862, wide; leaves 1–2 cm apart, thin and soft in texture; stipes 12888 (PE), T. Shimizu 11866 (TI); Mt. Ali (), S. Sasaki s. n. 3–10 cm long, 1.0–2.0 mm in diameter; laminae wide- (TI); Mt. Taiha, S. Suzuki s. n. (PE); Yilan ( ), C.C. Chuang 4157 (TI); Taizhong (), S. Okamoto s. n. (TI), H. Ito s. n. (TI). lanceolate, 20–30 cm long, 4–6 cm wide, middle widest, 5. Lepisorus megasorus (C. Chr.) Ching in Bull. Fan sometimes with long narrow tip; leaf scales not easily Mem. Inst. Biol. 4: 83. 1933; Devol in Fl. Taiwan 1: 491. seen; venation more or less distinct; sori subcostular, 1994; Y.X. Lin in Fl. Reip. Pop. Sin. 6 (2): 72. 2000. excl. 4–6 mm in diameter; paraphyses round, 0.4–0.8 mm in syn. P. kawakamii Hayata; Polypodium megasorum C. diameter. Chr in Index Filic. 544. 1906, nom. nov. for Polypodium A species distinct from others by the broad- macrosorum Baker in Journ. Bot. 23: 106. 1885, non Fee lanceolate leaves, large medial sori, large and pale (1850–52). Type: China. Taiwan (), W.Hancock 31 brown scales. (holotype, K; isotype, US). China (Guangxi, Guizhou, Yunnan, Taiwan). On Polypodium hypochrysum Hayata in Ic. Pl. Form. cliff or trunks in humid forest, alt. 1200–1700 m. Additional specimens examined: 5: 314, f. 127 A, B. 1915; C. Chr. in Index Filic. Suppl. China. Guangxi (): Jinxiu ( ), Dayaoshan Exped. ( 2: 26. 1917. Type: China. Taiwan (), Giran, T. ) 12874 (IBSC); Lingyun (), anonymous 64 (PE); Mt. Kawakami & S. Fujii s.n. (TI). Yaoshan (), S.S. Sin () 8980 (Type, PE), C. Wang () Fig. 1: J; Fig. 2: J, T; Fig. 4: A, B; Fig. 40271 (PE, IBSC); Guizhou (): Xingren (), Guizhou Exped. 6: I, J () 8600 (PE). Taiwan (): Taizhong (), Imaseki 70500 (TI). Yunnan (): Malipo (), K.M. Feng () 22641 Rhizome long-creeping, 2–4 mm in diameter; (KUN); Pingbian (), Sino-Russia Yunnan Exped. ( scales often fall off, ovate-lanceolate, 2.0–4.0 mm long, ) 3548 (PE), 3906 (PE, KUN), K.M. Feng () 5086 (PE, KUN); approximately 1.0–1.5 mm wide, a very rare scale shape, Xichou (), S.K. Wu () 4231 (PE), anonymous 62-K031 with wide ovate base and long acuminate apex, basi- (KUN), K.M. Feng () 11929 (PE, KUN), Q.A. Wu () ﬁxed, pale brown margin with small denticula; leaves 7285 (KUN). 0.5–2.0 mm apart, coriaceous or soft coriaceous; stipes 7. Lepisorus subsessilis Ching & Y.X. Lin in Y.X. 5–25 cm long, 1–2 mm in diameter; laminae lanceolate, Lin, Fl. Reip. Pop. Sin. 6 (2): 347. 2000. Type: China. 15–40 cm long, 1.5– 4.0 cm wide, near base widest; leaf Guangxi ( ): Fengshan ( ), Jingu ( ), Nahuai scales rounded or lanceolate, lumina brown and hyaline; ( ), Z.T. Li ( ) 601709 (PE). venation hidden; sori medial, 3.0–3.5 mm in diameter; Fig. 1: I; Fig. 2: I, S; Fig. 4: E, F; Fig. 6: paraphyses detachable, 0.5–0.8 mm in diameter. M, N

C 2009 Institute of Botany, Chinese Academy of Sciences QI & ZHANG: Taxonomic revision of Lepisorus sect. Lepisorus 597

Rhizome long-creeping, 2.0–2.5 mm in diame- physus are identical to these of L. gyirongensis. Unfor- ter; scales often deciduous, only present at the apex tunatly there are no mature spores available for com- or near phyllopodia; scales lanceolate, 2.5–4.0 mm parison. This species can be distinguished from others long, 0.8–1.5 mm wide, basifixed, margin entire and by the subcostule sori and lanceolate and translucent yellow-brown, central part dark brown, with nearly iso- scales. diameter lumina; leaves 0.5–1.5 mm apart, thick co- China (Yunnan, Xizang), India, Nepal, Sikkim and riaceous, pale green when fresh; stipes 2–6 cm long, Thailand. On rocks or trunks, alt. 1500–2300 m. 1–2 mm in diameter; lamina lanceolate, 18–30 cm long, Additional specimens examined: 1.0–2.5 cm wide, middle or inframedial widest, with China. Yunan (): Menghai ( ), W.M. Chu et al. ( ) 6830 (PYU). Xizang (): Gyirong (), Qinghai-Xizang gradual narrower apex; leaf scales rounded or narrow- Complex Exp. (Geobot. Sect.) () 4555 (PE), 5184 (Type, lanceolate, lumina brown, hyaline; venation hidden; PE). sori medial, 2–3 mm in diameter; paraphyses rounded, India. Mussoorie, S.P.Kullar 3 (PE); 110 (PE); 116 (PE); 118 (PE); with thick walled hyaline lumina, 0.3–0.5 mm in 124 (PE); 133 (PE); Simla, X. C. Zhang () 2754 (PE). Nepal. diameter. Kathmandu, H. Hara et al. 725526 (TI); Milke Danda, R.L. Fleming 2087 (PE); Shemma, H. Ohashi et al. 3187 (TI). Sikkim. S.P. Kullar 3 A rare species endemic to the limestone areas, (PE). scales with brown margin and darker central part, but 9. Lepisorus affinis Ching in Ching & Chu H. Wang this species with coriaceous and evergreen leaves can in Acta Phytotax. Sin. 8: 152, pl. 22, f. 28. 1959; Y.X. be distingushied from species of sect. Pachyphyllon by Lin in Fl. Reip. Pop. Sin. 6 (2): 79. 2000. Type: China. its bicolor scales. Hainan (): Mt. Wuzhi (), Fanye (), C.L. China (Guangxi, Guangdong). On limestone rocks, Tso () & N.K. Chun () 44067 (PE). alt. 800–1000 m. Lepisorus longus Ching in Acta Phytotax. Sin. 10: Additional specimens examined: China. Guangdong (): Huaiji (), W.T. Tsang ( ) 192. 1965; Y.X. Lin in Fl. Reip. Pop. Sin. 6 (2): 77. 23368 (IBSC). Guangxi (): Debao (), Baise Exped. () 2000; Lepisorus longifolius Ching in Ching & Chu H. 01558 (PE); Jingxi (), X.C. Zhang () 1075 (PE); Fengshan Wang in Acta Phytotax. Sin. 8: 170. 1959, non Holttum (), Z.T. Li () 601709 (Type, PE), 601733 (PE); Bama (), 1955. Type: China. Hainan (): Baoting (), Mt. R.C. Ching () 6670 (PE); Napo (), H.G. Zhou ( ) 2489 Diaoluo (), Xin’an Village (), Diaoluo (GAUA); 2490 (GAUA); 2491 (GAUA, PYU). Shan Exped. () 3302 (PE, IBSC). 8. Lepisorus nudus (Hook.) Ching in Bull. Fan Mem. Fig. 1: E; Fig. 2: F, R; Fig. 5: C, D; Fig. Inst. Biol. 4: 83. 1933; Pleopeltis nuda Hook. in Exot. 7: A, B Fl. pl. 63. 1823; Polypodium nudum Kunze in Linnaea Rhizome long-creeping, 2–4 mm in diameter; 23: 281. 1850; Drynaria nuda Fee,´ Gen. Fil 270. 1850– scales often deciduous; scales lanceolate, 2.5–4.0 mm 52; Phymatodes nuda J. Sm., Bot. Voy. Herald. 425. long, 1.0–1.5 mm wide, basifixed, margin entire, with 1857. Type: Nepal. Wallich s. n. (K). oblong luminae; leaves 0.5–2.0 cm apart, coriaceous, Lepisorus gyirongensis Ching & S.K. Wu in C.Y. sometimes soft coriaceous; stipes 2–15 cm long, 1.5– Wu, Fl. Xizang. 1: 304. 1983. Type: China. Xizang ( 2.0 mm in diameter; laminae lanceolate or linear- ): Gyirong (), Qinghai-Xizang Complex Exp. lanceolate, 20–65 cm long, 1–3 cm wide, middle or in- (Geobot. Sect.) () 5184 (PE). framedial widest; leaf scales round or lanceolate, lamina () Fig. 1: H; Fig. 2: H; Fig. 4: G, H; brown, hyaline; venation obscure, sori submarginal, 3– Fig. 6: O, P 4 mm in diameter; paraphyses detachable, 0.35–0.5 mm Rhizome long-creeping, 1.5–2.0 mm in diameter; in diameter. scales lanceolate, 2.0–3.0 mm long, 0.6–1.2 mm wide, Lepisorus longus is distinguished from L. affinis by basifixed, margin entire, with oblong lumina; leaves 0.5– its narrow leaves. The widths of leaves are variable even 1.0 cm apart, soft coriaceous or coriaceous; stipes 1– on one rhizome, but the rhizome scales, epidermis, and 5 cm long, 0.8–1.0 mm in diameter; laminae lanceolate, spore ornamentation have very little difference. 20–30 cm long, 1.0–2.0 cm wide, middle widest; leaf Endemic to Hainan Island. On trunks, alt. 900– scales hardly seen; venation hidden, sori costular, 2.0– 1700 m. 2.5 mm in diameter, extend to inframedial of leaves; Additional specimens examined: paraphyses rounded, 0.4–0.55 mm in diameter. China. Hainan (): Baisha (), S.K. Lau ( ) 26321, Lepisorus gyirongensis was published by Ching 26362 (PE), X.C. Zhang () 2827 (PE), E. Hainan Exped. ( and S.K. Wu (1983), with only two collections. Chan- ) 512 (PE); Changjiang (), X.C. Zhang () 1582 (PE); dra (2000) treated this species as a synonym of L. Lingshui (), S. Y. Dong () 475 (PE), X. C. Zhang () 1729 (PE), Hainan Exped. () 1903 (PE), Diaoluoshan Exped. ( nudus. We have studied a lot of specimens of L. nudus ) 3200 (PE), 3302 (PE, IBSC), F.C. How ( ) 75412 (PE), from India and Nepal and found the scales and para- Hainan Exped. () 1929 (PE); Qiongzhong (), S.Y. Dong (

C 2009 Institute of Botany, Chinese Academy of Sciences 598 Journal of Systematics and Evolution Vol. 47 No. 6 2009

) 824 (PE), 236–6 Exped. 01756 (PE); Mt. Wuzhi (), X.C. Lellinger DB. 2002. A modern multilingual glossary for taxo- Zhang et al. () 1650, 1679, 1681 (PE); Wuzhishan Fern Survey nomic pteridology. Pteridologia 3: 1–264. 151, 239, 630 (PE), C.L. Tso () & N.K. Chun () 44067 Lin Y-X.2000. Lepisorus. In: Flora Republicae Popularis Sinicae (Type, PE). (). Beijing: Science Press. 6(2): 43–93. Tsutsumi C, Kato M. 2008. Morphology and evolution of epi- Acknowledgements This study was supported by the phytic Davalliaceae scales. Botany 86: 1393–1403. National Natural Science Foundation of China (Grant Smith J. 1846. An enumeration of ferns cultivated at Kew, in Nos. 30470139 and 30770166). The authors of this pa- December 1845; with characters and observations on some of the genera and species. Botanical Magazine (Comp.) 72: per are grateful to the curators of the herbaria PE, KUN, 13. PYU, IMC, IBK, GXMI, GAUA, WUK, US, and TI for Sun Z-Y (), Zhang X-C (). 2009. Observation on checking specimens or sending specimens on loan. The the leaf epidermis of medicinal plants of Lepisorus from authors also thank R. VIANE for critical reading of the China. Chinese Bulletin of Botany ( ) 44(3): 282– manuscript and improving the English. 289. Tryon AF, Lugardon B. 1991. Spores of the Pteridophyta. New York: Springer-Verlag. 316–317. References Yu S-L ( ), Lin Y-X( ). 1996. Research on taxonomy of genus Lepisorus (Smith) Ching in China. Bulletin of Chandra S. 2000. The ferns of India. Dehra Dun: International Botanical Research () 16(1): 3–31. Book Distributors. 377–386. Yu S-L ( ), Lin Y-X( ). 1997. A study on systemat- Ching R-C (). 1933. The studies of Chinese ferns IX. ics of the genus Lepisorus (Polypodiaceae). Acta Phytotax- Bulletin of the Fan Memorial Institute of Biology ( onomica Sinica ( ) 35: 341–347. ) 4: 47–116. van Uffelen GA. 1985. Synaptospory in the fern genus Pyrrosia Ching R-C (). 1983. Lepisorus. In: Flora Xizangica ( (Polypodiaceae). Blumea 31: 57–64. ). Beijing: Science Press. 1: 300–312. van Uffelen GA. 1993. Sporogenesis in Polypodiaceae (Filiclaes). Dai X-L (), Cao J-G ( ), Wang Q-X (), III. Several species. Spore characters and their value in phy- Zhu R-L ( ). 2006. The structure and development logenetic analysis. Blumea 37: 529–561. of sporoderm of Lepisorus thunbergianus (Kaulf.) Ching van Uffelen GA. 1997. The spore wall in Polypodiaceae: (Polypodiaceae). Bulletin of Botanical Research ( development and evolution. In: Johns RJ ed. Holttum ) 26(5): 545–550. Memorial Volume. Kew: Royal Botanic Gardens. 95– Farr ER, Leussink JA, Stafleu FA. 1979. Index Nominum (Plan- 117. tarum). Vol. II (Regnum Vegetabile Vol. 101). Utrecht: Wang L (), Qi X-P (), Xiang Q-P ( ), Hein- Bohn, Scheltema & Holkema. richs J, Schneider H, Zhang X-C (). 2009. Phylogeny Hennipman E, Kramer KU, Veldhoen P. 1990. Polypodiaceae. In: of the paleotropical fern genus Lepisorus (Polypodiaceae, Kubitzki K ed. The families and genera of vascular plants. Polypodiopsida) inferred from four chloroplast DNA re- Berlin: Springer-Verlag. 1: 203–230. gions. Molecular Phylogenetics and Evolution (in press). Kong Y-N (), Liu X ( ), Ge Y (), Liu Q-R ( Wang Q-X (), Yu J ( ). 2003. Classification of spore ), Zhou Y-L ( ), Gai Q ( ). 2006. Stomatal ornamentation in filicales under SEM. Acta Botanica Yun- apparatus of ten species in the genus Lepisorus and its sys- nanica () 25(3): 313–320. tematic significance. Journal of Beijing Normal University Zhang Y-J( ), Yu H-F ( ), Lu Y-X ( ), Li H-D (Natural Science) ( ( )) 42: (). 1999. Stomatal apparatus of Chinese Polypodi- 291–294. aceae and its systematic significance. Journal of Lanzhou Kreier HS, Zhang X-C, Muth H, Schneider H. 2008. The mi- University (Natural Science) ( )( crosoroid ferns: inferring the relationships of a highly di- ) 35(1): 130–139. verse lineage of paleotropical epiphytic ferns (Polypodi- Zink MJ. 1993. Systematics of the fern genus Lepisorus (J. aceae, Polypodiopsida). Molecular Phylogenetics and Evo- Smith) Ching (Polypodiacea-Lepisoreae). Ph.D. Disertation. lution 48: 1155–1167. Zurich: University Zurich. 3–30, 55–119.

C 2009 Institute of Botany, Chinese Academy of Sciences