Thelypteridaceae (PDF)

This PDF version does not have an ISBN or ISSN and is not therefore effectively published (Melbourne Code, Art. 29.1). The printed version,however, was effectively published on 6 June 2013. Lin, Y. X., Z. Y. Li, K. Iwatsuki & A. R. Smith. 2013. Thelypteridaceae. Pp. 319–396 in Z. Y.Wu, P. H. Raven & D. Y. Hong, eds., Flora of China, Vol. 2–3 (Pteridophytes). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press.THELYPTERIDACEAE金星蕨科 jin xing jue keLin Youxing ( 林尤兴 ) 1 , Li Zhongyang ( 李中阳 ) 2 ; Kunio Iwatsuki 3 , Alan R. Smith 4Plants terrestrial or on rocks. Rhizomes stout, dictyostele radially symmetrical, branched or not, erect, ascending, or longcreeping, with scales at apices; scales basiflexed, lanceolate or nearly ovate, brown, thick, luminae elongate, usually with grayishwhite short setae on dorsal side or ciliate along margins. Fronds clustered, approximate, or remote; stipes slender, stramineous, notarticulate, with 2 crescent-shaped vascular bundles at base, usually scaly at bases, distally ± with grayish white unicellular acicularhairs, rarely with multicellular long hairs or stellate hairs. Fronds monomorphic, rarely subdimorphic, oblong-lanceolate or oblanceolate,sometimes ovate or ovate-triangular, usually pinnate-pinnatifid, sometimes 3- or 4-pinnate-pinnatifid, rarely 1-pinnate; pinnaesymmetrical at bases; costae grooved adaxially but grooves not confluent with rachial grooves, or raised and with dense grayishacicular hairs, with expanded tuberculate aerophores at bases of pinnae. Laminae herbaceous or papery, sometimes somewhatleathery, green or dark brown-green when dry, both sides (particularly rachises, costae, and main veins adaxially) with grayish whiteunicellular acicular hairs, rarely glabrous, usually with orange or reddish orange, stalked or sessile spherical or club-shaped glands,occasionally small scaly along rachises and costae abaxially. Sori orbicular, oblong, or shortly linear, dorsifixed on veins, indusiate orexindusiate; indusia orbicular-reniform, fixed by deep notch, most ± hairy, persistent or hidden in sori, caducous, or not concentratedinto sori but scattered along reticulate veins and exindusiate. Sporangia long stalked, usually with hairs or glandular hairs belowannuli and at distal end of sporangial stalks. Spores bilateral, rarely tetrahedral, tuberculate, echinate, granular, or usually with awinged perispore. Prothalli green, cordate or narrowly cordate, usually with broad wings, symmetrical, usually with hairs or glands. x= 27–36 (lacking 28).About 20 genera and ca. 1,000 species, more at lower elevations, very few tropical species above 4500 m: widespread in all tropical and subtropicalzones of the world, less common in temperate zone, particularly more common in Asia; 18 genera (one endemic) and 199 species (102endemic) in China.The family* is very natural and is distinguished from others by having grayish white unicellular acicular hairs and pubescence throughout theplant. However, there are many different viewpoints about generic circumscription in the family. Ching recognized 18 genera (including Hypodematium)in his 1963 treatment (Acta Phytotax. Sin. 8: 289–335); soon afterward, in 1978 (Ching, Acta Phytotax. Sin. 16(3): 12–13), the number ofrecognized genera in China grew to 20 (Hypodematium was removed and placed in its own family). In 1971, Holttum subdivided this family in thePaleotropics into 23 genera (Blumea 19(1): 17–52). In 1977, Pichi Sermolli, mainly following Holttum, circumscribed other genera for a total of 32genera (Webbia 3(2): 213–512). In 1990, A. R. Smith divided the family into five genera (in Kramer & Green, Fam. Gen. Vasc. Pl. 1: 263–272), i.e.,Thelypteris (including five subgenera), Phegopteris, Pseudophegopteris, Macrothelypteris, and Cyclosorus (including 20 subgenera). Of the manysystems, those of Holttum and Pichi Sermolli divide the family most finely, with the greatest number of genera. Holttum (loc. cit.) segregated thefollowing genera from Cyclosorus s.l.: Amphineuron, Christella, Pneumatopteris, and Sphaerostephanos. Recognition of these genera was based onseveral characters, including whether the proximal pinnae were shortened or not, and whether the sporangia and sporangial stalks bore hairs orglandular hairs. Holttum (loc. cit.) also segregated several genera (e.g., Parathelypteris and Coryphopteris) from the classical Thelypteris s.l., the freeveinedthelypteroids, by characters that included rhizome habit, laminar glands, and chromosome base number. Smith (loc. cit.: 265) noted: “Althoughmany of Holttum’s genera seem natural (i.e., monophyletic), a combination of characters must be used to circumscribe them. Some of the charactersconcern minute glands and hairs and require 30 × magnification or greater for observation. Others require that complete specimens be at hand(including lower part of blade and stem). Even then, identification to genus may be difficult, as generic lines are not always sharp.” The question ofgeneric delimitation within the family needs further study, but for now we adopt Ching’s system (1978), modified from Ching (1963).***In 1940, Ching established many new families (Sunyatsenia 5(4): 237), including Thelypteridaceae with 12 genera, but the names of these newfamilies are nomina nuda and were not therefore validly published (Melbourne Code, Art. 38.1(a)). Only in 1970 were these families formallypublished by Pichi Sermolli.**(1) Holttum set up the genus Amphineuron with mixed characters in 1971, which was also adopted by Ching in 1978. Among the 12recognized species in the genus, three occur in China: the type species A. opulentum, which is similar to Cyclosorus in venation and lemon-yellowglands; A. immersum, which is similar to Parathelypteris species in venation; and A. tonkinensis, which has already been removed to a new genus,Mesopteris. As Holttum himself considered Amphineuron to be a provisional treatment, this genus is not adopted here.(2) Trichoneuron Ching (Acta Phytotax. Sin. 10: 118, pl. 22. 1967) was based on a single gathering, and the collection locality was unknown.Later, this species was rediscovered by Prof. W. M. Chu in Pingbian, SE Yunnan. According to Prof. Chu’s study, it belongs to the genus Lastreopsis(Dryopteridaceae). It is, therefore, not included here.Using only venation, the family may be divided into 3 tribes: 1. Tribe Thelypterideae Ching: Veins free; with two subtypes: (1) all veinletsreaching margins or nearly reaching margins above sinuses between segments, with the bottom of sinuses not cartilaginous (e.g., Parathelypteris); (2)proximal pair of veinlets from adjacent segments reaching cartilaginous sinuses but not united; or acroscopic veinlet of basal pair of veinlets reachingcartilaginous sinus, basiscopic veinlet reaching margin above sinus (e.g., Pseudocyclosorus and Mesopteris); 2. Tribe Goniopterideae Ching: Veins1 Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.2 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’sRepublic of China.3 815-29 Kamoshida, Aoba-ku, Yokohama 227-0033, Japan.4 University Herbarium, University of California, 1001 Valley Life Sciences Building #2465, Berkeley, California 94720-2465, U.S.A.319

320THELYPTERIDACEAEpartly combined; at least proximal pair of veinlets from adjacent segments united to form triangular areoles and this vein union producing a long orshort excurrent veinlet; additional areoles may be produced in the same row, by subsequent vein unions of veinlets; excurrent veinlets may connectwith more distal vein unions, or with a translucent line leading to a sinus, thus forming oblique rhomboid areoles (goniopteroid venation) (e.g.,Cyclosorus, Ampelopteris), or squarish areoles (meniscioid venation) (e.g., Pronephrium, the areoles without included veinlets; 3. Tribe DictyoclineaeChing: Veinlets between lateral veins all combined into irregular squarish or pentagonal areoles and each areole sometimes with simple or forkedincluded veinlets.The following taxon is excluded from the present treatment, pending further research: Thelypteris calvata Ching (Bull. Fan Mem. Inst. Biol. Bot.ser. 2, 1: 313. 1949), described from Guangdong.Shing Kunghsia, Chiu Peishi, Yao Guanhu & Lin Youxin. 1999. Thelypteridaceae. In: Shing Kunghsia, ed., Fl. Reipubl. Popularis Sin. 4(1): 15–317, 319–353.Key 11a. Veins partly or entirely reticulate.2a. Veins entirely reticulate, with or without simple or forked included veinlets in areoles; sori scattered andattached along reticulate veins ............................................................................................................................... 18. Dictyocline2b. Veins partly reticulate (venation goniopteroid or meniscioid), areoles produced all without includedveinlets; sori orbicular or shortly linear.3a. Veins meniscioid, i.e., all veinlets joining into square or rectangular areoles; pinnae large, broadlylanceolate; sori orbicular when young and usually confluent when mature ............................................... 17. Pronephrium3b. Veins goniopteroid, i.e., proximal pair of veinlets from adjacent segments or one pair of veinletsbetween veinlets of pinna joined to form into triangular areoles and producing an excurrent veinletfrom these vein unions; excurrent veinlet or a line of translucent membrane at its end running toa sinus, several other pairs of veinlets connected to this excurrent vein or translucent membrane;pinnae small, narrowly lanceolate or triangular-lanceolate.4a. Pinnae pinnatifid; one long or short translucent membranous line at bottom of sinus on segmentsconnected with excurrent veinlet, veins except proximal pair from second to fifth pairs of veinletsconnected with excurrent veinlets or translucent membranous line and forming rhombic areoles,other veinlets reaching margins above sinus; laminae usually with orange or reddish orangespherical or club-shaped glands abaxially; sori indusiate ........................................................................ 14. Cyclosorus4b. Pinnae pinnatilobate or subentire, without translucent membranous line below sinus; exceptproximal pair of veins, other several pairs of veinlets connected with excurrent veinlet or reachingmargins above sinus; laminae without glands abaxially; sori exindusiate.5a. Plants with gemmae in pinna axils, these gemmae potentially developing into new plants;laminae covered with both simple and stellate hairs; sori orbicular or suborbicular;sporangia glabrous .......................................................................................................................... 16. Ampelopteris5b. Plants lacking gemmae; laminae with only simple hairs; sori thick and shortly linear;sporangia each with several setae at top ...................................................................................... 15. Stegnogramma1b. Veins free.6a. Sori indusiate.7a. Helophytes; lateral veins forked ....................................................................................................................... 1. Thelypteris7b. Terrestrial plants; veinlets usually simple (occasionally forked).8a. Costae glabrous adaxially (or occasionally with sparse caducous hairs); stipes with many scaleson proximal parts ......................................................................................................................................... 2. Oreopteris8b. Costae with dense persistent acicular hairs adaxially; stipes with sparse scales on proximal parts.9a. Costae rounded and raised adaxially; ends of veinlets not reaching margin; indusia small,or sometimes not developed.10a. Laminae oblong or broadly lanceolate, 2- or 3-pinnate, proximal pair of pinnae similarin shape and size or smaller than distal ones, throughout with unicellular hairs ............... 4. Metathelypteris10b. Laminae triangular-ovate, 3- or 4-pinnate, proximal pair largest, throughout withmulticellular hairs .............................................................................................................. 5. Macrothelypteris9b. Costae grooved adaxially; veinlets reaching margins; indusia large, brown.11a. Pinnae without tuberculate aerophores at pinna bases abaxially; proximal pairs ofveins reaching margins above sinus; laminae herbaceous, abaxially usually withreddish orange spherical glands .......................................................................................... 3. Parathelypteris11b. Pinnae each with a tuberculate aerophore at base abaxially; proximal pair of lateralveins from adjacent segments with acroscopic veinlet reaching cartilaginousbottom of sinus; laminae papery or leathery, abaxially without spherical glands ........ 12. Pseudocyclosorus6b. Sori exindusiate.12a. Sori orbicular.

THELYPTERIDACEAE 32113a. Laminae ovate-triangular, 3-pinnate; veinlets forked, not reaching margins; laminaethroughout with multicellular long hairs ..................................................................................... 5. Macrothelypteris13b. Laminae narrowly oblong or lanceolate, pinnate-pinnatifid; veinlets simple and reachingmargins; laminae throughout with unicellular short hairs.14a. Rachises at pinna bases abaxially with brown tuberculate aerophores; laminae ±brown when dry; segments without cartilaginous ridge at bottom of sinuses ....................... 9. Cyclogramma14b. Rachises at pinna bases abaxially without brown tuberculate aerophores; laminaegreen when dry; segments with a cartilaginous ridge at bottom of sinuses.15a. Plants ± with acicular hairs; pinnae pinnatifid nearly to costae; segmentsfalcate-lanceolate, proximal pair of veinlets nearly reaching translucentmembrane or margin above sinus; sori close to costules .................................. 11. Glaphyropteridopsis15b. Plants glabrous throughout; pinnae pinnatifid to 1/2 of distance to costae;segments triangular, proximal 3 pairs of veinlets reaching margin oftranslucent membrane below sinus but not joined; sori not close to costules ................. 13. Mesopteris12b. Sori thick and shortly linear, or oblong.16a. Sori thick and shortly linear; veins simple; segments entire.17a. Laminae brown or brownish green when dry, with full spreading acicular hairs on bothsurfaces; proximal 1 or 2 pairs of pinnae free, distal ones adnate to costae; veinletsexpanded at ends of veins and reaching margin; sori attached on middle of veinlets;sporangia each with 2–6 acicular hairs at tops ..................................................................... 10. Leptogramma17b. Laminae green when dry, subglabrous on both surfaces; pinnae completely free fromrachises; veinlets expanded at ends and not reaching margin; sori attached neartips of veinlets ........................................................................................................................ 7. Craspedosorus16b. Sori oblong or suborbicular; lateral veins ± forked; segments or ultimate pinnules pinnatifid.18a. Plants small; stipes stramineous, not polished; laminae ovate-triangular or narrowlylanceolate; lateral pinnae decurrent along both sides of rachises and connected to eachother; rachises and costae with more lanceolate scales, scales ciliate along margins;veinlets reaching margin ............................................................................................................. 6. Phegopteris18b. Plants usually taller and larger; stipes reddish brown or brownish stramineous, polished;laminae oblong, rarely broadly lanceolate; lateral pinnae not decurrent and free fromeach other; rachises and costae without scales; veinlets not reaching margin ............... 8. PseudophegopterisKey 2 (artificial)1a. Veins partly or almost fully connected.2a. Veins reticulate and forming regular areoles; sori scattered and attached along reticulate veins ........................ 18. Dictyocline2b. Veins partly joining; sori orbicular or shortly linear.3a. Veins meniscioid .......................................................................................................................................... 17. Pronephrium3b. Veins goniopteroid.4a. Plants of indeterminate growth, usually with gemmae in pinna axils, these gemmae potentiallydeveloping into new plants; laminae usually with mixed simple and stellate hairs ............................. 16. Ampelopteris4b. Plants of determinate growth; laminae with only simple hairs.5a. Segments with an elongate translucent membrane below sinus; laminae usually with orangeor reddish orange, spherical or clavate glands abaxially; sori orbicular, indusiate .......................... 14. Cyclosorus5b. Segments lacking translucent membranes below sinuses; laminae without glands abaxially;sori shortly linear, exindusiate ..................................................................................................... 15. Stegnogramma1b. Veins free.6a. Tuberculate aerophores present at pinna bases abaxially.7a. Proximal pair of veinlets on segments or acroscopic veinlet reaching sinus, others reachingmargin above sinus; laminae with only acicular hairs on both surfaces ............................................. 12. Pseudocyclosorus7b. Veinlets on segments all reaching margins above sinus; laminae ± with mixed hooked longhairs, except with acicular hairs ................................................................................................................... 9. Cyclogramma6b. Tuberculate aerophores lacking at pinna bases abaxially.8a. Segments each with a cartilaginous ridge at bottom of adjacent sinus and with a transparentmembranous line below that.9a. Plants ± with acicular hairs; pinnae pinnatifid nearly to costae; segments falcate-lanceolate,proximal pair of veinlets only reaching nearby line of transparent membrane below sinusor to margins above sinus; sori close to costules ...................................................................... 11. Glaphyropteridopsis9b. Plants glabrous throughout; pinnae pinnatifid to 1/2 of distance to costules; segmentstriangular; proximal 3 pairs of veinlets reaching transparent membranous margins belowsinus but not connected; sori not close to costules .................................................................................. 13. Mesopteris

322THELYPTERIDACEAE8b. Segments without cartilaginous ridge at bottom of sinus, lacking a translucent membranous linebelow that.10a. Laminae ovate-triangular or triangular, 3- or 4-pinnate; proximal pair of pinnae largest;veinlets not reaching margins; throughout with multicellular acicular hairs ................................. 5. Macrothelypteris10b. Laminae oblong or broadly lanceolate, most often pinnate-pinnatifid, to 3-pinnate-pinnatifid,rarely simple or bipinnatifid; proximal pair of pinnae similar in size or smaller than moredistal ones; throughout with unicellular acicular hairs.11a. Costae densely covered with persistent grayish white acicular hairs adaxially.12a. Costae rounded and raised adaxially; sporangia each sometimes with onemulticellular hair expanded at tip on distal part of sporangial stalks ............................ 4. Metathelypteris12b. Costae grooved adaxially; sporangial stalks sometimes each with a sessilespherical gland on distal part or sporangia each with one seta near annulus.13a. Veins not expanded at ends, reaching margins; sori orbicular, indusiate;sporangial stalks sometimes with 1–3 sessile spherical glands on distal parts .... 3. Parathelypteris13b. Veins with minute expanded hydathodes at ends and not reaching margins;sori oblong or shortly linear, exindusiate; sporangia each usually witha seta near annulus ................................................................................................. 7. Craspedosorus11b. Costae without unicellular acicular hairs adaxially (occasionally with sparse caducoushairs).14a. Rachises and costae with few small scales.15a. Scales on rachises and costae with glands at apices, without cilia; pinnae sessileat bases, not adnate to rachises ..................................................................................... 2. Oreopteris15b. Scales on rachises and costae without glands at tips but ciliate below tips; pinnabases adnate to rachises and decurrent ....................................................................... 6. Phegopteris14b. Rachises and costae without scales.16a. Plants of marshes; veinlets forked; sori indusiate; sporangia each with severalshort setae near top of annulus .................................................................................... 1. Thelypteris16b. Plants terrestrial; veinlets simple; sori exindusiate; sporangia each withseveral acicular hairs.17a. Sori orbicular; veinlets not reaching margins ....................................... 8. Pseudophegopteris17b. Sori oblong or shortly linear; veinlets reaching margins ............................ 10. Leptogramma1. THELYPTERIS Schmidel, Icon. Pl., ed. Keller, 3, 45. 18 Oct 1763, nom. cons.,not Adanson (Jul–Aug 1763).沼泽蕨属 zhao ze jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiLastrea Bory.Plants small to medium-sized, of marshes and meadows. Rhizomes long creeping, black, glabrous, sparsely covered with scalesat apices; scales ovate-lanceolate, with acicular hairs and unicellular glandular hairs on surfaces and along margins. Fronds remote orapproximate, stipitate; stipes nearly black at bases and slightly acicular hairy, distally stramineous, glabrous; laminae oblonglanceolate,not tapering or slightly tapering to bases, pinnate-pinnatifid, shortly acuminate at apices; pinnae mostly nearly flatspreading,lanceolate, bases truncate, symmetrical, pinnatifid, apices acute or shortly acuminate; segments ovate-triangular or oblong,shortly pointed at apices; veins free, pinnate on segments, veinlets forked or simple, reaching margins. Laminae thickly herbaceousor somewhat leathery, both surfaces with few acicular hairs when young, glabrescent when old, costae each with a groove adaxially,abaxially raised, sometimes with few membranous small scales. Sori orbicular, dorsifixed on veinlets, located between costules andmargins, in one line on each side of costules, usually ± covered by reflexed margins; indusia membranous, orbicular-reniform,greenish, deciduous or hidden in mature sori; sporangia each with 1 or 2 short capitate glandular hairs close to tops of annuli; sporesbilateral, reniform, perispores transparent, echinate, exospore smooth. x = 35.Four species: temperate regions of N Hemisphere, S tropical and S Africa, India, Madagascar, New Guinea, New Zealand; two species in China.In the past, Thelypteris has been defined to contain most or all species in the family. More recent studies show that, using multiple additionalcharacters, like types of hairs, glands, venation, lamina shape and dissection, indusial characters, aerophores, adaxial grooves, spores, and chromosomebase numbers, natural groups can be utilized to define smaller subsets of Thelypteris s.l. Here, we adopt a narrow concept for classification.1a. Plants 35–65 cm tall; fronds approximate, stipes 2–2.5 mm in diam. at bases; costae without scales abaxially ......... 1. T. palustris1b. Plants 14–26 cm tall; fronds remote, stipes slender, ca. 0.5 mm in diam. at bases; costae with sparse easilydeciduous, broadly ovate, brownish, membranous scales ........................................................................................... 2. T. fairbankii

THELYPTERIDACEAE 3231. Thelypteris palustris Schott, Gen. Fil. t. 10. 1834.沼泽蕨 zhao ze juePlants 35–65 cm tall. Rhizomes long creeping, black, glabrousor with sparse reddish brown, ovate-lanceolate scales.Fronds approximate; stipes 20–40 cm, bases black, distally darkstramineous, polished, usually glabrous, or white pubescentwhen young; laminae lanceolate, 22–28 × 6–9 cm or sometimesslightly wider, bases almost tapering, pinnate-pinnatifid, apicesshortly acuminate and pinnatifid; pinnae ca. 20 pairs, subopposite,flat- or obliquely spreading, usually slightly reflexed;proximal pair slightly shortened, middle pinnae lanceolate, 4–5× 1–1.2 cm, bases truncate, pinnatifid nearly to costae, apicesshortly acuminate; segments 5–7 × 3–5 mm, rounded-obtuse orobtuse-pointed at apices, fertile segments usually recurved toforming points along margin. Veins pinnate in segments, lateralveins 4–6 pairs, simple or forked and reaching margins, proximalpair arising from base of costa. Laminae papery, grassgreenor yellowish green when dry, glabrous on both surfaces,rachises and costae grooved adaxially, raised abaxially, glabrouson both sides or with acicular long hairs abaxially. Soriorbicular, dorsifixed at middle of veinlets, located betweencosta and margins; indusia small, orbicular-reniform, membranous,deciduous when mature. Spores smooth on surfaces ofexospore, perispores translucent, echinate. 2n = 70.Meadows, reed marshes, wet shaded places in forests; 200–800 m.Hebei, Heilongjiang, Henan, N Jiangsu, Jilin, Nei Mongol, Shandong,Sichuan, Xinjiang [widely distributed in temperate regions of the NHemisphere].1a. Rachises, costae, and veinlets glabrous ... 1a. var. palustris1b. Rachises, costae, and veinlets coveredwith multicellular acicular long hairsabaxially ................................................ 1b. var. pubescens1a. Thelypteris palustris var. palustris沼泽蕨 ( 原变种 ) zhao ze jue (yuan bian zhong)Acrostichum thelypteris Linnaeus, Sp. Pl. 2: 1071. 1753;Aspidium palustre A. Gray, nom. illeg. superfl.; A. thelypteris(Linnaeus) Swartz; Athyrium thelypteris (Linnaeus) Sprengel;Dryopteris thelypteris (Linnaeus) A. Gray; Lastrea thelypteris(Linnaeus) Bory; Nephrodium thelypteris (Linnaeus) Strempel;Polypodium palustre Salisbury (1796), not N. L. Burman(1768); P. pterioides Lamarck; P. thelypteris (Linnaeus) F. G.Weiss.Rachises, costae, and veins all glabrous.Meadows, reed marshes, wet shaded places in forests; 200–800m. Hebei, Heilongjiang, Henan, Jilin, Nei Mongol, Shandong, Sichuan,Xinjiang [widely distributed in temperate regions of N Hemisphere].1b. Thelypteris palustris var. pubescens (G. Lawson) Fernald,Rhodora 31: 34. 1929.毛叶沼泽蕨 mao ye zhao ze jueLastrea thelypteris var. pubescens G. Lawson, EdinburghNew Philos. J. 19: 277. 1864.Rachises, costae, and veinlets covered with multicellularacicular long hairs abaxially. 2n = 70.Wet meadows and marshes; below 800 m. Heilongjiang, N Jiangsu,Jilin, Shandong [temperate regions of E Asia and North America].2. Thelypteris fairbankii (Beddome) Y. X. Lin, K. Iwatsuki &M. G. Gilbert, comb. nov.鳞片沼泽蕨 lin pian zhao ze jueBasionym: Lastrea fairbankii Beddome, Ferns Brit. India,t. 254. 1866; Aspidium thelypteris (Linnaeus) Swartz var.squamigerum Schlechtendahl; L. thelypteris (Linnaeus) Boryvar. squamigera (Schlechtendahl) Beddome; Nephrodium thelypteris(Linnaeus) Strempel var. squamigerum (Schlechtendahl)Hooker [“squamulosum”]; Thelypteris squamigera (Schlechtendahl)Ching [“squamulosa”].Plants small, 14–26 cm tall. Rhizomes long creeping andbranched, black, apices including stipe bases with sparsemembranous, brownish ovate-lanceolate scales. Fronds remote;stipes slender, 6–18 cm, black at bases, distally stramineous,glabrous; laminae oblong-lanceolate or narrowly triangular-lanceolate,8–10 × 3–5 cm, sometimes slightly tapering at bases,pinnate-pinnatifid, shortly acuminate and pinnatifid at apices;pinnae 10–12 pairs, alternate or subopposite, middle ones linear-lanceolate,2–3 × 0.5–0.8 cm, truncate at bases, pinnatifid,shortly pointed at apices; segments oblong or triangular-lanceolate,usually reflexed at margins, entire, obtuse at apices.Veins pinnate in segments, lateral veins forked, veinletsreaching margins. Laminae somewhat leathery, glabrous, alongrachises and costae sparsely pubescent adaxially, costae withsparse brownish, membranous, easily deciduous, ovate-lanceolatescales. Sori not seen.Marshes. S Yunnan [S India; S Africa, Pacific islands (New Zealand)].2. OREOPTERIS Holub, Folia Geobot. Phytotax. (Praha) 4: 46. 1969.假鳞毛蕨属 jia lin mao jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants medium-sized, terrestrial. Rhizomes short, erect or ascending. Fronds clustered; stipes dark stramineous, densely coveredwith large, thin, brown, lanceolate scales, distally gradually sparsely so; laminae oblong-lanceolate, gradually tapering to bases, pinnate-pinnatifid;proximal pinnae gradually shortened, proximal ones triangular auriculate, middle ones lanceolate, pinnatifid to narrowwings on both sides of costae. Veins pinnate, free, reaching margins. Sori orbicular, attached above middle of veinlets, far fromcostules; indusia orbicular-reniform, usually with glands along margins. Sporangia usually with stalked glands near annuli. Spores bilateral,reniform, perispores not evident and easily deciduous, granular. x = 34.Three species: Europe, N India, E Asia to N America; two species in China.

THELYPTERIDACEAE 325shortened or 1 to several pairs of pinnae obviously shortened, even reduced to small auricles, pinnatifid; segments mostly oblong,rectangular, or subsquare, margins entire or ± serrate, apices rounded-obtuse, sometimes pointed or with sinuslike angles. Veinletspinnate, free, simple and all reaching margins. Laminae herbaceous or papery, when dry yellowish green, sometimes brownish greenor nearly black, both surfaces ± with pubescent or acicular hairs, rarely glabrous when dry, sometimes with orange-yellow or reddishpurple glands abaxially; costae grooved adaxially, densely covered with short setae, rounded and raised abaxially and usually ± withacicular hairs, rarely glabrous. Sori orbicular, medium-sized, dorsifixed at middle or near ends of veinlets, located between mainveins and margins or slightly closer to margins; indusia larger, orbicular-reniform, sometimes horseshoe-shaped, when dry brown,membranous, glabrous or hairy, usually persistent. Spores bilateral, orbicular-reniform, perispores thin and transparent, corrugate, ±finely reticulate on surfaces. x = 8, 9, 31.About 60 species: tropical and subtropical regions of E Asia, SE Asia to Pacific islands; 24 species (11 endemic) in China.Parathelypteris is very similar to Metathelypteris in shape, but it differs by the costae grooved adaxially; the veinlets usually simple, not forked,and reaching margins; and the indusia larger and readily discerned.The following taxa are excluded from the present treatment, pending further research: Parathelypteris auriculata H. G. Zhou & H. Li (Acta Bot.Yunnan. 14: 33. 1992), described from Guangxi (Wuming), and P. jinfoshanensis Ching & Z. Y Liu (Bull. Bot. Res., Harbin 4(3): 18. 1984), describedfrom Sichuan (Nanchuan).1a. Stipes stramineous, glabrous or sparsely shortly hairy near bases; sori usually attached near ends of veinlets and closeto margins.2a. Several pairs of pinnae on proximal part of lamina clearly shortened.3a. Plants short and small, usually no more than 40 cm; rhizomes long creeping; middle pinnae nomore than 3.5 cm; pinnae with more multicellular (usually 3–7 cells) acicular hairs abaxiallyalong costae ...................................................................................................................................................... 1. P. beddomei3b. Plants more than 40 cm tall; rhizomes stronger, creeping or ascending; middle pinnae usually over4 cm; pinnae usually with unicellular acicular hairs abaxially along costae, sometimes with 2- or3-celled hairs.4a. Pinnae 3 or 4 pairs on proximal part of lamina gradually shortened, proximal pair of pinnae1–2 cm, not auriculate ................................................................................................................ 2. P. changbaishanensis4b. Pinnae 5–8 pairs on proximal part of lamina gradually shortened, proximal pair of pinnaereduced to small auricles or lacking.5a. Laminae without glands or occasionally with few orange-yellow spherical glands abaxially;rhizomes long creeping, subglabrous ................................................................................................ 3. P. nipponica5b. Laminae with more orange-yellow spherical glands abaxially; rhizomes ascending orcreeping, densely rusty yellow pubescent.6a. Rhizomes ascending; rachises with glaucous fine acicular hairs abaxially; indusia withdense setae ................................................................................................................................ 4. P. qinlingensis6b. Rhizomes long creeping, densely rusty yellow hairy; rachises subentire abaxially;indusia glabrous or occasionally with 1 or 2 setae ........................................................................ 5. P. borealis2b. Proximal pinnae not shortened or slightly shortened.7a. Plants usually not more than 20 cm tall; middle pinnae 1.5–2 × 4–8 cm; segments 2–5 pairs;indusia horseshoe-shaped.8a. Pinnae subglabrous abaxially, nearly without glands; indusia larger and subglabrous,occasionally with few grayish white setae ........................................................................................ 6. P. cystopteroides8b. Costae and costules abaxially with sparse grayish white fine acicular hairs and orangeyelloworbicular glands; indusia small and with more setae ............................................................. 7. P. grammitoides7b. Plants 25–60 cm tall, or 2–3 m tall.9a. Plants 2–3 m tall; lateral pinnae 25–30 × 2–2.5 cm; segments 60–70 pairs per pinna; pinnaglands confined to veinlets .................................................................................................................. 11. P. subimmersa9b. Plants 25–60 cm tall; middle pinnae 2–6 × ca. 1 cm; segments 6–20 pairs; pinnae glandsscattered abaxially.10a. Costae glabrous abaxially; segments of fertile fronds crenate along margins ................................. 8. P. serrutula10b. Costae ± with acicular hairs; segments of fertile fronds entire along margins.11a. Middle pinnae 2–3 cm, acute at apices; segments 6–10 pairs; proximal several pairsof pinnae minutely shortened; costae with only few acicular hairs abaxially .................. 9. P. angustifrons11b. Middle pinnae ca. 4.5 cm, acuminate at apices; segments more than 15 pairs;proximal pinnae not shortened; costae with more acicular hairs abaxially ................... 10. P. glanduligera1b. Stipes in part or whole (usually up to rachis) castaneous or castaneous-brown, rarely stramineous, basesglabrous or with spreading grayish white acicular hairs; sori usually dorsifixed on middle of veinlets,located between costa and margins.

326THELYPTERIDACEAE12a. Laminae without reddish purple spherical glands abaxially.13a. Stipes with dense unicellular grayish white short acicular hairs; lamina with similar densehairs on both surfaces ................................................................................................................................ 20. P. castanea13b. Stipes with spreading multicellular acicular long hairs at bases.14a. Segments 2–4 sinuslike angular at apices.15a. Rhizomes short and erect; bases of stipes mixed with few spreading multicellularlong acicular hairs and, including above bases, with grayish white unicellular shortsetae; indusia sparsely pubescent .......................................................................................... 21. P. pauciloba15b. Rhizomes short, decumbent or ascending; stipes with dense spreading multicellularacicular long hairs, distally subglabrous; indusia densely shortly setaceous .................. 22. P. angulariloba14b. Segments not sinuslike angular at apices.16a. Laminae dark green or nearly black when dry; pinnae abaxially densely pubescentalong costae, veins and interval sparsely shortly hairy; indusia glabrous oroccasionally very sparsely pubescent when dry .................................................................. 23. P. nigrescens16b. Laminae grayish green when dry; pinnae with dense multicellular beardlike longacicular hairs; indusia densely pubescent ....................................................................... 24. P. indochinensis12b. Pinnae with reddish purple spherical large glands abaxially.17a. Stipes glabrous at bases.18a. Laminae lanceolate; middle pinnae narrower, 0.8–1.2 cm wide, glabrous or occasionallyvery sparsely grayish white pubescent abaxially; indusia glabrous or with sparse shorthairs; stipes castaneous-brown, never stramineous ........................................................................ 18. P. chinensis18b. Laminae ovate-oblong; middle pinnae 1.3–1.6 cm wide, usually grayish white pubescentabaxially, rarely glabrous; stipes usually coetaneous, occasionally stramineous ............................ 19. P. japonica17b. Stipes with spreading 2- or 3-celled grayish white acicular hairs at bases.19a. Laminae abaxially with long acicular hairs along costae.20a. Rachises usually hairy abaxially; lateral pinnae 12–15 pairs; segments ± angularat apices, 3–6 veinlets per segment; laminae with reddish purple glands abaxially ................. 12. P. chingii20b. Rachises usually glabrous abaxially, lateral pinnae 18–30 pairs; segmentsrounded-obtuse or rounded-truncate at apices and not angular, 7 or 8 pairsof veinlets per segment; laminae with denser reddish purple glands abaxially.21a. Indusia with dense grayish white acicular long hairs ..................................................... 13. P. petelotii21b. Indusia glabrous or occasionally with few short setae ................................................ 14. P. hirsutipes19b. Laminae with short hairs or subglabrous abaxially along rachises.22a. Laminae glabrous or with few short acicular hairs abaxially; indusia glabrous oroccasionally with short setae ................................................................................................... 15. P. caudata22b. Laminae with dense short acicular hairs abaxially; indusia hairy also.23a. Indusia large, close to each other, densely shortly setaceous ............................... 16. P. trichochlamys23b. Indusia medium-sized, separate from each other, sparsely pubescent ................... 17. P. caoshanensis1. Parathelypteris beddomei (Baker) Ching, Acta Phytotax.Sin. 8: 302. 1963.长根金星蕨 chang gen jin xing jueNephrodium beddomei Baker in Hooker & Baker, Syn. Fil.267. 1867; Aspidium beddomei (Baker) Prantl; Dryopteris beddomei(Baker) Kuntze; Lastrea beddomei (Baker) Beddome;Thelypteris beddomei (Baker) Ching; Wagneriopteris beddomei(Baker) A. Löve & D. Löve.Plants 20–30(–40) cm tall. Rhizomes extremely longcreeping, with sparse brown ovate small scales and densebrownish long hairs when young. Fronds remote or approximate;stipes slender, 4–10 cm, stramineous and glabrous; laminaeoblanceolate, 15–25(–30) × 3–4(–6) cm, gradually taperingto bases, pinnate-pinnatifid, acuminate and pinnatifid atapices; pinnae 20–24(–30) pairs, alternate, sessile, obliquelyspreading, proximal 7–9 pairs gradually shortened into smallauricles, proximal pair of pinnae only 1–2 mm, middle pinnaelanceolate, 1.5–3.5 × 0.4–0.5 cm, bases minutely broadened,symmetrical, rounded-truncate, pinnatifid to narrow wing onboth sides of costae, apices shortly acuminate; segments 10–14pairs, oblong, entire, rounded at apices. Veins visible on bothsides, veinlets pinnate, free, simple, reaching margins, 3 or 4pairs per segment, proximal pair arising from bases of costae.Laminae herbaceous, yellowish brown when dry, abaxially withgrayish white 3–7-celled fine long hairs along costae and veins,also with few orange-yellow spherical glands, adaxially withunicellular short acicular hairs along costae and veins. Sorismall, 2 or 3 pairs per segment, attached near ends of veinlets,close to margins; indusia orbicular-reniform, small, brown,thickly membranous, glabrous, persistent. 2n = 62.Mountain meadows, streamsides, wetlands; 600–2500 m. Taiwan,Zhejiang [S India, Indonesia, Japan, Malaysia, Philippines].Parathelypteris beddomei is similar to P. nipponica in outline butdiffers by plants thinner, rhizomes extremely slender, pinnae coveredwith more multicellular long acicular hairs, and growth in wet habitats.

THELYPTERIDACEAE 3272. Parathelypteris changbaishanensis Ching ex K. H. Shing,Fl. Reipubl. Popularis Sin. 4(1): 319. 1999.长白山金星蕨 chang bai shan jin xing juePlants 45–60 cm tall. Rhizomes long creeping, nearlyblack and glabrous. Fronds approximate; stipes 15–25 cm,bases nearly black, occasionally covered with few dark brownovate scales, distally stramineous, glabrous or sometimes withsparse grayish white fine and long hairs; laminae oblonglanceolate,25–35 × 8–12 cm, clearly tapering to bases, pinnatepinnatifid,acuminate and pinnatifid at apices; pinnae 20–25pairs, alternate, obliquely spreading, sessile, proximal 3 or 4pairs gradually shortened, proximal pair of pinnae 1–2 cm, lanceolateand reflexed proximally; middle pinnae linear-lanceolate,4–6 × 0.9–1.3 cm, bases minutely broadened, symmetrical,subtruncate, pinnatifid nearly to costae, apices acuminate; segmentsca. 15 pairs, proximal pair slightly longer, middle onesoblong, 4–6 × 2–3 mm, sparsely shallowly and thickly serratealong margins, rounded-obtuse at apices. Veinlets visible,oblique distally, 5(or 6) pairs per segment, proximal pair arisingfrom bases of costae. Laminae thinly herbaceous, grass-greenwhen dry, abaxially with orange-yellow spherical glands, costaevillous, sparsely so along costules; adaxially with denser appressedshort acicular hairs along grooves and veins. Sori small,orbicular, 3 or 4 pairs per segment, dorsifixed above middle ofveinlets, slightly closer to margins; indusia small, orbicularreniform,grayish yellow, membranous, hairy, persistent.● Meadows; 600–1400 m. Jilin (Changbai Shan).Parathelypteris changbaishanensis is similar to P. nipponica inoutline but differs by proximal 2–4 pairs of pinnae shortened, pinnaewith dense orange-yellow spherical glands and villous abaxially, andgrowth in meadows.3. Parathelypteris nipponica (Franchet & Savatier) Ching,Acta Phytotax. Sin. 8: 301. 1963.中日金星蕨 zhong ri jin xing jueAspidium nipponicum Franchet & Savatier, Enum. Pl. Jap.2: 242, 636. 1879; Dryopteris nipponica (Franchet & Savatier)C. Christensen; Lastrea nipponica (Franchet & Savatier) Copeland;Thelypteris nipponica (Franchet & Savatier) Ching; Wagneriopterisnipponica (Franchet & Savatier) A. Löve & D. Löve.Plants 40–60 cm tall. Rhizomes long creeping, subglabrous.Fronds approximate; stipes 10–20 cm, bases dark brown,± covered with reddish brown broadly ovate scales, distallybright stramineous, glabrous; laminae oblanceolate, 30–40 × 7–10 cm, gradually tapering to bases, pinnate-pinnatifid, acuminateand pinnatifid at apices; pinnae 25–33 pairs, proximal 5–7pairs subopposite, proximally gradually reduced into smallauricles, lowest ones tuberculate; middle pinnae alternate, sessile,spreading, lanceolate, 4–5 × 0.7–1.2 cm, bases slightlybroadened, symmetrical, truncate, pinnatifid nearly to costae,apices acuminate; segments ca. 18 pairs, slightly obliquelyspreading, oblong, 3–5 × ca. 2 mm, entire or shallowly thicklyserrate, rounded-obtuse at apices. Veinlets evident, simple, 4 or5 pairs per segment. Laminae herbaceous, grass-green whendry, abaxially with grayish white, spreading, unicellular andmixed with few multicellular acicular hairs along costae andmargins, interstitial region with dense micro-glandular hairs andfew orange-yellow spherical glands; adaxially subglabrousexcept costae and veins with short acicular hairs. Sori orbicular,medium-sized, 3 or 4 pairs per segment, dorsifixed abovemiddle of veinlets, far from costa; indusia medium-sized, orbicular-reniform,brown, membranous, with few grayish whitelong acicular hairs. Spores bilateral, orbicular-reniform, perisporescorrugate, exospore regularly finely reticulate. 2n = 124.On soil in open forests on hills, common; 400–2500 m. Fujian,Gansu, Guangxi, Guizhou, Henan, Hubei, Hunan, N Jiangsu, Jiangxi,Shaanxi, Shandong, Sichuan, Yunnan, Zhejiang [Japan, S Korea, Nepal].Parathelypteris nipponica is similar to P. novoboracensis (Linnaeus)Ching in outline, but the latter plant, which grows along theAtlantic coast, differs in segments acute at apices and indusia glabrousat the back.4. Parathelypteris qinlingensis Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 320. 1999.秦岭金星蕨 qin ling jin xing juePlants 45–60 cm tall. Rhizomes ascending. Fronds subclustered;stipes 10–20 cm, bases dark brown, with sparsebrown broadly ovate scales, distally stramineous, glabrous;laminae oblanceolate, 35–45 × 8–11 cm, gradually tapering tobases, pinnate-pinnatifid, acuminate and pinnatifid at apices;pinnae ca. 25 pairs, proximally gradually shortened intosmall auricles, lowest ones tuberculate; middle pinnae alternate,spreading, sessile, lanceolate, 4–6 × ca. 1 cm, bases symmetricaland slightly broader, subtruncate, pinnatifid nearly to costae,apices long acuminate; segments ca. 18 pairs, spreading, oblong-lanceolate,4–6 × ca. 2 mm, entire, obtuse at apices. Veinletsevident, simple, 5 or 6 pairs per segment, proximal pairarising from bases of costule. Laminae thickly herbaceous,grass-green when dry, abaxially with orange-yellow sphericalglands, costae, costules, and margins with dense grayish whitespreading fine acicular hairs, adaxially along grooves and veinswith appressed short setae, along rachises with sparse longhairs. Sori orbicular, 3–5 pairs per segment, dorsifixed nearends of veinlets and close to margins; indusia medium-sized,orbicular-reniform, brown, membranous, with more setae, persistent.Spores bilateral, orbicular-reniform, perispores corrugateand obviously finely reticulate on surfaces, exospore finelyreticulate.● Pinus armandii forests, common; ca. 1800 m. Gansu, Shaanxi.Parathelypteris qinlingensis differs from P. nipponica in havingrhizomes ascending, fronds subclustered, pinnae with more orange-yellowspherical glands, fine acicular long hairs abaxially, and indusiadensely covered with setae.5. Parathelypteris borealis (H. Hara) K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 37. 1999.狭脚金星蕨 xia jiao jin xing jueDryopteris nipponica (Franchet & Savatier) C. Christensenvar. borealis H. Hara, Bot. Mag. (Tokyo) 48: 695. 1934;Parathelypteris nipponica (Franchet & Savatier) Ching var.borealis (H. Hara) Nakaike; Thelypteris nipponica (Franchet &Savatier) Ching var. borealis (H. Hara) H. Hara.

328THELYPTERIDACEAEPlants 45–65 cm tall. Rhizomes long creeping, with denserusty yellow hairs. Fronds approximate; stipes ca. 13 cm, stramineous,glabrous, bases with sparse brown ovate scales; laminae25–50 × 5–11 cm; pinnae alternate, flatly spreading, sessile,proximal 7 or 8 pairs abruptly tapering into auricles, lowestones tuberculate; middle pinnae lanceolate, 3–6 × 0.6–0.9 cm,bases slightly broadened, symmetrical, subtruncate, pinnatifidnearly to costae, apices acuminate; segments 12–18 pairs, oblong,3–4.5 × 1.5–2 mm, entire, obtuse at apices. Veinlets evident,simple, 4 or 5 pairs per segment, proximal pairs arisingfrom above bases of costules. Laminae herbaceous, yellowishdark green when dry, with dense orange-yellow spherical glandsabaxially, with sparse grayish white fine acicular hairs alongcostae, costules, and margins, adaxially along grooves of costaewith short hairs, elsewhere glabrous. Sori orbicular, dorsifixednear ends of veinlets and close to margins, 1–4 pairs per segment;indusia small, orbicular-reniform, brown, membranous,subglabrous. Spores bilateral, orbicular-reniform, perisporesfew corrugate and finely reticulate on corrugations.Valley thickets, wet shaded places in forests, common; 400–1900m. N Anhui, N Fujian, Guangxi, Guizhou, Hunan, Jiangxi, Shaanxi, Sichuan[Japan].Parathelypteris borealis is very similar to P. nipponica in laminaroutline but differs by laminae with dense orange yellow sphericalglands abaxially and indusia subglabrous. It is also similar to P. qinlingensis,but the latter has rhizomes ascending, costae with denser longacicular hairs abaxially, and indusia with more setae.6. Parathelypteris cystopteroides (D. C. Eaton) Ching, ActaPhytotax. Sin. 8: 302. 1963.马蹄金星蕨 ma ti jin xing jueAthyrium cystopteroides D. C. Eaton, Proc. Amer. Acad.Arts 4: 110. 1858; Asplenium cystopteroides (D. C. Eaton)Hooker; Dryopteris abbreviatipinna Makino & Ogata; D. cystopteroides(D. C. Eaton) Kodama ex Tagawa; D. gracilescens(Blume) Kuntze var. abbreviata Kodama; Lastrea cystopteroides(D. C. Eaton) Copeland; Thelypteris cystopteroides (D. C.Eaton) Ching.Plants 7–20 cm tall. Rhizomes long creeping, branchingmixed and becoming feltlike, with sparse dark brown lanceolatesmall scales. Fronds approximate; stipes 3–5(–13) cm, slender,dark stramineous, subglabrous; laminae lanceolate, 4–7 × ca.1.5 cm, not tapering to bases, bipinnatipartite, acuminate andpinnatifid at apices; pinnae 7–10 pairs, alternate, spreading,stalks short, proximal pair of similar shape as distal ones, 7–10× 6–7 mm, subtruncate at bases, pinnatifid nearly to costae,obtuse at apices; segments 2 or 3 pairs, oblong, ca. 3 × ca. 1.5mm, proximal pair usually irregularly bilobate. Veinlets evident,simple, ca. 3 pairs per segment. Laminae herbaceous,dark green when dry, subglabrous abaxially, adaxially withsparse appressed short hairs along grooves on costae. Soriorbicular or oblong, medium-sized, 2–4 pairs per segment, dorsifixedslightly above middle of veinlets; indusia large, orbicular-reniform,brown, thickly membranous, occasionallywith few grayish white setae, persistent.On rocks in forests. Along the coast, offshore islands of Fujian,Taiwan [Japan, Korea].7. Parathelypteris grammitoides (Christ) Ching, Acta Phytotax.Sin. 8: 302. 1963.矮小金星蕨 ai xiao jin xing jueAspidium grammitoides Christ, Bull. Herb. Boissier 6:193. 1898; Athyrium hyalostegium Copeland; Dryopteris glanduligera(Kunze) Christ var. hyalostegia (Copeland) H. Itô; D.grammitoides (Christ) C. Christensen; Lastrea grammitoides(Christ) Copeland; Thelypteris glanduligera (Kunze) Ching var.hyalostegia (Copeland) H. Itô; T. grammitoides (Christ) Ching.Plants 10–18 cm tall. Rhizomes long creeping, apices includingbases of stipes with reddish brown narrowly lanceolatesmall scales. Fronds approximate; stipes 3–10 cm, dark stramineous,with grayish white fine acicular hairs; laminae lanceolate,5–8 × 2–3.5 cm, tapering to bases, pinnate-pinnatifid,acuminate and pinnatifid at apices; pinnae 6–8 pairs, alternate,subspreading, lanceolate, bases symmetrical and tapering,rounded-truncate, pinnatifid nearly to costae, apices obtuse oracute; segments 3–5 pairs, obliquely spreading, proximal pairoblong, 2–3 × 1.5–2 mm, entire, rounded at apices; distal segmentsgradually shortened. Veinlets visible abaxially, simple, 3or 4 pairs per segment, proximal pair arising above bases ofcostules. Laminae thickly leathery, dark green when dry, withorange-yellow spherical glands abaxially, costae and costuleswith sparse glaucous acicular long hairs, adaxially along costalgrooves with acicular hairs. Sori small, orbicular, 1–6 pairs persegment, dorsifixed near ends of veinlets, close to margins;indusia horseshoe-shaped, medium-sized, brown, thickly membranous,with more fine and long setae, persistent. Spores bilateral,orbicular-reniform, perispores corrugate, finely reticulateon corrugations.On rocks in forests; 1000–1400 m. Taiwan [Japan, Korea, Philippines].8. Parathelypteris serrutula (Ching) Ching, Acta Phytotax.Sin. 8: 303. 1963 [“serrulata”].有齿金星蕨 you chi jin xing jueThelypteris serrutula Ching, Bull. Fan Mem. Inst. Biol.,Bot. 6: 319. 1936.Plants ca. 50 cm tall. Rhizomes long creeping, glabrous,apices with sparse dark brown lanceolate scales. Fronds approximateor remote; stipes ca. 25 cm, dark stramineous, subglabrous;laminae broadly lanceolate, ca. 25 × 13 cm, not taperingto bases, pinnate-pinnatifid, acuminate and pinnatifid at apices;pinnae ca. 20 pairs, spreading, alternate or subopposite onproximal ones, subsessile, lanceolate, 5–7 × ca. 1.5 cm, basessymmetrical, subtruncate, apices acuminate; proximal pinnaeslightly tapering to bases, pinnatifid nearly to costae; segmentsca. 18 pairs, spreading, lanceolate, ca. 7 × 2 mm, acute at apices;segments on fertile laminae crenate along margins; sterile onessharply lobate along margins. Veinlets evident, simple, 6 or 7pairs per segment, proximal pair arising from bases of costules,with sparse spherical glands abaxially, glabrous, adaxial sideswith dense short hairs along costal grooves and on costae. Sorimedium-sized, orbicular, 5 or 6 pairs per segment, dorsifixednear ends of veinlets, close to margins; indusia small, orbicular-reniform,brown, membranous, glabrous or occasionally

THELYPTERIDACEAE 329sparsely hairy and deciduous when old. Spores bilateral, orbicular-reniform,perispores few, corrugate.● Streamsides in forests; ca. 100 m. Guizhou, SW Sichuan, Zhejiang.9. Parathelypteris angustifrons (Miquel) Ching, Acta Phytotax.Sin. 8: 302. 1963.狭叶金星蕨 xia ye jin xing jueAspidium angustifrons Miquel, Ann. Mus. Bot. Lugduno-Batavi 3: 178. 1867; Athyrium cystopteroides D. C. Eaton var.elatus D. C. Eaton; Lastrea miqueliana Tagawa; Thelypterisangustifrons (Miquel) Ching.Plants 25–35 cm tall. Rhizomes long creeping, apices includingstipe bases with sparse dark brown lanceolate thickscales. Fronds approximate; stipes 10–15 cm, stramineous, subglabrous,distal parts and rachises more grayish white pubescent;laminae lanceolate, 15–20 × 3–4 cm, slightly taperingto bases, pinnate-pinnatifid or subbipinnate, acuminate and pinnatifidat apices; pinnae 10–15 pairs, alternate, spreading, sessileor very shortly stalked, proximal pair of similar shape butslightly shorter than distal ones; middle pinnae lanceolate, 2–3× ca. 1 cm, bases ± symmetrical, truncate, pinnatifid nearly tocostae or subpinnate, apices obtuse or acute; segments or pinnules6–20 pairs, proximal 2 or 3 pairs usually free, proximalpair of pinnae larger, oblong, 4–5 × ca. 2 mm, bases cuneate,slightly adnate to costae, margins entire or thickly serrate,apices obtuse; distal pairs gradually reduced. Veinlets evidentabaxially, simple, 3 or 4 pairs per segment. Laminae herbaceous,when dry dark green, with orange-yellow sphericalglands abaxially, with few acicular hairs along costae andcostules, adaxial sides with sparse acicular short hairs alonggrooves. Sori orbicular, medium-sized, 3 or 4 pairs per segment,dorsifixed at distal parts of lateral veins, close to margins; indusiaorbicular-reniform, medium-sized, brown, membranous,with more setae, persistent.On rocks in forests or on forest floor; 200–1100 m. Fujian, Taiwan[Japan].10. Parathelypteris glanduligera (Kunze) Ching, Acta Phytotax.Sin. 8: 301. 1963.金星蕨 jin xing juePlants 35–50(–60) cm tall. Rhizomes long creeping, glabrous,apices with sparse lanceolate scales. Fronds approximate;stipes 15–20(–30) cm, stramineous, ± with short hairs orsometimes glabrous; laminae lanceolate or broadly lanceolate,18–30 × 7–13 cm, not tapering to bases, pinnate-pinnatifid,acuminate and pinnatifid at apices; pinnae ca. 15 pairs, flat orobliquely spreading, alternate or subopposite proximally, sessile,lanceolate or linear-lanceolate, 4–7 × 1–1.5 cm, basessymmetrical and slightly broadened, or proximal pair slightlytapering to bases, truncate, pinnatifid nearly to costae, apicesacuminate; segments 15–20 pairs or more, spreading, oblonglanceolate,5–6 × ca. 2 mm, entire, rounded-obtuse or obtuse-pointedat apices; proximal pair, particularly acroscopicone, longer. Veins evident, lateral veins simple, 5–7 pairs persegment, proximal pair arising from above base of costules.Laminae herbaceous, when dry grass-green or sometimes darkgreen; pinnae, except for orange-yellow spherical glands, glabrousor shortly pubescent abaxially, adaxially with denseacicular hairs along grooves on costae, veins occasionally withfew short acicular hairs, rachises ± grayish white pubescent.Sori small, orbicular, 4 or 5 pairs per segment, dorsifixed nearends of lateral veins, close to margins; indusia medium-sized,orbicular-reniform, brown, thickly membranous, with sparsegrayish white setae, persistent. Spores bilateral, orbicular-reniform,perispores corrugate and regularly finely reticulate. 2n =144.Open forests, beneath Phyllostachys pubescens; sea level to 1500m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan,Henan, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang[N India, Japan, S Korea, Nepal, Vietnam].1a. Pinnae abaxially subglabrous, exceptfor sparse grayish white acicular hairsalong costae, adaxially with shortacicular hairs along costae,occasionally with few appressedshort acicular hairs along veins ....... 10a. var. glanduligera1b. Pinnae abaxially more denselypubescent, along costae with fineacicular hairs, adaxially withsparse appressed fine acicularhairs along veins and inintercostal areas except foracicular hairs along costal grooves ....... 10b. var. puberula10a. Parathelypteris glanduligera var. glanduligera金星蕨 ( 原变种 ) jin xing jue (yuan bian zhong)Aspidium glanduligerum Kunze, Analecta Pteridogr. 44.1837; A. gracilescens Blume var. glanduligerum (Kunze) Franchet& Savatier; Christella glanduligera (Kunze) H. Léveillé;Dryopteris glanduligera (Kunze) Christ; D. gracilescens(Blume) Kuntze var. glanduligera (Kunze) C. Christensen; D.repentula C. B. Clarke ex Christ; Lastrea glanduligera (Kunze)T. Moore; L. gracilescens Hooker var. glanduligera (Kunze)Beddome; Nephrodium glanduligera (Kunze) Makino; N.gracilescens (Blume) Hooker var. glanduligera (Kunze) Baker;Thelypteris glanduligera (Kunze) Ching.Pinnae abaxially subglabrous, except for sparse grayishwhite acicular hairs along costae and main veins, adaxially withshort acicular hairs along costae, occasionally with few appressedshort acicular hairs along veins.Forests; sea level to 1500 m. Anhui, Chongqing, Fujian, Guangdong,Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangsu, Jiangxi,Sichuan, Taiwan, Yunnan, Zhejiang [N India, Japan, S Korea, Vietnam].10b. Parathelypteris glanduligera var. puberula (Ching)Ching ex K. H. Shing, Fl. Jiangxi 1: 199. 1993.微毛金星蕨 wei mao jin xing jueThelypteris glanduligera var. puberula Ching, Bull. FanMem. Inst. Biol., Bot. 6: 323. 1936.Pinnae abaxially densely pubescent, along costae with

330THELYPTERIDACEAEfiner acicular hairs, adaxially also with sparse appressed fineacicular hairs along veins and in intercostal areas, acicular hairsalong costal grooves.● Wet shaded places beneath Phyllostachys pubescens; 200–1000m. Anhui, Jiangsu, Jiangxi.11. Parathelypteris subimmersa (Ching) Ching, Acta Phytotax.Sin. 8: 303. 1963.海南金星蕨 hai nan jin xing jueThelypteris subimmersa Ching, Bull. Fan Mem. Inst. Biol.,Bot. 6: 306. 1936.Plants 2.6–3 m tall. Rhizomes strong, woody and ascending.Fronds approximate; stipes nearly to 1 m, thick, ca. 1cm in diam., bases with thick lanceolate scales, distally withsparse short setae, dark stramineous; laminae large, oblong-lanceolate,160–200 × 50–60 cm, bases nearly tapering or sometimesproximal pair of pinnae slightly shortened, pinnate-pinnatifid,acuminate and pinnatifid at apices; pinnae mostly subopposite,spreading, sessile; middle pinnae linear-lanceolate,25–30 × 2–2.5 cm, bases symmetrical, truncate, pinnatifid andreaching both narrow wings of costae, apices caudate-acuminate;segments 60–70 pairs, flatly spreading, pectinatelyarranged, linear, 8–10 × 2.2–2.5 mm, entire, rounded or acuteat apices. Veins evident, lateral veins simple, 12–14 pairs persegment, proximal pair arising from base of costules, allreaching margins above sinuses or basal acroscopic veinletclose to sinus. Laminae papery, when dry grass-green, pinnaeabaxially ± with yellow spherical small glands along veins,elsewhere glabrous, adaxially along grooves with dense grayishwhite fine and long acicular hairs and along veins sparselyshortly hairy. Sori orbicular, medium-sized, dorsifixed at middleof lateral veins, 8–10 pairs per segment; indusia orbicularreniform,somewhat leathery, glabrous, sulfur-colored glandsalong margins, persistent. Spores orbicular-reniform, perisporesechinate.● On wet sandy soils in forests on slopes. W Hainan.Parathelypteris subimmersa is similar to Amphineuron immersum,which grows in Malaysia. The latter differs in having pinnae broader(3–3.3 cm), segments with dense grayish white fine and long acicularhairs abaxially. Owing to more complex characters on Amphineuron,except that the plant of this species is particularly large and strong, othercharacters do not differ, so it is listed here.12. Parathelypteris chingii K. H. Shing & J. F. Cheng, JiangxiSci. 8(3): 44. 1990.秦氏金星蕨 qin shi jin xing juePlants 35–60(–75) cm tall. Rhizomes short and erect.Fronds clustered; stipes 15–30 cm, bases occasionally withlanceolate scales and sparsely grayish white multicellularspreading acicular hairs, dark brown proximally, distallybrownish stramineous, sparsely shortly hairy or subglabrous;laminae oblong-lanceolate, 20–30 × 10–13 cm, not tapering tobases, pinnate-pinnatifid, acuminate and pinnatifid at apices;pinnae 12–15 pairs, sessile, spreading, proximal ones not shortened,reflexed, pinnae from middle of lamina downward lanceolate,5–8 × 1.4–1.7 cm, bases slightly broadened, roundedtruncate,symmetrical, pinnatifid and reaching both narrowwings of costae, apices acuminate; segments 8–15 pairs, proximalones oblong, 4–8 × 3–4 mm, slightly tapering to androunded-truncate at apices, sometimes angular and entire onsides. Veins evident, lateral veins simple, 3–5 pairs per segment,proximal pair arising from high above base of costules. Laminaethickly herbaceous, when dry grass-green or dark green,abaxially with sparse reddish purple spherical glands, alongcostae and veins with more grayish white spreading acicularhairs, adaxially along grooves with dense acicular hairs and appressedshort setae, also mixed with multicellular spreadingacicular hairs. Sori orbicular, dorsifixed at middle of veinletsbetween costules and margins, 2–5 pairs per segment; indusialarge, orbicular-reniform, brown, thickly membranous and separatedfrom each other and with grayish white acicular hairs.● Wet places in dense valley forests, forests at foot of mountains;300–500 m. N Fujian, Guangdong, S Jiangxi.Parathelypteris chingii is similar to P. japonica but differs byplants shorter and smaller, stipes with spreading acicular hairs at bases,pinnae with spreading fine and long acicular hairs abaxially, and indusiaseparated from each other.1a. Plants 35–60 cm tall; pinnae sessile, abaxiallyalong costae and veins with grayish whiteacicular hairs, and adaxially along costalgrooves with appressed short setae andmulticellular acicular hairs, rachises withdenser acicular hairs .................................. 12a. var. chingii1b. Plants to 75 cm tall; proximal pinnae stalked,along costae and veins with dense shorthairs except long acicular hairs abaxially,rachises with dense setae adaxially, withsparse grayish white fine and longacicular hairs abaxially ............................... 12b. var. major12a. Parathelypteris chingii var. chingii秦氏金星蕨 ( 原变种 ) qin shi jin xing jue (yuan bian zhong)Plants 35–60 cm tall. Pinnae sessile, abaxially along costaeand veins with grayish white acicular hairs, adaxially along costalgrooves with appressed short setae and multicellular acicularhairs.● Wet places in dense valley forests; 300–500 m. N Fujian, SJiangxi.12b. Parathelypteris chingii K. H. Shing var. major (Ching)K. H. Shing, Fl. Reipubl. Popularis Sin. 4(1): 46. 1999.大羽金星蕨 da yu jin xing jueThelypteris angulariloba Ching var. major Ching, Bull.Fan Mem. Inst. Biol., Bot. 6: 325. 1936.Plants to 75 cm tall. Proximal pinnae stalked, along costaeand veins with dense short hairs except for long acicular hairsabaxially; rachises with dense setae adaxially, with sparsegrayish white fine and long acicular hairs abaxially.● Wet shaded places in forests at foot of mountains. Guangdong.

THELYPTERIDACEAE 33113. Parathelypteris petelotii (Ching) Ching, Acta Phytotax.Sin. 8: 303. 1963.长毛金星蕨 chang mao jin xing jueThelypteris petelotii Ching, Bull. Fan Mem. Inst. Biol.,Bot. 6: 326. 1963; Coryphopteris petelotii (Ching) Holttum;Lastrea petelotii (Ching) Tagawa.Plants to 70 cm tall. Rhizomes thick and erect, ± trunklike.Fronds clustered; stipes ca. 30 cm, with sparse scales at basesand with dense grayish white multicellular acicular hairs, distallycastaneous-brown and subglabrous; laminae oblong-lanceolate,ca. 40 × 15 cm, slightly tapering to bases, pinnatepinnatifid,acuminate and pinnatifid at apices; pinnae ca. 20pairs, alternate, sessile, proximal ones somewhat shortened, reflexed,lanceolate, ca. 10 × 2–2.5 cm, bases truncate, symmetrical,pinnatifid nearly to costae, apices long acuminate; segments20–40 pairs, spreading, segments below middle narrowlyrectangular, ca. 1 × 0.3 cm, shallowly undulate or usuallysparsely bluntly serrate along margins, slightly bent distally,rounded at apices. Veins evident on both sides, lateral veinssimple, 6–8 pairs per segment, proximal pair arising from baseof costules. Laminae herbaceous, dark green when dry, withdense reddish purple spherical glands abaxially, costae andveins with dense grayish white multicellular acicular hairs;adaxially spreading appressed pubescent throughout; rachisesdark stramineous, pubescent adaxially, glabrous abaxially. Soriorbicular, dorsifixed at middle of lateral veins, 3–6 pairs persegment; indusia large, orbicular-reniform, thickly membranous,brown, with spreading grayish white long acicular hairsthroughout, persistent.Evergreen forests; ca. 1500 m. S Guangxi [N Vietnam].14. Parathelypteris hirsutipes (C. B. Clarke) Ching, Acta Phytotax.Sin. 8: 301. 1963.毛脚金星蕨 mao jiao jin xing jueNephrodium gracilescens (Blume) Hooker var. hirsutipesC. B. Clarke, Trans. Linn. Soc. London, Bot. 1: 514. 1880;Coryphopteris hirsutipes (C. B. Clarke) Holttum; Dryopterisgracilescens (Blume) Kuntze var. chinensis Christ; D. hirsutipes(C. B. Clarke) C. Christensen; Lastrea hirsutipes (C. B.Clarke) Beddome; Thelypteris hirsutipes (C. B. Clarke) Ching.Plants 35–60 cm tall. Rhizomes erect, strong, cylindric.Fronds clustered; stipes 10–20(–30) cm, proximal part darkbrown, bases with dense grayish brown multicellular spreadingacicular long hairs (when dry easily fallen) and few dark brownlanceolate scales, distally dark stramineous and subglabrous;laminae narrowly oblong, 25–30 × 9–14 cm, slightly tapering tobases, pinnate-pinnatifid, acuminate and pinnatifid at apices;pinnae 20–25 pairs, alternate, spreading, proximal 3 or 4 pairsshortened; middle pinnae lanceolate, 5–8 × 1–1.5 cm, basestruncate, symmetrical, pinnatifid nearly to costae, apices caudate-acuminate;segments 15–18 pairs, spreading, rectangular,4.5–6.5 × 2–3 mm, entire or slightly undulate at margins,rounded at apices. Veins visible on both sides, lateral veinssimple, 5–8 pairs per segment, proximal pair arising from basesof or slightly above costules. Laminae herbaceous, dark greenwhen dry, abaxially with denser reddish purple spherical glandsand costae and veins with denser multicellular acicular hairs,adaxially spreading appressed pubescent throughout; rachisesstramineous and sparsely pubescent on both sides. Sori orbicular,dorsifixed at middle of lateral veins, 1–5 pairs per segment;indusia large, orbicular-reniform, thickly membranous,brown, glabrous or occasionally sparsely shortly setaceous.Seasonal rain forests or mixed forests on mountains; 1400–1600m. SE Yunnan [N India, Myanmar].15. Parathelypteris caudata Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 320. 1999.尾羽金星蕨 wei yu jin xing juePlants 50–80 cm tall. Rhizomes erect, thick and short.Fronds clustered; stipes 20–40 cm, bases nearly black and withgrayish brown, spreading multicellular acicular hairs and fewbrown lanceolate scales, distally castaneous-brown, subglabrousor grooves with brownish acicular hairs; laminae oblonglanceolate,30–40 × 10–20 cm, slightly tapering to bases, pinnate-pinnatifid,acuminate and pinnatifid at apices; pinnae 20–25 pairs, alternate, spreading, sessile, proximal ones shortened,reflexed; largest pinna pair lanceolate, 4.5–9 × 1.2–2 cm, basesslightly tapering, rounded-truncate, symmetrical, pinnatifid andreaching narrow wing on both sides of costae, apices caudateor acuminate; segments 20–28 pairs, spreading, linear-lanceolateto rectangular, 5–9 × 2.5–3 mm, undulate-crenate orthickly serrate at margins, rounded or rounded-truncate atapices. Veins evident abaxially, lateral veins simple, 6–9 pairsper segment, proximal pair arising from base of or slightlyabove costules. Laminae papery or thickly herbaceous, whendry dark green, abaxially with reddish purple spherical glands,usually glabrous or along rachises, costae, and veins sparselygrayish brown pubescent; adaxially with appressed reddishbrown hairs, rachises and costal grooves with grayish brownacicular hairs. Sori orbicular, medium-sized, dorsifixed at middleof veinlets, 5–7 pairs per segment; indusia large, orbicularreniform,brown, thickly membranous, glabrous or occasionallyshortly setaceous.● Mossy forests or bamboo forests in subalpine areas on mountains;1700–1900 m. Guangxi, S Yunnan, and E China (exact localityunknown).16. Parathelypteris trichochlamys Ching ex K. H. Shing, Fl.Reipubl. Popularis Sin. 4(1): 320. 1999.毛盖金星蕨 mao gai jin xing juePlants to 50 cm tall. Rhizomes erect, short. Fronds clustered;stipes 18–20 cm, with acicular hairs and dark brownlinear-lanceolate scales, distally castaneous-red and includingrachises and costae with dense short acicular hairs; laminae 26–30 × ca. 6 cm, not tapering to bases, apices acuminate andpinnatifid; middle pinnae narrowly lanceolate, 5–6 × ca. 1 cm,bases slightly tapering and truncate, symmetrical, slightly bentdistally, pinnatifid and reaching both narrow wings, apices acuminate;segments ca. 16 pairs, oblong-triangular, 4–4.5 × ca. 3mm, margins entire and usually recurved when dry, slightlytapering to apices, apices obtuse. Veins evident, lateral veinssimple, ca. 5 pairs per segment, proximal pair arising from

332THELYPTERIDACEAEslightly above base of costules. Laminae thickly herbaceous,when dry brownish green, abaxially with sparse reddish purplespherical glands, with dense short acicular hairs on both surfaces.Sori orbicular, dorsifixed at middle of lateral veins; indusiamedium-sized, orbicular-reniform, close to each other,brown, thickly membranous, with dense short setae, persistent.● Wet places in thickets, common. W Guangdong (Dang Shan,Huaiji).17. Parathelypteris caoshanensis Ching ex K. H. Shing, Fl.Reipubl. Popularis Sin. 4(1): 50, 321. 1999 [“Cyclosorus” caoshanensis,p. 321].草山金星蕨 cao shan jin xing juePlants ca. 38 cm tall. Rhizomes erect, black, short. Frondsclustered; stipes 14–18 cm, bases dark brown, with densespreading acicular long hairs and few lanceolate scales, distallycastaneous red and to rachises with dense unicellular shortacicular hairs; laminae 20–22 × ca. 9 cm, nearly tapering tobases, pinnate-pinnatifid, apices acuminate and pinnatifid; pinnae11–15 pairs, alternate (proximal ones subopposite), obliquedistally, shortly stalked, proximal pair shortened or slightlyshortened; middle ones lanceolate, ca. 6 × 1 cm, basesrounded-truncate, symmetrical, pinnatifid nearly to costae,apices acuminate; segments ca. 18 pairs, ones below middleoblong, ca. 5 × 2.5 mm, entire along margins, rounded atapices. Veins evident, lateral veins simple, ca. 6 pairs per segment,proximal pair arising from near base of costules. Laminaeherbaceous, dark brown when dry, abaxially with spreadingshort acicular hairs throughout and including veins with reddishpurple spherical glands, adaxially with spreading hairs throughout.Sori orbicular, dorsifixed slightly above bases of lateralveins; indusia medium-sized, orbicular-reniform, brown, thinlyherbaceous, separated from each other, sparsely pubescent, persistent.● Wet soil of valley forests. Taiwan (Caoshan, Taibei).On one page of the protologue (appendix, p. 321), the name iscited as “Cyclosorus” caoshanensis, but this is obviously a typographicalerror. It appears under the generic heading Parathelypteris with sixother species of Parathelypteris, whereas numerous species of Cyclosorusappear later (pp. 331–350); it is not included in the index underCyclosorus (p. 364); and the name appears as P. caoshanensis in thesynopsis (p. ii), key (p. 32), main text (p. 50), and index (p. 377).18. Parathelypteris chinensis (Ching) Ching, Acta Phytotax.Sin. 8: 303. 1963.中华金星蕨 zhong hua jin xing juePlants 57–80 cm tall. Rhizomes short and decumbent orascending. Fronds approximate; stipes 27–40 cm, bases nearlyblack, with sparse lanceolate scales, distally castaneous-brownor reddish brown, glabrous, polished; laminae lanceolate, 30–40× 8–12 cm, not tapering to bases, pinnate-pinnatifid, apicesacuminate and pinnatifid; pinnae ca. 18 pairs, opposite oralternate distally, obliquely spreading, sessile, proximal pair notshortened; middle ones narrowly lanceolate, 5–7 × 0.8–1.2 cm,bases truncate, symmetrical, pinnatifid and reaching both narrowwings on costae, apices acuminate; segments 18–24 pairs,spreading, oblong or triangular-oblong, 3–5 × 2–3 mm, entirealong margins, obtuse at apices. Veins evident, lateral veinssimple, 4 or 5(or 6) pairs per segment, proximal pair arisingfrom slightly above base of costules. Laminae herbaceous,brownish green when dry, usually glabrous except for reddishorange spherical glands; adaxially pubescent along costae. Soriorbicular, dorsifixed at middle of lateral veins, ca. 3 pairs persegment; indusia large, orbicular-reniform, brown, membranous,nearly touching one another, usually glabrous, persistent.● Valley forests, thickets; 700–2100 m. S Anhui, N Fujian,Guangdong, S Guangxi, Guizhou, W Hunan, Jiangxi, S Sichuan, Yunnan,Zhejiang.1a. Laminae glabrous except for reddish orangeglands abaxially; sori nearly touchingeach other, indusia glabrous .................. 18a. var. chinensis1b. Laminae with sparse glands abaxiallyon costae, costae, veins, and intercostalareas sparsely shortly hairy; sori ±separated from each other; indusiawith sparse short hairs ...................... 18b. var. trichocarpa18a. Parathelypteris chinensis var. chinensis中华金星蕨 ( 原变种 ) zhong hua jin xing jue (yuan bianzhong)Thelypteris chinensis Ching, Bull. Fan Mem. Inst. Biol.,Bot. 6: 311. 1936; Aspidium parathelypteris Christ; Dryopterisjaponica (Baker) C. Christensen var. elongata Rosenstock; D.parathelypteris (Christ) C. Christensen; T. parathelypteris(Christ) Ching.Laminae glabrous except for reddish orange glands abaxially;sori nearly touching each other, indusia glabrous.● Wet shaded places in valley forests; 700–1000 m. S Anhui, NFujian, Guangdong, S Guangxi, W Hunan, Jiangxi, S Sichuan, Zhejiang.18b. Parathelypteris chinensis var. trichocarpa Ching ex K.H. Shing & J. F. Cheng, Jiangxi Sci. 8(3): 44. 1990.毛果金星蕨 mao guo jin xing jueLaminae with sparse glands abaxially on costae, costae,veins, and intercostal areas with sparse short acicular hairs; sori± separated from each other; indusia with sparse short hairs.● Valley forests, thickets; 700–2100 m. Guizhou, Jiangxi, Yunnan.19. Parathelypteris japonica (Baker) Ching, Acta Phytotax.Sin. 8: 301. 1963.光脚金星蕨 guang jiao jin xing juePlants 55–70 cm tall. Rhizomes short, decumbent or ascending.Fronds approximate or subclustered; stipes 25–35 cm,bases nearly black, with sparse reddish brown lanceolate scales,distally dark brown-castaneous or castaneous-brown or whollystramineous, glabrous; laminae ovate-oblong, 30–35 × 17–20cm, not tapering at bases, pinnate-pinnatifid, apices acuminateand pinnatifid; pinnae 15–20 pairs, spreading, proximal 3 or 4pairs of pinnae longer, opposite or subopposite, sessile; middlepinnae lanceolate, 8–10 × 1.3–1.6 cm, bases subtruncate, symmetrical,pinnatifid and reaching both narrow wings on costae,

THELYPTERIDACEAE 333apices acuminate; segments 25–30 pairs, lanceolate and slightlyfalcate, 5–7 × ca. 2.6 mm, entire, apices obtuse or acute. Veinsevident, lateral veins simple, 8 or 9(or 10) pairs per segment,proximal pair arising from near base of costules. Laminae herbaceous,dark green when dry, abaxially along costae, costae(sometimes including lateral veins) and margins with grayishwhite sparse pubescence and more reddish brown globose largeglands, adaxially along costal grooves with dense acicular hairs,rachises with appressed short acicular hairs. Sori orbicular, dorsifixedslightly above middle of lateral veins, 3 or 4 pairs persegment; indusia large, orbicular-reniform, brownish, membranous,grayish white pubescent or glabrous, persistent. 2n = 124.Forests; 200–2000 m. Anhui, N Fujian, Guizhou, Hunan, N Jiangsu,Jiangxi, W Sichuan, Taiwan, N Yunnan, S and W Zhejiang [Japan, SKorea].1a. Stipes and rachises nearly black at bases,distally castaneous-brown; indusiasomewhat glaucous, pubescent .............. 19a. var. japonica1b. Stipes and rachises stramineous,pinnae abaxially and indusia glabrous ... 19b. var. glabrata19a. Parathelypteris japonica var. japonica光脚金星蕨 ( 原变种 ) guang jiao jin xing jue (yuan bianzhong)Nephrodium japonicum Baker, Ann. Bot. (Oxford) 5: 318.1891; Christella japonica (Baker) H. Léveillé; Coryphopterisjaponica (Baker) L. J. He & X. C. Zhang; Dryopteris japonica(Baker) C. Christensen; Lastrea japonica (Baker) Copeland;Parathelypteris japonica var. musashiensis (Hiyama) Ching;Thelypteris japonica (Baker) Ching; T. japonica var. musashiensisHiyama; Wagneriopteris japonica (Baker) A. Löve& D. Löve.Stipes and rachises nearly black at bases, distally castaneous-brown,pinnae abaxially and indusia sparsely pubescent, indusiamore glaucous, pubescent.Shaded places in forests; 200–2000 m. N Fujian, Guizhou, NJiangsu, Jiangxi, W Sichuan, Taiwan [Japan, S Korea].19b. Parathelypteris japonica var. glabrata (Ching) K. H.Shing, Fl. Jiangxi 1: 201. 1993.光叶金星蕨 guang ye jin xing jueThelypteris japonica var. glabrata Ching, Bull. Fan Mem.Inst. Biol., Bot. 6: 313. 1936.Stipes and rachises stramineous, pinnae abaxially and indusiaglabrous.Forests. Jiangxi [Japan, S Korea].20. Parathelypteris castanea (Tagawa) Ching, Acta Phytotax.Sin. 8: 302. 1963.台湾金星蕨 tai wan jin xing jueDryopteris castanea Tagawa, Acta Phytotax. Geobot. 4:132. 1935; Thelypteris castanea (Tagawa) Ching.Plants ca. 70 cm tall. Rhizomes short and ascending, black.Fronds clustered; stipes with bases black, distally castaneousbrown,polished, throughout covered with dense glaucous shortacicular hairs; laminae narrowly oblong, ca. 30 × ca. 14 cm,bases not tapering, pinnate-pinnatifid, apices acuminate andpinnatifid; pinnae 15–17 pairs, opposite or alternate distally,spreading, sessile, proximal several pairs not shortened, lanceolate,± falcate, 7–8 × ca. 1.4 cm, bases slightly tapering,truncate, pinnatifid and reaching narrow wings on both sides,apices acuminate; segments 15–18 pairs, ca. 6 × 2.8 mm, entire,apices rounded. Veins visible, lateral veins simple, 6 or 7 pairsper segment, proximal pair arising from base of costules.Laminae herbaceous, dark brown when dry, pinnae with densegrayish white short acicular hairs on both surfaces, hairs alongcostae denser and longer; rachises brown, grayish white pubescentabaxially. Sori orbicular, dorsifixed above middle oflateral veins, slightly closer to margins, 1 pair per segment; indusiasmaller, orbicular-reniform, brown, membranous, denselypubescent, persistent.Wet places in forests. Taiwan [Japan].Some authors treat this name as a synonym of the precedingspecies, Parathelypteris japonica (e.g., see Knapp, Ferns Fern AlliesTaiwan, 416. 2011, as Thelypteris japonica).21. Parathelypteris pauciloba Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 321. 1999.阔片金星蕨 kuo pian jin xing juePlants 30–35 cm tall. Rhizomes short and erect, black.Frond clustered; stipes 12–14 cm, slender, bases nearly black,mixed with grayish white setae and few spreading multicellularlong hairs, distally castaneous brown and with grayish whitesetae; laminae oblong, 17–22 × 9–12 cm, not tapering at bases,pinnate-pinnatifid, apices acuminate and pinnatifid; pinnae 10–12 pairs, alternate, sessile, proximal pinnae not shortened, 5–6.5× 1.2–1.6 cm, bases slightly tapering, truncate, symmetrical,pinnatifid and reaching broad wing on both sides of costae,apices acuminate; pinnae 5–8 pairs, approximate, rectangular,5–7 × ca. 5 mm, apices rounded or rounded-truncate and with2–4 angles. Veinlets evident, lateral veins simple, 3 or 4 pairsper segment, proximal pair arising from above base of costules.Laminae herbaceous, when dry deep green, abaxially withsparse short setae along costae, adaxially with sparse appressedshort hairs, hairs along costal grooves denser; rachises stramineous,with dense short setae. Sori orbicular, 1 or 2 per segment,dorsifixed at slightly high places at middle of lateralveins, in one row on each side of costules; indusia mediumsized,orbicular-reniform, brown, thickly membranous, slightlypubescent, persistent.● Wet soil at streamsides in forests; ca. 800 m. N Fujian.Parathelypteris pauciloba is similar to P. angulariloba, but thelatter differs in having rhizomes short and erect, stipes throughout withgrayish white short setae and mixed few spreading multicellular longacicular hairs, costae with sparse unicellular short setae abaxially, andindusia slightly pubescent.22. Parathelypteris angulariloba (Ching) Ching, Acta Phytotax.Sin. 8: 304. 1963.钝角金星蕨 dun jiao jin xing jueThelypteris angulariloba Ching, Bull. Fan Mem. Inst.

334THELYPTERIDACEAEBiol., Bot. 6: 323. 1936; Coryphopteris angulariloba (Ching) L.J. He & X. C. Zhang; Lastrea angulariloba (Ching) Tagawa;L. simozawae (Tagawa) Tagawa; Parathelypteris simozawae(Tagawa) Ching; T. simozawae Tagawa; Wagneriopteris angulariloba(Ching) A. Löve & D. Löve.Plants 30–60 cm tall. Rhizomes short, decumbent or ascending,nearly black. Fronds subclustered; stipes 10–30 cm,bases nearly black, with dense spreading multicellular acicularhairs, distally castaneous-red or castaneous-brown, subglabrous;laminae narrowly oblong, 17–30 × 6–12 cm, bases nottapering, pinnate-pinnatifid, apices acuminate and pinnatifid;pinnae ca. 20 pairs, alternate, proximal pair not reduced, ± reflexed;middle pinnae lanceolate or linear-lanceolate, 5–6 ×0.7–1.5 cm, bases truncate, ± symmetrical, sessile, pinnatifidand reaching 1/3–1/2, apices acuminate and pinnatifid or sometimesentire; segments 8–12 pairs, rectangular or subsquare, 3–5 × ca. 3.5 mm, entire, apices rounded or rounded-truncateand with 2–4 blunt angles. Veins evident, lateral veins simple,2 or 3(or 4) pairs per segment, proximal pair arising fromabove base of costules. Laminae thickly herbaceous, greenishwhen dry, abaxially along costae and main veins with multicellularshort acicular hairs, sometimes with mixed orangecapitate glands; adaxially along costal grooves with acicularhairs, elsewhere glabrous. Sori orbicular, dorsifixed at middleof lateral veins, 1 or 2 pairs per segment; indusia medium-sized,orbicular-reniform, brown, thickly membranous, with densegrayish white short setae, persistent.Streamsides in valley forests, shaded wet places in thickets; 400–1100 m. N and SE Fujian, N and SE Guangdong, E Guangxi, Taiwan[Japan].23. Parathelypteris nigrescens Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 321. 1999.黑叶金星蕨 hei ye jin xing juePlants 50–70 cm tall. Rhizomes short and erect, black.Fronds clustered; stipes 20–35 cm, nearly black on proximalpart and with spreading grayish white multicellular long acicularhairs, distally castaneous-brown and sparsely pubescent;laminae oblong or narrowly oblong, 30–38 × 12–15 cm, basesnot tapering, pinnate-pinnatifid, apices acuminate and pinnatifid;pinnae 15–18 pairs, spreading, alternate, proximal onessometimes stalked, proximal pair not shortened; middle pinnaelanceolate, slightly falcate, 8–11 × 1.2–1.5 cm, bases truncate,symmetrical, pinnatifid and reaching narrow wings on bothsides of costae, apices long acuminate; segments 15–20 pairs,obliquely spreading, narrowly ligulate, 5–7 × 3–3.5 mm, entirealong both sides, rounded-obtuse at apices. Veins visible, lateralveins simple, 5 or 6 pairs per segment, proximal pair arisingfrom base of costules. Laminae herbaceous, dark green whendry, or blackish brown, abaxially with sparse short hairs, costaedensely pubescent, adaxially along costae with dense short acicularhairs. Sori orbicular, dorsifixed at middle of lateral veins;indusia medium-sized, orbicular-reniform, brown, membranous,separated from each other, entire or occasionally with 1 or2 hairs, persistent or deciduous.● Streamsides in valley forests; 1000–1200 m. Guangxi, Yunnan.24. Parathelypteris indochinensis (Christ) Ching, Acta Phytotax.Sin. 8: 304. 1963.滇越金星蕨 dian yue jin xing jueDryopteris indochinensis Christ, J. Bot. (Morot), ser. 2, 1:263. 1908; Lastrea indochinensis (Christ) Tagawa; Thelypterisindochinensis (Christ) Ching.Plants ca. 60 cm tall. Rhizomes short and decumbent.Fronds subclustered; stipes ca. 30 cm, proximal part black,polished, with denser grayish white multicellular long acicularhairs, distally castaneous-brown, sparsely pubescent; laminaeoblong, ca. 30 × 20–40 cm, bases not tapering, pinnate-pinnatifid,apices acuminate and pinnatifid; pinnae ca. 15 pairs,spreading, alternate, sessile, proximal pair not shortened; middlepinnae lanceolate, 10–12 × ca. 2 cm, bases not tapering,truncate, symmetrical, pinnatifid and reaching narrow wings onboth sides of costae, apices acuminate; segments ca. 24 pairs,linear-lanceolate, slightly falcate, slightly tapering to apices,rounded-truncate, entire. Veins evident, lateral veins simple, 6or 7 pairs per segment, proximal pair arising above base of costules.Laminae herbaceous, when dry grayish green, abaxiallywith dense multicellular barblike long hairs, adaxially alongcostal grooves with dense long acicular hairs, main veinsslightly setaceous or subglabrous, rachises with dense longacicular hairs adaxially. Sori orbicular, dorsifixed at middle oflateral veins, 5 or 6 pairs per segment; indusia smaller, orbicular-reniform,brown, membranous, separated from each other,densely pubescent, persistent.Shaded wet places in valley forests. S Guangxi, S Yunnan [borderof Vietnam].4. METATHELYPTERIS (H. Itô) Ching, Acta Phytotax. Sin. 8: 305. 1963.凸轴蕨属 tu zhou jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiThelypteris sect. Metathelypteris H. Itô in Nakai & Honda, Nov. Fl. Jap. 4: 137. 1939.Plants small to medium-sized, terrestrial. Rhizomes short, decumbent, ascending or erect, rarely long creeping, covered withbrown lanceolate scales and glaucous short hairs or subglabrous. Fronds approximate or clustered; stipes ± dark brown at bases,distally stramineous, glabrous or sparsely hairy; laminae oblong, lanceolate, or ovate-triangular, pinnate-pinnatifid, rarely tripinnate,if latter then pinnules separated from each other, never connected with narrow wing along costae, apices acuminate and pinnatifid.Laminae herbaceous or thinly herbaceous, when dry usually green, both surfaces ± with grayish white unicellular (rarely multicellular)acicular hairs, hairs denser along rachises and costae, pinnae usually glandless abaxially, rarely orange-red sphericalglandular, costae rounded and raised adaxially, never grooved. Veinlets pinnate, simple or forked, oblique distally, not reaching

THELYPTERIDACEAE 335margins. Sori small, orbicular, attached above middle of veinlets; indusia medium-sized, orbicular-reniform, each attached at a sinus,membranous, usually green, when dry grayish yellow or brownish, persistent. Spores bilateral, perispores corrugate, usually foveolate;exospore with finely reticulate surfaces. x = 35.About 12 species: S Asia, S China to Japan, Madagascar, Malesia; 11 species (five endemic) in China.The following taxon is excluded from the present treatment, pending further research: Metathelypteris glandulosa H. G. Zhou & H. Li (ActaBot. Yunnan. 14: 34. 1992), described from Guangxi.1a. Pinnae with orange-red spherical glands or eglandular abaxially, costae with grayish white multicellularacicular hairs abaxially.2a. Pinnae glandular abaxially, abaxially with unicellular short acicular hairs along lateral veins andintercostal areas ............................................................................................................................................... 10. M. glandulifera2b. Pinnae without glands, with grayish white multicellular spreading long acicular hairs on both surfaces ........... 11. M. flaccida1b. Pinnae without orange-red spherical glands abaxially, costae with grayish white unicellular acicular hairs orsubglabrous abaxially.3a. Laminae ovate-triangular.4a. Laminae bipinnatipartite, segments entire .............................................................................................. 6. M. deltoideofrons4b. Laminae pinnate-pinnatifid or 3-pinnate to 3-pinnate-pinnatifid.5a. Laminae oblong, except for costae long villous adaxially, elsewhere glabrous; indusiaglabrous ................................................................................................................................................ 9. M. wuyishanica5b. Laminae ovate-triangular, hairy on both surfaces; also hairy at indusia.6a. Proximal pinnae sessile or with 0.5–1 mm stalk, pinnules rounded-obtuse or acute at apices,sessile .................................................................................................................................................... 7. M. hattorii6b. Proximal pinnae with 3.5–5 mm stalk, pinnules long acuminate-caudate at apices, stalk4–7 mm ........................................................................................................................................... 8. M. petiolulata3b. Laminae oblong or lanceolate.7a. Laminae thick, papery or thinly herbaceous; plants taller and larger, usually 75–95 cm tall or more;pinnae glabrous abaxially, or costae and costules occasionally with sparse acicular short hairs ........... 1. M. singalanensis7b. Laminae thinner, herbaceous or thinly herbaceous; plants smaller and shorter, no more than 65 cmtall; pinnae ± with short acicular hairs or glabrous abaxially.8a. Pinnae glabrous abaxially, or at most with hairs along costae, costae occasionally with very sparseacicular hairs.9a. Proximal pinnae usually not shortened, also not tapering to bases; laminae adaxially withdense grayish white acicular hairs along rachises and costae; veinlets usually simple,occasionally forked ....................................................................................................................... 2. M. gracilescens9b. Laminae with proximal 1 or 2 pairs of pinnae ± shortened, pinna bases clearly tapering;laminae adaxially with grayish white short acicular hairs along rachises and costae;veinlets on pinnules of proximal pinnae usually forked, simple distally ...................................... 3. M. adscendens8b. Pinnae abaxially at least with denser short acicular hairs along costae.10a. Pinnae arrangement sparser, proximal ones 2–4 cm from each other, bases clearly tapering;segments entire or usually with densely crenate sinuses, or pinnatifid .................................................. 4. M. laxa10b. Pinnae arrangement denser, proximal ones 1–2 cm from each other, bases nearly tapering,segments usually entire, at most shallowly undulate ...................................................................... 5. M. uraiensis1. Metathelypteris singalanensis (Baker) Ching, Acta Phytotax.Sin. 8: 306. 1963.鲜绿凸轴蕨 xian lü tu zhou jueNephrodium singalanense Baker, J. Bot. 18: 212. 1880;Dryopteris media Alderwerelt; D. singalanensis (Baker) C.Christensen; Lastrea singalanensis (Baker) Beddome; Thelypterissingalanensis (Baker) Ching.Plants 75–95 cm tall. Rhizomes thick and short, erect,apices including stipe bases with dense scales; scales brown,linear-lanceolate, caudate at apices. Fronds clustered; stipes 30–50 cm, stramineous, glabrous above bases, polished; laminaelanceolate to broadly lanceolate, 30–70 × 20–30 cm, bases ±tapering, rounded-truncate, pinnate-pinnatifid, apices acuminateand pinnatifid; pinnae 15–25 pairs, spreading or oblique distally,sessile, proximal pair usually slightly shortened, distalones linear-lanceolate, sometimes falcate, 12–18 × 2–3.5 cm,bases subtruncate, pinnatifid nearly to costae, apices caudateacuminate;segments 20–30 pairs, basiscopic ones on costaelonger than acroscopic ones, oblong, 8–15 × 3–5 mm, entire orserrate along margins, apices blunt-pointed. Veinlets evident,simple or forked on proximal segments, 6–10(–12) pairs, notreaching margins. Laminae papery, when dry green or yellowishgreen, glabrous abaxially, or sometimes with sparseshort acicular hairs along costae and costules, adaxially withgrayish white short acicular hairs along rachises and costae.Sori small, glandular, attached on ends of lateral veins or middleof acroscopic vein on forked lateral veins; indusia small,orbicular-reniform, membranous, when dry brownish yellow,sometimes sinuate along margins, glabrous or sparsely capitateshortly hairy. 2n = 144.

336THELYPTERIDACEAEStreamsides in forests; 800–1000 m. SE Hainan [Indonesia, Malaysia,Thailand].2. Metathelypteris gracilescens (Blume) Ching, Acta Phytotax.Sin. 8: 305. 1963.凸轴蕨 tu zhou jueAspidium gracilescens Blume, Enum. Pl. Javae 2: 155.1828; Dryopteris arisanensis Rosenstock; D. gracilescens(Blume) Kuntze; D. sublaxa Hayata; Lastrea gracilescens(Blume) T. Moore (1858), not Hooker (1857); Nephrodiumgracilescens (Blume) Hooker; Thelypteris gracilescens (Blume)Ching.Plants 40–60 cm tall. Rhizomes short, decumbent or ascending,apices including stipe bases covered with reddishbrown lanceolate small scales. Fronds subclustered; stipes 15–30 cm, stramineous or tan, glabrous or with sparse short hairs;laminae narrowly oblong, 20–30 × 6.5–10 cm, pinnate-pinnatifid,apices acuminate and pinnatifid; pinnae 15–18 pairs, suboppositeor alternate distally, spreading, proximal 1 or 2 pairs notshortened, tapering to bases, reflexed proximally, sessile; middleones linear-lanceolate, 4–6.5 × 1–1.4 cm, bases tapering,subtruncate, pinnatifid and reaching narrow wings on both sidesof costae, apices acuminate; segments 12–15 pairs, oblique distally,oblong, 4–6 × 2–4 mm, entire or slightly undulate-crenatedistally, apices rounded-obtuse. Veinlets evident abaxially, simple,occasionally forked, 5 or 6 pairs per segment, proximal pairarising from above base of costules. Laminae herbaceous, yellowishgreen when dry, abaxially glabrous or costae adaxiallywith sparse short hairs, adaxially with dense grayish white shortacicular hairs, also veins with sparse similar hairs adaxially.Sori small, orbicular, 3 or 4 pairs per segment, dorsifixed atmiddle of lateral veinlets, located between costules and margins;indusia small, orbicular-reniform, membranous, brownish,glabrous, persistent or sometimes deciduous.On soil in dense forests on mountains; 1000–2500 m. Taiwan,Yunnan [Indonesia, S Japan, Malaysia, Philippines; Pacific islands(Polynesia)].3. Metathelypteris adscendens (Ching) Ching, Acta Phytotax.Sin. 8: 306. 1963.微毛凸轴蕨 wei mao tu zhou jueThelypteris adscendens Ching, Bull. Fan Mem. Inst. Biol.,Bot. 6: 332. 1936.Plants 25–50 cm tall. Rhizomes short, decumbent, withsparse short hairs and brown ovate-lanceolate scales. Frondsclustered or approximate; stipes 10–25 cm, stramineous, glabrousabove bases; laminae 15–25 × 8–12 cm, ± tapering tobases, pinnate-pinnatifid, apices long acuminate and pinnatifid;pinnae 10–15 pairs, alternate, sessile, proximal 1 or 2 pairs ±shortened, bases slightly tapering; middle ones narrowly lanceolate,4–6 × 1–1.5 cm, pinnatifid and reaching narrow wingson both sides of costae, long acuminate at apices, sometimesslightly caudate; segments 10–14 pairs, spreading, approximate,oblong-lanceolate, 4–6 × ca. 2.5 mm, entire or densely serratein sinus along margins on proximal segments, rounded-obtuseat apices. Veins visible, veinlets usually forked on proximal segments,distal ones simple, 3–5 pairs per segment, proximal pairarising from slightly higher above base of costules, not reachingmargins. Laminae herbaceous, when dry green, glabrous onboth surfaces, only occasionally with sparse short hairs alongboth sides of costae. Sori small, orbicular, 2–4(or 5) pairs persegment, attached near ends of lateral veins, close to margins;indusia small, orbicular-reniform, membranous, green, orbrownish when dry, glabrous or occasionally with several shorthairs, usually deciduous when mature.● Valley forests; 200–700(–?1800) m. Fujian, Guangdong,Guangxi, Taiwan.4. Metathelypteris laxa (Franchet & Savatier) Ching, ActaPhytotax. Sin. 8: 306. 1963.疏羽凸轴蕨 shu yu tu zhou jueAspidium laxum Franchet & Savatier, Enum. Pl. Jap. 2:237. 1876; Dryopteris laxa (Franchet & Savatier) C. Christensen;D. macarthyi (Baker) C. Christensen; Lastrea laxa (Franchet& Savatier) Copeland; Nephrodium laxum (Franchet &Savatier) Diels; N. macarthyi Baker; Thelypteris laxa (Franchet& Savatier) Ching.Plants 30–60 cm tall. Rhizomes long, decumbent or ascending,with sparse grayish white short hairs and reddishbrown lanceolate scales, similar indument on stipe bases. Frondsapproximate; stipes 10–35 cm, stramineous, above bases subglabrous;laminae narrowly oblong, 15–35 × 10–18 cm, basesnearly tapering, pinnate-pinnatifid, apices acuminate and pinnatifid;pinnae 8–18 pairs, subopposite, linear-lanceolate, 5–9 ×1–2 cm, bases truncate, ± symmetrical, sessile, pinnatifid andreaching narrow wings on both sides of costae; segments oblong-lanceolate,middle ones 4–8 × 2–3 mm, entire or denselycrenate-sinuate, or parted into small segments, obtuse-pointedor acute at apices. Veins visible, veinlets forked on segments onproximal pinnae, others simple, 5–7 pairs per segment, proximalpair arising above base of costules, not reaching margins.Laminae herbaceous, green when dry, abaxially with spreadinghairs throughout, adaxially with acicular hairs along rachises,costae, and veins. Sori small, orbicular, 4–6 pairs per segment,attached on ends of veinlets or acroscopic vein on forked veinlets,close to margins; indusia small, orbicular-reniform, membranous,green, when dry grayish yellow, sparsely pubescent.Forests on mountains, dense valley forests; near sea level to 800m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan,Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang[Japan, S Korea].5. Metathelypteris uraiensis (Rosenstock) Ching, Acta Phytotax.Sin. 8: 306. 1963.乌来凸轴蕨 wu lai tu zhou juePlants 30–40 cm tall. Rhizomes short, decumbent or ascending,apices including stipe bases with sparse deep brownsmall scales and grayish white short acicular hairs. Fronds subclustered;stipes 14–20 cm, stramineous, glaucous, shortly hairy;laminae oblong-lanceolate, 16–22 × 8–15 cm, not tapering tobases, pinnate-pinnatifid, apices long acuminate and pinnatifid;pinnae 12–15 pairs, opposite or distal ones alternate, sessile,

THELYPTERIDACEAE 337proximal pair usually slightly shortened, bases slightly tapering,reflexed, above several pairs linear-lanceolate, 4–8 × 1.2–1.7cm, bases tapering, rounded-truncate, symmetrical, pinnatifidand reaching narrow wings on both sides of costae, apicescaudate-acuminate; segments 14–20 pairs, spreading, approximate,oblong-lanceolate, 4–7 × 2–3 mm, entire or sometimesundulate along margins, rounded-obtuse at apices. Veins evidentabaxially, veinlets usually forked, or distal ones simple, 5–7 pairs per segment, proximal pair arising from above base ofcostules. Laminae thinly herbaceous, when dry yellowish green,abaxially with grayish white short hairs, hairs along rachisesand costae denser, adaxially densely covered with grayish yellowacicular hairs along rachises and costae. Sori small, orbicular,2–4 pairs per segment, attached near ends of lateralveins, close to margins; indusia small, orbicular-reniform, membranous,green, when dry brownish, sometimes lacerate alongmargins and acicular hairy, persistent. 2n = 124.Streamsides in valley forests, broad-leaved forests on slopes; 400–1700 m. N Guangdong, Taiwan, SE Xizang, W Yunnan [Japan, Philippines].1a. Proximal pair of pinnae slightly shortened,bases slightly tapering, lateral veins forkedon proximal pinnae, intercostal areasabaxially with short hairs ......................... 5a. var. uraiensis1b. Proximal pair of pinnae of similar shapeand size as distal ones, lateral veinssimple, occasionally forked, pinnaeabaxially, except for grayish whiteshort acicular hairs along rachisesand costae, glabrous ................................... 5b. var. tibetica5a. Metathelypteris uraiensis var. uraiensis乌来凸轴蕨 ( 原变种 ) wu lai tu zhou jue (yuan bian zhong)Dryopteris uraiensis Rosenstock, Hedwigia 56: 341. 1915;D. hirsutisquama Hayata; Lastrea uraiensis (Rosenstock) Copeland;Macrothelypteris uraiensis (Rosenstock) A. Löve & D.Löve; Thelypteris uraiensis (Rosenstock) Ching.Proximal pair of pinnae slightly shortened, bases slightlytapering; lateral veins forked on segments of proximal pinnae,intercostal areas abaxially with short hairs.Streamsides in valley forests; 400–1600 m. N Guangdong, Taiwan,W Yunnan [Japan, Philippines].5b. Metathelypteris uraiensis var. tibetica (Ching & S. K.Wu) K. H. Shing, Fl. Reipubl. Popularis Sin. 4(1): 67. 1999.西藏凸轴蕨 xi zang tu zhou jueMetathelypteris tibetica Ching & S. K. Wu, Fl. Xizang. 1:166. 1983.Proximal pair of pinnae of similar shape and size as distalones; lateral veins simple, occasionally forked, pinnaeabaxially, except for grayish white short acicular hairs alongrachises and costae, glabrous.● Broad-leaved forests on slopes; ca. 1700 m. SE Xizang (Mêdog).6. Metathelypteris deltoideofrons Ching ex W. M. Chu & S.G. Lu, Fl. Yunnan. 20: 720. 2006.三角叶凸轴蕨 san jiao ye tu zhou juePlants 23–55 cm tall. Rhizomes short, ascending, includingstipe bases with grayish white acicular hairs and fewbrown linear-lanceolate small scales. Fronds approximate;stipes 10–30 cm, stramineous, subglabrous above bases; laminaeovate-triangular, 13–30 × 10–16 cm, broadest at bases, pinnate-pinnatifid,long acuminate and pinnatifid at apices; pinnae10–14 pairs, spreading, proximal ones subopposite, distal onesalternate, proximal pinnae largest, bent distally, linear-lanceolate,5–9.5 × 1.2–2.2 cm, bases truncate, sessile, pinnatifid andreaching narrow wings on both sides of costae, apices long acuminate;segments 12–20 pairs, 5–10 × 2–3 mm, entire, obtusepointedat apices. Veins evident abaxially, lateral veins simpleor forked, 5 or 6 pairs per segment, not reaching margins.Laminae thinly herbaceous, when dry yellowish green, withgrayish white short acicular hairs on both surfaces, hairs alongrachises and costae denser. Sori small, orbicular, attached nearends of lateral veins, close to margins; indusia small, orbicularreniform,thinly membranous, green, when dry grayish brown,with dense acicular hairs, persistent.● Rock crevices at streamsides, streamsides in bamboo forests;600–2200 m. Hunan, Yunnan.The FRPS account of Metathelypteris misidentified material of M.deltoideofrons as M. decipiens (C. B. Clarke) Ching. Here, we followFraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes, 592.2008) and treat the latter as a synonym of species no. 11, M. flaccida.7. Metathelypteris hattorii (H. Itô) Ching, Acta Phytotax. Sin.8: 306. 1963 [“hattori”].林下凸轴蕨 lin xia tu zhou jueDryopteris hattorii H. Itô, Bot. Mag. (Tokyo) 99: 359.1935 [“hattori”]; D. laxa (Franchet & Savatier) C. Christensenvar. dilatata Koidzumi; Lastrea hattorii (H. Itô) Tagawa; L.laxa (Franchet & Savatier) Copeland var. dilatata (Koidzumi)Honda; Thelypteris hattorii (H. Itô) Tagawa; T. nemoralisChing.Plants 30–60 cm tall. Rhizomes short, decumbent, apicesincluding stipe bases densely covered with reddish brown lanceolatescales and grayish white setae. Fronds approximate;stipes 15–30 cm, bases dark brown, stramineous above bases,subglabrous; laminae ovate-triangular, 15–35 × 14–26 cm,broadest at bases, bases rounded-truncate, 3-pinnate-pinnatifid,apices acuminate and pinnatifid; pinnae 12–16 pairs, proximalones subopposite, distal ones alternate, sessile, or proximalpinnae sometimes with 0.5–1 mm stalks; proximal pair notshortened, of similar shape and size as distal ones, lanceolate,10–15 × 2.5–3.5 cm, pinnae not tapering to base except forproximal 2 pairs, rounded-truncate at bases, pinnate-pinnatifid,acuminate at apices; pinnules ca. 16 pairs, subopposite, pinnaedistal to middle ones connected to each other by narrow wings,proximal ones oblong-lanceolate, 2.5–3 × ca. 1 cm, bases decurrent,sessile, separated from each other, pinnatifid to ca. 2/3,apices rounded-obtuse or acute; segments oblong, 3.5–4 × ca.2.4 mm, entire, rounded-obtuse at apices. Veins not evident,

338THELYPTERIDACEAElateral veins simple or forked, 2 or 3 pairs per segment, notreaching margins. Laminae herbaceous, when dry green,densely grayish white pubescent on both surfaces. Sori small,orbicular, usually 1 per segment, attached near ends of acroscopicveinlets on bases and close to margins; indusia small,orbicular-reniform, membranous, when dry grayish brown,sparsely pubescent, persistent.Valley forests; 100–1700 m. S Anhui, N Fujian, N Guangxi, Hunan,Jiangxi, SW Sichuan, Zhejiang [Japan].8. Metathelypteris petiolulata Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 321. 1999.有柄凸轴蕨 you bing tu zhou juePlants 55–65 cm tall. Rhizomes short and decumbent.Fronds subclustered; stipes 23–30 cm, dark brown, with densegrayish white acicular hairs, distally stramineous, subglabrous,polished. Laminae ovate-triangular, 30–40 cm, broadest atbases and similar in size as length, bases broadly cordate, 3-pinnate to 4-pinnatifid, apices acuminate and pinnatifid; pinnae10–12 pairs, ascending, proximal ones opposite or subopposite,stalk 3.5–5 cm, distal ones alternate, sessile; proximal pairlargest, triangular-lanceolate, 17–22 × 10–12 cm, bases subtruncate,2-pinnate to 2-pinnate-pinnatifid, apices caudateacuminate;pinnules 10–15 pairs, alternate or subopposite,approximate, short stalks of proximal several pairs 4–7 mm,distal ones sessile, basiscopic pinnules of costae longer thanacroscopic ones, but proximal pair of pinnules slightly shortenedsometimes, above ones lanceolate, 6–7 × ca. 2.5 cm, basesbroadly cuneate, pinnatifid, apices caudate-acuminate; pinnulesof 2-pinnate laminae ca. 10 pairs, proximal 2 or 3 pairs slightlyshortened, narrowly oblong, 6–15 × 3–4 mm, bases adnate tocostules, decurrent, connected to each other by narrow wings,margins lobed to 1/2 of distance to costule, apices roundedobtuseor acute; segments 4 or 5 pairs, triangular, entire, obtusepointedat apices. Veins not evident, lateral veins usuallyforked, 2–6 pairs per ultimate pinnule or segment and notreaching margins. Laminae thinly herbaceous, when dry yellowishgreen, with sparse grayish white short hairs on bothsurfaces. Sori orbicular, 1–5 pairs per pinnule or segment,attached near ends of acroscopic vein of forked veins, slightlycloser to margins; indusia small, orbicular-reniform, membranous,green, with short hairs, persistent.● Shaded wet places in valley forests; 800–1500 m. Anhui, Fujian,NW Jiangxi, S Zhejiang.Metathelypteris petiolulata is similar to M. hattorii but differs byits laminae broader, 3-pinnate to 3-pinnate-pinnatifid, proximal pinnaeand pinnules of 1-pinnate laminae stalked, and apices of pinnules of 1-pinnate laminae caudate-acuminate.9. Metathelypteris wuyishanica Ching, Wuyi Sci. J. 1: 5. 1981.武夷山凸轴蕨 wu yi shan tu zhou juePlants to 40 cm tall. Rhizomes short and erect. Frondsclustered; stipes ca. 20 cm, stramineous, bases with dense darkbrown lanceolate scales and glaucous acicular hairs, distally glabrous;laminae oblong, ca. 25 × 15 cm, 2-pinnate-pinnatifid,apices acuminate; pinnae ca. 10 pairs, spreading, proximal pairsimilar in size as above ones, oblong-lanceolate, ca. 7 × 3 cm,shortly stalked, pinnatifid; pinnules ca. 14 pairs, basiscopicones on costae longer than acroscopic ones, lanceolate, ca. 2 ×0.5 cm, bases connected by narrow wings, pinnatifid, apices obtuse-pointed;acroscopic segments ca. 1.2 × 0.3 cm, pinnatilobate.Laminae thinly herbaceous, when dry brownish, exceptcostae villous adaxially, elsewhere glabrous. Sori small, 4 or 5pairs per pinnule; indusia small, orbicular-reniform, brown, glabrous,deciduous.● Shaded places among rocks in mountain thickets. Fujian, Zhejiang.10. Metathelypteris glandulifera Ching ex K. H. Shing, Fl.Reipubl. Popularis Sin. 4(1): 322. 1999.有腺凸轴蕨 you xian tu zhou juePlants 50–60 cm tall. Rhizomes long creeping, apices includingstipe bases with deep brown linear-lanceolate scalesand grayish white acicular hairs. Fronds approximate; stipes20–30 cm, stramineous, above bases with similar hairs butsparsely so and with few scales; laminae oblong-lanceolate, ca.30 × 14–16 cm, nearly tapering to bases, pinnate-pinnatifid,apices acuminate and pinnatifid; pinnae 10–14 pairs, spreading,subopposite, sessile, lanceolate, 7–8 × 1–1.5 cm, bases slightlytapering, rounded-truncate, pinnatifid nearly to costae, apiceslong acuminate; segments ca. 17 pairs, spreading, lanceolate,sometimes falcate, 1–1.2 × ca. 0.3 cm, bases decurrent, connectedto each other by narrow wings, margins crenate, apicesrounded-obtuse or acute. Veins not evident, lateral veins forked,8–10 pairs per segment, proximal pair arising above bases ofcostae and not reaching margins. Laminae herbaceous, whendry dark green, abaxially covered with grayish white multicellularacicular hairs along rachises, costae, and costules, lateralveins and intercostal areas covered with short unicellularacicular hairs, mixed with small orange-red globose glands,adaxially densely grayish white acicular hairy along rachisesand costae, hairs along veins sparse. Sori small, orbicular, 6 or 7pairs per segment, attached near ends of acroscopic vein offorked lateral veins; sporangia each with 4 or 5 globose smallglands along both sides of annuli; indusia orbicular-reniform,greenish, when dry grayish brown, membranous, subentire, noteasily seen when mature.● Dense forests. N Guangxi.The lamina outline of Metathelypteris glandulifera is extremelysimilar to that of M. flaccida, but the pinnae are covered with orangeredglobose glands abaxially; the sporangia also have a few similarglands along both sides of the annuli, and the pinnae are covered withunicellular short acicular hairs along veinlets and intercostal areas.11. Metathelypteris flaccida (Blume) Ching, Acta Phytotax.Sin. 8: 306. 1963.薄叶凸轴蕨 bao ye tu zhou jueAspidium flaccidum Blume, Enum. Pl. Javae 2: 161. 1828;Dryopteris flaccida (Blume) Kuntze; D. gracilescens (Blume)Kuntze var. decipiens (C. B. Clarke) Alderwerelt; Lastrea flaccida(Blume) T. Moore; L. gracilescens Hooker var. decipiens(C. B. Clarke) Beddome; Metathelypteris decipiens (C. B.Clarke) Ching; Nephrodium flaccidum (Blume) Hooker; N.gracilescens (Blume) Hooker var. decipiens C. B. Clarke;

THELYPTERIDACEAE 339Thelypteris decipiens (C. B. Clarke) Ching; T. flaccida (Blume)Ching.Plants 40–60 cm tall. Rhizomes thick and short, erect, withdeep brown linear-lanceolate scales. Fronds clustered; stipes15–30 cm, bases dark brown, with similar scales and grayishwhite short acicular hairs, distally stramineous and sparselyshortly hairy; laminae oblong-lanceolate, 25–40 × 12–16 cm,nearly tapering to bases, 2-pinnate-pinnatifid, apices acuminateand pinnatifid; pinnae 10–15 pairs, spreading, subopposite,sessile; proximal ones lanceolate, 7–9 × 2–2.5 cm, bases ±tapering, rounded-truncate, pinnate-pinnatifid, apices caudateacuminate;pinnules 10–15 pairs, spreading, lanceolate, 1–1.5 ×0.3–0.5 cm, bases decurrent to form narrow wing along costae,pinnatifid to 1/2–2/3 distance to costae, apices rounded-obtuse;segments 5–7 pairs, triangular-lanceolate, 1–2 × ca. 1 mm,entire, obtuse-pointed at apices. Veins not evident, lateral veinssimple or forked, 2 or 3 pairs per segment, proximal pair arisingfrom base of costules, not reaching margins. Laminae thinlyherbaceous, when dry yellowish green, with dense spreadingmulticellular acicular hairs on both surfaces. Sori small, orbicular,1 per segment, attached above middle of acroscopic veinof forked lateral veins; indusia small, orbicular-reniform,membranous, greenish, when dry grayish brown, glabrous oroccasionally with few acicular hairs along margins, persistent.2n = 140.Streamsides in forests; 700–1800 m. Guizhou, SW Yunnan [Bhutan,India, Indonesia, Malaysia, Nepal, Philippines, Sri Lanka, Thailand,N Vietnam].5. MACROTHELYPTERIS (H. Itô) Ching, Acta Phytotax. Sin. 8: 308. 1963.针毛蕨属 zhen mao jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiThelypteris sect. Macrothelypteris H. Itô in Nakai & Honda, Nov. Fl. Jap. 4: 141. 1939.Plants medium-sized, terrestrial, sometimes treelike, to 4 m tall. Rhizomes thick and short, erect, ascending or decumbent, withbrown lanceolate long scales; scales thick, with acicular cilia along margins. Fronds clustered; stipes stramineous or reddish brown,glabrous, or with similar scales and after fallen with remaining lunate marks; laminae large, ovate-triangular, 3- or 4-pinnate-pinnatifid;pinnae and pinnules oblique or spreading and connected to each other by narrow wings along costae or costules; veinspinnate, free, lateral veins simple, sometimes forked. Laminae herbaceous or somewhat papery, yellowish green when dry, costae andcostules rounded and raised adaxially, ± hairy on both sides and intercostal areas, rarely glabrous, hairs slender, grayish whiteacicular, unicellular or consisting of several cells, except above hairs along rachises, usually also with brown multicellular acicularthick hairs and few lanceolate or subulate thick scales and remaining protruding marks after scales fallen. Sori small, attached nearends of veinlets, exindusiate or with small and usually deciduous indusia; sporangia sometimes with shortly stalked capitate hairsnear annuli. Spores bilateral, elliptic-reniform; perispores transparent and corrugate, echinate or minutely foveolate; exospore finelyreticulate. x = 31.About ten species: warmer parts of mainland Asia, NE Australia, Malesia, Pacific islands; seven species (one endemic) in China.1a. Rachises and costae covered with scales abaxially; scales inflated or incrassate at bases, leaving roughmarks after falling.2a. Plants 1–1.4 m tall; costae and costules with scales incrassate and not inflated at bases ...................................... 3. M. setigera2b. Plants 3–4 m tall, costae and costules covered with inflated bases of scales abaxially.3a. Scales on rachises denser, scales on costae entire .............................................................................................. 1. M. ornata3b. Scales on rachises sparse, scales on costae with dense acicular hairs along margins ........................... 2. M. polypodioides1b. Rachises and costae without scales abaxially, or with grayish white hairs, or glabrous.4a. Pinnae glabrous or with unicellular acicular hairs.5a. Laminae herbaceous, when dry yellowish green; pinnules oblique distally and intersecting withcostae at sharp angle; pinnae glabrous abaxially, at most with sparse acicular hairs along costae ......... 4. M. oligophlebia5b. Laminae thinly herbaceous, grass-green or dark green when dry; pinnules spreading andintersecting with costae at right angle; pinnae with more spreading acicular hairs abaxially ..................... 5. M. viridifrons4b. Pinnae with spreading multicellular acicular hairs abaxially.6a. Laminae herbaceous, pinnules oblique distally and intersecting with costae at sharp angle;proximal pinnae with 2–2.5 cm stalks; hairs abaxially denser and elongate and 2.5–3 mm ....................... 6. M. torresiana6b. Laminae thinly herbaceous, pinnules spreading and intersecting with costae at right angle;pinnae subsessile; hairs abaxially sparse and short and 1.5–2 mm ............................................................. 7. M. contingens1. Macrothelypteris ornata (Wallich ex J. Smith) Ching, ActaPhytotax. Sin. 8: 309. 1963.树形针毛蕨 shu xing zhen mao juePhegopteris ornata Wallich ex J. Smith, Hist. Fil. 233.1875, based on Polypodium ornatum Wallich ex Beddome,Ferns S. India 56, t. 171. 1864, not Klotzsch (1847); Dryopterisornata (Wallich ex J. Smith) C. Christensen; Lastrea ornata(Wallich ex J. Smith) Copeland; Nephrodium ornatum (Wallichex J. Smith) Christ; Thelypteris ornata (Wallich ex J. Smith)Ching.Plants 3–4 m or more tall. Rhizomes erect, cylindric, with

340THELYPTERIDACEAEdense scales; scales brownish, ciliate along margins, long subulateand thick. Fronds clustered; stipes 60–100 cm, thick, ca.2.5 cm in diam., stramineous, polished, bases with similarscales as on rhizomes, distally scales sparser, after fallingleaving tuberculate or lunate rough marks; laminae large, triangular-ovate,long and wide, to above 2 m, not tapering tobases, 4-pinnatifid, acuminate and pinnatifid at apices; pinnae18–25 pairs, subopposite, spreading, proximal ones stalked,oblong-lanceolate, ca. 60 × 30 cm, 3-pinnatifid, acuminate atapices; pinnules of 1-pinnate laminae large, spreading, shortlystalked, lanceolate, 10–16 × 3–4 cm, bases truncate, symmetrical,pinnate-pinnatifid, apices acuminate. Pinnules of 2-pinnate laminae 18–25 pairs, subopposite, spreading, sessile,lanceolate, slightly falcate, 1.5–2 × ca. 0.4 cm, bases decurrent,connected to each other by narrow wing along costae, pinnatifidto 1/2 or more of distance to costules, apices pointed; segments10–12 pairs, crenatelike or triangular, oblique distally, entire.Veins pinnate, veinlets 2 or 3 pairs per segment, forked, notevident. Laminae herbaceous, when dry yellowish green, abaxialsides with multicellular acicular hairs along costules andmain veins, costae with sparse linear-lanceolate scales abaxially,scales inflated at bases, persistent, adaxial sides with shortacicular hairs along costae. Sori small, orbicular, 1 pair persegment, attached near top of basal acroscopic veinlets; indusianot developed. 2n = 62.Forests in river valleys of subtropical regions. SE Xizang, WYunnan [Bhutan, India, Nepal, Thailand].The nomenclatural history of this taxon is confusing. The epithetwas introduced as a nomen nudum by Wallich (Numer. List, no. 327.1829). Fée (Mém. Foug. 5: 243. 1852) made the combination “Phegopterisornata” based on Wallich’s name but provided no further information,so this name is another nomen nudum and was likewise notvalidly published (Melbourne Code, Art. 38.1(a)). Beddome (1864) providedthe first actual description in his account of Polypodium ornatumin 1864, but unfortunately this name is a later homonym and is thereforeillegitimate. The earliest legitimate name appears to be that of J. Smithin 1875, as his combination can be indirectly associated with Beddome’sdescription through his citation of Wallich.2. Macrothelypteris polypodioides (Hooker) Holttum, Blumea17: 29. 1969.桫椤针毛蕨 suo luo zhen mao jueAlsophila polypodioides Hooker in Nightingale, OceanicSketches, 131. 1835; Cheilanthes gigantea Cesati; Dryopterisbrunneovillosa C. Christensen; D. leucolepis (C. Presl) Maxon;Lastrea leucolepis C. Presl; Macrothelypteris leucolepis (C.Presl) Ching; Phegopteris cheilanthoides (Baker) Alderwerelt;Polypodium cheilanthoides Baker; Thelypteris leucolepis (C.Presl) Ching.Plants ca. 1.5 m tall. Rhizomes short, decumbent, thick,with ascending apex. Fronds tufted; stipes ca. 80 cm, pale, dull,basal half at least bearing copious slender hairs and scales;scales thin, pale, linear, bearing copious very slender unicellularhairs and sometimes a few of more than one cell, base of scalesthick and dark, of larger ones flat, of smaller ones terete,leaving warts when scales fall; main rachises pale, hairy asstipe, with many small warts abaxially; laminae ca. 80 cm;basal 2 or 3 pairs of pinnae to 35 × 15 cm, upper ones graduallyshortened; basal pinnules of lower pinnae somewhat shortened;pinna rachis hairy adaxially, with narrow pale scales abaxially;largest pinnules sessile, ca. 10 × 2.5 cm, acuminate at apices,almost at right angle to pinna rachis, segments all with broadbases connected by a very narrow wing along costae; costaedensely hairy adaxially (sparsely hairy on costules), abaxiallywith very narrow pale scales, short capitate hairs also present;segments almost at right angles to costae, larger ones deeplylobed. Veinlets forked or shortly pinnate, laminae adaxiallybearing capitate hairs. Sori small, on acroscopic branch of avein; indusium small, persistent, bearing many short capitatehairs; sporangia with 2 or 3 similar capitate hairs near annulus;spores not winged, surface apparently minutely and irregularlygranular.Forest margins; 700–2100 m. Taiwan [Papua New Guinea, Philippines,Thailand; Australia, Pacific islands].3. Macrothelypteris setigera (Blume) Ching, Acta Phytotax.Sin. 8: 309. 1963.刚鳞针毛蕨 gang lin zhen mao jueCheilanthes setigera Blume, Enum. Pl. Javae 2: 138.1828; Aspidium setigerum (Blume) Kuhn (1869), not Swartz(1829); C. stenophylla Kunze; Dryopteris backeri Alderwerelt;D. setigera (Blume) Kuntze; Hypolepis setigera (Blume)Hooker; Nephrodium setigerum (Blume) Baker (1867), not C.Presl (1825); Thelypteris setigera (Blume) Ching.Plants 1–1.4 m tall. Rhizomes thick and ascending, withdense brownish linear-lanceolate scales; scales ca. 1 cm, thick,with sparse hairs along margins. Fronds clustered; stipes 50–60cm or more, stramineous, bases with similar scales as rhizomes,distally sparse, scales inflated at bases and remaining as tuberculaterough marks after falling; laminae oblong-lanceolate,50–80 × 30–50 cm, tapering at bases, 3-pinnate-pinnatifid, acuminateand pinnatifid at apices; pinnae 14–18 pairs, subopposite,spreading, sessile or subsessile, proximal pair larger,broadly lanceolate, 15–30 × 5–15 cm, bases truncate, symmetrical,bipinnate, apices acuminate; pinnules of 1-pinnate laminae20–30 pairs or more, sessile, spreading, lanceolate, 2.5–7 × 0.5–1.5 cm, symmetrical at bases, pinnatisect, acuminate at apices;pinnules of bipinnate laminae 12–20 pairs, lanceolate, 2.5–7 ×1.5–2.5 mm, bases decurrent, connected to each other by narrowwings, entire or sharply lobate, margins usually recurved,obtuse at apices. Veinlets not evident, simple or forked, 3–5 perultimate pinnule. Laminae herbaceous, brownish when dry,abaxially including costae and costules with more glaucousmulticellular acicular hairs, adaxially with acicular hairs alongcostae, rachises with hard scales. Sori small, orbicular, attachedabove middle of veinlets, wrapped by recurved margins; indusiasmall, deciduous or hidden in mature sori.Tropical rain forests. Taiwan [Indonesia, Malaysia].4. Macrothelypteris oligophlebia (Baker) Ching, Acta Phytotax.Sin. 8: 308. 1936.针毛蕨 zhen mao juePlants 60–150 cm tall. Rhizomes short and ascending,including stipe bases with deep brown lanceolate marginally

THELYPTERIDACEAE 341sparsely hairy scales. Stipes 30–70 cm, stramineous, glabrousabove bases; laminae triangular-ovate, similar in length tostipes, 30–45 cm wide at proximal part, not tapering at bases, 3-pinnatifid, acuminate and pinnatifid at apices; pinnae ca. 14pairs, oblique distally, alternate, or proximal ones opposite,stalks to 2 cm or more, proximal pair larger, oblong-lanceolate,ca. 20 × 5 cm, slightly tapering to bases, acuminate and pinnatifidat apices; second pair and above pinnae gradually reduceddistally, not tapering to bases, with stalks 1–1.5 cm, bipinnatifid;pinnules 15–20 pairs, 3.5–8 × 1–2.5 cm, lanceolate,bases rounded-truncate, symmetrical, sessile (proximal onesshortly stalked), ± decurrent (distal ones connected to eachother by narrow wing), pinnatifid nearly to costules, apicesacuminate; segments 10–15 pairs, spreading, 5–12 × 2–3.5 mm,bases connected to each other by narrow wings, margins entireor sharply lobate, apices obtuse or bluntly pointed. Veins evidentabaxially, veinlets simple or forked on sharply lobate segments,4–8 pairs per segment. Laminae herbaceous, when drygrass-green, glabrous on both surfaces, with orange-yellowtransparent capitate glandular hairs abaxially, or along costulesand near ends of costae with few unicellular acicular hairs,adaxially with grayish white short acicular hairs along costaeand costules, usually with reddish purple spots on costae. Sorismall, orbicular, 3–6 pairs per segment, attached near ends ofveinlets; indusia small, orbicular-reniform, grayish green, glabrous,deciduous when mature or hidden in sori.Streamsides in valleys, wet soil at forest margins. S Anhui, Fujian,Guangdong, N Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, Jiangsu,Jiangxi, Taiwan, Zhejiang [Japan, S Korea].1a. Pinnae glabrous on both surfaces ...... 4a. var. oligophlebia1b. Pinnae along costae and costules allwith grayish white unicellularacicular short hairs abaxially ..................... 4b. var. elegans4a. Macrothelypteris oligophlebia var. oligophlebia针毛蕨 ( 原变种 ) zhen mao jue (yuan bian zhong)Nephrodium oligophlebium Baker, J. Bot. 13: 291. 1875;Aspidium oligophlebium (Baker) Christ; Dryopteris oligophlebia(Baker) C. Christensen; Hypolepis punctata (Thunberg)Mettenius ex Kuhn var. henryi Christ; Lastrea oligophlebia(Baker) Copeland; Macrothelypteris torresiana (Gaudichaud)Ching var. calvata (Baker) Holttum; N. setigerum C. Presl var.calvatum Baker; Thelypteris oligophlebia (Baker) Ching; T. torresiana(Gaudichaud) Alston var. calvata (Baker) K. Iwatsuki;T. uliginosa (Kunze) Ching var. calvata (Baker) K. Iwatsuki.Pinnae glabrous on both surfaces.Streamsides in valleys, wet soil at forest margins; 400–800 m. SAnhui, N Guangxi, S Henan, Hubei, Hunan, S Jiangsu, Jiangxi, Zhejiang[Japan].4b. Macrothelypteris oligophlebia var. elegans (Koidzumi)Ching, Acta Phytotax. Sin. 8: 309. 1963.雅致针毛蕨 ya zhi zhen mao jueDryopteris elegans Koidzumi, Bot. Mag. (Tokyo) 38: 108.1924; Lastrea oligophlebia var. elegans (Koidzumi) Tagawa; L.uliginosa Newman var. elegans (Koidzumi) K. Iwatsuki;Macrothelypteris changshaensis Ching; M. oligophlebia var.changshaensis (Ching) K. H. Shing; Thelypteris oligophlebiavar. elegans (Koidzumi) Ching; T. uliginosa var. elegans (Koidzumi)K. Iwatsuki.Pinnae along costae and costules with grayish white unicellularshort acicular hairs abaxially. 2n = 62, 124.Streamsides in valleys, forest margins, on slopes and plains at lowelevations. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Henan, Hubei,Hunan, Jiangsu, Jiangxi, Taiwan, Zhejiang [Japan, S Korea].5. Macrothelypteris viridifrons (Tagawa) Ching, Acta Phytotax.Sin. 8: 310. 1963.翠绿针毛蕨 cui lü zhen mao jueThelypteris viridifrons Tagawa, J. Jap. Bot. 12: 747. 1936;Dryopteris elegans Koidzumi var. subtripinnata Tagawa; D.oligophlebia (Baker) C. Christensen var. subtripinnata (Tagawa)H. Itô; Lastrea oligophlebia (Baker) Copeland var. subtripinnata(Tagawa) Ohwi; L. viridifrons (Tagawa) Tagawa; T.oligophlebia (Baker) Ching var. subtripinnata (Tagawa) H. Itô.Plants 60–110 cm tall. Rhizomes short and erect, with reddishbrown hairy lanceolate scales at apices. Fronds clustered;stipes 30–50 cm, stramineous, bases with grayish white shortacicular hairs, distally glabrous; laminae similar in length tostipes or slightly longer, 20–50 cm wide, not tapering to bases,3-pinnate-pinnatifid, acuminate and pinnatifid at apices; pinnae10–12 pairs, alternate or subopposite, oblique distally, stalks1.5–5 cm, proximal pair largest, oblong-lanceolate, 24–30 × ca.10 cm, bases slightly tapering, rounded-truncate, apices acuminate;second pair of pinnae (distally all pinnae) of similarshape as proximal pair, but not tapering to bases, graduallyreduced, shortly stalked, 2-pinnate-pinnatifid; pinnules of 1-pinnate parts of laminae 10–15 pairs, alternate, spreading,proximal pair slightly shortened, distal ones oblong-lanceolate,5–6.5 × 2.5–3 cm, bases truncate, shortly stalked, pinnate-pinnatifid,apices acuminate; pinnules of 2-pinnate parts of lamina10–15 pairs, lanceolate, 1–1.5 × 0.4–0.7 cm, bases roundedtruncateand decurrent, connected to each other on both sides bynarrow wings, pinnate lobate or pinnatifid to 2/3 of distance tocostules, obtuse or bluntly pointed at apices; segments elliptic,ca. 2.5 × 1.5 mm, entire or slightly undulate along margins,rounded at apices. Veins visible, veinlets simple, 2 or 3 pairs persegment. Laminae thickly herbaceous, when dry grass-green,with more spreading acicular hairs abaxially, adaxially withmore short acicular hairs along costules; rachises stramineous,glabrous abaxially, polished. Sori small, orbicular, 1 or 2 persegment, attached near ends of proximal veinlets; indusia small,orbicular-reniform, green, membranous, with 1 or 2 villous-hairsand deciduous after maturity. 2n = 124.Shaded wet places in forests in mountain valleys; ca. 800 m. Anhui,N Fujian, Guizhou (Nayong), Hunan, S Jiangsu, Jiangxi, Zhejiang[Japan, S Korea].6. Macrothelypteris torresiana (Gaudichaud) Ching, ActaPhytotax. Sin. 8: 310. 1963.普通针毛蕨 pu tong zhen mao juePolystichum torresianum Gaudichaud, Voy. Uranie, Bot. 8:

342THELYPTERIDACEAE333. 1828; Aspidium mollissimum Christ; A. uliginosum Kunze;Dryopteris lasiocarpa Hayata; D. mollissima (Christ) C. Christensen;D. oligophlebia (Baker) C. Christensen var. lasiocarpa(Hayata) Nakai; D. tenericaulis (Wallich ex Hooker) Ching;D. uliginosa (Kunze) C. Christensen; Lastrea oligophlebia(Baker) Copeland var. lasiocarpa (Hayata) H. Itô ex M. Mizushima;L. tenericaulis (Wallich ex Hooker) T. Moore; L. torresiana(Gaudichaud) T. Moore; Nephrodium tenericaule(Wallich ex Hooker) Hooker; Polypodium tenericaule Wallichex Hooker; P. trichodes J. Smith; Thelypteris oligophlebia(Baker) Ching var. lasiocarpa (Hayata) H. Itô; T. torresiana(Gaudichaud) Alston; T. uliginosa (Kunze) Ching.Plants 60–150 cm tall. Rhizomes short, erect or ascending,with dense reddish brown hairy linear-lanceolate scales atapices. Fronds clustered; stipes 30–70 cm, grayish green, whendry stramineous, bases shortly hairy, distally subglabrous; laminaetriangular-ovate, 30–80 × 20–50 cm, not tapering at bases,3-pinnate, acuminate and pinnatifid at apices; pinnae ca. 15pairs, subopposite, stalks 2–2.5 cm; proximal pairs largest,oblong-lanceolate, 10–30 × 4–12 cm, bases slightly tapering,acroscopic sides parallel with rachises, 2-pinnate, apices acuminate;pinnules of 1-pinnate laminae 15–20 pairs, alternate,distal ones ± adnate to costae and decurrent and connected toeach other, proximal several pairs shortly stalked, lanceolate, 3–10 × 0.8–2 cm, rounded-truncate at bases, pinnatifid, acuminateat apices; segments 10–15 pairs, approximate, lanceolate, 4–12× 2–3 mm, bases connected to each other by narrow wings,margins entire or often sharply lobate, apices obtuse or bluntlypointed; second pair of pinnae of similar shape as proximal pair,but not tapering at bases and gradually shortened. Veins notevident, veinlets simple or forked on sharply lobate segments,3–7 pairs per segment. Laminae herbaceous, when dry darkbrownish green, with more grayish white multicellularspreading long acicular hairs and capitate short glands abaxially,adaxially with short hairs along costae and costules, rachisesand costae stramineous, glabrous abaxially, adaxiallywith multicellular slender hairs. Sori small, 2–6 pairs per segment,attached near ends of veinlets; indusia small, orbicularreniform,greenish, hidden in mature sori. Sporangia each with2 or 3 capitate short hairs at top. 2n = 144, 186.Wet places in mountain valleys; sea level to 1000 m. Anhui,Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Henan,Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang[Bhutan, India, Indonesia, Japan, Myanmar, Nepal, Philippines,Vietnam; tropical and subtropical regions of America, Australia, Pacificislands].7. Macrothelypteris contingens Ching, Acta Phytotax. Sin. 8:310. 1963.细裂针毛蕨 xi lie zhen mao juePlants ca. 1 m tall. Rhizomes short and erect, with darkbrown lanceolate scales. Fronds clustered; stipes ca. 50 cm,stramineous, distally glabrous; laminae ovate-lanceolate, similarin length as stipes, 15–30 cm proximally, not tapering tobases, 3-pinnate-pinnatifid, acuminate and pinnatifid at apices;pinnae ca. 15 pairs, opposite, distal ones alternate, spreading,subsessile; proximal pair of pinnae broadly lanceolate, 14–18 ×8–9 cm, bases slightly tapering, truncate, pinnate-pinnatifid,apices acuminate; pinnules of 1-pinnate laminae 15–20 pairs,connected to each other, spreading, proximal ones opposite, distalones alternate, oblong-lanceolate, 3–4 × ca. 1 cm, basesconnected along both sides by narrow wings, partite, acuminateat apices; segments 12–15 pairs, approximate, or obliquelyspreading, elliptic, 3.5–4.2 × ca. 2.2 mm, bases decurrent andconnected to each other by narrow wings, margins sharplylobate into 3 or 4 crenations, apices rounded; second pair ofpinnae of similar size as proximal pair, but not tapering atbases. Veins visible, veinlets 2- or 3-forked, occasionally simple,3 or 4 pairs per segment. Laminae thinly herbaceous, greenwhen dry, with sparse whitish multicellular spreading acicularhairs abaxially, adaxially with similar short acicular hairs butsparse, costae stramineous, glabrous or subglabrous abaxially,hairy adaxially. Sori small, orbicular, 3 or 4 pairs per segment,attached near ends on acroscopic veinlets forked; indusia small,not very prominent.● Wet soil in forests in mountain valleys; 900–1100 m. S Yunnan,S Zhejiang.6. PHEGOPTERIS (C. Presl) Fée, Mém. Foug. 5: 242. 1852.卵果蕨属 luan guo jue shuLin Youxing ( 林尤兴 ); Alan R. SmithPolypodium [par.] Phegopteris C. Presl, Tent. Pterid. 179. 1836.Plants mid- and small-sized, terrestrial. Rhizome long creeping or short and erect, densely covered with brown scales andwhitish acicular hairs. Fronds remote or clustered; stipe stramineous, shiny, slender, base scaly; scales brown, lanceolate and sparselylong hairy along margins; lamina bipinnatifid or pinnate-pinnatifid, ovate-triangular or narrowly lanceolate; pinnae connected to eachother by a narrow wing along rachis, or proximal 1–3 pairs free, proximal pinnae not shortened or basal pair only slightly shortened,or proximal several pairs gradually reduced to auricles; veins pinnate, lateral veins simple or forked, veinlets reaching margins;lamina herbaceous or soft papery, with whitish acicular hairs on both surfaces, rachis, costae, and costules rounded and raised onboth sides and similarly with dense acicular hairs, sometimes mixed with a few forked hairs, with more brownish hairs andlanceolate, ciliate scales abaxially. Sori orbicular to oblong, borne above middle of ultimate veins, exindusiate or indusia very smalland vestigial; sporangia often with a few short acicular hairs or capitate hairs near annulus. Spores bilateral, reniform, perisporeswinged, thin, and transparent, granular on surfaces. x = 30.Four species: widely distributed throughout the N temperate zone, one species in SE Asia; three species (one endemic) in China.Phegopteris is monophyletic and differs from Pseudophegopteris in its smaller stature, stramineous stipes, laminae usually triangular or

THELYPTERIDACEAE 343narrowly lanceolate, proximal pinna bases decurrent and adnate to each other by a rachis wing, veinlets reaching margin, sparsely ciliate scales onabaxial side of rachises, and costae sparsely ciliate. Phegopteris is primarily N temperate and circumboreal, while Pseudophegopteris is tropical andsubtropical and restricted to the Paleotropics.The following taxa are excluded from the present treatment, pending further research: Phegopteris amaurophylla Christ (Bull. Herb. Boissier7(1): 14. 1899), described from Yunnan, and P. somae (Hayata) Tagawa (Acta Phytotax. Geobot. 7: 75. 1938; Dryopteris somae Hayata, Icon. Pl.Formosan. 5: 287. 1915 [“somai”]), described from Taiwan.1a. Rhizome short, erect; lamina lanceolate, pinnae connected to each other by narrow wings; most pinnaebelow middle of blade gradually shortened, proximal pair of pinnae reduced to auricles ............................ 3. P. decursive-pinnata1b. Rhizome long creeping; lamina ± triangular, proximal 1–3 pairs of pinnae free, not tapering proximally,or basal pair slightly shortened, distal pinnae connected to each other by narrow rachis wing.2a. Lamina triangular, length and width subequal or length slightly longer than width, proximal pair ofpinnae largest and often reflexed downward ........................................................................................................ 1. P. connectilis2b. Lamina narrowly triangular, length almost 2 times width, proximal 1–3 pairs of pinnae free, proximalpair slightly shortened, spreading or slightly ascending ........................................................................................... 2. P. tibetica1. Phegopteris connectilis (Michaux) Watt, Canad. NaturalistGeol., n.s., 3: 159. 1866 [“connectile”].卵果蕨 luan guo juePolypodium connectile Michaux, Fl. Bor.-Amer. 2: 271.1803; Aspidium phegopteris (Linnaeus) Baumgarten; Dryopterisphegopteris (Linnaeus) C. Christensen; Gymnocarpiumphegopteris (Linnaeus) Newman; Nephrodium phegopteris(Linnaeus) Prantl; Phegopteris polypodioides Fée; P. vulgarisMettenius; Polypodium phegopteris Linnaeus; Polystichumphegopteris (Linnaeus) Roth; Thelypteris phegopteris (Linnaeus)Slosson.Plants 25–40 cm tall. Rhizome long creeping, with brightbrown, ovate-lanceolate thin scales at apex. Fronds remote;stipe dark brown at base, stramineous distally, 15–30 cm,sparsely scaly, nearly smooth; lamina bipinnatifid, deltoid, 13–20 × 10–18 cm, acuminate and pinnatifid at apex; pinnae ca. 10pairs, usually opposite, spreading, lanceolate, 5–9 × 1–2 cm,basal pair largest, bases slightly or not tapering, free fromsecond pair of pinnae, slightly deflexed, apices acuminate;segments oblong, entire, undulate, or lobed along margins,rounded or obtuse at apices; distal pinnae gradually shortened,bases connected along rachises by triangular wings. Veins pinnate,lateral veins simple or occasionally forked. Fronds herbaceousor papery, drying gray-green or yellowish green, withsparse gray-white acicular hairs on both surfaces, ± with smallscales along rachises and costae; scales brownish, ovate-lanceolateand ciliate along margins. Sori ovate-orbicular or orbicular,borne at or near ends of ultimate veins and close to margins.Sporangia with 1 or 2 hairs near annulus. 2n = 60, 90.Forests, shrublands; 1200–3600 m. Guizhou, Heilongjiang, Henan,Jilin, Liaoning, Shaanxi, Sichuan, Taiwan, Yunnan [widely distributedin temperate regions of the N Hemisphere, south to mountainsof C Asia and the Himalaya].Phegopteris connectilis is similar to P. hexagonoptera (Michaux)Fée, which occurs in North America only. In P. connectilis, the proximalpinnae taper to their bases, the proximal and subbasal pinnae are notconnected by decurrent wings along the rachis, the laminae are thinlyherbaceous and sparsely shortly hairy on both surfaces, and the lateralveins are mostly simple. Phegopteris hexagonoptera has the proximalpair of pinnae connected to the next pair by a wing along the rachis,shorter laminar hairs (less than 0.25 mm vs. mostly 0.3–0.5 mm orlonger in P. connectilis), and ultimate veins forked or pinnate.2. Phegopteris tibetica Ching, Fl. Xizang. 1: 161. 1983.西藏卵果蕨 xi zang luan guo juePlants ca. 45 cm tall. Rhizome not seen. Stipe dark brownat base, distally stramineous, 24–27 cm, with triangular-lanceolatescales, rounded abaxially, nearly smooth or with short hairs,grooved and with acicular hairs adaxially; lamina pinnate-pinnatifid,oblong, 18–20 × 10–14 cm, base not decurrent, apexacuminate and pinnatifid; pinnae 13–15 pairs, proximal 2 or 3pairs free, spreading or slightly obliquely spreading, 5–7 × ca.2 cm (basal one pair slightly shortened), oblong-lanceolate,slightly tapering to bases, ± adnate to rachis, pinnatifid; segmentsoblong, entire along margins (or occasionally undulatecrenateon long segments), sparsely ciliate, rounded-obtuse atapices; pinnae above middle gradually shortened, linear-lanceolate,bases decurrent and connected to each other along rachisby narrow wings. Veins pinnate, lateral veins forked or simpleand reaching margins. Fronds herbaceous, drying deep green,rachises raised abaxially, adaxially with grooves with denseacicular hairs, rachises and costae with acicular hairs on bothsides and mixed with a few forked hairs, with sparse brownishnarrowly lanceolate scales abaxially, scales sparsely long ciliatealong margins, subglabrous between veins on both surfaces.Sori orbicular or nearly so, borne subterminally on ultimateveins and close to margins, exindusiate. Sporangia each occasionallywith 1 or 2 hairs near annulus.● Abies forests; ca. 3600 m. Xizang (Bomi).Phegopteris tibetica is most similar to P. connectilis, but the laminarlength is 2 times the width, proximal 2 or 3 pairs of pinnae usuallynot connected at bases by a rachis wing, basal pair of pinnae slightlyshortened, and laminae glabrous between veins on both surfaces.3. Phegopteris decursive-pinnata (H. C. Hall) Fée, Mém.Foug. 5: 242. 1852.延羽卵果蕨 yan yu luan guo juePolypodium decursive-pinnatum H. C. Hall, Nieuwe Verh.Eerste Kl. Kon. Ned. Inst. Wetensch. Amsterdam 5: 204. 1836;Aspidium decursive-pinnatum (H. C. Hall) Kunze; Dryopterisdecursive-pinnata (H. C. Hall) Kuntze; Lastrea decurrens J.Smith; L. decursive-pinnata (H. C. Hall) J. Smith; Leptogrammadecursive-pinnata (H. C. Hall) J. Smith; Nephrodiumdecursive-pinnatum (H. C. Hall) Baker; Thelypteris decursivepinnata(H. C. Hall) Ching.

344THELYPTERIDACEAEPlants 30–60 cm tall. Rhizome short and erect, includingbase of stipe with reddish brown, ciliate, narrowly lanceolatescales. Fronds clustered; stipe stramineous, 10–25 cm; laminalanceolate, 20–50 × 5–12 cm, gradually tapering to base, bipinnatifidor pinnate and toothed along margins, acuminate andpinnatifid at apex; pinnae 20–30 pairs, alternate, obliquelyspreading, middle ones largest, narrowly lanceolate, 2.5–6 × ca.1 cm, broad and decurrent at bases, connected by orbicular auriclesor triangular wings between pinnae, pinnatifid to 1/3–1/2 ofdistance to costules, acuminate at apices; segments obliquelyspreading, ovate-triangular, entire, obtuse at apices; pinnaegradually shortened to both ends, basal one pair of pinnae oftenshortened into small auricles. Veins pinnate, lateral veins simpleand reaching margins. Fronds herbaceous, along rachis, costae,and veins with whitish unicellular acicular short hairs on bothsides, abaxial side with forked and stellate hairs, abaxial side ofrachises and costae with sparse brownish, hairlike or lanceolate,ciliate scales. Sori suborbicular, borne at or near ends of ultimateveins, 2 or 3 per segment, sometimes with tufts of stalkedforked hairs in center when young. Sporangia each sometimeswith 1 or 2 short hairs near annulus. 2n = 60, 90, 120.Along rivers on fluvial plains, lower montane regions of hills,forests by roadsides; sea level to 2000 m. Anhui, Chongqing, Fujian,Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu,Jiangxi, S Shaanxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, S Korea,N Vietnam].Phegopteris decursive-pinnata is very different from other speciesof the genus in having short, erect rhizomes and narrowly lanceolatelaminae, but it can spread by long-creeping rhizomes. Because of thesecharacters, Iwatsuki removed it from Thelypteris sect. Phegopteris andtreated it and some species that we place in Pseudophegopteris, as wellas other species, in T. sect. Lastrea (Bory) Alston. Three ploidal levelsare known, but these numbers do not clearly correlate with differingmorphologies.7. CRASPEDOSORUS Ching & W. M. Chu, Acta Phytotax. Sin. 16(4): 24. 1978.边果蕨属 bian guo jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants medium-sized, terrestrial. Rhizomes erect, including stipe bases with scales and short rough hairs; scales reddish brown,ovate-lanceolate, ciliate and densely shortly hairy. Fronds clustered; stipes greenish, with dense divided grayish white transparentacicular fine long hairs; laminae broadly lanceolate, not tapering at bases, pinnate-pinnatifid, acuminate and pinnatifid at apices;pinnae large, but proximal pair of pinnae not shortened, lanceolate, pinnatifid; segments with semitransparent membranous narrowedge and undulate along margins, when dry usually reflexed and not easily seen; costae grooved adaxially, abaxially rounded; veinsfree, veinlets on segments simple (occasionally forked at middle), slightly inflated at ends and not reaching margins; vein typesimilar to Pseudocyclosorus, i.e., acroscopic veinlet of basal pair of veins reaching excartilaginous sinuses, basiscopic veinletreaching margins above sinus. Laminae herbaceous, along rachises and costae setaceous and mixed with short rough hairs. Sorioblong, attached below ends of veinlets and close to margins, exindusiate. Sporangia glabrous or usually with only one short acicularhair near annulus. Spores echinate.● One species: China.Craspedosorus is clearly closely related to Leptogramma. They both are very similar in laminar outline, segment shape, venation, and sori, butCraspedosorus is larger, with more pinnae that are free from rachises (except for distal ones) and pinnatifid nearly to costae; veinlets not reachingmargins, segments slightly undulate along margins and with translucent membranous narrow edges (usually reflexed when dry and not easily seen);sori short and thick, oblong, attached near ends of veinlets and closer to margins; and sporangia glabrous or each with one short acicular hair. In theopinion of Thelypteridaceae co-author Alan R. Smith, these diagnostic characters are not enough to separate Craspedosorus from Leptogramma.1. Craspedosorus sinensis Ching & W. M. Chu, Acta Phytotax.Sin. 16(4): 26. 1978.边果蕨 bian guo jueStegnogramma sinensis (Ching & W. M. Chu) L. J. He &X. C. Zhang.Plants to 1.1 m tall. Stipes ca. 40 cm, glabrous abovebases; laminae broadly lanceolate, ca. 65 × 22 cm, not taperingat bases, pinnate-pinnatifid; pinnae ca. 25 pairs, sessile,alternate, middle ones lanceolate, ca. 14 × 2.5 cm, bases truncateon acroscopic sides and parallel with costae, basiscopicsides obliquely truncate, pinnatifid to above 2/3 of distance tocostule, apices acuminate; segments ca. 20 pairs, middle onesoblong-lanceolate, ca. 1 cm (proximal 3 or 4 pairs slightlyshorter), ca. 5 mm wide, with translucent membranous narrowedge along margins and slightly undulate, usually reflexedwhen dry, obtuse or bluntly pointed at apices. Veinlets in segments5–9 pairs, simple or occasionally forked at middle,slightly thickened at end and not reaching margins. Laminaeherbaceous, green when dry, abaxially glabrous, adaxially withlong setae and mixed short rough hairs on rachises and costae.Sori oblong, attached slightly below end of veinlets and close tomargins, costules with broad sterile space on both sides; sporangiaeach usually with one short acicular hair near annulus.● Shaded thickets; 1400–1500 m. NE Yunnan.8. PSEUDOPHEGOPTERIS Ching, Acta Phytotax. Sin. 8: 313. 1963.紫柄蕨属 zi bing jue shuLin Youxing ( 林尤兴 ); Alan R. SmithPlants medium-sized, terrestrial. Rhizome short and erect, long creeping to ascending, apex with brown lanceolate scales.

THELYPTERIDACEAE 345Fronds clustered, sparse or remote; stipe castaneous or red-brown, less often stramineous or tan, shiny, base sometimes with whitishacicular hairs, sometimes mixed with stellate hairs, distally often glabrescent and smooth; lamina pinnate-pinnatifid to bipinnate-pinnatifid,lanceolate, oblong-lanceolate, oblong, or ovate, tapering or not to base, acuminate and pinnatifid at apex; pinnae usuallyopposite or subopposite, spreading or obliquely spreading, pinnae below middle not adnate to rachis, sessile or shortly stalked, lanceolateor triangular-lanceolate, bases often broadened, truncate, symmetrical or not, sometimes hastate, apices acuminate; basiscopicsegments sometimes longer than acroscopic segments, and basal segments on both sides sometimes prolonged; costae raised on bothsides, usually same color as stipes and rachises or lighter colored, abaxially glabrous or with whitish acicular hairs, hairy adaxially;veins free, lateral veins simple or forked, each with a clavate hydathode at end and not reaching margin. Sori oblong, ovate, orsuborbicular, borne at middle or above middle, exindusiate. Sporangia glabrous or with short hairs just below annulus. Sporesbilateral, orbicular-reniform, perispores thin and transparent, reticulate or foveolate on surfaces, exospore smooth. x = 31.About 25 species: tropical and subtropical Asia, east to the Pacific islands, to the west reaching to W Africa; 12 species (four endemic) in China.Citations of Pseudophegopteris paludosa (Blume) Ching from China (Ching, Acta Phytotax. Sin. 8: 315. 1963; Tsai & Shieh, Fl. Taiwan, ed. 2,1: 401. 1994), a species known with certainty only from Malesia (Holttum, Blumea 17: 91. 1969; Fl. Males., Ser. 2, 1: 347. 1981), perhaps apply tospecimens of P. pyrrhorhachis or P. hirtirachis, which are two closely related and similar species.1a. Stipes stramineous, bases including abaxial sides of rachises and costae with whitish acicular hairs mixedwith irregularly forked or stellate hairs.2a. Stipes same size or slightly shorter than laminae, 15–30 cm; pinnae with sparse acicular hairs adaxially ............. 1. P. levingei2b. Stipes equal to 1/4 of laminae in length, 8–10 cm; pinnae except costae with sparse acicular hairsadaxially, otherwise glabrous .................................................................................................................................... 2. P. brevipes1b. Stipes red-castaneous or castaneous, or if stramineous or brown-stramineous then bases of stipe, abaxialsides of rachis, and costae never with mixed acicular hairs.3a. Basal pair of pinnules or segments on proximal pinnae, particularly basiscopic one, clearly longer thanadjacent one and also lobed or incised; pinna base asymmetrically hastate.4a. Laminae not tapering to base, i.e., proximal 1 or 2 pairs of pinnae largest; pinnae all sessile ................... 3. P. yigongensis4b. Laminae ± tapering to base, i.e., proximal 1 or 2 pairs of pinnae smaller than distal ones.5a. Pinnae below middle shortly stalked .......................................................................................................... 4. P. tibetana5b. Pinnae all sessile.6a. Rhizome long creeping; stipes nearly same length as or slightly shorter than laminae; laminaepapery, drying brownish green, glabrous on both surfaces ...................................................................... 5. P. aurita6b. Rhizome shortly creeping to ascending; stipes much shorter than laminae; laminae thinlyherbaceous, drying yellowish green, with fine acicular hairs on both surfaces ................................ 6. P. subaurita3b. Basal pair of pinnules or segments on proximal pinnae same shape and size as distal pinnules/segmentson same pinna, or at most slightly inflated; pinnae symmetrically hastate at base, or seemingly not hastate.7a. Stipes stramineous, occasionally tan; plants 90–120 cm tall; laminae 60–80 × 20–30 cm; segmentstoothed along margins ................................................................................................................................ 12. P. microstegia7b. Stipes red-castaneous, castaneous-brown, or reddish brown, never stramineous or brown-stramineous;segments entire along margins.8a. Laminae less than 20 cm wide, pinnate-pinnatifid.9a. Sori near end of acroscopic veinlet of a forked pair and closer to margin; sporangia lackinghairs; pinnae subglabrous on abaxial surfaces between veins ......................................................... 10. P. zayuensis9b. Sori at middle of ultimate veins, between costae and segment margins; sporangia with 1 or2 seta(e) below annulus; pinnae with sparse fine acicular hairs on abaxial surfaces betweenveins ............................................................................................................................................ 11. P. rectangularis8b. Laminae wider than 20 cm, pinnate-pinnatisect with pinnules often toothed or lobed, or2-pinnate-pinnatifid.10a. Laminae broadly ovate, proximal pair of pinnae largest, 30–45 × 13–20 cm, pinnulespinnatifid nearly to costules ....................................................................................................... 9. P. yunkweiensis10b. Laminae oblong or oblong-lanceolate, proximal pair of pinnae same size as ones aboveor slightly shorter, 10–20 × 2.5–7 cm, pinnules pinnatifid to 1/2 of distance to costule.11a. Rhizome long creeping; laminae with sparse acicular short hairs or sometimesglabrous on abaxial surfaces; sporangia glabrous .......................................................... 7. P. pyrrhorhachis11b. Rhizome short, ascending; laminae with dense acicular short hairs on abaxialsurfaces; sporangia each with 2 or more hairs ..................................................................... 8. P. hirtirachis1. Pseudophegopteris levingei (C. B. Clarke) Ching, Acta Phytotax.Sin. 8: 314. 1963.星毛紫柄蕨 xing mao zi bing jueGymnogramma aurita Hooker var. levingei C. B. Clarke,Trans. Linn. Soc. London, Bot. 1: 568. 1880 [“levingii”]; Dryopterislevingei (C. B. Clarke) C. Christensen; D. purdomii C.Christensen; G. levingei (C. B. Clarke) Baker; Lastrea levingei

346THELYPTERIDACEAE(C. B. Clarke) Copeland; Leptogramma aurita (Hooker) Beddomevar. levingei (C. B. Clarke) Beddome; Phegopteris levingei(C. B. Clarke) Tagawa; Thelypteris levingei (C. B. Clarke)Ching.Plants 60–80 cm tall. Rhizome long creeping, with reddishbrown, broadly lanceolate scales and whitish acicular hairs.Fronds remote; stipe stramineous, 15–30 cm, with sparse similarscales on proximal parts and denser whitish acicular hairsand few irregularly forked stellate hairs; lamina pinnate-pinnatisect,lanceolate or oblong-lanceolate, 35–60 × 5–15 cm,slightly tapering at base, acuminate and pinnatifid at apex;pinnae to ca. 20 pairs, opposite, sessile, pinnae above middleapproximate, proximal 2 or 3 pairs remote from each other,gradually shortened downward, basal pair of pinnae smallest,2–5 cm, others 3–8(–11) × 1.5–2.5 cm, lanceolate, bases truncate,± symmetrical, pinnatisect nearly to costae, apices acuminate;segments 8–15 pairs per pinna, opposite, obliquelyspreading, approximate, oblong, 1–1.3 × 0.3–0.5 cm, slightlybroader at base, connected to each other by a narrow wing,entire, crenate, or occasionally lobed along margins of largersegments, obtuse at apices. Veins visible on both sides, lateralveins simple or forked, 5–7 per segment, basal pair arising fromabove bases of main veins. Laminae herbaceous, drying darkgreen, abaxial surfaces with dense whitish acicular hairs and afew stellate short hairs along costae and veins, also ± hairybetween veins, adaxial surfaces with sparse acicular hairs, hairsdenser along rachises. Sori suborbicular or oblong, 3–5(–7)pairs per segment, borne at or above middle of ultimate veinsand closer to margin. Sporangia each with 1–3 hairs ca. 0.2 mmbelow annulus. 2n = 124.Forests beside streams, thickets; 1300–3100 m. SE Gansu, SCShaanxi, C Sichuan, Taiwan, E and S Xizang, NW Yunnan [Afghanistan,Bhutan, N India, Kashmir, Pakistan].Pseudophegopteris levingei is similar to P. bukoensis (Tagawa)Holttum, but the latter is larger in outline, with laminae 2-pinnate-pinnatifid,pinnae and pinnules oblong, pinnules incised to 2/3 of distanceto costules, and costae with more stellate hairs abaxially.Reviewer Ralf Knapp provided the record for Taiwan. Informationabout this taxon and a taxon previously cited for Taiwan, Pseudophegopterisaurita, will be included in Knapp’s Supplement to the Ferns andFern Allies of Taiwan, to be published in 2013.2. Pseudophegopteris brevipes Ching & S. K. Wu, Fl. Xizang.1: 163. 1983.短柄紫柄蕨 duan bing zi bing juePlants 50–60 cm tall. Rhizome long creeping, includingbase of stipe with sparse whitish long hairs and reddish brownscales; scales ovate-lanceolate, thinly membranous, apices hairlike,hairy on back and margins. Fronds remote; stipe stramineous,8–20 cm, with sparse whitish acicular and a few stellatehairs, hairs denser in grooves adaxially; lamina 40–45 × 12–16cm, gradually tapering to base, pinnate-pinnatifid, pinnatifid andacuminate at apex; pinnae ca. 20 pairs, opposite or subopposite,spreading, sessile, proximal 2 pairs of pinnae slightly shortened,basal pair shortest, ovate-lanceolate, 3.5–5 cm, above ones lanceolate,6–8 × ca. 2 cm, bases widest and ± symmetrical, truncate,pinnatisect nearly to costae, apices acuminate and entire;segments to ca. 12 pairs, approximate, spreading, oblong, 8–9 ×ca. 4.5 mm, hairy along margins, obtuse at apices. Veinsslender, visible adaxially, lateral veins forked, 4 or 5 pairs persegment, basal pair arising from above bases of costae. Laminaethinly herbaceous, drying deep green or dark brown-green,with sparse whitish hairs, hairs denser along costae and veinsand mixed with a few irregularly forked stellate hairs, adaxialsurfaces with sparse acicular hairs along costae, rachises stramineous,sparsely hairy. Sori oblong, 2–4 pairs per segment,borne near ends of forked veinlets and close to margins. Sporangiaeach with hairs below annulus.● In rock crevices in valleys; ca. 2300 m. SE Xizang (Zayü).3. Pseudophegopteris yigongensis Ching, Fl. Xizang. 1: 165.1983.易贡紫柄蕨 yi gong zi bing juePlants 60–75 cm tall. Rhizome long creeping, with lanceolatehairy scales. Fronds remote; stipe blackish brown at base,brown-stramineous and shiny distally, 30–40 cm, bases villous,distally glabrous; lamina narrowly oblong or narrowly lanceolate,25–35 × 6–10 cm, base not tapering, pinnate-pinnatifid(basal pinnules on proximal pair of pinnae often pinnate),apex pinnatifid and acuminate; pinnae 9–12 pairs, opposite,spreading, distal pinnae decurrent and connected, middle pinnae± decurrent and adnate to rachis but not connected to eachother, proximal 1 or 2 pairs of pinnae largest, sessile and notdecurrent, 7–9 × ca. 4.5 cm, bases asymmetrical, pinnatifid,apices long acuminate; basiscopic pinna segments longer thanacroscopic ones, particularly basal basiscopic one, to 3–4 cm,pinnatifid, other segments entire. Veins visible on both sides,lateral veins simple or forked, 4–6 pairs per segment. Laminaethinly herbaceous, drying yellowish green, abaxially villousalong veins, subglabrous adaxially, rachises stramineous andsparsely villous. Sori oblong, borne at middle or above middleof ultimate veinlets, 2–4 pairs per segment. Sporangia each with2–4 acicular hairs below annulus.● Dense forests; ca. 2500 m. SE Xizang (Bomi, Yigong).Pseudophegopteris yigongensis is somewhat similar to P. levingei,differing in the fewer lateral pinnae, proximal two pairs much largerthan distal ones, and laminae sparingly villous abaxially.4. Pseudophegopteris tibetana Ching & S. K. Wu, Fl. Xizang.1: 164. 1983.西藏紫柄蕨 xi zang zi bing juePlants 85–90 cm tall. Rhizome long creeping, with sparsedark brown lanceolate scales. Fronds remote; stipe red-castaneousat base, shiny, 15–40 cm, base with a few scales andacicular hairs; lamina lanceolate, (15–)45–50 × (4–)12–16 cm,slightly tapering proximally, pinnate-pinnatifid (pinnules onproximal pair of pinnae often lobed), pinnatifid and long acuminateat apex; pinnae 4–17 pairs, ± opposite or subopposite,pinnae above middle ± adnate to rachis, pinnae below middleshortly stalked, proximal 1 or 2 pairs of pinnae obliquely triangular,(2–)7–7.5 × 1.5–3 cm, bases asymmetrical, pinnatifid,apices acuminate; basiscopic segments largest, oblong-lanceolate,ca. 2 × 0.4–1 cm, pinnatifid, other segments entire; dis-

THELYPTERIDACEAE 347tal pinnae lanceolate, to 8 × 2–2.2 cm, symmetrical, pinnatifidto 1/2 distance to costae or less, acuminate at apices; segmentsoblong, entire. Veins visible abaxially, lateral veins forked orsimple, 2–6 pairs per segment, basal pair arising from bases ofcostae. Fronds papery, drying yellowish green, abaxial surfaceswith sparse hairs along veins and margins, rachises stramineousto brown, sparsely villous, hairs denser adaxially. Sori oblong,borne above middle of ultimate veinlets, 2–4 pairs per segment.Sporangia each with a seta below annulus.● Dense forests; ca. 2100 m. SE Xizang.The distinctness of Pseudophegopteris tibetana relative to P. auritaand P. yigongensis needs more study.5. Pseudophegopteris aurita (Hooker) Ching, Acta Phytotax.Sin. 8: 314. 1963.耳状紫柄蕨 er zhuang zi bing jueGymnogramma aurita Hooker, Icon. Pl. 10: t. 974. 1854;Aspidium auritum (Hooker) Christ; Dryopteris aurita (Hooker)C. Christensen; Grammitis aurita (Hooker) T. Moore; Leptogrammaaurita (Hooker) Beddome; Nephrodium auritum(Hooker) Handel-Mazzetti; Phegopteris aurita (Hooker) J.Smith; Polypodium auritum (Hooker) E. J. Lowe; Thelypterisaurita (Hooker) Ching.Plants 40–100 cm tall. Rhizome long creeping, with scalesat apex; scales brown, narrowly lanceolate, 3–4 mm, ciliatealong margins. Fronds remote; stipe red-castaneous or brownishat least toward base, shiny, 20–60 cm; lamina ovate-lanceolate,20–70 × 15–30 cm, slightly tapering to base, pinnate-pinnatifid,pinnatifid and acuminate at apex; pinnae 10–18 pairs, opposite,spreading, sessile, proximal 1 or 2 pairs of pinnae slightlyshortened, lanceolate, distal pinnae 7–15 × 2–4 cm, bases hastate,asymmetrical, pinnatifid nearly to costae, apices acuminate,pinnae above middle ± symmetrical at base and adnateto rachis; segments (10–)15–20 pairs, spreading, basiscopic segmentsof pinnae longer than acroscopic ones, basal pair largest,particularly basiscopic one more oblique, lanceolate, 2.5–4 ×0.7–1 cm, margins pinnately lobed or crenate, apices acuminate,acroscopic ones shorter, parallel to rachis, oblong, 1–2 cm,entire or shallowly undulate, obtuse at apices. Veins visibleabaxially, lateral veins forked or simple, 5–7 pairs per segment,basal pair arising from bases of costules. Laminae thicklyherbaceous, drying brown-green, with short hairs along bothsides of costae or only adaxially, laminae otherwise glabrous,rachises smooth abaxially, with dense short hairs adaxially. Sorioblong or sometimes ovate-orbicular, borne above middle ofultimate veinlets, far from costules, 2–5 pairs per segment.Sporangia each with hairs below annulus. 2n = 62, 124.Forests beside streams on high mountains; 1200–2000 m. Chongqing,C Fujian, C Guizhou, S and W Jiangxi, SE Xizang, W Yunnan[Bhutan, NE India, Indonesia, Japan, Malaysia, N Myanmar, Nepal,Papua New Guinea, Philippines, N Vietnam].A widespread species, Pseudophegopteris aurita varies considerablyin frond size, and hence in laminar dissection, over its range.Specimens outside of China tend to be considerably larger.6. Pseudophegopteris subaurita (Tagawa) Ching, Acta Phytotax.Sin. 8: 315. 1963.光轴紫柄蕨 guang zhou zi bing jueDryopteris subaurita Tagawa, Acta Phytotax. Geobot. 1:157. 1932; Phegopteris subaurita (Tagawa) Tagawa; Thelypterissubaurita (Tagawa) Ching.Plants 50–120 cm tall. Rhizome shortly creeping to ascending,apex and stipe base with dense brown linear-lanceolatehairy scales. Fronds clustered or slightly spaced; stipe red-castaneousat least at base, 10–35 cm, abaxially sparsely hairy, adaxiallywith dense hairs along groove; lamina oblong-lanceolate,40–55(–100) × 15–30 cm, tapering at base, pinnate-pinnatifid,pinnatifid and long acuminate at apex; pinnae 15–20 pairs,opposite, obliquely spreading, sessile, basal pair of pinnaesmallest, narrowly triangular, 2–6 × 3–6 cm, more distal pinnaegradually longer, second pair 8–15 × ca. 2 cm (excluding basalmostsegments), triangular-lanceolate, bases abruptly widenedto 7 cm, asymmetrically hastate, pinnatifid nearly to costules,apices acuminate; segments 15–25 pairs per pinna, opposite,obliquely spreading, basal pair clearly longer than distal pairs,particularly basiscopic one pinnatifid and more oblique, lanceolate,to ca. 4 × 1.2 cm, tapering to an acute apex, acroscopic oneparallel to rachis, to 4 cm, more distal segments to 3 × ca. 0.8cm, oblong, margins crenate to lobed, distal segments entire,obtuse or subacute at apices. Veins visible abaxially, lateralveins simple or forked on proximal larger segments, basal pairarising from bases of costae. Fronds herbaceous, drying yellowishgreen, with fine acicular hairs on both surfaces, hairsalong costae and veins denser abaxially. Sori suborbicular tooblong, borne at middle or above middle (closer to margin) ofultimate veins. Sporangia each often with 1 or 2 short hairs belowannulus. 2n = 62.Forests beside streams, open areas in thickets; 200–1000 m. N andS Taiwan [Japan (Ryukyu Islands)].Pseudophegopteris subaurita is closely related to P. pyrrhorhachisand to P. aurita but is seemingly absent from mainland China.7. Pseudophegopteris pyrrhorhachis (Kunze) Ching, ActaPhytotax. Sin. 8: 313. 1963 [“pyrrhorachis”].紫柄蕨 zi bing juePlants 80–100 cm tall. Rhizome long creeping, scaly atapex. Fronds approximate or sparse; stipe red-castaneous, shiny,20–40 cm, base with short hairs and few lanceolate scales,distally glabrous; lamina oblong-lanceolate, 60–70 × 20–35cm, somewhat tapered to base, pinnate-pinnatifid, acuminateat apex; pinnae 15–20 pairs, opposite, sessile, proximal pinnaenarrowly lanceolate, middle pinnae larger, 13–20 × 2.5–5 cm,bases slightly broadened, rounded-truncate, proximal 1–3 pairssometimes slightly shortened, pinnatifid, shortly acuminate atapices; pinnules 15–25 pairs, opposite, spreading, lanceolateand slightly falcate, 1.5–2.5 × 0.5–0.8 cm, bases slightlybroadened and adnate to costae, connected to each other by anarrow wing, lobed to 1/2 distance to costae, apices shortlyacuminate; segments triangular-oblong, acuminate and entire atapices. Veins hidden, pinnate on segments, 2–4 pairs per segment,basal pair arising from above base of costules. Frondsherbaceous, drying dark brown-green, abaxial surfaces withsparse short acicular hairs, hairs along costae, costules, and

348THELYPTERIDACEAEveins denser, adaxially with short setae along costules andveins; rachises and costae reddish brown, glabrous or sparselyshortly hairy. Sori suborbicular or ovate, 1 or 2 per segment,borne above middle of veinlets and close to margins, arrangedin an irregular row on each side of costules. Sporangia glabrousor each with 1 or 2 hairs below annulus. 2n = 62, 124, 186.Forests beside streams; 800–3000 m. Chongqing, Fujian, S Gansu,Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangxi, Sichuan,Taiwan, Yunnan [Bhutan, N India, Myanmar, Nepal, Sri Lanka,Vietnam].1a. Laminae abaxially with dense shortacicular hairs along costae, costules,and veins, sparsely hairy betweenveins ................................................ 7a. var. pyrrhorhachis1b. Laminae abaxially glabrous or atmost with extremely short capitatehairs along rachises, costae, andcostules, glabrous between veins ............. 7b. var. glabrata7a. Pseudophegopteris pyrrhorhachis var. pyrrhorhachis紫柄蕨 ( 原变种 ) zi bing jue (yuan bian zhong)Polypodium pyrrhorhachis Kunze, Linnaea 24: 257. 1851;Dryopteris laterepens (E. W. Trotter ex Hope) C. Christensen;Lastrea pyrrhorhachis (Kunze) Copeland; Macrothelypterispyrrhorhachis (Kunze) Pichi Sermolli; Nephrodium brunneumHandel-Mazzetti; N. distans Diels (1899), not Hooker (1862);Phegopteris brunnea Wallich ex J. Smith; P. distans Mettenius;P. pyrrhorhachis (Kunze) Tagawa; Polypodium distans D. Don(1825), not Kaulfuss (1824); P. distans var. adnatum C. B.Clarke; P. laterepens E. W. Trotter ex Hope; Thelypteris brunnea(Handel-Mazzetti) Ching; T. pyrrhorhachis (Kunze) B. K.Nayar & S. Kaur.Laminae abaxially with dense short acicular hairs alongcostae, costules, and veins, often hairy between veins.Forests beside streams; 800–2400 m. Chongqing, Fujian, S Gansu,Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangxi, Sichuan,Taiwan, Yunnan [Bhutan, N India, Myanmar, Nepal, Sri Lanka,Vietnam].“Polypodium brunneum” (Wallich, Numer. List, no. 333. 1829)belongs here but is a nomen nudum and was not therefore validly published(Melbourne Code, Art. 38.1(a)).7b. Pseudophegopteris pyrrhorhachis var. glabrata (C. B.Clarke) Holttum, Blumea 17: 24. 1969.光叶紫柄蕨 guang ye zi bing juePolypodium distans var. glabrata C. B. Clarke, Trans.Linn. Soc. London, Bot. 1: 544. 1880; Phegopteris distans var.glabrata (C. B. Clarke) Beddome; P. pyrrhorhachis var. glabrata(C. B. Clarke) Tagawa; Thelypteris brunnea var. glabrata(C. B. Clarke) Ching.Lamina abaxially glabrous or with extremely short capitatehairs along rachises, costae, and costules, lacking hairsbetween veins.Forests beside streams; below 3000 m. N Guizhou, W Hubei,Sichuan, Yunnan [N India, Myanmar, SW regions of Himalaya].8. Pseudophegopteris hirtirachis (C. Christensen) Holttum,Blumea 17: 22. 1969.密毛紫柄蕨 mi mao zi bing jueDryopteris hirtirachis C. Christensen in H. Léveillé, Fl.Kouy-Tchéou, 49. 1915; Dryopteris christii H. Léveillé (1915),not C. Christensen (1905); Phegopteris pyrrhorhachis (Kunze)Tagawa var. hirtirachis (C. Christensen) Tagawa; Pseudophegopterispyrrhorhachis (Kunze) Ching var. hirtirachis (C.Christensen) Ching; Thelypteris brunnea (Handel-Mazzetti)Ching var. hirtirachis (C. Christensen) Ching; T. pyrrhorhachis(Kunze) B. K. Nayar & S. Kaur subsp. hirtirachis (C. Christensen)Fraser-Jenkins.Plants to 1 m. Rhizome ascending, stipe base with brownlanceolate scales. Fronds somewhat clustered; stipe red-castaneous,shiny, 20–55 cm, base with scales and whitish hairs;lamina oblong, mostly 50–80 × 15–35 cm, slightly tapering atbase, 2-pinnate-pinnatifid, acuminate and pinnatifid at apex;pinnae opposite to subopposite, sessile, lanceolate, middle pinnaelargest, 9–20 cm or more, mostly 2–6 cm wide, slightlyasymmetrical, acute at apices, proximal 1 or 2 pairs shortened,proximal pair 3–10 cm, pinnatifid or pinnatisect; pinnules 12–20 pairs per pinna, opposite, spreading, basiscopic ones onproximal pinnae slightly longer than acroscopic ones, 0.8–3.5 ×0.5–0.8 cm, bases broadened and adnate to costae, connected toeach other by narrow wings, apices shortly acuminate toobtuse, basal pair of pinnules same size or slightly longer,toothed to pinnatifid to 1/2 distance to costules; segments triangular-oblong,entire to dentate, obtuse to acute at apices. Veinsvisible on both sides, lateral veins simple or forked, mostly 4–15 or more pairs per pinnule, basal pair arising above bases ofcostules. Laminae herbaceous, drying deep green or brownishgreen, abaxial surfaces with dense short whitish hairs along costae,costules, veins, and between veins, adaxially with appressedthick short hairs along rachises, costae, and costules, rachisescastaneous or red-castaneous and with sparse to dense, short,spreading hairs abaxially. Sori suborbicular to oblong, near endsof ultimate veinlets. Sporangia each usually with 2 or more shorthairs below annulus.Forests beside streams; 1500–2000 m. N Guangdong, N Guangxi,S Guizhou, C Sichuan, Taiwan, W Yunnan [NE India, Nepal].Pseudophegopteris hirtirachis is similar to P. pyrrhorhachis inlaminar outline but differs in its rhizomes shorter and ascending,rachises with short, spreading hairs abaxially, costules and costae withdense whitish short acicular hairs abaxially, and sporangia with 2 ormore short hairs.9. Pseudophegopteris yunkweiensis (Ching) Ching, Acta Phytotax.Sin. 6: 315. 1963.云贵紫柄蕨 yun gui zi bing jueThelypteris yunkweiensis Ching, Bull. Fan Mem. Inst.Biol., Bot. 6: 274. 1936; Phegopteris yunkweiensis (Ching)Tagawa.Plants to 1.5 m tall. Rhizome stout (short and ascending?,not seen). Stipe red-castaneous, shiny, 40–55 cm, base withdense brown, ovate-lanceolate, hairy scales, distally glabrous;lamina ovate, ca. 100 × 40–60 cm, not tapering to base, 2-

THELYPTERIDACEAE 349pinnate-pinnatifid, apex acuminate and pinnatifid; pinnae 10 ormore pairs, opposite, or distal ones subopposite and moreoblique, sessile or proximal ones shortly stalked, proximal pairlargest, 30–45 × 13–20 cm, oblong-lanceolate, bases symmetricalor basal several pinnules shortened acroscopically, pinnate-pinnatifid,acuminate at apices, basiscopic pinnules notelongate relative to more distal pinnules; pinnules 15–25 pairsper pinnae, 2.5–3 cm apart, alternate, spreading, mostly freefrom each other, sessile or narrowly adnate to costae, distalmostones connected by a narrow wing, lanceolate, 6–13 × 1.4–2.4cm, truncate at bases, pinnatifid 3/4–4/5 distance to costules,acute at apices; segments 15–20 pairs, falcate-lanceolate, marginscrenate or entire distally, apices acute or obtuse; veinsvisible on both sides, especially adaxially, lateral veins forked,4–7 pairs per segment. Laminae herbaceous, drying dark green,rachises and costae reddish, costules stramineous, abaxiallywith sparse acicular hairs or glabrescent along rachises, costae,and costules, with denser acicular hairs adaxially along costaeand costules, glabrous on both surfaces on and between veins.Sori suborbicular or slightly oblong, borne at middle of lateralveins between costae and margins, 3–6 pairs per segment. Sporangiaglabrous.Forests beside streams. N Guizhou (Zunyi), SE Yunnan (Mengzi)[N Vietnam].10. Pseudophegopteris zayuensis Ching & S. K. Wu, Fl. Xizang.1: 164. 1983 [“zayüensis”].察隅紫柄蕨 cha yu zi bing juePlants ca. 85 cm tall. Rhizome not seen. Stipe reddishbrown, shiny, ca. 30 cm, scaly at base; scales reddish brown,linear-lanceolate, glabrous, or occasionally with a few longhairs dorsally, including rachises with sparse whitish acicularhairs; lamina to ca. 55 × 16 cm, slightly tapering at base, pinnate-pinnatifid,shortly acuminate and pinnatifid at apex; pinnaeca. 25 pairs, opposite or subopposite, proximal 1 or 2 pairs sessile,spreading, proximal pair shortest, ca. 5 cm, middle pinnaeca. 8 × 1.5–2 cm, linear-lanceolate, truncate and symmetrical atbases, pinnatifid to 3/4 of distance to costae, acute at apices;segments ca. 20 pairs per pinna, rectangular, 7–8 × 4–5 mm,entire. Veins visible abaxially, 4 or 5 pairs per segment, basalpair arising from above base of costules. Laminae thinly papery,drying green, subglabrous on both surfaces between veins, costaeat bases red-castaneous abaxially, distally stramineous, shiny,with sparse acicular hairs. Sori orbicular, borne near tips ofacroscopic vein of forked lateral vein pair and close to margin.Sporangia glabrous.● Broad-leaved forests; ca. 2100 m. S Xizang.This species is known only from the type.Pseudophegopteris zayuensis is closely related to P. tibetana, differingprimarily by the basiscopic basal pinnule on proximal pinnae notbeing strongly elongate and the glabrous sporangia. More collectionsare needed to evaluate the importance of these differences.11. Pseudophegopteris rectangularis (Zollinger) Holttum,Blumea 17: 19. 1969.对生紫柄蕨 dui sheng zi bing juePolypodium rectangulare Zollinger, Syst. Verz. 37, 48.1854; Dryopteris moussetii Rosenstock; Phegopteris moussetii(Rosenstock) Alderwerelt; P. oppositipinna Alderwerelt; Polypodiumdistans Kaulfuss var. minor C. B. Clarke; Pseudophegopterisoppositipinna (Alderwerelt) Ching; Thelypteris oppositipinna(Alderwerelt) Ching; T. rectangularis (Zollinger) B. K.Nayar & S. Kaur.Plants 40–70 cm tall. Rhizome short, suberect. Frondsclustered; stipe red-castaneous, shiny throughout, 10–30 cm,with short spreading acicular hairs, base with narrowly lanceolate,brownish, sparsely hairy, thin scales; lamina narrowly oblong-lanceolate,30–50 × 7–12 cm, slightly tapering and withpinnae more spaced proximally, pinnate-pinnatifid; pinnae ca.20 pairs, opposite to subopposite, spreading, sessile and distalones becoming narrowly adnate to rachis, proximal 1–3 pairsslightly shortened, to 3 cm, basal segments of proximal pinnaenot significantly elongate or lobed, distal pinnae lanceolate, 4–7× 0.8–1.8 cm, pinnately lobed to 3/4 of distance to costae,acuminate at apices; segments 12–20 pairs per pinna, obliquelyspreading, oblong-triangular, entire along margins, obtuse atapices. Veins visible, lateral veins simple or occasionally forked,3–5 pairs per segment, basal pair arising from above base ofcostules, acroscopic one of a pair running toward sinus but notreaching margin. Laminae herbaceous, drying greenish, adaxialsurfaces nearly smooth except for hairs along costae, abaxialsurfaces with sparse to many acicular hairs to 0.5 mm, hairsalong costae denser, also sometimes with a few capitate shorthairs between veins, rachises red-castaneous and with denseracicular hairs. Sori suborbicular, borne at middle or near tips oflateral veins, 2–4 pairs per segment. Sporangia each usuallywith 1 or 2 acicular hairs below annulus. 2n = 124.Forests beside streams; 1000–1500 m. N Guangxi, SE Xizang, SEand W Yunnan [Bhutan, NE India, Indonesia, Malaysia, Nepal].“Dryopteris oppositipinnaAlderwerelt” (Bull. Jard. Bot. Buitenzorg,sér. 2, 16: 24. 1914) is invalid (Melbourne Code, Art. 36.1(c)).12. Pseudophegopteris microstegia (Hooker) Ching, Fl. Xizang.1: 162. 1983.禾杆紫柄蕨 he gan zi bing jueNephrodium microstegium Hooker, Sp. Fil. 1: 119. 1862;Lastrea microstegia (Hooker) Beddome; Pseudophegopterispallida (Ching) Ching; Thelypteris brunnea (Handel-Mazzetti)Ching var. pallida Ching.Plants 90–120 cm tall. Rhizome long creeping, with sparse,appressed, brownish, broadly lanceolate scales. Fronds remote;stipe stramineous, occasionally brown-stramineous, 30–40 cm,base with sparse brownish lanceolate scales, distally glabrous;lamina 60–80 × 20–30 cm, tapering proximally, pinnate-pinnatifid,pinnatifid and acuminate at apex; pinnae 20–25 pairs,subopposite, obliquely spreading or spreading, sessile, proximalpinnae linear-lanceolate, 10–15 × 2–3 cm, bases truncate,apices acute, proximal 2 or 3 pairs sometimes slightly shortened,7–10 cm, pinnatifid nearly to costae; segments ca. 25pairs per pinna, opposite, spreading, basal pair of segmentssame shape and size as more distal ones, oblong, 1–1.5 × ca. 0.5cm, coarsely dentate or entire along margins, sparsely ciliate,

350THELYPTERIDACEAEobtuse or truncate at apices. Veins visible abaxially, lateral veinsforked, 6 or 7 pairs per segment, basal pair arising from abovebase of costules. Laminae thinly herbaceous, drying greenish,both surfaces with short hairs along costae and veins, hairsdenser adaxially, rachises stramineous, adaxially densely hirsutealong groove. Sori orbicular to slightly oblong, borne at middleof acroscopic vein of lateral vein pair. Sporangia glabrous.Evergreen broad-leaved forests; 2300–2400 m. Chongqing (Nanchuan),C Sichuan, S and SE Xizang, NW and SE Yunnan [NE India].Pseudophegopteris microstegia is very similar to P. pyrrhorhachis(and treated as a synonym of that by Holttum, 1969), differing mainly inthe stramineous stipes and rachises and laminae abaxially glabrous excepton rachises and costae. The purported differences need furtherstudy.9. CYCLOGRAMMA Tagawa, Acta Phytotax. Geobot. 7: 52. 1938.钩毛蕨属 gou mao jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants medium-sized, terrestrial. Rhizomes short and strong, erect or long creeping, with grayish white unicellular short acicularhairs and few thick scales; scales brown, broadly lanceolate and with acicular and hooked hairs or along margins. Fronds clustered orsparse; stipes ± hairy or subglabrous; laminae oblong or broadly lanceolate, acuminate or pinnatifid at apices; pinnae large, alternateor opposite, lanceolate, proximal several pairs sometimes shortened into auricles, sessile or occasionally shortly stalked, middle pinnaepinnatifid; segments large, lanceolate or suboblong, entire, rounded-obtuse or obtusely pointed at apices. Veins pinnate, free,lateral ones simple, reaching margins above sinuses. Laminae herbaceous or papery, when dry brownish green or nearly brown, bothsurfaces ± with grayish white unicellular short acicular hairs and few hooked thick long hairs, aerophores present at bases of pinnaeand abaxially on rachises. Sori small, consisting of few sporangia, orbicular, dorsifixed at middle or below middle of veinlets,arranged in one row on each side of costules, exindusiate; sporangia shortly stalked and with 1–3 erect short setae or hooked hairs onboth sides of annuli near top. Spores bilateral, orbicular-reniform, perispores clearly echinate or corrugate on surfaces, exosporesmooth. x = 9.About ten species: mainly in mountains of subtropical regions; nine species (seven endemic) in China.1a. Laminae clearly tapering to bases, i.e., proximal 1 to several pairs of pinnae shortened, proximal one auriculate.2a. Proximal 2–5 pairs of pinnae gradually shortened, proximal 1 or 2 pairs of pinnae auriculate, less than 1 cm.3a. Plants to more than 100 cm tall; rhizomes short and erect; adaxially with sparse appressed short hairson intercostal areas, rachises with thick and long acicular hairs and with remaining marks after hairsfallen; sporangia each with 1 or 2 setae near top of annulus ......................................................................... 1. C. auriculata3b. Plants 60–70 cm tall; rhizomes long creeping; adaxially subglabrous on intercostal area, hairson both sides of rachises leaving tubercular marks after fallen; sporangia glabrous ..................................... 2. C. omeiensis2b. Proximal 1–3 pairs of pinnae gradually shortened or abruptly shortened, proximal pair of pinnaenot auriculate, 2–4 cm.4a. Proximal pair of pinnae abruptly shortened; adaxially subglabrous on intercostal areas; sori dorsiflexedat middle of veinlets, located between costae and margins and not confluent when mature ........................... 3. C. leveillei4b. Proximal 2 or 3 pairs of pinnae gradually shortened; adaxially with sparse appressed short acicularhairs; sori dorsifixed near bases of veinlets and close to costules, ± confluent when mature ................. 4. C. costularisora1b. Laminae not tapering to bases, i.e., proximal pair of pinnae similar in size to distal ones.5a. Smaller aerophores not clearly present at bases of pinnae on rachises.6a. Rhizomes long creeping or ascending; pinnae with sparse short acicular hairs abaxially, adaxiallywith dense short acicular hairs along costae; sori dorsifixed below middle of veinlets and slightlyclose to costules .................................................................................................................................................... 8. C. flexilis6b. Rhizomes short and erect; pinnae with dense short acicular hairs abaxially, adaxially with dense shortacicular hairs along costae and mixed with few long acicular hairs; sori dorsifixed above middle ofveinlets and far from costules ............................................................................................................................. 9. C. tibetica5b. Aerophores linear-lanceolate to triangular-lanceolate at bases of pinnae on rachises.7a. Pinnae from middle clearly shortly stalked; sori dorsiflexed at middle or above middle and far fromcostules ...................................................................................................................................................... 5. C. maguanensis7b. Pinnae not stalked; sori dorsifixed below middle of veinlets and slightly close to costules.8a. Rhizomes short and erect; laminae papery; sporangia usually glabrous; plants 1.3–2.2 m tall ...... 6. C. neoauriculata8b. Rhizomes long creeping; laminae herbaceous; sporangia each with 1 or 2 short setae; plantsless than 1 m tall ............................................................................................................................................. 7. C. chunii1. Cyclogramma auriculata (J. Smith) Ching, Acta Phytotax.Sin. 8: 317. 1963.耳羽钩毛蕨 er yu gou mao juePhegopteris auriculata J. Smith, Hist. Fil. 4: 233. 1875;Cyclogramma himalayensis (C. Christensen) Tagawa; C. simulans(Ching) Tagawa; Cyclosorus auriculatus (J. Smith) C. M.Kuo; Dryopteris auriculata (J. Smith) Ching (1931), not (Lin-

THELYPTERIDACEAE 351naeus) Kuntze (1891); D. himalayensis C. Christensen; D.squamistipes (C. B. Clarke) C. Christensen; Glaphyropterissimulans (Ching) H. Itô; Lastrea himalayensis (C. Christensen)Copeland; L. simulans (Ching) Copeland; Polypodium appendiculatumHoffmann var. squamistipes C. B. Clarke; P. auriculatum(J. Smith) Wallich ex Hooker (1863), not Linnaeus (1753);P. subvillosum T. Moore; Thelypteris auriculata (J. Smith) K.Iwatsuki; T. simulans Ching; T. squamistipes (C. B. Clarke)Ching; T. subvillosa (T. Moore) Ching.Plants to more than 1 m tall. Rhizomes short and erect,black, with scales or glabrous when old. Fronds clustered; stipesstrong, 10–20(–30) cm, bases black, with grayish white acicularhairs and scales; scales dark brown, oblong-triangular, bothsurfaces and margins setaceous, distally deep stramineous andto rachises with similar hairs; laminae oblong-lanceolate, (60–)90–130 × 20–30 cm, gradually tapering to bases, pinnate-pinnatifid,acuminate and pinnatifid at apices; pinnae 30–50 pairs,opposite or distal ones sometimes alternate, spreading, sessile,proximal 3–5 pairs of pinnae gradually shortened, proximalones auriculate, only ca. 1 cm; middle pinnae linear-lanceolate,12–20 × 1.5–2.5 cm, bases not tapering, rounded-truncate, pinnatifidto 3/4–4/5 of distance to costae, apices acuminate; segments20–30 pairs, oblong, 5–10 × 4–6.5 mm, margins entireand sparsely acicular hairy, apices rounded-obtuse. Veinlets visible,simple, 10–12 pairs per segment, proximal pair arisingfrom bases of costules. Laminae herbaceous or subpapery,when dry brownish green, abaxially with dense hooked longhairs along costae and slightly pubescent along costules,elsewhere subglabrous, costae grooved adaxially and with denselong acicular hairs, with sparse appressed short hairs on intercostalareas, abaxial sides of rachises with hairy brown aerophores.Sori orbicular and dorsifixed below middle of veinletsand close to costules, 8–10 pairs per segment; sporangia eachwith 1 or 2 setae near top. 2n = 144.Streamsides in evergreen broad-leaved forests; 1600–2800 m. ETaiwan, Yunnan [Bhutan, N India, Indonesia, Myanmar, Nepal].2. Cyclogramma omeiensis (Baker) Tagawa, Acta Phytotax.Geobot. 7: 53. 1938.峨眉钩毛蕨 e mei gou mao juePolypodium omeiense Baker, J. Bot. 26: 229. 1888; Cyclosorusomeiensis (Baker) C. M. Kuo; Dryopteris omeiensis(Baker) C. Christensen; Glaphyropteris omeiensis (Baker) H.Itô; Lastrea omeiensis (Baker) Copeland; Leptogramma omeiensis(Baker) Tagawa; Nephrodium omeiense (Baker) Diels;Thelypteris omeiensis (Baker) Ching.Plants 60–70 cm tall. Rhizomes long creeping. Frondsremote; stipes 15–20 cm, subglabrous above bases; laminae oblong-lanceolate,45–50 × ca. 20 cm, tapering to bases, pinnatepinnatifid,acuminate and pinnatifid at apices; pinnae 25–28pairs, alternate, obliquely spreading, sessile, proximal 2 or 3pairs of pinnae clearly shortened, proximal pair of pinnae onlyca. 1 cm or shorter; middle pinnae linear-lanceolate, 9–12 × ca.2 cm, bases not tapering and rounded-truncate, symmetrical,pinnatifid nearly to costae, apices long acuminate with 1.5–2cm tail; segments 15–22 pairs, falcate-lanceolate, 7–12 × 3–4mm, entire, obtuse or acute at apices. Veins evident abaxially,veinlets simple, 11 or 12 pairs per segment, proximal pairarising above base of costules and all reaching margins abovesinus. Laminae papery, brownish when dry, abaxially withsparse thick long acicular hairs along costae, slightly shortlyhairy along costules, adaxially with dense short acicular hairsalong grooves and mixed hooked thick long hairs, with similarhairs along costules; rachises with denser thick long hairs, pinnaewith brown aerophores at bases abaxially. Sori small, orbicular,dorsifixed below middle of veinlets and close to costules,10 or 11 pairs per segment; sporangia glabrous.● Grassy slopes, streamsides in forests; 900–1700 m. Sichuan,Yunnan.3. Cyclogramma leveillei (Christ) Ching, Acta Phytotax. Sin.8: 317. 1963.狭基钩毛蕨 xia ji gou mao jueDryopteris leveillei Christ, Bull. Acad. Int. Géogr. Bot. 20:176. 1909; D. izuensis Kodama; Leptogramma izuensis (Kodama)H. Itô; Thelypteris leveillei (Christ) C. M. Kuo.Plants 45–100 cm tall. Rhizomes long creeping, includingstipe bases with lanceolate hairy brown thick scales andgrayish white acicular hairs. Fronds approximate; stipe 15–45cm, bases brown, distally stramineous, with sparse short hairsor subglabrous; laminae oblong-lanceolate, 30–55 × 12–20 cm,abruptly tapering to bases, pinnate-pinnatifid, acuminate andpinnatifid at apices; pinnae 12–20 pairs, proximal ones usuallyopposite, spreading or sometimes reflexed; middle pinnae alternate,obliquely spreading; proximal pair clearly shortened, oblong-lanceolate,2–4 × 1–1.5 cm, bases clearly tapering, apicesshortly caudate-acuminate; second pair and more distal pinnaeof similar length or slightly shortened; middle ones 7–13 × 1.5–2 cm, pinnatifid to 3/4 of distance to costae; segments 12–18pairs, oblong or subelliptic, 6–8 × 4–5 mm, entire, rounded atapices. Veins evident abaxially, veinlets simple, 6–10 pairs persegment, proximal pairs arising above bases of costules and allreaching margins above sinus. Laminae herbaceous, brownishgreen after drying, abaxially with denser spreading grayishwhite acicular setae along costae and costules, slightly villousor glabrous on intercostal areas, adaxially with dense short hairsalong costal grooves, sparsely shortly hairy along costules andwith few long acicular hairs near apices, rachises with denseshort acicular hairs on both sides and at intervals mixed withhooked thick long hairs, abaxially with brown aerophores. Sorismall, orbicular, dorsifixed at middle of veinlets, 5–7 pairs persegment; sporangia each usually with 2 or 3 setae near top.Perispores irregularly spinulose, exospore smooth.Humus on rocks in forests; 600–2100 m. Fujian, N Guangdong,Guizhou, Sichuan, Taiwan, Yunnan [Japan].Cyclogramma leveillei is similar to C. omeiensis, both have beentreated as one species in the past. However, the former species differs inhaving only the proximal pair of pinnae shortened, 2–4 cm, neverauriculate, adaxial sides with only short acicular hairs along costae,abaxial sides with spreading acicular hairs, and everywhere withouthooked hairs; and the sporangia each with 2 or 3 setae near tops. Weconclude that it is appropriate to treat the two as distinct species.

352THELYPTERIDACEAE4. Cyclogramma costularisora Ching ex K. H. Shing, Fl.Reipubl. Popularis Sin. 4(1): 322. 1999.无量山钩毛蕨 wu liang shan gou mao juePlants ca. 65 cm tall. Rhizomes short and thick, decumbentor ascending, including stipe bases with sparse brownovate-triangular hairy scales and grayish white dense shorthairs. Fronds approximate; stipes 25–28 cm, bases blackishbrown, distally stramineous, polished and sparsely grayishwhite shortly hairy; laminae oblong-lanceolate, 35–40 × 10–16cm, tapering to bases, pinnate-pinnatifid, acuminate and pinnatifidat apices; pinnae 14–20 pairs, alternate or proximal onessubopposite, proximal 2 or 3 pairs of pinnae graduallyshortened and reflexed, proximal pair shortest, but similarlyshaped, ca. 2 cm; middle pinnae linear-lanceolate, 6.5–9.5 ×1.5–2 cm, bases not tapering, rounded-truncate, pinnatifid to 4/5of distance to costae, apices caudate or entire; segments ca. 14pairs, oblong, 6–8 × 3–3.5 mm, entire along margins and ±recurved proximally, rounded at apices. Veins evident, veinletssimple, ca. 8 pairs per segment, proximal pair arising from baseof costules and all reaching margins above sinus. Laminaeherbaceous, glabrous or brownish green, abaxially includingcostae and costules with dense short acicular hairs, adaxiallywith dense short hairs along costal grooves and with sparselong acicular hairs along costules, with sparse appressed shorthairs on intercostal areas, rachises with dense short acicularhairs and sparse long acicular hairs, abaxially with brown aerophores.Sori orbicular, attached near bases of veinlets and extremelyclose to costules, 6 or 7 pairs per segment and confluentwhen mature. Sporangia each with 1 or 2 setae at tops.● Shaded wet streamsides in forests; ca. 2400 m. SW Yunnan(Jingdong).5. Cyclogramma maguanensis Ching ex K. H. Shing, Fl.Reipubl. Popularis Sin. 4(1): 322. 1999.马关钩毛蕨 ma guan gou mao juePlants ca. 80 cm tall. Rhizomes creeping. Fronds approximate;stipes ca. 37 cm, bases blackish brown, with densegrayish white short hairs and occasionally with brown hairythick small scales, distally stramineous, with sparse acicularhairs or subglabrous when old; laminae lanceolate, ca. 43 × 25cm, not tapering to bases, pinnate-pinnatifid, acuminate andpinnatifid at apices; pinnae ca. 12 pairs, alternate, or severalproximal pairs opposite or subopposite, pinnae below middle1–3 mm stalked, lanceolate, 12–14 × 2.5–3 cm, subtruncate atbases, pinnatifid to 4/5 of distance to costae, long acuminate atapices; segments ca. 20 pairs, oblong, 8–12 × 5–6 mm, entirealong margins and slightly recurved when dry, rounded-obtuseat apices. Veins evident abaxially, veinlets simple, ca. 12 pairsper segment, proximal pair arising from above base of costulesand all reaching margins above sinus. Laminae herbaceous,when dry dark green abaxially, with grayish white acicular hairsalong costae and veins, adaxially yellowish green, with denseappressed short acicular hairs along costal grooves, with densehooked hairs on both sides of rachises, abaxially with brown triangular-lanceolateaerophores. Sori orbicular, dorsifixed atmiddle or above middle of veinlets, slightly above costules, 7–9pairs per segment; sporangia each with 2–4 setae near annuli,setae hooked.● Shaded wet places in forests on slopes; ca. 1000 m. SE Yunnan.6. Cyclogramma neoauriculata (Ching) Tagawa, Acta Phytotax.Geobot. 7: 53. 1938.滇东钩毛蕨 dian dong gou mao jueDryopteris neoauriculata Ching, Bull. Fan Mem. Inst.Biol. 2: 196. 1931; Thelypteris neoauriculata (Ching) Ching.Plants 1.3–2.2 m tall. Rhizomes thick and short, erect.Fronds clustered; stipes 70–120 cm, bases brown, with deepbrown triangular-lanceolate shortly hairy scales and glaucousshort acicular hairs, distally dark stramineous, distally withgrayish white hooked long hairs, rachises similarly hairy; laminaenarrowly oblong, 60–100 cm or more, 25–40 cm wide,bases not tapering, pinnate-pinnatifid, apices acuminate and pinnatifid;pinnae 25–30 pairs, distal ones alternate, proximal onessubopposite, spreading, linear-lanceolate, 15–20 × 3–3.5 cm,broadly cuneate at bases, pinnatifid nearly to costae, caudateacuminateat apices; segments ca. 23 pairs, spreading, middleones oblong-lanceolate, 11–14 mm (proximal 1 or 2 pairsslightly shortened), 5–6 mm wide, entire, rounded-obtuse atapices. Veinlets evident abaxially, simple, 10–15 pairs persegment, proximal pair arising from slightly higher above baseof costules and all reaching margins above sinuses. Laminaepapery, brownish green when dry, subglabrous on both surfacesor with sparse short acicular hairs abaxially, rachises groovedadaxially, with dense hooked long hairs on both sides, abaxiallywith occasional small brown lanceolate scales, pinna basesabaxially with brown linear-lanceolate aerophores. Sori orbicular,smaller, attached below middle of veinlets and close tocostules, 7–12 pairs per segment; sporangia glabrous.● Open forests on mountain slopes; ca. 1800 m. Yunnan.7. Cyclogramma chunii (Ching) Tagawa, Acta Phytotax. Geobot.7: 55. 1938.焕镛钩毛蕨 huan yong gou mao jueThelypteris chunii Ching, Bull. Fan Mem. Inst. Biol., Bot.6: 284. 1936.Plants 70–105 cm tall. Rhizomes strong, long creeping,nearly black, including stipe bases with brown triangular-lanceolatescales and grayish white short hairs. Fronds subapproximate;stipes 30–50 cm, bases dark brown, distally stramineous,subglabrous; laminae oblong-lanceolate, 40–55 × 20–25 cm,pinnae not deflexed at bases, pinnate-pinnatifid, acuminate andpinnatifid at apices; pinnae 12–18 pairs, alternate, obliquelyspreading, or proximal ones opposite, spreading, sessile, linearlanceolate,10–15 × 2–3 cm, bases slightly tapering, roundedtruncate,pinnatifid to 3/4 of distance to costae, apices shortlycaudate-acuminate; segments 16–20 pairs, oblong-lanceolate,8–12 × 4–7 mm, entire, rounded-obtuse at apices. Veins not evident,veinlets simple, 10–11 pairs per segment, proximal pairarising from slightly higher above base of costules, or basiscopicveinlets arising from base of costules and all reachingmargins above sinuses. Laminae thinly herbaceous, when dry

THELYPTERIDACEAE 353brownish green, abaxially with acicular short hairs along costaeand costules, adaxially with short hairs along costal grooves, rachiseswith dense hooked hairs, pinna bases abaxially with deepbrown, linear-lanceolate or linear-bent aerophores. Sori small,orbicular, attached below middle of veinlets and slightly closeto costules, 7–10 pairs per segment; sporangia each with 1 or 2short setae near top of annuli.● On rocks in forests in mountain valleys. SW Guangdong (Xinyi).8. Cyclogramma flexilis (Christ) Tagawa, Acta Phytotax. Geobot.7: 55. 1938.小叶钩毛蕨 xiao ye gou mao jueAspidium flexile Christ, Bull. Acad. Int. Géogr. Bot. 11:252. 1902; A. melanorhizum Christ (1901), not Desvaux(1827); Dryopteris flexilis (Christ) C. Christensen; D. omeiensis(Baker) C. Christensen var. flexilis (Christ) C. Christensen; D.subthelypteris Christ ex C. Christensen; Thelypteris flexilis(Christ) Ching.Plants 30–60 cm tall. Rhizomes long creeping or ascending,black, with sparse black-brown hairy lanceolate scales.Fronds approximate; stipes 10–30 cm, bases black, with sparseblackish brown triangular-lanceolate scales and dense glaucousshort acicular hairs, distally stramineous and subglabrous; laminaenarrowly lanceolate, 20–30(–40) × 6–10(–14) cm, nottapering to bases, pinnate-pinnatifid, caudate and long acuminateand pinnatifid at apices; pinnae 12–20 pairs, alternate orproximal ones opposite, obliquely spreading, sessile; proximalones linear-lanceolate, 3.5–8 × 0.8–2.5 cm, bases symmetrical,rounded-truncate, pinnatifid and reaching narrow wings of bothsides, apices shortly caudate-acuminate; segments 7–13(–15)pairs, oblong, 4–12 × 2–4 mm, margins entire, apices roundedobtuse.Veins evident abaxially, veinlets simple, 4–9 pairs persegment, proximal pair arising from above base of costules andall reaching margins above sinuses. Laminae papery, when drydark green, abaxially with glaucous short hairs, hairs along costaeand costules dense and mixed with few acicular long hairs,adaxially with dense short acicular hairs along costal grooves,rachises with dense short acicular hairs on both sides, abaxialsides mixed with few thick and long acicular hairs and withsmaller brownish tuberculate aerophores. Sori small, orbicular,attached below middle of veinlets and slightly closer to costules,4–6 pairs per segment; sporangia each with 1 or 2(or 3)short setae near tops.● Forests on limestone; 300–1400 m. Guizhou, Sichuan.9. Cyclogramma tibetica Ching & S. K. Wu, Fl. Xizang. 1:157. 1983.西藏钩毛蕨 xi zang gou mao juePlants ca. 75 cm tall. Rhizomes short and erect, includingstipe bases with sparse scales and short hairs. Fronds clustered;stipes ca. 30 cm, dark brown proximally, distally to rachisesbrownish stramineous, shortly hairy distally; laminae lanceolate,ca. 45 × 18 cm, nearly tapering to bases, pinnate-pinnatifid,acuminate and pinnatifid at apices; pinnae ca. 20 pairs,above middle alternate, spreading or proximal ones subopposite,reflexed, sessile, linear-lanceolate, 8–9.5 × 1.4–1.7 cm,nearly flat-truncate and slightly narrowed at base, pinnatifidnearly to costae, acuminate or caudate at apices; segments ca.20 pairs, oblong, entire along margins. Veins evident abaxially,veinlets simple, 8 or 9 pairs per segment, proximal pair arisingfrom base of costules and all reaching margins above sinuses.Laminae papery, when dry dark green abaxially, adaxiallysomewhat brown, both surfaces including rachises with denseshort acicular hairs and few thick and long acicular hairs. Soriorbicular, dorsifixed slightly above middle of veinlets; sporangiaeach with 1(sometimes 2) hooked hairs near top.● Broad-leaved forests; ca. 1500 m. SE Xizang (Mêdog).10. LEPTOGRAMMA J. Smith, J. Bot. (Hooker) 4: 51. 1841.茯蕨属 fu jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants medium-sized, terrestrial. Rhizomes short and erect or ascending, with sparse scales; scales ovate-oblong or lanceolate,reddish brown, hairy. Fronds clustered; stipes deep stramineous, with sparse scales proximally, throughout with grayish white, mostlyunicellular acicular long hairs and unicellular short setae; laminae oblong, hastate, or lanceolate, pinnate-pinnatifid; pinnae 7 or 8pairs (or more), oblique or spreading, lanceolate, bases rounded or truncate, symmetrical, usually sessile, apices obtuse or acuminate;proximal 1 or 2 pairs or several pairs of pinnae free, distally ± adnate to rachises, proximal pair not or slightly shortened, sometimesprolonged, costae grooved adaxially, pinnae pinnatifid and usually reaching 1/2–2/3 of distance to costae; segments orbicular oroblong, entire and rounded at apices. Veinlets free, 3–6 pairs per segment, not forked, reaching margins, rarely reaching sinus.Laminae herbaceous or papery, when dry dark brown or brownish green, usually with acicular hairs or short setae on both surfaces,or with both mixed. Sori elongate, borne along veinlets, length slightly shorter than veinlets, exindusiate; sporangia each with 2–6erect setae at top; spores bilateral, reniform, echinate on surfaces. x = 36.About 15 species: tropical and subtropical areas of Asia, west to Africa; nine species (six endemic) in China.“Leptogramma intermedia” (Ching in C. F. Zhang, Fl. Zhejiang 1: 154. 1993) was not validly published because no Latin description ordiagnosis, or reference to such, was provided (Melbourne Code, Art. 39.1). It does not appear to represent a distinct species.1a. Proximal pair of pinnae longer than distal ones, hence laminae hastate ....................................................................... 1. L. tottoides1b. Proximal pair of pinnae similar length as distal ones or slightly shorter, laminae not hastate.2a. Proximal free pinnae with short stalks.

354THELYPTERIDACEAE3a. Laminae not tapering to bases; free pinnae 5 pairs; sporangia each with 3 or 4 setae near top ...................... 2. L. scallanii3b. Laminae slightly tapering to bases; free pinnae 9 pairs; sporangia each with 1 or 2 setae near top .................... 3. L. pozoi2b. All pinnae sessile.4a. Stipes dark brown; laminae somewhat leathery and brown when dry; free pinnae 7–9 pairs .................. 4. L. huishuiensis4b. Stipes stramineous or deep stramineous; laminae herbaceous and greenish to deep green when dry(rarely dark brown); free pinnae 1–7 pairs.5a. Laminae dark brown when dry; pinnae without hairs on intercostal areas ........................................... 5. L. yahanensis5b. Laminae greenish to deep green when dry, with hairs on intercostal areas.6a. Rhizomes ascending; laminae yellowish green when dry ......................................................... 6. L. jinfoshanensis6b. Rhizomes erect; laminae greenish to deep green when dry.7a. Laminae greenish when dry; sporangia subglabrous near tops ......................................... 7. L. centrochinensis7b. Laminae deep green when dry; sporangia each with 2–4 setae near top.8a. Pinnae with sparse acicular hairs adaxially, abaxially pubescent throughout ..................... 8. L. himalaica8b. Pinnae with dense acicular hairs on both surfaces ..................................................................... 9. L. sinica1. Leptogramma tottoides H. Itô, Bot. Mag. (Tokyo) 49: 434.1935.小叶茯蕨 xiao ye fu jueCyclosorus tottoides (H. Itô) C. M. Kuo; Leptogrammacaudata Ching; L. totta (Schlechtendahl) J. Smith var. tottoides(H. Itô) H. Itô; Stegnogramma tottoides (H. Itô) K. Iwatsuki;Thelypteris caudata (Ching) C. F. Reed.Plants 17–32 cm tall. Rhizomes short and erect, includingstipe bases with sparse reddish brown broadly lanceolate scalesand grayish white acicular hairs. Fronds clustered; stipes 10–17cm, slender, dark stramineous, throughout with sparse unicellularacicular long hairs; laminae 1-pinnate, hastate-lanceolate,14–20 × 4–6 cm, bases hastate and broadest, apices acuminate;pinnae 16–20 pairs, subopposite and spreading, subsessile,proximal 2 or 3 pairs free, distal ones ± adnate torachises, proximal pair largest, spreading, oblong-lanceolate, 2–3 × ca. 1 cm, truncate and symmetrical at bases, pinnatifid to1/2 distance to costae, obtuse or shortly acute at apices; segments4–6 pairs, ovate-orbicular, entire; second pair of pinnaeabruptly shortened, 1.5–2 cm, of similar shape as proximal pair,middle pairs of pinnae of similar size and shape, bases adnate torachises but free from each other, distal pinnae slightly shorterthan middle ones, bases connected to each other by broadwings, entire or crenate proximally. Veinlets evident, 3 or 4pairs per segment, simple, proximal pair arising from far abovebase of costules and acroscopic one reaching sinus or marginsof sinus, basiscopic one reaching margin far above sinus. Laminaethinly herbaceous, dark brown when dry, pinnae throughoutwith acicular hairs adaxially, abaxially including rachises withspreading grayish white acicular fine hairs, veins sparsely puberulent.Sori linear, usually attached on proximal half on proximalpair of veinlets; sporangia each with 3 or 4 setae near top.● On rocks in forests; 800–2500 m. Fujian, Guizhou, Jiangxi, Taiwan,Zhejiang.2. Leptogramma scallanii (Christ) Ching, Sinensia 7: 101.1936.峨眉茯蕨 e mei fu jueAsplenium scallanii Christ, Boll. Soc. Bot. Ital. 1901: 296.1901; Dryopteris scallanii (Christ) C. Christensen; Stegnogrammascallanii (Christ) K. Iwatsuki.Plants 20–30 cm tall. Rhizomes short and erect, includingproximal parts of stipes with sparse reddish brown lanceolate,hairy scales and dense acicular hairs. Fronds clustered; stipes 5–10 cm, dark stramineous, with dense acicular long hairs distally;laminae 1-pinnate, oblong, 14–20 × 5–7 cm, bases nottapering, apices acuminate and pinnatifid; pinnae 10–14 pairs,alternate, proximal 3–5 pairs shortly stalked, similar in size todistal ones; middle pinnae lanceolate, 2.5–4 × 0.7–1 cm, basessubtruncate, symmetrical, pinnatifid to 1/3–1/2 of distance tocostae, apices acuminate or shortly acuminate; segments ca. 10pairs, ovate-orbicular, entire, separated from each other byobtriangular sinuses. Veins evident, 4 or 5 pairs of veinlets persegment, proximal pair arising from above base of costules,acroscopic veinlet reaching sinuses. Laminae papery, when drybrownish green, adaxially with 1 or 2 acicular hairs alongcostae and veinlets, abaxially with sparse acicular thick hairsalong costae and costules. Sori oblong or linear, attached onabaxial sides along veinlets, 1 or 2 pairs per segment. Sporangiaeach with 2 or 3 setae near top.Wet places in forests, on rocks in ravines; 400–1400 m. Fujian,Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Yunnan, Zhejiang[N Vietnam].3. Leptogramma pozoi (Lagasca) Heywood, Repert. Spec.Nov. Regni Veg. 64: 19. 1961.毛叶茯蕨 mao ye fu jueHemionitis pozoi Lagasca, Nov. Gen. Pl. 33. 1816; Cyclosoruspozoi (Lagasca) C. M. Kuo; Gymnogramma tottaSchlechtendahl; G. totta var. mollissima Kunze; Lastrea africana(Desvaux) Ching; Leptogramma africana (Desvaux)Nakai ex Mori; L. mollissima (Kunze) Ching; L. pozoi subsp.mollissima (Kunze) Nakaike; L. totta (Schlechtendahl) J. Smith;Polypodium africanum Desvaux; P. totta Willdenow (1810), notThunberg (1800); Stegnogramma mollissima (Kunze) Fraser-Jenkins; S. pozoi (Lagasca) K. Iwatsuki; Thelypteris mollissima(Kunze) Thapa; T. pozoi (Lagasca) C. V. Morton.Plants 35–45 cm tall. Rhizomes short and erect, withbrown lanceolate hairy scales. Fronds clustered; stipes ca. 13cm, stramineous, bases with brown scales, distally with grayishwhite acicular hairs; laminae lanceolate, 20–28 × 10–13 cm,slightly tapering proximally, pinnate-pinnatifid, pinnate-acuminateat apices; lateral free pinnae 9 pairs, proximal pinnae

THELYPTERIDACEAE 355shortly stalked, proximal pair of similar shape and size and reflexed;middle pinnae lanceolate, ca. 6 × 1.5 cm, bases truncate,pinnatifid to 1/2 of distance to costae, apices long caudate;segments rectangular, entire, obtuse-rounded at apices. Veinletsevident, simple, basiscopic vein of proximal pair arising fromcostae, acroscopic vein arising from base of costules andreaching margin. Laminae herbaceous, deep green when dry,costae and veins with acicular hairs, also with acicular shorthairs on intercostal areas of both surfaces. Sori narrowly ovate,attached on middle or proximal parts of veinlets and slightlycloser to bases. Sporangia each with 1 or 2 setae near annuli. 2n= 144.Lower mountain areas; 300–600 m. SE Taiwan (Lan Yu) [S India,Indonesia, Japan, Sri Lanka; Africa].Fraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes,195. 2008) believes Thelypteris mollissima is a strictly Asiantaxon that should be separated from the Macronesian and African T.pozoi. This is not accepted here.4. Leptogramma huishuiensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 323. 1999.惠水茯蕨 hui shui fu juePlants to 33 cm tall. Rhizomes not seen. Stipes 10–13 cm,dark brown, bases with sparse brown lanceolate scales, distallywith dense grayish white acicular long hairs; laminae lanceolate,12–25 × 5–10 cm, slightly tapering to bases, pinnate-pinnatifid,pinnatifid-acuminate at apices; lateral free pinnae 7–9pairs, sessile, distal pinnae adnate to rachises, proximal pair ofsimilar shape and size as distal ones; middle pinnae lanceolate,ca. 5 cm, truncate at bases, pinnatifid to 1/2 of distance to costae,acuminate at apices; segments ca. 12 pairs per pinna, oblong,entire, separated from each other by obtriangular sinuses,obtuse at apices. Laminae somewhat leathery, when dry darkbrown, rachises, costae, and veinlets with dense acicular hairsabaxially, adaxial sides with short setae, abaxially with sparseshort setae on intercostal areas. Sori narrowly ovate, attachedon middle to proximal parts along veinlets. Sporangia each withmore than 3 setae near top.● Guizhou (Huishui).5. Leptogramma yahanensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 324. 1999.雅安茯蕨 ya an fu juePlants to 18 cm tall. Rhizomes short and erect, includingstipe bases with sparse reddish brown lanceolate, hairy scalesand acicular hairs. Stipes 5–6 cm, throughout with acicularhairs; laminae broadly lanceolate, 10–12 × ca. 4 cm, nottapering to bases, 1-pinnate, acuminate at apices; pinnae ca. 10pairs, alternate except for proximal pair, others ± adnate torachises; middle pinnae lanceolate and slightly falcate, 2–2.5 ×0.7–0.8 cm, undulate or crenate, shortly acuminate at apices.Veins evident, veinlets 2 pairs per group, proximal pair arisingfrom far above base of costules, acroscopic veinlet reachingsinus. Laminae herbaceous, when dry dark brown, adaxiallysubglabrous except with sparse, short hairs along costae, abaxiallywith spreading acicular long hairs along costae and costules.Sori linear, attached on middle on acroscopic veinlet ofeach group. Sporangia each with 2 or 3 setae near top.● On rocks in forests; ca. 1000 m. SW Sichuan (Ya’an).6. Leptogramma jinfoshanensis Ching & Z. Y. Liu, Bull. Bot.Res., Harbin 4(3): 17. 1984.金佛山茯蕨 jin fo shan fu juePlants 30–35 cm tall. Rhizomes ascending. Fronds clustered;stipes ca. 22 cm, stramineous, with dense grayish whitelanceolate acicular hairs; laminae lanceolate, ca. 30 × 8–18 cm,1-pinnate, pinnatifid-acuminate at apices; free-pinnae 3–7 pairs,sessile, truncate, proximal pair of similar shape and size as distalones, bases slightly tapering, apices caudate; middle pinnaelanceolate, 7–8 × ca. 1.5 cm, truncate at bases, pinnatifid to1/2–2/3 of distance to costae, long caudate at apices; segmentsrectangular, with acicular hairs on intercostal areas of both surfaces,with dense acicular setae along rachises and veins, entire,rounded at apices. Veins evident, veinlets reaching marginsabove sinuses, 4 or 5 pairs per segment. Laminae herbaceous,when dry yellowish green, all with acicular hairs on both surfaces.Sori ovate, attached near bases of veinlets. Sporangiaeach with 2–4 acicular hairs near top.● On wet rocks in broad-leaved forests; ca. 1800 m. Chongqing(Nanchuan).7. Leptogramma centrochinensis Ching ex Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 323. 1999.华中茯蕨 hua zhong fu juePlants 30–35 cm tall. Rhizomes short and erect, includingstipe bases with sparse broadly lanceolate hairy scales andacicular short hairs. Fronds clustered; stipes 8–14 cm, deepstramineous, above bases with dense short hairs and mixed withfew acicular long hairs; laminae lanceolate, 18–24 × ca. 6 cm,1-pinnate, acuminate at apices; pinnae 14–16 pairs, alternate,spreading, sessile, proximal pair not shortened and of similarshape and size as distal ones, proximal 2 pairs free, distal ones± adnate to rachises; middle ones broadly lanceolate, 3–3.5 ×ca. 1 cm, bases truncate, symmetrical (proximal pair of pinnaeslightly broad and basal basiscopic one cuneate), pinnatifid to1/2 of distance to costae, apices shortly acuminate; segments 7or 8 pairs, broadly ovate, entire, separated by obtriangularsinuses. Veins evident, (3 or)4 pairs of veinlets per segment,proximal pair (basiscopic veinlet) arising from bases of costaeand all reaching margins above sinuses. Laminae herbaceous,when dry greenish, adaxially throughout with appressed acicularhairs, abaxially pubescent along costae and costules andwith acicular short hairs along rachises. Sori orbicular or oblong,attached on middle of veinlets. Sporangia subglabrous.● Wet places in open forests. W Hubei.8. Leptogramma himalaica Ching, Sinensia 7: 100. 1936.喜马拉雅茯蕨 xi ma la ya fu jueLeptogramma yunnanensis Ching; Stegnogramma himalaica(Ching) K. Iwatsuki; Thelypteris himalaica (Ching) C. F.Reed.

356THELYPTERIDACEAEPlants 30–35 cm tall. Rhizomes short and erect. Frondsclustered; stipes to 20 cm, grayish stramineous proximally,densely pubescent and with 1 or 2 reddish brown lanceolatehairy scales, distally stramineous, polished and subglabrous;laminae lanceolate, 22–26 × 6.5–8 cm, not tapering to bases,1-pinnate, acuminate at apices; pinnae 12–16 pairs, opposite,distal ones alternate, sessile, proximal 2 or 3 pairs free, distalones ± adnate to rachises; middle pinnae lanceolate, 3–3.5 × ca.1 cm, bases truncate, symmetrical, pinnatifid to 1/3 of distanceto costae, apices shortly pointed or acute; segments 6–8 pairs,ovate, separated by obtriangular sinuses. Veins evident, 3 pairsper segment, proximal pair arising above base of costules andall reaching sinuses. Laminae herbaceous, when dry dark green,pinnae with sparse acicular hairs adaxially, abaxially pubescentthroughout. Sori linear, attached on middle of veinlets. Sporangiaeach with 3 or 4 setae near top.Shaded places by rocks or slopes; 2100–2500 m. Xizang, NWYunnan [N India].9. Leptogramma sinica Ching ex Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 323. 1999.中华茯蕨 zhong hua fu juePlants 16–23 cm tall. Rhizomes short and erect, includingstipe bases with sparse reddish brown ovate-lanceolate hairyscales and grayish white unicellular acicular long hairs. Frondsclustered; stipes 5–7 cm, slender, deep stramineous; laminaeoblong-ovate, 11–16 × 5–7 cm, not tapering to bases, 1-pinnate,apices pinnatifid-acuminate; pinnae 7–14 pairs, alternate, proximalpair not shortened, sessile, second pair and more distalpairs ± adnate to rachis; middle pinnae broadly lanceolate, 2.6–4 × ca. 1 cm, acroscopic bases truncate, basiscopic basesrounded, pinnatifid to 1/2 distance to costae, apices shortlyacuminate; segments 7–11 pairs, oblong, entire, separated byobtriangular-sinuses. Veins evident, 4 or 5 pairs per segments,proximal pair arising above base of costules, acroscopic veinreaching sinuses. Laminae herbaceous, when dry dark green,both surfaces with dense acicular hairs on intercostal areas,except with acicular hairs along costae and veins. Sori linear,attached along veinlets, 2 or 3 pairs per segment, free from eachother. Sporangia each with 2 or 3 setae near top.● In shade and on wet rocks in forested ravines; ca. 800 m. Guizhou,Hunan.11. GLAPHYROPTERIDOPSIS Ching, Acta Phytotax. Sin. 8: 320. 1963.方杆蕨属 fang gan jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants medium-sized to large, terrestrial, ± with acicular hairs. Rhizomes short and thick, decumbent or ascending, sessile, withsparse scales. Fronds clustered or approximate; stipes strong, with sparse brown lanceolate scales at bases; laminae elliptic, nottapering to bases, pinnate-pinnatifid; pinnae large, linear-lanceolate, sessile, free, opposite or subopposite, pinnatifid nearly to costae;rachises rectangular abaxially, flat, glabrous or with sparse short hairs, when dry reddish, costae grooved adaxially and densely hairyabaxially; segments large, falcate-lanceolate. Veins free, pinnate on segments, veinlets simple, prominent and reaching margins,proximal pair only reaching nearby line of transparent membrane below sinus or to margins above sinuses. Laminae herbaceous,papery or leathery, when dry yellowish green, rachises and costae ± with grayish white long hairs on both sides, not glandular. Soriorbicular, attached at bases of veinlets, in a row on each side close to costules, usually confluent into a line when mature, exindusiateor indusiate. Sporangia usually with acicular hairs near tops. Spores elliptic, bilateral, irregularly echinate or small tuberculate onsurfaces. x = 12(36).Twelve species: Bhutan, China, N India, S Japan, N Myanmar, Nepal, Philippines, N Vietnam; 11 species (ten endemic) in China.1a. Sori exindusiate.2a. Sporangia glabrous .............................................................................................................................................. 1. G. erubescens2b. Sporangia bearing hairs .................................................................................................................................... 2. G. sichuanensis1b. Sori indusiate.3a. Sori obviously indusiate.4a. Indusia glabrous ....................................................................................................................................... 8. G. jinfushanensis4b. Indusia with acicular hairs.5a. Rachises, costae, veins, and intercostal areas all glabrous abaxially ......................................................... 9. G. glabrata5b. Rachises, costae, veins, and intercostal areas abaxially all with dense acicular hairs.6a. Laminae glabrous adaxially on intercostal area, sporangia glabrous .................................................. 10. G. villosa6b. Laminae adaxially with short setae on intercostal area, sporangia bearing denseacicular hairs ............................................................................................................................. 11. G. rufostraminea3b. Sori with small scale-shaped indusia, usually obscured by mature sporangia and not easily seen.7a. Sporangia glabrous.8a. Plants over 1 m tall; proximal pair of veins from adjacent segments reaching bottom of sinuses ............... 3. G. mollis8b. Plants normally 50–60 cm tall; proximal pair of lateral veins on segments reaching marginsabove sinuses ............................................................................................................................................ 4. G. emeiensis7b. Sporangia bearing hairs.9a. Proximal pair of veinlets on segments reaching both sides of bottom of sinuses; rachises, veins,and intercostal areas glabrous .................................................................................................................. 5. G. eriocarpa

THELYPTERIDACEAE 3579b. Proximal pair of veinlets on segments reaching margins above sinuses; rachises, veins, andintercostal areas ± with short setae or acicular hairs.10a. Rachises, costae, and veins all with dense long acicular hairs, laminae only with short setaeabaxially, glabrous on intercostal areas; Sichuan ........................................................................... 6. G. splendens10b. Rachises with sparse short setae abaxially, costae and veins with sparse long setae, laminaewith sparse setae on veins and intercostal areas; Yunnan ................................................................... 7. G. pallida1. Glaphyropteridopsis erubescens (Wallich ex Hooker)Ching, Acta Phytotax. Sin. 8: 320. 1963.方杆蕨 fang gan juePolypodium erubescens Wallich ex Hooker, Sp. Fil. 4:236. 1862; Asplenium distans D. Don var. cadieri Christ; Cyclosoruserubescens (Wallich ex Hooker) C. M. Kuo; Dryopterisbraineoides (Baker) C. Christensen; D. erubescens(Wallich ex Hooker) C. Christensen; D. reflexa Ching; Glaphyropteriserubescens (Wallich ex Hooker) Fée; Lastrea erubescens(Wallich ex Hooker) Copeland; Nephrodium braineoides(Baker) Diels; N. erubescens (Wallich ex Hooker) Diels; Phegopteriserubescens (Wallich ex Hooker) J. Smith; Polypodiumbraineoides Baker; Thelypteris erubescens (Wallich ex Hooker)Ching.Plants 2–3 m tall or more. Rhizomes stout, decumbent,woody and glabrous. Fronds clustered; stipes 1–2 m, thickerthan 1 cm, ribbed, throughout glabrous, stramineous and oftenreddish; laminae 100–200 × 25–50 cm, not tapering to bases,pinnate-pinnatifid or subbipinnate, acuminate and pinnatifid atapices; pinnae 40–50 pairs per frond, opposite, sessile, proximalseveral pairs strongly oblique distally, bases tapering; middlepinnae spreading, linear, (10–)20–30 or more × (1.5–)2.5–4 cm,bases truncate and close to costae, pinnatifid nearly to bothlateral narrow wings of costae, apices acuminate; segmentsmany (ca. 50 pairs), pectinately arranged, spreading, linear-lanceolate,slightly falcate, 1.4–2 × ca. 0.4 cm, slightly broadenedto bases, entire, separated by narrow sinuses, pointed at apices.Veins evident, costae grooved adaxially, pubescent, abaxiallyrounded and raised, glabrous or sometimes sparsely hairy, veinlets12–23 pairs per segment, simple, proximal pair arising frombase of costules and reaching both sides of rounded sinuses.Laminae papery, when dry greenish or yellowish green, pinnaeglabrous adaxially, with 1 or 2 acicular hairs along margins, orsparsely hairy abaxially; rachises rectangular abaxially, flat,stramineous or reddish, with grayish white acicular hairs whenyoung and then deciduous. Sori orbicular, 10–15 pairs per segment,attached at bases of veinlets, close to both sides of costulesand in 1 row on each side, confluent into lines when mature,exindusiate. Sporangia glabrous.Forests in ravines of low mountains; 800–2000 m. Guizhou, Sichuan,Taiwan, Yunnan [Bhutan, N India, S Japan, Kashmir, N Myanmar,Nepal, Pakistan, Philippines, N Vietnam].“Glaphyropteridopsis erubescens var. mollis” (Ching, Vasc. Pl.Hengduan Mount. 1: 97. 1993) was not validly published because noLatin description or diagnosis, or reference to such, was provided andno type was indicated (Melbourne Code, Art. 39.1 and 40.1). The name“Christella erubescens” (H. Léveillé, Fl. Kouy-Tchéou, 474–476.1915) belongs here but is a nomen nudum and was not therefore validlypublished (Art. 38.1(a)).Thelypteris ×erubesquirolica W. C. Shieh & J. L. Tsai (J. Sci.Engin. 24: 7. 1987), described from Taiwan (Nantou), appears to be apresumed hybrid between Glaphyropteridopsis erubescens and Pseudocyclosorusesquirolii.2. Glaphyropteridopsis sichuanensis Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 324, 131. 1999 [“sichanensis,” p. 324].四川方杆蕨 si chuan fang gan juePlants to 1.2 m tall. Rhizomes strong, creeping, darkbrown, woody, hard. Fronds approximate; stipes ca. 60 cm,dark brown, with brown lanceolate scales, distally stramineous,glabrous; laminae broadly lanceolate, ca. 60 × 30 cm, slightlytapering to bases, pinnate-pinnatifid, pinnatifid-acuminate atapices; lateral free pinnae to 25 pairs, proximal ones subopposite,distal ones alternate, sessile, spreading; middle pinnaeoblong-lanceolate, ca. 17 × 1.5 cm, flatly truncate at bases, pinnatifidnearly to costae, pinnatifid-acuminate at apices; segmentsca. 40 pairs, lanceolate, ca. 1 × 0.4 cm, entire, separatedby small interval, apices slightly bent distally and acute; proximalpair extremely elongated, acroscopic one to 2 cm. Veinsevident, raised abaxially, proximal pair of veinlets arising frombase of costules, acroscopic veins reaching bottom of sinuses,basiscopic one reaching margin above sinus, 15 or 16 pairs ofveinlets per segment. Laminae papery, when dry brownish, withacicular long hairs on intercostal areas abaxially, glabrous onintercostal areas adaxially, setose along margins; rachises withsparse acicular hairs abaxially, costae and veins with dense longacicular hairs, with dense acicular setae along both sides ofgrooves of rachises and costae, with sparse appressed shortsetae along both sides of longitudinal grooves adaxially of costae,abaxial sides of veins subglabrous. Sori orbicular, attachedat bases of veinlets and close to costules, 8–12 pairs per segment,exindusiate. Sporangia bearing fine acicular hairs.● Sichuan.3. Glaphyropteridopsis mollis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 324. 1999.柔弱方杆蕨 rou ruo fang gan juePlants ca. 1.5 m tall. Rhizomes creeping, thick, ca. 1.5 cm,dark brown, woody and hard. Fronds approximate; stipes ca. 80cm, dark brown, distally deep stramineous and glabrous; laminaebroadly lanceolate, ca. 62 × 38 cm, pinnatifid-acuminate atapices; proximal pinnae nearly not shortened, pinnate-pinnatifid;free pinnae ca. 31 pairs, opposite, spreading, sessile; middlepinnae lanceolate, ca. 20 × 2.5 cm, pinnatifid and subcuneate atbases, pinnatifid and long acuminate at apices; segments ca. 31pairs, lanceolate, ca. 12 × 4–4.5 mm, entire, acute at apices;proximal pair extremely elongated to 2 cm. Veins visible onboth sides, costae raised abaxially, adaxially impressed, 15–17pairs of veinlets per segment, proximal pair arising from base ofcostules and all reaching bottom of sinuses. Laminae darkbrown when dry, abaxially with acicular hairs along rachises,

358THELYPTERIDACEAEcostae, veins, and intercostal areas, adaxially with dense shortsetae only along costae and elsewhere glabrous. Sori orbicularand attached at bases of veinlets and close to costules; indusiasmall, bearing hairs. Sporangia glabrous.● Broad-leaved forests; 800–900 m. Sichuan (Emei Shan).4. Glaphyropteridopsis emeiensis Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 324. 1999.峨眉方杆蕨 e mei fang gan juePlants ca. 80 cm tall. Rhizomes not seen. Stipes ca. 40 cm,stramineous, glabrous; laminae lanceolate, ca. 70 × 20–25 cm,pinnate-pinnatifid, pinnatifid-acuminate at apices; lateral freepinnae ca. 25 pairs, obliquely spreading, proximal ones subopposite,distal ones alternate, subsessile; middle pinnae lanceolate,ca. 13 × 1.7 cm, truncate at bases, pinnatifid, pinnatifidacuminateat apices; segments ca. 30 pairs, ca. 3 mm wide,entire, obtuse-pointed at apices; proximal pair (particularlyacroscopic one) extremely elongated. Veins visible on bothsides, adaxially impressed, costules raised abaxially, veinlets 12pairs per pinna, proximal pair arising from closely above baseof costules and all reaching margins above sinuses. Laminaedark green or green when dry, papery, rachises, costae, veins,and abaxial sides of intercostal areas all with dense acicularhairs, adaxially with dense short hairs along grooves of costae,veins, intercostal areas, and margins with short setae. Sori orbicularand attached on proximal parts of veinlets, close to costules,indusiate and hairy. Sporangia hairy.● Mixed forests at roadsides, rock crevices by streams; 1800–1900 m. Sichuan (Emei Shan).5. Glaphyropteridopsis eriocarpa Ching, Acta Phytotax. Sin.8: 321. 1963.毛囊方杆蕨 mao nang fang gan juePlants to 1.2 m tall. Rhizomes strong and decumbent, subwoodyand glabrous. Fronds subapproximate; stipes ca. 50 cm,stramineous, glabrous; laminae narrowly oblong, ca. 70 × 27cm, not tapering to bases, pinnate-pinnatifid and reachingboth lateral narrow wings of costae, acuminate and pinnatifid atapices; pinnae 25–30 pairs, opposite or subopposite, sessile,spreading, proximal several pairs not shortened, reflexed, basestapering; middle pinnae linear-lanceolate, ca. 17 × 2 cm, basestruncate and to 3 cm wide, symmetrical, pinnatifid to both lateralnarrow wings of costae, apices long acuminate; segments25–30 pairs, spreading, lanceolate, ca. 9 mm (proximal pair to1.5 cm), ca. 3 mm wide, gradually broadened to bases, entire,acute at apices. Veins evident, 11 or 12 pairs of veinlets per segment,proximal pair arising from base of costules and allreaching both sides near sinuses. Laminae papery, greenish ordark green when dry, rachises adaxially with hairs, costae, veins,and abaxial surfaces of intercostal areas all glabrous. Sori orbicular,6–9 pairs per segment, attached at bases of veinlets and inrows close to costules, indusia scaly and small. Sporangia eachwith acicular hairs near top.● Forests in ravines; 700–1500 m. Chongqing (Nanchuan).6. Glaphyropteridopsis splendens Ching, Acta Phytotax. Sin.8: 322. 1963.大叶方杆蕨 da ye fang gan juePlants ca. 1.2 m tall. Rhizomes strong, decumbent, subwoodyand glabrous. Fronds subapproximate; stipes ca. 50 cm,deeply grooved, glabrous, stramineous; laminae narrowly oblong,ca. 80 × 28 cm, not tapering to bases, pinnate-pinnatifidnearly to costae, acuminate and pinnatifid at apices; pinnaeca. 30 pairs or more, opposite or subopposite, sessile, proximalseveral pairs not shortened; middle pinnae linear-lanceolate, ca.21 × 2.5 cm, (ca. 3.5 cm wide at bases), bases symmetrical,acroscopic sides truncate and close to rachises, basiscopic sidesrounded, pinnatifid nearly to costae, apices long acuminate;segments 35–40 pairs, spreading, lanceolate and slightly falcate,ca. 1.2 × 0.4 cm (proximal pair 1.6–2 cm), gradually broadenedto bases and gradually tapering and pointed at apices and entire,separated by obtriangular sinuses. Veins visible, ca. 14 pairs persegment, simple, proximal pair arising from base of costulesand reaching margins above sinuses. Laminae thinly herbaceous,green when dry, pinnae adaxially with short setae alongcostules, abaxially with dense grayish white acicular hairs alongrachises, costae, and veins; rachises stramineous, rectangular.Sori orbicular, 10–12 pairs per segment, attached at bases ofveinlets, in one row on each side of costules; indusia scalelike,small, hairy. Sporangia hairy near tops.● On rocks in forests; ca. 600 m. Sichuan (Emei Shan).7. Glaphyropteridopsis pallida Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 325. 1999.灰白方杆蕨 hui bai fang gan juePlants ca. 82 cm tall. Rhizomes decumbent, woody, with 1or 2 brown broadly lanceolate thick scales. Fronds sparse; stipesca. 28 cm, stramineous, grooved adaxially, glabrous throughout;laminae oblong, ca. 56 × 22 cm, slightly tapering to bases,pinnate-pinnatifid, acuminate and pinnatifid at apices; pinnaeca. 26 pairs, sessile, opposite, distal ones alternate, spreading,close, without interval, proximal pair slightly shortened andslightly reflexed, ca. 10 cm, tapering to bases; second pair ofpinnae falcate-lanceolate, ca. 12 × 2.4 cm, bases truncate andclose to rachises, pinnatifid nearly to costae, apices acuminate;segments ca. 28 pairs, spreading, separated by narrow interval,falcate-lanceolate, ca. 1.3 cm (basal acroscopic one longer), ca.0.3 cm wide, entire, pointed at apices. Veins fine, not evident,veinlets 10–12 pairs per segment, proximal pair reaching marginsabove sinuses. Laminae papery when dry, greenish, adaxiallywith dense setae along grooves of rachises, veins and intercostalareas with sparse hairs, abaxially with dense long setaealong costae and veins, occasionally sparsely hairy on intercostalareas. Sori orbicular, 4 or 5 pairs per segment, attached atbases of veinlets and in one row on each side of costules; indusiasmall and hairy. Sporangia each with 1 or 2 acicular hairsnear tops.● Shaded wet places in mixed forests on limestone mountains; ca.1500 m. NE Yunnan.8. Glaphyropteridopsis jinfushanensis Ching ex Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 325. 1999.金佛山方杆蕨 jin fo shan fang gan juePlants ca. 1.5 m tall. Rhizomes decumbent. Fronds ap-

THELYPTERIDACEAE 359proximate; stipes ca. 54 cm, ca. 1 cm in diam. at bases, darkbrown, distally grayish brown and glabrous; laminae broadlylanceolate, ca. 100 × 40 cm, pinnate-pinnatifid, pinnatifid atapices; free lateral pinnae usually ca. 32 pairs, spreading, sessile,proximal 5 pinnae abruptly reduced and hastate; middlepinnae narrowly lanceolate, ca. 25 × 3 cm, rounded-cuneate atbases, pinnatifid nearly to costae, acuminate at apices; segmentsmore than 40 pairs, lanceolate, ca. 1.5 × 0.4 cm, entire, acute atapices. Veins evident, costae raised on both sides, veinlets 10 or11 pairs per segment, proximal pair arising from base of costules,acroscopic veinlet reaching bottom of sinuses, basiscopicone reaching margins above sinuses. Laminae papery when dry,deep green, glabrous on both surfaces of intercostal areas, abaxiallywith long hairs along apices of costae, costules, and veinletswith sparse appressed short setae, adaxially with densestrong setae along rachises, costae, and veins. Sori orbicular,attached above middle part of veinlets and close to margins; indusialarge, leathery, brown, glabrous, persistent.● Mixed forests by streams; ca. 800 m. Chongqing (Jinfo Shan).9. Glaphyropteridopsis glabrata Ching & W. M. Chu ex Y. X.Lin, Fl. Reipubl. Popularis Sin. 4(1): 325. 1999.光滑方杆蕨 guang hua fang gan juePlants ca. 85 m tall. Rhizomes creeping, glabrous. Frondssparse; stipes ca. 35 cm, bases dark brown, distally stramineous,grooved adaxially, glabrous throughout; laminae oblong, ca. 50× 20 cm, slightly tapering to bases, pinnate-pinnatifid, acuminateand pinnatifid at apices; pinnae ca. 30 pairs, opposite,sessile, proximal 2 pairs slightly shortened, 7–10 cm, taperingto bases, third pair of pinnae linear-lanceolate, ca. 12 × 1.5 cm,bases widest, truncate, pinnatifid nearly to costae, apicesacuminate; segments ca. 35 pairs, spreading, proximal pairlongest (9–11 mm), distal ones triangular-lanceolate, ca. 7 ×3 mm, entire and pointed at apices. Veins evident, veinlets 9 or10 pairs per segment, proximal pair reaching margins abovesinuses. Laminae herbaceous, green when dry, glabrous exceptfor adaxially along grooves of rachises. Sori orbicular, attachedat bases of veinlets and in one row on each side of costules;indusia bearing acicular hairs. Sporangia each with 1–3 acicularhairs near top.● Mixed forest margins in valleys; 1500–1800 m. NE Yunnan.10. Glaphyropteridopsis villosa Ching & W. M. Chu ex Y. X.Lin, Fl. Reipubl. Popularis Sin. 4(1): 325. 1999.柔毛方杆蕨 rou mao fang gan juePlants ca. 65 cm tall. Rhizomes not seen. Stipes ca. 34 cm,stramineous, grooved adaxially, with sparse brown scales, glabrous;laminae broadly lanceolate, ca. 32 × 20 cm, not taperingto bases, pinnate-pinnatifid, acuminate at apices; pinnae24 pairs, sessile, pinnae below middle opposite, spreading,distal ones alternate, obliquely spreading, proximal 2 pairs notshortened, reflexed, of similar size as distal ones, but cuneate atbases; distal pinnae lanceolate, ca. 10 × 1.6 cm, bases widest,rounded-cuneate, pinnatifid nearly to costae, apices acuminate;segments ca. 25 pairs, falcate-lanceolate, ca. 7 mm (basalacroscopic one ca. 1 cm), ca. 2.5 mm wide, pointed at apices.Veins not evident, veinlets ca. 9 pairs per segment, basalacroscopic veinlet reaching bottom of sinuses, basiscopic onereaching margins above sinuses. Laminae papery, grayish greenwhen dry, abaxially with dense grayish white acicular hairsalong costae, veins, and intercostal areas, adaxially denselyhairy along costae, glabrous on intercostal areas. Sori orbicular,5–7 pairs per segment, attached near bases of veinlets and inone row on each side of costules; indusia bearing long hairs.Sporangia glabrous.● In shade and wet limestone crevices; ca. 700 m. Yunnan.11. Glaphyropteridopsis rufostraminea (Christ) Ching, ActaPhytotax. Sin. 8: 321. 1963.粉红方杆蕨 fen hong fang gan jueAspidium rufostramineum Christ, Bull. Soc. Bot. France52(Mém. 1): 36. 1905; Dryopteris rufostraminea (Christ) C.Christensen; Thelypteris rufostraminea (Christ) Ching.Plants 50–100 cm tall. Rhizomes creeping, glabrous.Fronds approximate; stipes 20–40 cm, stramineous, glabrousand usually reddish; laminae oblong-lanceolate, 40–50 × 18–25cm, not tapering to bases, pinnate-pinnatifid nearly to costae,acuminate and pinnatifid at apices; pinnae 20–28 pairs, oppositeor distally subalternate, sessile, spreading or slightly obliquelybent distally, linear-lanceolate, long acuminate at apices; proximal1 or 2 pairs slightly shortened and reflexed, bases tapering;middle pinnae 10–16 × 1.2–2 cm, wider at bases and subtruncate,pinnatifid nearly to costae; segments 30–35 pairs,spreading, linear-falcate, 6–10 × 2–2.5 mm; proximal pairlonger, entire, obtuse-pointed or acute at apices. Veins evident,veinlets 8–10 pairs per segment, proximal pair arising frombase of costules and reaching margins above sinuses. Laminaepapery, when dry yellow-green, abaxially with dense long acicularhairs along rachises, costae, veins, and intercostal areas,adaxially only with short setae. Sori orbicular, 3–5 pairs persegment, attached at bases of veinlets and in one row on eachside of costules; indusia bearing acicular hairs. Sporangiabearing acicular hairs.● Forest margins, roadsides; 1300–1500 m. Chongqing, Guizhou,Hubei, Sichuan, Yunnan.12. PSEUDOCYCLOSORUS Ching, Acta Phytotax. Sin. 8: 322. 1963.假毛蕨属 jia mao jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiTrigonospora Holttum.Plants medium-sized. Rhizomes creeping, decumbent or erect, bases with sparse lanceolate brown scales. Fronds remote,

360THELYPTERIDACEAEapproximate or clustered; stipes usually with sparse short hairs, stramineous; laminae pinnate-pinnatifid, proximal pinnae graduallyshortened into auricle, hastate or abruptly contracted into tubercule, pinnae attached to rachises and at base abaxially each usuallywith a dark brown tuberculate aerophore. Veinlets free, costules raised on both sides, proximal pair on adjacent segmentssometimes reaching cartilaginous sinuses, rarely connivent, but usually acroscopic veinlet reaching sinus and basiscopic onereaching margin above sinus. Laminae usually deep green when dry, usually papery, sometimes herbaceous or leathery, laminaeadaxially with dense appressed setae along costal grooves, sparsely setaceous also along veins, abaxially mostly with acicular hairson intercostal areas, rarely glabrous. Sori orbicular, usually attached at middle of veinlets, sometimes at distal or proximal parts;indusia orbicular-reniform, thick, usually brown, persistent, with fine hairs or glabrous, sometimes glandular. Spores normallymonolete, rarely trilete, exospore echinate, perispore cristate. x = 35, 36.About 50 species: tropical and subtropical regions; 38 species (31 endemic) in China.Pseudocyclosorus tibeticus Ching & Y. X. Lin (Acta Phytotax. Sin. 22: 201. 1984) was described from Xizang (Mêdog) but is excluded from thepresent treatment, pending further studies. It is similar to P. tylodes.1a. Proximal pinnae not gradually reduced, not tuberculate.2a. Spores monolete.3a. Laminae abaxially glabrous on intercostal areas ............................................................................................ 35. P. latilobus3b. Laminae abaxially with fine and long acicular hairs on intercostal areas ........................................... 36. P. guangxianensis2b. Spores trilete.4a. Proximal pair of veinlets on segments reaching bottom of sinus ..................................................................... 37. P. ciliatus4b. Proximal acroscopic veinlet on segment reaching bottom of sinus, others reaching marginabove sinus ................................................................................................................................................. 38. P. caudipinnus1b. Proximal pinnae abruptly reduced into aerophores or auricles or hastate.5a. Proximal pinnae abruptly reduced to brown aerophores.6a. Basal acroscopic veinlet on segments of middle pinnae reaching bottom of sinus, basiscopic onereaching margin above sinus.7a. Plants over 100 cm tall; laminae abaxially with acicular fine hairs on intercostal areas;indusia hairy ......................................................................................................................................... 4. P. xinpingensis7b. Plants only 25–58 cm tall; laminae abaxially lacking hairs on intercostal areas; indusia glabrous .......... 5. P. torrentis6b. Proximal pair of veinlets on segments of middle pinnae reaching bottom of sinus.8a. Sori attached on distal parts of veinlets and close to margins ................................................................. 3. P. cavaleriei8b. Sori attached on proximal parts of veinlets and close to costules.9a. Plants small; lateral pinnae ca. 13 × 1–1.4 cm, segments ca. 2 mm wide ............................................. 1. P. tylodes9b. Plants large; lateral pinnae length and width over 15 cm and 2 cm respectively, segmentsca. 4 mm wide ................................................................................................................................. 2. P. tuberculifer5b. Proximal pinnae gradually reduced into auricles or hastate.10a. Rachises, costae, and veins with only fine hairs abaxially, at most with 1 or 2 acicular hairs atapices.11a. Pinnae obliquely spreading.12a. Plants ca. 1.2 m tall; fronds ca. 100 × 30 cm; middle pinnae ca. 25 × 3 cm;segments over 35 pairs, more than 1.2 cm ................................................................................ 33. P. obliquus12b. Plants 90–110 cm tall; fronds 70–85 × ca. 20 cm; middle pinnae 10–15 × 1.2–2 cm;segments fewer than 30 pairs, length less than 9 mm ....................................................... 34. P. subochthodes11b. Pinnae spreading.13a. Indusia with fine hairs .................................................................................................. 32. P. damingshanensis13b. Indusia glabrous.14a. Sori attached on middle of veinlets ........................................................................................... 29. P. tsoi14b. Sori attached above middle of veinlets.15a. Veins simple, not forked and glandular abaxially .................................................. 30. P. emeiensis15b. Veins forked, without glands abaxially ...................................................... 31. P. furcatovenulosus10b. Rachises ± with long acicular hairs abaxially, costae and veins usually with acicular hairsabaxially, sometimes glabrous.16a. Indusia glabrous.17a. Pinnae linear and oblique distally.18a. Pinnae extremely oblique distally, basal acroscopic segments clearly elongated;indusia glandular ............................................................................................................ 19. P. falcilobus18b. Pinnae slightly oblique distally, basal acroscopic segments not clearly elongated;indusia without glands ................................................................................................. 20. P. lushanensis

THELYPTERIDACEAE 36117b. Pinnae not linear, spreading, at least proximal pinnae spreading.19a. Costae and veins not glandular abaxially.20a. Laminae abaxially with fine hairs on intercostal areas ....................................... 26. P. stramineus20b. Laminae abaxially glabrous on intercostal areas.21a. Basal segments on proximal pinnae entire along margins ........................... 27. P. duclouxii21b. Basal segments on proximal several pairs of pinnae lobed alongmargins ....................................................................................................... 28. P. jijiangensis19b. Costae and veins glandular abaxially.22a. Laminae abaxially hairy on intercostal areas.23a. Plants to 140 cm tall; laminae ca. 100 cm, pinnae ca. 17 × 2–2.5 cm;segments separated from each other by broad intervals, abaxiallywith sparse short hairs along costae and veins ................................... 24. P. qingchengensis23b. Plants ca. 90 cm tall; laminae ca. 50 cm; pinnae 9–13 × ca. 1.5 cm;segments separated from each other by narrow intervals, abaxiallywith mixed sparse setae and fine hairs along costae and veins ............. 25. P. angustipinnus22b. Laminae abaxially glabrous on intercostal areas.24a. Sori attached above middle of veinlets and close to margins .............. 23. P. submarginalis24b. Sori attached on middle of veinlets.25a. Pinnae lanceolate, basal acroscopic veinlet reaching bottomof sinus, basiscopic one reaching margin above sinus ........................ 21. P. esquirolii25b. Pinnae linear-lanceolate, basal pair of veinlets all reachingbottom of sinus ....................................................................................... 22. P. linearis16b. Indusia with hairs.26a. Pinnae extremely oblique distally, basal basiscopic segments clearly contracted or notcontracted.27a. Pinna bases contracted basiscopically.28a. Plants 40–110 cm tall; pinnae 1.4–1.5 cm wide in middle, costae slightlygrooved adaxially and with appressed acicular hairs, not shiny ....................... 6. P. subfalcilobus28b. Plants only 25–40 cm tall; pinnae 9–10 mm wide in middle, costaerounded and raised adaxially, subglabrous and shiny ................................. 7. P. pseudofalcilobus27b. Pinna bases not contracted.29a. Rachises, costae, and veins with mixed sparse long acicular hairs andshort fine hairs abaxially, and glabrous on intercostal areas abaxially ................ 8. P. dehuaensis29b. Rachises, costae, and veins with dense long acicular hairs abaxiallyonly and hairy on intercostal area ...................................................................... 9. P. guangxiensis26b. Pinnae spreading or slightly obliquely spreading, proximal pair of segments oracroscopic one clearly elongated.30a. Sori attached below middle of veinlets and close to costules.31a. Fronds dimorphic; pinna bases with segments not clearly elongate,acroscopic veinlet of pinna basal pair reaching bottom of sinus ................. 10. P. shuangbaiensis31b. Fronds monomorphic; pinna bases with segments clearly elongate,proximal veinlets of segments all reaching bottom of sinuses .................. 11. P. dulongjiangensis30b. Sori attached on middle of veinlets.32a. Laminae abaxially with fine setae on intercostal areas ............................................... 12. P. canus32b. Laminae abaxially lacking fine setae on intercostal areas.33a. Indusia with only few fine hairs.34a. Proximal 7–9 pairs of pinnae abruptly reduced into linearauricles, proximal ones tuberculate; laminae abaxially glabrouson intercostal areas ..................................................................... 18. P. paraochthodes34b. Proximal 3 pairs of pinnae abruptly reduced and hastate.35a. Pinnae (particularly acroscopic ones) obviouslybroadened at their bases, margins pinnatifid; segments11–14 × 5–6 mm; laminae herbaceous; sori attachedabove middle of veinlets ................................................... 16. P. gongshanensis35b. Pinnae narrowed at their bases; segments ca. 9 × 3 mm;laminae papery; sori attached on middle of veinlets .......... 17. P. pseudorepens33b. Indusia with dense hairs.36a. Proximal pinnae mostly reduced and hastate or auriculate ................ 15. P. zayuensis36b. Proximal 2 or 3 pairs of pinnae reduced and hastate.

362THELYPTERIDACEAE37a. Lateral pinnae to 30 pairs, spreading ..................................... 13. P. fugongensis37b. Lateral pinnae ca. 17 pairs, proximal pair slightlyreflexed, others ascending distally .......................................... 14. P. lushuiensis1. Pseudocyclosorus tylodes (Kunze) Ching, Acta Phytotax.Sin. 8: 323. 1963.假毛蕨 jia mao jueAspidium tylodes Kunze, Linnaea 24: 281. 1851[“xylodes”]; Cyclosorus tylodes (Kunze) Panigrahi; Dryopterisochthodes (Kunze) C. Christensen var. tylodes (Kunze) C.Christensen; D. tylodes (Kunze) Christ; Lastrea ochthodes(Kunze) T. Moore var. tylodes (Kunze) Beddome; L. tylodes(Kunze) T. Moore [“xylodes”]; Nephrodium prolixum (Willdenow)Desvaux var. tylodes (Kunze) Baker; N. tylodes (Kunze)Baker; Thelypteris tylodes (Kunze) Ching.Plants to 1.2 m tall. Rhizomes erect, apices and bases ofstipes with sparse brown lanceolate scales. Fronds clustered;stipes 25–40 cm, grayish brown at bases, distally deep stramineousand glabrous; laminae oblong-lanceolate, 45–80 × ca. 24cm, slightly tapering to bases, pinnate-pinnatifid, acuminatepinnatifidat apices; proximal 5–10 pairs of pinnae abruptly reducedinto tuberculate aerophores; normal pinnae ca. 34 pairs,alternate, oblique distally, sessile; middle pinnae lanceolate, ca.13 × 1.2–1.4 cm, long acuminate at apices, proximal pinnaetapering to bases, pinnae above middle slightly tapering tobases and broadly cuneate, pinnatifid; segments 40–45 pairs,obliquely spreading, ca. 5 × 1.5–2 mm, entire, pointed orobtuse-pointed at apices. Veinlets evident on both sides, 9 or 10pairs per segment, proximal pair arising from base of costulesand all reaching sinuses. Laminae firmly papery when dry,brownish, costae with acicular hairs, elsewhere glabrous. Soriorbicular, attached below middle of veinlets and close to costules;indusia orbicular-reniform, thick, glabrous, persistent.Forests by streams or on rocks; 800–4300 m. Guangdong, Guangxi,Guizhou, Hainan, Sichuan, E and S Taiwan, E and S Xizang, Yunnan[India, Myanmar, N Philippines, Sri Lanka, N Thailand, Vietnam].In Kunze’s protologue, the epithet “xylodes” is considered to be amisprint; it was printed as “tylodes” on pages 244 and 283. For furtherdiscussion on the spelling, see Holttum and Grimes (Kew Bull. 34: 504.1980).The name Nephrodium prolixum (Willdenow) Desvaux has beenmisapplied to material of this taxon by Dunn and Tutcher (Bull. Misc.Inform. Kew, Addit. Ser. 10: 348. 1920).2. Pseudocyclosorus tuberculifer (C. Christensen) Ching, ActaPhytotax. Sin. 8: 324. 1963 [“tuberculiferus”].瘤羽假毛蕨 liu yu jia mao jueDryopteris tuberculifera C. Christensen, Contr. U.S. Natl.Herb. 2: 275. 1931; Thelypteris tuberculifera (C. Christensen)Ching.Plants more than 2 m tall. Rhizomes erect, firm, apiceswith dense brownish lanceolate scales. Fronds clustered; stipes100–130 cm, brownish, sparsely scaly, distally glabrous; laminaebroadly oblong-lanceolate, ca. 100 × 45 cm, tapering tobases, pinnate-pinnatifid, acuminate at apices; proximal to 15pairs of pinnae abruptly reduced into brown aerophores, middlenormal pinnae 28–30 pairs, proximal ones opposite or subopposite,spreading, distally alternate, obliquely spreading, sessile;middle pinnae linear-lanceolate, 20–30 × 2.2–3 cm, linearacuminateat apices; proximal pinnae slightly tapering, cuneate,distal pinnae slightly broadened to bases, rounded-cuneate, pinnatifidand reaching broad wings on both sides; segments 35–45 pairs, oblique distally, slightly bent, subfalcate, 8–14 × ca. 4mm, entire, acute at apices. Veins evident, raised on both sides,veinlets 11–15 pairs per segment, proximal pair reaching bottomof sinuses. Laminae papery when dry and brownish green,rachises and costae with sparse acicular hairs at apices abaxially,costae with dense appressed setae along grooves abaxially,elsewhere glabrous. Sori orbicular, attached below middleof veinlets and close to costules; indusia orbicular-reniform,thick, brown, glabrous, persistent.Gravelly soil by streams; 600–1900 m. Guangdong, Guangxi,Yunnan [N India (Sikkim)].3. Pseudocyclosorus cavaleriei (H. Léveillé) Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 143. 1999.青岩假毛蕨 qing yan jia mao jueChristella cavaleriei H. Léveillé, Fl. Kouy-Tchéou, 474.1915.Rachises subrectangular; middle pinnae spreading, alternate,ca. 18 × 3 cm, pinnate-pinnatifid, pinnatifid-acuminate atapices; segments ca. 35 pairs, separated by broad sinuses, lanceolate,ca. 1 × 0.4 cm, acute at apices; proximal pair clearlyelongated, to 1.5 cm, proximal pair on segments reaching bottomof sinuses. Sori orbicular, attached above middle of veinletsand close to margins, ca. 10 pairs per segment.● Guizhou (Guiyang).The present authors have not seen a complete specimen ofPseudocyclosorus cavaleriei. The type of P. cavaleriei was collected byCavalerie on 5 August 1903, from Guiyang Ging-yan (Tsin Gai), Guizhouprovince, no. 1244 (E). It has the middle part of the lamina andseven pairs of pinnae. Ching examined the specimen and identified it asDryopteris tylodes in 1931. The species was included within Thelypterisesquirolii (Christ) Ching by Holttum in 1936 and later in P. esquirolii(species no. 21 in this account) (Holttum & Grimes, Kew Bull. 34: 514–515. 1980).The subrectangular rachises, opposite pinnae, and proximal pair ofveinlets reaching bottom of sinuses suggest that the species should notbe included in Pseudocyclosorus esquirolii, but is instead closer to P.tylodes. However, the sori attached on distal parts of veinlets and closeto margins are different from those of specimens of P. tylodes. So, it ishere treated as an independent species, and we hope that more specimensof this species will be collected for study.4. Pseudocyclosorus xinpingensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 326. 1999.新平假毛蕨 xin ping jia mao juePlants to 110 cm tall. Rhizomes ascending, brown. Fronds

THELYPTERIDACEAE 363clustered; stipes ca. 54 cm, deep stramineous, with fine hairs;lamina lanceolate, ca. 57 × 22 cm, slightly tapering to bases,pinnate-pinnatifid, acuminate-pinnatifid at apices; lateral pinnaeca. 25 pairs, alternate, spreading, shortly stalked, proximal 3pairs of pinnae abruptly reduced into black aerophores; middlepinnae narrowly lanceolate, ca. 14 × 1.4 cm, slightly tapering tobases, acuminate at apices; lowest several pinnae clearlyshortened, pinnatifid; segments ca. 24 pairs, ca. 9 × 5 mm, entire,acute at apices. Veins clearly raised on both sides, veinlets9 pairs per segment, proximal pair arising from base of costules,basiscopic veinlet reaching margin of sinus, acroscopic onereaching bottom of sinus. Laminae green when dry, papery, rachiseswith sparse short setae abaxially, adaxially with denseshort setae, costae and veins with sparse short setae abaxially,costae with sparse appressed short hairs adaxially, veinlets with1 or 2 setae, intercostal areas acicular hairy abaxially of andglabrous adaxially, setaceous along margins. Sori attached onmiddle of veinlets, 6 or 7 pairs per segment; indusia large,brown, membranous, hairy, persistent.● Shade of mossy forests on mountain summits; ca. 2300 m. Yunnan(Xinping).5. Pseudocyclosorus torrentis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 326. 1999.急梳假毛蕨 ji shu jia mao juePlants 25–58 cm tall. Rhizomes erect. Fronds clustered;stipes 13–23 cm, stramineous, bases with brown scales, distallywith acicular hairs abaxially. Laminae oblong-lanceolate, 10–15× 7–10 cm, bases slightly tapering, proximal 2 or 3 pairsabruptly reduced to auricles, or tuberculate, pinnate-pinnatifid,apices pinnatifid-acuminate; lateral normal pinnae ca. 10 pairs;middle pinnae lanceolate, 4–6 × ca. 0.5 cm, bases broadeneddue to one pair of segments elongated, pinnatifid, pinnatifidacuminateat apices; segments ca. 11 pairs, lanceolate, ca. 4 ×2.5 mm, entire, acute at apices. Laminae dark brown when dry,papery; abaxial sides of rachises with sparse setae on distal half,adaxial sides densely setaceous along grooves, abaxial sides ofcostae and veins glabrous; adaxial sides of costules with appressedshort setae, glabrous on both veinlets and intercostalareas, with few short setae along margins. Veins evident, raisedon both sides, veinlets 4 or 5 pairs per segment, basal basiscopicveinlet arising from above base of costules and reachingmargins above sinuses, acroscopic veinlet arising from base ofcostules and reaching bottom of sinuses. Sori orbicular andattached on proximal parts of veinlets and close to main veins,1–3 pairs per segment; indusia brown, thick, glabrous, persistent.● 800–1500 m. Yunnan (on route from Yingjiang to Ruili).6. Pseudocyclosorus subfalcilobus Ching ex K. H. Shing, ActaPhytotax. Sin. 31: 571. 1993.光脉假毛蕨 guang mai jia mao juePlants 40–55(–110) cm tall. Rhizomes short and erect ordecumbent, dark brown, sparsely scaly. Fronds clustered; stipes10–20 cm, bases with sparse lanceolate brown scales and denseacicular hairs, stramineous; laminae narrowly ovate, 15–30 ×5–13 cm, pinnatifid; proximal 2 or 3 pairs of pinnae abruptlyreduced and hastate, normal lateral pinnae extremely obliquedistally, narrowly lanceolate, 11–13 pairs, alternate, shortlystalked; middle pinnae 5–8 × 1–1.5 cm, bases cuneate, notsymmetrical, pinnatifid nearly to narrow wings on both sides ofcostae, pinnatifid and long caudate at apices; segments 11–15pairs, ligulate, 5–8 × 2–3 mm, entire, shortly pointed at apices;basal basiscopic segments much smaller than acroscopic ones.Costules raised adaxially, visible abaxially, veinlets not evident,5–8 pairs per segment, basal acroscopic veinlet reaching bottomof sinus, basiscopic one reaching margin above sinus. Laminaepapery when dry, dark brown, both surfaces with dense longacicular hairs along rachises and costae, abaxial sides of costulesand margins acicular hairy, adaxial sides of costules appressedshortly hairy. Sori orbicular, attached below middle ofveinlets and close to costules, 6–8 pairs per segment; indusiaorbicular-reniform, thickly membranous, brown, shortly hairy,persistent.● Shaded wet rocks or sandy beaches by margins of evergreenbroad-leaved forests on slopes; 1300–1500 m. NW Yunnan (Gongshan:Dulongjiang Valley).7. Pseudocyclosorus pseudofalcilobus W. M. Chu, Acta Bot.Yunnan., Suppl. 5: 46. 1992.似镰羽假毛蕨 si lian yu jia mao juePlants 25–40 cm tall. Rhizomes creeping, firm. Frondsclustered; stipes 10–15 cm, stramineous, glabrous; laminae narrowlylanceolate, 20–35 × ca. 8 cm, proximal 1 or 2 pairs ofpinnae abruptly reduced into auricles, pinnate-pinnatifid, pinnatifid-acuminateat apices; normal free pinnae 10–15 pairs,subopposite, oblique distally, sessile; middle pinnae narrowlylanceolate, ca. 5 × 1 cm, tapering to bases, 1-pinnatifid, pinnatifidand acuminate-caudate at apices; segments ca. 12 pairs,ligulate, ca. 5 × 2 mm, acute at apices. Veins evident on bothsides, veinlets 6–8 pairs per segment, basal acroscopic veinletreaching bottom of sinus, basiscopic one reaching margin abovesinus. Laminae somewhat leathery, deep green when dry; rachisesand veins all with setae and short hairs abaxially, adaxialsides of costae subglabrous with dense appressed setae alonggrooves, elsewhere glabrous. Sori attached below middle ofveinlets and close to costules, ca. 8 pairs per segment; indusiaorbicular-reniform, brown, papery, persistent, densely finelyhairy.● Sandy beaches at margins of evergreen broad-leaved forests onslopes; ca. 1300 m. NW Yunnan (Gongshan: Dulongjiang).8. Pseudocyclosorus dehuaensis Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 326. 1999.德化假毛蕨 de hua jia mao juePlants 85–95 cm tall. Rhizomes decumbent. Fronds few;stipes 20–25 cm, glabrous; laminae lanceolate, 60–70 × ca. 15cm, proximal 4–10 pairs reduced and hastate; normal pinnae20–25 pairs, pinnate-pinnatifid; middle pinnae narrowly lanceolate,ca. 10 × 1.5 cm, oblique distally, sessile, alternate, caudateacuminateat apices; proximal several pairs slightly narrowed atbases, pinnae above middle slightly broadened at bases androunded-cuneate, 1-pinnatifid; segments 20–25 pairs, subrectangular,ca. 6 × 3 mm, entire, obtuse-pointed at apices. Costae

364THELYPTERIDACEAEraised on both sides, veinlets not evident, 8 or 9 pairs persegment and all arising above base of costules, basal acroscopicvein reaching bottom of sinus, basiscopic vein reaching marginabove sinus. Laminae dark green when dry, papery, with denseappressed acicular hairs along rachises, abaxial sides of costaeand veins with mixed spreading acicular hairs and short setae,adaxial sides of costae densely setaceous along grooves, veinletssparsely setaceous, glabrous on intercostal areas of bothsides. Sori orbicular, attached on middle of veinlets; indusiaorbicular-reniform, brown, thick, with few fine long hairs, persistent.● Streamsides in forests; ca. 700 m. Fujian (Dehua).9. Pseudocyclosorus guangxiensis Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 327. 1999.广西假毛蕨 guang xi jia mao juePlants ca. 60 cm tall. Rhizomes not seen. Stipes ca. 8.5cm, deep stramineous, bases sparsely scaly, distally denselyacicular hairy; laminae narrowly lanceolate, ca. 50 × 10 cm,proximal 4 or 5 pairs of pinnae gradually reduced and hastate;middle normal pinnae ca. 25 pairs, alternate, extremely obliquedistally, sessile, pinnate-pinnatifid; middle pinnae narrowlylanceolate, ca. 7 × 1 cm, pinnatifid nearly to costae; segmentsca. 18 pairs, acroscopic one of basal pair of pinnae slightlyelongated, ca. 6 mm, others ca. 4 × 2 mm, bent, entire, acuminateat apices. Costules raised on both sides, veinlets not evident,5 or 6 pairs per segment. Laminae dark green when dry,papery; rachises, costae, and veinlets all with dense long acicularhairs abaxially, adaxially densely setaceous along costalgrooves, costules occasionally with 1 or 2 setae, finely hairy onintercostal areas abaxially. Sori orbicular, attached on middle ofveinlets; indusia orbicular-reniform, papery, brown, persistent,acicular hairy.● On shaded rocks by streamsides; ca. 300 m. Guangxi (Fusui).10. Pseudocyclosorus shuangbaiensis Ching ex Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 327. 1999.双柏假毛蕨 shuang bai jia mao juePlants ca. 90 cm tall. Rhizomes creeping. Fronds remote;dimorphic; fertile stipes to 43 cm, stramineous, sparsely brownscaly and glabrous; laminae oblong-lanceolate, ca. 50 × 18 cm,proximal 1 or 2 pairs of pinnae slightly shortened, pinnatepinnatifid,acuminate at apices; lateral pinnae ca. 20 pairs,spreading, alternate, subsessile; middle pinnae narrowly lanceolate,ca. 9 × 1.4 cm, not narrowed to bases, pinnatifid, pinnatifidand long caudate-acuminate at apices; segments ca. 25 pairs,lanceolate, slightly bent, ca. 4 × 3 mm, entire, obtuse-pointed atapices; sterile fronds smaller, proximal 4 pairs of pinnaeabruptly reduced into auricles. Veins evident on both sides, 7 or8 pairs of veinlets per segment, proximal pair arising fromabove base of costules, acroscopic veinlet reaching bottom ofsinus, basiscopic one reaching margin above sinus. Laminaedark green when dry, papery; rachises and costae with denseacicular hairs on both sides, acicular hairy on veins and abaxialsurface of intercostal areas, adaxially occasionally with 1 or 2acicular hairs and glabrous on intercostal areas, shortly setaceousalong margins. Sori orbicular and attached below middleof veinlets and close to costules, 7 or 8 pairs per segment;indusia brown, membranous and hairy, persistent.● Evergreen broad-leaved forests; ca. 2100 m. Yunnan (Shuangbai).11. Pseudocyclosorus dulongjiangensis W. M. Chu, Acta Bot.Yunnan., Suppl. 5: 44. 1992.独龙江假毛蕨 du long jiang jia mao jueRhizomes creeping, dark brown and firm. Stipes ca. 8 cm,stramineous, sparsely scaly proximally. Laminae lanceolate, ca.110 cm, pinnatifid-acuminate at apices; proximal 12 pairs reducedand hastate; middle normal pinnae ca. 23 pairs, spreading,alternate, sessile, ca. 15 × 2.6 cm, pinnatifid, acuminate atapices; segments more than 30 pairs, subrectangular, proximalpair of pinnae elongate, spreading, others slightly bent distally;proximal segments 1.2–1.5 × ca. 0.4 cm, entire, obtuse-pointedat apices. Costules raised on both sides, 8 or 9 pairs per segment,proximal pair reaching bottom of sinus. Laminae greenwhen dry, thinly papery, abaxially with acicular hairs alongrachises, costae, veins, and intercostal areas, adaxially withappressed setae along costal grooves, costules and veinletssparsely hairy, setaceous along margins. Sori orbicular, attachedbelow middle of veinlets and close to costules; indusia orbicular-reniform,persistent, glandular and finely hairy.● Margins of evergreen broad-leaved forests on mountain slopes;1200–1500 m. NW Yunnan (Gongshan: Dulongjiang).12. Pseudocyclosorus canus (Baker) Holttum & Jeff W.Grimes, Kew Bull. 34: 509. 1980.长根假毛蕨 chang gen jia mao jueNephrodium canum Baker in Hooker & Baker, Syn. Fil.267. 1867; Cyclosorus canus (Baker) S. Lindsay; Dryopteriscana (Baker) Kuntze; D. repens C. Christensen; N. repens C.Hope (1899), not Baillon (1874); Pseudocyclosorus medogensisChing & S. K. Wu; P. repens (C. Christensen) Ching; Thelypteriscana (Baker) Ching; T. repens (C. Christensen) Ching.Plants more than 1 m tall. Rhizomes suberect and subglabrous.Fronds clustered; stipes 15–20 cm, stramineous orbrownish, with sparse brown scales and dense setae; laminaeoblong-lanceolate, 40–80 × 13–25 cm, pinnatifid-acuminate atapices; proximal pairs of pinnae reduced and hastate or auriculate;normal pinnae 20–25 pairs, alternate, sessile, spreading,pinnate-pinnatifid; middle pinnae narrowly lanceolate, pinnatifidand long acuminate; proximal pair of segments slightlyelongate, spreading, 7–14 × 1.6–2.4 cm, pinnatifid nearly tocostae; segments 20–28 pairs, subligulate, slightly bent, 7–12 ×2.5–4 mm, entire, acute. Veins raised on both sides, visible, 8–12 pairs of veinlets per segment, proximal pair arising frombases or above base of costules, acroscopic vein reaching bottomof sinus, basiscopic one reaching margin above sinus. Laminaethinly papery or herbaceous, dark green or greenish whendry; abaxial sides of rachises, costae, and veins all with acicularhairs, adaxially with dense appressed setae along grooves,sparsely setaceous along veins, shortly hairy on intercostalareas on both surfaces. Sori orbicular, attached on middle veinlets;indusia orbicular-reniform, brown, persistent, hairy.

THELYPTERIDACEAE 365Broad-leaved forests on mountain slopes; 900–2800 m. SE Xizang(Mêdog) [Bhutan, N India, Kashmir, Nepal].“Pseudocyclosorus pectinatus” (Ching ex K. H. Shing, Vasc. Pl.Hengduan Mount. 1: 98. 1993 [“pectinata”]) is conspecific with P. canusbut was not validly published because no Latin description or diagnosis,or reference to such, was provided (Melbourne Code, Art. 39.1).13. Pseudocyclosorus fugongensis Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 327. 1999.福贡假毛蕨 fu gong jia mao juePlants to 1.3 m tall. Rhizomes not seen. Stipes ca. 50 cm,dark brown, deep stramineous distally and glabrous; laminaebroadly lanceolate, ca. 85 × 20 cm, abruptly tapering to bases,pinnatifid-acuminate at apices; proximal 2 pairs of pinnae reducedand hastate; middle normal pinnae ca. 30 pairs, alternate,spreading, sessile, pinnate-pinnatifid; middle pinnae lanceolate,12–15 × ca. 1.5 cm, bases not narrowed, rounded-truncate, pinnatifidnearly to costae, apices long acuminate; segments ca. 25pairs, lanceolate, ca. 7 × 3 mm, entire, acute at apices. Veinsevident, costules raised on both sides, veinlets 8(or 9) pairs persegment, proximal pair arising from base of costules, acroscopicvein reaching bottom of sinus, basiscopic one reachingmargin above sinus. Laminae dark green when dry, papery,abaxially along rachises, costae, veins, and intercostal areas allwith long acicular hairs, adaxially along costal grooves withappressed setae, costae appressed shortly hairy, veinlets eachwith 1 or 2 setae, glabrous on intercostal areas. Sori orbicular,attached on middle of veinlets; indusia orbicular-reniform,thickly membranous, brown, densely shortly hairy, persistent.● Evergreen broad-leaved forests on mountain slopes; ca. 1800 m.Yunnan (Fugong).Pseudocyclosorus fugongensis differs from P. canus mainly inintercostal areas glabrous adaxially and laminae with 2 pairs of reducedpinnae proximally.14. Pseudocyclosorus lushuiensis Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 327. 1999.泸水假毛蕨 lu shui jia mao juePlants to 70 cm tall. Rhizomes erect. Fronds clustered;stipes to 32 cm, stramineous; laminae ovate-lanceolate, ca. 38 ×18 cm, pinnatifid-acuminate at apices; proximal 3 pairs ofpinnae reduced into auricles, one pair reduced and hastate;middle normal pinnae ca. 17 pairs, proximal ones subopposite,distal ones alternate, shortly stalked, except for proximal pairslightly reflexed, others ascending distally, pinnate-pinnatifid;pinnae lanceolate, bent or falcate distally; middle pinnae ca. 13× 2 cm, pinnatifid; segments ca. 26 pairs per pinna, subrectangular,bent distally, ca. 6 × 3 mm, entire, acute at apices.Costules raised on both sides, ca. 10 pairs of veinlets per segment,clearly visible. Laminae dark green when dry, papery;rachises, costae, and costules all with dense acicular hairs andpubescence abaxially, adaxial side of costae with dense appressedsetae along grooves, veinlets and along margins sparselysetaceous, costules shortly hairy adaxially, intercostal areas withdense acicular fine hairs abaxially. Sori orbicular, attached onmiddle of veinlets; indusia orbicular-reniform, brown, hairy andpersistent.● Evergreen broad-leaved forests; ca. 2200 m. Yunnan (Lushui).15. Pseudocyclosorus zayuensis Ching & S. K. Wu, Fl. Xizang.1: 168. 1983.察隅假毛蕨 cha yu jia mao juePlants 1–1.5 m tall. Rhizomes short and decumbent, subglabrous.Fronds clustered; stipes 10–30 cm, stramineous, withsparse brown scales proximally, distally shortly setaceous andeasily deciduous when brushed; laminae oblong-lanceolate, 90–120 × 20–30 cm, proximal pairs of pinnae reduced and hastateor auriculate, pinnatifid-acuminate at apices; middle normalpinnae ca. 22 pairs, spreading, alternate, linear-lanceolate, basesnot tapering, truncate, sessile or extremely shortly stalked, pinnatifidnearly to costae, apices acuminate; segments ca. 30pairs, obliquely spreading, subligulate, 8–9 × ca. 3 mm, slightlybent, entire, obtuse-pointed at apices. Veins evident, costulesevident and raised on both sides, 9 or 10 pairs of veinlets persegment, simple, proximal pair arising above base of costules,acroscopic veinlet reaching bottom of sinus, basiscopic onereaching margin above sinus. Laminae green or dark greenwhen dry, papery; abaxially with dense acicular hairs alongrachises, costae, costules, and on intercostal areas, densely setaceousalong costal grooves adaxially, hairs sparser on veins.Sori orbicular, attached on middle of veinlets, ca. 9 pairs persegment; indusia orbicular-reniform, brown, with acicular hairs,persistent.● Evergreen broad-leaved forests, streamsides; 2000–2200 m. SEXizang (Zayü), Yunnan (Gongshan).16. Pseudocyclosorus gongshanensis Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 328. 1999.贡山假毛蕨 gong shan jia mao juePlants to 1 m tall. Rhizomes not seen. Stipes ca. 23 cm,bases brownish, distally stramineous, glabrous; laminae broadlylanceolate, ca. 70 × 30 cm, proximal 3 pairs of pinnae reducedand hastate, pinnate-pinnatifid, pinnatifid-acuminate at apices;middle normal pinnae ca. 20 pairs, alternate, spreading, lanceolate,15–17 × 2.5–3 cm, bases truncate, sessile, pinnatifid,apices acuminate; segments more than 25 pairs, broadly ligulate,11–14 × 5–6 mm, rounded-obtuse at apices; proximalpair of segments (particularly acroscopic one) clearly elongated,slightly lobed along margins. Veins evident abaxially,raised on both sides, 10–12 pairs of veinlets per segment,acroscopic veinlet of basal pair reaching bottom of sinus, basiscopicone reaching margin above sinus. Laminae greenishwhen dry, herbaceous, abaxially with dense acicular hairs alongrachises, costae, veins, and intercostal areas and mixed glands;adaxially densely setaceous along costal grooves, appressedshortly hairy on costules, glabrous on intercostal areas. Sori orbicular,attached above middle of veinlets, 7 or 8 pairs per segment;indusia orbicular-reniform, membranous, brown, with 1or 2 short hairs, persistent.● Margins of evergreen broad-leaved forests on slopes; ca. 1400m. Yunnan (Gongshan).Pseudocyclosorus gongshanensis is similar to P. pseudorepens butdiffers in the proximal pair of segments on each pinna much larger thanmore distal ones, lobed along margins, and the sori attached on middleof veinlets.

366THELYPTERIDACEAE17. Pseudocyclosorus pseudorepens Ching ex K. H. Shing,Acta Phytotax. Sin. 31: 571. 1993.毛脉假毛蕨 mao mai jia mao juePlants 90–110 cm tall. Rhizomes decumbent, woody.Fronds approximate; stipes ca. 40 cm, stramineous, subglabrous;laminae lanceolate, 40–70 × 15–20 cm, pinnate-pinnatifid,pinnatifid-acuminate at apices; proximal 3 pairs of pinnaereduced and hastate; middle normal pinnae ca. 20 pairs,obliquely spreading, alternate, sessile, ca. 14 × 2 cm, slightlytapering to bases, pinnatifid nearly to costae, pinnatifid andlong caudate at apices; segments ca. 25 pairs, sublanceolate, ca.9 × 3 mm, entire, rounded-obtuse at apices. Veins visible,costules raised on both sides, 9–11 pairs of veinlets per segment,oblique distally, proximal pair arising above base of costules,acroscopic veinlet reaching bottom of sinus, basiscopicone reaching margin above sinus. Laminae dark green whendry, papery; abaxially with long acicular hairs along rachises,costae, veins, and on intercostal areas, adaxially with dense setaealong costal grooves, costules finely hairy adaxially, onveinlets and along margins with 1 or 2 setae. Sori orbicular,attached on middle of veinlets; indusia orbicular-reniform,brown, occasionally finely hairy, persistent.● On rocks by streams; 2200–2500 m. Yunnan (Gongshan).18. Pseudocyclosorus paraochthodes Ching ex K. H. Shing &J. F. Cheng, Jiangxi Sci. 8(3): 43. 1990.武宁假毛蕨 wu ning jia mao juePlants to 1.2 m tall. Rhizomes decumbent, dark brown.Fronds approximate; stipes 8–10 cm, dark brown proximally,distally stramineous, glabrous; laminae oblong-lanceolate orbroadly lanceolate, ca. 110 × 26–35 cm, abruptly narrowed atbases, pinnate-pinnatifid, pinnatifid-acuminate at apices; normallateral pinnae 28–30 pairs, extremely shortly stalked, alternate,obliquely spreading, proximal 7–9 pairs abruptly reducedinto small linear auricles, proximal one reduced, tuberculate;middle pinnae linear-lanceolate, ca. 20 cm, broadly cuneate atbases, pinnatifid nearly to costae, long acuminate-caudate atapices; segments ca. 35 pairs, oblique distally, sublanceolate, 8–10 × ca. 3 mm, entire, acute at apices. Veins evident abaxially, 9or 10 pairs of veinlets per segment, acroscopic veinlet of proximalpair arising above base of costules and reaching bottom ofsinus, basiscopic veinlet arising from base of costule andreaching margin above sinus. Laminae green when dry, papery;abaxially with sparse acicular hairs along rachises, along costaeand veins with mixed acicular setae and fine hairs; adaxiallywith dense acicular setae along grooves of rachises and costae,costules with dense appressed short hairs adaxially, veinlets andalong margins with 1 or 2 setae. Sori orbicular, attached onmiddle of veinlets, 7 or 8 pairs per segment; indusia orbicularreniform,thick, brown, finely long hairy, persistent.● Rock crevices. Jiangxi (Wuning).19. Pseudocyclosorus falcilobus (Hooker) Ching, Acta Phytotax.Sin. 8: 324. 1963.镰片假毛蕨 lian pian jia mao jueLastrea falciloba Hooker, Hooker’s J. Bot. Kew Gard.Misc. 9: 337. 1857; Aspidium falcilobum (Hooker) Hooker;Cyclosorus falcilobus (Hooker) L. J. He & X. C. Zhang;Dryopteris falciloba (Hooker) C. Christensen; Glaphyropterisfalciloba (Hooker) H. Itô; Nephrodium falcilobum (Hooker)Hooker; Thelypteris falciloba (Hooker) Ching.Plants 65–80 cm tall. Rhizomes erect, woody, apices andbases of stipes with brown lanceolate scales. Fronds clustered;stipes 6–10 cm, bases dark brown, distally stramineous, glabrous;laminae lanceolate, 60–70 × 14–18 cm, proximallyabruptly narrowed, pinnate-pinnatifid, pinnatifid-acuminate atapices; proximal 3–6 pairs of pinnae reduced into smallauricles, middle normal pinnae 36–38 pairs, very oblique distally,alternate or subopposite, sessile, linear-lanceolate, 12–13× 1–1.2 cm, cuneate at bases, pinnatifid nearly to costae, longacuminate at apices; segments 22–25 pairs, falcate-lanceolate,oblique distally, 5–7 × 2–2.5 mm, entire, acute at apices; basalacroscopic one particularly elongated to 1 cm. Veins visibleadaxially, costules raised on both sides, veinlets extremelyoblique distally, 9 or 10 pairs per segment, proximal pair arisingfrom base of costule, acroscopic veinlet reaching bottom sinus,basiscopic one reaching margin above sinus. Laminae darkgreen when dry, papery; abaxially with acicular setae along rachises,costae, and veins, glabrous on intercostal areas, adaxiallywith appressed setae along grooves of costae and veins,margins subglabrous. Sori orbicular, attached on middle of veinlets;indusia orbicular-reniform, thick, brown, glandular, persistent.On rocky soil at watersides in valleys; 300–1100 m. Fujian,Guangdong, Guangxi, Hainan, S and SW Yunnan, Zhejiang [India,Japan, Laos, Myanmar, Thailand, Vietnam].20. Pseudocyclosorus lushanensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 328. 1999.庐山假毛蕨 lu shan jia mao juePlants 40–50 cm tall. Rhizomes decumbent, with sparsescales. Fronds approximate; stipes 13–18 cm, bases sparselybrown scaly, distally glabrous, deep stramineous; laminae lanceolate,20–32 × 10–13 cm, tapering proximally, pinnate-pinnatifid,pinnatifid-acuminate at apices; proximal 2 pairs of pinnaereduced into auricles; middle normal pinnae 16–18 pairs,proximal ones subopposite, distal ones alternate, obliquelyspreading, sessile, narrowly lanceolate, 6.7–7.5 × ca. 1.2 cm,bases not narrowed, truncate, pinnatifid nearly to costae, apiceslong acuminate; segments ca. 17 pairs, obliquely spreading,lanceolate, 4–5 × ca. 7 mm, entire, obtuse-pointed at apices.Veins evident, costules raised on both sides, ca. 6 pairs of veinletsper segment. Laminae dark green when dry, papery; abaxiallywith sparse setae and fine short hairs along rachises, costae,and veins, glabrous on intercostal areas, adaxially densely setaceousalong costal grooves, hairs sparser along veins but mixedwith few glands. Sori orbicular, attached on middle of veinlets;indusia orbicular-reniform, brown, glandular and with 1 or 2fine hairs, persistent.● Streamsides in forests; ca. 600 m. Fujian, Jiangxi (Jinggangshan,Lushan).

THELYPTERIDACEAE 36721. Pseudocyclosorus esquirolii (Christ) Ching, Acta Phytotax.Sin. 8: 324. 1963.西南假毛蕨 xi nan jia mao jueDryopteris esquirolii Christ, Bull. Acad. Int. Géogr. Bot.17: 144. 1907; Cyclosorus esquirolii (Christ) C. M. Kuo; D.eberhardtii Christ; Thelypteris esquirolii (Christ) Ching.Plants to 1.5 m tall. Rhizomes creeping. Fronds remote;stipes deep stramineous, glabrous above bases. Laminaebroadly oblong-lanceolate, ca. 130 × 30 cm, tapering to bases,pinnate-pinnatifid, pinnatifid-acuminate at apices; pinnae proximal9–11 pairs alternate, gradually reduced to triangular auricles,distal pairs alternate, sessile, spreading, lanceolate, 15–20 × 2–2.3 cm, bases rounded-truncate, symmetrical, pinnatifidnearly to costae, apices long caudate-acuminate; segments 30–35 pairs, spreading, lanceolate, 9–10 × 2.5–3 mm, entire, obtuseor acute at apices; proximal pair (particularly acroscopic one)clearly elongated. Veins visible, costae raised on both sides,8–12 pairs of veinlets per segment, proximal pair arising frombase of costules, acroscopic veinlet reaching bottom of sinus,basiscopic one reaching margin above sinus. Laminae darkgreen when dry, papery, glabrous on both surfaces of intercostalareas, abaxially acicular hairy along rachises and costae, adaxiallydensely appressed-setaceous along costal grooves, with 1or 2 setae on veins and margins. Sori orbicular, attached onmiddle of veinlets, 10–12 pairs per segment; indusia orbicularreniform,thickly membranous, brown, glabrous, persistent.On rocks by streams in valleys, in bamboo by streams; 400–2100m. Fujian, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan[N India, Myanmar, Nepal, Thailand].“Christella esquirolii” (H. Léveillé, Fl. Kouy-Tchéou, 474. 1915,nom. nud.) belongs here.Thelypteris ×erubesquirolica W. C. Shieh & J. L. Tsai (J. Sci. Engin.24: 7. 1987), described from Taiwan (Nantou), appears to be a presumedhybrid between Pseudocyclosorus esquirolii and Glaphyropteridopsiserubescens.22. Pseudocyclosorus linearis Ching & K. H. Shing ex Y. X.Lin, Fl. Reipubl. Popularis Sin. 4(1): 328. 1999.线羽假毛蕨 xian yu jia mao juePlants more than 1.6 m tall. Rhizomes not seen. Stipes ca.30 cm, bases dark brown, distally stramineous, glabrous; laminaelanceolate, ca. 140 × 40 cm, pinnate-pinnatifid, pinnatifidacuminateat apices; proximal 9 pairs of pinnae abruptlyreduced into auricles, middle normal pinnae more than 30 pairs,alternate, obliquely spreading, sessile, linear-lanceolate, ca. 20× 2.3 cm, not broadened at bases, pinnatifid, pinnatifid-acuminateat apices; segments ca. 40 pairs, ligulate, 9–11 × ca. 5mm, acute at apices. Veins not evident, ca. 12 pairs of veinletsper segment, proximal pair reaching bottom of sinus. Laminaedark green when dry, herbaceous; abaxially setaceous alongrachises, glabrous along costae, veins densely glandular, adaxiallydensely setaceous along costal grooves, with 1 or 2 setaeon veinlets, with few fine hairs along costules. Sori orbicular,attached on middle of veinlets; indusia orbicular-reniform,brown, thick, persistent, glabrous.● Mixed forests by streams; ca. 900 m. Sichuan (Emei Shan).Pseudocyclosorus linearis is similar to P. esquirolii but differs inthe pinnae linear-lanceolate and the proximal pair of veinlets on segmentsreaching bottom of sinus.23. Pseudocyclosorus submarginalis Ching ex Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 328. 1999.边囊假毛蕨 bian nang jia mao juePlants to 75 cm tall. Rhizomes creeping, woody, firm.Fronds remote; stipes ca. 23 cm, deep stramineous, bases withsparse deep brown lanceolate scales, distally glabrous; laminaebroadly lanceolate, ca. 50 × 25 cm, abruptly narrowed proximally,pinnate-pinnatifid, pinnatifid-acuminate at apices; proximal3 pairs of pinnae reduced and hastate; middle normal pinnaeca. 21 pairs, alternate, spreading, sessile, 12–14 × 1.5–1.7cm, bases nearly tapering, truncate, pinnatifid nearly to costae,apices pinnatifid and long acuminate; segments ca. 22 pairs,obliquely spreading, lanceolate, ca. 7 × 4 mm, entire, roundedobtuseat apices. Veins raised on both sides, 7–9 pairs of veinletsper segment, proximal pair arising above base of costules,acroscopic veinlet reaching bottom of sinus, basiscopic onereaching margin above sinus. Laminae greenish when dry, thinlypapery; abaxially sparsely long acicular hairy along rachises,glabrous along costae, sparsely finely hairy and glandularalong veins, adaxially with dense appressed setae along costalgrooves, sparsely setaceous along veinlets. Sori orbicular,attached above middle of veinlets or closer to margins; indusiaorbicular-reniform, brown, glabrous, persistent.● Sichuan.Pseudocyclosorus submarginalis is similar to P. esquirolii but differsin the plants smaller, laminae ca. 50 cm, proximal pair of veinletson segment arising above base of costules, and sori attached abovemiddle of veinlets and close to margins.24. Pseudocyclosorus qingchengensis Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 329. 1999.青城假毛蕨 qing cheng jia mao juePlants to 1.4 m tall. Rhizomes creeping, woody, firm, darkbrown, sparsely scaly; scales lanceolate, deep brown, fragileand easily deciduous. Fronds remote; stipes 32–37 cm, basesdark brown, distally stramineous, glabrous; laminae lanceolate,ca. 100 × 30 cm, abruptly narrowed at bases, pinnate-pinnatifid,pinnatifid-acuminate at apices; proximal 3–6 pairs of pinnaereduced and hastate; middle normal pinnae ca. 30 pairs, proximalones subopposite, distal ones alternate, sessile, narrowlylanceolate, 12–19 × ca. 2.5 cm, truncate at bases, pinnatifid,pinnatifid-acuminate at apices; segments ca. 37 pairs, 8–10 ×ca. 3 mm, entire, obtuse-pointed at apices. Veins evident, costulesraised on both sides, ca. 12 pairs of veinlets per segment,proximal pair arising above base of costules, acroscopic veinletreaching bottom of sinus, basiscopic one reaching margin abovesinus. Laminae dark green when dry, papery; abaxially withlong acicular hairs along rachises, sparsely shortly hairy alongcostae and veins, densely shortly hairy on intercostal areas,adaxially with dense appressed setae along costal grooves, with1 or more setae along veins and glabrous on intercostal areas.Sori orbicular, attached on middle of veinlets; indusia orbicularreniform,brown, thick, glabrous, persistent.

368THELYPTERIDACEAE● Forests on slopes, wet shaded forests by streams. Guangxi(Longsheng), Sichuan (Guanxian, Qingchengshan).Pseudocyclosorus qingchengensis is similar to P. esquirolii inlaminar outline but differs by the proximal pair of segments on largerpinnae clearly elongate, proximal pair of veinlets arising from abovebase of costules, and laminae with dense short hairs on intercostalareas abaxially.25. Pseudocyclosorus angustipinnus Ching ex Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 159, 329. 1999 [“angustpinnus,”p. 329].狭羽假毛蕨 xia yu jia mao juePlants to 90 cm tall. Rhizomes creeping, dark brown,woody, apices with brown scales. Fronds remote; stipes ca. 35cm, bases dark brown, sparsely scaly, distally stramineous andglabrous; laminae lanceolate, ca. 50 × 16 cm, pinnate-pinnatifid,pinnatifid-acuminate at apices; proximal pair of pinnae reducedinto auricles, second pair slightly shorter than abovenormal ones; normal pinnae ca. 25 pairs, spreading, alternate,sessile, lanceolate, 8–9 × ca. 1.5 cm, rounded-truncate at bases,pinnatifid nearly to costae, long acuminate at apices; segmentsca. 20 pairs, ca. 6 × 3 mm, entire, rounded-obtuse at apices.Veins evident on both sides, costules raised on both sides, ca. 10pairs of veinlets per segment, proximal pair arising above baseof costules, acroscopic vein reaching bottom of sinus, basiscopicones reaching margin above sinus. Laminae grayishgreen when dry, papery; abaxially with dense long acicularhairs along rachises, sparsely setaceous and with fine short hairsalong costae and veins, also finely hairy on intercostal areas,adaxially densely appressed-setaceous along costal grooves,sparsely setaceous along veins, glabrous on intercostal areas.Sori orbicular, attached above middle of veinlets and close tomargins, brown, thick, glabrous, persistent.● Wet forests; ca. 1000 m. Guizhou (Fanjing Shan).Pseudocyclosorus angustipinnus is similar to P. esquirolii butdiffers by the laminae hairy on intercostal area abaxially and the sori ondistal parts of veinlets.26. Pseudocyclosorus stramineus Ching ex Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 159, 329. 1999 [“straminea,” p.329].禾杆假毛蕨 he gan jia mao juePlants ca. 1.2 m tall. Rhizomes not seen. Stipes ca. 36 cm,with dense acicular hairs. Laminae lanceolate, ca. 65 × 20 cm,tapering to both ends, pinnate-pinnatifid; proximal ca. 8 pairs ofpinnae reduced and hastate; normal pinnae more than 25 pairs,alternate, spreading, sessile, narrowly lanceolate, ca. 13 × 1.7cm, not tapering to bases, 1-pinnatifid, pinnatifid-acuminate atapices; segments more than 30 pairs, lanceolate, ca. 7 × 4 mm,entire, acute at apices. Veins raised on both sides, ca. 9 pairs ofveinlets per segment, proximal pair arising above base of costules,basiscopic veinlet reaching to margin above sinus, acroscopicone reaching bottom of sinus. Laminae dark brown whendry, papery; rachises with dense long acicular hairs on bothsides, along abaxial sides of costae sparsely acicular hairy, withappressed short setae along veins adaxially, with 1 or 2 setaealong veinlets and margins. Sori orbicular and attached onmiddle of veinlets, ca. 7 pairs per segment; indusia orbicularreniform,brown, thick, glabrous, persistent.● W Yunnan.27. Pseudocyclosorus duclouxii (Christ) Ching, Acta Phytotax.Sin. 8: 324. 1963.苍山假毛蕨 cang shan jia mao jueDryopteris duclouxii Christ, Bull. Acad. Int. Géogr. Bot.17: 139. 1907; Thelypteris duclouxii (Christ) Ching.Plants to 2 m tall. Rhizomes creeping. Fronds remote;stipes ca. 55 cm, with sparse appressed ovate scales, distallywith sparse grayish white short hairs; laminae ca. 150 × 35 cm,abruptly narrowed to bases, pinnate-pinnatifid, pinnatifid-acuminateat apices; proximal pairs of pinnae reduced to triangularauricles; middle normal pinnae ca. 45 pairs, obliquelyspreading, alternate, sessile, lanceolate, 18–20 × 3–3.5 cm,truncate at bases, pinnatifid to broad wing on both sides of costae,long pinnatifid-acuminate at apices; segments ca. 40 pairs,spreading, proximal one 10–12 × ca. 4 mm, entire or not clearlyundulate-crenate, acuminate at apices. Veins evident on bothsides, 11–13 pairs of veinlets per segment, proximal pair arisingfrom base of costules, acroscopic veinlet reaching bottom ofsinus, basiscopic one reaching margin above sinus. Laminaedark brown when dry, herbaceous; abaxially with dense acicularhairs along rachises, sparsely shortly hairy along costaeand veins, adaxially densely appressed-setaceous along costalgrooves, sparsely setaceous along veins and margins, glabrouson both surfaces on intercostal areas. Sori orbicular, attachedabove middle of veinlets and close to margins; indusia small,orbicular-reniform, brownish, thickly membranous, glabrous,persistent.● Streamsides. W Yunnan (Dali: Cangshan).28. Pseudocyclosorus jijiangensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 330. 1999.綦江假毛蕨 qi jiang jia mao juePlants to 40 cm tall. Rhizomes decumbent, sparsely brownscaly. Fronds approximate; stipes ca. 14 cm, stramineous,sparsely brown scaly on bases, distally glabrous; laminae ovatelanceolate,ca. 28 × 17 cm, abruptly narrowed at bases, pinnatepinnatifid,pinnatifid-acuminate at apices; proximal 2 pairs ofpinnae reduced and hastate, reflexed, middle normal pinnae ca.16 pairs, proximal ones opposite, distal ones alternate, sessile,slightly oblique distally, lanceolate, 8.5–9 × ca. 2 cm, broadenedand truncate at bases, pinnatifid, long pinnatifid-acuminateat apices; segments 22–24 pairs, pinnatifid nearly to costae,lanceolate, except for proximal pair, all obliquely spreading distally,proximal pair clearly elongated, ca. 6 × 3 mm, proximalpair of segments on lowest 2 pairs of pinnae pinnate-lobate andundulate, acuminate at apices, others all obtuse-pointed and entire.Veins evident, 6 or 7 pairs of veinlets per segment, proximalpair arising above base of costules, acroscopic veinletreaching bottom of sinus, basiscopic one reaching margin abovesinus. Laminae dark green when dry, papery; abaxially sparselong acicular hairy along rachises, sparsely setaceous alongcostae and veins, adaxially densely appressed-setaceous along

THELYPTERIDACEAE 369costal grooves, sparsely setaceous along veins and margins,glabrous on intercostal areas on both surfaces. Sori orbicular,attached above middle of veinlets or slightly closer to margins;indusia orbicular-reniform, brownish, membranous, persistent,glabrous.● Raised land by wells. Chongqing (Qijiang).29. Pseudocyclosorus tsoi Ching, Fl. Fujian. 1: 619. 1982.景烈假毛蕨 jing lie jia mao juePlants 75–150 cm tall. Rhizomes ascending, with densebrown broadly lanceolate scales on apices. Fronds subclustered;stipes 20–50 cm, dark brown at bases and sparsely scaly, distallystramineous and glabrous; laminae oblong-lanceolate, 50–100 × 20–30 cm, abruptly narrowed at bases, pinnate-pinnatifid,pinnatifid-acuminate at apices; proximal pairs of pinnae becomingauricles or hastate; middle normal pinnae 20–25 pairs,spreading, sessile, alternate, narrowly lanceolate, 15–18 × 1.5–3cm, slightly broadened at bases and broadly cuneate, pinnatifidnearly to costae, pinnatifid and long acuminate at apices; segments20–30 pairs, oblique distally, lanceolate, proximal pairclearly elongated to ca. 15 mm, distal ones 7–12 × 2–4 mm,entire, obtuse-pointed at apices. Veins evident on both sides,costules raised, 9–12 pairs of veinlets per segment, proximalpair arising from base of costules, acroscopic veinlet reachingbottom of sinus, basiscopic one reaching margin above sinus.Laminae dark green when dry, papery; abaxially finely hairyalong rachises, costae, and veins, adaxially densely appressedsetaceousalong costal grooves, sparsely setaceous on apices ofcostae, veinlets, and margins, glabrous on intercostal areas. Soriorbicular, attached on middle of veinlets; indusia orbicularreniform,brownish, thickly membranous, glabrous, persistent.● Wetlands in valleys, streamsides; 500–700 m. Fujian, Guangdong,Guangxi, Hunan, Jiangxi, Zhejiang.30. Pseudocyclosorus emeiensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 330. 1999.峨眉假毛蕨 e mei jia mao juePlants to 100 cm tall. Rhizomes not seen. Stipes 15–20cm, stramineous, glabrous; laminae lanceolate, 80–90 × ca. 20cm, pinnate-pinnatifid; proximal 4 pairs of pinnae reduced andhastate; middle normal pinnae ca. 20 pairs, spreading, alternate,narrowly lanceolate, 12–14 × ca. 1.4 cm, bases rounded-truncate,sessile, pinnatifid, remote from each other; segments ca.30 pairs, narrowly ligulate, ca. 5 × 2 mm, margins undulate ondistal half; proximal pair of segments not clearly elongated.Veins evident on both sides, ca. 8 pairs of veinlets per segment,simple, basal acroscopic veinlet reaching bottom of sinus, basiscopicone reaching margin above sinus. Laminae dark greenwhen dry, firmly papery; rachises subglabrous proximally, withfew setae distally, setae denser adaxially on rachises, abaxiallyfinely shortly hairy on costae, costules with appressed softhairs and glands, adaxially with dense setae along grooves ofcostae, sparsely hairy on veins, glabrous on intercostal areas onboth surfaces. Sori orbicular and attached above middle ofveinlets; indusia orbicular-reniform, brown, thick, glabrous,persistent.● Streamsides, under Larix trees; ca. 700 m. Sichuan (EmeiShan).31. Pseudocyclosorus furcatovenulosus Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 330. 1999 [“furcato-venulosus”].叉脉假毛蕨 cha mai jia mao juePlants to 130 cm tall. Rhizomes creeping. Stipes ca. 20cm, bases brown and sparsely scaly, distally stramineous andglabrous; laminae ca. 100 × 40 cm, pinnate-pinnatifid; proximal6 pairs of pinnae abruptly reduced and hastate; middle normalpinnae to 28 pairs, spreading, alternate, sessile, lanceolate, ca.20 × 2.5–3 cm, truncate at bases, acuminate at apices, pinnatifid;segments more than 30 pairs, lanceolate, to ca. 16 × 7mm, shallowly pinnatifid. Veins evident abaxially, 11–13 pairsof veinlets per segment and usually furcate, basal acroscopicveinlet reaching bottom of sinus, basiscopic one reaching marginabove sinus. Laminae dark green when dry, papery; abaxiallysubglabrous along rachises and costae, sparsely shortlyhairy along veins, adaxially densely setaceous along costalgrooves, finely hairy along veins and with 1 or 2 setae alongveinlets, glabrous on intercostal areas. Sori nearly orbicular,attached above middle of veinlets and closer to margins; indusiaorbicular-reniform, papery, brown, glabrous, persistent.● Forests of Phyllostachys pubescens on S slopes. Sichuan(Junlian).32. Pseudocyclosorus damingshanensis Ching ex Y. X. Lin,Fl. Reipubl. Popularis Sin. 4(1): 162, 330. 1999 [“daminshanensis,”p. 330].大明山假毛蕨 da ming shan jia mao juePlants ca. 64 cm tall. Rhizomes decumbent, dark brown.Fronds clustered; stipes ca. 14 cm, bases brown and sparselybrown scaly, distally stramineous and glabrous; laminaelanceolate, ca. 32 × 23 cm, tapering to both ends, pinnate-pinnatifid;proximal 3 pairs of pinnae gradually reduced and hastate;middle normal pinnae ca. 17 pairs, spreading, alternate,sessile, narrowly lanceolate, ca. 11 × 1.7 cm, pinnatifid; segmentsca. 20 pairs, lanceolate, ca. 8 × 3 mm, entire, acute atapices. Veins raised on both sides, ca. 8 pairs of veinlets persegment, proximal pair arising above base of costules, acroscopicveinlet reaching bottom of sinus, basiscopic onereaching margin above sinus. Laminae dark green when dry,thinly papery; abaxially with few fine short hairs along costaeand veins and mixed with 1 or 2 setae, adaxially densely longsetaceous along rachises and costal grooves, except for fewshort hairs also with few setae along costae, glabrous on intercostalareas. Sori orbicular, attached on middle of veinlets, 4or 5 pairs per segment; indusia brown, thick, with fine hairs,persistent.● Guangxi (Daming Shan).33. Pseudocyclosorus obliquus Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 331. 1999.斜展假毛蕨 xie zhan jia mao juePlants ca. 1.2 m tall. Rhizomes not seen. Stipes ca. 30 cm,brown and glabrous; laminae oblong-lanceolate, ca. 100 × 30

370THELYPTERIDACEAEcm, tapering to both ends, pinnate-pinnatifid; lower 4 pairs ofpinnae abruptly reduced and hastate; middle normal pinnae ca.20 pairs, extremely oblique distally, alternate, narrowly lanceolate,ca. 25 × 3 cm, acuminate at apices, entire, pinnatifid;segments more than 35 pairs per pinna, lanceolate, 1.2–1.5 ×ca. 0.3 cm, entire, acuminate at apices, basal acroscopic oneslightly elongated. Veins visible on both sides and all raised, ca.11 pairs of veinlets per segment, proximal veinlet arising frombase of costules and reaching margin above sinus, acroscopicone arising above base of costules and reaching bottom ofsinus. Laminae grayish green when dry, papery; abaxially withshort setae along rachises, subglabrous along costae and veins,adaxially densely setaceous along costal grooves, sparsely hairyalong veins and margins, glabrous on intercostal areas on bothsides. Sori orbicular and attached on distal parts of veinlets, ca.9 pairs per segment; indusia orbicular-reniform, brown, thick,glabrous, persistent.● Streamsides. Guangxi (Lingui).34. Pseudocyclosorus subochthodes (Ching) Ching, Acta Phytotax.Sin. 8: 325. 1963.普通假毛蕨 pu tong jia mao jueThelypteris subochthodes Ching, Bull. Fan Mem. Inst.Biol., Bot. 6: 305. 1936; Cyclosorus subochthodes (Ching) L. J.He & X. C. Zhang; Dryopteris eberhardtii Christ var. glabrataChrist; Lastrea subochthodes (Ching) Tagawa.Plants 90–110 cm tall. Rhizomes short and decumbent,dark brown, sparsely scaly. Fronds approximate or clustered;stipes 20–25 cm, bases dark brown and sparsely scaly, distallystramineous and glabrous; laminae oblong-lanceolate, 70–85 ×ca. 20 cm, abruptly narrowed at bases, pinnate-pinnatifid,pinnatifid-acuminate at apices; proximal 3 or 4 pairs of pinnaeabruptly reduced into triangular auricles; middle normal pinnae26–28 pairs, subopposite or alternate, obliquely spreading, sessile,lanceolate, 10–15 × 1.2–2 cm, not or slightly narrowed tobases, rounded-cuneate, pinnatifid nearly to costae, pinnatifidand long acuminate at apices; segments 28–30 pairs, lanceolate,acroscopic one on proximal pair slightly elongated, others 7–9× 2–3.5 mm, entire, acute or acuminate at apices. Veins evidenton both sides, costules raised, 9 or 10 pairs of veinlets per segment,proximal pair arising above bases of costae, acroscopicveinlet reaching bottom of sinus, basiscopic one reaching marginabove sinus. Laminae grayish green when dry, papery, glabrouson both intercostal areas, abaxially subglabrous orsparsely shortly hairy along rachises, costae, and veins, adaxiallydensely appressed-setaceous along costal grooves and with1 or 2 setae along veins. Sori orbicular, attached above middleof veinlets and slightly closer to margin; indusia orbicular-reniform,thickly membranous, glabrous, persistent.Wet places in mixed forests, rocks in valleys; 200–2000 m. SAnhui, Fujian, Guangdong, SE Guangxi, Guizhou, Hunan, Jiangxi, Sichuan,Yunnan, Zhejiang [Japan, Korea].35. Pseudocyclosorus latilobus (Ching) Ching, Acta Phytotax.Sin. 8: 324. 1963.阔片假毛蕨 kuo pian jia mao jueThelypteris latiloba Ching, Bull. Fan Mem. Inst. Biol.,Bot. 6: 303. 1936.Caudex not seen; base of stipes missing, remainderslightly flushed red, glabrous, ca. 25 cm. Laminae ca. 80 cm;pinnae over 20 pairs; basal pinnae slightly reduced and morewidely spaced, frond apex not seen. Largest pinnae ca. 16 × 3cm; aerophores not enlarged; basal acroscopic segments sometimeselongate, basal basiscopic segments of several lower pinnaereduced, edges lobed to 2 mm from costae; segments hardlyfalcate, oblong or oblong-lanceolate, 1.2–1.7 × ca. 0.6 cm,entire or shallowly undulate, acute at apices; costules ca. 7 mmapart. Veins ca. 11 pairs per segments, concolorous, slender, notprominent. Laminae herbaceous; abaxially rachis with sparsehairs, costae with short spreading hairs near base only, shortcapitate hairs on distal part, this distribution repeated on costuleswhich also bear some much-reduced scales consisting of 2cells; adaxially costae densely hairy, costules with sparse shorthairs and scattered long ones. Sori small, attached above middleof veinlets and close to margins; indusia pale, firm, with veryshort capitate hairs on edge; stalks of sporangia slender, sometimeswith a sessile spherical cell.● C Guizhou (Ziyun).36. Pseudocyclosorus guangxianensis Ching ex Y. X. Lin, Fl.Reipubl. Popularis Sin. 4(1): 331. 1999.灌县假毛蕨 guan xian jia mao juePlants to 80 cm tall. Rhizomes decumbent, dark brown,woody, firm. Fronds approximate; stipes ca. 30 cm, brownproximally, distally stramineous and glabrous; laminae lanceolate,ca. 55 × 24 cm, tapering proximally, pinnate-pinnatifid,pinnatifid-acuminate at apices; free pinnae ca. 20 pairs, slightlyobliquely spreading, sessile, subopposite, proximal 2 or 3 pairsof pinnae slightly shortened; middle ones lanceolate, ca. 12 ×1.9 cm, bases subcuneate, pinnatifid nearly to costae, pinnatifidacuminateat apices; segments ca. 23 pairs, lanceolate, ca. 7 × 4mm, entire, obtuse-pointed at apices; basal acroscopic segmentparticularly elongated (to 1 cm). Veins evident on bothsides, costules raised abaxially, ca. 9 pairs of veinlets, proximalpair arising above bases of costae, acroscopic veinlet reachingbottom of sinus, basiscopic one reaching margin above sinus.Laminae greenish red when dry, subherbaceous; abaxially withsparse acicular long hairs along rachises, hairs denser alongcostae and veins, fine and long acicular hairy on intercostalarea, adaxially with dense appressed short setae along rachisesand costal grooves, subglabrous along veins, hairy along margins,glabrous on intercostal areas. Sori orbicular and attachedbelow middle of veinlets and closer to costules; indusia orbicular-reniform,hairy, brown, persistent. Sporangia each with 2 or3 erect acicular hairs.● Shaded places in forests; ca. 1200 m. Sichuan (Guanxian: QinchengShan).37. Pseudocyclosorus ciliatus (Wallich ex Bentham) Ching,Acta Phytotax. Sin. 8: 324. 1963.溪边假毛蕨 xi bian jia mao jueAspidium ciliatum Wallich ex Bentham, Fl. Hongk. 455.

THELYPTERIDACEAE 3711861; Dryopteris calcarata (Blume) Kuntze var. sericea (J.Scott ex Beddome) C. Christensen; D. ciliata (Wallich ex Bentham)C. Christensen; D. pseudocalcarata C. Christensen; Lastreacalcarata (Blume) T. Moore var. ciliata (Wallich ex Bentham)Beddome; L. calcarata var. sericea (J. Scott ex Beddome)Beddome; L. ciliata (Wallich ex Bentham) Hooker(1857), not Liebmann (1849), nor C. Presl (1851); L. sericea J.Scott ex Beddome; Nephrodium calcaratum (Blume) Hookervar. ciliatum (Wallich ex Bentham) Baker; N. calcaratum var.sericeum (J. Scott ex Beddome) Alderwerelt; Polypodium tenerumRoxburgh; Thelypteris ciliata (Wallich ex Bentham)Ching; T. sericea (J. Scott ex Beddome) C. F. Reed; T. tenera(Roxburgh) C. V. Morton ex Fraser-Jenkins; Trigonospora ciliata(Wallich ex Bentham) Holttum.Plants 20–40 cm tall. Rhizomes short and erect, subglabrous.Fronds clustered; stipes 8–25 cm, dark brown, withsparse ovate-lanceolate scales, distally stramineous, throughoutdensely grayish white acicular hairy; laminae lanceolate, 12–15× 7–8 cm, slightly narrowed to bases, pinnate-pinnatifid, pinnatifid-acuminateat apices; pinnae ca. 15 pairs, proximal pairslightly shortened, opposite, reflexed, others ascending distally,alternate, sessile, lanceolate, 3.5–5 × 0.8–1.5 cm, broadlycuneate at bases, pinnatilobate to 1/4–1/3 of distance to costae,pinnatifid-acuminate at apices; segments 9–12 pairs, subtriangular-lanceolate,1.5–4 × 1–1.5 mm (basal acroscopic one to6 mm), entire, obtuse at apices. Veins evident on both sides, 4–6pairs of veinlets per segment, proximal pair arising above baseof costules, acroscopic veinlet reaching bottom of sinus, basiscopicone reaching margin above sinus, sometimes 2 veinletsjoined and elongated to bottom of sinus. Laminae dark brownwhen dry, firmly papery, glabrous on both intercostal areas,with dense acicular hairs on both sides of rachises and costae,sparsely setaceous along both sides of veinlets. Sori orbicularand attached on proximal parts of veinlets and close to costules;indusia orbicular-reniform, dark brown, thickly membranous,with dense acicular hairs, persistent. Spores trilete.Wetlands in mountain valleys, rock crevices by streams; 100–900m. Guangdong, Guangxi, Hainan, Yunnan [Bhutan, N India, Indonesia,Malaysia, Myanmar, Nepal, Singapore, Sri Lanka, Thailand, Vietnam].38. Pseudocyclosorus caudipinnus (Ching) Ching, Acta Phytotax.Sin. 8: 324. 1963.尾羽假毛蕨 wei yu jia mao jueThelypteris caudipinna Ching, Bull. Fan Mem. Inst. Biol.,Bot. 6: 288. 1936.Plants ca. 55 cm tall. Rhizomes not seen. Stipes ca. 28 cm;laminae broadly lanceolate, ca. 27 × 15 cm, pinnatifid-acuminateat apices, bases not narrowed, pinnate-pinnatifid; pinnae11–13 pairs, proximal 8 or 9 pairs opposite, distal onesalternate, proximal pair reflexed proximally, middle pinnaespreading, distal ones obliquely spreading, sessile; middle pinnaelanceolate, 7–8 × ca. 1.5 cm, not narrowed to bases, pinnatifidnearly to costae, pinnatifid and long acuminate at apices;segments ca. 15 pairs, broadly lanceolate, basal acroscopic oneslightly elongated, others ca. 5 × 4 mm, entire, obtuse-pointedat apices. Veins evident, basal acroscopic veinlet on segmentsreaching bottom of sinus, basiscopic one reaching marginabove sinus. Laminae dark brown when dry, papery; abaxiallyrachises, costae, and veins with long hairs, finely hairy onintercostal areas, adaxially with dense, appressed setae alongcostal grooves, sparsely setaceous along veins, glabrous onintercostal areas. Sori orbicular, attached above middle of veinletsand closer to margins; indusia orbicular-reniform, brown,hairy, persistent. Spores trilete.● Hainan.13. MESOPTERIS Ching, Acta Phytotax. Sin. 16(4): 21. 1978.龙津蕨属 long jin jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants large, terrestrial, glabrous throughout. Rhizomes long creeping, dark brown, woody, including stipe bases with densedark brown lanceolate scales. Fronds remote; stipes strong, glabrous above bases, without tuberculate aerophores; laminae pinnatepinnatifid;terminal pinna of similar shape and size as proximal lateral ones, stalked; segments pointed at apices, proximal 2.5 pairsforming sharp triangular elongate and translucent membranes at sinuses, connivent at or just below sinuses, keels protruding(arching) on abaxial laminar surfaces. Laminae firmly papery when dry, dark brown, subglabrous on both surfaces. Sori orbicular,exindusiate, not close to costules; sporangia glabrous.One species: China, N Vietnam.Thelypteridaceae author Alan R. Smith prefers to follow Holttum and treat the one species of Mesopteris within Amphineuron (here subsumedwithin Cyclosorus). The present authors note that from the recent molecular phylogeny (He Lijuan, Molec. Phylogen. Evol., ined.) Mesopteris is notclustered together with Amphineuron species or other genera. While it is clearly a member of Cyclosorus s.l., it should be recognized as a separategenus, just as Glaphyropteridopsis and Pseudocyclosorus are here treated as separate genera.1. Mesopteris tonkinensis (C. Christensen) Ching, Acta Phytotax.Sin. 16(4): 22. 1978.龙津蕨 long jin jueDryopteris tonkinensis C. Christensen, Bull. Mus. Natl.Hist. Nat., sér. 2, 6: 102. 1934; Amphineuron tonkinense (C.Christensen) Holttum; Cyclosorus tonkinensis (C. Christensen)L. J. He & X. C. Zhang; Thelypteris tonkinensis (C. Christensen)Ching.Plants to 2 m tall. Stipes angular, to 1.2 m; laminae large,pinnate-pinnatifid, narrowly elliptic, 50–80 × ca. 40 cm; lateralpinnae to 30 pairs, 30–40 × ca. 2 cm, pinnatifid, proximal ones

372THELYPTERIDACEAEsubopposite, distal ones alternate, obliquely spreading, abaxially± with minute golden, spherical glands; veinlets 8–10 pairsper segment. Sori orbicular, in 1 or more lines along each sideof costae, usually arranged in 2–5 irregular lines, not close tocostules, or attached on proximal pair of veinlets and close tocostules; sterile on distal veinlets.On wet rocks in open forests on limestone mountains; ca. 100 m.SW Guangxi (Longjin) [N Vietnam].14. CYCLOSORUS Link, Hort. Berol. 2: 128. 1833.毛蕨属 mao jue shuLin Youxing ( 林尤兴 ), Li Zhongyang ( 李中阳 ); Kunio IwatsukiAmphineuron Holttum; Christella H. Léveillé; Pneumatopteris Nakai; Sphaerostephanos J. Smith.Plants usually medium-sized, terrestrial. Rhizomes creeping to erect, scaly (sometimes also hairy). Fronds distant to clustered;laminae usually oblong-lanceolate, narrowed or not to bases, pinnate to pinnate-pinnatifid; middle pinnae usually linear-lanceolate,subentire to deeply lobed, sometimes with swollen aerophores at bases abaxially; apical pinnae usually more lobed; proximal pinnaesometimes shortened; segments entire or rarely crenate; veinlets simple or rarely forked, proximal one or more pairs on adjacentsegments anastomosing with an excurrent veinlet from uniting point to sinus membrane, other veinlets to sinus membrane or marginabove sinus. Laminae herbaceous to papery, both surfaces usually hairy along costae and veins, abaxial surface sometimes glandular.Sori orbicular, usually at middle of veinlets; indusia orbicular-reniform, membranous, glabrous or hairy, sometimes glandular, persistent.Sporangia usually bearing hairs or glands. Spores oblong-reniform, cristate, with wings or echinate. x = 36.About 250 species: distributed throughout tropical and subtropical regions of the Old World, most in Asia; several species also in the NewWorld; 40 species (ten endemic) in China.“Christella sadlerioidea” (H. Léveillé, Fl. Kouy-Tchéou, 475. 1915) is a nomen nudum and was not therefore validly published (MelbourneCode, Art. 38.1(a)). Léveillé recorded C. urophylla (Mettenius) H. Léveillé (Fl. Kouy-Tchéou, 476. 1915; Phegopteris urophylla Mettenius, Abh.Senckenberg. Naturf. Ges. 2: 310. 1858; Aspidium urophyllum (Mettenius) Christ; Dryopteris urophylla (Mettenius) C. Christensen; Goniopterisurophylla (Mettenius) Beddome; Nephrodium urophyllum (Mettenius) Beddome; Polypodium urophyllum (Mettenius) Wallich ex Hooker; Thelypterisurophylla (Mettenius) K. Iwatsuki) from Guizhou, but there is no other record of this Malaysian species from China.The following taxa are excluded from the present treatment, pending further research: Cyclosorus lunganensis Ching (Wuyi Sci. J. 1: 5. 1981),described from Fujian, C. oppositipinnus Ching & Z. Y. Liu (Bull. Bot. Res., Harbin 4(3): 15. 1984), described from Sichuan, and C. subdentatusChing (Wuyi Sci. J. 1: 4. 1981), described from Fujian.1a. Aquatic plants; costae with ovate scales abaxially .................................................................................................... 1. C. interruptus1b. Terrestrial plants; costae without scales or rarely with lanceolate scales abaxially.2a. Pinnae with sessile spherical glands abaxially, or pustular between veins when dried; sporangia bearingsessile glands on capsules; spores yellow to light brown.3a. Pinnae without glands but pustular between veins when dried; segments truncate at apices .............................. 2. C. truncatus3b. Pinnae with sessile spherical glands abaxially and not pustular; segments acute or obtuse at apices.4a. Sporangia bearing hairs on capsules; rhizomes creeping .................................................................................. 3. C. productus4b. Sporangia glabrous; rhizomes erect or suberect.5a. Proximal pinnae gradually shortened; pinnae almost glabrous abaxially ..................................................... 4. C. latebrosus5b. Proximal pinnae abruptly shortened; pinnae hairy abaxially.6a. Proximal pinnae narrowed at bases; middle pinnae with basal segments not elongate; indusialarge, thickly membranous ..................................................................................................................... 5. C. heterocarpus6b. Proximal pinnae not narrowed at bases; middle pinnae with basal segments elongate; indusiasmall, membranous .................................................................................................................................. 6. C. taiwanensis2b. Pinnae without glands or with ellipsoid/clavate glands abaxially; sporangia bearing glands on stalksor without glands; spores brown to black.7a. Proximal veinlets on each segment sterile; veinlets with minute yellow glands abaxially.8a. Veinlets 1 pair beneath sinus; pinnae lobed 1/2–2/3 toward costae .................................................................. 7. C. opulentus8b. Veinlets ca. 2.5 pairs beneath sinus; pinnae lobed 1/3–1/2 toward costae ....................................................... 8. C. terminans7b. Proximal veinlets on each segment usually fertile; veinlets without such minute yellow glands.9a. Veinlets 1–1.5(–2) pairs beneath sinus; laminae herbaceous to papery.10a. Rachises distally with scales; lowest pinnae shortened to ca. 1 cm ............................................................... 9. C. crinipes10b. Rachises distally without scales; lowest pinnae much longer than 1 cm.11a. Proximal pinnae narrowed at bases, with shortened basal segments ................................................. 10. C. shimenensis11b. Proximal pinnae not narrowed or slightly so at bases, with basal segments not shortened.12a. Basal pinnae ± reduced; pinnae lanceolate or oblanceolate.13a. Plants 20–80 cm tall; pinnae without glands abaxially; veinlets ca. 1.5 pairs beneath sinus ........... 11. C. dentatus13b. Plants to 20 cm tall; pinnae with glands abaxially; veinlets 1 pair beneath sinus.

THELYPTERIDACEAE 37314a. Pinnae with dense, short acicular hairs; hairs on indusia no longer than diam. of sori;glands abaxially small .................................................................................................................. 12. C. subacutus14b. Pinnae with sparse acicular hairs; hairs on indusia longer than diam. of sori; glandsabaxially large ............................................................................................................................. 13. C. parvifolius12b. Proximal pinnae not reduced or slightly so; pinnae linear-lanceolate.15a. Pinnae without glands abaxially; rhizomes creeping or erect.16a. Rhizomes erect; excurrent veinlet short, sometimes obscure ...................................................... 14. C. siamensis16b. Rhizomes creeping; excurrent veinlet long and evident.17a. Sporangia bearing glands on stalks; acicular hairs on indusia shorter than diam. of sori ...... 15. C. procurrens17b. Sporangia lacking conspicuous glands; acicular hairs on indusia longer than diam.of sori .................................................................................................................................... 16. C. molliusculus15b. Pinnae glandular abaxially; rhizomes creeping.18a. Pinnae with long clavate glands; proximal pinnae narrowed at bases .................................... 17. C. cylindrothrix18b. Pinnae with ellipsoid glands or capitate glandular hairs; proximal pinnae not narrowedat bases.19a. Laminae much longer than stipes; veinlets 10–16 pairs per segment ....................................... 18. C. procerus19b. Laminae no longer than stipes; veinlets 4–8 pairs per segment.20a. Pinnae with glands abaxially; veinlets 5–8 pairs per segment; pinnae wider than 1 cm .... 19. C. parasiticus20b. Pinnae with glandular hairs abaxially; veinlets 4–6 pairs per segment; pinnae usuallynarrower than 1 cm ................................................................................................................ 20. C. pygmaeus9b. Veinlets 2 or more pairs beneath sinus; laminae papery.21a. Sporangia bearing hairs on capsules and eglandular.22a. Middle pinnae 1–2 cm wide; proximal pinnae not decrescent at bases ................................................. 21. C. hirtisorus22b. Middle pinnae 2–3.5 cm wide; proximal pinnae decrescent at bases ................................................... 22. C. attenuatus21b. Sporangia lacking hairs but with glands (rarely eglandular).23a. Rhizomes and stipe bases with ovate scales; pinnae with setae along costules; veinlets usuallyover 6 pairs beneath sinus ..................................................................................................................... 23. C. articulatus23b. Rhizomes and stipe bases with ovate-lanceolate to linear-lanceolate scales; pinnae subglabrousor with acicular hairs along costules; veinlets 2–5 pairs beneath sinus.24a. Sporangia bearing large spherical red glands on stalks; pinnae with minute hairs or subglabrousabaxially.25a. Rhizomes erect; veinlets ca. 2 pairs beneath sinus .............................................................................. 24. C. papilio25b. Rhizomes creeping; veinlets ca. 3 pairs beneath sinus ...................................................................... 25. C. evolutus24b. Sporangia bearing ellipsoid or clavate, golden or reddish orange glands on stalks(rarely eglandular); pinnae usually ± hairy abaxially.26a. Lateral pinnae lobed to 1/3 toward costae, sometimes subentire; pinnae with dense,capitate glandular hairs abaxially .................................................................................................... 26. C. latipinnus26b. Lateral pinnae usually more lobed; pinnae usually without glandular hairs.27a. Pinnae eglandular abaxially (sometimes with glandular hairs).28a. Veinlets ca. 2 pairs beneath sinus; middle pinnae with elongate basal acroscopicsegment ................................................................................................................................... 27. C. acuminatus28b. Veinlets more numerous; middle pinnae with normal or shortened basal acroscopicsegment.29a. Rhizomes shortly creeping; proximal pinnae not shortened or 1 or 2 pairs slightlyshortened.30a. Veinlets 5–8 pairs per segment; middle pinnae usually less than 1.5 cm wide ............... 28. C. nanxiensis30b. Veinlets 8–11 pairs per segment; middle pinnae usually more than 2 cm wide.31a. Lateral pinnae usually more than 12 pairs; sori orbicular ............................................. 29. C. scaberulus31b. Lateral pinnae usually less than 12 pairs; sori usually slightly elongate ...................... 30. C. calvescens29b. Rhizomes long creeping; proximal 3 or more pairs of pinnae shortened.32a. Pinnae with long acicular hairs abaxially; proximal shortened pinnae caudate atapices ................................................................................................................................. 31. C. hokouensis32b. Pinnae without hairs abaxially; proximal shortened pinnae acute or obtuse at apices.33a. Sporangia without glands; proximal shortened pinnae with basal segments notenlarged ............................................................................................................................... 32. C. gustavii33b. Sporangia bearing glands on stalks; proximal shortened pinnae with enlargedbasal segments ...................................................................................................... 33. C. wulingshanensis27b. Pinnae glandular abaxially.34a. Veinlets ca. 2 pairs beneath sinus; rhizomes erect to suberect ........................................... 34. C. jinghongensis

374THELYPTERIDACEAE34b. Veinlets more than 2 pairs beneath sinus; rhizomes shortly to long creeping.35a. Pinnae with ellipsoid glands throughout (along and between veins) abaxially .................... 35. C. jaculosus35b. Pinnae with clavate glands abaxially.36a. Veinlets ca. 3 pairs beneath sinus; lateral pinnae oblanceolate; glands sparse ....................... 36. C. ensifer36b. Veinlets more than 3 pairs beneath sinus; lateral pinnae linear-lanceolate orlanceolate; glands dense.37a. Veinlets straight and oblique; pinnae linear-lanceolate, lobed to 1/3 towardcostae, sometimes only serrate ............................................................................................. 37. C. aridus37b. Veinlets usually spreading and curved; pinnae lanceolate, usually morelobed than above.38a. Pinnae subglabrous abaxially; proximal pinnae decurrent toward bases ..................... 38. C. cuneatus38b. Pinnae densely hairy abaxially; proximal pinnae truncate or rounded at bases.39a. Rhizomes 2–3 mm in diam.; fronds distant; proximal shortened pinnae notor slightly narrower than middle ones .................................................................... 39. C. fukienensis39b. Rhizomes 5–10 mm in diam.; fronds approximate; proximal shortenedpinnae much narrower than middle ones .................................................................. 40. C. subelatus1. Cyclosorus interruptus (Willdenow) H. Itô, Bot. Mag. (Tokyo)51: 714. 1937.毛蕨 mao juePteris interrupta Willdenow, Phytographia, 13. 1794;Aspidium gongylodes Schkuhr; A. unitum (Linnaeus) Swartzvar. glabrum Mettenius; A. unitum var. hirsutum Mettenius;Cyclosorus gongylodes (Schkuhr) Link; C. gongylodes var.glaber (Mettenius) Ching; C. gongylodes var. hirsutus (Mettenius)Farwell; Dryopteris gongylodes (Schkuhr) Kuntze; D.gongylodes var. hirsuta (Mettenius) C. Christensen; D. interrupta(Willdenow) Ching; Nephrodium gongylodes (Schkuhr)Schott; Thelypteris gongylodes (Schkuhr) Small; T. interrupta(Willdenow) K. Iwatsuki.Plants 40–100 cm tall. Rhizomes long creeping, black,including bases of stipes with sparse ovate-lanceolate scales.Fronds distant; stipes 20–60 cm, bases black, distally stramineous;laminae 20–50 × 10–20 cm, bases not narrowed, apicescaudate with apical pinna similar to lateral ones; lateral pinnae10–25 pairs, linear-lanceolate, 5–10 × ca. 1 cm, shortly stalked,bases rounded-truncate, lobed 1/4–1/2 toward costae, apicesacuminate; segments 20–30 pairs on middle pinnae, triangular,3–5 × 2–4 mm, apices pointed; veinlets 6–10 pairs, proximalpair anastomosing, next 0.5–1 pair running to sinus membrane.Laminae somewhat leathery, reddish when dried, subglabrousadaxially, abaxially with acicular hairs and reddish orange sessilespherical glands along veins and several membranousbroadly ovate scales along costae. Sori orbicular, medial to submarginal,proximal 1 or 2 pairs of veins sterile; indusia sparselyhairy. Sporangia bearing spherical reddish orange glands onstalks. Spores sparsely minutely echinulate. 2n = 72, 144.Swamps, wetlands; near sea level to 500 m. Fujian, Guangdong,Guangxi, Hainan, Jiangxi, Taiwan, S Yunnan [throughout tropical andsubtropical regions of the world].According to recent molecular study by Smith and Cranfill (Amer.Fern J. 92(2): 131–149. 2002), Cyclosorus interruptus is distantly relatedto other species in this genus but is quite close to Ampelopterisprolifera. This relationship requires further study.2. Cyclosorus truncatus (Poiret) Farwell, Amer. Midl. Naturalist12: 259. 1931.截裂毛蕨 jie lie mao juePlants 0.6–2 m tall. Rhizomes erect, stipes throughout withbrown ovate-lanceolate scales. Fronds clustered; stipes 20–50cm; laminae oblong-lanceolate, 40–150 × 20–40 cm, basesabruptly narrowed, apices acuminate; pinnae 20–35 pairs, proximal2–6 pairs abruptly shortened, middle pinnae linear, 12–30× 1–2.5 cm, bases rounded-truncate and usually slightly narrowedon proximal pinnae, lobed 1/4–1/2 toward costae, apiceslong acuminate; aerophores beneath pinna bases swollen; segments25–40 pairs, rectangular, 3–6 × 2–4 mm, margins usuallydentate, apices truncate or rounded-truncate; veinlets 4–10pairs, proximal 1 or 2 pairs anastomosing, next pair running tosinus membrane. Laminae papery, yellowish green or grayishgreen and pustular between veins when dried, glabrous on bothsurfaces. Sori orbicular, medial; indusia membranous, glabrous.Sporangia bearing light yellow glands on capsules. Spores darkyellow, irregularly thinly cristate or with small wings.Streamsides, wet places in forests; 100–1300 m. S Fujian, Guangdong,Guangxi, S Guizhou, Hainan, S Hunan, Taiwan, SE Xizang, SYunnan [India, Indonesia, Japan, Laos, Malaysia, Myanmar, Philippines,Sri Lanka, Thailand, Vietnam; N Australia, Pacific islands (Polynesia)].1a. Pinnae 1.5–2.5 cm wide, lobed 1/3–1/2toward costae ........................................... 2a. var. truncatus1b. Pinnae 1–1.5 cm wide, lobed to 1/3toward costae ................................... 2b. var. angustipinnus2a. Cyclosorus truncatus var. truncatus截裂毛蕨 ( 原变种 ) jie lie mao jue (yuan bian zhong)Polypodium truncatum Poiret, Encycl. 5: 534. 1804;Cyclosorus pustulifer Ching ex K. H. Shing [“pustuliferus”]; C.truncatus var. acutiloba Ching; C. truncatus f. kwashotensis(Hayata) H. Itô; C. truncatus f. laevifrons (Hayata) H. Itô; C.truncatus f. sublaevifrons (Tagawa) H. Itô; Dryopteris kwashotensisHayata; D. laevifrons Hayata; D. laevifrons var. kwashotensis(Hayata) Tagawa; D. sublaevifrons Tagawa; D. truncata(Poiret) Kuntze; Nephrodium truncatum (Poiret) C. Presl;Pneumatopteris truncata (Poiret) Holttum; Polystichum truncatum(Poiret) Gaudichaud; Thelypteris truncata (Poiret) K.Iwatsuki; T. truncata f. kwashotensis (Hayata) C. F. Reed; T.

THELYPTERIDACEAE 375truncata f. laevifrons (Hayata) C. F. Reed; T. truncata f. sublaevifrons(Tagawa) C. F. Reed.Plants 0.6–2 m tall. Middle pinnae 18–30 × 1.5–2.5 cm,lobed 1/3–1/2 toward costae; veinlets 6–10 pairs per segment,proximal 1 or 2 pairs anastomosing, next pair running to sinusmembrane. 2n = 72, 144.Streamsides, wet places in forests; 100–1300 m. S Fujian, Guangdong,Guangxi, S Guizhou, Hainan, S Hunan, Taiwan, SE Xizang, SYunnan [India, Indonesia, Japan, Laos, Malaysia, Myanmar, Philippines,Sri Lanka, Thailand, Vietnam; N Australia, Pacific islands (Polynesia)].In FRPS (4(1): 257. 1999), K. H. Shing mentioned that manyspecimens from China with broadened pinna bases differed from thetype. Although both broadened and narrowed pinna bases exist, all ofthe Chinese specimens are ± different in general outline from the type.More detailed studies are needed.2b. Cyclosorus truncatus var. angustipinnus Ching, Bull. FanMem. Inst. Biol., Bot. 8: 218. 1938.线羽截裂毛蕨 xian yu jie lie mao jueCyclosorus angustipinnus (Ching) K. H. Shing.Plants 0.6–1 m tall. Middle pinnae 12–18 × 1–1.5 cm,lobed to 1/3 toward costae; veinlets 4–6 pairs per segment,proximal pair anastomosing, next pair running to sinus membrane.● Wet places in forests; 400–600 m. Hainan.Compared with the typical variety, Cyclosorus truncatus var.angustipinnus has a smaller laminar outline, narrower pinnae, and feweranastomosing veinlets. According to Holttum (Blumea 21(2): 314.1973), it might be a diploid.3. Cyclosorus productus (Kaulfuss) Ching, Bull. Fan Mem.Inst. Biol., Bot. 10: 248. 1941.兰屿大叶毛蕨 lan yu da ye mao jueAspidium productum Kaulfuss, Enum. Filic. 237. 1824;Cyclosorus kotoensis (Hayata) W. C. Shieh; C. truncatus(Poiret) Farwell var. kotoensis (Hayata) H. Itô; Dryopteris kotoensisHayata; D. producta (Kaulfuss) C. Christensen; Sphaerostephanoskotoensis (Hayata) Holttum ex C. M. Kuo; S. productus(Kaulfuss) Holttum; Thelypteris kotoensis (Hayata) K.Iwatsuki; T. producta (Kaulfuss) C. F. Reed.Plants ca. 100 cm tall. Rhizomes shortly creeping, includingbases of stipes with dark brown linear-lanceolate scales.Fronds approximate; stipes ca. 20 cm, stramineous; laminae ca.100 × 30 cm, bases narrowed, apices acuminate; pinnae ca. 30pairs, proximal ca. 10 pairs abruptly shortened with lowest 1pair ca. 5 mm; middle pinnae linear-lanceolate, ca. 20 × 1.5–2cm, bases truncate, lobed 1/3–1/2 toward costae, apices longacuminate; segments 30–40 pairs on middle pinnae, oblong, ca.3 × 3 mm, subacute or obtuse at apices; veinlets 6–8 pairs persegment, proximal 1.5 pairs anastomosing, next pair running tosinus membrane. Laminae papery, shortly pubescent and withseveral acicular hairs on both surfaces along costae and veins,abaxially also with sessile spherical yellow glands. Sori orbicular,medial; indusia glandular and shortly hairy. Sporangiabearing several hairs and sessile spherical glands on capsules,and similar glands on stalks. Spores light brown, echinate orcristate. 2n = 72.Forest margins, slopes of ravines; 100–400 m. SE Taiwan (LanYu) [Philippines].4. Cyclosorus latebrosus (Kunze ex Mettenius) Ching, Bull.Fan Mem. Inst. Biol., Bot. 10: 245. 1941.阴生毛蕨 yin sheng mao jueAspidium latebrosum Kunze ex Mettenius, Abh. Senckenberg.Naturf. Ges. 2: 388. 1858; Sphaerostephanos latebrosus(Kunze ex Mettenius) Holttum.Plants 50–100 cm tall. Rhizomes erect. Fronds clustered;stipes 5–10 cm, dark stramineous, bases with sparse brownlanceolate scales; laminae 40–90 × 20–30 cm, bases graduallynarrowed, apices acuminate; pinnae 30–40 pairs, proximal ca.10 pairs gradually shortened with lowest one ca. 5 mm, slightlyreflexed; middle pinnae lanceolate, 10–18 × 1–2 cm, basesbroadly cuneate to truncate, lobed 1/2–2/3 toward costae, apicesacuminate; aerophores beneath pinna bases slightly swollen;segments 20–30 pairs on middle pinnae, oblong, 5–8 × 3–4mm, entire, obtuse at apices; veinlets 6–8 pairs, proximal pairanastomosing, next 0.5–1 pair running to sinus membrane.Laminae papery, reddish brown when dried, adaxially withsparse short acicular hairs along costae and veins, abaxial surfacewith yellow sessile spherical glands throughout and almostglabrous. Sori orbicular, medial; indusia glabrous, sometimesglandular. Sporangia bearing yellow spherical glands on capsules.Spores light brown, densely echinulate.Wet places in dense forests; 300–400 m. S Yunnan (Jinping)[Bangladesh, Indonesia, Malaysia, Philippines].“Cyclosorus glabellus” (Ching ex W. M. Chu & S. G. Lu, Fl.Yunnan. 20: 608. 2006) belongs here but was not validly published becauseno Latin description or diagnosis, or reference to such, was providedand no type was indicated (Melbourne Code, Art. 39.1 and 40.1).5. Cyclosorus heterocarpus (Blume) Ching, Bull. Fan Mem.Inst. Biol., Bot. 8: 180. 1938.异果毛蕨 yi guo mao jueAspidium heterocarpon Blume, Enum. Pl. Javae 2: 155.1828; Dryopteris heterocarpa (Blume) Kuntze; Nephrodiumheterocarpum (Blume) T. Moore; Sphaerostephanos heterocarpus(Blume) Holttum; Thelypteris heterocarpa (Blume) C.V. Morton.Plants 60–120 cm tall. Rhizomes erect, with brown lanceolatescales. Fronds clustered; stipes 10–20 cm, bases darkbrown and scaly, dark stramineous distally; laminae 40–100 ×20–30 cm, bases abruptly narrowed, apices acuminate to caudate;pinnae 20–40 pairs, sessile, proximal 5–10 pairs shortenedand auriculate with lowest 1 pair tuberculate; proximal pinnaenarrowed toward bases; middle pinnae linear-lanceolate, 10–16× 1–1.5 cm, bases truncate, lobed 1/2–2/3 toward costae, apiceslong acuminate; aerophores beneath pinna bases slightly swollen;segments 20–30 pairs, 4–5 × 2–2.5 mm, entire, apicesobtuse; veinlets 6–9 pairs, proximal pair anastomosing, some-

376THELYPTERIDACEAEtimes next vein running to sinus membrane. Laminae herbaceous,brownish green when dried, with short hairs alongcostae on both surfaces and along veins adaxially, also goldensessile spherical glands abaxially (sometimes also adaxially).Sori orbicular, medial; indusia slightly hairy, sometimesglandular. Sporangia bearing spherical glands on capsules.Spores light brown, densely echinulate. 2n = 72.Wet places in forests; 100–1100 m. Guangdong, Guangxi, Hainan[Indonesia, Malaysia, Philippines, Thailand, Vietnam; Australia, Pacificislands (Polynesia)].6. Cyclosorus taiwanensis (C. Christensen) H. Itô, Bot. Mag.(Tokyo) 51. 728. 1937.台湾毛蕨 tai wan mao jueDryopteris taiwanensis C. Christensen, Index Filic. 297.1905; Aspidium lobulatum Christ (1904), not Blume (1828); D.subhispidula Rosenstock; Sphaerostephanos taiwanensis (C.Christensen) Holttum ex C. M. Kuo; Thelypteris taiwanensis(C. Christensen) K. Iwatsuki.Plants 60–120 cm tall. Rhizomes suberect to erect, includingbases of stipes with brown ovate-lanceolate scales.Fronds clustered; stipes 10–20 cm, dark stramineous; laminaebroadly lanceolate, 50–100 × 20–30 cm, bases abruptly narrowed,apices acuminate; pinnae 30–45 pairs, proximal 4–6pairs abruptly shortened and auriculate with lowest 1 pair subtuberculate;middle pinnae linear-lanceolate, 10–18 × 0.8–1.2cm, bases truncate, apices long acuminate; lobed ca. 1/2 towardcostae; aerophores beneath pinna bases slightly swollen; segments25–35 pairs, 3–5 × 2–2.5 mm, basal acroscopic oneslonger, subrectangular, entire, obtuse at apices; veinlets 5–8pairs per segment, proximal pair anastomosing, next 0.5–1 pairrunning to sinus membrane. Laminae papery, brownish greenwhen dried, both surfaces with short acicular hairs along costaeand veins, adaxially with short hairs and yellow sessile sphericalglands throughout. Sori orbicular, medial or inframedial; indusiashortly hairy and glandular. Sporangia bearing sessilespherical glands on capsules. Spores yellow to light brown,densely echinulate. 2n = 72.Streamsides, wet places in dense forests; 200–700 m. Fujian,Guangdong, Guangxi, Jiangxi, Taiwan [S Japan].7. Cyclosorus opulentus (Kaulfuss) Nakaike, Enum. PteridJap., Filic. 277. 1975.腺脉毛蕨 xian mai mao jueAspidium opulentum Kaulfuss, Enum. Filic. 238. 1824;Amphineuron opulentum (Kaulfuss) Holttum; Aspidium extensumBlume; Cyclosorus extensus (Blume) H. Itô; Dryopterisextensa (Blume) Kuntze; Nephrodium extensum (Blume) T.Moore; N. opulentum (Kaulfuss) C. Presl; Thelypteris extensa(Blume) C. V. Morton; T. opulenta (Kaulfuss) Fosberg.Plants 60–100 cm tall. Rhizomes shortly creeping, apicesand bases of stipes with dark brown linear-lanceolate scales.Fronds approximate; stipes ca. 30 cm, brownish; laminae 30–60× 20–30 cm, bases not narrowed or slightly so, apices caudate;pinnae 10–20 pairs, almost sessile; proximal pair of pinnaeslightly shortened; middle pinnae linear-lanceolate, 15–25 × 1–2 cm, bases rounded-truncate (or slightly cuneate on proximalpinnae), lobed 1/2–2/3 toward costae, apices long acuminate;segments 25–40 pairs, middle ones subfalcate, 4–8 × 2–3 mm,entire, subacute to obtuse at apices; veinlets 8–10 pairs, proximalpair anastomosing or only connivent, sometimes next veinrunning to sinus membrane. Laminae herbaceous, yellowishgreen when dried, with several acicular hairs along veins onboth surfaces, minute hairs and many minute yellow sphericalglands along veins abaxially, also minute hairs between veinsabaxially. Sori orbicular, submarginal, usually sterile on proximal1 or 2 pairs of veins; indusia glabrous or glandular alongmargins. Sporangia bearing golden spherical glands on stalks.Spores cristate.Streamsides in forests or thickets; 100–300 m. Hainan [S India,Indonesia, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand; EAfrica, tropical America, N Australia, Micronesia].8. Cyclosorus terminans (J. Smith ex Hooker) K. H. Shing, Pl.Sci. Res. (India) 20(1–2): 25. 1998.顶育毛蕨 ding yu mao jueNephrodium terminans J. Smith ex Hooker, Sp. Fil. 4: 73.1862; Amphineuron terminans (J. Smith ex Hooker) Holttum;Thelypteris terminans (J. Smith ex Hooker) Panigrahi.Plants 40–100 cm tall. Rhizomes long creeping, includingbases of stipes with brown linear-lanceolate scales. Fronds distant;stipes 20–55 cm, dark stramineous; laminae deltoid-lanceolate,20–45 × 20–40 cm, bases not narrowed or slightly so,apices caudate with an apical pinna similar to lateral ones;lateral pinnae 10–15 pairs, subsessile; middle pinnae linearlanceolate,15–25 × 1–1.8 cm (usually narrowed toward bases),bases rounded-truncate, lobed 1/3–1/2 toward costae, apiceslong acuminate; segments 20–35 pairs, triangular, 3–4 × 3–4mm, subacute or obtuse at apices; veinlets 6–8 pairs, proximalpair anastomosing, next 1–1.5 pairs running to sinus membrane.Laminae papery, brownish green or yellowish green when dried,adaxially shortly hairy along costae and veins, abaxial surfacewith short hairs along costae and veins, also with minute sphericalsessile yellow glands along veins. Sori orbicular, marginal,sterile on proximal 2 or 3 pairs of veins, sometimes only top 2or 3 veins fertile; indusia glabrous. Sporangia bearing goldenglands on stalks. Spores cristate. 2n = 144.Wet places in forests or thickets; 200–500 m. Hainan [India,Indonesia, Japan, Malesia, Myanmar, Thailand, Vietnam; Africa, N Australia,Pacific islands (Micronesia, Polynesia)].The epithet is derived from “Aspidium terminans” (Wallich, Numer.List, no. 386. 1829; Kunze, Linnaea 23: 230. 1850), which is a nomennudum and was not therefore validly published (Melbourne Code,Art. 38.1(a)).9. Cyclosorus crinipes (Hooker) Ching, Bull. Fan Mem. Inst.Biol., Bot. 8: 179. 1938.鳞柄毛蕨 lin bing mao jueNephrodium crinipes Hooker, Sp. Fil. 4: 71. 1862; Christellacrinipes (Hooker) Holttum; Dryopteris crinipes (Hooker)Kuntze; Thelypteris crinipes (Hooker) K. Iwatsuki.

THELYPTERIDACEAE 377Plants 0.5–1.5 m tall. Rhizomes erect, apices with densedark brown lanceolate scales. Fronds clustered; stipes 20–40cm, dark stramineous, with dense scales from bases to rachises;laminae 30–120 × 25–45 cm, bases abruptly narrowed, apicesacuminate; pinnae 20–45 pairs, proximal 3–5 pairs shortened,triangular-auriculate, lowest one ca. 1 × 1 cm; middle pinnaelinear-lanceolate, 10–25 × 1–1.5 cm, bases truncate, lobed 1/2–2/3 toward costae, apices long acuminate; segments ca. 30pairs, oblong, 3–5 × ca. 3 mm, entire, apices obtuse; veinlets 7–9 pairs, proximal 1–1.5 pairs anastomosing, next pair running tosinus membrane. Laminae papery, grayish green when dried,with several acicular hairs along veins adaxially, with shortacicular hairs along costae and veins abaxially, also with minutehairs and minute glands throughout. Sori orbicular, medial; indusiashortly hairy. Sporangia bearing reddish orange glands onstalks. Spores dark brown, echinate. 2n = 72.Wet forest margins, wet places in forests; 100–1300 m. S Guangdong,S Guangxi, S Guizhou, Hainan, SE Xizang, S Yunnan [Bhutan,India, Indonesia, Laos, Myanmar, Nepal, Thailand, Vietnam].10. Cyclosorus shimenensis K. H. Shing & C. M. Zhang, KeysVasc. Pl. Wuling Mts. 565. 1995.石门毛蕨 shi men mao jueCyclosorus wangmoensis K. H. Shing & P. S. Wang.Plants 40–100 cm tall. Rhizomes shortly creeping, withdark brown lanceolate scales and setae. Fronds approximate;stipes 20–50 cm, dark stramineous; laminae 20–50 × 12–30 cm,bases slightly narrowed, apices acuminate to caudate; pinnae10–15 pairs, shortly stalked, proximal 1 or 2 pairs sometimesslightly shortened; middle pinnae falcate-lanceolate, 8–15 ×1.5–2.5 cm, bases rounded-cuneate (decurrent on proximal pinnae),lobed 2/3–3/4 toward costae, apices long acuminate;segments 20–30 pairs on middle pinnae, proximal segmentsshortened or missing on proximal pinnae; middle segments falcate-lanceolate,4–10 × 3–4 mm, obtuse or subacute at apices;veinlets 6–12 pairs, proximal pair anastomosing, excurrentveinlets usually obsolete on basal segments, next 0.5–1 pairrunning to sinus membrane. Laminae papery, grayish greenwhen dried, with short acicular hairs along costae and veins andminute hairs between veins adaxially, shortly hairy throughoutabaxially. Sori small, medial or supramedial; indusia shortlyhairy. Sporangia with reddish orange glands on stalks. Sporesdark brown, sparingly cristate.● Wet places in forests, semi-open forest margins; 300–1000 m.Chongqing, Guizhou, Hunan.The epithet was misspelled as “simenensis” in FRPS (4(1): 171,195. 1999).The types of Cyclosorus sinodentatus Ching & Z. Y. Liu and C.macrophyllus Ching & Z. Y. Liu both have features intermediatebetween C. shimenensis and C. dentatus. There are only a fewgatherings of C. sinodentatus other than the type. They are all rarelyfertile. Cyclosorus macrophyllus is represented by only one gathering(the type), lacking fertile sori. These two taxa might be hybrids.11. Cyclosorus dentatus (Forsskål) Ching, Bull. Fan Mem.Inst. Biol., Bot. 8: 206. 1938.齿牙毛蕨 chi ya mao juePolypodium dentatum Forsskål, Fl. Aegypt.-Arab. 185.1775; Christella dentata (Forsskål) Brownsey & Jermy; Cyclosorusangustus Ching (1982), not (Copeland) Ching (1941); C.fengii Ching ex K. H. Shing; C. jiulungshanensis P. S. Chiu &G. Yao ex Ching; C. mekongensis Ching ex K. H. Shing; C.mianningensis Ching ex K. H. Shing; C. paradentatus Ching exK. H. Shing; C. pingshanensis Ching & H. S. Kung ex K. H.Shing; C. proximus Ching; C. shapingbaensis Ching ex K. H.Shing; C. stenopes Ching & K. H. Shing; C. wangii Ching;Dryopteris dentata (Forsskål) C. Christensen; D. oblancifoliaTagawa; Thelypteris dentata (Forsskål) E. P. St. John.Plants (20–)40–60(–100) cm tall. Rhizomes suberect toshortly creeping, apices including bases of stipes with brownlinear-lanceolate scales. Fronds subclustered; stipes (5–)10–20(–35) cm, bases dark brown, stramineous distally; laminae(10–)15–40(–60) × 10–20 cm, bases slightly narrowed, apicesacuminate; lateral pinnae 10–20 pairs, proximal 2 or 3 pairsslightly shortened; middle pinnae lanceolate to oblanceolate, 5–10 × (0.8–)1–2 cm, bases rounded-truncate, lobed 1/2–2/3toward costae, apices acuminate; segments 10–20 pairs, rectangularor oblong, 3–5 × 3–4 mm (basal acroscopic one slightlylonger, sometimes crenate), rounded-obtuse at apices; veinlets5–8 pairs per segment, proximal pair anastomosing, next 0.5–1pair running to sinus membrane. Laminae herbaceous to papery,brownish green when dried, shortly hairy adaxially, along veinswith several acicular hairs, densely puberulent abaxially, sometimeswith glandular hairs. Sori orbicular, medial; indusiashortly hairy. Sporangia bearing minute golden glands on stalks.Spores brown, irregularly cristate. 2n = 72, 144, 288.Habitat variable, usually semi-open to open places; near sea levelto 2800 m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan,Hunan, Jiangxi, Sichuan, Taiwan, SE Xizang, Yunnan, S Zhejiang [NAfrica, tropical America, tropical and subtropical Asia].The type of Cyclosorus longqishanensis K. H. Shing (FRPS 4(1):337. 1999) has features intermediate between C. dentatus and C. acuminatus.Only one gathering (the type) with few sori has been collected.This taxon might be a hybrid.The type of Cyclosorus wenzhouensis K. H. Shing & C. F. Zhang(FRPS 4(1): 341. 1999) resembles C. dentatus but differs in having alarger laminar outline and 2 pairs of veinlets joined beneath the sinuses.Only one gathering (the type) with irregular spores has been collected.This taxon might be a hybrid.12. Cyclosorus subacutus Ching, Fl. Fujian. 1: 598. 1982.短尖毛蕨 duan jian mao juePlants 15–20 cm tall. Rhizomes short, suberect, with densedark brown lanceolate scales at apices. Fronds clustered; stipes3–7 cm, stramineous; laminae 8–14 × 3.5–6 cm, bases slightlynarrowed, apices acuminate; pinnae 6–12 pairs, sessile, proximal2 or 3 pairs slightly shortened; middle ones oblong-lanceolateto oblong-oblanceolate, 2–3 × ca. 1 cm, bases truncate,lobed 1/2–2/3 toward costae, apices acuminate or acute; segments6–10 pairs on middle pinnae, oblong, 2–3 × ca. 2 mm(basal acroscopic one slightly longer), entire, obtuse at apices;veinlets 3–5 pairs, proximal pair anastomosing with an excurrentveinlet running to sinus membrane. Laminae herbaceous,grayish green or brownish green when dried, adaxially with

378THELYPTERIDACEAEdense short acicular hairs along costae, with sparse acicularhairs along veins and shorter hairs between veins; abaxial surfacewith dense short thin acicular hairs and golden to orangesmall glands. Sori small, orbicular, medial; indusia denselyhairy. Sporangia bearing golden to orange glands on stalks.Spores with long to short wings.● Semi-open places by forest margins; near sea level to 100 m.Fujian, Guangdong, Jiangxi, ?Taiwan, S Zhejiang.Cyclosorus subacutus differs from C. dentatus in being muchsmaller plants and having glandular hairs abaxially, but in other aspects,the two species are quite similar. Knapp (Ferns Fern Allies Taiwan,2011) did not record this species for Taiwan.13. Cyclosorus parvifolius Ching, Fl. Fujian. 1: 598. 1982.小叶毛蕨 xiao ye mao juePlants 10–20 cm tall. Rhizomes suberect to shortlycreeping, with sparse brown lanceolate scales at apices. Frondsclustered; stipes 3–8 cm, stramineous; laminae 6–10 × 3–5 cm,bases slightly narrowed, apices acuminate; pinnae 5–8 pairs,sessile, proximal 1 or 2 pairs slightly shortened; middle onesoblong-lanceolate to broadly lanceolate, 1.5–2.5 × 0.5–1 cm,bases broadly cuneate to subtruncate, slightly projected on acroscopicside, lobed 1/3–1/2 toward costae, apices obtuse or subacute;segments 5 or 6 pairs on middle pinnae, subrectangular,2–3 × 2–3 mm, entire, obtuse at apices; veinlets 3 or 4 pairs,proximal 1 pair anastomosing with an excurrent veinlet runningto sinus membrane. Laminae herbaceous, yellowish green whendried, with several thin short acicular hairs on both surfaces,and reddish orange glands along veins abaxially. Sori orbicular,at end of basal veins and usually confluent; indusia with shortthin acicular hairs. Sporangia bearing reddish orange glands onstalks. Spores with short wings.● Fujian.Cyclosorus parvifolius resembles C. parasiticus and differs onlyin being much smaller in stature. The spores are not well developed.14. Cyclosorus siamensis (Tagawa & K. Iwatsuki) Panigrahi,Res. J. Pl. Environm. 9: 67. 1993.泰国毛蕨 tai guo mao jueThelypteris siamensis Tagawa & K. Iwatsuki, Acta Phytotax.Geobot. 22: 101. 1967; Christella siamensis (Tagawa & K.Iwatsuki) Holttum.Plants 70–100 cm tall. Rhizomes erect, apices along withstipe bases with dense linear-lanceolate brown scales. Frondsclustered; stipes 20–40 cm, stramineous, subglabrous or withsparse pale long acicular hairs distally; laminae 50–70 × 30–40cm, bases not narrowed or slightly so, apices acuminate; lateralpinnae 25–35 pairs, middle ones linear-lanceolate, 10–20 × 1.5–2 cm, bases broadly cuneate or truncate, sessile, lobed 2/3–4/5toward costae, apices long acuminate; segments 20–30 pairs onmiddle pinnae, oblong, 5–8 × ca. 3 mm, obtuse at apices, basalacroscopic ones usually longer and crenate; veinlets 6–12 pairs,proximal pair anastomosing or sometimes only connivent, distalveins ending before margin and above sinus. Laminae herbaceous,yellowish green when dried, with pale long acicular hairsalong rachises, costae, and veins on both surfaces. Sori orbicular,medial; indusia with dense pale acicular hairs. Sporangiabearing golden glands on stalks. Spores echinate or tuberculate.Wet places in forests; 1300–1400 m. ?Taiwan, S Yunnan (Jinping)[?NE India, Thailand].The character of veinlets not reaching the margin is unique in thisspecies of Cyclosorus but also occurs in Metathelypteris. The conniventveinlets resemble those of Pseudocyclosorus. The systematic position ofthis species requires more study.15. Cyclosorus procurrens (Mettenius) Copeland, Fern Fl.Philipp. 2: 340. 1960.无腺毛蕨 wu xian mao jueAspidium procurrens Mettenius, Ann. Mus. Bot. Lugduno-Batavi 1: 231. 1864; Cyclosorus kweichowensis Ching ex K.H. Shing; C. laui Ching; Dryopteris procurrens (Mettenius)Kuntze; Nephrodium procurrens (Mettenius) Baker; Thelypterisprocurrens (Mettenius) C. F. Reed.Plants 35–100 cm tall. Rhizomes shortly creeping, includingbases of stipes with sparse brown lanceolate scales.Fronds approximate; stipes 15–40 cm, brown at bases, distallydark stramineous; laminae 20–60 × 15–25 cm, bases not narrowedor slightly so, apices long acuminate; pinnae 15–25pairs, sessile, proximal 1 or 2 pairs sometimes slightlyshortened and reflexed; middle pinnae linear-lanceolate, 10–18× 1.5–2 cm, bases truncate, lobed ca. 2/3 toward costae, apiceslong acuminate; segments 20–30 pairs on middle pinnae, subfalcate-oblong,4–8 × 3–4 mm (basal acroscopic one longer),obtuse at apices; veinlets 6–10 pairs, proximal pair anastomosing,next 0.5–1 pairs running to sinus membrane. Laminaeherbaceous to papery, yellowish green to brownish green whendried, with sparse short acicular hairs adaxially, abaxial surfacewith dense pale acicular hairs. Sori orbicular, medial; indusiadensely hairy. Sporangia bearing reddish orange glands onstalks. Spores winged or otherwise ornamented.Semi-open places by forest margins, thickets; 300–1400 m.Guangdong, Guangxi, S Guizhou, Hainan, Taiwan, S Yunnan [S India,Indonesia, Malaysia, Myanmar, Philippines].Cyclosorus procurrens is quite similar to C. parasiticus and differsonly in being eglandular on the pinnae. But there are many intermediatespecimens. Holttum (Kew Bull. 31: 309. 1976) regarded C. procurrensas a synonym of C. parasiticus. This treatment seems reasonable butwe cannot confirm it. Based on similar morphology, these two speciesprobably belong to the same complex.16. Cyclosorus molliusculus (Wallich ex Kuhn) Ching, Bull.Fan Mem. Inst. Biol., Bot. 8: 196. 1938.美丽毛蕨 mei li mao jueAspidium molliusculum Wallich ex Kuhn, Bot. Zeitung(Berlin) 26: 41. 1868; Cyclosorus acutilobus Ching ex K. H.Shing; C. densissimus Ching ex K. H. Shing; Thelypteris molliuscula(Wallich ex Kuhn) K. Iwatsuki.Plants 40–100 cm tall. Rhizomes shortly to long creeping,including bases of stipes with dark brown lanceolate scales.Fronds approximate to distant; stipes 20–40 cm, stramineous;laminae 30–80 × 15–30 cm, bases not narrowed or slightly so,apices acuminate; lateral pinnae 10–20 pairs, sessile, proximal

THELYPTERIDACEAE 3791–3 pairs slightly shortened and reflexed; middle pinnae linearlanceolate,8–15 × 1–2.5 cm, bases truncate, lobed 2/3–4/5toward costae, apices long acuminate; segments 20–25 pairs,middle ones lanceolate, 5–11 × 3–4 mm (basal acroscopic onelonger and crenate), obtuse at apices; veinlets simple (occasionallyforked on basal segment), 6–12(–14) pairs, proximalpair anastomosing, next vein running to sinus membrane. Laminaeherbaceous, yellowish green when dried, both surfaces particularlyalong costae and veins with sparse slender acicularhairs, adaxially with several short acicular hairs between veins.Sori orbicular, supramedial, proximal pair usually confluent; indusiasmall, with acicular hairs. Sporangia without conspicuousglands. Spores brown, with long fimbriate wings.Semi-open places by forest margins, roadsides, thickets; 100–1600 m. Guangxi, S Guizhou, S Yunnan [India, Myanmar, Nepal, Thailand].According to Iwatsuki (Fl. Thailand 3: 424. 1988) and K. H.Shing (FRPS 4(1): 198. 1999), Cyclosorus molliusculus is distributedfrom SW China to Thailand. However, the present authors found thatthe specimens from S Yunnan, although similar in outline, are quitedifferent from those from SE Xizang, Nepal, and N India. It may bequestionable to use this name to represent the plants from Guangxi,Guizhou, and S Yunnan. As this problem cannot be resolved here, forthe purposes of this treatment, we temporarily follow the treatments ofShing (loc. cit.) and Lu (Fl. Yunnan. 20: 605. 2006).17. Cyclosorus cylindrothrix (Rosenstock) Ching, Bull. FanMem. Inst. Biol., Bot. 8: 199. 1938.柱腺毛蕨 zhu xian mao jueDryopteris cylindrothrix Rosenstock, Repert. Spec. Nov.Regni Veg. 12: 246. 1913; Christella cylindrothrix (Rosenstock)Holttum; Cyclosorus parasiticus (Linnaeus) Farwell var. cylindrothrix(Rosenstock) Tardieu & C. Christensen; Thelypteriscylindrothrix (Rosenstock) K. Iwatsuki.Plants 40–80 cm tall. Rhizomes shortly to long creeping.Fronds usually distant; stipes 20–40 cm, stramineous or reddish,proximal part with linear-lanceolate scales; laminae deltoid-lanceolate,30–40 × ca. 20 cm, bases not narrowed, apicesacuminate; lateral pinnae 15–25 pairs, sessile; proximal pinnaeslightly reflexed, not or slightly shortened, proximal severalpinnae narrowed toward their bases; middle pinnae linear-lanceolate,9–15 × 1.5–2.5 cm, slightly auriculate on acroscopicbases, bases truncate, lobed 2/3–3/4 toward costae, apices longacuminate; segments ca. 30 pairs on middle pinnae, 4–6 × 2–3mm, obtuse at apices; veinlets 6–10 pairs, proximal pair anastomosingwith a short excurrent veinlet running to sinusmembrane. Laminae herbaceous to papery, yellowish green orgrayish green when dried, abaxially with short acicular hairsand many golden thin clavate glands throughout, with shortacicular hairs along costae and veins and between veins adaxially.Sori orbicular, medial or supramedial; indusia with sparseshort hairs and golden clavate glands. Sporangia bearing similarglands on stalks. Spores with wings.SE Xizang (Mêdog) [Bhutan, N India, N Myanmar, Nepal, Thailand].This is a new record for China.Cyclosorus cylindrothrix has only one gathering from China, andthe rhizome is lacking. The description of rhizomes, stipe bases, andscales are according to Holttum (Kew Bull. 31: 308. 1976).18. Cyclosorus procerus (D. Don) S. Lindsay & D. J. Middleton,Nordic J. Bot. 30: 308. 2012.高毛蕨 gao mao jueNephrodium procerum D. Don, Prodr. Fl. Nepal. 6. 1825;Aspidium procerum (D. Don) Sprengel; Christella appendiculataHolttum; Cyclosorus appendiculatus Panigrahi (1993), not(Blume) Ching (1941); C. dulongjiangensis W. M. Chu; N. appendiculatumC. Presl (1851), not Schott (1834); Thelypterisprocera (D. Don) Fraser-Jenkins.Plants 60–120 cm tall. Rhizomes long creeping, includingstipe bases with brown lanceolate scales. Fronds distant; stipes20–40 cm, stramineous; laminae 40–90 × 20–30 cm, basesslightly narrowed, apices acuminate; lateral pinnae 20–30 pairs,proximal 2–5 pairs slightly shortened; middle pinnae lanceolate,8–18 × 1.5–4 cm, bases truncate, lobed 4/5 or more towardcostae, apices long acuminate; segments 20–30 pairs, lanceolate,10–15 × 2–3 mm, entire (basal acroscopic segment longerand crenate), pointed or acute at apices; veinlets 10–15 pairs,basal pair anastomosing. Laminae herbaceous, yellowish greenwhen dried, with sparse long pale acicular hairs along abaxialsurface of rachises and both sides of veins, and minute yellowglands along veins abaxially, glabrous between veins on bothsurfaces. Sori orbicular, submarginal; indusia very small, glabrousor with several long acicular hairs. Sporangia bearingsmall orange glands on stalks. Spores variously winged, tuberculate,or echinate.Semi-open places in sparse forests, roadsides; 1100–2400 m. SEXizang, W Yunnan [Bhutan, N India, Nepal, Thailand].K. H. Shing (FRPS 4(1): 202. 1999) mentioned that the type ofCyclosorus dulongjiangensis might be a hybrid and needed furtherresearch. We have examined the type in PYU, and the spores appearedwell developed; thus, the spores of this species do not support its putativehybrid status.19. Cyclosorus parasiticus (Linnaeus) Farwell, Amer. Midl.Naturalist 12: 259. 1931.华南毛蕨 hua nan mao juePolypodium parasiticum Linnaeus, Sp. Pl. 2: 1090. 1793;Aspidium parasiticum (Linnaeus) Swartz; Christella parasitica(Linnaeus) H. Léveillé; Cyclosorus aureoglandulosus Ching &K. H. Shing; C. damingshanensis Ching ex K. H. Shing; C.excelsior Ching & K. H. Shing; C. hainanensis Ching; C.orientalis Ching ex K. H. Shing; C. rupicola Ching & K. H.Shing; C. xunwuensis Ching ex K. H. Shing & J. F. Cheng; C.yandongensis Ching & K. H. Shing; Dryopteris parasitica(Linnaeus) Kuntze; Nephrodium parasiticum (Linnaeus) Desvaux;Thelypteris parasitica (Linnaeus) Tardieu.Plants (30–)50–70(–100) cm tall. Rhizomes shortly to longcreeping, including stipe bases with dark brown lanceolatescales. Fronds approximate to distant; stipes (10–)20–30(–40)cm, stramineous; laminae (20–)30–50(–60) × (8–)15–25(–35)cm, bases not narrowed (sometimes slightly narrowed), apices

380THELYPTERIDACEAEcaudate-acuminate; lateral pinnae 10–15(–20) pairs, proximal 1or 2 pairs reflexed; middle pinnae lanceolate, (5–)10–15(–20) ×(0.5–)1–1.5 cm, bases truncate, lobed 1/2–2/3 toward costae,apices long acuminate; segments 20–25 pairs, 3–4 × ca. 3 mm(basal acroscopic one longer, sometimes crenate), obtuse tosubacute at apices; veinlets 5–8 pairs, simple (ca. 10 pairs onbasal acroscopic segment, occasionally forked), proximal pairanastomosing, sometimes next vein running to sinus membrane.Laminae herbaceous, brownish green or yellowish green whendried, with thin acicular hairs throughout on both surfaces, andreddish orange glands throughout abaxially. Sori orbicular,medial; indusia densely hairy. Sporangia bearing reddish orangeglands on stalks. Spores with wings. 2n = 72, 108, 144.Semi-open places in thickets, roadsides; near sea level to 1900 m.Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan,Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [N India, Indonesia, Japan,Korea, Laos, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].The authors have not seen material of Cyclosorus parasiticus var.formosanus Ching (Bull. Fan Mem. Inst. Biol., Bot. 8: 205. 1938;Thelypteris parasitica var. formosana (Ching) C. F. Reed), describedfrom Taiwan.The type of Cyclosorus brevipes Ching ex K. H. Shing (FRPS4(1): 336. 1999) is partly bipinnate-pinnatifid with veins and glandssimilar to those of C. parasiticus. It has no sori and only one gatheringhas been collected. This taxon might be a hybrid.The type of Cyclosorus pauciserratus Ching & C. F. Zhang (Bull.Bot. Res., Harbin 3(3): 8. 1983) has no mature sori. Only one gatheringhas been made. This taxon might represent a young frond of C. parasiticus.The types of Cyclosorus contractus Ching ex K. H. Shing (FRPS4(1): 333. 1999) and C. parvilobus Ching ex K. H. Shing (FRPS 4(1):338. 1999) both have the features intermediate between C. acuminatusand C. parasiticus. Each species has only one gathering (the type) withno sori collected. These two taxa might be hybrids.Cyclosorus ×intermedius W. C. Shieh & J. L. Tsai (J. Sci. Engin.24: 8. 1987) is endemic to Taiwan. It is the putative hybrid between C.dentatus and C. parasiticus.20. Cyclosorus pygmaeus Ching & C. F. Zhang, Bull. Bot.Res., Harbin 3(3): 5. 1983.矮毛蕨 ai mao jueCyclosorus chengii Ching ex K. H. Shing & J. F. Cheng.Plants 15–40 cm tall. Rhizomes suberect, with brown lanceolatescales. Fronds clustered; stipes 5–15 cm, stramineous;laminae equal to stipes or slightly shorter in length, 5–15 × 3–7cm, bases not narrowed, apices acuminate; pinnae ca. 10 pairs,sessile, proximal pinnae lanceolate, 1.5–4 × 0.3–0.7 cm, basestruncate, lobed ca. 1/2 toward costae, apices acuminate; segmentsoblong, obtuse or subtruncate at apices; veinlets 4–6pairs per segment, proximal pair anastomosing, others to marginabove sinus. Laminae herbaceous when dry, yellowishgreen, with short acicular hairs on both surfaces, and manyglandular hairs abaxially. Sori small, submarginal, proximalpair usually confluent; indusia with short thin acicular hairs.Sporangia bearing reddish orange glands on stalks. Spores withshort wings.● Among rocks along streams, wet forest margins; below 100–800m. Jiangxi, Zhejiang.21. Cyclosorus hirtisorus (C. Christensen) Ching, Bull. FanMem. Inst. Biol., Bot. 8: 221. 1938.毛囊毛蕨 mao nang mao jueDryopteris hirtisora C. Christensen, Contr. U.S. Natl.Herb. 26: 277. 1931; D. hirticarpa Ching; Sphaerostephanoshirtisorus (C. Christensen) Holttum; Thelypteris hirtisora (C.Christensen) K. Iwatsuki.Plants 60–100 cm tall. Rhizomes long creeping, withovate-lanceolate dark brown scales. Fronds distant; stipes 20–50 cm, dark brown and sparsely scaly at bases, dark stramineousdistally; laminae 40–70 × 20–40 cm, bases slightly or notnarrowed, apices caudate with a large apical pinna; pinnae 10–15 pairs, stalked, proximal pair abruptly reduced to 2–5 cm orsometimes longer; middle pinnae linear, 10–25 × 1–2 cm, basesbroadly cuneate to truncate, lobed to 1/3 toward costae or onlyserrate, apices long acuminate; segments 30–45 pairs, triangularor oblong, 2–4 × ca. 4 mm, subacute or pointed at apices; veinlets7–10 pairs, quite oblique, proximal 1 or 2 pairs anastomosing,next 1.5–2.5 pairs running to sinus membrane. Laminaepapery, brownish green when dried, on both surfaces withacicular hairs along costae and veins, also with minute hairsbetween veins abaxially. Sori orbicular, medial; indusia denselyhairy. Sporangia bearing several hairs on capsules. Spores withfimbriate wings.Wet places in forests, semi-open forest margins; 500–1800 m.Guangxi, S Yunnan [Laos, Myanmar, Thailand].22. Cyclosorus attenuatus Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 341. 1999.下延毛蕨 xia yan mao jueCyclosorus mollissimus Ching ex K. H. Shing.Plants 50–100 cm tall. Rhizomes shortly creeping, includingstipe bases with lanceolate dark brown scales. Frondsapproximate; stipes 25–45 cm, stramineous; laminae 30–70 ×20–40 cm, bases not narrowed or slightly so, apices caudate;lateral pinnae 10–20 pairs, lanceolate, middle pinnae 8–20 ×2–3.5 cm, bases cuneate to truncate, apices long acuminate;proximal pinnae slightly shortened, shortly decurrent towardbases, shortly stalked; apical pinnae similar to lateral ones withseveral longer segments at bases; middle pinnae lobed to 1/3toward costae, sometimes only crenate; segments 25–40 pairs,falcate-triangular, 2–5 × ca. 3 mm, entire, obtuse or acute atapices; veinlets 6–10 pairs, proximal 2–5 pairs anastomosing,next 1–1.5 pairs running to sinus membrane. Laminae papery,grayish green when dried, with several acicular hairs alongcostae adaxially, abaxial surface with dense long acicular hairs,and short hairs between veins. Sori orbicular, medial; indusiadensely hairy. Sporangia bearing several hairs on capsules.Spores shortly and thickly cristate.

THELYPTERIDACEAE 381● Wet places in forests, forest margins; 300–1100 m. SE Yunnan.23. Cyclosorus articulatus (Houlston & T. Moore) Panigrahi,Res. J. Pl. Environ. 9: 66. 1993.节状毛蕨 jie zhuang mao jueNephrodium articulatum Houlston & T. Moore, Gard.Mag. Bot. 3: 293. 1851; Christella euphlebia (Ching) Holttum;Cyclosorus euphlebius Ching; C. laetestrigosus (C. B. Clarke)Ching; C. nigrescens Ching ex K. H. Shing; C. transitoriusChing ex K. H. Shing; C. yunnanensis Ching ex K. H. Shing; N.glandulosum J. Smith var. laetestrigosum C. B. Clarke; Pronephriumarticulatum (Houlston & T. Moore) Holttum; Thelypterisarticulata (Houlston & T. Moore) Tagawa & K. Iwatsuki;T. euphlebia (Ching) C. F. Reed.Plants 0.6–1.5 m tall. Rhizomes massive, shortly creeping,including stipe bases with brown ovate-lanceolate scales andbrown setae. Fronds clustered to approximate; stipes 20–60 cm,bases blackish brown, dark stramineous distally with setae;laminae 40–100 × 20–40 cm, bases slightly narrowed, apicesacuminate to caudate; lateral pinnae 10–25 pairs, proximal 1 or2 pairs slightly shortened, shortly stalked, bases cuneate anddecurrent; middle pinnae almost sessile, sterile pinnae 15–20 ×2–5 cm (fertile ones narrower), lanceolate, bases rounded-truncate,lobed to 1/4 toward costae or shallowly serrate, apicesacuminate or caudate; aerophores beneath pinna bases slightlyswollen; segments/serrations 25–40 pairs on middle pinnae, triangular,1–3 × ca. 3 mm, obtuse at apices; veinlets 6–12 pairs,proximal 2–6 pairs anastomosing with usually interrupted excurrentveinlet, next 1–3 pairs to prominent sinus membrane.Laminae papery, brownish green or reddish green when dried,adaxially with brown or pale setae along costae, abaxial surfacewith sparse setae along costae and veins (sometimes also betweenveins). Sori orbicular, medial or supramedial; indusiaglabrous. Sporangia bearing minute light yellow glands onstalks. Spores brown, echinate.Wet places in forests, forest margins; 200–1100 m. Guangxi, SGuizhou, SE Xizang, S Yunnan [India, Sri Lanka, Thailand, Vietnam].24. Cyclosorus papilio (C. Hope) Ching, Bull. Fan Mem. Inst.Biol., Bot. 8: 214. 1938.蝶状毛蕨 die zhuang mao jueNephrodium papilio C. Hope, J. Bombay Nat. Hist. Soc.12: 625. 1899; Christella papilio (C. Hope) Holttum; Dryopterispapilio (C. Hope) C. Christensen; Thelypteris papilio (C.Hope) K. Iwatsuki.Plants 60–120 cm tall. Rhizomes erect, including stipebases with brown ovate-lanceolate scales. Fronds clustered;stipes 10–20 cm, dark stramineous or reddish; laminae 40–110× 20–30 cm, bases gradually narrowed, apices acuminate tocaudate; lateral pinnae 30–40 pairs, sessile, proximal 5–10 pairsgradually shortened, triangular, often hastate with basal segmentelongate on both sides of rachises; middle pinnae linearlanceolate,10–18 × 1.5–2 cm, bases truncate, lobed 2/5–1/2toward costae, apices long acuminate; segments 20–25 pairs onmiddle pinnae, oblong, 3–5 × 3–4 mm (basal acroscopic oneslightly longer), entire, obtuse at apices; veinlets 5–8 pairs,proximal pair anastomosing, next 1–1.5 pairs running to sinusmembrane. Laminae papery, grayish green when dried, adaxiallywith sparse acicular hairs along costae and veins, abaxialsurface subglabrous. Sori orbicular, medial; indusia shortlyhairy. Sporangia bearing large reddish orange spherical glandson stalks. Spores echinate. 2n = 72, 216.Wet places in forests, on rocks in streams or beside streams; 500–2400 m. S Sichuan, Taiwan, SE Xizang, Yunnan [N India, Kashmir,Nepal, Sri Lanka].25. Cyclosorus evolutus (C. B. Clarke & Baker) Ching, Bull.Fan Mem. Inst. Biol., Bot. 8: 219. 1938.展羽毛蕨 zhan yu mao jueNephrodium amboinense C. Presl var. evolutum C. B.Clarke & Baker, J. Linn. Soc., Bot. 24: 417. 1888 [“evoluta”];Christella evoluta (C. B. Clarke & Baker) Holttum; Cyclosoruschingii Z. Y. Liu ex Ching & Z. Y. Liu; C. flaccidus Ching & Z.Y. Liu; Dryopteris evoluta (C. B. Clarke & Baker) C. Christensen;N. evolutum (C. B. Clarke & Baker) Beddome; Thelypterisevoluta (C. B. Clarke & Baker) Tagawa & K. Iwatsuki.Plants 60–120 cm tall. Rhizomes shortly to long creeping,including stipe bases with dark brown lanceolate scales. Stipes15–40 cm, dark stramineous; laminae 40–80 × 20–40 cm,bases abruptly narrowed, apices caudate with a large apicalpinna; lateral pinnae 10–20 pairs, proximal 1–5 pairs abruptlyshortened, proximal pair triangular-auriculate, ca. 3 × 2 cm orlonger, reflexed, sometimes hastate; middle pinnae lanceolate,12–20 × (1.5–)2–4 cm, bases truncate to rounded-truncate, sessileor subsessile, lobed 1/5–1/3 toward costae, sometimes onlycrenate, apices long acuminate or caudate; segments 20–30pairs per pinna on middle pinnae, ovate or oblong, 2–6 × 5–7mm, obtuse at apices; veinlets 7–9 pairs per segment, proximal1.5–3 pairs anastomosing, next 1–3 pairs running to sinus membrane.Laminae papery, grayish green when dried, subglabrousor glabrous on both surfaces (rarely shortly hairy). Sori orbicular,inframedial or medial; indusia glabrous or shortly hairy.Sporangia bearing large reddish orange spherical glands onstalks. Spores thickly cristate.Wet places in dense forests; 300–1600 m. Chongqing, Guangxi,Guizhou, Hunan, Yunnan [India, Thailand].There are several specimens of Cyclosorus evolutus with shorthairs on the abaxial surface; but other characteristics are similar to thetype, so no infraspecific name is given here.26. Cyclosorus latipinnus (Bentham) Tardieu, Notul. Syst.(Paris) 7: 73. 1938.宽羽毛蕨 kuan yu mao jueAspidium molle Swartz var. latipinnum Bentham, Fl.Hongk. 455. 1861; A. latipinnum (Bentham) Hance; Christellalatipinna (Bentham) H. Léveillé; Cyclosorus decipiens Ching;C. grossodentatus Ching ex K. H. Shing; C. nanpingensisChing; C. oblanceolatus K. H. Shing & C. F. Zhang; C. papilionaceusK. H. Shing & C. F. Zhang; C. paralatipinnus Ching exK. H. Shing; Dryopteris latipinna (Bentham) Kuntze; D. parasitica(Linnaeus) Kuntze var. latipinna (Bentham) C. Christensen;Nephrodium latipinnum (Bentham) Hooker ex Baker;Thelypteris latipinna (Bentham) K. Iwatsuki.

382THELYPTERIDACEAEPlants (15–)20–40(–60) cm tall. Rhizomes shortlycreeping, dark brown, apices and stipe bases with sparse brownlinear-lanceolate scales. Fronds approximate; stipes 5–20 cm;laminae 10–30 × 5–15 cm, bases narrowed, apices caudate witha large and deeper lobed apical pinna; lateral pinnae 4–10 pairs,proximal 1–4 pairs shortened with lowest pair ca. 1 × 0.5 cm,triangular-auriculate; middle pinnae lanceolate or oblanceolate,3–10 × 1.5–2 cm, bases rounded-truncate, lobed 1/4–1/3 towardcostae, sometimes subentire, apices acuminate or acute; segments10–20 pairs, 1–3 × ca. 3 mm, entire, obtuse at apices;veinlets 5 or 6 pairs, proximal pair anastomosing, next 0.5–1pair running to sinus membrane, excurrent veinlet usually interrupted.Laminae herbaceous, yellowish green when dried,shortly hairy along costae and veins adaxially, abaxial surfacewith very minute hairs and several reddish orange glands alongveins, also with capitate glandular hairs throughout. Sori orbicular,medial, proximal pair usually confluent; indusia shortlyhairy. Sporangia bearing reddish orange glands on stalks. Sporesbrown, large, irregularly small tuberculate and echinate.Wet or semi-open forest margins, beside streams; near sea level to1300 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, N andS Taiwan, S Yunnan, S Zhejiang [India, Indonesia, Malaysia, Myanmar,Philippines, Sri Lanka, Thailand, Vietnam; Australia, Pacific islands(Polynesia)].According to the author’s study, Cyclosorus latipinnus is a speciesof variable size, not always a small plant as described in FRPS (4(1):227. 1999), so many names are treated as synonyms here. Characterssuch as venation, capitate glandular hairs, and spore ornamentation aresimilar.Holttum (Kew Bull. 31: 324. 1976; Fl. Males., Ser. 2, 1. 1982)treated both Cyclosorus latipinnus and C. jaculosus as synonyms ofChristella subpubescens (Blume) Holttum (Aspidium subpubescensBlume; Cyclosorus subpubescens (Blume) Ching; Thelypteris subpubescens(Blume) K. Iwatsuki), which makes C. subpubescens acomplex and confused species. We found that C. jaculosus and C.latipinnus are two distinct species, so the application of the name C.subpubescens is questionable. Pending detailed studies of the type, wemaintain the use of the name C. latipinnus.27. Cyclosorus acuminatus (Houttuyn) Nakai in Thunberg,Misc. Pap. Japan. Pl. 15. 1935.渐尖毛蕨 jian jian mao juePlants (20–)40–60(–80) cm tall. Rhizomes long creeping,apices including stipe bases with brown lanceolate scales.Fronds distant; stipes (10–)20–30(–40) cm, stramineous tobrown; laminae (10–)30–50(–60) × 10–25 cm, bases not narrowedor slightly so, apices caudate to acuminate; lateral pinnae(5–)10–20 pairs, shortly stalked; middle pinnae linear-lanceolateto lanceolate, (5–)8–15(–18) × (0.5–)0.8–1.5(–2) cm, basestruncate or shallowly cordate, lobed 1/2–2/3 toward costae,apices acuminate to acute; segments (10–)15–30 pairs, 2–5 × 2–4 mm, basal acroscopic one (sometimes also basiscopic one)longer, sharply pointed or acute at apex; veinlets 6–10 pairs,simple (sometimes forked on basal acroscopic segment), proximalpair anastomosing, next (0.5–)1–1.5 pairs running to sinusmembrane. Laminae papery to subleathery, grayish green whendried, both surfaces with short acicular hairs along costae andveins, also with minute hairs throughout. Sori orbicular, submarginal;indusia shortly hairy or subglabrous. Sporangiabearing reddish orange glands on stalks. Spores dark brown,densely cristate. 2n = 72, 108, 144, 216.Semi-open places in thickets, grasslands, farmland margins, roadsides;near sea level to 1700 m. Anhui, Chongqing, Fujian, S Gansu,Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangsu,Jiangxi, S Shaanxi, Shandong, Sichuan, Taiwan, Yunnan, Zhejiang[Japan, Korea, Philippines].Two forms have been recognized from Taiwan: Cyclosorus acuminatusf. ensipinnus (Hayata) H. Itô (Bot. Mag. (Tokyo) 51: 712. 1937;Dryopteris sophoroides (Thunberg) Kuntze f. ensipinna Hayata, Icon.Pl. Formosan. 4: 180–181. 1914) and C. acuminatus f. pilosus H. Itô(Bot. Mag. (Tokyo) 51: 712. 1937).1a. Proximal pinnae not shortened,acuminate at apices ............................ 27a. var. acuminatus1b. Proximal several pinna pairs slightlyshortened, acute at apices ................ 27b. var. kuliangensis27a. Cyclosorus acuminatus var. acuminatus渐尖毛蕨 ( 原变种 ) jian jian mao jue (yuan bian zhong)Polypodium acuminatum Houttuyn, Nat. Hist. 2(14): 181.1783; Aspidium sophoroides (Thunberg) Swartz; Christellaacuminata (Houttuyn) Holttum; C. sophoroides (Thunberg) H.Léveillé; Cyclosorus cangnanensis K. H. Shing & C. F. Zhang;C. ciliensis K. H. Shing; C. dissitus Ching ex K. H. Shing; C.glabrescens Ching ex K. H. Shing; C. kuizhouensis K. H.Shing; C. nanchuanensis Ching & Z. Y. Liu; C. sinoacuminatusChing & Z. Y. Liu; C. sophoroides (Thunberg) Tardieu; C. subacuminatusChing ex K. H. Shing & J. F. Cheng; C. subcoriaceousChing ex K. H. Shing; Dryopteris acuminata (Houttuyn)Nakai; D. sinica Christ; D. sophoroides (Thunberg) Kuntze;Nephrodium sophoroides (Thunberg) Desvaux; P. sophoroidesThunberg; P. unitum Thunberg (1784), not Linnaeus (1759);Thelypteris acuminata (Houttuyn) C. V. Morton.Proximal pinnae usually not shortened, acuminate atapices; laminae with usually 2 pairs of veinlets beneath sinus.Semi-open places in thickets, grasslands, farmland margins, roadsides;near sea level to 1700 m. Anhui, Chongqing, Fujian, S Gansu,Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangsu,Jiangxi, S Shaanxi, Shandong, Sichuan, Taiwan, Yunnan, Zhejiang[Japan, Korea, Philippines].The type of Cyclosorus abbreviatus Ching & K. H. Shing ex K.H. Shing & J. F. Cheng (Jiangxi Sci. 8(3): 45. 1990) has features intermediatebetween C. acuminatus var. acuminatus and C. aridus. Thistaxon might be a hybrid.The type of Cyclosorus yuanjiangensis Ching ex K. H. Shing(FRPS 4(1): 335. 1999) has features intermediate between C. acuminatusvar. acuminatus and C. procurrens. Only one gathering withrare mature sori has been collected. This taxon might be a hybrid.The type of Cyclosorus zhangii K. H. Shing (FRPS 4(1): 340.1999) has features intermediate between C. acuminatus var. acuminatusand C. parasiticus. Only one gathering (the type) without mature sorihas been collected. This taxon might be a hybrid.The type of Cyclosorus tarningensis Ching (Wuyi Sci. J. 1: 4.1981) might be an irregular plant of C. acuminatus var. acuminatus. Thespores of the two species appear similar.

THELYPTERIDACEAE 383The type of Cyclosorus nanchuanensis Ching & Z. Y. Liu (Bull.Bot. Res., Harbin 4(3): 14. 1984) has not been examined by us, but itseems quite similar to C. acuminatus var. acuminatus according to thephoto. Thus, this taxon is synonymized here, pending further study.27b. Cyclosorus acuminatus var. kuliangensis Ching, Bull.Fan Mem. Inst. Biol., Bot. 8: 192. 1938.鼓岭渐尖毛蕨 gu ling jian jian mao jueChristella acuminata monstr. kuliangensis (Ching) Nakaike;C. acuminata var. kuliangensis (Ching) C. M. Kuo;Cyclosorus kuliangensis (Ching) K. H. Shing; Thelypteris acuminatavar. kuliangensis (Ching) K. Iwatsuki.Pinnae usually 4–8 pairs, oblanceolate; proximal severalpinnae enlarged at apices and abruptly acute; veinlets usually1.5 pairs below sinuses.● Semi-open places in thickets, grasslands, farmland margins,roadsides; near sea level to 1200 m. Anhui, Fujian, Guangdong, Guangxi,Hunan, Jiangxi, Yunnan, Zhejiang.The types of Cyclosorus paracuminatus Ching ex K. H. Shing &J. F. Cheng (Jiangxi Sci. 8(3): 46. 1990) resemble this variety in generaloutline but have orange glands abaxially and lack sori. Only onegathering (the type) without sori has been collected. This taxon might bea hybrid.Cyclosorus acuminatus var. acuminatoides W. C. Shieh & J. L.Tsai (J. Sci. Engin. 24: 8. 1987) is endemic to Taiwan. This variety is aprobable hybrid between C. acuminatus var. acuminatus and C. acuminatusvar. kuliangensis.28. Cyclosorus nanxiensis Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 343. 1999.南溪毛蕨 nan xi mao juePlants 40–100 cm tall. Rhizomes shortly creeping, includingstipe bases with dense dark brown linear-lanceolatescales. Fronds approximate; stipes 15–30 cm, dark stramineousor reddish; laminae 25–70 × 20–30 cm, proximally slightly narrowed,apices acuminate to caudate; lateral pinnae 10–18 pairs,subsessile, proximal 1 or 2 pairs shortened; middle pinnae 5–15× 1–2 cm, sterile ones usually broader, lanceolate, bases subtruncateor broadly cuneate, lobed to 1/2 toward costae or onlyserrate, apices long acuminate; segments 15–25 pairs on middlepinnae, oblong, 2–5 × ca. 3 mm, subacute or obtuse at apices;veinlets 5–8 pairs, proximal 1.5–2.5 pairs anastomosing, next1–1.5 pairs running to sinus membrane. Laminae papery,brownish green or grayish green when dried, adaxially shortlyhairy along costae and veins, abaxially with sparse short hairsalong costae and veins, with minute hairs throughout. Sori orbicular,medial, proximal pair sometimes confluent; indusiasubglabrous or minutely hairy. Sporangia bearing orange glandson stalks. Spores shortly cristate or echinate.● Wet places in forests; 100–700 m. SE Yunnan (Hekou).29. Cyclosorus scaberulus Ching, Bull. Fan Mem. Inst. Biol.,Bot. 8: 223. 1938.糙叶毛蕨 cao ye mao jueChristella scaberula (Ching) Holttum; Thelypteris scaberula(Ching) C. F. Reed.Plants 70–100 cm tall. Rhizomes shortly creeping, withdark brown linear-lanceolate scales. Fronds approximate; stipes30–45 cm, dark stramineous, subglabrous or with sparse scalesto middle of rachises; laminae 40–60 × 15–30 cm, bases notnarrowed or slightly narrowed, apices caudate to acuminate;pinnae 10–20 pairs, subsessile; middle pinnae 8–15 × 1.5–2.5cm, lobed 1/3–2/5 toward costae, lanceolate, bases roundedtruncate,apices long acuminate; segments 20–35 pairs on middlepinnae, triangular, 3–4 × 2–4 mm, obtuse or acute at apices;veinlets 8–11 pairs, proximal 2 pairs anastomosing, next 1–1.5 pairs running to sinus membrane. Laminae papery, grayishgreen to brownish green when dried, adaxially with severalacicular hairs along costae and veins, also with minute hairsbetween veins, abaxial surface with minute hairs throughout(sometimes with several scales on costae). Sori orbicular, medialto supramedial; indusia glabrous or shortly hairy. Sporangiabearing reddish orange glands on stalks. Spores echinate orshortly cristate. 2n = 72.● Nearby streams, wet places in dense forests; 700–1200 m.Hainan.30. Cyclosorus calvescens Ching, Bull. Fan Mem. Inst. Biol.,Bot. 8: 225. 1938.光羽毛蕨 guang yu mao jueChristella calvescens (Ching) Holttum; Cyclosorus sanduensisK. H. Shing & P. S. Wang; Thelypteris calvescens (Ching)C. F. Reed.Plants 60–100 cm tall. Rhizomes shortly creeping, includingstipe bases with dark brown lanceolate scales. Frondsapproximate; stipes 20–50 cm, dark stramineous; laminae ovatelanceolate,40–60 × 20–35 cm, bases not narrowed or slightlyso, apices caudate with a long apical pinna; lateral pinnae 10–15 pairs, middle ones 15–20 × 2–3 cm, bases cuneate (decurrenton proximal pinnae), shortly stalked, lobed 1/4–1/3 towardcostae, apices long acuminate; proximal 1 or 2 pairs shortenedto less than 1/2 length of middle pinnae or sometimes longer;segments 20–30 pairs on middle pinnae, triangular, entire, obtuseor subacute at apices; veinlets 8–10 pairs, proximal 1.5–2pairs anastomosing, next 1.5–2.5 pairs running to sinus membrane.Laminae papery, grayish green or brownish greenwhen dried, adaxially with sparse acicular hairs along costaeand veins, abaxial surface subglabrous. Sori orbicular or elongate,medial; indusia subglabrous. Sporangia bearing orange orgolden glands on stalks. Spores irregularly cristate or echinate.Wet places in forests; 300–1600 m. Guangxi, S Guizhou, S Yunnan[Vietnam].31. Cyclosorus hokouensis Ching, Bull. Fan Mem. Inst. Biol.,n.s., 1: 289. 1949.河口毛蕨 he kou mao jueChristella hokouensis (Ching) Holttum; Thelypteris hokouensis(Ching) C. F. Reed.Plants 0.8–1.3 m tall. Rhizomes long creeping, darkbrown, including stipe bases with sparse dark brown lanceolatescales and brown setae. Fronds distant; stipes 10–15 cm, darkstramineous; laminae 70–120 × 20–35 cm, bases abruptly nar-

384THELYPTERIDACEAErowed, apices caudate; pinnae 20–35 pairs, proximal 5–15 pairsabruptly reduced into hastate pinnae with caudate apices,proximal pair ca. 1 × 0.5 cm; middle pinnae linear-lanceolate,10–18 × 1–2 cm, bases truncate, lobed 1/4–1/3 toward costae,apices long acuminate; segments 20–30 pairs on middle pinnae,oblong, 2–5 × ca. 3 mm, entire, subacute at apices; veinlets 8–11 pairs, proximal 2 or 3 pairs anastomosing, next 1–1.5 pairsrunning to sinus membrane. Laminae papery, brownish greenwhen dried, with dense acicular hairs along costae on both surfaces,with several acicular hairs along veins adaxially, withdense acicular hairs throughout and several capitate glandularhairs abaxially. Sori orbicular, medial; indusia hairy. Sporangiawith small reddish orange glands on stalks. Spores with shortwings.Semi-open forests, wet places in forests, forest margins; 100–1100m. SW Guangxi, SE Yunnan [E India].The type of Cyclosorus caii Ching ex K. H. Shing (FRPS 4(1):346. 1999) has features intermediate between C. hokouensis and C.aridus. Only one gathering (3 sheets of the type) with rare fertile sorihas been collected. This taxon might be a hybrid.32. Cyclosorus gustavii (Beddome) Ching, Bull. Fan Mem.Inst. Biol., Bot. 8: 215. 1938.古斯塔毛蕨 gu si ta mao jueNephrodium gustavii Beddome, J. Bot. 31: 227. 1893;Christella gustavii (Beddome) Holttum.Plants 50–100 cm tall. Rhizomes long creeping, includingstipe bases with linear-lanceolate dark brown scales, similarscales on stipe bases. Fronds distant; stipes 15–25 cm, darkstramineous; laminae 40–80 × 25–35 cm, bases gradually narrowed,apices caudate with a large apical pinna; lateral pinnae10–15 pairs, lanceolate, 12–18 × 2–3 cm, lobed ca. 1/3 towardcostae, bases truncate to broadly cuneate, almost sessile, apicesacuminate; apical pinnae similar to lateral ones; proximal 3–5pairs of lateral pinnae gradually shortened, lowest pair ca. 2 cm;segments 20–30 pairs on middle pinnae, 3–4 × 3–4 mm, entire,obtuse at apices; veinlets 6–12 pairs, proximal 2 or 3 pairsanastomosing, next 1–1.5 pairs running to sinus membrane.Laminae papery, grayish green when dried, with several acicularhairs along costae and veins adaxially, with short acicularhairs along veins and minute hairs between veins abaxially. Soriorbicular, medial; indusia shortly hairy. Sporangia lacking conspicuousglands. Spores brown with long fimbriate wings.Wet places in forests; 600–1000 m. W Yunnan [India, Thailand].Cyclosorus gustavii has only two gatherings from W Yunnan, soour knowledge about it is quite limited. The distinction between thisspecies and C. assamicus (Beddome) Ching, which also occurs in NEIndia, is not obvious. Here, we follow the treatment in Fl. Yunnan. (20:613. 2006).33. Cyclosorus wulingshanensis C. M. Zhang, Keys Vasc.Pl. Wuling Mts. 567. 1995.武陵毛蕨 wu ling mao jueCyclosorus leipoensis Ching & H. S. Kung ex K. H.Shing.Plants 70–120 cm tall. Rhizomes long creeping, with lanceolatedark brown scales and setae. Stipes 10–30 cm, darkstramineous; laminae 60–100 × 15–30 cm, bases gradually narrowed,apices acuminate to caudate; lateral pinnae 15–25 pairs,proximal 3–6 pairs gradually shortened, sometimes hastate withlonger basal acroscopic segments, lowest pair 1.5–3 × ca. 2 cm;middle pinnae lanceolate, 10–18 × 1.5–2.5 cm, bases roundedtruncate,lobed 1/4–2/5 toward costae, apices long acuminate;segments 25–35 pairs on middle pinnae, 2–5 × ca. 3 mm (basalacroscopic segment longer), obtuse at apices; veinlets 6–9 pairs,proximal 1.5–2 pairs anastomosing, next 0.5–1 pair running tosinus membrane. Laminae papery, grayish green when dried,adaxially with sparse short acicular hairs along costae (occasionallyalong veins), abaxial surface subglabrous except forseveral short acicular hairs on costae. Sori orbicular, medial;indusia subglabrous. Sporangia bearing golden to orange glandson stalks. Spores thickly cristate.● Wet places in forests, forest margins; 300–2100 m. Chongqing,Guangxi, W Hunan, S Sichuan, SE Xizang, Yunnan.34. Cyclosorus jinghongensis Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 337. 1999.景洪毛蕨 jing hong mao jueCyclosorus baiseensis Ching ex K. H. Shing; C. oppositusChing ex K. H. Shing; C. pumilus Ching ex K. H. Shing.Plants 30–80 cm tall. Rhizomes erect to suberect, apicesincluding stipe bases with dark brown lanceolate scales. Frondsclustered; stipes 5–20 cm, stramineous; laminae 25–60 × 8–18cm, bases gradually narrowed, apices acuminate to caudate;pinnae 10–20 pairs, proximal 3–5 pairs gradually shortened,proximal pair ca. 1 × 1 cm; middle pinnae lanceolate, 5–10 × 1–2 cm, bases subtruncate, lobed ca. 1/3 toward costae, apicesacuminate; segments 15–20 pairs, subsquare, 2–4 × ca. 3 mm,entire, obtuse at apices; veinlets 5–7 pairs, proximal 1–1.5 pairsanastomosing, next 1–1.5 pairs running to sinus membrane.Laminae herbaceous, grayish green when dried, veins withsparse short hairs on both surfaces, abaxial surface with reddishorange glands along and between veins. Sori orbicular, medial;indusia shortly hairy. Sporangia bearing reddish orange glandson stalks. Spores thinly cristate.Wet places in forests, semi-open places by forest margins; 200–1000 m. Guangxi, Hainan, S Yunnan [Thailand, Vietnam].35. Cyclosorus jaculosus (Christ) H. Itô, Bot. Mag. (Tokyo)51: 725. 1937.闽台毛蕨 min tai mao jueAspidium jaculosum Christ, Bull. Herb. Boissier, sér. 2, 4:615. 1904; Christella jaculosa (Christ) Holttum; C. subarida(Tatewaki & Tagawa) Holttum ex C. M. Kuo; Cyclosorusaureoglandulifer Ching ex K. H. Shing; C. houi Ching; C.pararidus Ching ex K. H. Shing; C. simillimus Ching ex K. H.Shing; C. sparsisorus Ching ex K. H. Shing; C. subaridus Tatewaki& Tagawa; Dryopteris jaculosa (Christ) C. Christensen;Nephrodium jaculosum (Christ) Hayata; Thelypteris houi(Ching) C. F. Reed; T. jaculosa (Christ) Panigrahi; T. subarida(Tatewaki & Tagawa) C. F. Reed.

THELYPTERIDACEAE 385Plants 40–100 cm tall. Rhizomes long creeping, darkbrown, including stipe bases with dark brown lanceolate scales.Fronds distant; stipes 5–30 cm, dark stramineous; laminae 30–90 × 10–30 cm, bases gradually narrowed, apices caudate toacuminate; pinnae 20–30 pairs, proximal 3–8 pairs graduallyshortened with lowest pair ca. 1.5 × 1.5 cm; middle pinnaelanceolate, 6–18 × 1–2 cm, bases rounded-truncate, lobed 1/3–1/2 toward costae, apices long acuminate; segments 15–30pairs, triangular-oblong, 3–5 × ca. 3 mm, margins sometimescrenate on proximal ones, rounded-obtuse to subacute at apices;veinlets 5–8 pairs, proximal 1–1.5 pairs anastomosing, next1–1.5 pairs running to sinus membrane. Laminae papery,brownish green to grayish green when dried, adaxially withseveral acicular hairs along costae and veins and minute hairsbetween veins, abaxial surface with dense minute hairs andmany reddish orange glands throughout. Sori orbicular, medial;indusia glabrous or with sparse minute hairs. Sporangia bearingreddish orange glands on stalks. Spores shortly cristate andechinate. 2n = 72.Streamsides, wet places in forests, semi-open places; near sealevel to 1200 m. Fujian, Guangdong, Guangxi, Guizhou, Hunan, SJiangxi, Taiwan, Yunnan, S Zhejiang [Bhutan, India, Japan, Nepal, Vietnam].36. Cyclosorus ensifer (Tagawa) W. C. Shieh, J. Sci. Engin.13: 42. 1976.广叶毛蕨 guang ye mao jueDryopteris ensifera Tagawa, Acta Phytotax. Geobot. 6: 89.1937; Christella ensifera (Tagawa) Holttum ex C. M. Kuo;Cyclosorus gaoxiongensis Ching ex K. H. Shing; C. subnamburensisChing ex K. H. Shing; Thelypteris ensifera (Tagawa)K. Iwatsuki.Plants 25–40 cm tall. Rhizomes shortly to long creeping,with dark brown linear-lanceolate scales. Fronds approximate todistant; stipes 10–15 cm, dark stramineous; laminae 15–30 × 5–10 cm, bases slightly narrowed, apices caudate; lateral pinnae4–10 pairs, proximal 1 or 2 pairs shortened, ca. 1 cm or longer;middle pinnae oblanceolate, 3–10 × 1–2 cm, bases truncate toshallowly cordate, lobed 1/3–1/2 toward costae, apices acuminate;segments 8–20 pairs, 3–8 × ca. 3 mm (basal acroscopicone longer and crenate), acute at apices; veinlets 6–10 pairs,simple (sometimes forked on longer segments), proximal 2 pairsanastomosing, next pair running to sinus membrane. Laminaepapery, adaxially with several acicular hairs along costae andveins, also with dense minute hairs along and between veins,abaxial surface with several acicular hairs and several clavateglands along veins, and dense minute hairs along and betweenveins. Sori orbicular, medial; indusia shortly hairy. Sporangiabearing clavate glands on stalks. Spores cristate. 2n = 72.● Wet places in forests; near sea level to 700 m. S and SE Taiwan.37. Cyclosorus aridus (D. Don) Ching, Bull. Fan Mem. Inst.Biol., Bot. 8: 194. 1938.干旱毛蕨 gan han mao jueAspidium aridum D. Don, Prodr. Fl. Nepal. 4. 1825;Christella arida (D. Don) Holttum; Cyclosorus acutissimusChing ex K. H. Shing & J. F. Cheng; C. serrifer Ching ex K. H.Shing; Dryopteris arida (D. Don) Kuntze; Nephrodium aridum(D. Don) J. Smith; Thelypteris arida (D. Don) C. V. Morton.Plants (30–)50–100(–150) cm tall. Rhizomes longcreeping, including stipe bases with sparse brown lanceolatescales. Fronds distant; stipes 10–35 cm; laminae (20–)40–80(–120) × (10–)15–35 cm, bases abruptly or gradually narrowed,apices caudate to acuminate; pinnae 15–40 pairs, proximal2–10 pairs shortened; middle pinnae linear-lanceolate,(5–)10–18 × 1–2 cm, bases truncate, lobed to 1/3 toward costaeor sometimes only dentate, apices long acuminate; segments20–40 pairs on middle pinnae, triangular, 1–3 × ca. 3 mm,entire, acute or pointed at apices; veinlets 6–12 pairs, muchoblique, proximal 2 or 3 pairs anastomosing, next 1 or 2 pairsrunning to sinus membrane. Laminae papery to somewhatleathery, brownish green or yellowish green when dried, adaxiallysubglabrous except for several short acicular hairs alongcostae, abaxial surface with short acicular hairs along costaeand veins, also with yellow or orange clavate glands alongveins. Sori orbicular, medial; indusia glandular, sometimeshairy. Sporangia bearing yellow or orange clavate glands onstalks. Spore with long wings or ridged folds. 2n = 72.Wet or semi-open places, usually among tall grasses; near sealevel to 2500 m. S Anhui, Chongqing, Fujian, Guangdong, Guangxi,Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Taiwan, SE Xizang, Yunnan,Zhejiang [Bhutan, India, Indonesia, Kashmir, Malesia, Nepal,Philippines, Vietnam; Australia, Pacific islands].K. H. Shing mentioned in FRPS (4(1): 278. 1999) that Cyclosorussubaridus represented a small form of C. aridus. However, after carefulexamination, we found that C. subaridus resembles the type of C.jaculosus, and it is here included in synonymy of that species.The type of Cyclosorus medogensis Ching & S. K. Wu (Fl.Xizang. 1: 171. 1983) resembles this species but differs by deeper lobedsegments and less anastomosing veins. Its type has quite rare fertilesori. Only one gathering (the type) has been collected. This taxonmight be a hybrid.The type of Cyclosorus omeigensis Ching (Bull. Fan Mem. Inst.Biol., n.s., 1: 289. 1949; Christella omeigensis (Ching) Holttum; Thelypterisomeigensis (Ching) C. F. Reed) has features intermediate betweenCyclosorus aridus and Pronephrium penangianum. Only one gathering(the type) lacking fertile sori has been collected. This taxon might be ahybrid.The type of Cyclosorus pseudoaridus Ching ex K. H. Shing(FRPS 4(1): 278. 1999) has features intermediate between C. aridus andC. acuminatus. Only one gathering (the type) with rare fertile sori hasbeen collected. This taxon might be a hybrid.38. Cyclosorus cuneatus Ching ex K. H. Shing, Fl. Reipubl.Popularis Sin. 4(1): 347. 1999.狭基毛蕨 xia ji mao jueCyclosorus clavatus K. H. Shing.Plants 0.8–1.5 m tall. Rhizomes long creeping, includingstipe bases with sparse dark brown lanceolate scales and brownshort setae. Fronds distant; stipes 30–60 cm, dark stramineous;laminae 50–90 × 30–40 cm, bases not or slightly narrowed,apices caudate with a large apical pinna; lateral pinnae 4–12

386THELYPTERIDACEAEpairs, proximal ones stalked; proximal pair sometimes shortenedto ca. 3 cm; middle pinnae lanceolate, 15–22 × 2.5–3.5 cm,bases cuneate, lobed 1/4–1/3 toward costae, apices long acuminate;segments 30–40 pairs on middle pinnae, falcate-triangular,3–5 × 4–5 mm, obtuse or subacute at apices; veinlets 8–12pairs, proximal 2 pairs anastomosing with usually an interruptedexcurrent veinlet, next 2 or 3 pairs running to sinusmembrane. Laminae papery, grayish green when dried, acicularhairy along costae (sometimes also veins) adaxially, abaxiallywith golden clavate glands and minute hairs along veins. Soriorbicular or slightly elongate, medial or inframedial; indusiawith sparse short hairs. Sporangia bearing clavate glands onstalks. Spores with fimbriate wings.Wet places in dense forests; 600–700 m. Chongqing, Guangxi,Guizhou [Vietnam].39. Cyclosorus fukienensis Ching, Bull. Fan Mem. Inst. Biol.,Bot. 8: 209. 1938.福建毛蕨 fu jian mao jueChristella fukienensis (Ching) Holttum; Cyclosorus dehuaensisChing & K. H. Shing; C. fraxinifolius Ching & K. H.Shing; C. luoqingensis Ching & C. F. Zhang; C. nanlingensisChing ex K. H. Shing & J. F. Cheng; C. paucipinnus Ching &C. F. Zhang ex K. H. Shing; Thelypteris fukienensis (Ching) C.F. Reed.Plants 40–100 cm tall. Rhizomes long creeping, includingstipe bases with dark brown lanceolate scales and setae. Frondsdistant; stipes 10–30 cm; laminae 30–80 × 15–30 cm, basesgradually narrowed or slightly so, apices caudate with amedium-sized to large apical pinna; pinnae 5–15 pairs, shortlystalked or almost sessile, proximal 1–5 pairs graduallyshortened, or sometimes only 1 pair slightly shortened; middlepinnae lanceolate, 10–18 × 1.5–3 cm, bases truncate or broadlycuneate, lobed 1/5–1/3 toward costae, apices acuminate; segments20–30 pairs on middle pinnae, 2–4 × 3–5 mm, obtuse orsubtruncate at apices; veinlets 5–10 pairs, proximal 2 pairsanastomosing, next 1.5–2 pairs running to sinus membrane.Laminae papery, grayish green or brownish green when dried,with short acicular hairs along costae and veins and minutehairs between veins on both surfaces, also with golden clavateglands along veins abaxially. Sori orbicular, medial; indusiashortly hairy, sometimes glandular. Sporangia bearing golden toreddish orange clavate glands on stalks. Spores with short fimbriatewings.● Wet places in forests, forest margins; near sea level to 1100 m.Fujian, N Guangdong, S Hunan, S Jiangxi, S Zhejiang.The type of Cyclosorus grandissimus Ching & K. H. Shing (Fl.Fujian. 1: 599. 1982) has features intermediate between C. fukienensisand C. aridus. Only one gathering (the type) with rare sori has beencollected. This taxon might be a hybrid.The type of Cyclosorus hirtipes K. H. Shing & C. F. Zhang (FRPS4(1): 345–346. 1999) has features intermediate between C. fukienensisand C. acuminatus. Only one gathering (the type) with rare fertile sorihas been collected. This taxon might be a hybrid.40. Cyclosorus subelatus (Baker) Ching, Bull. Fan Mem. Inst.Biol., Bot. 8: 224. 1938.巨型毛蕨 ju xing mao jueNephrodium subelatum Baker, Bull. Misc. Inform. Kew1906: 11. 1906; Christella subelata (Baker) Holttum; Dryopterissubelata (Baker) C. Christensen; Thelypteris subelata(Baker) K. Iwatsuki.Plants 0.6–1.5 m tall. Rhizomes shortly or long creeping,massive and woody, including stipe bases with dark brownlanceolate scales. Fronds approximate or distant; stipes 10–30cm, bases dark brown, stramineous distally; laminae 50–120 ×30–40 cm, bases narrowed, apices caudate; pinnae 15–30 pairs,shortly stalked or almost sessile, proximal 2–10 pairs graduallyshortened and narrowed, proximal pair ca. 1 cm; middle pinnaelanceolate, 10–20 × 2–3.5 cm, bases rounded-truncate, lobed1/4–1/3 toward costae, apices long acuminate; segments 20–35pairs on middle pinnae, oblong, 2–5 × 3–5 mm, entire, obtuseto subacute at apices; veinlets 8–12 pairs, proximal 1 or 2 pairsanastomosing, next 1–3 pairs running to sinus membrane.Laminae papery, brownish green when dried, with short acicularhairs along costae and veins on both surfaces, also withclavate glands along veins and also minute hairs between veinsabaxially. Sori orbicular, medial or supramedial; indusia shortlyhairy. Sporangia bearing orange to golden clavate glands onstalks. Spores with medium-sized to long fimbriate wings.Wet places in forests, semi-open forest margins; 200–1500 m.Guangxi, S Yunnan [India, Laos, Myanmar, N Thailand].The type of Cyclosorus elatus Ching ex K. H. Shing (FRPS 4(1):342. 1999), not (Mettenius ex Kuhn) Alston (1956), has features intermediatebetween C. subelatus and C. dentatus. Only one gathering (thetype) with mostly sterile sori has been collected. This taxon might be ahybrid.15. STEGNOGRAMMA Blume, Enum. Pl. Javae 2: 172. 1828.溪边蕨属 xi bian jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants medium-sized, on soil. Rhizomes short, erect or ascending, with dense unicellular or multicellular long hairs and sparsebrown setaceous lanceolate scales. Fronds clustered; stipes dark brown, bases scaly and with grayish white unicellular or multiseparatedacicular long hairs; laminae oblong-lanceolate or broadly lanceolate, not narrowed to bases or slightly so, 1-pinnate, pinnatifid-acuminateat apices; pinnae ca. 10 pairs, spreading, sessile or very shortly stalked on proximal several pairs, distal ones ±adnate to rachises, lanceolate or ovate-lanceolate, rounded-cuneate or truncate at bases, subopposite, undulate or crenate along margins,or lobed to 1/3 of distance to costule, shortly acute or rounded-obtuse at apices, sometimes acuminate, both surfaces ± withhairs, costae obviously raised abaxially, grooved adaxially, with dense acicular hairs. Veins goniopteroid, 3–5 pairs of veinlets per

THELYPTERIDACEAE 387segment, proximal 1–3(–5) pairs joining to form triangular areoles, proximal pair arising above bases of costules, distally veins allreaching margin above sinuses. Laminae herbaceous, dark brown-green when dry, abaxially with grayish white acicular hairs, adaxiallysparsely setaceous; usually with multicellular long hairs and short hairs along rachises. Sori linear, attached along veins, exindusiateor sometimes indusiate; sporangia bearing erect short hairs, sometimes glabrous; spores bilateral, elliptic, sharply echinate.More than ten species: SW China, India, Malesia, Myanmar, Vietnam; six species (five endemic) in China.1a. Veinlets 3 or 4 pairs, joined on both sides of costules and forming 2 lines of tetragonal areoles ....................... 1. S. dictyoclinoides1b. Veinlets 1 or 2 pairs joined on both sides of costules and forming 1–3 triangular or irregular areoles.2a. Sporangia glabrous ........................................................................................................................................... 2. S. xingwenensis2b. Sporangia bearing short hairs.3a. Laminae with many short setae remaining at attachments of sori after sori fallen ................................... 3. S. cyrtomioides3b. Laminae with remaining glandlike objects at attachments of sori after sori fallen.4a. Pinnae broadly ovate-lanceolate, margins lobed to 1/3 of distance to costule, bases truncate,apices shortly acute, proximal 3 or 4 pairs of pinnae shortened and reflexed; endemic toYunnan (Gongshan) .................................................................................................................................... 4. S. latipinna4b. Pinnae lanceolate, acuminate at apices, or ovate-lanceolate, rounded-obtuse, margins undulateor lobate, bases rounded-cuneate, proximal pinnae not or slightly shortened.5a. Proximal pinnae slightly shortened, lanceolate, broadest on mid to lower part, acuminateand bent distally, glabrous on intercostal areas of abaxial surface of pinnae; endemicto Chongqing (Jinyun Shan) ........................................................................................................... 5. S. diplazioides5b. Proximal several pinnae shortened, narrowly ovate-lanceolate, broadest at bases,rounded-obtuse at apices, setaceous on intercostal areas of abaxial surface of lamina;Chongqing, Sichuan, Yunnan ..................................................................................................... 6. S. jinfoshanensis1. Stegnogramma dictyoclinoides Ching, Sinensia 7: 92. 1936.屏边溪边蕨 ping bian xi bian jueCyclosorus dictyoclinoides (Ching) C. M. Kuo; Thelypterisdictyoclinoides (Ching) C. M. Kuo.Plants ca. 50 cm tall. Rhizomes short and thick, ascending,almost scaly. Fronds clustered; stipes 15–23 cm, dark stramineous,glabrous on bases, throughout with spreading grayishwhite acicular hairs and short hairs; laminae broadly lanceolate,15–25 × 7–10 cm, 1-pinnate, pinnatifid-acuminate at apices;pinnae 7 or 8 pairs, spreading, proximal 1 or 2 pairs free, sessile,slightly shortened, distal ones adnate to rachises and connectedby narrow wings, apical ones connected by broad wings;proximal pinnae oblong-lanceolate, 3–4 × ca. 1.5 cm, basestruncate or rounded-cuneate, symmetrical, margins crenate, apicesshortly acuminate; middle pinnae of similar shape as proximalones, ca. 5 × 2 cm, bases adnate to rachises; costae raisedon both sides. Veins evident, 3 or 4 pairs of veinlets joining intoareoles, areoles in 2 lines, 4 or 5 per line, subsquare or pentagonaland with an excurrent veinlet arising from joining point ±tortuous, proximal pair of veinlets arising from far above basesof costules. Laminae herbaceous, dark brown-green when dry,both surfaces with dense grayish white acicular long hairs alongrachises, costae, and veins. Sori linear, attached along veinlets(sometimes attached along excurrent veins), exindusiate. Sporangiaeach with 1 or 2 erect acicular hairs near top.Streamsides in forests, steep forested slopes; 1200–2000 m. S andSE Taiwan, SE Yunnan [N Vietnam].2. Stegnogramma xingwenensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 286, 350. 1999 [“xinwenensis,” p.350].兴文溪边蕨 xing wen xi bian juePlants ca. 50 cm tall. Rhizomes erect, with dense brownshort hairs at margins, lanceolate scales at apices. Frondsclustered; stipes 14–20 cm, dark stramineous, with scales andthroughout with mixed grayish white different-sized acicularhairs; laminae lanceolate, 30–40 × ca. 10 cm, slightly narrowedto bases, 1-pinnate, pinnatifid-acuminate at apices; pinnae 10–14 pairs, alternate, obliquely spreading, almost sessile, exceptfor proximal several pairs of pinnae slightly reduced others ±adnate to rachises; proximal pinnae slightly reduced, ovate-lanceolate,ca. 4 × 1.5 cm, bases rounded-cuneate, not symmetrical,apices shortly acute; middle pinnae lanceolate, ca. 6 × 1.8cm, bases rounded-cuneate, margins pinnately lobed, apicesacuminate; costae raised on both sides. Veins evident, veinletsobliquely spreading, 1 or 2 veinlets joining into 2 or 3 triangularor subrhomboid areoles. Laminae grayish green when dry, herbaceous,both surfaces with dense appressed setae along veins,with sparse short hairs abaxially, glabrous adaxially. Sori linear,attached along veinlets, exindusiate; sporangia glabrous.● Forests at roadsides; ca. 1300 m. Sichuan (Xingwen).3. Stegnogramma cyrtomioides (C. Christensen) Ching, Sinensia7: 95. 1936.贯众叶溪边蕨 guan zhong ye xi bian jueDryopteris stegnogramme (Blume) C. Christensen var.cyrtomioides C. Christensen, Acta Horti Gothob. 1: 56. 1924.Plants 28–50 cm tall. Rhizomes short and erect, with densehairy brown narrowly lanceolate scales and multicellularacicular long hairs. Fronds clustered; stipes 8–25 cm, stramineous,with sparse scales on bases, throughout with densegrayish white multicellular long acicular hairs when young,hairs gradually fallen when old; laminae lanceolate, 15–25 × 4–8 cm, not narrowed to bases, 1-pinnate, pinnatifid-acuminate atapices; pinnae 8–10 pairs, alternate, spreading, proximal pairslightly shortened, proximal 3 or 4 pairs free, sessile, distal ones± adnate to rachises; middle pinnae ovate-oblong, 2–3.5 × ca.

388THELYPTERIDACEAE1.5 cm, bases broader, rounded-truncate, symmetrical, marginssubentire or slightly undulate, apices shortly acute. Veins evident,2 or 3 pairs of veinlets between veinlets, proximal pairjoining, sometimes an excurrent veinlet arising at this union andconnected to an acroscopic veinlet of second pair, forming atriangular plus a rhomboid areole. Laminae yellow-green, herbaceousor papery, abaxial surface on intercostal areas shortlyhairy, adaxially setaceous along margins and apices; along abaxialside of rachises with dense multicellular long acicular hairs,shortly hairy along abaxial sides of costae and veins, setaceousadaxially. Sori linear, attached along veinlets, exindusiate, remainingclustered and with erect short hairs at attached placesof sori after fallen; sori with 2 or 3 short hairs.● Scrub; 600–1500 m. Guizhou, Sichuan.4. Stegnogramma latipinna Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 350. 1999.阔羽溪边蕨 kuo yu xi bian juePlants ca. 70 cm tall. Rhizomes short, erect, includingstipe bases with dense setaceous brown lanceolate scales andspreading multicellular grayish white acicular long hairs. Frondsclustered; stipes 20–24 cm, dark stramineous, sparsely setaceousat bases; laminae 30–35 × 10–13 cm, slightly narrowedto bases, 1-pinnate, pinnatifid-acuminate at apices; pinnae 10–12 pairs, alternate, spreading, sessile, proximal ones free fromrachises, distal ones ± adnate to rachises and completely combinednear apical part; middle pinnae broadly lanceolate, ca. 7 ×2.2 cm, bases rounded-truncate and slightly broadened, notsymmetrical, lobed to 1/4 of distance to costule, apices shortlyacuminate; segments ovate-triangular, ca. 4 × 4 mm, entire, obtuse-roundedat apices. Veins evident, 4 or 5 pairs of veinletsper segment, proximal 2 pairs (sometimes 1.5 pairs) joining bytheir ends, veinlets of proximal pair arising far above base ofcostules. Laminae dark green when dry, herbaceous, abaxial surfacewith acicular hairs along rachises, costae, veins, and intercostalareas, adaxially with dense setae along rachises, costae,and veins, sparsely setaceous along margins, glabrous on intercostalareas. Sori linear, attached along veinlets, exindusiate;sporangia each with 3 or 4 erect acicular hairs near apices.● Streamsides in forests; 2300–2500 m. NW Yunnan (Gongshan).5. Stegnogramma diplazioides Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 351. 1999.缙云溪边蕨 jin yun xi bian juePlants ca. 60 cm tall. Rhizomes strong, suberect, with darkbrown shortly hairy lanceolate scales and few multicellulartranslucent acicular hairs at apices. Fronds clustered; stipes 25–30 cm, dark stramineous on proximal part and sparsely scaly,distally stramineous, throughout shortly setaceous; laminaeoblong-lanceolate, 25–34 × 10–12 cm, almost or slightly narrowedto bases, 1-pinnate, pinnatifid-acuminate at apices; pinnae10–14 pairs (proximal 5 or 6 pairs free from rachises),subopposite, sessile, spreading, proximal pair slightly shortened(ca. 4.5 cm), above one lanceolate, 5–6 × ca. 1.5 cm, basesrounded-cuneate and slightly narrowed, sessile, distal ones ofsimilar shape, but ± adnate to rachises at bases, distal onesgradually reduced and completely adnate to rachises at bases,entire or slightly undulate. Veins evident, veinlets oblique distally,veinlets ca. 3 pairs, bent, proximal 2 pairs (sometimes 1.5pairs) joining by their ends, proximal pair arising far above baseof costules. Laminae greenish when dry, somewhat papery,abaxially with sparse short hairs on laminae, adaxially glabrous,setaceous along margins, with unicellular long setae alongabaxial side of rachises, shortly setaceous along costae andveins, with dense appressed setae along adaxial side of costae,sparsely setaceous along veins. Sori linear, attached along veinlets;indusia shortly setaceous near apices of sporangia.● Beneath bamboo on sunny slopes. Chongqing (Beibei, JinyunShan).6. Stegnogramma jinfoshanensis Ching & Z. Y. Liu, Bull.Bot. Res., Harbin 3(4): 13. 1983.金佛山溪边蕨 jin fo shan xi bian juePlants 35–40 cm tall. Rhizomes short and ascending, includingstipe bases with dense multicellular acicular hairs andhairy reddish brown lanceolate scales. Fronds clustered; stipes10–20 cm, grayish brown, with sparse scales and spreadingmulticellular acicular hairs mixed with unicellular setae onproximal part; laminae lanceolate, 18–27 × 5.2–7 cm, not orslightly narrowed to bases, 1-pinnate, pinnatifid-acuminate atapices; pinnae 8–12 pairs, alternate, obliquely spreading, sessile,proximal 3 or 4 pairs free, distal ones ± adnate to rachises;middle pinnae broadly lanceolate, 4–5 × ca. 1.6 cm, basesbroadened, subtruncate, symmetrical, margins crenate or pinnatilobate,apices acute or obtuse. Veins evident, veinlets 3 or 4pairs, proximal 1.5 pairs joining by their ends, proximal pairarising from far above base of costules. Laminae grayish greenor greenish when dry, thinly papery, abaxially with short acicularhairs on intercostal areas, adaxially usually shortly hairy onintercostal areas, with unicellular long setae along both sides ofrachises, shortly hairy along costae and veins, with dense appressedsetae along adaxial side of costae and sparsely setaceousalong veins. Sori linear, attached along veinlets, usuallyconfluent at apices, exindusiate, with remaining glandlike materialat attachment of sori; sporangia setaceous near apices.● Shaded thickets at foot of limestone mountains; ca. 2500 m.Chongqing, Sichuan, Yunnan.16. AMPELOPTERIS Kunze, Bot. Zeitung (Berlin) 6: 114. 1848.星毛蕨属 xing mao jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants trailing, in soil. Rhizomes long creeping, stipe bases with sparse hairy dark brown scales. Fronds clustered or approximate;stipes subglabrous; laminae lanceolate; rachises usually prolonged, whiplike and rooting on soil and forming a new plant, 1-

THELYPTERIDACEAE 389pinnate; pinnae ca. 30 pairs; pinna axils often with gemmae producing a 1-pinnate small lamina. Veins evident, veinlets obliquelyspreading and joining by their ends, a tortuous excurrent veinlet connected with veinlets of every pair to marginal sinus and forminga line of oblique square areoles. Laminae papery, greenish or dark brown-green, with forked or simple short hairs along both sides ofrachises and intercostal areas, glabrous when old. Sori suborbicular or oblong, exindusiate. Sporangia glabrous. Spores elliptic,monolete, perispore thin and translucent, minutely reticulate and with small spines.One species: tropical and subtropical regions of the Old World.1. Ampelopteris prolifera (Retzius) Copeland, Gen. Fil. 144.1947.星毛蕨 xing mao jueHemionitis prolifera Retzius, Observ. Bot. 6: 36. 1791;Abacopteris prolifera (Retzius) W. C. Shieh; Ampelopteriselegans Kunze; Aspidium proliferum (Retzius) Hieronymus(1895), not R. Brown (1810); Cyclosorus prolifer (Retzius)Tardieu & C. Christensen; Dryopteris prolifera (Retzius) C.Christensen; Goniopteris prolifera (Retzius) C. Presl; Menisciumproliferum (Retzius) Swartz; Phegopteris luxurians(Kunze) Mettenius; P. prolifera (Retzius) Kuhn (1879), not(Kaulfuss) Mettenius (1856); Polypodium luxurians Kunze;P. proliferum (Retzius) Hooker (1864), not Kaulfuss (1824);Thelypteris prolifera (Retzius) C. F. Reed.Plants ca. 1 m tall. Stipe bases with sparse dark brownstellate-hairy lanceolate scales. Stipes stramineous, firm, ca.40 cm; laminae lanceolate, bases slightly narrowed; pinnaespreading, subopposite, subsessile, lanceolate, 5–10(–15) × ca.2 cm, bases rounded-truncate, margins undulate, apices shortlypointed. Sori attached on middle of veinlets, usually confluentwhen mature. n = 36.On sandy floodplains by streams in sunny areas; 100–1000 m.Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan,Taiwan, Yunnan [tropical and subtropical regions of the worldexcept the Americas].The young fronds are eaten as a vegetable.Abacopteris Fée.17. PRONEPHRIUM C. Presl, Epimel. Bot. 258. 1851.新月蕨属 xin yue jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants, medium-sized, in soil. Rhizomes long creeping, or short and decumbent, with sparse usually hairy brown scales. Frondsremote or approximate; stipes glabrous except at bases, but often (particularly when young) ± with unicellular acicular hairs; laminaeusually 1-imparipinnate, sometimes simple or ternate; pinnae large, usually 3–10(–15) pairs, terminal pinna free, of similar shape aslateral ones, proximal pair not or slightly shortened, lanceolate, bases rounded or cuneate, subsessile or shortly stalked, not adnate torachises, margins entire or thickly serrate, apices acuminate; costae evident, veinlets mostly obliquely spreading; venation meniscioid,i.e., veinlets joining into oblique square areoles between veinlets, an excurrent veinlet arising from joining point of every pairof veinlets continuous or interrupted and with hydathodes at apices. Laminae herbaceous or papery, sometimes somewhat leathery,green or dark brown when dry, often reddish (at least abaxially along rachises, costae and veinlets), usually ± with acicular andhooked hairs (at least abaxially along along rachises and costae), abaxially usually foveolate on intercostal areas. Sori orbicular, in 2lines between veinlets, one per veinlet, if attached on distal part of veinlets then usually confluent when mature, rarely spreadingthroughout abaxial surface of pinnae, exindusiate or indusiate, hairy or glabrous; sporangia glabrous or with acicular hairs. Sporesbilateral, reniform, perispore transparent or translucent, perispore ridged corrugate, verrucate, or echinate. x = 36.Sixty-one species: tropical and subtropical regions of Asia; 18 species (eight endemic) in China.The following taxon is excluded from the present treatment, pending further research: Pronephrium fengkaiensis B. S. Wang & S. H. Shi (ActaSci. Nat. Univ. Sunyatseni 29: 72. 1990).1a. Plants with hooked hairs throughout.2a. Laminae simple, strongly dimorphic, sterile laminae cordate or hastate (i.e., occasionally with one pair ofsmall auricles proximally), sori spreading throughout abaxially on mature laminae ............................................... 1. P. simplex2b. Laminae ternate or pinnate (occasionally simple, proximally rounded or cuneate), monomorphic, sometimesdimorphic, sori arranged in lines when mature.3a. Laminae ternate (sometimes with 2 pairs of lateral pinnae), green when dry ............................................... 2. P. triphyllum3b. Laminae pinnate and with multiple pairs of lateral pinnae, green or reddish when dry.4a. Rhizomes short and decumbent; fronds approximate, pinnae distally usually with axillary budsin their axils ............................................................................................................................................ 3. P. cuspidatum4b. Rhizomes long creeping; fronds remote, pinnae distally lacking buds in their axils.5a. Proximal pinnae obviously reduced; pinna apices abruptly narrowed into a long tail 2–4 cm ...... 4. P. megacuspe5b. Proximal pinnae usually largest; pinna apices not narrowed or caudate.6a. Apical pinnae lobed throughout .................................................................................................... 5. P. insularis

390THELYPTERIDACEAE6b. Apical pinnae entire (at most crenate along margins).7a. Laminae oblong to ovate-rounded, lateral pinnae smaller than ca. 9 × 2 cm ............. 6. P. longipetiolatum7b. Laminae triangular, lateral pinnae ca. 15 × 3 cm, proximal pair of pinnae largest ................. 7. P. parishii1b. Plants without hooked hairs on every part.8a. Sori exindusiate or with very small indusia.9a. Laminae normally greenish when dry; pinnae linear-lanceolate, margins regularly and sharplyserrate, pinna lobes triangular ................................................................................................................... 8. P. penangianum9b. Laminae ± purplish red when dry; pinnae ovate-lanceolate, margins entire or slightly undulate.10a. Pinnae narrowly ovate, broadest at bases and rounded-cuneate; laminae completely glabrouson both surfaces; sori ovate and confluent when mature; Xizang ..................................................... 9. P. medogensis10b. Pinnae lanceolate, broadest at middle, costae and veins with short hairs; sori orbicular andnot confluent when mature; throughout China.11a. Laminae abaxially with only sparse short hairs along rachises and costae, elsewhereglabrous; sori with small indusia ......................................................................................... 10. P. lakhimpurense11b. Laminae abaxially with acicular hairs along rachises, costae, veins, and intercostalareas; sori exindusiate ................................................................................................................... 11. P. hirsutum8b. Sori obviously indusiate.12a. Pinnae broadly linear-lanceolate, bases not or slightly narrowed, margins regularly serrate.13a. Sporangia glabrous; indusia occasionally with one or two short hairs .............................................. 12. P. nudatum13b. Sporangia and indusia all with hairs.14a. Stipes throughout with acicular setae, laminae abaxially with dense acicular long hairsalong costae, veins, and on intercostal areas .............................................................................. 13. P. setosum14b. Stipes glabrous, laminae abaxially with only sparse hairs along costae and veins ............. 14. P. yunguiensis12b. Pinnae ovate-lanceolate or falcate, bases cuneate, margins entire or irregularly undulate-crenate.15a. Rhizomes short and decumbent; plants smaller; lateral pinnae 2 or 3 pairs, pinnaeca. 7 × 2–2.5 cm .................................................................................................................................... 15. P. gracilis15b. Rhizomes long creeping; plants much larger; lateral pinnae more than 4 pairs,more than ca. 15 × 3 cm.16a. Laminae abaxially with sparse short hairs along costae and veins ............................ 16. P. gymnopteridifrons16b. Laminae abaxially with dense long hairs along costae, veins, and intercostal areas.17a. Pinnae ovate-lanceolate, bases rounded-cuneate; laminae not foveolate onintercostal areas, with unicellular acicular hairs; sori attached on middleof veinlets, not confluent when mature ................................................................... 17. P. macrophyllum17b. Pinnae oblong-lanceolate, bases cuneate; laminae obviously foveolateabaxially on intercostal areas, with dense multicellular articulate longhairs; sori attached on distal part of veinlets and confluentwhen mature .................................................................................................................. 18. P. hekouensis1. Pronephrium simplex (Hooker) Holttum, Blumea 20: 122.1972.单叶新月蕨 dan ye xin yue jueMeniscium simplex Hooker, London J. Bot. 1: 294.1842; Abacopteris simplex (Hooker) Ching; Aspidium simplex(Hooker) Hance; Asplenium simplex (Hooker) Hance; Cyclosorussimplex (Hooker) Copeland; Dryopteris simplex (Hooker)C. Christensen; Nephrodium simplex (Hooker) Diels; Phegopterissimplex (Hooker) Mettenius; Polypodium simplex(Hooker) E. J. Lowe (1858), not N. L. Burman (1768), norSwartz (1801); Thelypteris simplex (Hooker) K. Iwatsuki.Plants 30–40 cm tall. Rhizomes long creeping, with sparse,dark brown lanceolate scales and hooked hairs at apices. Frondsremote, simple, dimorphic; sterile stipes 14–18 cm, stramineous,occasionally with 1 or 2 scales at bases, distally withdense hooked short hairs, sometimes with acicular long hairs;laminae elliptic-lanceolate, 15–20 × 4–5 cm, entire or undulate.Veins visible, oblique distally, parallel to each other, onesubrectangular areole between veinlets, above with 2 lines ofsubsquare areoles. Laminae papery when dry, with hookedshort hairs on both surfaces, denser hairs along rachises andveins sometimes with long acicular hairs. Fertile fronds muchtaller than sterile ones; stipes 30–35 cm; laminae lanceolate, 5–10 × 8–15 cm, entire, bases cordate, apices long acuminate;veins and hairs same as those on sterile fronds. Sori attached onveinlets, orbicular when young, exindusiate, spreading throughoutabaxial surface of pinnae when mature.Forests by streams, forests in valleys; sea level to 1500 m. Fujian,Guangdong, Hainan, Taiwan, SE Yunnan [Japan, Vietnam].The correct position of Abacopteris simplex var. trifoliata Ching(Bull. Fan Mem. Inst. Biol., Bot. 10: 10. 1940; Cyclosorus simplex var.trifoliata (Ching) T. J. Liu; Thelypteris simplex var. trifoliata (Ching) C.F. Reed), described from Fujian, is not known.2. Pronephrium triphyllum (Swartz) Holttum, Blumea 20:122. 1972.三羽新月蕨 san yu xin yue jueMeniscium triphyllum Swartz in Schrader, J. Bot. 1800(2):16. 1801; Abacopteris triphylla (Swartz) Ching; Cyclosorus

THELYPTERIDACEAE 391triphyllus (Swartz) Tardieu; Dryopteris triphylla (Swartz) C.Christensen; Nephrodium triphyllum (Swartz) Diels; Phegopteristriphylla (Swartz) Mettenius; Thelypteris triphylla(Swartz) K. Iwatsuki.Plants 20–50 cm tall. Rhizomes long creeping, blackishbrown, with dense white hooked short hairs and brown hairylanceolate scales. Fronds monomorphic or dimorphic; stipes10–40 cm, dark stramineous, bases with sparse scales andthroughout with dense curved short hairs; laminae ovate-triangular,12–20 × 7–11 cm, rounded at bases, ternate, long acuminateat apices; lateral pinnae 1 pair (rarely 2 pairs), obliquedistally, opposite, oblong-lanceolate, 5–9 × 1.5–2.5 cm,bases rounded or rounded-cuneate, stalk 1–2 mm, margins entire,apices shortly acuminate; terminal pinna very large, lanceolate,15–18 × 3–3.5 cm, bases rounded or rounded-cuneate,stalk 6–12 mm, margins entire or undulate, apices acuminate.Veins evident abaxially, veinlets obliquely spreading and parallel,veins in middle of pinnae usually 8 or 9 pairs oblique orspreading, veinlet pairs joining by their ends and forming triangularareoles, an excurrent veinlet arising from joining pointconnected with veinlets of others forming subsquare areoles.Laminae firmly papery, adaxially glabrous except for densehooked hairs in costal grooves, abaxially with hooked hairsalong costae and veinlets and also with scattered hooked hairson intercostal areas. Fertile fronds slightly taller than sterileones, stipes longer, pinnae narrower. Sori attached on veinlets,orbicular when young, becoming narrowly ovate and confluent,exindusiate; sporangia each with 2 hooked hairs.Forests; 100–600 m. Fujian, Guangdong, Guangxi, Taiwan, SEYunnan [India, Indonesia, Japan, S Korea, Sri Lanka, Malaysia, Myanmar;NE Australia].The correct position of Abacopteris triphylla var. simplicifoliaChing (Bull. Fan Mem. Inst. Biol., Bot. 8: 243. 1938) is not known.Cyclosorus ×pseudoliukiuensis (Serizawa) Ralf Knapp (Ferns FernAllies Taiwan, 445. 2011; Thelypteris ×pseudoliukiuensis Serizawa)is the putative hybrid between Pronephrium triphyllum (as C. triphyllus)and the following species, P. cuspidatum (as C. liukiuensis).3. Pronephrium cuspidatum (Blume) Holttum, Blumea 20:123. 1972.顶芽新月蕨 ding ya xin yue jueMeniscium cuspidatum Blume, Enum. Pl. Javae 2: 114.1828; Abacopteris cuspidata (Blume) Ching; A. liukiuensis(Christ ex Matsumura) Tagawa; Cyclosorus cuspidatus (Blume)Copeland; C. liukiuensis (Christ ex Matsumura) Masamune;Dryopteris cuspidata (Blume) Christ; D. liukiuense (Christ exMatsumura) C. Christensen; M. liukiuense Christ ex Matsumura;Nephrodium clavivenum Yabe ex Matsumura & Hayata;Phegopteris cuspidata (Blume) Mettenius; Thelypteris liukiuensis(Christ ex Matsumura) K. Iwatsuki.Plants ca. 1 m tall. Rhizomes short and decumbent, withdark brown scales; scales lanceolate, sparsely shortly hairy.Fronds approximate; stipes 15–35 cm, bases with dark brownscales and hooked hairs, distally glabrous; laminae ovate, 25–30 cm, 1-imparipinnate; lateral pinnae 2–4 pairs, oblanceolate,8–14 × 2–3.5 cm, cuneate or narrowly rounded at bases, shortlystalked, usually with a gemma in axil, entire or undulate-crenateat margins, caudate-acuminate at apices; terminal pinnalarger than lateral ones. Laminae papery when dry, brown andtinged scarlet. Veinlets obvious, 6–8 pairs regularly joining intoareoles, excurrent veinlets usually not reaching next pair ofjoined veins. Sori orbicular or elongate, attached on middle ofveinlets, usually confluent and spreading throughout pinna surface,exindusiate.Dense forests on low mountains. Taiwan [Japan, Malaysia; Pacificislands (Solomon Islands)].Knapp (Ferns Fern Allies Taiwan, 444. 2011) does not treat Cyclosoruscuspidatus and C. liukiuensis as conspecific, accepting only C.liukiuensis for Taiwan. Dryopteris cuspidata var. epigea Copeland(Philipp. J. Sci. 3: 278. 1908), described from Guangdong (Tai MoShan), may belong here.4. Pronephrium megacuspe (Baker) Holttum, Blumea 20: 122.1972.微红新月蕨 wei hong xin yue juePolypodium megacuspe Baker, J. Bot. 28: 266. 1890;Abacopteris sampsonii (Baker) Ching; P. sampsonii Baker;Pronephrium sampsonii (Baker) Ching ex K. H. Shing.Plants 50–70 cm tall. Rhizomes creeping, blackish brown,with dense hooked hairs and hairy brown lanceolate scales.Fronds remote; stipes 25–35 cm, stramineous, sparsely hairy atbases, distally sparsely setaceous; laminae oblong, 25–35 × ca.30 cm, slightly narrowed to bases, 1-imparipinnate, caudatepointedat apices; lateral pinnae 5 or 6 pairs, alternate, proximalpair slightly shortened, shortly stalked, others sessile, obliquedistally, lanceolate, 12–14 × ca. 2.5 cm, cuneate at bases, entireor undulate, caudate-acuminate at apices; terminal pinna of similarshape as lateral ones, but very large, stalk 2–4 mm. Veinsevident, veinlets oblique distally and parallel, veinlets subobliquelyspreading and forming triangular areoles at basesbetween veinlets, distally forming a line of V-shaped areoles,with an expanded hydathode at excurrent vein arising fromjoining point. Laminae papery when dry, reddish, abaxial surfacewith more hooked hairs along and short hairs when young,fallen when old and with few remaining on rachises and veinsonly. Sori attached above middle of veinlets and confluentwhen mature and forming a horizontal, equidistant row betweenveinlets, exindusiate; sporangia bearing hairs when young.Dense forests, along streams, wetlands; 100–400 m. Guangdong,Guangxi, Jiangxi, S Taiwan, Yunnan [Japan, Thailand, Vietnam].5. Pronephrium insularis (K. Iwatsuki) Holttum, Blumea 20:123. 1972.岛生新月蕨 dao sheng xin yue jueAbacopteris insularis K. Iwatsuki, Acta Phytotax. Geobot.18: 6. 1959; Cyclosorus insularis (K. Iwatsuki) C. M. Kuo;Thelypteris insularis (K. Iwatsuki) K. Iwatsuki.Rhizomes long creeping, with dense hooked hairs andsparsely hairy triangular-lanceolate scales. Fronds remote; stipes10–15 cm, with linear-lanceolate scales at bases; laminae triangularor ovate, 20–35 × 15–25 cm, terminal pinna very large,linear-lanceolate, bases pinnatifid; lateral pinnae 4–7 pairs, lanceolate,bases rounded-obtuse, not symmetrical, apices acu-

392THELYPTERIDACEAEminate; proximal pinnae stalked. Sori orbicular, well separated,attached on middle of veinlets, exindusiate.Forests; ca. 1000 m. E Taiwan [Japan].6. Pronephrium longipetiolatum (K. Iwatsuki) Holttum,Blumea 20: 123. 1972.长柄新月蕨 chang bing xin yue jueAbacopteris longipetiolata K. Iwatsuki, Acta Phytotax.Geobot. 18: 11. 1959; Cyclosorus longipetiolatus (K. Iwatsuki)C. M. Kuo; Thelypteris longipetiolata (K. Iwatsuki) K. Iwatsuki.Plants 30–50 cm tall. Rhizomes long and decumbent, withdense hooked hairs and scales; scales narrowly triangular-lanceolate,sparsely hairy on surface and margins, acuminate atapices. Fronds remote; stipes 10–30 cm, stramineous, withhooked hairs, bases slightly scaly; laminae subdimorphic, oblongto ovate-oblong, 15–20 × 9–14 cm, imparipinnate; terminalpinna very large; lateral pinnae 2–4 pairs, oblong to lanceolate,5–9 × 1.5–2 cm, bases rounded-cuneate, obviously shortlystalked, margins entire or irregularly undulate, apices acuminateto shortly caudate-acuminate; terminal pinna oblong-lanceolate,9–13 × 2.5–4 cm, usually not symmetrical, long stalked, shortlycaudate-acuminate at apices; rachises and costae with sparsehooked hairs on both sides. Veinlets not obviously raised abaxiallyand slightly oblique distally, veinlets forming areolesand spreading an excurrent veinlet from joining point. Soriattached on middle of veinlets, confluent when mature, exindusiate,hooked-hairy on sporangia.● Forests; 200–500 m. S and SE Taiwan.7. Pronephrium parishii (Beddome) Holttum, Blumea 20:123. 1972.羽叶新月蕨 yu ye xin yue jueMeniscium parishii Beddome, Ferns Brit. Ind. t. 184.1866; Abacopteris triphylla (Swartz) Ching var. parishii (Beddome)Ching; Cyclosorus parishii (Beddome) Tardieu; M. triphyllumSwartz var. parishii (Beddome) Beddome; Pronephriumtriphyllum (Swartz) Holttum var. parishii (Beddome)Nakaike; Thelypteris triphylla (Swartz) K. Iwatsuki var. parishii(Beddome) K. Iwatsuki.Plants 30–50 cm tall. Rhizomes long creeping, blackishbrown, with dense white hooked short hairs and brown hairylanceolate scales. Fronds monomorphic or dimorphic; stipes10–40 cm, dark stramineous, bases with sparse scales andthroughout with dense hooked hairs; laminae ovate-triangular,25–30 × 10–15 cm, long acuminate at apices; lateral pinnae 2–5pairs (fertile laminae sometimes ternate), oblique distally, subopposite,oblong-lanceolate; proximal pair of pinnae longest, 6–15 × 2–3 cm, stalk 1–2 mm, rounded or rounded-cuneate atbases, entire, shortly acuminate at apices; distal pinnae adnateto rachises and decurrent; terminal pinna ca. 20 × 3–4 cm,margins undulate, usually with 1 or 2 free small auricles atbases, acuminate at apices. Veins evident abaxially, veinletsobliquely spreading and parallel, in middle of lateral pinnaeusually 8 or 9 pairs, oblique or spreading, veinlet pairs joiningby theirs ends and forming triangular areoles, an excurrentveinlet arising from joining point connected with veinlets ofothers forming subsquare areoles. Laminae firmly papery, adaxiallyglabrous except for dense hooked hairs in costal grooves,abaxially with hooked hairs along costae and veinlets and alsowith scattered hooked hairs on intercostal areas. Sori attachedon veinlets, orbicular when young, becoming narrowly ovateand confluent, exindusiate; sporangia each with 2 hooked hairs.Forests; 200–500 m. Taiwan [S India, Japan, Malaysia, Myanmar,Sri Lanka].8. Pronephrium penangianum (Hooker) Holttum, Blumea 20:110. 1972.披针新月蕨 pi zhen xin yue juePolypodium penangianum Hooker, Sp. Fil. 5: 13. 1863[“panangianum”]; Abacopteris penangiana (Hooker) Ching;Aspidium porphyrophlebium Christ; A. rampans (Baker) Christ;Christella porphyrophlebia (Christ) H. Léveillé; Dryopterisporphyrophlebia (Christ) C. Christensen; D. rampans (Baker)C. Christensen; Goniopteris penangiana (Hooker) Beddome;Nephrodium rampans Baker; Thelypteris penangiana (Hooker)C. F. Reed; T. porphyrophlebia (Christ) C. F. Reed; T. rampans(Baker) C. F. Reed.Plants 1–2 m tall. Rhizomes long creeping, dark brown,ca. 1.2 cm in diam., occasionally with 1 or 2 brown lanceolatescales. Fronds remote; stipes ca. 1 m, dark brown, distally reddishbrown, glabrous; laminae oblong-lanceolate, 40–80 × 25–40 cm, 1-imparipinnate; lateral pinnae 10–15 pairs, obliquelyspreading, alternate, stalked, broadly linear, pinnae from proximalmiddle part 20–30 × 2–2.7 cm, broadly cuneate at bases,sharply cartilaginous-serrate or teethlike along margins, acuminateat apices; pinnae distally slightly shortened; terminal pinnaof similar shape and size as lateral pinnae, stalks ca. 1 cm. Veinletsevident abaxially, spreading and parallel to each other, 9 or10 pairs, joining by theirs ends and forming triangular areolesbetween veinlets, an excurrent veinlet arising from joining pointconnected with more distal veinlets (sometimes interrupted) andforming 2 rows of elongated square areoles, distal 2 or 3 pairsof veinlets free and reaching margins. Laminae papery whendry, dark brown or reddish brown, glabrous throughout. Soriorbicular, attached on middle or proximal middle of veinletsand in 2 rows between veinlets, 6 or 7 per row, exindusiate.Sparse forests, shaded streamsides; 900–3600 m. Guangdong,Guangxi, Guizhou, S Henan, Hubei, Hunan, Jiangxi, Sichuan, Zhejiang[Bhutan, India, Kashmir, Nepal, Pakistan].Pronephrium penangianum is used as a medicinal herb at EmeiShan, Sichuan. The fronds are used in traditional Chinese medicine forirregular menstruation.9. Pronephrium medogensis Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 351. 1999.墨脱新月蕨 mo tuo xin yue juePlants to 2 m tall. Rhizomes long creeping. Fronds remote;stipes ca. 1 m, occasionally sparsely scaly at bases, dark stramineous;laminae oblong-lanceolate or ovate-oblong, 80–100cm, 1-imparipinnate, acuminate at apices; lateral pinnae 8–12pairs, subobliquely spreading, narrowly lanceolate, ca. 25 × 3–

THELYPTERIDACEAE 3933.5 cm, broadest at bases and subrounded, long caudateacuminateat apices, sessile; terminal pinna of similar shape asproximal ones. Veins visible adaxially, raised abaxially, costaeprominent; lateral veinlets subobliquely spreading and parallelto each other, neighboring ones joining with each other into V-shaped areoles, excurrent veins very short. Laminae somewhatleathery when dry, reddish, rachises, costae, and intercostalareas glabrous throughout. Sori orbicular, attached above middleof veinlets and confluent when mature, exindusiate.● Monsoon forests; ca. 700 m. Xizang (Beibeng, Mêdog).10. Pronephrium lakhimpurense (Rosenstock) Holttum, Blumea20: 110. 1972.红色新月蕨 hong se xin yue jueDryopteris lakhimpurensis Rosenstock, Meded. Rijks-Herb. 31: 7. 1917; Abacopteris rubra (Ching) Ching; Cyclosorusrubrus (Ching) Tardieu ex Tardieu & C. Christensen; D.rubra Ching; Meniscium cuspidatum Blume var. longifronsWallich ex C. B. Clarke; Polypodium urophyllum Wallich exHooker & Baker var. khasianum C. B. Clarke; Thelypterislakhimpurensis (Rosenstock) K. Iwatsuki; T. rubra (Ching) K.Iwatsuki.Plants ca. 1.5 m tall. Rhizomes long creeping. Fronds remote;stipes 80–90 cm, occasionally with 1 or 2 scales at bases,dark stramineous; laminae oblong-lanceolate or ovate-oblong,60–85 cm, 1-imparipinnate, acuminate at apices; lateral pinnae8–12 pairs, subobliquely spreading, pinnae on proximal to middlepart broadly lanceolate, 24–32 × 4–6 cm, stalk ca. 2 mm,somewhat rounded at bases, entire or undulate, shortly caudatepointedat apices; terminal pinna of similar shape as proximalones, stalk 1.5–2 cm. Veins slender, evident abaxially, veinletssubobliquely spreading and parallel to each other, veinlets 13–17 pairs, subobliquely spreading, proximal pairs joined to formtriangular areoles, distally every pair of veinlets connected withexcurrent veinlet forming 2 rows of rhomboid areoles, excurrentveinlets reaching or near joining point of above pair ofveinlets. Laminae thinly papery or herbaceous, dark brown, glabrouson both surfaces, occasionally with 1 or 2 short setaeabaxially, rachises, costae, and veins with sparse short hairs.Sori orbicular, attached on middle or above middle part of veinletsand arranged in 2 rows, occasionally confluent when mature,exindusiate.Valleys, streamsides in forests; 300–1600 m. Fujian, Guangdong,Guangxi, Jiangxi, Sichuan, Yunnan [Bhutan, N India, Nepal, N Thailand,Vietnam].11. Pronephrium hirsutum Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 305, 351. 1999 [“Pronrphrium,” p. 351].针毛新月蕨 zhen mao xin yue juePlants 60–100 cm tall. Rhizomes long creeping, withsparse ovate-lanceolate brown scales. Fronds remote; stipes 25–70 cm, sparsely brown scaly at bases, stramineous; laminae lanceolate,30–50 × 15–30 cm, 1-imparipinnate; lateral pinnae 4–8pairs, alternate, sessile, proximal pair reduced; middle pinnaeoblong-lanceolate, 10–25 × 2–5 cm, bases rounded-cuneate,margins entire or slightly undulate, apices acuminate; terminalpinna much larger than lateral ones, ovate-lanceolate, 15–25(–30) × 2.5–5.5(–6.5) cm, stalk 1–4 cm, bases not symmetricaland slightly decurrent on one lateral side, roundedcuneate,margins undulate or crenate, apices long acuminate.Veins evident abaxially, costules thick and raised, veinletsobliquely spreading and parallel to each other, veinletsobliquely spreading, proximal pair joining into triangularareoles between veinlets, distally every pair joining into areoles,excurrent veinlet not reaching joining point of above pair, withdense short setae along veins. Laminae papery or herbaceouswhen dry, dark green, reddish, or dark red, veins reddish abaxially,rachises, costae, and veins and abaxial intercostal areas allwith dense acicular hairs, abaxial surface with sparse short setaealong costae and veins, with 1 or 2 occasional setae on intercostalareas. Sori orbicular, attached on middle or above middlepart of veinlets, not confluent when mature, exindusiate.● Gregarious on shaded wet precipitous slopes, wetlands byrivers, marshlands. Chongqing, Fujian, Guangdong, Guizhou, Yunnan.12. Pronephrium nudatum (Roxburgh) Holttum, Blumea 20:111. 1972.大羽新月蕨 da yu xin yue juePolypodium nudatum Roxburgh, Calcutta J. Nat. Hist. 4:491. 1844; Abacopteris multilineata (Wallich ex Hooker)Ching; Aspidium moulmeinense (Beddome) Christ; A. multilineatum(Wallich ex Hooker) Christ (1897), not Mettenius(1858); Christella moulmeinense (Beddome) H. Léveillé; Cyclosorusmoulmeinensis (Beddome) Tardieu & C. Christensen;C. multilineatus (Wallich ex Hooker) Tardieu & C. Christensen;Dryopteris moulmeinensis (Beddome) C. Christensen; Goniopterislineata (Colebrook ex Hooker) Beddome; G. multilineata(Wallich ex Hooker) Beddome; Nephrodium moulmeinenseBeddome; N. multilineatum (Wallich ex Hooker) Beddome;Phegopteris lineata (Colebrook ex Hooker) Mettenius;P. multilineata (Wallich ex Hooker) Luerssen; Polypodiumlineatum Colebrook ex Hooker; P. multilineatum Wallich exHooker; Thelypteris multilineata (Wallich ex Hooker) C. V.Morton; T. nudata (Roxburgh) C. V. Morton.Plants to 2.5 m tall. Rhizomes strong, creeping, woody,dark brown, with sparse broadly lanceolate scales. Fronds remote;stipes 50–80(–140) cm, bases with brown scales, distallyglabrous, dark brown, distally brownish; laminae broadly ovateoblong,60–90 × 26–40(–60) cm, 1-imparipinnate; lateral pinnae8–14(–16) pairs, obliquely spreading, alternate, subsessile,pinnae on low and middle parts broadly linear-lanceolate, 26–30(–35) × 3–4(–5) cm, bases somewhat rounded or cuneate,margins regularly shortly and sharply serrate, apices long acuminate;distal pinnae slightly shortened; terminal pinna of similarshape as middle ones, slightly shorter, bases with both sidesnot symmetrical, stalk ca. 1 cm. Veins evident, veinlets raisedon both sides, spreading or obliquely spreading and parallel toeach other, veinlets obliquely spreading distally, raised abaxiallyand forming triangular areoles between veinlets, distallyforming juxtaposed rhomboid areoles. Laminae herbaceouswhen dry, green or grayish green, abaxial surface with sparseshort setae along veins, also along both sides of rachises andcostae, abaxial surface foveolate on intercostal areas. Sori

394THELYPTERIDACEAEorbicular, attached on middle of veinlets, arranged in 2 rowsbetween costules; indusia small, shortly hairy adaxially. Sporangiaglabrous.Shaded sparse forests on slopes; 100–1600 m. Guizhou, Xizang,Yunnan [Bhutan, N India, Indonesia, Myanmar, Nepal, Philippines,Vietnam].“Polypodium lineatum Colebrook ex Wallich” (Numer. List, no.300. 1829, nom. nud.) and “Goniopteris lineata C. Presl” (Tent. Pterid.183. 1836, nom. nud.) belong here. The taxon was validated by Hookerin 1864 as P. lineatum. Beddome named the taxon Nephrodium moulmeinensebecause of the blocking name N. lineatum (Blume) Beddome.13. Pronephrium setosum Y. X. Lin, Fl. Reipubl. PopularisSin. 4(1): 352. 1999.刚毛新月蕨 gang mao xin yue juePlants 50–100 cm or taller. Rhizomes creeping, ca. 1 cmin diam., with dense lanceolate brown scales. Fronds remote;stipes 30–100 cm, woody, dark stramineous, bases with darkbrown lanceolate scales, throughout with acicular setae; laminaeelliptic, 50–80 × 30–60 cm, 1-imparipinnate; lateral pinnae8–11 pairs, subobliquely spreading, alternate, proximal pairslightly reduced; middle ones narrowly ovate, 15–40 × 3–3.5cm, bases rounded-cuneate, shortly stalked, margins regularlyserrate, apices acuminate; terminal pinna of similar shape andsize as lateral ones, long stalked. Veins evident, costae groovedadaxially, raised abaxially, veinlets raised on both sides, proximalpair of veinlets forming a triangular areole, distally everypair with excurrent veinlets joining into multipaired subrectangularareoles. Laminae papery when dry, yellowishbrown, adaxially with dense appressed setae along costalgrooves, occasionally with 1 or 2 setae along veinlets, elsewhereglabrous, abaxial surface with dense long acicular hairsalong veins and intercostal areas. Sori orbicular, attached onmiddle of veinlets, with dense short hairs on indusia, alsoshortly hairy on sporangia.● Dense forests; 500–1300 m. Yunnan.14. Pronephrium yunguiensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 352. 1999.云贵新月蕨 yun gui xin yue juePlants 70–150 cm tall. Rhizomes creeping, thick, ca. 1 cmin diam., with narrowly lanceolate scales. Fronds remote; stipes40–100 cm, woody and glabrous; laminae ovate-lanceolate, 55–80 × 40–60 cm, 1-imparipinnate; lateral pinnae 8–16 pairs,proximal pair slightly reduced; middle pinnae narrowly ovate,30–50 × 3.5–4.5 cm, bases rounded-cuneate, shortly stalked,margins regularly serrate, apices long caudate; terminal pinnalarger than proximal ones, of similar shape, long stalked. Veinsevident, raised abaxially, costae grooved adaxially. Laminaedark brown-green when dry, papery, adaxially with denseappressed setae along costal grooves, abaxial surface with verysparse short hairs along costae and veinlets, elsewhere glabrous,usually not foveolate on intercostal areas. Sori orbicular,attached on middle of veinlets; indusia and sporangia hairy.● Sparse forests on slopes; 200–800 m. Guizhou, Yunnan.15. Pronephrium gracilis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 308, 352. 1999 [“gracillis,” p. 352].小叶新月蕨 xiao ye xin yue juePlants 25–40 cm tall. Rhizomes short and decumbent, withsparse narrowly lanceolate brown scales. Fronds remote; stipes20–30 cm, bases sparsely scaly, distally shortly setaceous,stramineous; laminae elliptic, 12–18 × 11–14 cm, 1-imparipinnate;lateral pinnae 2 or 3 pairs, alternate, slightly bent distally,subsessile, proximal pair not reduced, ovate-lanceolate, 7–8 ×2–2.5 cm, not symmetrical, bases cuneate, margins slightly undulate,apices acuminate or acute; terminal pinna very large,symmetrical, bases cuneate, long stalked, apices acuminate.Veins evident, raised abaxially, costae grooved adaxially, proximalpair of veinlets forming a triangular areole, distally everypair joining with an excurrent veinlet into square or rectangularor subrhomboid areoles. Laminae dark brown-green whendry, papery, adaxially with dense appressed setae along costalgrooves, veinlets and tissue between veins with sparse shorthairs; abaxially veins with short hairs, mesophyll with longacicular hairs and foveolate. Sori orbicular, attached near endsof veinlets and often confluent, hairy on indusia and also onsporangia.● Yunnan.16. Pronephrium gymnopteridifrons (Hayata) Holttum, Blumea20: 112. 1972.新月蕨 xin yue jueDryopteris gymnopteridifrons Hayata, Icon. Pl. Formosan.8: 148. 1919; Abacopteris gymnopteridifrons (Hayata) Ching;Cyclosorus gymnopteridifrons (Hayata) C. M. Kuo; C. pustulosusCopeland; D. glandulosa C. Christensen; Polypodiumurophyllum Wallich ex Hooker & Baker var. uniseriale Hooker;Thelypteris gymnopteridifrons (Hayata) C. M. Kuo; T. pustulosa(Copeland) C. F. Reed.Plants 80–120 cm tall. Rhizomes long creeping, withdense brown lanceolate scales. Fronds remote; stipes 28–80 cm,bases scaly, distally with dense short hairs, stramineous; laminaebroadly ovate or ovate-oblong, 40–80 × 15–30 cm, 1-imparipinnate; lateral pinnae 3–8 pairs, rarely more, sessile,oblique distally, proximal pair shorter, subopposite, distal onesalternate; middle pinnae oblong-lanceolate, 15–30 × 3.5–5.5cm, rounded-cuneate at bases, entire or serrate along margins,shortly caudate at apices; pinnae distally slightly smaller, terminalpinna of similar shape as middle ones, slightly larger, notsymmetrical at bases and long stalked. Veins visible adaxially,obviously raised abaxially, costae grooved adaxially, veinletsparallel to each other, veinlets oblique distally, proximal pairjoining into a triangular areole, distally every pair joining withexcurrent veinlet forming subsquare or rectangular areoles.Laminae greenish when dry, papery, adaxially with appressedshort hairs along costal grooves, elsewhere glabrous, abaxiallywith sparse short hairs along veins, occasionally with 1 or 2hairs and slightly foveolate on intercostal areas. Sori orbicular,attached on middle of veinlets and arranged in 2 rows betweencostules and not confluent; indusia small, shortly hairy, sporangiaalso hairy.Dense forests by streams in valleys, sparse forests on slopes;

THELYPTERIDACEAE 395100–500 m. Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan[Philippines].W. C. Shieh & J. L. Tsai (Fl. Taiwan, ed. 2, 1: 394. 1994) includedPronephrium gymnopteridifrons within P. asperum (C. Presl) Holttum(Blumea 20(1): 112. 1972; Goniopteris aspera C. Presl, Tent. Pterid.183. 1836, based on Polypodium asperum C. Presl, Reliq. Haenk. 1: 24.1825, not Linnaeus (1753); Abacopteris aspera (C. Presl) Ching; Dryopterispresliana Ching; Thelypteris aspera (C. Presl) K. Iwatsuki).17. Pronephrium macrophyllum Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 352. 1999.硕羽新月蕨 shuo yu xin yue juePlants ca. 1.5 m tall. Rhizomes not seen. Stipes 70–90 cm,stramineous, with appressed setae; laminae oblong, ca. 90 ×40–50 cm, 1-imparipinnate; lateral pinnae 8–11 pairs, alternate,obliquely spreading, sessile, proximal pair slightly reduced;middle pinnae oblong-lanceolate, 25–35 × ca. 6 cm, broadlycuneate at bases, obviously crenate along margins, acuminate atapices; distal pinnae becoming smaller; terminal pinna of similarshape and size as middle ones, bases not symmetrical andlong stalked. Veins visible adaxially, obviously raised abaxially,costae grooved adaxially, veinlets slightly raised, obliquelyspreading and parallel, proximal pair of veinlets joining intotriangular areoles, distally every pair joining with excurrentveinlet into subrectangular areoles. Laminae herbaceous whendry, green, greenish, or yellow-green, adaxially glabrous exceptfor appressed setae along grooves of costae, elsewhere, abaxialsurface with dense acicular long hairs along veins and alsoslightly foveolate on intercostal areas. Sori orbicular, attachedon middle of veinlets; indusia and sporangia hairy.● Forests by streams; 500–800 m. NW Guangxi, S Yunnan.18. Pronephrium hekouensis Ching ex Y. X. Lin, Fl. Reipubl.Popularis Sin. 4(1): 353. 1999.河口新月蕨 he kou xin yue juePlants 40–110 cm tall. Rhizomes long creeping, blackish,woody, with brown lanceolate scales. Fronds remote; stipes 25–60 cm, scaly at bases, distally with acicular hairs, stramineous;laminae broadly ovate or ovate-oblong, 20–50 × 15–30 cm, 1-imparipinnate; lateral pinnae 3–6 pairs, alternate, obliquelyspreading, shortly stalked, proximal pair shortened; middle pinnaeovate-lanceolate or narrowly lanceolate, 15–25 × 3–6.5 cm,cuneate at bases, undulate at margins, acuminate at apices; distalpinnae becoming smaller; terminal pinna slight larger thanmiddle ones, not symmetrical at bases, long stalked. Veins notevident adaxially, obviously raised abaxially, costae groovedadaxially, veinlets obliquely spreading and parallel to eachother, proximal pair of veinlets forming a triangular areole, nextpair joining with excurrent veinlet into subsquare or rectangularareoles. Laminae papery or herbaceous when dry, green,greenish, or grayish green, adaxially with dense appressed setaealong grooves of costae, sparsely hairy along veins and veinlets,abaxial surface with acicular hairs and multicellular articulatehairs along veins and intercostal areas, ± foveolate on intercostalareas. Sori orbicular, attached on middle of veinlets andusually confluent when mature; indusia and sporangia denselyhairy.● On slopes or in forests by streams; 100–500 m. Hainan, Yunnan.18. DICTYOCLINE T. Moore, Gard. Chron. 1855: 854. 1855.圣蕨属 sheng jue shuLin Youxing ( 林尤兴 ); Kunio IwatsukiPlants medium-sized, terrestrial. Rhizomes short and erect or ascending, including stipe bases with sparse scales; scales lanceolate,dark brown, thick, margins setaceous. Fronds clustered; stipes grayish stramineous, grooved adaxially, throughout with hairs;laminae elliptic or triangular, cordate at bases, 1-pinnate or pinnatifid, or simple, acuminate at apices; laminae if pinnate then with 1–6 pairs of lateral pinnae, pinnae broadly lanceolate, bases rounded, symmetrical, margins entire, free or adnate to rachises, obliquelyspreading, apices acuminate; costae raised on both sides, veinlets evident, oblique distally and reaching margins, reticulate, thick andevident, areoles in 3 or 4 rows, slightly tetragonal or pentagonal, without included veinlet or simple or forked included veinlets. Laminaepapery, dark brown when dry, rough, with dense hooked thick hairs on both surfaces. Sori scattered on included veinlets, exindusiate;sporangia bearing erect acicular hairs near annuli; spores elliptic, echinate. x = 36.Four species: Bhutan, China, N India, Japan, Myanmar, Nepal, Thailand, Vietnam; four species (two endemic) in China.1a. Laminae pinnate, with 1–6 pairs of free lateral pinnae, without included veinlet in areoles.2a. Laminae ternate at top, free lateral pinnae 1–3(or 4) pairs, 3–4 cm wide, entire ................................................... 1. D. griffithii2b. Laminae pinnatifid at top, free lateral pinnae 4–6 pairs, ca. 2 cm wide, usually undulate ......................... 2. D. mingchegensis1b. Laminae pinnatifid or not lobed, ± with simple or forked included veinlets in areoles.3a. Laminae with long acicular hairs abaxially, transverse veins between lateral veins indistinct, few includedveinlets in rectangular areoles on both sides of rachises and costae, rarely joining into small square areoles ...... 3. D. wilfordii3b. Laminae puberulent abaxially only, or very few acicular hairy, transverse veins between lateral veinsdistinct, more included veinlets in larger areoles on both sides of rachises or costae, these includedveinlets joining again into smaller square areoles ............................................................................................... 4. D. sagittifolia1. Dictyocline griffithii T. Moore, Index Fil. 59. 1857.圣蕨 sheng jueAspidium griffithii (T. Moore) Diels (1899), not (Baker)Beddome (1876); Cyclosorus griffithii (T. Moore) C. M. Kuo;Gymnogramma griffithii (T. Moore) Hance; Hemionitis griffi-

396THELYPTERIDACEAEthii (T. Moore) J. D. Hooker & Thomson; H. griffithii var. pinnataHooker; Stegnogramma griffithii (T. Moore) K. Iwatsuki;Thelypteris griffithii (T. Moore) C. F. Reed.Plants 40–70 cm tall. Rhizomes short and ascending, includingstipe bases with sparse scales and dense acicular longsetae; scales lanceolate, reddish brown, thick, cetaceous alongmargins. Laminae narrowly elliptic, 20–35 × 12–19 cm, notnarrowed to bases, imparipinnate, caudate at apices; lateral pinnae2 or 3 pairs (sometimes 1 pair only), free, almost sessile,proximal pair not shortened and of similar shape and size asabove ones, narrowly lanceolate, bent distally, 10–15 × 3–3.5cm, bases rounded-cuneate or rounded, symmetrical, marginsentire, apices acuminate; terminal pinna ternate, bases cuneateor rounded-cuneate, stalk ca. 2 cm, one pair of lateral segmentof similar shape as proximal pinnae, middle segment larger, entire,acuminate at apices. Costae straight, raised on both sides,with dense thick setae, veinlets evident, oblique distally andreaching margins, veinlets reticulate between veinlets; areolesin 2 or 3 rows, subtetragonal or oblique-square, rarely pentagonal,without included vein. Laminae papery when dry, darkbrown, throughout with hairs, abaxial surface with acicularthick hairs along veins, adaxially sparsely setaceous. Sori scatteredalong reticulate veins; sporangia shortly stalked, each with3 or 4 erect setae near annuli; spores elliptic, echinate.Dense forests, shaded wet streamsides; 600–1400 m. Fujian,Guangxi, Guizhou, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [NIndia, Japan, Myanmar, Vietnam].2. Dictyocline mingchegensis Ching, Acta Phytotax. Sin. 8:334. 1963.闽浙圣蕨 min zhe sheng jueDictyocline griffithii T. Moore var. tenuissima Ching.Plants ca. 50 cm tall. Rhizomes short and ascending, withdense reddish brown setaceous lanceolate scales and grayishwhite acicular hairs. Fronds clustered; stipes ca. 20 cm, stramineous,with sparse acicular hairs, with 1 or 2 scales on bases;laminae narrowly oblong, 26–30 × 12–14 cm, not narrowed tobases, 1-pinnate, acuminate at apices; lateral pinnae 4–6 pairs,opposite, almost sessile, spreading, proximal pair not reduced,broadly lanceolate, 7–8 × ca. 2 cm, rounded at bases, entire or ±undulate, acuminate at apices; terminal pinna particularly large,bases decurrent, margins pinnatifid, apices acuminate; proximalsegments of similar shape as lateral pinnae. Costae raised onboth sides and with acicular hairs, veinlets evident, obliquedistally, veinlets reticulate between veinlets, areoles in 2 rows,subtetragonal, without included veinlets. Laminae papery, grassgreenwhen dry, abaxial surface with acicular setae along veins,adaxially glabrous or with 1 or 2 short sparse hairs along veins.Sori sparsely attached along veins.● Shaded wet places in valleys or forests; 300–900 m. Fujian,Jiangxi, Zhejiang.3. Dictyocline wilfordii (Hooker) J. Smith, Hist. Fil. 149. 1875.羽裂圣蕨 yu lie sheng jueHemionitis wilfordii Hooker, Fil. Exot. t. 93. 1859; Dictyoclinegriffithii T. Moore var. pinnatifida Beddome; D. griffithiivar. wilfordii (Hooker) T. Moore; H. griffithii (T. Moore) J. D.Hooker & Thomson var. pinnatifida Hooker; Stegnogrammagriffithii (T. Moore) K. Iwatsuki var. wilfordii (Hooker) K. Iwatsuki;Thelypteris griffithii (T. Moore) C. F. Reed var. wilfordii(Hooker) C. M. Kuo.Plants 30–50 cm tall. Rhizomes short and thick, ascending,with dense blackish brown lanceolate scales; scales acicularlong hairy along margins. Fronds clustered; stipes 17–30 cm,dark stramineous, firm, with dense scales and short setae andacicular long hairs on proximal part; laminae triangular, ca. 20× 17 cm, cordate at bases, pinnatifid nearly to rachises on proximalpart, distally lobate, undulate and acuminate at apices; lateralsegments 3 pairs, proximal pair largest, broadly lanceolate,ca. 9 × 2.5–3.5 cm, slightly bent distally, entire or undulate,acuminate at apices, connected with above pair (sometimesnearly free) by broad wing; other segments of similar shape butshortened distally, distalmost triangular; costae of segments allraised on both sides and with dense acicular hairs; lateral veinsdistinct, veinlets between lateral veins reticulate, areoles in 3rows, subtetragonal or pentagonal, usually with simple or forkedincluded veinlets. Laminae papery when dry, dark brown, abaxialsurface with acicular hairs along veins, adaxially with denseappressed setae. Sori sparsely attached along reticulate veinlets.Shaded wet places in valleys or forests; 100–1100 m. Fujian,Guangdong, Guangxi, Guizhou, Jiangxi, Sichuan, Taiwan, SE Yunnan,Zhejiang [Japan, Vietnam].4. Dictyocline sagittifolia Ching, Acta Phytotax. Sin. 8: 335.1963.戟叶圣蕨 ji ye sheng jueStegnogramma sagittifolia (Ching) L. J. He & X. C.Zhang.Plants 30–40 cm tall. Rhizomes short and ascending, withsparse linear-lanceolate scales; scales ciliate along margins.Fronds clustered; stipes 15–30 cm, with dense brown shortsetae; laminae hastate, ca. 17 × 11–13 cm, deeply cordate atbases, entire or sometimes undulate, shortly acuminate at apices;costae raised on both sides, lateral veins distinct, obliquelyspreading, with 5–7 distinct transverse veins between lateralveins divided into large rectangular areoles and divided againinto ca. 2 × 4 subtetragonal smaller areoles, with simple orforked included veinlets in smallest areoles. Laminae paperywhen dry, dark brown, adaxially with dense short pubescencealong costae and appressed short hairs on intercostal areas, abaxialsurface with dense short pubescence along costae andveinlets, sparsely pubescent along reticulate veinlets. Sori scatteredand attached along reticulate veinlets.● Evergreen forests, rock crevices; 400–700 m. Guangdong,Guangxi, Hunan, Jiangxi.

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