TECTARIACEAE [Draft: 24 Apr 2012]

三叉蕨科  san cha jue ke

Xing Fuwu (邢福武)[1], Yan Yuehong (严岳鸿)[2], Dong Shiyong (董仕勇)1,
Wang Faguo (王发国)1; Maarten J. M. Christenhusz[3], Peter Hovenkamp[4]

Ferns, terrestrial, 10–300 cm tall. Rhizome erect or ascending to creeping, short or long, stout or slender, scaly at apex; rhizome and basal stipe scales brown, linear or lanceolate, margins entire, finely toothed, or ciliate, membranous. Stipe yellow, brown, or black, scaly at base or sometimes throughout. Fronds tufted or approximate, monomorphic to strongly dimorphic, simple or pinnate to 4 times pinnate-pinnatifid, often triangular or pentagonal, usually decompound toward apices; rachises and costae usually (in most genera) covered with articulate multicellular (ctenitoid) hairs; veins free or variously anastomosing, included veinlets if present single or forked. Sori terminal on included free veins, dorsal on veins or compital on connected veins, usually orbicular, sometimes elongate, anastomosing in lines, in some species throughout abaxial surface of lamina when mature; indusiate or exindusiate; indusia if present orbicular-reniform, persistent or caducous. Spores ovoid or elliptic, monolete, perispore with winglike folds, cristate, echinate, verrucose, rugose, or spinose. n = 10, 40, 41.

Eight to 15 genera and ca. 300 species: pantropical; four genera and 41 species (seven endemic) in China.

The number of genera and species in this family has been in considerable flux. Molecular studies (Hasebe et al. 1995) have increased our understanding of many fern families, and the placement of many uncertainly placed genera has recently become more evident. The genera Ctenitis (incl. Ataxipteris), Dryopsis, and Lastreopsis, which were previously placed in Tectariaceae are now known to belong to the Dryopteridaceae (Hasebe et al., 1995; Smith et al., Taxon 55: 717. 2006; Liu et al., 2007) and are treated in that family in this flora. The two families have always been difficult to separate on a morphological basis, but molecular evidence shows that Tectariaceae in its strict sense is sister to a clade uniting Polypodiaceae, Oleandraceae, and Davalliaceae. The group directly sister to Tectariaceae is Lomariopsidaceae (incl. Nephrolepidaceae), which in turn forms a sister clade to Dryopteridaceae, together forming the Eupolypods I clade (Smith et al., loc. cit.: 707).

Few molecular studies have so far addressed the Tectariaceae as a whole and the generic delimitations are, therefore, far from established. Recent studies suggest that a number of segregate genera is deeply embedded within Tectaria, making it a somewhat polymorphic genus. Here, we have followed this broad concept of Tectaria, and included the genera Ctenitopis, Hemigramma, and Quercifilix. This is admittedly a conservative approach, but it will have to suffice for the purposes of the present treatment until further studies provide a better understanding and better established delimitation of the genera within Tectariaceae.

Pleocnemia and Pteridrys are maintained as separate from Tectaria s.l., because they show sufficient molecular and morphological distinctness. Pleocnemia was tentatively placed in Dryopteridaceae by Liu et al. (2007), but evidence for this is scarce, and we, therefore, maintain it here in Tectariaceae following Smith et al. (loc. cit.: 718), until further data are available.

Ching Ren-chang, Fu Shu-hsia, Wang Chu-hao & Shing Gung-hsia. 1959. Arthropteris. In: Ching Ren-chang, ed., Fl. Reipubl. Popularis Sin. 2: 318–319; Wu Shiewhung. 1999. Arthropteris. In: Wu Shiewhung, ed., Fl. Reipubl. Popularis Sin. 6(1): 151–153; Wang Chuhao. 1999. Aspidiaceae (excluding Ctenitis and Lastreopsis). In: Wu Shiewhung, ed., Fl. Reipubl. Popularis Sin. 6(1): 37–56, 59–103.

1a.       Plants epiphytic; rhizome long creeping; stipes articulate to a long stipelike phyllopodium  1. Arthropteris

1b.       Plants terrestrial; rhizome shortly creeping to erect; stipes tufted, not articulate.

2a.       Sinuses of lobes lacking teeth .............................................................................  4. Tectaria

2b.       Each sinus between two pinna- or pinnule lobes bearing a small tooth.

3a.       Veins along costae and costules anastomosing; fronds mostly 2- or 3-pinnate, cylindric glands many on abaxial surface of lamina .............................................................................  2. Pleocnemia

3b.       Veins free; fronds 1-pinnate, glands absent on adaxial surface of lamina ...........  3. Pteridrys

1. ARTHROPTERIS J. Smith in J. D. Hooker, Fl. Nov.-Zel. 2: 43. 1854.

爬树蕨属  pa shu jue shu

Xing Fuwu (邢福武), Wang Faguo (王发国); Peter Hovenkamp

Plants epiphytic, with long twining, wiry rhizomes. Rhizomes with peltate scales, with 2 rows of fronds. Fronds distant, articulate to phyllopodium; lamina pinnate; pinnae close together, subsessile. Venation free, 2 or 3 Χ forked. Sori orbicular, in a single row between costa and margin; indusia reniform; sporangia long stipitate, annulus with 10–13 thickened cells. Spores elliptic; perispore granulate. n = 41.

About 20 species: Africa, SW to E Asia, Australia, Madagascar, Pacific islands; one species in China.

1. Arthropteris palisotii (Desvaux) Alston, Bol. Soc. Brot., sιr. 2, 30: 6. 1956.

爬树蕨  pa shu jue

Aspidium palisotii Desvaux, Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 5: 320. 1811; Arthropteris guinanensis H. G. Zhou & Y. Y. Huang.

Rhizome long creeping, up to 2–3 m, ca. 2 mm in diam., covered with ovate, dark brown, fimbriate scales. Fronds 5–10 cm apart; stipe 1–2 cm; lamina lanceolate, 15–40 Χ 4–8.5 cm; pinnae 30–40 pairs, alternate, 2–4 Χ ca. 1 cm, articulate to rachis, sessile, base asymmetrical, usually auriculate on upper side, margin undulate or crenate, apex rounded; lower pinnae shortened, deflexed; rachis densely covered with brown glandular hairs and scales abaxially, shallowly grooved adaxially. Sori nearer margin than costa; indusia brown, orbicular-reniform, glabrous.

Climbing on tree trunks or rocks in forests; 200–600 m. Guangxi, Hainan, Taiwan, Yunnan [India, Indonesia, Japan, Malaysia, Philippines, Vietnam; Africa, Australia, Pacific islands (Polynesia)].

Hovenkamp and Miyamoto (Blumea 50: 306. 2005) noted that, although Nephrolepis obliterata (R. Brown) J. Smith (in Hooker, J. Bot. 4: 197. 1841; Nephrodium obliteratum R. Brown, Prodr. Fl. Nov. Holland. 148. 1810; Arthropteris obliterata (R. Brown) J. Smith) has often been treated as a synonym of A. palisotii, the type is clearly a Nephrolepis species.

2. PLEOCNEMIA C. Presl, Tent. Pterid. 183. 1836.

黄腺羽蕨属  huang xian yu jue shu

Dong Shiyong (董仕勇); Maarten J. M. Christenhusz

Plants terrestrial, (60–)100–200 cm tall. Rhizome erect or rarely creeping, rhizome apex and stipe base densely covered with linear scales; scales on stipe bases narrow, 1.5–3 cm, usually twisted, their edges entire or finely toothed. Fronds clustered; stipe usually dark brown at base and light brown distally. Lamina 2-pinnate to 4-pinnatifid, ovate to subpentagonal, widest at base; basal pinnae always largest, triangular, with greatly enlarged basiscopic pinnules; distal pinnae or pinnules adnate to rachis or costae and ± decurrent at their bases, pinnae and pinnules ± deeply lobed, with a tooth in each sinus between two lobes; veins along costae anastomosing, without free veinlets within areoles; lamina papery or rarely membranous, usually glabrous on both surfaces and with cylindric glands along costules and veins on abaxial surfaces; rachises and costae usually raised on both sides, ctenitoid hairs present on adaxial surfaces of rachises. Sori dorsal on free veins; indusia present or not; spores ellipsoidal to spheroidal, perispore compressed in winglike folds or cristate with echinulations. n = 41.

About 20 species: tropical Asia, from NE India, SE China, throughout Malaysia and the W Pacific to Samoa; two species in China.

1a.       Stipe base scales with toothed margin; sori indusiate; perispore sharply echinate
                                                                                                                                                             1. P. leuzeana

1b.       Stipe base scales with entire margin; sori usually exindusiate; perispore forming winglike folds  2. P. winitii

1. Pleocnemia leuzeana (Gaudichaud) C. Presl, Tent. Pterid. 184. 1836.

台湾黄腺羽蕨  tai wan huang xian yu jue

Polypodium leuzeanum Gaudichaud, Voy. Uranie, Bot. 361. 1829; Aspidium leuzeanum (Gaudichaud) Kunze; A. rufinerve Hayata; Pleocnemia cumingiana C. Presl; P. rufinervis (Hayata) Nakai; Tectaria leuzeana (Gaudichaud) Copeland.

Rhizome prostrate; stipe 90–100 cm; rhizome and stipe base scales 2.5–3 cm, ca. 2 mm wide at base, margins conspicuously toothed. Lamina 3- or 4-pinnatifid, 80–90 Χ ca. 100 cm; basal pinnae largest, ca. 50 Χ 30 cm, its basal basiscopic primary pinnule elongate, ca. 30 cm; pinnules of middle pinnae lobed more than 3/4 toward costules, lobes crenate at margin, sinuses between two lobes 3–5 mm wide, with a triangular tooth in each sinus; lamina papery, veins forming in one or two rows of costal areoles and one row of costular areoles, other veins free; costules and veins with yellow glandular hairs on abaxial surfaces. Sori medial, dorsal on free veins; indusia persistent; perispore densely spiny.

Forests. Taiwan [Indonesia, Philippines; W Pacific islands].

2. Pleocnemia winitii Holttum, Reinwardtia 1: 181. 1951.

黄腺羽蕨  huang xian yu jue

Aspidium submembranaceum Hayata; Pleocnemia hamata Ching & Chu H. Wang; P. kwangsiensis Ching & Chu H. Wang; P. submembranacea (Hayata) Tagawa & K. Iwatsuki.

Rhizome prostrate; stipe (20–)60–100 cm; rhizome and basal stipe scales ca. 1.5 cm, 0.5–0.1 mm wide at base, margins entire. Lamina 3- or 4-pinnatifid, (50–)120–200 Χ (50–)100–130 cm; basal pinnae largest, 30–60 Χ 20–40 cm, basal basiscopic primary pinnule elongate, 20–30 cm; pinnules of middle pinnae lobed 1/2–2/3 toward costules, lobes entire or crenate at margin, sinuses between two lobes 1–2(–4) mm wide, with a triangular tooth in each sinus; lamina papery, veins along costae and those at base of costules anastomosing, other veins free; costules and veins with many orange cylindric glands on abaxial surface. Sori medial, dorsal on free veins; indusia absent or rarely present and persistent; perispore having a continuous wing with some cross wings. 2n = 82*.

Forests; 100–1000 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan, Yunnan [NE India, Myanmar, Thailand, Vietnam].

Pleocnemia conjugata (Blume) C. Presl was reported from Hong Kong (Holttum, 1974) and later also reported from Hainan (Wu, Fl. Guangdong. 7: 248. 2006). While studying these specimens on which these reports were based, it was noticed that these specimens belong to P. winitii. They were confused with P. conjugata due to the presence of indusia. The presence or absence of indusia in P. winitii does not appear to be constant; specimens are not always exindusiate. Besides a difference in the perispore, the most obvious character lies in the stipe scales to distinguish between P. winitii (entire scales) and P. conjugata (conspicuously dentate scales).

3. PTERIDRYS C. Christensen & Ching, Bull. Fan Mem. Inst. Biol. Bot. 5: 129. 1934.

牙蕨属  ya jue shu

Dong Shiyong (董仕勇); Maarten J. M. Christenhusz

Plants terrestrial, 1–2 m tall. Rhizome creeping, ascending, or erect; rhizome apex and stipe base covered with lanceolate scales; scales brown, 0.8–1.5 cm, margins entire. Fronds close or clustered; stipe light to dark stramineous or brown. Lamina 2-pinnatifid at base, elliptic, oblong, or ovate-lanceolate, widest at or below middle; basal pair of pinnae often largest, lanceolate or broadly lanceolate, basiscopic lobes elongate or not; lateral pinnae not adnate to rachis, ± deeply lobed, with a tooth in each sinus between two lobes; veins free; lamina papery or herbaceous, both surfaces usually glabrous; rachises grooved adaxially, glabrous; costae raised on adaxial surface, glabrous or sometimes with sparse short hairs on adaxial surface. Sori terminal or dorsal on veins, medial, one line on either side of costules; indusia present, persistent or fugacious; perispore consisting of anastomosing wings. n = 41.

Seven species: tropical Asia, from NE India, Sri Lanka, S China, to Indochina, Malaysia, Borneo, Indonesia, and the Pacific islands; three species (one endemic) in China.

1a.       Basal pinnae stalked; stalks 1–4 cm; pinna lobes 3–5 cm; sori dorsal on veins, (10–)15–17 pairs on each lobe ............................................................................................................  1. P. cnemidaria

1b.       Basal pinnae subsessile; stalks 0.1–0.4 cm; pinna lobes 2–3 cm; sori terminal on veins, 4–8 pairs on each lobe.

2a.       Basal basiscopic lobes of basal pinnae not elongate, entire; costae with short hairs on abaxial surface    2. P. australis

2b.       Basal basiscopic lobes of basal pinnae elongate and pinnatifid; costae glabrous on abaxial surface         3. P. lofouensis

1. Pteridrys cnemidaria (Christ) C. Christensen & Ching, Bull. Fan Mem. Inst. Biol. Bot. 5: 136. 1934.

薄叶牙蕨  bao ye ya jue

Dryopteris cnemidaria Christ, Bull. Acad. Int. Gιogr. Bot. 20: 140. 1910.

Plants 1.2–1.5 m tall. Rhizome creeping or erect. Stipe stramineous, 0.6–1 m, sparsely scaly at base; rhizome and basal stipe scales appressed, lanceolate, 10–15 Χ 1–2 mm. Lamina 2-pinnatifid, ca. 1.2 Χ 0.6 m; lateral pinnae ca. 30 pairs; basal pinnae usually largest, broadly lanceolate, 25–46 Χ 4–10 cm, deeply lobed nearly to costae, stalks 1–4 cm, basal basiscopic lobes slightly elongate; pinna lobes 30–34 pairs, (2–)3–5 Χ 0.6–0.8 cm, margins ± crenate to crenate-dentate, with a sharp tooth in each sinus, apex acute; lamina papery, greenish when dry; costae glabrous; veins free, veinlets 15–18 pairs on each lobe, 2–4 times furcate. Sori dorsal on veinlets, (10–)15–17 pairs on each lobe; indusia persistent, glabrous.

Forests; 100–1300 m. Guangxi, Guizhou, Taiwan, Yunnan [India, Laos, Myanmar, Vietnam].

2. Pteridrys australis Ching, Bull. Fan Mem. Inst. Biol. Bot. 5: 142. 1934.

毛轴牙蕨  mao zhou ya jue

Pteridrys nigra Ching & Chu H. Wang.

Plants 0.7–1.5 m tall. Rhizome creeping. Stipe dark stramineous or brown, 0.3–0.6 m, densely scaly at base; rhizome and basal stipe scales lanceolate, 5–7 Χ 1–2 mm. Lamina 2-pinnatifid, 0.4–0.8 Χ 0.2–0.3 m; lateral pinnae 12–15 pairs; basal pinnae longest or shortest, lanceolate, 9–27 Χ 3–5 cm, lobed 1/2–2/3 toward costule, stalks 1–4 mm, basal basiscopic lobe slightly shortened; pinna lobes 20–26 pairs, 1.5–2 Χ 0.5–0.8 cm, margins slightly serrate, with a sharp tooth in each sinus, obtuse at apex; lamina thickly papery, brown when dried; costae with short hairs on abaxial surface; veins free, veinlets 8–10 pairs on each lobe, veinlets 2- or 3-furcate. Sori terminal on veinlets, 5–7 pairs on each lobe; indusia with long hairs, fugacious.

Forests along streams; 100–700 m. Guangdong, Yunnan [Laos, Malaysia, Myanmar, Thailand, Vietnam].

3. Pteridrys lofouensis (Christ) C. Christensen & Ching, Bull. Fan Mem. Inst. Biol. Bot. 5: 141. 1934.

云贵牙蕨  yun gui ya jue

Dryopteris lofouensis Christ, Bull. Acad. Int. Gιogr. Bot. 20: 143. 1910.

Plants 0.8–1.3 m tall. Rhizome ascending or suberect. Stipe brown, ca. 0.5 m, densely scaly at base; rhizome and basal stipe scales lanceolate, 6–8 Χ ca. 1 mm. Lamina 2-pinnatifid, 0.7–1 Χ 0.3–0.6 m; lateral pinnae ca. 15 pairs; basal pinnae usually largest, lanceolate, usually 20–25 Χ 3–4 cm, lobed 1/2–2/3 toward costule, stalks 1–2 mm, basal basiscopic lobes very elongate; pinna lobes 20–25 pairs, 2–3 Χ 0.5–0.8 cm, margins slightly serrate at edge, with a sharp tooth in each sinus, obtuse at apex; lamina herbaceous or thinly papery, brown when dry; costae glabrous; veins free, veinlets 7 or 8 pairs on each lobe, veinlets 2- or 3-furcate. Sori (sub-)terminal on veinlets, 5–7 pairs on each lobe; indusia persistent, glabrous.

* Forests, scrub; ca. 1200 m. Guizhou, Yunnan.

4. TECTARIA Cavanilles, Ann. Hist. Nat. 1: 115. 1799.

叉蕨属  cha jue shu

Xing Fuwu (邢福武), Yan Yuehong (严岳鸿); Maarten J. M. Christenhusz

Bathmium C. Presl ex Link; Cardiochlaena Fιe; Ctenitopsis Ching ex Tardieu & C. Christensen; Fadyenia Hooker; Hemigramma Christ; Lenda Koidzumi; Podopeltis Fιe; Quercifilix Copeland; Sagenia C. Presl.

Plants terrestrial. Rhizome erect or ascending to creeping, short or long, stout or slender, scaly at apex; rhizome and basal stipe scales brown, lanceolate, margins entire or ciliate, membranous. Stipe stramineous or brown to black, scaly at base or sometimes throughout. Fronds monomorphic to strongly dimorphic, simple or pinnate to multiple times pinnate-pinnatifid, often triangular or pentagonal, pinnae and lobes usually entire, never serrate; all axes hairy with articulate multicellular hairs; veins variously anastomosing, included veinlets if present single or forked. Sori terminal on included free veins, dorsal on veins or compital on connected veins, usually orbicular, sometimes elongate, anastomosing in lines, in some species throughout abaxial surface of lamina when mature; indusiate or exindusiate; indusia if present orbicular-reniform, persistent or caducous. Spores ovoid or elliptic, perispore verrucose, rugose, or spinose. n = 10, (40).

About 230 species: pantropical; 35 species (six endemic) in China.

1a.       Fertile lamina much contracted; sporangia not forming sori but dense along veinlets.

2a.       Lamina glabrous; plants usually large (Hemigramma group) ........................  13. T. harlandii

2b.       Lamina hairy; plants minute, fronds less than 20 cm (Quercifilix group) ........  35. T. zeilanica

1b.       Fertile fronds not or only slightly contracted; sori orbicular, separate, usually indusiate.

3a.       Veins free or partly anastomosing, but then only those veins along costae or costules connecting, areoles without included veinlets (Ctenitopsis group).

4a.       Basiscopic pinnules or lobes of basal pinnae not elongated, often shortened.

5a.       Fronds bipinnate at base; veinlets anastomosing and forming obvious areoles
................................................................................................................  2. T. chinensis

5b.       Fronds bipinnatifid at base; veinlets free or occasionally anastomosing.

6a.       Stipes and rachises deep castaneous to dark brown, glossy; basal pinnae not shortened; sori intramarginal, indusiate .....................................................................  26. T. sagenioides

6b.       Stipes and rachises dark brown, dull; basal pinnae shortened; sori terminal, exindusiate  30. T. subsageniacea

4b.       Basiscopic pinnules or lobes of basal pinnae obviously elongated.

7a.       Plants ca. 2 m or taller; basal lamina 4-pinnatifid to 4-pinnate, 2-pinnate or 3-pinnatifid toward apices.

8a.       Fronds glabrescent on both surfaces, dimorphic ........................................  16. T. ingens

8b.       Fronds densely hairy on both surfaces, homomorphic .............................  27. T. setulosa

7b.       Plants up to 1 m tall; basal lamina 3-pinnatifid to 3-pinnate, 2-pinnatifid upward.

9a.       Veinlets free.

10a.     Fronds triangular-ovate to ovate; pinnae 6–8 pairs, stipes and rachis glabrous or nearly so           6. T. dissecta

10b.     Fronds elliptic lanceolate to elliptic; pinnae 9–12 pairs, stipes and rachis densely hairy  17. T. kusukusensis

9b.       Veinlets anastomosing and forming obvious areoles.

11a.     Fronds homomorphic, 3-pinnate at base; stipes stramineous or brown to castaneous, scales deep brown, concolorous ...............................................................................  5. T. devexa

11b.     Fronds dimorphic, 2-pinnate at base; stipes stramineous, scales dark brown with a paler margin.

12a.     Sori only at tip of lobes ...................................................................  1. T. acrocarpa

12b.     Sori throughout .................................................................................  10. T. fuscipes

3b.       Veins fully anastomosing, areoles with included veinlets (Tectaria group).

13a.     Sori large, terminal on (included) veinlets, in 2 rows between lateral veins; indusia large, persistent.

14a.     Veins forming a single row of narrow areoles along pinna rachises; included veinlets simple (not forked) when present.

15a.     Fronds dimorphic; length of stipes up to ca. 2 Χ length of lamina ............  15. T. impressa

15b.     Fronds homomorphic; length of stipes equal to or shorter than length of lamina.

16a.     Stipes scaly throughout .......................................................................  11. T. griffithii

16b.     Stipes scaly only at base.

17a.     Rhizome and stipe base with blunt spines; pinnae distant ..............  24. T. remotipinna

17b.     Rhizome and stipe base without spines; pinnae closer ......................  3. T. coadunata

14b.     Veins inconspicuous; included veinlets forked.

18a.     Bases of lamina decurrent; stipes broadly winged nearly to base.

19a.     Lamina deeply pinnately lobed, oblong ovate ......................................  4. T. decurrens

19b.     Lamina simple, not lobed, lanceolate ...............................................  18. T. leptophylla

18b.     Bases of lamina not decurrent; stipes not winged.

20a.     Stipes and rachises black, glossy ...........................................................  8. T. ebenina

20b.     Stipes and rachises brown or stramineous, slightly glossy or dull.

21a.     Stipes stramineous; fronds small, deltoid-ovate, palmately 3–5-parted
.....................................................................................................  29. T. subpedata

21b.     Stipes brown to dark brown; fronds large, ovate-lanceolate, pinnately 1–3-parted.

22a.     Stipes densely scaly throughout .............................................................  7. T. dubia

22b.     Stipes subglabrous throughout except at base.

23a.     Pinnae 5 pairs or more; asymmetrical at base of basal pinnae; middle pinnae lanceolate, ca. 3 cm wide, pinnate-lacerate ......................................................  21. T. phaeocaulis

23b.     Pinnae 4–6 pairs; symmetrical at base of basal pinnae; middle pinnae triangular, ca. 8 cm wide, pinnatipartite .........................................................................  34. T. yunnanensis

13b.     Sori small, on anastomosing veinlets, in irregular rows between lateral veins; indusium small, caducous.

24a.     Stipes winged at least at apex.

25a.     Stipes winged almost throughout, almost to base; lateral pinnae 3 or 4 pairs, without gemmae         33. T. vasta

25b.     Stipes winged at apex at most up to middle; lateral pinnae 1 or 2 pairs, gemmate or not  9. T. fauriei

24b.     Stipes not winged.

26a.     Lamina pinnatifid at apex, different from basiscopic pairs.

27a.     Stipes and rachises castaneous to dark brown, slightly glossy ...................  25. T. rockii

27b.     Stipes and rachises stramineous, dull.

28a.     Rhizome long creeping; sori well distributed over pinnae ..............  31. T. subtriphylla

28b.     Rhizome short, ascending or erect; sori only near margins .........  12. T. grossedentata

26b.     Terminal pinna lanceolate or elliptic, separate, simple or trifurcate, similar to basiscopic pairs.

29a.     Rhizome long creeping; pinnae never forked.

30a.     Rhizome scales dark brown; stipes hairy; terminal pinna bases cuneate; lateral pinna bases asymmetrical ...........................................................................  14. T. herpetocaulos

30b.     Rhizome scales light brown; stipes glabrous, terminal pinna bases decurrent; lateral pinna bases symmetrical ....................................................................................  32. T. variabilis

29b.     Rhizome short, ascending or erect; pinnae usually forked.

31a.     Lamina once pinnate (basal pinnae forked).

32a.     Base of terminal pinna rounded or cordate ................................  22. T. polymorpha

32b.     Base of terminal pinna decurrent ...............................................  23. T. quinquefida

31b.     Lamina twice pinnate.

33a.     Stipes and rachis stramineous to light brown, dull .......................  19. T. luchunensis

33b.     Stipes and rachises brown or deep castaneous to black, glossy.

34a.     Stipes and rachis deep castaneous to black; exindusiate ................  28. T. simonsii

34b.     Stipes and rachis dark brown to deep castaneous, never black; indusiate  20. T. media

1. Tectaria acrocarpa (Ching) Christenhusz, Phytotaxa in press.

顶囊轴脉蕨  ding nang zhou mai jue

Ctenitopsis acrocarpa Ching, Acta Phytotax. Sin. 19: 124. 1981.

Plants terrestrial, ca. 50 cm tall. Rhizome erect, short, 1–1.5 cm in diam., densely scaly at apex; scales dark brown with narrow paler margin, narrowly lanceolate, ca. 5 mm, membranous, entire, acuminate. Fronds clustered; stipe dark stramineous, ca. 20 cm, ca. 3 mm in diam. at base, grooved above, with sparse scales at base, glabrous upward. Lamina subdimorphic, base tripinnatifid, bipinnatipartite toward apex, sage green when dried, elliptic-lanceolate, ca. 30 Χ 20 cm, herbaceous, abaxially glabrous, adaxially pubescent, apex acuminate; rachises stramineous, with sparse dark brown linear scales, rachises and costae densely clothed with articulate brown hairs; pinnae 7–10 pairs, subopposite, interval 2–3 cm; basal pinna pairs largest, obliquely triangular, 13–15 Χ 8–10 cm, acuminate, stalks ca. 1 cm, pinnatipartite incised to 3/4 toward costa, with a pair of separate large pinnules; middle pinnae lanceolate, ca. 12 Χ 4 cm, bases cuneate, apices acuminate, pinnatipartite to 3/4 toward costa; basal basiscopic pinnules lanceolate, ca. 8 Χ 2 cm, bases cuneate, apices acuminate, pinnatipartite to 2/3 toward costa; lobes 8–10 pairs, interval 1–1.5 mm, falcate-oblong, 8–10 Χ 6–7 mm, mucronate. Veins pinnate, branches 8 or 9 pairs, single or forked, free or forming angular areoles, conspicuous on both sides. Sori orbicular, 3 or 4 pairs terminal on veinlets at apical lobes; indusia brown, orbicular-reniform, membranous, caducous.

* Dense forests in valleys on limestone; 300–500 m. Yunnan (Hekou, Jinping, Mengzi).

2. Tectaria chinensis (Ching & Chu H. Wang) Christenhusz, Phytotaxa in press.

中华轴脉蕨  zhong hua zhou mai jue

Ctenitopsis chinensis Ching & Chu H. Wang, Acta Phytotax. Sin. 19: 124. 1981.

Plants terrestrial, 60–80 cm tall. Rhizome erect, short, stout, 1–1.5 cm in diam. Stipe deep brown, 35–40 cm, grooved above, sparsely scaly; scales light brown, lanceolate, 4–6 mm, membranous, entire, acuminate at apex. Lamina bipinnatifid to pinnate toward apex, light brown when dried, triangular-ovate, 35–50 Χ 25–30 cm, papery, with sparse articulate off-white hairs adaxially, occasionally with lanceolate brown scales, base cuneate, apex acuminate; rachises and costae stramineous, densely clothed with articulate off-white hairs; free lateral pinnules 4–6 pairs, subopposite, interval 2–3 cm, upward; basal pinnules ovate-lanceolate, large, ca. 20 Χ 7–8 cm, bases cuneate, apices acuminate, stalks 0.8–1 cm, pinnatipartite with a single pair of separate lobes, lobes 8–10 pairs, alternate, slightly oblique, broadly lanceolate, 5–8 Χ 1–1.5 cm, acuminate, basal pairs nearly free, serrate upward, interval 2–3 cm, conjoined with broad wings; middle pinnules lanceolate, 12–15 Χ 2.5–3 cm, bases adnate to rachis, decurrent ± beneath, apices acuminate, pinnatipartite, lobes falcate-lanceolate, 1–1.5 Χ 0.8–1 cm, acuminate, entire. Veinlets forming one row of areoles along rachises and costae, veins free toward apex, raised on both surfaces. Sori orbicular, central on veinlets, in one or two rows beside main veins.

* Dense forests; 100–1200 m. Yunnan (Hekou, Xichou).

3. Tectaria coadunata (J. Smith) C. Christensen, Contr. U.S. Natl. Herb. 26: 331. 1931.

大齿叉蕨  da chi cha jue

Sagenia coadunata J. Smith in Hooker, J. Bot. 4: 184. 1841; Aspidium coadunatum Wallich ex Hooker & Greville (1831), not Kaulfuss (1824); A. kwanonense Hayata; A. macrodontum (Fιe) Ching; A. pinfaense Christ; Pleocnemia kwantungensis (Ching) Ching; Sagenia macrodonta Fιe, nom. superfl.; Tectaria coadunata var. hirsuta Holttum; T. coadunata var. minor Holttum; T. consimilis Ching & Chu H. Wang; T. junlianensis Ching & Chu H. Wang; T. kwantungensis Ching; T. macrodonta (Fιe) C. Christensen.

Plants terrestrial, 30–100 cm tall. Rhizome shortly creeping or erect, thick, densely scaly at apex and stipe bases; scales stiff, dark brown with a light margin, lanceolate, 6–7 mm, entire. Fronds clustered; stipe stramineous to pale castaneous, glossy, 20–40 cm, glabrescent above. Lamina pinnatifid to quadripinnatifid, light green to brown when dried, deltoid, 20–40 Χ 20–30 cm, herbaceous to thickly papery, apex acute; rachises, costae, midribs of pinnules, adaxial surface of lamina and segments ± hairy; hairs articulate, coarse; basal pinnae asymmetrical, oblong-subdeltoid, subopposite, up to 10–25 cm, bipinnate to tripinnatifid, long stalked, middle pinnae alternate, 10–15 cm, 5–7 pairs below deeply lobed apical pinnae, oblong to oblong-lanceolate; pinnules of middle pinnae stalked, oblong-subtriangular, ca. 15 Χ10 cm, apices acuminate, deeply lobed; secondary pinnules falcate-lanceolate, bases crenate or pinnatifid, apices rounded; segments falcate-lanceolate, entire, obtuse. Veins copiously anastomosing, with included free veinlets. Sori orbicular, at apex of included veinlets, in a single row on each side of midrib of ultimate lobes, medial indusiate; indusia rather large, brown, entire, clypeate, membranous, glabrous or hairy. 2n = 80.

Dense forests; 500–2500 m. Guangdong, Guangxi, Guizhou, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, India, Kashmir, Laos, Madagascar, Malaysia, Nepal, Sri Lanka, Thailand, Vietnam].

Because Tectaria coadunata is variable in size, stipe color, and lamina shape, this has resulted in a large number of synonyms.

4. Tectaria decurrens (C. Presl) Copeland, Leafl. Philipp. Bot. 1: 234. 1907.

下延叉蕨  xia yan cha jue

Aspidium decurrens C. Presl, Reliq. Haenk. 1: 28. 1825; A. copelandii C. Christensen; A. heterodon Copeland (1905), not Schrader (1824), nor Blume (1828); A. pteropus Kunze; Cardiochlaena alata Fιe; Nephrodium decurrens (C. Presl) Baker; Sagenia decurrens (C. Presl) T. Moore; S. mammillosa T. Moore; S. pteropus (Kunze) T. Moore; Tectaria peralata Copeland; T. simulans Ching.

Plants terrestrial, 50–100 cm tall. Rhizome erect, short, thick, 1.5–2 cm in diam., densely scaly; scales stiff, brown, with pale ferruginous margins, 8–12 cm, entire, apices acute. Fronds clustered, dimorphic; stipe brown at base, stramineous upward, 35–60 cm, glabrescent above, winged nearly to base. Lamina deeply pinnately incised, deep green, oblong-ovate, 30–80 Χ 30–40 cm, thickly papery, both surfaces glabrous, base cuneate and long decurrent, apex acute, lobes narrowed in fertile fronds; rachises winged throughout, stramineous; costae and main veins raised abaxially; apical pinnae broadly lanceolate, 20–25 Χ 5–8 cm, gradually narrowing toward base, subentire or crenulate, apices caudate; lateral pinnae 3–8 pairs, opposite, ascending, broadly lanceolate, 15–20 Χ 3–5 cm, subentire or crenulate, apices caudate, interval 3–4 cm, narrowing toward base and coadnate to rachis, sessile, usually forked in each basal pinna. Veinlets forming hexagonal areoles, cross veins not distinct, included veinlets forked. Sori large, close to main veins, located at coupling veinlets, in two rows between adjacent main veins; indusia up to 1.2 mm in diam., glabrous, entire, persistent. 2n = 160.

On rocks or near streams in dense forests; below 1200 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Taiwan, Yunnan [India, Indonesia, Japan, Myanmar, Nepal, Philippines, Thailand, Vietnam].

5. Tectaria devexa (Kunze ex Mettenius) Copeland, Philipp. J. Sci., C 2: 415. 1907.

毛叶轴脉蕨  mao ye zhou mai jue

Aspidium devexum Kunze ex Mettenius, Ann. Mus. Bot. Lugduno-Batavi 1: 237. 1864; A. intermedium Mettenius (1858), not Willdenow (1810); A. membranaceum Hooker; Ctenitopsis devexa (Kunze) Ching & Chu H. Wang; Pleocnemia devexa (Kunze) Alderwerelt; P. kwangtungensis (Ching) Ching; P. membranacea (Hooker) Beddome; Sagenia gigantea (Blume) T. Moore var. minor Beddome; Tectaria kwangtungensis Ching.

Plants terrestrial, 50–70 cm tall. Rhizome erect, short, ca. 6 mm in diam., densely scaly at apex and stipe bases; scales deep brown, light glossy, lanceolate, 4–5 mm, membranous, entire, fibriform and curly at apices. Fronds clustered; stipe stramineous or brown to castaneous, 25–30 cm, 2–3 mm in diam. at base, grooved above, with sparse brown articulate hairs above, glabrous below. Lamina tripinnate-pinnatifid, bipinnatifid toward apex, green when dried, triangular, 25–40 Χ 20–25 cm, thinly papery, both surfaces with sparse brown articulate hairs, base subcordate, apex acuminate; rachises and main veins stramineous to brown stramineous, densely clothed with brown articulate hairs; pinnae 3–5 pairs, subopposite, interval 1–1.5 cm; basal pinnae largest, obliquely triangular, 12–14 Χ 7–9 cm, stalks ca. 1 cm, bases rounded-cuneate, apices long acuminate, pinnatipartite to form 2–4 pairs of separate large pinnules at base; middle pinnae lanceolate, 7–9 Χ 2–2.5 cm, bases cuneate and nearly symmetrical, apices long acuminate, pinnatipartite to 2/3 way toward costa, often with some subseparate basiscopic pinnules; basal basiscopic pinnules broadly lanceolate, ca. 8 Χ 2 cm, bases broadly cuneate, apices long acuminate, pinnatipartite to broadly winged along costa; lobes 8–15 pairs, interval 2–3 mm, upward, falcate-lanceolate, 10–15 Χ 4–5 mm, margin mucronate or undulate to toothed, ciliate. Anastomosing veins forming 1 row of areoles along pinna rachis and main vein, other veins free, conspicuous on both sides. Sori orbicular, terminal on veinlets, submarginal; indusia orbicular-reniform, membranous, hairy, entire, persistent.

On rocky patches in forest gaps, especially in limestone regions. Chongqing, Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, Yunnan [Indonesia, Japan (Ryukyu Islands), Malaysia, Philippines, Sri Lanka, Thailand, Vietnam; Pacific islands (Polynesia)].

6. Tectaria dissecta (G. Forster) Lellinger, Amer. Fern J. 58: 156. 1968.

薄叶轴脉蕨  bao ye zhou mai jue

Polypodium dissectum G. Forster, Fl. Ins. Austr. 81. 1786; Aspidium membranifolium (C. Presl) Kunze; Ctenitis angustodissecta H. Itτ; C. dissecta (G. Forster) H. Itτ; C. sasaki (Hayata) Ching; C. tenuifrons (Hayata) Ching; Ctenitopsis angustodissecta (Hayata) Ching; C. dissecta (G. Forster) Ching; C. membranifolia (C. Presl) Ching; C. sasaki (Hayata) Ching & Chu H. Wang; C. subfuscipes Tagawa; Dryopteris angustodissecta Hayata; D. dissecta (G. Forster) Kuntze; D. sasaki Hayata; D. tenuifrons Hayata (1914), not C. Christensen (1905); Lastrea dissecta (G. Forster) Carruthers; Nephrodium dissectum (G. Forster) Desvaux; N. membranifolium C. Presl.

Plants terrestrial, ca. 1 m tall. Rhizome erect, short, ca. 1 cm in diam., densely scaly at apex; scales brown with narrow paler margin, glossy, lanceolate, ca. 4 mm, membranous, entire, apices acuminate. Fronds clustered; stipe deep stramineous to brown, ca. 40 cm, 4–5 mm in diam. at base, grooved above, with sparse light brown articulate hairs, lower part clothed with brown scales similar to those of rhizome. Lamina tripinnatifid, bipinnatifid toward apex, brown when dried, triangular-ovate or elliptic-ovate, 50–60 Χ 30–40 cm, thinly papery, adaxially with sparse brown articulate hairs, base subcordate, apex acuminate; rachises, costae, and main veins brown, clothed with brown articulate hairs; pinnae 6–8 pairs, subopposite on lower part and alternate upward, interval ca. 2 cm; basal pair of pinnae largest, obliquely triangular, 16–20 Χ 12–16 cm, acuminate, stalks 8–10 mm (subsessile upward), pinnatifid to forming pairs of separate large pinnules; middle pinnae lanceolate, 15–18 Χ 5–6 cm, bases cuneate, apices long acuminate, pinnatipartite to 3/4 way toward costa, occasionally with subseparate pinnules; basal basiscopic pinnules lanceolate, 10–12 Χ 2.5–3 cm, bases cuneate, apices acuminate, pinnatipartite; lobes 10–12 pairs, interval 2–2.5 mm, light upward, oblong, 15–17 Χ 6–7 mm, obtuse, scalloped. Veins pinnate, free, veinlets forked. Sori orbicular, terminal on veinlets; indusia light brown, orbicular-reniform, hairy, entire, persistent.

Forest understories. Taiwan [India; SE Asia to Polynesia].

7. Tectaria dubia (C. B. Clarke & Baker) Ching, Sinensia 2(2): 23. 1931.

大叶叉蕨  da ye cha jue

Nephrodium cicutarium (Linnaeus) Baker var. dubia C. B. Clarke & Baker, J. Linn. Soc., Bot. 24: 417. 1888; Aspidium dubium (C. B. Clarke & Baker) Beddome; Tectaria jinpingensis Ching & Chu H. Wang.

Plants terrestrial, 1.5–2 m tall. Rhizome erect, short, stout, ca. 3 cm in diam., densely scaly at apex and stipe bases; scales stiff, dark brown, linear-lanceolate, 8–10 mm, thickly membranous, entire, apices long acuminate. Stipes clustered, dark brown to castaneous, ca. 1 m, 1–1.5 cm in diam., sparsely covered with light brown articulate hairs. Lamina tripinnatifid, dark green when dried, deltoid, ca. 1 Χ 1 m, papery, base slightly cordate, apex acuminate; rachises and costae dark brown, densely covered with articulate hairs; pinnae 6 pairs, oblique, opposite at base and alternate upward, interval ca. 10 cm; basal pinnae 40–50 cm, rather large, ovate-deltoid, stalks 5–6 cm, pinnules 8–10 pairs, broadly falcate-lanceolate, 20–25 cm, alternate, upward, interval 1.5–2 cm, pinnate with 1–3 separate pinnules, pinnatifid to broad wings of costae upward, segments 10–12 pairs, falcate-lanceolate, 3–5 Χ 0.8–1.5 cm, undulate or entire; lateral pinnae broadly lanceolate, subsessile, ca. 30 cm, bases cuneate, apices long acuminate, pinnate-lacerate to lanceolate acute lobes. Veinlets forming copious subhexagonal areoles, main veins and veinlets clothed with brown articulate hairs, included veinlets forked. Sori orbicular, on anastomosing veins, irregularly arranged beside main veins; indusia brown, reniform, membranous, entire, persistent.

Near streams in dense forests; 600–1000 m. Yunnan [Bhutan, NE India, Nepal, Vietnam].

8. Tectaria ebenina (C. Christensen) Ching, Sinensia 2(2): 18. 1931.

黑柄叉蕨  hei bing cha jue

Aspidium ebeninum C. Christensen, Bull. Acad. Int. Gιogr. Bot. 23: 138. 1913.

Plants terrestrial, ca. 1.5 m tall. Rhizome erect, short, stout, 2–3 cm in diam., scaly at apex and stipe bases; scales brown, lanceolate, 5–6 mm, entire, apices acuminate. Fronds clustered; stipe black, glossy, ca. 60 cm, ca. 1 cm in diam., upper side sparsely covered with light brown articulate hairs and glabrous beneath. Lamina tripinnate or quadripinnatifid, dark green when dried, deltoid, ca. 80 Χ60 cm, thinly papery, glabrous on both sides, apex acuminate; rachises, costae, and main veins black and glossy, densely covered with light brown articulate hairs; pinnae 6 pairs, oblique, basal ones opposite and alternate upward, interval ca. 6 cm; basal pinnae rather large, deltoid, 40–50 Χ 25–30 cm, stalks 3–5 cm, pinnules ca. 8 pairs, broadly lanceolate, 10–20 cm, alternate, upward, sessile, lower 1 or 2 pairs separate and adnate to costae, confluent with broad wings upward, interval 1–3 cm, pinnatifid, segments 5–10 pairs, falcate-lanceolate, 3–5 Χ1–1.5 cm, undulate or entire; lateral pinnae lanceolate, 12–15 cm, bases attenuate, narrowed to and adnate with rachises, apices acuminate, pinnatifid to falcate-lanceolate lobes. Veinlets copiously forming subhexagonal areoles, main veins and veinlets clothed with brown articulate hairs, included veinlets single or forked. Sori orbicular, terminal on included veinlets, in a single row along main veins; indusia brown, reniform, membranous, entire, persistent.

Dense forests; 1300–1600 m. Guizhou, Yunnan (Malipo) [N Vietnam].

9. Tectaria fauriei Tagawa, J. Jap. Bot. 14: 102. 1938.

芽孢叉蕨  ya bao cha jue

Anapausia bonii (Christ) Nakai; Gymnopteris bonii Christ; Tectaria fengii Ching & Chu H. Wang; T. gemmifera Ching & Chu H. Wang (1981), not (Fιe) Alston (1939); T. gymnosora Holttum; T. hainanensis Ching & Chu H. Wang; T. simulans Ching.

Plants 0.6–1 m tall. Rhizome erect, short, stout, 1–1.5 cm in diam., scaly at apex and stipe bases; scales (dark) brown, (linear-lanceolate or) lanceolate, 6–8 mm, thickly membranous, entire, apices long acuminate. Fronds clustered; stipe deep stramineous to brown, 40–70 cm, 4–5 mm in diam., bases densely scaly, often clothed with brown articulate hairs, glabrescent upward, winged in upper half or very narrowly nearly to base. Lamina simple or pinnatisect to odd-pinnate, green to deep green, deltoid-ovate, 30–40 Χ 20–30 cm, base slightly cordate, herbaceous, both surfaces glabrous; rachises and costae dark stramineous, costa raised and pubescent on abaxial surface, main veins raised abaxially, glabrous adaxially but sparsely covered with light brown articulate hairs abaxially, rachises winged throughout; terminal pinna oblanceolate or oblong, 15–18(–25) Χ 5–10 cm, larger than lateral ones, base attenuate, narrowed into winged rachis, entire or 3-lobed, apex acuminate to caudate; lateral pinnae 1–3 pairs, subopposite, sessile, ovate-lanceolate, 15–18 Χ 3–5 cm, oblique, subentire, interval 4–6 cm, bases cuneate-attenuate, apices acuminate to caudate; basal pinnae rather large, often with a gemma often forming broadly lanceolate basiscopic lobes; gemmae (when present) scaly, one or more, axillary on both sides of lamina from base to apex; rachises and costae deep stramineous, pubescent abaxially, glabrescent adaxially. Cross veins distinct, veinlets forming inconspicuous subhexagonal areoles, included veinlets simple or forked. Sori orbicular, terminal on included veinlets or anastomosing veins, in two irregular rows beside lateral veins, often contiguous-confluent at maturity; indusia brown, reniform, membranous, glabrous, entire, caducous.

Dense valley forests; 500–1000 m. Hainan, Taiwan, Yunnan [India (Assam), Japan (Ryukyu Islands), Peninsular Malaysia, Thailand, Vietnam].

The pinna shape of Tectaria fauriei varies greatly from lanceolate or broadly lanceolate to ovate or oblong. It usually bears one or more scaly gemma axillary on either or both sides of the lamina from the base to the apex. Ching described plants without gemmae as Tectaria hainanensis Ching & C. H. Wang, but gemmae are present in the type specimen of that species.

10. Tectaria fuscipes (Wallich ex Beddome) C. Christensen, Contr. U.S. Natl. Herb. 26: 290. 1931.

黑鳞轴脉蕨  hei lin zhou mai jue

Aspidium fuscipes Wallich ex Beddome, Ferns Brit. Ind. Suppl. 15. 1876; A. membranifolium (C. Presl) Kunze; A. membranifolium var. dimorphum (C. B. Clarke) Christ; Ctenitopsis glabra Ching & Chu H. Wang; C. fuscipes (Wallich ex Beddome) C. Christensen; C. membranifolia (C. Presl) Ching; Lastrea fuscipes (Wallich ex Beddome) T. Moore; Nephrodium fuscipes (Wallich ex Beddome) C. B. Clarke; N. membranifolium C. Presl; N. membranifolium var. dimorpha C. B. Clarke.

Plants terrestrial, 40–70 cm tall. Rhizome suberect or ascending, short, 1–1.5 cm in diam., densely scaly at apex; scales dark brown with narrow paler margins, glossy, lanceolate, 7–8 mm, membranous, entire, apices fibriform. Fronds clustered, dimorphic or subdimorphic, fertile fronds taller and narrower; stipe deep stramineous, 20–30 cm, ca. 3 mm in diam. at base, grooved above, sparsely covered with articulate light brown hairs, lower part clothed with dark brown scales similar to those of rhizome, becoming shorter and narrower upward. Lamina base tripinnatifid, bipinnatipartite upward, dark green when dried, elliptic-ovate, 25–40 Χ 15–20 cm, herbaceous, glabrous or pubescent, margins ciliate, apex acuminate; rachises stramineous, with sparse linear dark brown scales, rachises and costae densely clothed with articulate brown hairs or glabrescent abaxially; pinnae 3–10 pairs, subopposite toward base, apical ones alternate, oblique, interval 2–5 cm; basal pair of pinnae largest, obliquely triangular, 8–15 Χ 6–10 cm, acuminate, stalks ca. 1 cm, pinnatipartite to 3/4 way toward costa, bearing a pair of separate large pinnules; middle pinnae lanceolate, 6–10 Χ 3–4 cm, bases cuneate, apices acuminate, pinnatipartite to 3/4 way toward costa, occasionally with subseparate pinnules; basal basiscopic pinnules lanceolate, 6–8 Χ 1.5–2 cm, bases cuneate, apices acuminate, pinnatipartite to 1/2 to costa; lobes 8–12 pairs, interval 1–2 mm, falcate-oblong, 7–8 Χ 5–6 mm, mucronate. Veins pinnate, 8 or 9 pairs, forked, free or forming angular areoles, conspicuous on both sides. Sori orbicular, 5–7 pairs terminal on veinlets; indusia brown, orbicular-reniform, membranous, entire, persistent.

Forests; 100–400 m. Guangxi, Guizhou, Hainan, Taiwan, Xizang, Yunnan [N India, Indonesia, Myanmar, N Vietnam].

11. Tectaria griffithii (Baker) C. Christensen, Index Filic., Suppl. 3, 180. 1934.

鳞柄叉蕨  lin bing cha jue

Nephrodium griffithii Baker, Syn. Fil. 300. 1867; Aspidium griffithii (Baker) Beddome; A. malayense Christ; A. multicaudatum (C. B. Clarke) Beddome; Nephrodium multicaudatum C. B. Clarke; Sagenia griffithii (Baker) Beddome; Tectaria malayense (Christ) Copeland; T. multicaudata (C. B. Clarke) Ching.

Plants terrestrial, 1–1.2 m tall. Rhizome erect, short, thick, ca. 1.5 cm in diam., densely scaly at apex and stipes; scales stiff, dark brown, linear-lanceolate, up to 2 cm, entire, apices long acuminate. Fronds clustered; stipe castaneous, glossy, up to 60 cm, 8–10 mm in diam. Lamina 2-pinnate to 3-pinnatifid, brown when dried, pentagonal, ca. 60 Χ 60 cm, thinly papery, both surfaces glabrous, base cordate, apex acuminate; pinnae 4 or 5 pairs, subopposite, slightly oblique; rachises and costae brown-castaneous, occasionally with linear scales on abaxial side of rachises, densely hairy adaxially, sparsely hairy abaxially; basal pinnae largest, oblique-deltoid, 20–25 Χ ca. 20 cm, stalks ca. 3 cm, bases asymmetrical, rounded-cuneate, apices long acuminate, basiscopic pinnules obviously elongated and forming 1 pair of separate pinnules at base, pinnatipartite to broadly winged along costae; middle pinnae deltoid-lanceolate, 15–20 Χ 10–12 cm, bases symmetrical, rounded-cuneate, apices long acuminate, pinnatipartite to broadly winged along costae and forming falcate-lanceolate lobes; pinnules or lobes 8–10 pairs, interval 0.5–1 cm, sessile, basiscopic pinnules largest, falcate-lanceolate, 12–14 Χ 4.5–5 cm, pinnatipartite to 2/3; lobules 10 pairs, slightly oblique, interval ca. 2 mm, elliptic, 1.5–2 Χ ca. 1 cm, obtuse, entire. Veins copiously anastomosing, with narrow areoles beside rachises and costae, included veinlets single. Sori orbicular, terminal on included veinlets, in one row on each side of midrib of ultimate lobes; indusia brown, large, clypeate, membranous, entire, persistent. 2n = 80.

Forest understories in valleys; 100–800 m. Guizhou, ?Yunnan [Borneo, Cambodia, India (Assam, Sikkim), Indochina, Indonesia, Malaysia, Myanmar, Philippines, Singapore, Thailand, Vietnam].

Tectaria griffithii is listed for China here with doubt. One of us (Christenhusz) studied a single specimen from China without precise locality (Matthew 110, K). Wu et al. (Bull. Dept. Biol. Sun Yatsen Univ. 3: t. 23. 1932) recorded the species from Guangxi, but the description and plates represent Dictyocline wilfordii (Thelypteridaceae) and not a Tectaria. Ching (1931) recorded the species from Guizhou (Tatsing, Esquirol 2253), but we have not found these specimens. The Flora of Taiwan does not list this species, and Holttum (Fl. Malesiana, ser. 2, 2: 54–55. 1991) does not list it for China.

12. Tectaria grossedentata Ching & Chu H. Wang, Acta Phytotax. Sin. 19: 127. 1981.

粗齿叉蕨  cu chi cha jue

Plants terrestrial, 60–90 cm tall. Rhizome erect or ascendant, short, stout, densely scaly at apex and stipe bases; scales dark brown, linear-lanceolate, ca. 5 mm, membranous, entire, apices capillaceous. Fronds clustered; stipe stramineous, 30–50 cm, 5–6 mm in diam. at base, grooved above, sparsely covered with light brown and articulate hairs, glabrous beneath. Lamina 2-pinnate to 2-pinnatifid at base, pinnate upward, dark green when dried, deltoid-ovate, 35–45 Χ 25–40 cm, papery, abaxially glabrous, adaxially hairy, base slightly cordate, apex acuminate; rachises and costae stramineous, densely hairy; lateral veins slightly flexuose, raised below, sparsely covered with light brown articulate hairs; pinnae 2 or 3 pairs, subopposite at base, alternate upward; basal pinnae largest, stalks 2.5–4 cm, deltoid, ca. 22 Χ 10–12 cm, pinnate with 2 or 3 separate pinnules, basal basiscopic pinnules up to 12 cm, terminal pinnules 13–15 Χ ca. 6 cm, bases cuneate, apices long acuminate; secondary pinnae broadly falcate-lanceolate, 15–20 Χ 3–5 cm, stalk ca. 1 cm, bases cordate, apices long acuminate, margins coarsely sinuate to sinuate-dentate, dentation (lobes) triangular; terminal pinna hastate, up to 20 cm at apex, base cuneate to slightly decurrent, apex long acuminate, pinnatilobed, basal lobes ovate-lanceolate. Veinlets forming subhexagonal areoles with cross veins, included veinlets forked, obscure adaxially, slightly raised abaxially. Sori orbicular, small, located at coupling veinlets, in irregular rows between adjacent lateral veins, near margin of lobes; indusia brown, small, clypeate, entire, persistent.

* Wet forests beside streams; sea level up to ca. 400 m. Guizhou, Yunnan.

13. Tectaria harlandii (Hooker) C. M. Kuo, Taiwania 47(2): 173. 2002.

沙皮蕨  sha pi jue

Acrostichum harlandii Hooker, Sp. Fil. 5: 274. 1864; Anapausia bonii (Christ) Nakai; A. harlandii (Hooker) Nakai; Gymnopteris bonii Christ; G. decurrens Hooker (non Aspidium decurrens C. Presl); Hemigramma decurrens (Hooker) Copeland; H. distinctipetiolata Ching; Leptochilus harlandii (Hooker) C. Christensen; Polypodium hainanense C. Christensen.

Plants terrestrial, 30–70 cm tall. Rhizome ascending to creeping, short, 1.5–2 cm in diam., roots lignified, densely scaly at apex and stipe bases; scales brown, glossy, linear-lanceolate, up to 1 cm, membranous, margins densely ciliate, apices long acuminate. Fronds clustered, dimorphic; sterile fronds in rosettes in juvenile stage, stipe dark stramineous or brown to castaneous, slightly glossy, 10–25 cm, 3–4 mm in diam. at base, glabrous, grooved above, narrowly winged toward apex. Lamina simple, trifoliate, or odd-pinnate, dark green when dried, ovate, 20–35 Χ 20–25 cm, thickly papery, both surfaces glabrous, base decurrent or not, apex acuminate; rachises and costae dark stramineous, slightly grooved above, glabrous on both sides; terminal pinna larger, broadly lanceolate, ca. 20 Χ 5–6 cm, subsessile or with a stalk up to 1 cm, base cuneate and decurrent, apex long acuminate, entire to undulate; lateral pinnae 1–3 pairs, opposite, interval 5–6 cm, slightly oblique, subsessile, lanceolate, 15–20 Χ 3–4 cm, bases cuneate and decurrent and forming narrow wings, margin entire or slightly undulate, apices long acuminate. Fertile fronds elongate and narrow; stipe up to 40 cm, glabrous; lamina narrow, trifoliate or odd-pinnate, pinnae linear-lanceolate, 8–10 Χ ca. 2 mm, sessile, entire or undulate. Veins anastomosing forming copious areoles, including veinlets forked, raised on both sides, glabrous. Sori orbicular, anastomosing in line along areoles, throughout abaxial surface when mature; exindusiate.

On wet mossy rocks in forests or along streams; 100–700 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan, Yunnan [Japan (Ryukyu Islands), Vietnam].

Tectaria harlandii is commonly segregated into the genus Hemigramma, because of its dimorphic leaves with contracted fertile fronds and linear sori that cover the entire abaxial surface of the blade when mature. Molecular evidence has shown however that this species is part of a greater Tectaria clade and cannot be maintained in a separate genus. The characters separating Hemigramma from Tectaria are minor and are known to occur in other species of Tectaria in other parts of the world.

14. Tectaria herpetocaulos Holttum, Dansk. Bot. Ark. 23: 241. 1965.

思茅叉蕨  si mao cha jue

Tectaria polymorpha (Wallich ex Hooker) Copeland var. subcuneata Ching & Chu H. Wang; T. simaoensis Ching & Chu H. Wang.

Plants terrestrial, 0.8–1 m tall. Rhizome creeping, long, 0.8–1 cm in diam., scaly at apex and stipe bases; scales dark brown and light glossy, pale along margins, lanceolate, 2–3 mm, thickly membranous, subentire, apices acuminate, caducous downward. Fronds widely spaced; stipe stramineous, 30–60 cm, 4–5 mm in diam., clothed with light brown articulate hairs above, glabrescent beneath. Lamina odd-pinnate, dark green when dried, ovate to oblong, 30–45 Χ 25–30 cm, papery, both surfaces glabrous, margin cartilaginous; rachises, costae, and lateral veins stramineous, pubescent adaxially, glabrescent abaxially, grooved adaxially and raised abaxially; terminal pinna ovate to oblong, entire, ca. 20 Χ 7–9 cm, stalk 2–3 cm, base broadly cuneate, apex caudate; lateral pinnae 2–4 pairs, simple, ovate-lanceolate, 18–20 Χ 5–6 cm, slightly oblique, opposite, subentire, interval 2–5 cm, bases slightly asymmetrically cuneate, apices abruptly caudate; basal pinnae shortly stalked, pinnae sessile toward apices. Veinlets forming conspicuous subhexagonal areoles, raised on both sides, included veinlets single or forked. Sori orbicular, on anastomosing veins, in irregular 4–6 rows between lateral veins; indusia brown, orbicular, membranous, glabrous, entire, caducous.

Understories of dense forests; 600–1100 m. Yunnan [India, Malaysia, Myanmar, Thailand, Vietnam].

Tectaria herpetocaulos is very similar to T. polymorpha, but can be easily recognized by its creeping rhizomes and abaxially glabrous laminae; its pinnae are simple and broadly rounded at the base and somewhat reminiscent of a Cyclosorus (Thelypteridaceae).

15. Tectaria impressa (Fιe) Holttum, Kew Bull. 43: 483. 1988.

疣状叉蕨  you zhuang cha jue

Phlebiogonium impressum Fιe, Mιm. Foug. 5: 314. 1852; Aspidium immersum Hooker (1862), not Blume (1828); A. prominens Alderwerelt; A. variolosum Wallich ex Hooker; A. zollingerianum Beddome (1868), not Kunze (1846); Nephrodium variolosum (Wallich ex Hooker) Baker; Sagenia variolata T. Moore; Tectaria prominens (Alderwerelt) C. Christensen; T. variolosa (Wallich ex Hooker) C. Christensen.

Plants terrestrial, 40–80 cm tall. Rhizome creeping to suberect, short, stout, densely scaly at apex and stipe bases; scales stiff, brown, linear-lanceolate, 6–8 mm, membranous, ciliate, apices long acuminate. Fronds clustered, subdimorphic, fertile fronds rather tall but narrowed; stipe stramineous, 30–50 cm, throughout covered with light brown hairs. Sterile lamina bipinnatifid to tripinnatifid, deep green or brown when dried, pentagonal, 30–35 Χ 30–35 cm, thickly herbaceous to leathery, both surfaces pubescent; rachises and costae stramineous, densely covered with articulate hairs; terminal pinna pinnatifid, base cuneate to decurrent, apex acuminate; lateral pinnae 1–5 pairs, opposite, interval 4–6 cm, oblique; basal pinnae pinnate with 1–3 undulate or pinnatifid lobes, 15–20 cm, rather large, stalks 2–3 cm, lower lobes larger than upper ones; middle pinnae falcate-lanceolate, undulate to pinnatifid, stalks becoming shorter, oblique, 6–8 cm, apices long caudate. Veinlets forming copious subhexagonal areoles with cross veins, included veinlets simple or not, distinctly forming long areoles with arcuate veinlet beside pinna rachis and main veins. Sori orbicular, terminal on veinlets, in one row beside main veins; indusia brown, orbicular or reniform, firm, persistent.

Usually on limestone in dense forests; 100–500 m. Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [India, Indonesia, Laos, Malaysia, Nepal, Thailand, Vietnam].

16. Tectaria ingens (Atkinson ex C. B. Clarke) Holttum, Rev. Fl. Mal. 2: 503. 1955.

西藏轴脉蕨  xi zang zhou mai jue(C. ingens) or 粤北轴脉蕨 yue bei zhou mai jue(C. matthewii)

Nephrodium ingens Atkinson ex C. B. Clarke, Trans. Linn. Soc. London, Bot. 1: 526. 1880; Aspidium cadieri Christ; Ctenitopsis cadieri (Christ) C. Christensen ex Tardieu & C. Christensen; C. ingens (Atkinson ex C. B. Clarke) Ching; C. matthewii (Ching) Ching; Dryopteris dissecta (G. Forster) Kuntze var. ingens (Atkinson ex C. B. Clarke) C. Christensen; D. ingens (Atkinson ex C. B. Clarke) Ching; Tectaria matthewii Ching.

Plants terrestrial, up to 3 m tall. Rhizome erect, short, stout, 1.5–2.5 cm in diam., densely scaly at apex; scales brown, (broadly lanceolate or) lanceolate, 6–9 mm, membranous, entire, apices acuminate. Fronds clustered, usually dimorphic, fertile fronds obviously narrowed; stipe dark stramineous to light brown, 30–80 cm, 5–9 mm in diam. at base, grooved above, with sparse (light) brown articulate hairs, lower part densely clothed with brown scales, similar to those of rhizome, few upward. Lamina tripinnate to quadripinnatifid, bipinnatifid upward, green or brownish when dried, triangular-ovate, 50–120 Χ 40–80 cm, thinly papery to membranous, both surfaces glabrescent, adaxially sometimes with a few brown articulate hairs, base cordate, apex (long acuminate or) acuminate; rachises, costae, and main veins dark stramineous to brown, densely clothed with brown articulate hairs; pinnae 10–15 pairs, lower part subopposite, oblique, interval 2–2.5 cm; basal pinnae largest, triangular, 22–42 Χ 15–30 cm, stalks 2–5 cm (with shorter stalks or subsessile upward), bases (rounded-cuneate or) cuneate and asymmetrical to nearly symmetrical, apices (long acuminate or) acuminate, basiscopic pinnules longer; middle pinnae elliptic-lanceolate, 20–35 Χ 8–20 cm, terminal pinna (rounded-cuneate or) cuneate at base, apex (long acuminate or) acuminate, largest basal basiscopic pinnules (broadly lanceolate or) lanceolate, 12–20 Χ 3.5–7 cm, bases oblique cuneate, apices (long acuminate or) acuminate, bipinnate at base, bipinnatipartite to broad wings beside costa upward; ultimate pinnules 10–14 pairs, alternate, interval 1.5–4 mm, light upward, sessile and adnate to pinnule rachises, pair of basal pinnae larger, lanceolate, 2.5–4 Χ 1–1.5 cm, mucronate, basiscopic lobes decurrent at base, pinnatifid at base, scalloped or subentire upward; lobes 5–7 pairs, interval 1–1.5 mm, oblique, falcate-triangular, 3–4 Χ 3–4 mm, obtuse, entire, brown articulate ciliate on margin. Veins pinnate, free or anastomosing below indenture, veinlets 5–8 pairs, single or forked, inconspicuous above, conspicuous and with sparse brown articulate hairs below. Sori orbicular, 4–8 pairs, terminal on veinlets, in center of midrib and margin; indusia orbicular-reniform, membranous, glabrous, entire, persistent.

Forests; 1000–2500 m. Guangdong (Lianxian), Xizang [NE India, Malaysia, Vietnam].

Tectaria matthewii only differs in its smaller size and does not seem to form a separate species. It is therefore (tentatively) united here with T. ingens. The isotype of T. matthewii in K is sterile.

Holttum (1991) wrote that this species “is perhaps the nearest existing species to a prototype for the genus.”

17. Tectaria kusukusensis (Hayata) Lellinger, Amer. Fern J. 58: 157. 1968.

台湾轴脉蕨  tai wan zhou mai jue

Dryopteris kusukusensis Hayata, Icon. Pl. Formosan. 4: 157. 1914; Ctenitis kusukusensis (Hayata) H. Itτ; Ctenitopsis hainanensis Ching & Chu H. Wang; C. kusukusensis (Hayata) C. Christensen ex Tardieu & C. Christensen; C. kusukusensis var. crenatolobata Tagawa; C. tamdaoensis Ching; D. membranoides Hayata.

Plants terrestrial, 40–100 cm tall. Rhizome erect, short, 1–1.5 cm in diam., densely scaly at apex; scales dark brown with narrow paler margin, broadly lanceolate, 4–6 mm, rigidly membranous, entire, apices acuminate. Fronds clustered; stipe dark brown to dark stramineous, 20–50 cm, 3–5 mm in diam. at base, grooved above, sparsely to densely clothed with articulate brown to light gray hairs, lower part with sparse dark brown scales like on rhizome. Lamina tripinnatifid, bipinnatipartite upward, dark brown or green when dried, elliptic to elliptic-lanceolate, 30–50 Χ 15–25 cm, (thickly) papery, abaxially glabrous, adaxially with sparse caducous brown or light gray articulate hairs, base subcordate, apex long acuminate to caudate (then tail 2.5–3 cm, subentire); rachises, costae and main veins brown to dark stramineous, densely clothed with long, brown or gray articulate hairs; pinnae 10–12 pairs, subopposite on lower part, alternate upward, interval 4–5 cm; basal pinnae largest, obliquely triangular, 10–20 Χ 6–15 cm, stalk 3–5 mm (subsessile upward), (long) acuminate, pinnatipartite to forming 1–3 pairs of separate large pinnules at base; middle pinnae (linear) lanceolate, 8–17 Χ 2.5–3 cm, bases cuneate, apices long acuminate to caudate, pinnatipartite to 2/3–3/4 toward costa, often with subseparate pinnules below; basal basiscopic pinnules lanceolate, 4–12 Χ 1–2.5 cm, bases rounded-cuneate, mucronate, apices long acuminate to caudate, pinnatipartite to 2/3 toward costa; lobes 8–15 pairs, interval 1–3 mm, oblique, triangular-elliptic to broadly lanceolate, 6–10 Χ 3–7 mm, margin mucronate, entire or undulate to toothed, ciliate. Veins pinnate, free, forked or simple, inconspicuous above, raised below. Sori orbicular, 3–8 pairs terminal on veinlets near margin, regular in 1 or 2 rows beside main veins; indusia dark brown, orbicular-reniform, membranous, glabrous, entire, persistent.

Near streams in valley forests. Guangdong, Guangxi, Hainan, Taiwan, Yunnan [Vietnam].

Ctenitopsis hainanensis (in the above synonymy) appears to differ from Tectaria kusukusensis only in its smaller stature and caudate pinna apices. It may well be an unusual population in Hainan, but one of us (Christenhusz) has not found sufficient evidence for its status as a separate species and thus refrains from making a nomen novum in Tectaria for this taxon.

18. Tectaria leptophylla (C. H. Wright) Ching, Sinensia 2(2): 22. 1931.

剑叶叉蕨  jian ye cha jue

Nephrodium leptophyllum C. H. Wright, Bull. Misc. Inform. Kew 1906: 11. 1906; Aspidium leptophyllum (C. H. Wright) C. Christensen.

Plants terrestrial, 25–40 m tall. Rhizome erect, short, ca. 1 cm in diam., scaly at apex and stipe bases; scales brown, linear-lanceolate, 3–4 mm, entire, membranous, acuminate. Fronds clustered; stipe stramineous to light brown, 2–5 cm, ca. 2 mm in diam., grooved, broadly winged nearly to base, sparsely scaly toward apex. Lamina simple, dimorphic; sterile fronds broad, fertile fronds homomorphic but taller and narrowed, 35–40 Χ 3–4 cm, papery, deep green when dried; main veins and lateral veins stramineous, glabrous on both sides. Veinlets forming inconspicuous subhexagonal areoles, included veinlets forked. Sori orbicular, terminal on included veinlets, in 3–5 irregular rows between lateral veins; indusia brown, orbicular, membranous, entire, persistent or revolute.

Forests; 300–400 m. Yunnan [N Vietnam].

19. Tectaria luchunensis S. K. Wu, Acta Phytotax. Sin. 40: 535. 2002.

绿春叉蕨  lό chun cha jue

Plants terrestrial, 60–100 cm tall. Rhizome erect, short, thick, densely scaly at apex and stipe bases; scales dark brown, linear-lanceolate, ca. 10 Χ 1 mm, membranous, entire. Fronds clustered; stipe stramineous, 40–50 cm, 5–6 mm in diam. at base, grooved, pubescent throughout, densely scaly at base, sparsely scaly upward. Lamina bipinnate at base, pinnate upward, green when dried, ovate or deltoid-lanceolate, 30–55 Χ 20–25 cm, thinly papery, both surfaces glabrous, base cordate, apex acuminate to caudate; rachises and costae deep brown, densely clothed with brown articulate hairs above, sparsely so abaxially; pinnae 3 or 4 pairs, subopposite, interval 10–15 cm, terminal pinna 3-lobed or simple, terminal lobe oblong-lanceolate, 10–20 Χ 2–4 cm, undulate to pinnatilobed, base cordate to cuneate, shortly stalked, lateral lobes like terminal lobe, opposite, smaller; basal pinnae largest, 15–20 Χ 10–20 cm, ovate-lanceolate, stalk 1–3 cm, pinnate or 3-lobed, pinnules 1 or 2 pairs, alternate or opposite, interval 1–3 cm, lanceolate or falcate-lanceolate, margins undulate to pinnatilobed; middle pinnae pinnate or trifurcate to simple, stalks becoming shorter to sessile upward, margins pinnatilobed to undulate, broadly lanceolate or falcate-lanceolate, 19–21 Χ 2–4 cm, bases cuneate, apices caudate. Lateral veins pinnate, veinlets forming subhexagonal areoles, raised on both surfaces, included veinlets single or forked, glabrous. Sori small, orbicular, at middle of coupling veinlets, in 2 or 3 irregular rows between adjacent lateral veins; exindusiate.

* Evergreen broad-leaved forests; 800–1300 m. Yunnan (Lόchun).

20. Tectaria media Ching, Acta Phytotax. Sin. 8: 169. 1959.

中型叉蕨  zhong xing cha jue

Plants terrestrial, ca. 60 cm tall. Rhizome ascending or erect, short, densely scaly at apex and stipe bases; scales linear-lanceolate, 3–4 mm, thickly membranous, margins entire, apices acuminate. Fronds clustered; stipe dark brown to castaneous, glossy, 35–40 cm, 3–4 mm in diam. at base, grooved above, sparsely pubescent at base. Lamina bipinnatifid at base, pinnate upward, light brown when dried, deltoid, ca. 30 Χ 25 cm; terminal pinna trifurcate, ovate-deltoid, ca. 18 Χ 5–6 cm, papery, both surfaces glabrous, base rounded, margins lobed, apex acuminate; rachises castaneous, pubescent adaxially, glabrescent abaxially; costae castaneous at base, stramineous upward, densely hairy adaxially, glabrescent abaxially; terminal lobe ovate-lanceolate, lobed, basal lateral lobes longer, ca. 8 cm, falcate-lanceolate, opposite; lateral pinnae 2 pairs, opposite, slightly oblique, interval 2–3 cm, stalks ca. 2 cm, lateral pinnae subsessile upward; basal pinnae largest, deltoid, pinnatifid, ca. 18 Χ 12 cm, bases cordate, apices long acuminate, lobes falcate-lanceolate, undulate to slightly lobed; middle pinnae broadly falcate-lanceolate, ca. 15 Χ 3.5 cm, bases cuneate, margins undulate to slightly lobed, apices acuminate. Veinlets forming subhexagonal areoles with cross veins, raised on both surfaces, included veinlets single or forked. Sori small, orbicular, located at coupling veinlets, in irregular rows between adjacent lateral veins; indusia small, brown, membranous, persistent.

* Dense forests; ca. 700 m. Fujian, Guangdong, Guangxi, Hainan.

21. Tectaria phaeocaulis (Rosenstock) C. Christensen, Index Filic., Suppl. 3, 183. 1934.

条裂叉蕨  tiao lie cha jue

Aspidium phaeocaulon Rosenstock, Hedwigia 56: 345. 1915; Tectaria laciniata Ching.

Plants terrestrial, 56–140 cm tall. Rhizome erect, short, thick, ca. 1.5 cm in diam., densely scaly at apex and stipe bases; scales stiff, brown with narrow paler ciliate margins, linear-lanceolate, ca. 6 mm, apices acute. Fronds clustered; stipe dark brown to castaneous, ± glossy, 30–80 cm, 4–5 mm in diam. at base, glabrescent upward. Lamina bipinnate at base, deep green when dried, oblong-ovate, 45–60 Χ 30–40 cm, leathery, both surfaces glabrous; costa and costules raised and pubescent on both surfaces, castaneous; main veins paler, pubescent abaxially; terminal pinna acuminate and pinnate-lacerate, basal pinnae bipinnate to tripinnatifid; lateral pinnae 5–8 pairs, broadly lanceolate, subopposite but alternate upward, ascending, 15–20 Χ 3–5 cm, apices caudate, pinnate-lacerate, interval 2–3 cm; basal pinnules largest, triangular-lanceolate with stalk ca. 2 cm, pinnate or bipinnate with 2 or 3 pairs of separate lobes at base and pinnatifid-lacerate at apex; middle pinnules lanceolate, coadnate, pinnatifid-lacerate to lanceolate or triangular lobes. Veinlets forming subhexagonal areoles with cross veins, included veinlets forked. Sori large, terminal on included free veinlets, orbicular, in two irregular rows between adjacent main veins; indusia reflexed, small, pubescent, entire, persistent.

Near streams or in dense forests; up to ca. 800 m. Fujian, Guangdong, Guangxi, Hainan, Jiangxi, Taiwan, Yunnan [Indonesia, Japan (Ryukyu Islands), Thailand, Vietnam].

22. Tectaria polymorpha (Wallich ex Hooker) Copeland, Philipp. J. Sci., C, 2: 413. 1907.

多形叉蕨  duo xing cha jue

Aspidium polymorphum Wallich ex Hooker, Sp. Fil. 4: 54. 1862; A. nantoense Hayata.

Plants terrestrial, 0.5–1 m tall. Rhizome erect, stout, 1.5–2 cm in diam., scaly at apex and stipe bases; scales dark brown and light glossy, pale at margins, linear-lanceolate, 6–7 mm, membranous, ciliate, apices long acuminate and curled. Fronds clustered, subdimorphic; fertile fronds rather tall, slightly narrowed; stipe stramineous, 20–60 cm, 4–5 mm in diam., grooved, glabrescent. Lamina simple or trifurcate to odd-pinnate, dark green or brown when dried, ovate to oblong, 30–60 Χ 25–30 cm, papery, abaxially pubescent, adaxially glabrous; rachises, costae, and lateral veins stramineous, glabrous adaxially, densely hairy abaxially, grooved adaxially and raised abaxially; terminal pinna simple or trifurcate, oblong, 18–20 cm, 7–9 cm wide at middle, entire or undulate, stalks 2–3 cm, bases slightly narrowed, rounded-cordulate, apices abruptly caudate; lateral pinnae 1–4 pairs, simple or basal pairs lobed, ovate-lanceolate, 18–20 Χ 6–8 cm, oblique, opposite, undulate to entire, interval 2–8 cm, bases gradually tapering, asymmetrical, cuneate, apices abruptly caudate; basal pinnae shortly stalked, sessile upward. Veinlets forming conspicuous subhexagonal areoles, raised on both sides, included veinlets forked. Sori orbicular, terminal on included veinlets or anastomosing veins, in 3–5 irregular rows between lateral veins; indusia brown, orbicular, membranous, entire, caducous.

On rocks or wet soil in forested valleys; 800–1500 m. Guangxi, Guizhou, Hainan, Taiwan, Yunnan [Bhutan, Cambodia, India, Indonesia, Malaysia, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].

Tectaria polymorpha is diverse in lamina shape, varying from simple, ternate to odd-pinnate. It is similar to T. herpetocaulos, but can be readily distinguished from that species by its erect rhizomes.

23. Tectaria quinquefida (Baker) Ching, Sinensia 2(2): 26. 1931.

五裂叉蕨  wu lie cha jue

Nephrodium quinquefidum Baker, J. Bot. 28: 265. 1890; Aspidium esquirolii (Christ) C. Christensen; A. polysorum Rosenstock; A. quinquefidum (Baker) Diels; Sagenia esquirolii Christ; Tectaria hekouensis Ching & Chu H. Wang.

Plants terrestrial, 60–80 cm tall. Rhizome erect, short, stout, 1–2 cm in diam., scaly at apex and stipe bases; scales dark brown, lanceolate, 5–7 mm, membranous, entire, apices long acuminate. Fronds clustered; stipe deep stramineous to brown or reddish, 25–40 cm, 3–4 mm in diam., grooved, sparsely hairy. Laminae subdimorphic, fertile fronds rather tall and slightly narrowed, odd-pinnate, brown when dried, deltoid-ovate, 20–35 Χ 18–30 cm, herbaceous, both surfaces glabrous; rachises and costae light brown, glabrous adaxially, pubescent and raised abaxially; terminal pinna simple or 3-lobed; middle lobes broadly lanceolate, 16–22 Χ 4–8 cm, bases narrowed, slightly cordulate, apices acuminate to caudate, margins undulate to dentate, stalk ca. 1.5 cm, lateral lobes falcate-lanceolate, rather small; lateral pinnae of a single pair, opposite, subsessile, ovate-lanceolate, 14–15 Χ 4–5 cm, slightly oblique, subentire to undulate, bases asymmetrical and cuneate, apices acuminate; gemmae present, axillary. Veinlets forming subhexagonal areoles, inconspicuous adaxially, raised and pubescent abaxially, included veinlets forked. Sori orbicular, on anastomosing veins, in irregular rows between lateral veins; indusia dark brown, reniform, membranous, entire, caducous.

Understories of dense forests; 300–600 m. Guangxi, Guizhou, Yunnan [N Vietnam].

From Yunnan (Hekou), a form of Tectaria quinquefida is known without gemmae and glabrescent pinna rachises, which was described as Tectaria hekouensis Ching & Chu H. Wang. These characters are a regional form that may be recognizable at the variety level.

24. Tectaria remotipinna Ching & Chu H. Wang, Acta Phytotax. Sin. 19: 129. 1981.

疏羽叉蕨  shu yu cha jue

Tectaria viridifrons Ching & Chu H. Wang.

Plants terrestrial, 80–90 cm tall. Rhizome erect, thick, ca. 2 cm in diam., dilated and bluntly spiny, scaly on apex and stipe bases; scales dark brown, lanceolate, ca. 6 mm, subleathery, margins entire, apices long acuminate. Fronds clustered, stipe stramineous, up to 50 cm, obviously dilated and spiny at base, grooved above. Lamina 3-pinnatifid, 2-pinnatifid upward, light brown when dried, deltoid-ovate, 40–50 Χ 25–30 cm, thinly papery, base slightly cordate, apex acuminate; pinnae ca. 4 pairs, opposite, interval up to 8–9 cm; rachises stramineous, glabrous on both surfaces, hairy on adaxial side of costae and midribs of pinnules, glabrous abaxially; basal pinnae largest, asymmetrically deltoid, ca. 20 Χ 15 cm, bases rounded-cuneate, apices acuminate, long stalked (up to 3 cm), shortly stalked upward; bipinnate to tripinnatifid, with 1 or 2 pairs of separate pinnules at base; middle pinnae broadly lanceolate, 12–15 Χ 5–7 cm, bases cuneate, apices acuminate, pinnatipartite to broadly winged along costa; pinnules of basal pinnae ca. 8 pairs, opposite at base, alternate toward apex, interval 1–2 cm, shortly stalked at base, sessile or adnate to costa upward, basal basiscopic pinnules larger, broadly lanceolate, 10–12 Χ 4–5 cm; lobes ca. 10 pairs, slightly oblique upward, interval 2–3 mm, elliptic-lanceolate, 1.5–2 Χ ca. 1.2 cm. Veins copiously anastomosing to hexagonal areoles, included veinlets single if present inconspicuous and hairy adaxially and slightly raised and glabrescent abaxially, margins ciliate. Sori orbicular, terminal on included veinlets, in one row at each side of midrib of ultimate lobes; indusia brown, clypeate, membranous, entire, persistent. 2n = 80*.

* Along wet streams; ca. 1700 m. Yunnan (Jingdong, Wuliang Shan).

Tectaria remotipinna is similar to T. coadunata but can be easily differentiated by the blunt spines on its rhizome.

25. Tectaria rockii C. Christensen, Contr. U.S. Natl. Herb. 26: 331. 1931.

洛克叉蕨  luo ke cha jue

Tectaria burmanica Ching; T. kweichowensis Ching & Chu H. Wang; T. linloensis Ching & Chu H. Wang.

Plants terrestrial, 60–120 cm tall. Rhizome decumbent or ascending, short, densely scaly at apex and stipe bases; scales stiff, dark brown, linear-lanceolate, 10–15 mm, membranous, entire, apices long acuminate. Fronds clustered or close together; stipe glossy, 40–80 cm, 5–8 mm in diam. at base, grooved, scaly toward base, covered throughout with dark brown, articulate hairs. Lamina bipinnate to tripinnatifid, dark green when dried, deltoid-ovate, 40–60(–80) cm, thickly papery, abaxially glabrous, adaxially pubescent, base cordate, apex acuminate; rachises castaneous, costae and midribs brown to deep stramineous, densely covered with articulate hairs; terminal pinna acuminate, deltoid, ca. 20 Χ 15 cm, base broadly cuneate, pinnatifid or pinnate-lacerate, lobes falcate-lanceolate, subentire, apices caudate; basal pinnae bipinnate, ca. 10 Χ 20–30 cm, rather large, stalks 2–3 cm, bases acuminate with 2 or 3 separate lobes, lobes lanceolate, rather long beneath, undulate or pinnatifid; middle pinnae 3 or 4 pairs, lanceolate, subopposite or alternate upward, interval 8–12 cm, oblique; fertile pinnae homomorphic but smaller, stalks short below, sessile upward, pinnatifid, bases cordate, apices caudate. Veinlets forming irregular areoles, included veinlets forked, inconspicuous adaxially and raised abaxially. Sori small, orbicular, located at coupling veinlets, in irregular rows between adjacent main veins; indusia brown, orbicular, small, membranous, entire, persistent.

Limestone areas under dense forests; 700–1200 m. Guangxi, Guizhou, Hainan, Taiwan, Yunnan [Indochina, Myanmar, Thailand, Vietnam].

Tectaria rockii is characterized by its dark brown to castaneous stipe and pinnatifid terminal pinna, however, the lateral pinnae are variable. We confirm that T. linloensis and T. kweichouensis are synonyms, the types (in PE and K) of these both representing T. rockii. Tectaria linloensis has in the past been treated as a synonym of T. simonsii (Wang, FRPS 6(1): 89. 1999; Wu, Fl. Yunnan. 13: 2004), which is not correct.

26. Tectaria sagenioides (Mettenius) Christenhusz, Phytotaxa in press.

轴脉蕨  zhou mai jue

Aspidium sagenioides Mettenius, Abh. Senckenberg. Naturf. Ges. 2: 397. 1858; Ctenitis sagenioides (Mettenius) Copeland; Ctenitopsis obscura (Hooker) C. Christensen; C. sagenioides (Mettenius) Ching; C. sagenioides var. glabrescens Ching & Chu H. Wang; Dryopteris laokaiensis C. Christensen; D. sagenioides (Mettenius) Kuntze; D. sagenioides subsp. obscura (Hooker) C. Christensen; D. schizoloma (Alderwerelt) C. Christensen; Heterogonium sagenioides (Mettenius) Holttum; Lastrea melanopus (Hooker) Beddome; L. sagenioides (Mettenius) T. Moore; Nephrodium melanopus Hooker; N. obscurum Hooker; N. sagenioides (Mettenius) Baker; Phegopteris obscura Fιe ex Christ; P. schizoloma Alderwerelt; Polypodium obscurum Hooker; P. viscosum C. H. Wright.

Plants terrestrial, 70–80 cm tall. Rhizome erect, short, ca. 1 cm in diam., densely scaly at apex; scales brown, broadly lanceolate, 4–5 mm, membranous, margins entire, apices acuminate. Fronds clustered; stipe deep castaneous to black, glossy, 30–40 cm, grooved above, occasionally clothed with lanceolate brown scales at base, with sparse light brown articulate hairs upward. Lamina bipinnatifid, dark brown when dried, elliptic, 40–50 Χ 20–25 cm, herbaceous, with sparse articulate brown hairs on both surfaces or abaxially glabrescent and adaxially with a few hairs, base retuse, margins ciliate, apex acuminate; rachises deep castaneous to black, glossy, ± clothed with articulate brown hairs; costae brown at base, stramineous upward, clothed with brown, articulate hairs; lateral pinnae 12–15 pairs, subopposite or alternate upward, interval 1.5–2 cm, applanate, subsessile; basal pinnae not shortened, reflexed, falcate-lanceolate, 10–12 Χ 4–4.5 cm, bases cuneate, apices acuminate, pinnatipartite nearly to costa; lobes 12–15 pairs, undulate or serrate, basal lobes shortened, middle lobes elongate toward base of lamina, ultimate ones acuminate; upper pinnae lanceolate, 10–12 Χ ca. 3 cm at base, pinnatipartite to 2/3, lobes ca. 15 pairs, subapplanate, interval 2–3 mm, ovate, 1.2–1.5 Χ 4–5 mm, obtuse, entire to undulate. Veins free, pinnate, 5 or 6 pairs, veinlets forked. Sori orbicular, terminal on veinlets, in one row between main veins and margin; indusia brown, orbicular-reniform. 2n = 80.

Wet dense forests; 100–300 m. Guangxi, Hainan, Yunnan [India, Indonesia, Malaysia, Myanmar, Thailand, Vietnam].

A more glabrous form of Tectaria sagenioides was found in Guangxi (Jinxiu) and Yunnan (Hekou), as Ctenitopsis sagenioides var. glabrescens Ching & Chu H. Wang, but this form does not merit taxonomic recognition. Holltum (Kalikasan 4: 205–231. 1975) stated that this species (the type of Ctenitopsis) belongs to the genus Heterogonium, but no evidence is provided for this and the species (and the related T. subsageniacea) fits morphologically well in Tectaria.

27. Tectaria setulosa (Baker) Holttum, Kew Bull. 43: 479. 1988.

棕毛轴脉蕨  zong mao zhou mai jue

Nephrodium setulosum Baker, J. Bot. 28: 265. 1890; Ctenitopsis setulosa (Baker) C. Christensen ex Tardieu & C. Christensen; Dryopteris dissecta (G. Forster) Kuntze var. lepidota Christ; D. setulosa (Baker) C. Christensen.

Plants terrestrial, up to 2 m tall. Rhizome erect, short, stout, ca. 2 cm in diam., densely scaly at apex; scales deep brown with paler margin, broadly lanceolate to ovate-lanceolate, 1–1.5 cm, entire, apices acuminate. Fronds clustered; stipe dark brown, 60–80 cm, ca. 1 cm in diam. at base, grooved above, densely clothed with light brown articulate hairs, lower part densely clothed with brown scales like on rhizome, narrower and sparser upward. Lamina tripinnate to quadripinnatifid, brown when dried, triangular-ovate, 100–150 Χ 70–80 cm, papery, abaxially glabrous, adaxially hairy, base cordate, apex long acuminate; rachises, costae, and main veins dark brown, densely clothed with brown articulate hairs; pinnae ca. 15 pairs, lower part subopposite, oblique upward, interval 2–3 cm; basal pinnae largest, obliquely triangular, 40–45 Χ 30–35 cm, stalks 4–5 cm (with shorter stalks upward), bases cuneate and nearly symmetrical, apices long acuminate, basiscopic pinnules slightly longer; middle pinnae triangular-ovate, 35–40 Χ 25–30 cm, bases rounded-cuneate, apices long acuminate, basal basiscopic largest pinnules broadly lanceolate, 16–20 Χ 7–9 cm, bases cuneate, apices long acuminate, bipinnate at base, bipinnatipartite to broad wings beside costa upward; ultimate pinnules ca. 15 pairs, alternate, interval 6–8 mm, oblique, sessile and adnate to pinnule rachises, basal pair larger, lanceolate, 4–5 Χ 1–1.5 cm, basiscopic lobes decurrent at base, apices long acuminate, pinnatifid at base, scalloped or subentire upward; lobes 8–10 pairs, interval 1–1.5 mm, oblique, falcate-ovate, 4–6 Χ 3–4 mm, obtuse, entire, margins ciliate with brown articulate hairs. Veins pinnate, free, veinlets 5–7 pairs, simple or forked, slightly raised and with sparse brown articulate hairs on both sides. Sori orbicular, 4–6 pairs, terminal on veinlets, near margin; indusia brown, orbicular-reniform, membranous, glabrous, entire, persistent.

Forests; 300–600 m. Guangdong, Guangxi, Yunnan [N India, Malaysia, Myanmar, Vietnam].

28. Tectaria simonsii (Baker) Ching, Sinensia 2(2): 32. 1931.

燕尾叉蕨  yan wei cha jue

Nephrodium simonsii Baker, Syn. Fil., ed. 2, 504. 1874; Aspidium kwarenkoense Hayata; A. longicrure (Christ) Christ; A. simonsii (Baker) Beddome; A. subtriphyllum (Hooker & Arnott) Hooker var. ebenosum Nakai; Dryomenis kwarenkoensis (Hayata) Nakai; Sagenia longicruris Christ; Tectaria kwarenkoensis (Hayata) C. Christensen; T. longicruris (Christ) C. Christensen; T. subtriphylla (Hooker & Arnott) Copeland var. ebenosa (Nakai) Nemoto.

Plants terrestrial, 60–120 cm tall. Rhizome creeping, ascending, or erect, short, thick, densely scaly at apex and stipe bases; scales stiff, dark brown to nearly purple, 8–10 mm, linear-lanceolate, entire, apices long acuminate. Fronds clustered; stipe dark brown or castaneous to black, glossy, 40–60 cm, 4–5 mm in diam. at base, minutely brown pubescent throughout. Lamina bipinnate to tripinnatifid, deep green when dried, subpentagonal or triangular-ovate, 30–60 Χ 25–40 cm, papery, both surfaces glabrous; rachises castaneous to black, pubescent, costa and costules raised abaxially, castaneous, glabrescent; terminal pinna 3-lobed or simple; terminal lobe ovate-lanceolate, entire or undulate to pinnatifid, base cordate to cuneate, lateral lobes like terminal lobe, opposite, rather small; lateral pinnules 2 or 3 pairs, opposite, interval 4–6(–10) cm, oblique; basal pinnules pinnate or bipinnatifid, 10–15 cm, rather large, stalks long; middle pinnules 3-lobed to simple, becoming sessile toward apices, entire or undulate, broadly lanceolate, 8–10 cm, bases cordate, apices caudate. Veinlets forming subhexagonal areoles with cross veins, included veinlets single or forked. Sori small, orbicular, located at coupling veinlets, irregular rows between adjacent main veins; exindusiate.

Dense forests; 200–1300 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [India, Japan (Ryukyu Islands), Thailand, Vietnam].

Tectaria simonsii is characterized by its castaneous to black stipe and ternate terminal pinna. We do not follow the treatment in FRPS (6(1): 89. 1999) in accepting T. linloensis (see T. rockii, p.) and T. subtriphylla var. ebenosa. The only differences lie in minor characters of the terminal pinna, which are variable even within the same plant.

29. Tectaria subpedata (Harrington) Ching, Sinensia 2(2): 23. 1931.

掌状叉蕨  zhang zhuang cha jue

Nephrodium subpedatum Harrington, J. Linn. Soc., Bot. 16: 30. 1877; Aspidium morsei (Baker) C. Christensen; A. subpedatum (Harrington) Diels; N. morsei Baker; Sagenia subpedata (Harrington) Nakai.

Plants terrestrial, 30–45 cm tall. Rhizome creeping to ascending, short, thick, 1–2 cm in diam., densely scaly at apex and stipe bases; scales brown, lanceolate, 4–5 mm, entire, apices long acuminate. Fronds clustered; stipe brown at base, stramineous upward, 20–30 cm, ca. 3 mm in diam., sparsely hairy. Lamina palmately 3–5-lobed, dark green when dried, deltoid-ovate, 15–20 Χ 12–16 cm, papery, abaxially densely hairy, adaxially glabrous, base cordate; costae and lateral veins stramineous, glabrous adaxially, densely hairy abaxially; terminal lobe elliptic-lanceolate, 15–18 Χ 5–7 cm, base slightly narrowed, apex long acuminate, entire or undulate to pinnatilobed; secondary pairs of lobes broadly lanceolate, 8–9 Χ 3–4 cm, bases not narrowed, apices long acuminate, undulate or subentire; basal lobes falcate-lanceolate, 5–6 Χ 2–3 cm, bases rounded-cuneate, apices long acuminate, entire. Veinlets forming subhexagonal areoles, raised on both surfaces, hairy, included veinlets forked. Sori rather large, close to lateral veins, locate at coupling veinlets of areoles, in 2 regular rows between adjacent lateral veins; indusia brown, clypeate, membranous, entire, slightly involute, persistent.

Forests on limestone. Guangxi, Taiwan [Myanmar, Vietnam].

Tectaria subpedata was treated as a synonym of T. polymporpha in Flora Malesiana (Holttum, 1991). We have not seen the type from Taiwan, but we checked the specimens from Guangxi in IBSC. The characteristic of these specimens is obviously different from T. polymorpha in the size and location of sori, and the rhizome of T. polymorpha is never creeping, which is usually the case in T. subpedata.

30. Tectaria subsageniacea (Christ) Christenhusz, Phytotaxa in press.

无盖轴脉蕨  wu gai zhou mai jue

Aspidium subsageniaceum Christ, Bull. Acad. Int. Gιogr. Bot. 16: 240. 1906; Ctenitopsis austrosinensis (Christ) C. Christensen & Tardieu; C. subsageniacea (Christ) Ching; Dryopteris austrosinensis Christ; D. subsageniacea (Christ) C. Christensen; Heterogonium austrosinense (Christ) Tagawa; H. subsageniaceum (Christ) Holttum; Tectaria austrosinensis (Christ) C. Christensen.

Plants terrestrial, 150–200 cm tall. Rhizome erect, short, 2–2.5 cm in diam., densely scaly at apex and stipe bases; scales dark brown, broadly lanceolate, 10–12 mm, membranous, entire, apices long acuminate. Fronds clustered; stipe deep brown, glossy, 40–50 cm, ca. 1 cm in diam. at base, grooved above, with sparse broadly lanceolate brown scales upward. Lamina brown when dried, oblong, gradually narrowed toward apex, 100–150 Χ 30–45 cm, herbaceous, both surfaces glabrous, margins ciliate on teeth; rachises deep brown, with sparse linear scales adaxially, densely clothed with articulate brown hairs abaxially; costae and main veins densely clothed with articulate brown hairs adaxially, glabrescent abaxially; bases subcuneate, apices acuminate, bipinnatifid; pinnules ca. 20 pairs, alternate, applanate, subsessile, linear-lanceolate, middle pinnules 18–22 Χ 3–4 cm, pinnatipartite to 2/3, basal 2 or 3 pairs of pinnules slightly shortened and deflexed; lobes ca. 20 pairs, subapplanate, interval 2–3 mm, falcate-oblong, 1–1.5 Χ 7–8 mm, entire, obtuse or mucronate. Veins pinnate, 7–9 pairs, free or connected with veinlets from costa. Sori orbicular to elliptic, at middle of veinlets, in 2 irregular rows beside main veins; exindusiate.

Limestone gaps in dense forests; 100–1500 m. Guangxi, Guizhou, Yunnan [N Vietnam].

31. Tectaria subtriphylla (Hooker & Arnott) Copeland, Philipp. J. Sci., C, 2: 410. 1907.

三叉蕨  san cha jue

Polypodium subtriphyllum Hooker & Arnott, Bot. Beechey Voy. 256. 1838; Aspidium hokutense Hayata; A. subtriphyllum (Hooker & Arnott) Hooker; Lenda subtriphylla (Hooker & Arnott) Koidzumi; Nephrodium subtriphyllum (Hooker & Arnott) Baker; Sagenia subtriphylla (Hooker & Arnott) Beddome.

Plants terrestrial, 20–70 cm tall. Rhizome long creeping or ascendant, stout, densely scaly at apex and stipe bases; scales stiff, dark brown, linear-lanceolate, 3–4 mm, membranous, bases rounded or cordate, entire, apices long acuminate. Fronds widely spaced; stipe dark stramineous, 10–40 cm, ca. 3 mm in diam. at base, covered throughout with light brown jointed hairs. Laminae subdimorphic, fertile fronds rather tall but narrowed, pinnate to bipinnatifid at base, deep green when dried, triangular-pentagonal, 10–35 Χ 10–25 cm, papery, both surfaces glabrescent to densely hairy; rachises stramineous, densely covered with articulate hairs; costae and costules raised on both surfaces, pubescent abaxially; terminal pinna trifurcate or acuminate, base cordate or cuneate to decurrent, terminal lobe ovate-lanceolate, pinnatifid or pinnate-lacerate, lateral lobes opposite, lanceolate, undulate, apices caudate; lateral pinnae 1–3 pairs, opposite, interval 1.5–3 cm, oblique; basal pinnae pinnate, ca. 15 cm, rather large, stalks 1–2 cm, lobes undulate or with orbicular segments; middle pinnae simple, becoming sessile upward, entire, undulate or with orbicular segments, broadly lanceolate, 10–12 cm, bases cordate, apices caudate. Veinlets forming subhexagonal areoles with cross veins, included veinlets forked. Sori small, orbicular, located at coupling veinlets, in irregular rows between adjacent main veins; indusia brown, reniform, small, deciduous.

Dense forests, near streams; below 400 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [India, Indonesia, Japan (Ryukyu Islands), Myanmar, Sri Lanka, Thailand, Vietnam; Pacific islands (Polynesia)].

A hairier form of Tectaria subtriphylla is found on limestone rocks in Hainan (Changjiang, Danxian, San Ya).

32. Tectaria variabilis Tardieu & Ching in Lecomte, Notul. Syst. (Paris) 5(2): 81. 1936.

多变叉蕨  duo bian cha jue

Plants terrestrial, 50–60 cm tall. Rhizome long creeping, ca. 0.5 cm in diam., scaly at apex and stipe bases; scales light brown, linear-lanceolate, 4–5 mm, membranous, entire, apices acuminate. Fronds subclustered; stipe stramineous, 30–35 cm, ca. 3 mm in diam., glabrous, not winged. Lamina simple or odd-pinnate, light brown when dried, deltoid, 20–25 Χ 10–15 cm, papery, both surfaces glabrous; rachises, costae, and lateral veins stramineous, glabrous, raised on both surfaces; terminal pinna lanceolate, ca. 20 Χ 5–10 cm, stalks ca. 2 cm, base decurrent, apex long acuminate, subentire; lateral pinnae 1 or 2 pairs, opposite, oblique, sessile, without gemmae, ovate-lanceolate, lateral pinnae 1 or 2 pairs, opposite, upward, interval 2–3 cm, stalks 3–5 mm, lanceolate, 15–17 Χ 2–4 cm, bases cuneate, apices caudate, entire. Veinlets forming inconspicuous subhexagonal areoles, included veinlets single or forked. Sori orbicular, on anastomosing veins, in 2 or 3 irregular rows between lateral veins; indusia light brown, reniform, membranous, entire, persistent.

On rocks in dense forests; ca. 300 m. Hainan (Baoting) [N Vietnam].

33. Tectaria vasta (Blume) Copeland, Philipp. J. Sci., C, 2: 411. 1907.

翅柄叉蕨  chi bing cha jue

Aspidium vastum Blume, Enum. Pl. Javae 2: 142. 1828; A. alatum Hooker & Greville; Sagenia alata (Hooker & Greville) Beddome; S. vasta (Blume) T. Moore; Tectaria decurrentialata Ching & Chu H. Wang.

Plants terrestrial, 60–150 cm tall. Rhizome erect or suberect. Stipes clustered, densely scaly at base; scales deep stramineous to brown, dark brown, broadly lanceolate, 40–45 cm, ca. 1 cm in diam., membranous, with light brown articulate hairs, broadly winged nearly to base, entire, apices acuminate. Lamina odd-pinnatipartite, deep green when dried, oblong, 30–60 Χ 30–35 cm, thinly herbaceous, both surfaces glabrous; rachises and costae stramineous, glabrous adaxially, slightly pubescent abaxially; terminal lobe ovate-lanceolate, 25–30 Χ ca. 10 cm, base narrowed and decurrent to broad wings along entire rachis, apex acuminate, undulate to toothed, sessile; lateral lobes 1–4 pairs, homomorphic but smaller; opposite, oblique upward, sessile, without gemmae, broadly lanceolate, ca. 20 Χ 8 cm, bases narrowed and adnate to rachis, apices caudate, subentire or undulate, basal lobes lobed to form smaller lanceolate lobes. Veinlets forming conspicuous subhexagonal areoles, included veinlets forked, glabrous on both sides. Sori orbicular, on anastomosing veins, in irregular rows between lateral veins; indusia brown, orbicular, membranous, caducous.

Near streams in dense forests; 600–800 m. Yunnan (Cangyuan, Jinping, Luchuan) [N India, Indonesia, Malaysia, Thailand].

34. Tectaria yunnanensis (Baker) Ching, Sinensia 2(2): 24. 1931.

云南叉蕨  yun nan cha jue

Nephrodium yunnanense Baker, Bull. Misc. Inform. Kew 1906: 11. 1906; Aspidium yunnanense Christ.

Plants terrestrial, 1.5–2 m tall. Rhizome ascending, short, stout, densely scaly at apex and stipe bases; scales stiff, dark brown to castaneous, linear-lanceolate, entire, apices long acuminate. Stipes clustered, castaneous, glossy, 60–80 cm, 0.7–0.8 cm in diam. at base, glabrous. Lamina pinnatifid, dark green when dried, deltoid, 60–80 Χ ca. 50 cm, papery, base broadly cuneate, apex acuminate; rachises castaneous, glabrous adaxially and pubescent abaxially, with broad wings upward, costae and midvein castaneous to brown, with brown articulate hairs on both sides; pinnae 4–6 pairs, oblique, lower pinnae opposite, upper pinnae alternate, interval 6–8 cm; basal pinnae 30–35 cm, deltoid, stalks 4–5 cm, pinnatifid to broad wings along costae, lobes 8–10 pairs, sometimes with a pair of separate lobes, sessile, falcate-lanceolate, entire or undulate; 10–12 Χ 3–4 cm, alternate, interval ca. 1 cm; lateral pinnae broadly lanceolate, sessile, 25–30 Χ 7–8 cm, bases narrowed, apices acuminate, pinnatifid to broad wings along costae and forming falcate-lanceolate acute lobes, lower lobes decurrent and adnate to rachis. Veinlets forming copious subhexagonal areoles, included veinlets forked. Sori orbicular, terminal on veinlets, regularly beside main veins, near to main veins; indusia brown, orbicular, membranous, entire, often revolute, persistent.

Near streams in dense forests; 100–1400 m. Guangxi, Hainan, Sichuan, Taiwan, Yunnan [NE Vietnam].

Tectaria yunnanensis is similar to T. dubia, but it differs in the scaliness and color of the stipes and the lamina division.

35. Tectaria zeilanica (Houttuyn) Sledge, Kew Bull. 27: 422. 1972.

地耳蕨  di er jue

Ophioglossum zeilanicum Houttuyn, Nat. Hist. 14: 43. 1783; Acrostichum quercifolium Retzius; Gymnopteris quercifolia Bernhardi; Leptochilus zeilanicus (Houttuyn) C. Christensen; Quercifilix zeilanica (Houttuyn) Copeland.

Plants terrestrial, 10–20 cm tall. Rhizome ascending to long creeping, slender, 2–3 mm in diam., densely scaly; scales brown, glossy, lanceolate, ca. 3 mm, membranous, margins ciliate, apices fibriform. Fronds distant, interval 5–10 mm, obviously dimorphic; sterile fronds usually in rosette, not erect; stipe dark stramineous, 3–8 cm, slender, ca. 1.5 mm in diam. at base, grooved above, densely scaly at base; densely hairy throughout with long brown articulate hairs. Lamina green, triangular-ovate, 6–9 Χ 2.5–3.5 cm, herbaceous, hairy abaxially and along margins, base cordate to hastate, apex obtuse, undulate, trifoliate to pinnatifid, usually with 1 pair of separate pinnae; rachises and costae stramineous, hairy; costae and main veins stramineous, glabrous adaxially, hairy abaxially; lateral pinnae opposite, applanate, subsessile or shortly stalked, triangular, 1–2.5 Χ 1–2 cm, bases rounded cuneate to subcordate, asymmetrical, apices rounded, margins subentire or slightly undulate, sometimes bearing auricles basiscopically at base; terminal pinna oblong, base broadly cuneate, apex rounded, lobed at margins to 1/3 toward costa, lobes rounded-subdeltoid, entire. Fertile fronds erect; stipe slender, 20–25 cm, very sparsely scaly or glabrescent; lamina trifoliate, very contracted; terminal pinna linear, 50–70 Χ 2–3 mm, stalk ca. 1 cm, base cuneate, apex obtuse, margins undulate or lobed; lateral pinnae opposite, oblique, shortly stalked, linear, 10–20 Χ 2–3 mm, bases bearing short lobe basiscopically, apices obtuse, margins undulate. Veins anastomosing, forming copious areoles, with or without included veinlets, these simple or forked. Sori orbicular, anastomosing in line, throughout abaxial surface when mature; exindusiate.

On muddy rocks of forest understories, near streams, on steep banks; 300–1000 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [S India, Indonesia, Laos, Malaysia, Philippines, Sri Lanka, Thailand, Vietnam; Indian Ocean islands (Mauritius), Pacific islands (Polynesia)].

The unusual spelling “zeilanica” is published as such and is to be pertained. It should not be corrected into the more common “zeylanica” or “ceylanica.” Tectaria zeilanica was previously most commonly treated as the separate monotypic genus Quercifilix, but molecular evidence shows that this genus is embedded within the greater Tectaria clade. The species is easily recognized by its small size, its coarsely crenate sterile leaves (that somewhat resemble those of Quercus, hence the name), and its trifoliate, strongly contracted fertile leaves, with anastomosing sori that cover the entire lower side of the lamina when mature.



[1] South China Botanical Garden, Chinese Academy of Sciences, Longdong, Shahe, Guangzhou, Guangdong 510650, People’s Republic of China.

[2] Herbarium, School of Life Sciences, Hunan University of Science and Technology, Xiangtan 411201, People’s Republic of China.

[3] Botanical Garden and Herbarium, Finnish Museum of Natural History, PO Box 7, University of Helsinki, 00010 Helsinki, Finland.

[4] Nationaal Herbarium Nederland, P.O. Box 9514, 2300 RA Leiden, Netherlands.