DRYOPTERIDACEAE, pro parte [Draft]

CTENITIS (C. Christensen) C. Christensen in Verdoorn, Man. Pteridol. 544. 1938 [??Notul. Syst. (Paris) 7: 86. 1938].

肋毛蕨属  lei mao jue shu [??which]

三相蕨属  san xiang jue shu

Dryopteris subg. Ctenitis C. Christensen, Biol. Arb. Tilegn. E. Warming 77. 1911; Ataxipteris Holttum.

Dong Shiyong (董仕勇)1; Xing Fuwu (邢福武)1, Yan Yuehong (严岳鸿)2; Maarten J. M. Christenhusz3

Plants terrestrial, 10–140 cm tall. Rhizome short, erect to ascending, densely scaly at apex and at stipe bases. Fronds clustered; stipe dark brown to stramineous, scaly throughout; scales of two different types: (1) lanceolate, membranous, appressed and iridescent scales, and (2) subulate, stiff, spreading scales without obvious iridescence. Lamina oblong-lanceolate to ovate-triangular, broadest at base, 1–4-pinnate; basal pair of pinnae always largest, triangular or broadly lanceolate, basal basiscopic pinnules of basal pinnae usually much enlarged; distal pinnae or pinnules ± adnate to rachis or costae and ± decurrent at their bases; veins all free, veinlets on ultimate pinnules or lobes simple or (rarely) bifurcate to pinnate; lamina texture herbaceous or papery, rarely leathery, with ctenitoid hairs on adaxial surface and short glandular hairs on abaxial surface, or rarely glabrous; costae of pinnules prominent, always covered with ctenitoid (multicellular) hairs. Sori medial or rarely submarginal; indusia present or not, sometimes very small and hidden by maturing sporangia. Perispore echinate, coarsely folded, or tuberculate. x = 41.

100–150 species in tropics and subtropics of America, Africa, Asia, and Australia; 10 species in China (four endemic).

The genus Ataxipteris is sometimes kept separate (based on the absence of unicellular, pallid, opaque, cylindric glands on the indusia and lamina), separating Ataxipteris distinctly from SE Asian Ctenitis, but molecular and morphological studies have suggested this genus to be embedded in a larger Ctenitis (Hasebe et al. 1995 [??page citation], as C. sinii). Ctenitis is part of Dryopteridaceae and not closely related to Tectaria, which is more closely allied to Oleandraceae (Hasebe et al. 1995[??page citation]).

1a.     Scales on stipe, rachis, or at least those on abaxial surface of costae (ovate-)lanceolate.

2a.    Pinnae (except basal one) 1-pinnatifid; veinlets obscure, some veinlets between pinna lobes arising from costae; perispore coarsely folded .......................................................................  9. C. sinii

2b.    Pinnae (except basal one) 2-pinnatifid; veinlets distinct, no veinlets arising from costae; perispore coarsely folded or echinate.

3a.     Basal pinnae broadly lanceolate with symmetrical bases; perispore coarsely folded  5. C. guidianensis

3b.     Basal pinnae deltoid with asymmetrical bases, basiscopic 2 or 3 pinnules much enlarged; perispore echinate.

4a.     Sori indusiate, indusia usually fugacious or not well-developed; widespread in SE Asia  10. C. subglandulosa

4b.     Sori exindusiate, usually covered by scales; Hunan and Sichuan (endemic)  8. C. pseudorhodolepis

1b.     Scales on stipe, rachis, and on abaxial surface of costae linear-lanceolate to subulate (inrolled and hairlike) or nearly so.

5a.    Lamina with 2 or 3 pairs of free pinnae; pinnae shallowly lobed; some veinlets between pinna lobes arising from costae; perispore coarsely folded ..........................  2. C. dianguiensis

5b.    Lamina with 5–15 pairs of free pinnae; pinnae 1–3-pinnate; no veinlets or rarely a few veinlets arising from costae; perispore coarsely folded, tuberculate or echinate.

6a.     Stipe scales 1–3 mm, linear to subulate; sori exindusiate; perispore echinate.

7a.     Sori usually covered by scales .......................................  3. C. dingnanensis

7b.     Sori naked, not scaly ..................................................  6. C. jinfoshanensis

6b.     Stipe scales 3–8 mm, subulate; indusia persistent or partly fugacious; perispore coarsely folded or tuberculate.

8a.     Free pinnae (except basal ones) of same size, 1-pinnatifid, pinnae bases not broadened  7. C. mannii

8b.     Free pinnae gradually reduced toward lamina apex, 1- or 2-pinnatifid, nearly all pinnae with broadened bases.

9a.     Fronds > 1 m, 4-pinnate at base; stipe scales and rachis rufescent, 6–8 mm    1. C. decurrentipinnata

9b.     Fronds < 60 cm, 2- or 3-pinnate at base; stipe scales and rachis dark brown, 3–4 mm .........................................................................................  4. C. eatonii

 

1. Ctenitis decurrentipinnata (Ching) [Tardieu & C. Christensen, Notul. Syst. (Paris) 7: 86. 1938]; Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 291. 1938.

海南肋毛蕨  hai nan lei mao jue

Dryopteris decurrentipinnata Ching, Bull. Fan Mem. Inst. Biol. Bot. 2: 195. 1931.

Rhizomes erect or suberect; rhizome scales brown, narrowly lanceolate to linear-subulate, 1.5–2 mm. Stipe stramineous, 40–70 cm; stipe scales spreading, reddish brown, subulate, 6–8 mm; scales on rachises and costae alike but smaller. Lamina 3-pinnate or 4-pinnate-pinnatifid, triangular-lanceolate, 60–80 Χ 30–50 cm; pinnae 12–18 pairs; basal pinnae largest, triangular-lanceolate, 20–30 Χ 15–25 cm; pinnules 10–13 pairs; basal basiscopic pinnules of basal pinnae longest, 10–20 Χ 3–6 cm; base of upper pinnae or lobes decurrent with veinlets arising from rachis. Lamina herbaceous to thinly papery; veins free, distinct, no veinlets arising from costae except a few in distal part of costae. Sori medial or submedial, indusiate, indusia usually persistent. Perispore coarsely folded or tuberculate.

Forests; 500–1400 m. Hainan [Philippines, Vietnam].

2. Ctenitis dianguiensis (W. M. Chu & H. G. Zhou) S. Y. Dong, Novon 18: 35. 2008.

滇桂肋毛蕨  dian gui lei mao jue [??which]

滇桂三相蕨  dian gui san xiang jue

Ataxipteris dianguiensis W. M. Chu & H. G. Zhou, Acta Phytotax. Sin. 42: 376. 2004.

Plants 60–80 cm tall. Rhizome short, suberect or prostrate; scales at apex of rhizomes yellowish brown to dark brown, opaque, linear-lanceolate, 10–20 mm, margins entire, fimbriform and curly at apex. Stipe brown at base, dark stramineous toward apex, 25–40 cm, 2–3 mm in diam. at base, grooved above, densely clothed with subulate, 3–5 mm, dark brown, spreading scales; scales on rachises and main veins alike but smaller. Lamina bipinnatifid, ovate-triangular, 25–40 Χ 15–25 cm, decurrent toward apex; pinnae 2–4(or 5) pairs; basal pinnae largest, subopposite, triangular, 12–18 Χ 4–6 cm, bases asymmetrically rounded-cuneate, margins deeply lobed without free pinnules, apices caudate-acuminate; basal basiscopic lobes of basal pinnae longest, 4–5 Χ 1–1.5 cm; base of upper pinnae or lobes decurrent with veinlets arising from rachis. Lamina thickly papery to leathery, nearly glabrous on both surfaces, rachises and main veins stramineous, densely clothed with long, brown, articulate hairs and spreading, linear squamules; veins pinnate, free, distinct, with some veinlets arising from costae, simple or forked, rarely a few veinlets anastomosing. Sori round, medial or submedial, in two irregular rows between main veins, exindusiate. Perispore coarsely folded.

Dense broad-leaved forests on limestone; 500–1400 m. Guangxi, Hainan, Yunnan [Vietnam].

3. Ctenitis dingnanensis Ching, Acta Phytotax. Sin. 19: 122. 1981.

二型肋毛蕨  er xing lei mao jue

Rhizomes erect or suberect; rhizome scales reddish brown, narrowly lanceolate, 10–20 mm. Stipe stramineous, 13–21 cm; stipe scales spreading, brown, linear-subulate, 2–3 mm; scales on rachis and costae alike but smaller. Lamina 3- or 4-pinnatifid, triangular, 15–30 Χ 10–25 cm; pinnae 6–8 pairs; basal pinnae largest, triangular, 5–15 Χ 5–10 cm; pinnules 6–10 pairs; basal basiscopic pinnules of basal pinnae longest, 4–9 Χ 1.5–3 cm; base of upper pinnae or upper lobes not obviously decurrent, without veinlets arising from rachises. Lamina herbaceous; veins free, distinct, no veinlets arising from costae. Sori submedial, exindusiate, some sori covered by scales. Perispore echinate and thinly winged.

* Forests; 300–700 m. Guangdong, Jiangxi.

4. Ctenitis eatonii (Baker) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 291. 1938.

直鳞肋毛蕨  zhi lin lei mao jue

Nephrodium eatonii Baker, Syn. Fil. (Hooker & Baker) 276. 1867; Aspidium eatonii (Baker) H. Christ; A. leucostipes (Baker) H. Christ; Ctenitis changanensis Ching; C. confusa Ching; Dryopteris eatonii (Baker) Kuntze; D. leucostipes (Baker) C. Christensen; Nephrodium leucostipes Baker.

Rhizomes erect or suberect; rhizome scales dark brown, linear-subulate, 5–12 mm. Stipe stramineous, 5–35 cm; stipe scales spreading, dark brown, subulate, 2–4 mm; scales on rachis and costae alike but smaller. Lamina 2-pinnate to 3-pinnatifid, triangular-lanceolate, 7–35 Χ 5–15 cm; pinnae 5–12 pairs; basal pinnae largest, triangular-lanceolate, 2.5–13 Χ 1.5–7 cm; pinnules 2–10 pairs; basal basiscopic pinnules or lobes of basal pinnae longest, 0.8–4.5 Χ 0.4–1.5 cm; base of upper pinnae or upper lobes not obviously decurrent without veinlets arising from rachis. Lamina herbaceous; veins free, distinct, no veinlets arising from costae. Sori near central, near costa, indusiate, indusia usually persistent. Perispore coarsely folded or tuberculate.

Wet places in forests, usually in limestone areas; sea level up to 400 m. Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Taiwan [S Japan].

5. Ctenitis guidianensis H. G. Zhou & W. M. Chu, Acta Bot. Yunnan. 17: 424. 1995.

桂滇肋毛蕨  gui dian lei mao jue

Rhizomes suberect; rhizome scales dark brown, narrowly lanceolate, ca. 12 mm. Stipe brown to stramineous, 50–90 cm; stipe scales appressed, brown, ovate-lanceolate, ca. 3 mm; scales on rachis and costae alike but smaller. Lamina 2-pinnate to 3-pinnatifid, ovate-lanceolate, 50–80 Χ 20–50 cm; pinnae 6–11 pairs; basal pinnae as large as second, broadly lanceolate, 15–30 Χ 8–15 cm; pinnules 10–12 pairs; basal basiscopic pinnules of basal pinnae as long as acroscopic ones, 2–8 Χ 1–3 cm; bases of upper pinnae or lobes not obviously decurrent, without veinlets arising from rachis. Lamina thickly papery; veins free, distinct, no veinlets arising from costae except a few in distal part of costae. Sori medial or submedial, indusiate; indusia usually fugacious. Perispore with inflated folds.

* Forests on limestone; 800–1100 m. Guangxi, Yunnan.

6. Ctenitis jinfoshanensis Ching & Z. Y. Liu, Bull. Bot. Res., Harbin 4: 14. 1984.

金佛山肋毛蕨  jin fo shan lei mao jue

Rhizomes suberect or prostrate; rhizome scales reddish brown, narrowly lanceolate, ca. 10 mm. Stipe brown to stramineous, ca. 60 cm; stipe scales spreading, brown, subulate, 5–6 mm; scales on rachis and costae alike but smaller. Lamina 2-pinnate to 3-pinnatifid, broadly ovate, ca. 70 Χ 50 cm; pinnae 12 pairs; basal pinnae largest, triangular, 20–25 Χ 13–15 cm; pinnules 12–15 pairs; basal basiscopic pinnules of basal pinnae longest, 10–12 Χ ca. 3 cm, ultimate pinnule or pinna lobe oblong, obtuse at apex; bases of upper pinnae or lobes not obviously decurrent, without veinlets arising from rachis. Lamina herbaceous; veins free, distinct, no veinlets arising from costae. Sori medial, exindusiate, some covered by scales. Perispore coarsely echinate.

* Rock crevices in forests; 700–800 m. Sichuan.

7. Ctenitis mannii (C. Hope) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 289. 1938.

银毛肋毛蕨  yin mao lei mao jue

Nephrodium mannii C. Hope, J. Bot. 28: 145. 1893[??1890]; Ctenitis fulgens Ching & Chu H. Wang; Dryopteris intermedia (Muhlenberg ex Willdenow) A. Gray var. mannii (C. Hope) H. Christ; D. mannii (C. Hope) Alderwerelt.

Rhizomes erect; rhizome scales brown, narrowly lanceolate, 5–7 mm. Stipe dark stramineous, 15–40 cm; stipe scales spreading, dark brown, subulate, 3–5 mm; scales on rachis and costae alike but smaller. Lamina 2- or 3-pinnatifid, broadly lanceolate, 20–30 Χ ca. 10 cm; pinnae 10–12 pairs; basal pinnae largest, triangular, 5–6 Χ ca. 4 cm; pinnules 5–8 pairs; basal basiscopic pinnules or lobes of basal pinnae longest, ca. 3 Χ 1 cm; bases of upper pinnae or lobes not obviously decurrent, without veinlets arising from rachis. Lamina herbaceous; veins free, distinct, no veinlets arising from costae. Sori medial, indusiate, indusia partly persistent. Perispore coarsely folded or tuberculate.

Forests; 100–200 m. Yunnan (Hekou) [NE India].

In Malesia, Ctenitis mannii is often confused with C. subobscura (H. Christ) Holttum (Holttum 1991: 31 [??citation])

8. Ctenitis pseudorhodolepis Ching & Chu H. Wang, Acta Phytotax. Sin. 19: 121. 1981.

棕鳞肋毛蕨  zong lin lei mao jue

Rhizomes erect; rhizome scales reddish brown, linear, 1.5–1.8 mm. Stipe dark stramineous, 50–60 cm; stipe scales appressed, dark brown, narrowly lanceolate, 5–6 mm; scales on rachis and costae alike but smaller. Lamina 3-pinnate to 4-pinnatifid, triangular, 40–50 Χ 35–45 cm; pinnae 8–10 pairs; basal pinnae largest, triangular, 20–30 Χ 10–15 cm; pinnules 8–10 pairs; basal basiscopic pinnules of basal pinnae longest, 8–10 Χ 3–4 cm; bases of upper pinnae or lobes not obviously decurrent, without veinlets arising from rachis. Lamina papery; veins free, distinct, no veinlets arising from costae. Sori submedial or near midvein, exindusiate, some sori covered by scales. Perispore coarsely echinate.

* Near streams in bamboo forests; 600–800 m. Hunan, Sichuan (Emei Shan).

9. Ctenitis sinii (Ching) Ohwi, Fl. Japan Pterid. 92. 1957.

厚叶肋毛蕨  hou ye lei mao jue, 三相蕨  san xiang jue [??which]

Tectaria sinii Ching, Bull. Dept. Biol. Sun Yatsen Univ. 6: 22. 1933; Ataxipteris sinii (Ching) Holttum; Ctenitopsis sinii (Ching) Ching.

Rhizomes erect, short, 2–2.5 cm in diam., densely scaly at apex and at base of stipes; rhizome scales reddish brown, narrowly lanceolate, 1–2 cm, thinly membranous, entire at margins, fibriform and curly at apex. Stipe dark brown at base, stramineous toward apex, 30–55 cm, 3.5–5 mm in diam. at base, grooved above, densely clothed with scales; scales appressed, reddish brown, lanceolate to ovate-lanceolate, 2–3 cm. Lamina bipinnatifid to tripinnatifid at base, triangular-lanceolate, 30–60 Χ 20–45 cm; pinnae (3–)7 or 8 pairs; basal pinnae largest, subopposite, triangular, 14–25 Χ 9–16 cm; pinnules 1–3 pairs; basal basiscopic pinnules or lobes of basal pinnae longest, 8–15 Χ 2.3–3.5 cm, bases asymmetrically rounded-cuneate, margins crenate-lobed, apices long acuminate; base of upper pinnae or lobes decurrent, with veinlets arising from rachis; texture thickly papery to leathery, dark brown when dried, glabrous on both surfaces; rachises and main veins deeply stramineous and densely clothed with brown, long-articulate hairs and with sparse squamules. Veins free, pinnate, 6–10 pairs per lobe, inconspicuous, simple or 2–4-forked, basiscopic veinlets anastomosing, forming a single row of narrow areoles along costae. Sori round, medial along veinlets, in 2 irregular rows between main veins; exindusiate. Perispore coarsely folded.

Densely forested valleys; 300–500 m. Fujian, Guangdong, Guangxi, Hunan, Jiangxi, Zhejiang [Japan].

10. Ctenitis subglandulosa (Hance) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 302. 1938 [??okay, see IPNI].

亮鳞肋毛蕨  liang lin lei mao jue

Alsophila subglandulosa Hance, Ann. Sci. Nat., Bot., sιr. 5, 5: 253. 1866; Aspidium subtripinnatum Miquel; Ctenitis anyuanensis Ching & Chu H. Wang; C. calcarea Ching & Chu H. Wang; C. chungyiensis Ching & Chu H. Wang; C. costulisora Ching; C. dumrongii Tagawa & K. Iwatsuki; C. maolanensis P. S. Wang; C. membranifolia Ching & Chu H. Wang; C. rhodolepis (C. B. Clarke) Ching; C. subtripinnata (Miquel) H. Itτ; C. yunnanensis Ching & Chu H. Wang; Cyathea subglandulosa (Hance) Copeland; Dryopteris oldhamii (Baker) C. Christensen; D. rhodolepis (C. B. Clarke) C. Christensen; D. subglandulosa (Hance) Hayata (1917), not Kuntze (1891); D. subtripinnata (Miquel) Kuntze; Lastrea intermedia [??C. Presl] var. rhodolepis (C. B. Clarke) Beddome; Nephrodium rhodolepis C. B. Clarke; Polypodium oldhamii Baker.

Rhizomes erect, suberect, or prostrate; rhizome scales reddish brown, linear, 1.5–3 mm. Stipe dark stramineous, 40–50 cm; stipe scales appressed, reddish brown, lanceolate or ovate-lanceolate, 2–3 mm; scales on rachis and costae alike but smaller. Lamina 3-pinnate to 4-pinnatifid, ovate-triangular, 40–60 Χ 30–40 cm; pinnae 12–14 pairs; basal pinnae largest, triangular, 18–22 Χ 12–16 cm; pinnules 10–12 pairs; basal basiscopic pinnules of basal pinnae longest, 8–10 Χ 2.5–3.5 cm; bases of upper pinnae or lobes not obviously decurrent, without veinlets arising from rachis. Lamina herbaceous; veins free, distinct, no veinlets arising from costae. Sori medial or submedial, often near costae, indusiate; indusia usually fugacious, glabrous. Perispore coarsely echinate.

Usually on limestone; sea level up to 1700 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, [??Xizang], Yunnan, Zhejiang [Bhutan, India, Malaysia, Philippines, Vietnam; SE Asia].

Ctenitis subglandulosa is a widespread and morphologically variable species. The indusia of this species are thin and colorless at an early stage, but stop growing midway through its development. Therefore in most herbarium specimens, the indusia are difficult to observe. However, sometimes the indusia are distinct and persistent. The size and abundance of the scales on stipes and rachises, the color and abundance of glands on the lamina beneath, as well as the abundance of hairs on the lamina, are all variable characters. It is observed that when plants grow in open and dry habitats, they develop hairs on the lower leaf surface, which are absent when growing in shade. Spore ornamentation of many samples of this species was examined, and we found most having echinate perispores, except one specimen from Hainan and one from Yunnan having folded perispores. The occurrence of these two different spore types may indicate that this is a species complex that needs more detailed taxonomic revision, preferably using molecular techniques.

Dryopsis Holttum & P. J. Edwards, Kew Bull. 41: 179. 1986.

轴鳞蕨属  zhou lin jue shu

Dong Shiyong (董仕勇)1; Maarten J. M. Christenhusz3

Plants terrestrial, 25–150 cm. Rhizomes short, erect or rarely prostrate, apex covered by lanceolate scales. Fronds tufted; stipe stramineous, brown, or castaneous, scaly throughout; stipe scales brown or blackish brown, narrowly to ovate lanceolate, never clathrate or iridescent (as those in Ctenitis). Lamina lanceolate to ovate, once pinnate with deeply lobed pinnae to amply[??ok] 3-pinnate; basal pinnae oblong, lanceolate, or triangular, usually not or slightly longer than next pair, basal basiscopic lobe or pinnule mostly not enlarged; distal pinnae adnate to rachis and ± decurrent at their bases; veins free; lamina papery, adaxial surface with thick multicellular hairs; rachises and costae grooved and densely covered in ctenitoid hairs or sometimes with hair-scales (structure intermediate between hairs and scales) on adaxial surface, various scales types present or not on abaxial surface. Sori terminal or dorsal on veins, medial to submarginal, one line on either side of costules; indusia present, persistent; spores ellipsoidal to spheroidal, perispore echinate or coarsely tuberculate. x = 41.

Sixteen species in tropical and subtropical Asia, southwest to S India and Sri Lanka, east to Japan and the Philippines, south to Malaysia and Indonesia, most diverse in the southern and southeastern Himalaya; 10 species in China (five endemic).

1a.     Fronds 3-pinnate to 4-pinnatifid, with many free pinnules present; basal stipe scales broadly lanceolate or ovate lanceolate, generally 2–5 mm wide; abaxial surface of costae with few scales except in D. Χ fauriei; perispore coarsely tuberculate.

2a.    Rachis sparsely scaly; indusia poorly developed and difficult to see, but sometimes fully developed and persistent or fugacious .................................................................  9. D. sphaeropteroides

2b.    Rachis densely scaly; indusia fully developed and persistent.

3a.     Sori throughout length of pinnules; abaxial surface of costae sparsely scaly  7. D. maximowicziana

3b.     Sori confined to distal parts of pinnules; abaxial surface of costae copiously scaly  4. D. Χ fauriei

1b.     Fronds 2-pinnate or rarely 3-pinnatifid, if present pinnules usually adnate to costae; scales on stipe base narrowly lanceolate, usually less than 1 mm wide; abaxial surface of costae copiously scaly; perispore sharply echinate.

4a.    Scales on abaxial surface of costae flat.

5a.     Pinnules or pinna lobes with conspicuous hyaline margins .............  2. D. clarkei

5b.     Pinnules or pinna lobes lacking hyaline margins ...............................  8. D. nidus

4b.    Scales on abaxial surface of costae ± bullate.

6a. Scales on abaxial surface of costae slightly bullate; sori confined to distal parts of pinnules  1. D. apiciflora

6b.     Scales on abaxial surface of costae obviously bullate; sori throughout length of pinnules.

7a.     Stipe stramineous or rarely dark brown; pinna lobes entire, crenate, or shallowly lobed, rarely pinnatifid; abaxial surfaces of lobe-costae without or with a few scales ............  6. D. kawakamii

7b.     Stipe castaneous and shiny; pinna lobes usually shallowly pinnate to deeply lobed, rarely entire; abaxial surfaces of lobe-costae scaly.

8a.    Basal stipe scales few, ca. 0.5 cm; sori throughout length of pinnules  5. D. heterolaena

8b.    Basal stipe scales copious, ca. 1 cm; sori usually confined to distal part of pinnules  3. D. Χ dulongensis

 

1. Dryopsis apiciflora (Wallich ex Mettenius) Holttum & P. J. Edwards, Kew Bull. 41: 189. 1986.

顶果轴鳞蕨  ding guo zhou lin jue

Aspidium apiciflorum Wallich ex Mettenius, Abh. Senckenberg. Naturf. Ges. 4: 54. 1858[??2: 338]; Ctenitis apiciflora (Wallich ex Mettenius) Ching; Dryopteris apiciflora (Wallich ex Mettenius) Kuntze; Lastrea apiciflora (Wallich ex Mettenius) C. Presl; Nephrodium apiciflorum (Wallich ex Mettenius) Hooker.

Rhizomes erect. Stipe stramineous to dark brown, 20–40 cm; stipe scales blackish brown, lanceolate, 10–12 Χ 1(–2.5) mm. Lamina 2-pinnate, lanceolate, 40–80 Χ 14–40 cm; free lateral pinnae 18–35 pairs, continuous; basal pinnae slightly shortened, 7–15 Χ 1.5–2.5 cm, not reflexed; middle pinnae 7–18 Χ 2–4 cm; pinnules entire or rarely crenate, obtuse and entire at apex, hyaline margins lacking. Rachises and costae abaxially covered with many scales, scales on costae lanceolate near rachises and ± bullate distally, sometimes scales absent or few on costules. Veins of ultimate pinnules pinnate, with brown hairs and few glandular hairs on abaxial surface. Sori only on distal part of pinnules, contiguous, medial. Indusia distinct and persistent, entire or nearly so. Perispore echinate.

Forests; 1800–3200 m. Taiwan, Xizang, Yunnan [Bhutan, NE India, N Myanmar, Nepal].

2. Dryopsis clarkei (Baker) Holttum & P. J. Edwards, Kew Bull. 41: 181. 1986.

膜边轴鳞蕨  mo bian zhou lin jue

Nephrodium clarkei Baker, Syn. Fil. (Hooker & Baker), ed. 2, 497. 1874; Ctenitis clarkei (Baker) Ching; Dryopteris clarkei (Baker) Kuntze.

Rhizomes erect. Stipe dark brown, 3–18 cm; stipe scales blackish brown, lanceolate, 5–12 Χ 0.5–1 mm. Lamina 2-pinnatifid to 2-pinnate, lanceolate, 25–90 Χ 7–17 cm; free lateral pinnae 20–40 pairs, ± spaced; basal pinnae distinctly shortened, 1–4 Χ 0.5–1.5 cm, reflexed; middle pinnae 3.5–8.5 Χ 1–2 cm; pinnules entire or nearly so, hyaline margins present, obtuse or nearly truncate and entire at apex. Rachises and costae abaxially covered with scales, scales on costae not very abundant, lanceolate, costules without scales. Veins of ultimate pinnules pinnate and veinlets once forked, without glands on abaxial surface. Sori on whole length of pinnules, separate, medial to submarginal. Indusia distinct, persistent or partly fugacious, entire or nearly so. Perispore echinate.

Mossy forests; 1300–3800 m. Guangxi, Guizhou, Sichuan, Xizang, Yunnan [NE India (Sikkim), N Myanmar, Nepal].

Dryopsis clarkei is easily confused with D. nidus, because D. clarkei is the perfect 'nest-plant,' whereas D. nidus is not, despite its name. Dryopsis clarkei is also easily recognized by its hyaline pinna margins.

3. Dryopsis Χ dulongensis (S. K. Wu & X. Cheng) S. Y. Dong, Acta Bot. Yunnan. 23: 187. 2001.

独龙江轴鳞蕨  du long jiang zhou lin jue

Ctenitis dulongensis S. K. Wu & X. Cheng, Acta Phytotax. Sin. 23: 402. 1985.

Rhizomes erect. Stipe reddish brown or castaneous, ca. 50 cm; stipe scales blackish brown, lanceolate, 10–12 Χ 0.8–1 mm. Lamina 2-pinnate to deeply 3-pinnatifid, linear-lanceolate, ca. 98 Χ 34 cm; free lateral pinnae 33–38 pairs, continuous; basal pinna not shortened, ca. 20 Χ 6 cm, not reflexed; middle pinnae ca. 20 Χ 4 cm; pinnules deeply lobed, entire at obtuse apex, hyaline margins lacking. Rachises abaxially covered by many scales, scales on costae broadly lanceolate or ovate-lanceolate and bullate at distal part, many scales on abaxial surfaces of costules. Veins of ultimate pinnules pinnate and veinlets single or forked, with light yellow glands on abaxial surfaces. Sori generally confined to distal part of pinnules, medial. Indusia distinct and persistent, entire or nearly so. Spores not developed.

* Evergreen broad-leaved forests; ca. 1700 m. Yunnan (Gongshan).

Dryopsis Χ dulongensis is only known from the type specimen, which is morphologically intermediate between D. apiciflora and D. heterolaena. Its abortive spores indicate that this taxon may be of hybrid origin, probably between these species.

4. Dryopsis Χ fauriei Holttum & P. J. Edwards, Kew Bull. 41: 198. 1986.

傅氏轴鳞蕨  fu shi zhou lin jue

Rhizomes erect. Stipe castaneous, > 30 cm; stipe scales dark brown, lanceolate, ca. 8 Χ 1.5 mm. Lamina 2-pinnate to 3-pinnatifid, lanceolate, ca. 90 Χ 50 cm; free lateral pinnae ca. 20 pairs, continuous; basal pinnae not shortened, ca. 26 Χ 4.5 cm, not reflexed; middle pinnae ca. 26 Χ 4.5 cm; pinnules entire to deeply lobed, hyaline margins lacking, obtuse at apex. Rachises and costae abaxially scaly, scales on costae ovate, costules without scales. Veins of ultimate pinnules pinnate. Sori confined to distal parts of pinnules. Indusia distinct and persistent. Spores not developed?.

* Forests; ca. 2500 m. Taiwan (Arisan).

Dryopsis Χ fauriei is known from the type specimen only. It is presumably a hybrid between D. apiciflora and D. maximowicziana (Holttum & Edwards 1986 [??page citation]), although the latter 'parent' is not recorded from Taiwan.

5. Dryopsis heterolaena (C. Christensen) Holttum & P. J. Edwards, Kew Bull. 41: 185. 1986.

异鳞轴鳞蕨  yi lin zhou lin jue

Dryopteris heterolaena C. Christensen, Acta Horti Gothob. 1(2): 62. 1924; Ctenitis chingii Z. Y. Liu & J. I. Chang; C. fengiana Ching; C. heterolaena (C. Christensen) Ching; C. kwangsiensis Ching & P. S. Chiu; C. omeiensis Ching & Chu H. Wang; C. tibetica Ching & S. K. Wu; C. zhejiangensis Ching & C. F. Zhang; Dryopsis fengiana (Ching) Holttum & P. J. Edwards; D. kwangsiensis (Ching & P. S. Chiu) Holttum & P. J. Edwards.

Rhizomes erect. Stipe castaneous, 20–60 cm; stipe scales blackish brown, lanceolate, 5–8 Χ ca. 0.5 mm. Lamina 2-pinnate to 3-pinnatifid, lanceolate, 35–85 Χ 20–30 cm; free lateral pinnae 20–30 pairs, ± spaced; basal pinnae not shortened or slightly shorter than next pair, 10–15 Χ 2.5–5 cm, not reflexed; middle pinnae 10–15 Χ 2–4 cm; pinnules usually deeply lobed, or rarely entire, hyaline margins lacking, obtuse and serrate at apex. Rachises and costae abaxially very scaly, scales on costae distinctly bullate, abaxial surfaces of costules also covered in bullate scales. Veins of ultimate pinnules pinnate, with dark brown or castaneous glands on abaxial surfaces. Sori not confined to distal part of pinnules, medial. Indusia distinct and persistent, entire or nearly so. Perispore echinate. 2n = 82*.

* Forests; 800–2400 m. Guangdong, Guangxi, Guizhou, Hunan, Sichuan, Xizang, Yunnan, Zhejiang.

6. Dryopsis kawakamii (Hayata) Holttum & P. J. Edwards, Kew Bull. 41: 186. 1986.

泡鳞轴鳞蕨  pao lin zhou lin jue [??which]

截头轴鳞蕨  jie tou zhou lin jue

Dryopteris kawakamii Hayata, J. Coll. Sci. Imp. Univ. Tokyo 30: 416. 1911; Ctenitis jiulungshanensis P. C. Chiu & G. H. Yao ex Ching, [‘jiulunghsanensis’]; C. kawakamii (Hayata) Ching; C. mariformis (Rosenstock) Ching; C. sacholepis (Hayata) H. Itτ; C. truncata Ching & H. S. Kung[??Ching & Chu H. Wang]; Dryopsis mariformis (Rosenstock) Holttum & P. J. Edwards; D. truncata (Ching & H. S. Kung) Holttum & P. J. Edwards; Dryopteris mariformis Rosenstock; D. sacholepis Hayata.

Rhizomes erect. Stipe dark brown to stramineous, 8–25 cm; stipe scales (blackish) brown, lanceolate, 3–6 Χ ca. 0.5 mm. Lamina 2-pinnatifid to 2-pinnate, (linear) lanceolate, 20–60 Χ 8–15 cm; free lateral pinnae 20–32 pairs, ± spaced, or sometimes continuous; basal pinnae distinctly shortened, 3–5 Χ 1.2–1.8 cm, slightly reflexed; middle pinnae 3.5–10 Χ 0.9–2 cm; pinnules entire to distinctly crenate, obtuse, entire, or sometimes only slightly crenate at apex, hyaline margins lacking. Rachises and costae abaxially covered in many scales, scales of costae distinctly bullate distally, costules without scales. Veins of ultimate pinnules pinnate and veinlets once forked, with pale yellow to castaneous glands on abaxial surfaces. Sori separate, distributed throughout, not confined to distal part of pinnules, medial to submarginal. Indusia distinct and persistent, entire or nearly so. Perispore echinate. 2n = 82*.

* (Bamboo) forests; 400–3100 m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang.

7. Dryopsis maximowicziana (Miquel) Holttum & P. J. Edwards, Kew Bull. 41: 197. 1986.

阔鳞轴鳞蕨  kuo lin zhou lin jue

Aspidium maximowiczianum Miquel, Ann. Mus. Bot. Lugduno-Batavi 3: 178. 1867; Ctenitis aureovestita (Rosenstock) Ching; C. maximowicziana (Miquel) Ching; C. whankanshanensis Ching & Chu H. Wang; Dryopteris aureovestita Rosenstock; D. matsumurae (Makino) C. Christensen; D. maximowicziana (Miquel) C. Christensen; Nephrodium matsumurae Makino.

Rhizomes prostrate or ascending. Stipe stramineous, 18–50 cm; stipe scales brown, lanceolate or broadly lanceolate, ca. 15 Χ 2–4 mm. Lamina 3-pinnate to 4-pinnatifid, ovate, 40–70 Χ 30–50 cm; free lateral pinnae 15–20 pairs, continuous; basal pinnae not shortened, 18–30 Χ 7–11 cm, not reflexed; middle pinnae 18–25 Χ 5–6 cm; pinnules entire to deeply lobed, hyaline margins lacking, obtuse and entire at apex. Rachises and costae abaxially covered in many scales, scales on costae lanceolate; costules without scales. Veins of ultimate pinnules pinnate and veinlets single or forked, with brown hairs on abaxial surfaces. Sori on basal half part of pinnules, or at least not on distal part, medial. Indusia distinct and persistent, eroded and with many minute glands. Perispore coarsely tuberculate. 2n = 82*.

Forests; 100–1500 m. [??Anhui], Chongqing, Fujian, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, [??Taiwan], Zhejiang [Japan, Korea].

8. Dryopsis nidus (Baker) Holttum & P. J. Edwards, Kew Bull. 41: 192. 1986.

巢形轴鳞蕨  chao xing zhou lin jue

Nephrodium filix-mas (Linnaeus) Richard var. nidus Baker, Syn. Fil. (ed. 2): 498. 1874; Ctenitis contigua Ching; C. crassirachis Ching; C. crenata Ching; C. dentisora Ching; C. nidus (Baker) Ching; C. silaensis Ching; C. submariformis Ching & Chu H. Wang; C. transmorrisonensis (Hayata) Tagawa; C. wantsingshanica Ching & K. H. Hsing; C. zayuensis Ching & S. K. Wu; Dryopsis contigua (Ching) Holttum & P. J. Edwards; D. crassirachis (Ching) Holttum & P. J. Edwards; D. crenata (Ching) Holttum & P. J. Edwards; D. dentisora (Ching) Holttum & P. J. Edwards; D. silaensis (Ching) Holttum & P. J. Edwards; D. submariformis (Ching & Chu H. Wang) Holttum & P. J. Edwards; D. transmorrisonensis (Hayata) Holttum & P. J. Edwards; D. wantsingshanica (Ching & K. H. Hsing) Holttum & P. J. Edwards; Polystichum transmorrisonensis Hayata.

Rhizomes erect. Stipe stramineous to reddish brown, 15–26 cm; stipe scales blackish brown, lanceolate, 5–12 Χ 0.5–1 mm. Lamina 2-pinnate, lanceolate, 28–65 Χ 10–19 cm; free lateral pinnae 16–35 pairs, ± spaced; basal pinnae shortened or not, 4–10 Χ 1.5–2 cm, reflexed or not; middle pinnae 5.5–10 Χ 1.2–2.2 cm; pinnules entire to crenate at lateral margins, hyaline margins lacking, obtuse or nearly truncate and crenate to serrate at apex. Rachises and costae abaxially covered in many scales, scales on costae lanceolate or ovate; costules without scales. Veins of pinnules pinnate, hairs, glands, or gland-hairs on veins present or not. Sori in middle, or sometimes confined to distal half of pinnules, separate, medial. Indusia distinct and persistent, entire or erose at margin, glands on indusia present or not. Perispore echinate.

Forests, thickets; 2000–3900 m. Guizhou, Sichuan, Taiwan, Xizang, Yunnan [NE India (Sikkim), E Nepal].

Some characters of Dryopsis nidus vary immensely, e.g., pinnules entire to distinctly crenate or lobed halfway, pinnules closely neighboring to separated 1–2 mm apart, sori medial to marginal on pinnules, indusia entire to erose, glands present on abaxial surface of veinlets and indusia or absent. We have tried to maintain D. silaensis and D. transmorrisonensis at the varietal level, but we could not find any discontinuities in these characters, making a subclassification of this species not applicable. There are however morphological trends but these do not warrant taxonomic recognition.

Despite the name Dryopsis nidus, a more perfect 'nest-plant' is D. clarkei. As a result, the two names are easily confused. Dryopsis clarkei is easily recognized by its hyaline pinna margins.

9. Dryopsis sphaeropteroides (Baker) Holttum & P. J. Edwards, Kew Bull. 41: 199. 1986.

大鳞轴鳞蕨  da lin zhou lin jue

Polypodium sphaeropteroides Baker, Bull. Misc. Inform. Kew 1895: 55. 1895; Aspidium sphaeropteroides (Baker) H. Christ; Athyrium sphaeropteroides (Baker) C. Christensen; Ctenitis sphaeropteroides (Baker) Ching; Dryopteris sphaeropteroides (Baker) C. Christensen; Phegopteris sphaeropteroides (Baker) H. Christ.

Rhizomes erect. Stipe stramineous, 30–45 cm; stipe scales brown, ovate to broadly lanceolate, ca. 10 Χ 3–5 mm. Lamina 3-pinnate to 4-pinnatifid, ovate, 30–45 Χ 20–30 cm; free lateral pinnae 18–21 pairs, contiguous except basal ones; basal pinnae not shortened, 12–16 Χ 6–8 cm, not reflexed; middle pinnae 13–16 Χ 4–6 cm; pinnules entire or sometimes distinctly serrate, obtuse and serrate at apex, hyaline margins lacking. Rachises and costae abaxially covered in hairs but no scales; costules without scales. Veins of ultimate pinnules pinnate, with brown hairs on abaxial surface. Sori not confined to distal part of pinnules, medial to submarginal. Indusia distinct, persistent or partly fugacious, entire or nearly so. Perispore coarsely tuberculate.

* Forests; 2100–2800 m. Sichuan, Yunnan.

LASTREOPSIS Ching, Bull. Fan Mem. Inst. Biol. 8: 157. 1938.

节毛蕨属  jie mao jue shu

Parapolystichum (Keyserling) Ching.

Dong Shiyong (董仕勇)1; Maarten J. M. Christenhusz3

Plants terrestrial, 40–120 cm tall. Rhizomes creeping or ascending, rhizome apices and stipe bases densely covered in lanceolate scales. Fronds approximate or 1–3 cm apart; stipe brown to stramineous, with scales only at base; scales lanceolate, brown or dark brown, without iridescence. Lamina ovate to pentagonal, 3–5-pinnate; basal pinna pairs largest, triangular, basal basiscopic pinnules of basal pinnae longest; distal pinnae or pinnules adnate to rachis or costae and decurrent at their bases; veins all free, veinlets on ultimate pinnules or lobes simple or forked, usually not reaching margin of leaf segments; lamina texture herbaceous, hairy on both surfaces; costae of pinnules prominent, densely covered in hairs. Sori usually terminal on veins and near margins of pinna lobes; indusia present or not; spores ellipsoidal to spheroidal, perispore with inflated folds or tuberculate, sometimes broadly winged. x = 41.

About 35 species: pantropical, extending into south temperate regions, most diverse in Australasia; three species in China (two endemic).

1a.     Sori exindusiate ......................................................................................... 2. L. subrecedens

1b.     Sori indusiate.

2a.    Rhizomes less than 0.5 cm in diam., stipes approximate; lamina 3-pinnate to 4-pinnatifid at base, with abundant glands on abaxial surfaces; hairs on lamina consisting of 3–6 cells ......  3. L. tenera

2b.    Rhizomes up to 1.5 cm in diam., stipes distant (1–3 cm apart); lamina 4-pinnate to 5-pinnatifid at base, without glands on abaxial surfaces; hairs on lamina consisting of 10–25 cells  1. L. microlepioides

 

1. Lastreopsis microlepioides (Ching) W. M. Chu & Z. R. He, Acta Bot. Yunnan. 22: 259. 2000.

云南节毛蕨  yun nan jie mao jue

Trichoneuron microlepioides Ching, Acta Phytotax. Sin. 10: 119. 1965.

Plants 80–120 cm tall. Rhizomes creeping, ca. 1.5 cm in diam., sparsely covered in scales. Stipes distant, 1–3 cm apart on rhizome, dark stramineous, 40–60 cm, sparsely scaly at base; stipe and other parts densely clothed in soft hairs; hairs consisting of 10–25 cells, ca. 3 mm. Lamina 3- or 4-pinnate and 5-pinnatifid at base, narrowly ovate, 40–60 Χ 40–60 cm; pinnae 10–15 pairs, 2 pairs of basal pinnae petiolate and others subsessile to sessile, pinnae subopposite at base and alternate upward, primary pinnules anadromous at basal pinnae and catadromous at upper pinnae; basal pinnae largest, 25–30 Χ 15–20 cm, obliquely deltoid; petiolule 1.5–2 cm, with 14–20 pairs of primary pinnules and basal basiscopic pinnule longest; veins prominent on abaxial surface, obviously not reaching margins of pinna lobes, without glands on abaxial surface. Sori at ends of veins, usually medial between margin and midrib of pinna lobes; indusia large, glabrous but with short glandular hairs at margins.

* Forests; ca. 1600 m. Yunnan.

2. Lastreopsis subrecedens Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 160. 1938.

海南节毛蕨  hai nan jie mao jue

Plants 40–75 cm tall. Rhizomes creeping, 1–1.5 cm in diam., covered in lanceolate scales. Stipe approximate, dark stramineous to brown, 18–38 cm, densely scaly at base; stipe and other parts densely clothed in soft hairs; hairs consisting of 5–8 cells, ca. 0.7 mm. Lamina 3-pinnate and 4-pinnatifid at base, pentagonal, 25–35 Χ 25–40 cm; pinnae 8–10 pairs, only basal pinnae shortly petiolulate, all pinnae usually opposite to subopposite, basal primary pinnules anadromous or subopposite at basal pinnae and catadromous at upper pinnae; basal pinna largest, 15–25 Χ 10–18 cm, obliquely deltoid; stipe 0.7–1.5 cm, with 8–12 pairs of primary pinnules and basal basiscopic pinnules longest; veins prominent on adaxial surface, not reaching margin of pinna lobes, with sphaeroid yellow glands on abaxial surface. Sori terminal on veins and near margin of pinna lobes; exindusiate.

* Forests; 700–1000 m. Hainan.

3. Lastreopsis tenera (R. Brown) Tindale, Vict. Nat. 73: 181. 1957.

台湾节毛蕨  tai wan jie mao jue

Nephrodium tenerum R. Brown, Prod. Fl. Nov. Holland. 149. 1810; Aspidium gardnerianum Mettenius; Ctenitis tenera (R. Brown) Copeland; Dryopteris simozawae Tagawa; D. tenera (R. Brown) C. Christensen; Lastreopsis simozawae (Tagawa) Tagawa.

Plants 25–50 cm tall. Rhizomes creeping, ca. 0.5 cm in diam., covered in lanceolate scales. Stipes approximate, dark stramineous, 10–30 cm, sparsely scaly at base, stipe and other parts densely clothed in soft hairs; hairs consisting of 3–6 cells, ca. 0.5 mm. Lamina 2- or 3-pinnate and at most 4-pinnatifid at base, ovate, 15–20 Χ 20–25 cm; pinnae ca. 10 pairs, usually only basal pinnae shortly petiolate, all pinnae usually opposite to subopposite, primary pinnules anadromous at basal pinnae and catadromous at upper pinnae with rare exceptions; basal pinnae largest, 7–10 Χ 10–18 cm, obliquely deltoid; petiolule 0.3–0.4 cm, with 8–10 pairs of primary pinnules and basal basiscopic pinnule longest; veins prominent on adaxial surface, not reaching or nearly reaching margins of pinna lobes, with abundant sphaeroid or shortly clavate orange glands on abaxial surfaces. Sori terminal or subterminal on veins and usually near margins of pinna lobes; indusia small or large, glabrous but with abundant sphaeroid glands at margin.

Forests. Taiwan [India, Indonesia, Malaysia, Philippines, Sri Lanka; Australia].