PTERIDACEAE [Draft: 03 Aug 2012]

凤尾蕨科  feng wei jue ke

Ding Mingyan (丁明艳), Dong Shiyong (董仕勇)[1], Liao Wenbo (廖文波), Lin Youxing
(林尤兴)[2], Wu Zhaohong (吴兆洪 Wu Shiew-hung), Zhang Gangmin (张钢民)[3], Zhang Xianchun (张宪春)[4];
Edward R. Alverson[5], A. Michele Funston, Michael G. Gilbert[6], Masahiro Kato[7],
Shigeo Masuyama, Jordan S. Metzgar[8], Jefferson Prado[9], Tom A. Ranker[10]

Ferns, terrestrial or rupestral, or aquatic (Ceratopteris), mostly small, exceptionally fronds to 6 m. Rhizome erect to long creeping, with a medullated protostele, a solenostele, or a dictyostele, bearing trichomes or scales or both. Stipe with 1–4 vascular bundles (or to many in Ceratopteris) near base, terete to adaxially sulcate or 2-ridged; lamina usually pinnate, or entire, pedate, palmate, radiate, or stipe furcate apically into 2 strongly recurved rachises; veins usually free, if anastomosing then areolae without free included veinlets. Sori exindusiate, in short to long lines along veins or marginal, or sori borne at vein tips and then usually covered by a modified marginal indusium, with or without special trichomes among sporangia. Sporangia usually long stalked, annulus vertical or rarely oblique, interrupted by stalk. Spores trilete, without chlorophyll.

Ching Ren-chang, Fu Shu-hsia, Wang Chu-hao & Shing Gung-hsia. 1959. Taenitis. In: Ching Ren-chang, ed., Fl. Reipubl. Popularis Sin. 2: 279–280.

Lin Youxing. 1990. Adiantaceae. In: Ching Renchang & Shing Kunghsia, eds., Fl. Reipubl. Popularis Sin. 3(1): 173–216.

Shing Kunghsia. 1990. Acrostichaceae. In: Ching Renchang & Shing Kunghsia, eds., Fl. Reipubl. Popularis Sin. 3(1): 92–94.

Shing Kunghsia. 1990. Coniogramme and Cryptogramma. In: Ching Renchang & Shing Kunghsia, eds., Fl. Reipubl. Popularis Sin. 3(1): 98–103, 228–274.

Shing Kunghsia. 1990. Hemionitidaceae. In: Ching Renchang & Shing Kunghsia, eds., Fl. Reipubl. Popularis Sin. 3(1): 216–278.

Shing Kunghsia & Wu Sukung. 1990. Sinopteridaceae. In: Ching Renchang & Shing Kunghsia, eds., Fl. Reipubl. Popularis Sin. 3(1): 97–173.

Wu Shiewhung. 1990. Pteridaceae. In: Ching Renchang & Shing Kunghsia, eds., Fl. Reipubl. Popularis Sin. 3(1): 10–91.

Zhang Xian-chun. 1999. Antrophyaceae and Vittariaceae. In: Chu Wei-ming, ed., Fl. Reipubl. Popularis Sin. 3(2): 1–31.

1. Subfam. CRYPTOGRAMMOIDEAE

珠蕨亚科  zhu jue ya ke

Zhang Gangmin (张钢民); Tom A. Ranker

Plants terrestrial or on rocks, small (Cryptogramma) or large (Coniogramme) plants. Rhizomes erect or creeping, scaly; scales colorless or brownish. Fronds monomorphic (Coniogramme) or dimorphic (Cryptogramma), distant, closely spaced, or clustered; stipe usually straw-colored, with a vascular bundle near base, adaxially sulcate; lamina 1- or 2-imparipinnate, with an odd terminal segment, or 2–4-pinnate. Veins free or rarely anastomosing near midrib, vein tips enlarged forming hydathodes. Sori along veins, exindusiate; or sori borne at vein tips, covered by a modified marginal indusium. Spores tetrahedral-globose, with prominent angles. x = 30.

Three genera and ca. 40 species: worldwide; two genera and 25 species in China.

1a.       Large plants, to 1 m tall or so; fronds monomorphic; sori borne along veins; margin of fertile segments not modified .....................................................................................................  1. Coniogramme

1b.       Small plants, usually less than 20 cm tall; fronds dimorphic; sori borne at vein tips; fertile segments with a modified marginal indusium ........................................................................  2. Cryptogramma

1. CONIOGRAMME Fιe, Mιm. Foug. 5: 167. 1852, nom. cons.

凤了蕨属  feng liao jue shu

Zhang Gangmin (张钢民); Tom A. Ranker

Plants terrestrial. Rhizomes decumbent or creeping, sparsely scaly; scales brown, lanceolate, margin entire. Fronds monomorphic, distant or closely spaced; stipe straw-colored or with brown spots, or chestnut-brown, glabrous distally, with a single U-shaped vascular bundle; lamina 1- or 2-imparipinnate, rarely trifoliate or 3-pinnate, sometimes entire in juvenile plants, usually herbaceous to papery, glabrous or with hairs on one or both surfaces. Pinnae usually ca. 5 pairs, stalked; in 1-pinnate species terminal pinna same as lateral pinnae; if 2-pinnate then only proximal pinnae imparipinnate or trifoliate (occasionally basal pair only bifurcate), and terminal pinna same as terminal pinnules of proximal pinnae. Pinnules (or distal simple pinnae) lanceolate or oblong-lanceolate, base rounded to cuneate, rarely cordate, margin entire or serrate. Veins free or rarely anastomosing near midrib, then free, vein tips enlarged forming hydathodes. Indusia absent. Sori along veins, intermixed with short, hairy paraphyses. Spores tetrahedral, plain. x = 30.

About 25–30 species: Africa, E and SE Asia, North America; 22 species (11 endemic) in China.

Morphologically, the species in the genus Coniogramme are poorly differentiated. Intermediate individuals exist in some groups, increasing the difficulty of discriminating species. On Emei Shan of Sichuan Province, juvenile plants of some species have simple, lanceolate, or trifoliate to 1-pinnate fronds, and only after several years’ development are the fronds 2- or 3-pinnate. The stipe color is also variable, even in the same plant; some stipes are entirely green, and others have brownish purple spots abaxially. The following characters generally seem to be stable: glabrous or hairy lamina surfaces, hair form (jointed hairs or seta), lamina margin (entire or serrate), and the shape and locality of the hydathodes (far from the tooth at lamina margin, extending into the teeth, or even fusing with the cartilaginous tooth margin). Further field work and cytological study of the genus are still needed.

The fiddlehead of the species of Coniogramme is an edible vegetable, and the rhizome is known to store starch.

1a.       Veins anastomosing to form at least some areoles on each side of midrib.

2a.       Veins anastomosing regularly to form 1 or 2(or 3) continuous rows of areoles on each side of midrib     22. C. japonica

2b.       Veins anastomosing irregularly to form 1 discontinuous row of areoles on each side of midrib, occasionally with only 1 or 2 areoles on each side.

3a.       Veins forming a discontinuous row of areoles on each side of midrib; stipe straw-colored  21. C. wilsonii

3b.       Veins forming only 1 or 2 areoles on each side of midrib; stipe chestnut-colored.

4a.       Pinnules lanceolate, broadest at base, base rounded; hydathodes linear (Yunnan)  19. C. fauriei

4b.       Pinnules elliptic, broadest at middle, base cuneate; hydathodes shortly clavate (Fujian, Guizhou, Hunan, Jiangxi, Zhejiang) ........................................................  20. C. jinggangshanensis

1b.       Veins all free.

5a.       Margins of pinnules entire or at most undulate (C. fraxinea s.l.).

6a.       Hydathodes extending to cartilaginous lamina margin, margins often undulate and revolute when dry    4. C. merrillii

6b.       Hydathodes spindle-shaped, not extending to lamina margin.

7a.       Base of pinnules rounded or slightly cordate ...............................................  1. C. petelotii

7b.       Base of pinnules cuneate or rounded-cuneate.

8a.       Stipe and rachis abaxially straw-colored .................................................  2. C. fraxinea

8b.       Stipe and rachis abaxially reddish purple ...............................................  3. C. rubicaulis

5b.       Margins of pinnules serrate or with irregular short obtuse teeth.

9a.       Hydathodes usually not extending to base of marginal tooth.

10a.     Lamina 1-pinnate; stipe and rachis usually dark brown to purplish black
..................................................................................................................  5. C. robusta

10b.     Lamina 2-pinnate; stipe and rachis straw-colored.

11a.     Pinnules abaxially shortly hairy, apex caudate .........................................  6. C. serrulata

11b.     Pinnules abaxially glabrous, apex acuminate .............................................  7. C. venusta

9b.       Hydathodes extending into marginal tooth or to tooth base.

12a.     Pinnules abaxially papillose, each papilla tipped with a short and stiff hair
...............................................................................................................  8. C. rosthornii

12b.     Pinnules abaxially not papillose, glabrous or pubescent with hairs curved and flat.

13a.     Lamina 3-pinnate in mature plants, lowermost pinnae 2-pinnate.

14a.     Lamina with middle pinnules broadly lanceolate, widest at base, at ca. 90Ί to costae  9. C. procera

14b.     Lamina with middle pinnules narrowly oblong-lanceolate, sides subparallel; pinnules oblique         10. C. affinis

13b.     Lamina 1- or 2-pinnate, lowermost pinnae simple pinnules or pinnate, never 2-pinnate.

15a.     Pinnules lanceolate or oblanceolate; hydathodes extending into teeth and close to or fusing with tooth margin.

16a.     Pinnules lanceolate or oblanceolate, base narrowly cuneate, apex abruptly caudate; hydathodes fusing with cartilaginous tooth margin
.....................................................................................................  17. C. pubescens

16b.     Pinnules lanceolate, base rounded to rounded-cuneate, apex long acuminate or caudate-acuminate; hydathodes extending into teeth or close to tooth margin.

17a.     Teeth of pinnule margin dense, sharp, and fine; hydathodes slightly thicker than veins, extending to tips of teeth and fused with teeth ....................................................  10. C. affinis

17b.     Teeth of pinnule margin coarse, spreading; hydathodes twice as thick as veins, extending into teeth or close to tooth margin .....................................................  18. C. intermedia

15b.     Pinnules often broadly lanceolate to oblong; hydathodes extending to tooth base or slightly into teeth.

18a.     Pinnules abaxially glabrous.

19a.     Pinnules ovate or ovate-oblong; hydathodes extending slightly into teeth  13. C. ovata

19b.     Pinnules broadly lanceolate; hydathodes extending to tooth base.

20a.     Pinnules straight ........................................................................  11. C. emeiensis

20b.     Pinnules arcuate or falcate ........................................................  12. C. falcipinna

18b.     Pinnules abaxially hairy.

21a.     Stipe and rachis reddish purple .......................................................  14. C. sinensis

21b.     Stipe and rachis straw-colored or purplish brown proximally.

22a.     Pinnules adaxially with short and sunken jointed hairs ..............  15. C. suprapilosa

22b.     Pinnules adaxially glabrous .....................................................  16. C. caudiformis

1. Coniogramme petelotii Tardieu, Bull. Mus. Natl. Hist. Nat., sιr. 2, 6: 334. 1933.

心基凤了蕨  xin ji feng liao jue

Coniogramme subcordata Ching, Bull. Fan Mem. Inst. Biol. 2: 213. 1931, not Copeland (1910), nor Maxon (1913).

Plants 30–50 cm tall. Rhizomes long creeping; scales brownish, narrowly lanceolate. Fronds closely spaced; stipe straw-colored, 10–40 cm Χ 2–4 mm; lamina dark green, ovate or narrowly ovate, nearly as long as stipe, 10–25 cm wide, trifoliolate or 1-pinnate, thinly papery or herbaceous when dry, both surfaces glabrous or nearly so. Lateral pinnae 2–5 pairs, oblong or oblong-lanceolate, 16–28 Χ 5–7 cm, with stalks 3–5 mm, base slightly cordate or broadly rounded, margin entire, apex shortly caudate; terminal pinna larger than lateral ones, with long stalk ca. 4 cm. Veins distinct abaxially, lateral veins 2-forked, oblique; hydathodes spindle-shaped, far from lamina margin. Sori extending to 2/3–3/4 of veins.

Rocky cliffs in dense forests; 700–1400 m. Yunnan [Vietnam].

The type specimen of Coniogramme petelotii was collected from N Vietnam.

Fraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes, 135. 2008) included Coniogramme petelotii within the following species, C. fraxinea.

2. Coniogramme fraxinea (D. Don) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 262. 1899.

全缘凤了蕨  quan yuan feng liao jue

Diplazium fraxineum D. Don, Prodr. Fl. Nepal. 12. 1825; Coniogramme fraxinea f. connexa Ching; C. gigantea Ching; C. javanica (Blume) Fιe; Gymnogramma fraxinea (D. Don) Beddome; G. javanica Blume; Neurogramma fraxinea (D. Don) Christ.

Plants up to 2.5 m tall. Rhizomes stout, shortly creeping; scales dark brown, narrowly lanceolate. Fronds scattered; stipe straw-colored, 90–100 Χ 0.5–1 cm; lamina narrowly ovate or ovate-oblong, 80–150 Χ 40–50 cm, pinnate or sometimes 2-pinnate, papery, thin, both surfaces glabrous or sometimes with sparse short hairs abaxially. Pinnae 6–12 pairs; basal pair simple or pinnate, pinnules broadly lanceolate, broadly oblanceolate, or oblong-lanceolate, stalked, base cuneate to rounded, margin entire or occasionally undulate with narrow, colorless, membranous side, apex abruptly caudate. Veins distinct on both surfaces, 2-forked; hydathodes much enlarged, spindle-shaped, not extending to lamina margin. Sori extending to 2/3–4/5 of veins.

Evergreen forests; 800–2000 m. Taiwan, SE Xizang (Mκdog), Yunnan [India, Indonesia, Malaysia, Nepal, Pakistan, Philippines, Vietnam].

3. Coniogramme rubicaulis Ching in K. H. Shing, Acta Bot. Yunnan. 3: 219. 1981.

紫杆凤了蕨  zi gan feng liao jue

Plants up to 1.5 m tall. Stipe reddish purple, 60–70 Χ ca. 1 cm, with sparse dark brown, broadly lanceolate scales proximally, glabrous distally, rounded abaxially, grooved adaxially; lamina brownish green when dry, ovate-deltoid, 80–90 Χ ca. 40 cm, 2-pinnate, herbaceous, thin, both surfaces glabrous; rachis and part of costae reddish purple. Pinnae 8–10 pairs; proximal pinnae ca. 30 Χ 12–15 cm, with stalks 3–3.5 cm. Pinnules 3–5 pairs, ovate-lanceolate, basal pair 8–10 Χ ca. 3.5 cm, sessile, slightly decurrent, base broadly rounded, apex acuminate or caudate-acuminate; terminal pinnule ca. 15 Χ 3.5–5 cm, long stalked, stalk 2–3 cm, base rounded-cuneate, apex caudate-acuminate. Simple pinnae and pinnules entire along margins. Veins distinct, 2-forked; hydathodes much enlarged, not extending to lamina margin. Sori extending to 3/4 of veins.

* Forests in deep valleys; below 800 m. N Guangxi.

Coniogramme rubicaulis resembles C. fraxinea but differs by its reddish purple (not straw-colored) stipe and rachis. More research is needed to clarify their relationship.

4. Coniogramme merrillii Ching, Sinensia 1: 49. 1930.

海南凤了蕨  hai nan feng liao jue

Coniogramme fraxinea (D. Don) Diels var. coriacea Merrill; C. lanceolata Ching.

Plants up to 1 m tall. Rhizomes decumbent, stout; scales broadly lanceolate. Stipe straw-colored or dark straw-colored, 20–50 cm Χ 3–8 mm, abaxially rounded, adaxially grooved; lamina ovate-oblong, 50–70 Χ 30–40 cm, 1-pinnate (sometimes basal pair of pinnae bifurcate), papery or thickly leathery when dry, both surfaces glabrous or sparsely pubescent abaxially. Pinnae 5–8 pairs, broadly lanceolate, broadly oblanceolate, or oblong, 20–35 Χ 3–5 cm, stalked, base cuneate or rounded-cuneate, margin entire, with cartilaginous sides, often undulate and revolute when dry, apex acuminate or caudate-acuminate. Veins distinct, 2-forked; hydathodes much enlarged, extending to cartilaginous lamina margin. Sori extending nearly to vein tips.

Forests in valleys; below 1400 m. Hainan, W Yunnan [NE India, Indonesia, Nepal, Philippines].

Fraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes, 135. 2008) included Coniogramme merrillii and C. lanceolata within C. fraxinea.

5. Coniogramme robusta (Christ) Christ, Bull. Acad. Int. Gιogr. Bot. 19: 175. 1909.

黑轴凤了蕨  hei zhou feng liao jue

Plants 50–80 cm tall. Rhizomes creeping, 3–5 mm in diam.; scales brown, lanceolate. Fronds scattered; stipe purplish black, lustrous, 25–40 cm Χ 2–3 mm, abaxially rounded, adaxially grooved; lamina oblong or broadly ovate, nearly as long as stipe, 15–40 cm wide, 1-pinnate, herbaceous or papery when dry, both surfaces glabrous; rachis and costae purplish black, brown, or straw-colored abaxially, usually lustrous. Pinnae 2–4 pairs, nearly of same shape and size, oblong or oblong-lanceolate, shortly stalked, base rounded or rounded-cuneate, somewhat unequal, margin with short and obtuse teeth, cartilaginous, apex acuminate-caudate to caudate; terminal pinna slightly larger than lateral pinnae. Veins 1- or 2-forked; hydathodes clavate or ovate, not extending to tooth base. Sori extending to bases of hydathodes.

* Forests in valleys, shaded places by roadsides; 600–1500 m. Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan.

1a. Rachis and costae purplish black abaxially ...........................................................  5a. var. robusta

1b. Rachis and costae brown or straw-colored.

2a. Stipe, rachis, and costae straw-colored abaxially .......................................  5b. var. rependula

2b. Stipe and rachis brown; costae straw-colored or sometimes brown proximally  5c. var. splendens

5a. Coniogramme robusta var. robusta

黑轴凤了蕨(原变种)  hei zhou feng liao jue (yuan bian zhong)

Gymnogramma javanica Blume var. robusta Christ, Bull. Acad. Int. Gιogr. Bot. 11: 202. 1902; Coniogramme neorobusta Ching & K. H. Shing; C. pseudorobusta Ching & K. H. Shing.

Stipe, rachis, and costae purplish black abaxially.

* Forests in valleys; 700–1000 m. Guangxi, Guizhou, Hunan.

5b. Coniogramme robusta var. rependula Ching & K. H. Shing, var. nov.

黄轴凤了蕨  huang zhou feng liao jue

Type: China. Jiangxi: Without locality, Xiong Yao-guo 06276 (holotype, PE).

Validating Latin diagnosis: that of “Coniogramme robusta var. rependula Ching et Shing” (Ching & K. H. Shing, Fl. Reipubl. Popularis Sin. 3(1): 279. 1990).

Stipe, rachis, and costae straw-colored abaxially.

* Valleys, shaded places by roadsides; below 800 m. Guizhou (Duyun), Jiangxi.

This name was not validly published by Ching and K. H. Shing in 1990 because the herbarium in which the type is conserved was not specified (Vienna Code, Art. 37.7).

5c. Coniogramme robusta var. splendens Ching & K. H. Shing in S. Y. Jin & Y. L. Chen, Cat. Type Spec. Herb. China (Suppl.), 17. 1999.

棕轴凤了蕨  zong zhou feng liao jue

Stipe and rachis brown; costae straw-colored or sometimes brown (at proximal portion) abaxially.

* Forests; 600–1500 m. Guangdong (Ruyuan), Guizhou, Jiangxi (Wugong Shan), Sichuan (Pingshan).

6. Coniogramme serrulata (Blume) Fιe, Mιm. Foug. 5: 167. 1852.

澜沧凤了蕨  lan cang feng liao jue

Gymnogramma serrulata Blume, Enum. Pl. Javae 2: 113. 1828; Coniogramme fraxinea (D. Don) Diels var. serrulata (Blume) Hieronymus; C. lancangensis Ching & K. H. Shing.

Plants up to 1 m tall. Rhizomes decumbent, stout and fleshy; scales grayish brown, narrowly lanceolate. Fronds scattered; stipe straw-colored with purple spots, 60–75 cm Χ ca. 5 mm, abaxially rounded, adaxially grooved; lamina abaxially grayish green, adaxially green, broadly ovate-lanceolate, 55–70 Χ 22–36 cm, 2-pinnate, herbaceous and wrinkled when dry, abaxially pubescent, adaxially glabrous. Lateral pinnae 5–7 pairs; proximal 1 or 2 pairs of pinnae oblong, 30–35 Χ ca. 20 cm, with stalks ca. 3 cm, pinnate; medial pinnae trifoliolate or bifurcate, distal pinnae simple. Pinnules lanceolate or oblanceolate, ca. 14 Χ 3–3.5 cm, stalked (distal ones sessile), base cuneate, apex long caudate (narrowed tip 2–4 cm), margin obviously with sparse short teeth. Veins distinct abaxially, 1- or 2-forked; hydathodes spindle-shaped, far from base of teeth. Sori extending to 2/3 of veins.

Forests; 900–1300 m. SW Yunnan [NE India, Indonesia, Nepal, Philippines].

“Coniogramme lantsangensis” (in FRPS 3(1): 242. 1990) is an orthographic error for C. lancangensis.

7. Coniogramme venusta Ching, Acta Bot. Yunnan. 3: 222. 1981.

美丽凤了蕨  mei li feng liao jue

Plants 0.7–1.2 m tall. Stipe light straw-colored, 30–50 cm Χ 1.5–3 mm; lamina deltoid-oblong or oblong, 45–65 Χ ca. 25 cm, 2-pinnate, herbaceous when dry, both surfaces glabrous. Lateral pinnae 4–7 pairs, basal pair largest, ovate-deltoid, 20–30 Χ 10–14 cm, with stalk 1.5–3 cm, trifoliolate or pinnate; upper pairs trifoliolate or simple; third pair of pinnae lanceolate, 16–22 Χ 2.5–3 cm, shortly stalked, base rounded or rounded-cuneate, margin with sparse, short, blunt teeth, apex caudate-acuminate or somewhat acute; terminal pinna larger than adjacent pinnae, often divided with a small segment at base. Pinnules of basal pinnae 1–3 pairs, basal basiscopic pinnule longer, lanceolate, somewhat arcuate, 10–15 Χ 1.5–2.5 cm, base rounded-cuneate, margin with sparse short teeth, apex caudate-acuminate; terminal pinnule largest. Veins 2-forked; hydathodes spindle-shaped, not extending to base of teeth. Sori extending to 3/4 of veins.

* Mixed woodlands by streams; 1600–2000 m. Yunnan.

Coniogramme venusta is similar to C. intermedia var. glabra, but its hydathodes are spindle-shaped and do not extend into the teeth.

8. Coniogramme rosthornii Hieronymus, Hedwigia 57: 307. 1916 [“rosthorni”].

乳头凤了蕨  ru tou feng liao jue

Coniogramme rubescens Ching & K. H. Shing; C. taipaishanensis Ching & Y. T. Hsieh.

Plants 60–140 cm tall. Rhizomes long creeping, ca. 5 mm in diam.; scales brown, lanceolate. Fronds widely spaced; stipe straw-colored to reddish purple, 40–85 cm Χ 2.5–5 mm; lamina abaxially greenish, adaxially brownish green, narrowly ovate or ovate-deltoid, as long as stipe or shorter, 18–26 cm wide, 2-pinnate, herbaceous when dry, abaxially densely papillose, with a short and stiff hair on each papilla, adaxially glabrous or sometimes with a few jointed hairs. Lateral pinnae 3–6 pairs, basal pair of pinnae largest, oblong-lanceolate to broadly ovate, stalked, pinnate; lateral pinnules 1–3 pairs, lanceolate to broadly lanceolate, 6–15 Χ 1.5–3.5 cm, base rounded-cuneate or subrounded, margin with short, spreading deltoid teeth, apex acuminate or caudate-acuminate. Medial pinnae as terminal pinnule, lanceolate or narrowly lanceolate, 10–20 Χ 2–4.5 cm, shortly stalked, base rounded-cuneate, margin sharply serrate, apex long acuminate. Veins free; hydathodes slightly thickened, linear, extending to bases of teeth or slightly into teeth. Sori extending nearly to pinna margin.

Forests, rock crevices; 1000–3000 m. Gansu, Guizhou, Henan, Hubei, Shaanxi, Sichuan, Yunnan [Vietnam].

9. Coniogramme procera Fιe, Mem. Soc. Sci. Nat. Strasbourg 6(1): 22. 1865.

直角凤了蕨  zhi jiao feng liao jue

Coniogramme parvipinnula Hayata.

Plants up to 1.8 m tall. Stipe grayish brown proximally, straw-colored distally, 60–90 Χ 0.5–1 cm; lamina abaxially grayish green, adaxially brownish green, narrowly ovate or ovate-oblong, 60–100 Χ 40–60 cm, 2-pinnate (sometimes 3-pinnate), papery when dry, both surfaces glabrous; rachis straw-colored or abaxially purple. Pinnae 10–15 pairs; proximal pairs oblong-lanceolate, 30–50 Χ 15–30 cm, with stalks 2–3 cm, pinnate or sometimes 2-pinnate; pinnules 10–13 pairs, at ca. 90Ί to costae, proximal pinnules lanceolate, 8–12 Χ 2–2.5 cm, base rounded-truncate or truncate (sometimes slightly cordate), apex caudate or acuminate; distal pinnules gradually reduced; terminal pinnule larger than adjacent pinnules, base unequal. Supramedial pinnae smaller, with few pairs of pinnules. Distal pinnae simple, margins serrate, teeth broad, slightly ascending. Hydathodes slender, extending into teeth or only to base of teeth. Sori extending to 1/2–2/3 of veins.

Streamsides in woodlands; 1400–3600 m. Taiwan (Jiayi), S Xizang, NW Yunnan [Bhutan, India, Myanmar, Nepal, Thailand, Vietnam].

10. Coniogramme affinis (C. Presl) Hieronymus, Hedwigia 57: 279. 1916.

尖齿凤了蕨  jian chi feng liao jue

Gymnogramma affinis C. Presl, Tent. Pterid. 218. 1836; Coniogramme affinis var. pilosa H. S. Kung; C. argutiserrata Ching & K. H. Shing.

Plants 60–120 cm tall. Stipe straw-colored, or sometimes brown proximally, 30–70 cm Χ 3–7 mm; lamina brownish green, narrowly ovate or ovate-oblong, 25–85 Χ 15–50 cm, 2-pinnate, or 3-pinnate at base (rarely 1-pinnate), thinly herbaceous when dry, both surfaces glabrous or rarely hairy abaxially. Pinnae 5–8 pairs, basal pair ovate or narrowly ovate, 20–35 Χ 12–20 cm, with stalks 2–3 cm, pinnate (or 2-pinnate with ultimate pinnules 1 or 2 pairs); lateral pinnules 3–6 pairs, lanceolate, 8–15 Χ 1.5–3 cm, shortly stalked or sessile, base cuneate, rounded-cuneate, or subtruncate, somewhat unequal, apex long acuminate or caudate-acuminate; terminal pinnule larger than lateral pinnules, sometimes bifid. Pinnae of second pair pinnate or trifoliolate. Distal pinnae simple and shortened, lanceolate or broadly lanceolate, 10–17 Χ 2–3 cm, margins somewhat irregularly serrate, teeth fine and sharp, cartilaginous. Hydathodes slightly thickened, extending to tips of teeth and fused with teeth. Sori extending to 2/3 of veins.

Forests; 1600–3600 m. Chongqing, Gansu, Heilongjiang, Henan, Liaoning, Shaanxi, Sichuan, Xizang, Yunnan [India, Myanmar, Nepal].

11. Coniogramme emeiensis Ching & K. H. Shing, Acta Bot. Yunnan. 3: 223. 1981.

峨眉凤了蕨  e mei feng liao jue

Coniogramme crenatoserrata Ching & K. H. Shing; C. emeiensis var. lancipinna Ching & K. H. Shing; C. emeiensis var. salicifolia Ching & K. H. Shing; C. longissima Ching & H. S. Kung; C. nanchuanensis Ching & K. H. Shing; C. xingrenensis Ching & K. H. Shing.

Plants up to 1.5 m tall. Rhizomes decumbent, short, stout; scales dark brown, lanceolate. Fronds closely spaced; stipe straw-colored to chestnut brown, 40–90 cm Χ 4–5 mm; lamina broadly ovate-oblong, 30–70 Χ 20–40 cm, 2-pinnate, herbaceous when dry, sometimes with irregular yellow stripes along veins, both surfaces glabrous. Pinnae 6–10 pairs, basal 1–3 pairs of pinnae largest, subovate or ovate-lanceolate, 15–35 Χ 10–20 cm, stalked, pinnate; upper pairs trifoliolate or simple. Pinnules broadly lanceolate, 7–15 Χ 1.5–3 cm, base cuneate or rounded-cuneate, margin with coarse teeth, apex caudate-acuminate or long acuminate. Veins 1- or 2-forked; hydathodes clavate, extending to base of teeth but not into teeth.

* Broad-leaved forests or shaded places by roadsides; 600–1800 m. Chongqing, Guangdong, Guangxi, Guizhou, Hubei, Sichuan, Yunnan, Zhejiang.

The simple pinnae of Coniogramme emeiensis sometimes vary in their length, width, and base shape. This species can be separated from C. intermedia by the clavate hydathodes, which extend only to the tooth base (not into the teeth).

12. Coniogramme falcipinna Ching & K. H. Shing, Acta Bot. Yunnan. 3: 224. 1981.

镰羽凤了蕨  lian yu feng liao jue

Coniogramme falcipinna var. pilocostata P. S. Wang & X. Y. Wang; C. latipinna Ching & K. H. Shing; C. sichuanensis H. S. Kung.

Plants up to 1 m tall. Rhizomes decumbent, ca. 4 mm in diam.; scales brown, lanceolate. Fronds closely spaced; stipe straw-colored with brown spots, 35–60 cm Χ 2–4 mm, glabrous; lamina green or with yellow stripes along veins, ovate-lanceolate or ovate-oblong, 35–50 Χ 18–22 cm, pinnate or proximal 1 or 2 pairs of pinnae trifoliolate or bifurcate, herbaceous when dry, both surfaces glabrous. Pinnae 4–8 pairs; medial pinnae broadly lanceolate, somewhat arcuate, 12–17 Χ 3–4 cm, shortly stalked, base rounded or rounded-cuneate, margin with coarse, ascending teeth, apex caudate-acuminate; distal pinnae similar to medial ones but gradually reduced, sessile; terminal pinna larger than adjacent pinnae, long stalked or bifid at base. Veins distinct, 1- or 2-forked; hydathodes clavate, extending to base of teeth. Sori extending to 2/3–3/4 of veins.

* Forests; 1000–1800 m. Chongqing, Sichuan, Zhejiang.

The broadly lanceolate and somewhat falcate simple pinnae, the rough, ascending teeth along pinna margins, and the hydathodes extending to the base of the teeth distinguish Coniogramme falcipinna from C. intermedia.

13. Coniogramme ovata S. K. Wu, Acta Bot. Yunnan. 3: 230. 1981.

卵羽凤了蕨  luan yu feng liao jue

Plants ca. 80 cm tall. Stipe straw-colored, 28–32 cm Χ 3–4 mm; lamina brownish green, ovate-oblong, ca. 45 Χ 18–20 cm, 2-pinnate, herbaceous when dry, both surfaces glabrous. Lateral pinnae ca. 7 pairs, basal pair larger, ovate-deltoid, ca. 16 Χ 8–10 cm, with stalks ca. 1 cm, trifoliolate; lateral pinnules ovate, ca. 8 Χ 3–3.5 cm, sessile, base rounded-cuneate, apex shortly caudate; terminal pinnule much larger than lateral pinnules, base bifid, with stalk ca. 1 cm. Pinnae of second pair bifurcate. Distal pinnae simple, elliptic or ovate-oblong, 10–12 Χ 3.5–4 cm, shortly stalked or sessile, base rounded-cuneate, apex caudate. Terminal pinna larger than adjacent pinnae, 5–5.5 cm wide, base unequal; pinna and pinnule margins serrate; teeth rough, deltoid, ascending. Hydathodes clavate, somewhat extending into teeth. Sori extending to 1/2–2/3 of veins.

* On rocks in broad-leaved forests; below 1400 m. Yunnan (Yanjin).

14. Coniogramme sinensis Ching, Fl. Tsinling. 2: 210. 1974.

紫柄凤了蕨  zi bin feng liao jue

Plants 60–90 cm tall. Stipe reddish purple, 30–45 cm Χ 3–4 mm, lustrous; scales brownish, lanceolate; lamina oblong-ovate, 30–45 Χ 16–26 cm, 2-pinnate, herbaceous when dry, abaxially sparsely pilose, adaxially glabrous; rachis reddish purple. Pinnae 4 or 5 pairs, basal pair largest, oblong, 15–25 Χ 7–15 cm, stalks 2–3 cm, pinnate; lateral pinnules 2 or 3 pairs, broadly lanceolate or oblong-lanceolate, stalked or sessile, base rounded, apex caudate-acuminate; terminal pinnule larger, with stalk ca. 1 cm. Second pair of pinnae trifoliolate, bifurcate, or simple. Third pair of pinnae lanceolate, 15–20 Χ 2.5–3.8 cm, base rounded, apex caudate-acuminate. Margins of pinnae and pinnules with fine, ascending teeth. Hydathodes slightly thickened, linear, extending to bases of teeth or slightly into teeth. Sori extending to 3/4 of veins.

* Forests or thickets; 400–1600 m. Gansu, Henan, Shaanxi, Sichuan, Zhejiang.

15. Coniogramme suprapilosa Ching, Fl. Tsinling. 2: 209. 1974.

上毛凤了蕨  shang mao feng liao jue

Plants 45–60 cm tall. Stipe straw-colored to light chestnut-colored, 25–30 cm Χ ca. 2.5 mm, lustrous; scales dark brown, lanceolate; lamina abaxially grayish green, adaxially brownish green, broadly ovate, nearly as long as wide, 23–28 cm, 2-pinnate, herbaceous when dry, abaxially pilose, adaxially sparsely pilose along costae and costules, strigose between veins (hairs sunken, leaving small pits on lamina when shed); rachis straw-colored or with occasional brown spots abaxially. Pinnae 2 or 3(or 4) pairs, basal pair larger than adjacent ones, deltoid-ovate, 15–19 Χ 8–14 cm; distal pinnae simple (rarely second pair bifurcate), gradually somewhat reduced; second pair of pinnae oblong or oblong-lanceolate, 15–20 Χ 3–4 cm, base rounded or rounded-cuneate, somewhat unequal, apex abruptly caudate-acuminate. Pinnules 8–15 Χ 2–3 cm, shortly stalked, base rounded, apex caudate-acuminate. Margins of pinnae and pinnules serrate, teeth sharp and deltoid. Veins free; hydathodes slightly thickened, extending to base of teeth. Sori extending to 3 mm from pinna margin.

* Forests or thickets in valleys; 1400–1900 m. Chongqing (Chengkou), Shaanxi, Yunnan (Daguan).

16. Coniogramme caudiformis Ching & K. H. Shing, Acta Bot. Yunnan. 3: 233. 1981.

尾尖凤了蕨  wei jian feng liao jue

Plants 70–100 cm tall. Stipe straw-colored, or purplish brown proximally, 30–55 cm Χ 3–4 mm; lamina dull green, ovate-oblong, 40–50 Χ 20–25 cm, 2-pinnate, herbaceous when dry, abaxially pilose, adaxially glabrous. Pinnae 4–7 pairs, basal pair largest, with stalks 1–1.5 cm, pinnate; lateral pinnules 1 or 2 pairs, lanceolate to oblong-lanceolate, 6–12 Χ 2–3 cm, shortly stalked, base subrounded, apex caudate; terminal pinnule oblong or oblong-lanceolate, as medial pinnae in shape and size, 15–20 Χ 3–4 cm. Pinnae of second pair trifoliolate or bifurcate, other pinnae simple and gradually shortened distally. Terminal pinna larger than adjacent ones. Pinna and pinnule margins densely and sharply serrate. Hydathodes clavate, slightly extending into teeth.

* Forests; 800–1800 m. Sichuan, Zhejiang.

17. Coniogramme pubescens Hieronymus, Hedwigia 57: 314. 1916.

骨齿凤了蕨  gu shi feng liao jue

Coniogramme caudata (Ettingshausen) Ching var. salwinensis Ching & K. H. Shing; C. spinulosa (Christ) Hieronymus; Gymnogramma javanica Blume var. spinulosa Christ, Mιm. Soc. Bot. France 1: 55. 1905.

Plants 60–80 cm tall. Stipe straw-colored, 30–45 cm Χ 3–4 mm; lamina brownish green, ovate-oblong or narrowly ovate, 30–40 Χ 20–30 cm, usually 1-pinnate (rarely basal pair of pinnae bifurcate or trifoliolate), thickly papery when dry, abaxially densely hairy or sometimes glabrous, hairs gray or grayish brown and short, adaxially glabrous. Pinnae 2–4 pairs, basal pair lanceolate or oblanceolate, 15–22 Χ 2–4 cm, stalked (stalks 1–1.5 cm), base narrowly cuneate, margin serrate with fine cartilaginous sides, apex abruptly caudate to caudate-acuminate; distal pairs of pinnae as basal ones but slightly reduced, shortly stalked. Hydathodes extending to and fused with cartilaginous sides of teeth. Sori extending nearly to pinna margins.

Streamsides in mixed forests; 1600–3300 m. SE Xizang, W Yunnan [India, Myanmar, Nepal].

 This species was included in Coniogramme caudata in FRPS, but C. caudata has been reduced to a synonym of C. fraxinea. Coniogramme caudata var. salwinensis was separated from C. caudata sensu FRPS on the basis of its narrowly lanceolate pinnules with gradually caudate-acuminate apices and glabrous abaxial surfaces.

18. Coniogramme intermedia Hieronymus, Hedwigia 57: 301. 1916.

普通凤了蕨  pu tong feng liao jue

Plants 60–120 cm tall. Stipe straw-colored, or with brownish spots, 24–60 cm Χ 2–3 mm; lamina dull green, ovate-deltoid or ovate-oblong, as long as stipe or shorter, 15–25 cm wide, 2-pinnate, herbaceous to papery when dry, both surfaces glabrous or pilose abaxially. Lateral pinnae 3–9 pairs, basal pair largest, deltoid-oblong, 18–28 Χ 8–17 cm, stalks 1–2.5 cm, pinnate; lateral pinnules 1–3 pairs, lanceolate, 6–12 Χ 1.4–2 cm, shortly stalked, base rounded to rounded-cuneate, apex long acuminate or caudate-acuminate; terminal pinnule much larger than lateral pinnules, base very unequal, frequently bifid. Pinnae of second pair trifoliolate or simple (rarely pinnate). Simple pinnae lanceolate, 12–18 Χ 2–3 cm, shortly stalked to sessile, base rounded-cuneate, somewhat unequal, apex long acuminate or caudate-acuminate; terminal pinna often bifid at base; pinna and pinnule margins serrate. Veins 1- or 2-forked; hydathodes linear, extending into teeth or rarely into tooth margin. Sori extending close to lamina margin.

Forests, grasslands, streamsides; 300–2800 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jiangxi, Jilin, Liaoning, Ningxia, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, Korea, Nepal, Pakistan, Russia, Vietnam].

Populations of Coniogramme intermedia represent two taxa treated here as varieties: one is hairy on the abaxial lamina surface, and the other is glabrous on both surfaces. Hieronymus did not designate a type when he published the species. Indian scholars Dixit and Das (Proc. Ind. Acad. Sci. 88(B): II. 263. 1979) chose the duplicate of Meebold 2587, a hairy lamina specimen, as the lectotype of var. intermedia.

1a. Lamina abaxially pilose .................................................................................  18a. var. intermedia

1b. Lamina abaxially glabrous ...................................................................................  18b. var. glabra

18a. Coniogramme intermedia var. intermedia

普通凤了蕨(原变种)  pu tong feng liao jue (yuan bian zhong)

Coniogramme guangdongensis Ching; C. intermedia var. pulchra Ching & K. H. Shing; C. intermedia f. striata H. G. Zhou; C. intermedia f. villosa (Ching) Sa. Kurata; C. intermedia var. villosa Ching; C. latibasis Ching; C. maxima Ching & K. H. Shing; C. simillima Ching [“simileima”].

Lamina abaxially pilose.

Forests, grasslands, streamsides. Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hunan, Jiangxi, Shaanxi, Sichuan, Yunnan, Zhejiang [India, Japan, Korea].

18b. Coniogramme intermedia var. glabra Ching, Icon. Filic. Sin. 3: t. 143. 1935.

无毛凤了蕨  wu mao feng liao jue

Coniogramme guizhouensis Ching & K. H. Shing; C. taibeiensis Ching; C. taiwanensis Ching.

Lamina abaxially glabrous.

Forests; 300–2800 m. Fujian, Gansu, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jilin, Liaoning, Ningxia, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, Korea, Nepal, Pakistan, Russia, Vietnam].

19. Coniogramme fauriei Hieronymus, Hedwigia 57: 320. 1916.

单网凤了蕨  dan wang feng liao jue

Coniogramme simplicior Ching; C. simplicior f. concinna H. G. Zhou.

Rhizomes decumbent, stout; scales dark brown, lanceolate. Stipe chestnut-colored, ca. 35 cm Χ 3 mm; lamina brownish green, ovate-deltoid or ovate-oblong, as long as stipe, 18–20 cm wide, pinnate, herbaceous when dry, abaxially brown pilose, adaxially glabrous. Lateral pinnae ca. 5 pairs, basal pair largest, lanceolate, 14–18 Χ ca. 2.5 cm, stalks ca. 1 cm, base unequal, basiscopically divided, small segment 6–9 cm, apex caudate-acuminate; other pinnae simple, somewhat shortened distally, shortly stalked or sessile; pinnae of second pair broadly lanceolate, 13–16 Χ 2.5–3 cm, base broadly rounded, margin serrate, teeth sharp, ascending, apex caudate-acuminate; terminal pinna larger than adjacent ones, base bifid. Veins 1- or 2-forked, occasionally connected forming 1 or 2 areoles; hydathodes linear, only extending to tooth base. Sori extending to 2/3 of veins.

Forests; below 2300 m. W Yunnan [Korea].

Coniogramme fauriei is very similar to C. wilsonii but differs by its veins occasionally connected and forming 1 or 2 areoles.

20. Coniogramme jinggangshanensis Ching & K. H. Shing, Acta Bot. Yunnan. 3: 238. 1981.

井岗山凤了蕨  jing gang shan feng liao jue

Coniogramme tsingkangshanensis Ching.

Stipe brownish purple, ca. 70 cm Χ 5 mm; lamina dull green when dry, ovate-oblong, somewhat longer than stipe, ca. 30 cm wide, 2-pinnate, herbaceous, both surfaces glabrous; rachis brownish purple abaxially, brown adaxially. Lateral pinnae ca. 8 pairs, basal pair largest, ovate-deltoid, ca. 35 Χ 15 cm, stalks ca. 3 cm, pinnate; lateral pinnules 3 pairs, well separated, lanceolate, ca. 15 Χ 2 cm, base broadly cuneate, apex long acuminate; terminal pinnule much longer than lateral ones, as medial pinnae in shape and size. Pinnae of second pair trifoliolate. Medial pinnae simple, linear-lanceolate, ca. 25 Χ 3 cm, stalks ca. 1 cm, base broadly cuneate, apex long acuminate; distal pinnae gradually shortened. Terminal pinna larger than adjacent ones. Pinna and pinnule margins serrate, teeth shallow, incised. Veins 1- or 2-forked, occasionally connected forming 1 or 2 areoles; hydathodes extending to tooth base. Sori extending close to lamina margin.

* Evergreen forests by streams or in ravines; 500–1300 m. Fujian, Guizhou, Hunan, Jiangxi, Zhejiang.

Coniogramme jinggangshanensis looks much like C. intermedia and C. japonica. It differs from the former by its larger stature (to 1.5 m tall), the stout teeth of the pinna margin, and the hydathodes reaching only to the tooth bases; it differs from the latter by the pinnae somewhat linear-lanceolate and the veins not forming 2 or 3 rows of areoles along each side of the midrib.

21. Coniogramme wilsonii Hieronymus, Hedwigia 57: 321. 1916.

疏网凤了蕨  shu wang feng liao jue

Stipe straw-colored or dull straw-colored, ca. 40 cm Χ 3–5 mm; lamina abaxially grayish green, adaxially brownish green, ovate-deltoid or ovate-oblong, 28–50 Χ 20–25 cm, 2-pinnate, herbaceous when dry, both surfaces glabrous. Lateral pinnae 3–5 pairs, basal pair largest, deltoid-ovate, 18–25 Χ 10–15 cm, stalks ca. 1.5 cm, pinnate; lateral pinnules 1–3 pairs, lanceolate, 8–12 Χ 2–2.5 cm, shortly stalked or adnate to costa, base unequal, somewhat cordate or rounded-cuneate, apex caudate-acuminate; terminal pinnule much larger, as medial pinnae in shape and size. Medial pinnae simple, 15–20 Χ 2–3 cm. Pinna and pinnule margins serrate, teeth sparse and shallow. Veins free except forming a few discontinuous areoles along each side of costae; hydathodes linear, not extending to tooth base. Sori extending close to lamina margin.

* Forests; 1000–1600 m. Gansu, Guizhou, Henan, Hubei, Hunan, Shaanxi, Sichuan.

Coniogramme wilsonii resembles C. japonica, but it differs in its pinnules with unequal and somewhat cordate bases and its areoles fewer and discontinuous along the midribs.

22. Coniogramme japonica (Thunberg) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 262. 1899.

凤了蕨  feng liao jue

Hemionitis japonica Thunberg, Fl. Jap. 333. 1784; Coniogramme ankangensis Ching & Y. P. Hsu; C. centrochinensis Ching; C. gracilis Ogata; C. japonica subsp. gracilis (Ogata) Nakaike; C. japonica var. gracilis (Ogata) Tagawa; Dictyogramme japonica (Thunberg) Fιe; Gymnogramma japonica (Thunberg) Diels; Notogramme japonica (Thunberg) C. Presl.

Stipe straw-colored or chestnut brown, 30–50 cm Χ 3–5 mm; lamina oblong-deltoid or ovate-deltoid, 20–40 cm wide, 2-pinnate, papery or herbaceous when dry, both surfaces glabrous. Lateral pinnae usually 3–5 pairs, basal pair largest, ovate-deltoid, 20–35 Χ 10–15 cm, stalks 1–2 cm, pinnate (occasionally bifurcate); lateral pinnules 1–3 pairs, lanceolate or broadly lanceolate, 8–15 Χ 1.5–3.5 cm, stalked or distal pinnules sessile, base cuneate, rounded, or rarely cordate, apex acuminate, long acuminate, or shortly caudate; terminal pinnule broadly lanceolate, 20–28 Χ 2.5–4 cm, usually attenuate toward base. Pinnae of second pair trifoliolate, bifurcate, or simple; simple pinnae as terminal pinnule. Pinna or pinnule margins serrate, teeth sparse and short. Veins anastomosing to form 1–3 rows of areoles along each side of midrib, then free to margin; hydathodes spindle-shaped, extending below base of teeth. Sori extending nearly to lamina margin.

Forests, shaded wet soil in ravines; 100–2000 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, S Jiangsu, Jiangxi, Sichuan, Shaanxi, Taiwan, Yunnan, Zhejiang [Japan (including Ryukyu Islands), Korea].

2. CRYPTOGRAMMA R. Brown in Franklin, Narr. Journey Polar Sea, 767. 1823.

珠蕨属  zhu jue shu

Zhang Gangmin (张钢民); Edward R. Alverson, Jordan S. Metzgar

Plants small, usually on rocks. Rhizomes short, erect or ascending with dictyostele, or occasionally long creeping with siphonostele; scales colorless or brownish, thin. Fronds strongly dimorphic, clustered or rarely scattered, sterile fronds shorter than fertile ones; stipe dark brown proximally, light brown to straw-colored distally, grooved adaxially, scaly. Sterile fronds: lamina broadly ovate or oblong, 2–4-pinnate, membranous to leathery, glabrous. Fertile fronds: lamina 2- or 3-pinnate. Ultimate segments of sterile lamina ovate, spatulate, elliptic, or fan-shaped; fertile segments linear or elongate-oblong. Veins free, pinnately branched, single or forked distally; hydathodes often sunken below surface on dried specimens. False indusia broad, clearly marginal, continuous, making a single fertile segment podlike. Sori borne at vein tips, orbicular or elliptic, confluent when mature. Spores yellow, tetrahedral, trilete, verrucose. x = 30.

About ten species: temperate and subtropical alpine regions, Asia, Europe, North and South America; two species in China.

Plants of Cryptogramma resemble those of Onychium but are smaller, strongly dimorphic, and have fertile fronds markedly taller than sterile fronds. There is not a commisural vein connecting veinlet tips, so the sori are borne at veinlet tips and become confluent only when mature.

1a.       Rhizomes slender and creeping; fronds sparse and scattered; sterile lamina 1- or 2-pinnate, herbaceous to membranous, thin; hydathodes absent .................................................................  1. C. stelleri

1b.       Rhizomes stout and erect; fronds densely tufted; sterile lamina 3- or 4-pinnate, papery or somewhat leathery, obscure; hydathodes present and conspicuous.

2a.       Sterile lamina 3-pinnate-pinnatifid, ultimate segments triangular, with an acute apex  2. C. brunoniana

2b.       Sterile lamina 4-pinnate-pinnatifid, ultimate segments linear, with a rounded apex  3. C. raddeana

1. Cryptogramma stelleri (S. G. Gmelin) Prantl in Engler, Bot. Jahrb. Syst. 3: 413. 1882.

稀叶珠蕨  xi ye zhu jue

Pteris stelleri S. G. Gmelin, Novi Comment. Acad. Sci. Imp. Petrop. 12: 519. 1768; Allosorus gracilis (Michaux) C. Presl; A. stelleri (S. G. Gmelin) Ruprecht; Cheilanthes gracilis (Michaux) Kaulfuss; Pellaea gracilis (Michaux) Hooker; P. stelleri (S. G. Gmel.) Baker; Pteris gracilis Michaux.

Rhizomes slender, long creeping; scales light brown, lanceolate or ovate-lanceolate. Fronds scattered along rhizomes; stipe brownish straw-colored, glabrous, stipe of sterile frond 3–8 cm, that of fertile frond 6–12 cm. Sterile fronds: lamina yellowish green, ovate or ovate-oblong, 3–6 Χ 1.5–3 cm, 1- or 2-pinnate, herbaceous to membranous, glabrous on both surfaces, apex obtuse; hydathodes poorly developed or absent. Fertile fronds: lamina broadly lanceolate or oblong, 4–7 Χ 1.8–4 cm, 2-pinnate. Sterile pinnae 3 or 4 pairs, subrounded, margins entire or slightly undulate, apex obtuse. Fertile pinnae 4 or 5 pairs, shortly stalked or subsessile, basal pair largest, 1-pinnate. Pinnules 1 or 2 pairs, anadromous, broadly lanceolate, shortly stalked or sessile, base cuneate, apex acute or obtuse. Sori borne at vein tips, discrete, often confluent at maturity. False indusia pale green, membranous, margins somewhat irregular, not reaching to main vein.

Rock crevices in Abies or Rhododendron woodlands; 1700–4200 m. Gansu, Hebei, Qinghai, Shaanxi, Shanxi, Sichuan, Taiwan, Xinjiang, SE Xizang, NW Yunnan [Afghanistan, Bhutan, India, Japan, Kashmir, Nepal, Russia; North America].

2. Cryptogramma brunoniana Wallich ex Hooker & Greville, Icon. Filic. 2: t. 158. 1829.

高山珠蕨  gao shan zhu jue

Cryptogramma crispa var. brunoniana (Wallich ex Hooker & Greville) Baker; C. crispa f. indica Hooker; C. emeiensis Ching & K. H. Shing; C. shensiensis Ching; Phorolobus brunonianus (Wallich ex Hooker & Greville) Fιe.

Rhizomes stout and erect; scales brown, lanceolate, membranous. Fronds tufted; stipe straw-colored, sparsely scaly proximally, stipe of sterile frond 4–5 cm, that of fertile frond 7–16 cm. Sterile fronds: lamina broadly ovate or ovate-deltate, 3–4 Χ 1.5–4 cm, 3- or 4-pinnate, herbaceous; hydathodes spindle-shaped, slightly sunken below frond surface on dried specimens. Fertile fronds: lamina ovate or ovate-oblong, 4–8 Χ 1.5–3.5 cm, finely 3-pinnate to 3-pinnate-pinnatifid. Sterile pinnae 6 or 7 pairs, basal pair largest, ovate, 1.6–2.2 Χ 1.2–1.7 cm. Ultimate sterile segments deltate or oblong, apex acute, one veinlet in each segment. Ultimate fertile segments linear at first, elliptic when mature, 3–5 Χ 1–2 mm, apex obtuse, or sometimes acute. Sori borne near vein tips, somewhat rounded, confluent and spreading over abaxial surface of fertile lamina at maturity. False indusia brown, linear, margins entire.

Rock crevices; 3300–3700 m. Sichuan, E and C Taiwan, NW Yunnan [Bhutan, N India, Japan, Nepal].

Cryptogramma brunoniana has been treated as a variety of the C. crispa (Linnaeus) R. Brown ex Hooker from Europe and Asia, but an analysis of chloroplast and nuclear DNA variation shows that the two species are only distantly related (Metzgar, unpubl. data).

3. Cryptogramma raddeana Fomin, Izv. Kievsk. Bot. Sada 10: 3. 1929.

珠蕨  zhu jue

Allosorus raddeanus (Fomin) Ching; Cryptogramma crispa (Linnaeus) R. Brown ex Hooker var. sinensis Christ; C. brunoniana Wallich ex Hooker & Greville var. raddeana (Fomin) Fraser-Jenkins; C. brunoniana var. sinensis (Christ) G. M. Zhang.

Rhizomes stout and erect; scales brown, lanceolate, membranous. Fronds tufted; stipe straw-colored, sparsely scaly proximally, stipe of sterile frond shorter, 4–5 cm, that of fertile frond 7–16 cm. Sterile fronds: lamina brownish green when dry, broadly ovate, 3–5 Χ 1.5–2 cm, 4-pinnate-pinnatifid, papery, glabrous on both surfaces; hydathodes obovate and not sunken below frond surface on dried specimens. Fertile fronds: lamina ovate or ovate-oblong, 4–6 Χ 1.5–3.5 cm, finely 3-pinnate to 3-pinnate-pinnatifid. Sterile pinnae 6 or 7 pairs, basal pair largest, ovate, 1.6–2.2 Χ 1.2–1.7 cm. Ultimate sterile segments linear or spatulate, apex rounded, one veinlet in each segment. Ultimate fertile segments linear at first, elliptic when mature, 3–5 Χ 1–2 mm, apex obtuse. Sori confluent and spreading over abaxial surface of fertile frond when mature. False indusia brown, linear, margins entire.

On rocks; 2600–4600 m. W Hubei, Shaanxi, NW and W Sichuan, Xizang, NW Yunnan [NW Nepal, Russia].

Cryptogramma raddeana has been treated as a variety or subspecies of C. brunoniana (Zhang & Zhang, Acta Phytotax. Sin. 41: 475–482. 2003), but plastid DNA data resolve each of the two taxa as well-supported clades that are reciprocally monophyletic to one another and have a level of genetic differentiation similar to other interspecific divergences in the genus (Metzgar, unpubl. data).  They also differ in slight but consistent morphological differences (e.g., lamina dissection, pinnule shape). Combined, this suggests that they are distinct though closely related species. Perceived intergradation between related taxa in Cryptogramma can be due either to formation of sterile hybrids (Alverson, Biosyst. Parsley-Ferns, MS thesis, Oregon State University. 1989) or to the presence of fronds that are developmentally intermediate between sterile and fertile fronds, which obscure the differences between taxa in frond dissection and segment shape (Hultιn, Fl. Aleutian Islands. 397pp. 1937).

2. Subfam. CERATOPTERIDOIDEAE

水蕨亚科  shui jue ya ke

Two genera and ca. ten species: tropics and subtropics; two genera and four species in China.

1a.       Lamina 1-pinnate with distinct terminal pinna; sporangia scattered on whole of abaxial lamina surface       3. Acrostichum

1b.       Lamina finely pinnate or pinnatifid; sori borne at vein tips, covered by a modified marginal indusium          4. Ceratopteris

3. ACROSTICHUM Linnaeus, Sp. Pl. 2: 1067. 1753.

卤蕨属  lu jue shu

Dong Shiyong (董仕勇); A. Michele Funston

Plants terrestrial or in coastal swamps. Rhizome stout, creeping or erect, dictyostelic with additional strands; scales dark brown to black, large, broadly lanceolate, entire. Fronds clustered or approximate, stipe stout, glabrous; lamina 1-pinnate with distinct terminal pinna; pinnae stalked, tongue-shaped to narrowly oblong, thickly papery to thickly leathery or fleshy, entire, apex obtuse to acuminate; veins anastomosing, without included free veinlets. Fertile pinnae distal on frond or most or all of lamina fertile, slightly smaller. Sporangia scattered on whole of abaxial surface; paraphyses capitate, apex lobed and multicellular. Spores tetrahedral-globose, perispore papillate to tuberculate, with rodlets or sparse strands. x = 30.

Three species: pantropical; two species in China.

1a.       Sterile pinnae rounded to retuse and shortly mucronate at apex ...........................  1. A. aureum

1b.       Sterile pinnae narrowly acuminate at apex ......................................................  2. A. speciosum

1. Acrostichum aureum Linnaeus, Sp. Pl. 2: 1069. 1753.

卤蕨  lu jue

Acrostichum inaequale Willdenow; Chrysodium aureum (Linnaeus) Mettenius; C. inaequale (Willdenow) Fιe; C. vulgare Fιe.

Plants 1–2 m tall. Rhizome erect; scales dark brown to black, broadly lanceolate, 1–2[–4] Χ 0.2–0.6[–2] cm. Stipe straw-colored to pale castaneous, 10–70[–100] cm, 0.5–1 cm in diam., glabrous upward; lamina 40–50[–300] Χ 20–40 cm, lateral pinnae to 5–12 pairs; sterile pinnae narrowly oblong, (10–)15–35[–80] Χ 1.8–5.5[–8] cm, with stalk 0.5–1.5 cm, thickly leathery, both surfaces glabrous, base cuneate to rounded, margin entire and cartilaginous, apex rounded to retuse and shortly mucronate; costae strongly raised abaxially, flat or slightly grooved adaxially; veins raised abaxially, hardly visible adaxially, closely spaced, copiously anastomosing to form irregular areoles. Fertile pinnae distal on frond, like sterile pinnae but slightly smaller. n = 30, 120.

Coastal areas, often in mangrove swamps; near sea level to 100 m. Guangdong, Guangxi, Hainan, Taiwan (Hualian, Pingdong), Yunnan [pantropical].

2. Acrostichum speciosum Willdenow, Sp. Pl. 5: 117. 1810.

尖叶卤蕨  jian ye lu jue

Plants 1–1.5 m tall. Rhizome scales dark brown to black, broadly lanceolate, ca. 1 Χ 0.2–0.3 cm. Stipe straw-colored, 30–40 cm, 0.3–0.5 cm in diam., glabrous; lamina 35–50 Χ 20–30 cm; lateral pinnae to 5–8 pairs; sterile pinnae lanceolate, (8–)15–20 Χ 2–2.5 cm, with stalk 0.5–1 cm, thickly leathery, both surfaces glabrous, base narrowly cuneate, margin entire and cartilaginous, apex narrowly acuminate; costae strongly raised abaxially, flat or slightly grooved adaxially; veins raised abaxially, hardly visible adaxially, closely spaced, anastomosing. Fertile pinnae distal on frond, like sterile pinnae but somewhat abruptly attenuate and shortly caudate.

Mangrove swamps; near sea level. Hainan (Wenchang) [Malaysia, Thailand, Vietnam; tropical Asia, Australia].

4. CERATOPTERIS Brongniart, Bull. Sci. Soc. Philom. Paris, sιr. 3, 8: 186. 1821[1822 per ING].

水蕨属  shui jue shu

Lin Youxing (林尤兴); Shigeo Masuyama

Parkeria Hooker.

Ferns annual, juicy, aquatic. Rhizome erect, short, with thick roots, dictyostele, with scales at apex; scales broadly ovate or cordate to peltate, entire, thin, brownish hyaline. Fronds clustered, dimorphic; stipe green, ± expanded, semicylindrical, fleshy, smooth, with sparse scales, with many longitudinal ridges on surface and many small vascular bundles inside. Sterile lamina green, ovate- to lanceolate-triangular, thinly herbaceous, simple or pinnate; ultimate lobe broadly lanceolate or loriform, entire, acute at apex; veins anastomosing; gemma occasionally formed in axil of pinnae, brownish, small, ovate, resulting in juveniles through asexual propagation. Fertile lamina similar in morphology to sterile lamina but normally taller, divided more deeply and finely; ultimate lobe reflexed toward costa to enclose sori, green when young and brownish when old, linear to siliquiform; rachis green, with longitudinal ridges, deplanate when dry. Sori attached along costa, narrowly linear, covered with reflexed margin of lobe. Sporangium large, subsessile; annulus broad, vertical, consisting of 0–70 incrassate cells; trilete mark obvious or not. Spores 16 or 32 per sporangium, large, tetrahedral, trilete, with fine, parallel ridgelike ornamentations. x = 13(39).

Four to seven species: tropics and subtropics; two species in China.

In China, the rhizomes and fronds are used medicinally for treating fetal toxins and phlegm buildup. The young fronds are used as a vegetable.

1a.       Plants rooting in silt; sterile frond varied in form, pinnate to 3-pinnate, tall or short depending on environment; stipe 3–30 cm, ca. 1 cm in diam., base unexpanded; fertile fronds taller than sterile ones, lamina oblong or ovate-triangular ...............................................................  1. C. thalictroides

1b.       Plants usually floating; sterile frond simple to pinnatifid to pinnate, broadly triangular; stipe 5–8 cm, 1–3 cm in diam., base much expanded; fertile fronds not taller than sterile ones, lamina broadly triangular             2. C. pteridoides

1. Ceratopteris thalictroides (Linnaeus) Brongniart, Bull. Sci. Soc. Philom. Paris, sιr. 3, 8: 186. 1821.

水蕨  shui jue

Acrostichum thalictroides Linnaeus, Sp. Pl. 2: 1070. 1753; A. siliquosum Linnaeus; Ceratopteris siliquosa (Linnaeus) Copeland; Ellobocarpus oleraceus Kaulfuss; Furcaria thalictroides (Linnaeus) Desvaux; Pteris siliquosa (Linnaeus) P. Beauvois; P. thalictroides (Linnaeus) Swartz (1800), not Muhlenberg (1793); Teleozoma thalictroides (Linnaeus) R. Brown ex H. Richards.

Plants green, 5–70 cm tall, juicy and soft. Rhizome erect, short. Fronds clustered and dimorphic. Sterile fronds: stipe green, semicylindrical, 3–30 cm, ca. 1 cm in diam. or less, fleshy, not expanded, sparsely scaly; lamina erect or floating when young, ovate to lanceolate, 6–30 Χ 3–15 cm, base rounded-cuneate, apex acuminate, 2–4-pinnate; pinnae 5–8 pairs, alternate; lower 1 or 2 pairs larger, ovate to oblong, up to 10(–35) Χ 7 cm, base subrounded to subtruncate, apex acute to acuminate, 1–3-pinnate; pinnules 2–5 pairs, alternate, stalk short and with narrow wings on both sides, blade broadly ovate or ovate-triangular, up to 4 Χ 3 cm, deeply divided, base rounded-truncate, apex obtuse to acuminate; ultimate lobe linear-oblong or linear-lanceolate, up to 2 Χ 0.5 cm, entire, base decurrent along rachis forming broad wing, apex obtuse to acute; upper pair of pinnae similar in shape to basal pair of pinnae but gradually smaller. Fertile fronds: stipe same as in sterile fronds; lamina oblong or ovate-triangular, 15–40 Χ 10–22 cm, base rounded-cuneate or rounded-truncate, 2- or 3-pinnate, apex acuminate; pinna 3–8 pairs, alternate, lower 1 or 2 pairs of pinnae larger, ovate or narrowly triangular, up to 14 Χ 6 cm, stalked; ultimate lobe linear to siliquiform, 1–4 Χ ca. 0.2 cm, margin thin, strongly reflexed toward costa, like false indusium, apex acuminate. Veins anastomosing. Lamina softly herbaceous, green when young and brownish when old, glabrous; rachis and costa same color as stipe, smooth. Sporangia attached to veinlets on both sides of main vein, covered with reflexed margin of lobe, brown, with 30–70 annulus cells, with 32 spores inside. Spores tetrahedral, more than 100 μm in diam., with granulate perine and thick exine forming rich parallel ridges on surface. 2n = 154, 156 (tetraploid).

Ponds, ditches, rice fields, taro patches, usually rooting. Anhui, Fujian, Guangdong, Guangxi, Hainan, Hubei, Jiangsu, Jiangxi, Shandong, Sichuan, Taiwan, Yunnan, Zhejiang [India, Indonesia, Japan, Malaysia, Myanmar, Nepal, New Guinea, Philippines, Sri Lanka, Thailand, Vietnam; Africa, Australia, Central, North, and South America, Madagascar, Pacific islands, West Indies].

2. Ceratopteris pteridoides (Hooker) Hieronymus, Bot. Jahrb. Syst. 34: 561. 1905 [“pteroides”].

粗梗水蕨  cu geng shui jue

Parkeria pteridoides Hooker, Exot. Fl. 2: t. 147. 1825; Ceratopteris parkeria J. Smith.

Plants usually floating, 20–30 cm tall. Stipe, rachis, and costa of lower pinnae all obviously expanded toward base, base of stipe narrowly cuneate, covered with roots. Fronds dimorphic. Sterile fronds green, smooth; stipe semicylindrical, 5–8 cm, ca. 1.5 cm in diam.; lamina ovate-triangular, simple and deeply divided, sometimes opposite-pinnate; lobes triangular to broadly loriform. Fertile fronds green when young and brownish when old, smooth; stipe 5–8 cm, 1–3 cm in diam.; lamina broadly triangular, 15–25 cm, 2–4-pinnate; ultimate lobe linear or siliquiform, 2–6 Χ ca. 0.2 cm, margin thin, strongly reflexed toward main vein to cover sori, apex acuminate. Sporangia attached to veinlets on both sides of main vein, covered with reflexed margin of lobe, brown, with 0–40 annulus cells, with 32 spores inside. Spore tetrahedral, below 100 μm in diam., with few parallel ridges. 2n = 78 (diploid).

Marshes, ponds, ditches, usually floating on water. Anhui, Hubei, Jiangsu, Jiangxi, Shandong [Bangladesh, India, Vietnam; Central, North, and South America].

3. Subfam. PTERIDOIDEAE

凤尾蕨亚科  feng wei jue ya ke

Zhang Gangmin (张钢民), Liao Wenbo (廖文波), Ding Mingyan (丁明艳), Dong Shiyong (董仕勇), Wu Zhaohong (吴兆洪 Wu Shiew-hung); Jefferson Prado, Tom Ranker

Plants terrestrial, small, medium, or large. Rhizome erect or ascending, rarely long creeping, with solenostele or dictyostele, scaly, or rarely covered with bristles (Taenitis). Fronds monomorphic, rarely dimorphic or subdimorphic; stipe often straw-colored, seldom castaneous-reddish or brown, glabrous, rarely hispid or scaly; lamina 1-pinnate or 2–5-pinnate-pinnatifid, or rarely digitate or pedate, occasionally simple or 3-forked, oblong or ovate-triangular, rarely pentagonal in outline, herbaceous, papery or leathery, mostly glabrous, rarely with white or yellow farina abaxially; venation free or rarely areolate, veinlets not included in areoles. Sori submarginal, linear, on a vascular commissure joining apices of veins, with linear, membranous indusia, or sori borne along veins, or rarely forming a narrow longitudinal band between midrib and margin and exindusiate (Taenitis); spores trilete, hyaline, usually with an equatorial flange.

About 14 genera: distributed in the tropics or subtropics, especially in tropical America; five genera and 83 species (36 endemic) in China.

1a.       Lamina with white farina abaxially ...............................................................  8. Pityrogramma

1b.       Lamina without white farina abaxially.

2a.       Rhizome covered with bristles; sori linear and forming a narrow longitudinal band between midrib and margin ................................................................................................................  7. Taenitis

2b.       Rhizome covered with scales; sori borne along veins or on a marginal commissure.

3a.       Sori borne along veins, margin not modified ...............................................  6. Anogramma

3b.       Sori borne on a marginal commissure, covered by a modified marginal indusium.

4a.       Rhizomes long creeping; ultimate segments or lobes narrow and small, usually less than 2 mm wide  9. Onychium

4b.       Rhizomes short and erect; ultimate segments or lobes usually more than 5 mm wide  5. Pteris

5. PTERIS Linnaeus, Sp. Pl. 2: 1073. 1753.

凤尾蕨属  feng wei jue shu

Liao Wenbo (廖文波), Ding Mingyan (丁明艳), Wu Zhaohong (吴兆洪 Wu Shiew-hung); Jefferson Prado

Plants terrestrial. Rhizome erect or ascending (rarely short or prostrate), vascular system a complex solenostele or dictyostele, scaly; scales brown, narrowly lanceolate or linear, membranous, firm. Fronds clustered; stipe deeply grooved adaxially, with one vascular bundle V-shaped in cross section; fronds mostly 1- or 2-pinnate, rarely 3-pinnate (Pteris cryptogrammoides), pedate, or sometimes 3-forked, pinnae entire or pectinately divided into segments, sometimes asymmetrical; basal pinnae often with 1 (or more) pinnule(s) near base on basiscopic side, this similar to main part of pinna but smaller; apical pinnae similar to lateral pinnae; costa or midvein deeply grooved adaxially, often with awns on base of each costa; venation free or areolate, veins simple or 2-forked, if areolate then with regular rows of narrow areoles along costa (sometimes also along segment costules), a few species with veinlike heterocells (“false veins”) below epidermis (P. cadieri, P. grevilleana, P. multifida, etc.); lamina herbaceous or papery when dried, sometimes subleathery, glabrous or rarely pubescent. Sori linear, along margin, except at base or basal sinus and apex, with paraphyses; annulus consisting of 16–34 incrassate cells; spores trilete (a few species with monolete spores), gray or black, surfaces scabrous or verrucose. x = 29.

About 250 species: distributed in tropical and subtropical areas, southward to New Zealand, Australia, and South Africa, north to Japan and North America; 71 species (33 endemic) in China.

Of the 71 species of Pteris, most occur mainly in S and SW China, with a few species in E China and south of the Qin Ling.

Apogamy is frequent in Pteris; the gametophytes bud off embryos without fertilization.

Key to sections

1a.       Venation areolate, veins ± anastomosing along costules; segments without cartilaginous margins; veins forming 1 or more series of areoles along costa (sometimes along midvein); costa with spines along adaxial groove (species nos. 67–75) ................................................................  3. P. sect. Campteria

1b.       Venation free.

2a.       Fronds often dimorphic or subdimorphic; pinnae/pinnules entire, with cartilaginous margins, basal pair(s) sometimes forked near base, but never pectinately divided; sterile margins often acutely serrate, rarely entire, costae without spines, grooves not erose (species nos. 1–35) ...........  1. P. sect. Pteris

2b.       Fronds monomorphic; pinnae/pinnules pectinately divided or lobed on at least one side, basal pair(s) of pinna often with 1–3(or 4) pinnules near base on basiscopic side; segments lanceolate or ± oblong-falcate, often obtuse or acute, without cartilaginous margins, entire or rarely serrate; adaxial grooves of costae with spines or erose margins (species nos. 36–66) ............  2. P. sect. Quadriauricula

Key to Pteris sect. Pteris

1a.       Fronds simple, digitate or pedate with pinnae clustered at apex of stipe and no discernable main rachis.

2a.       Fronds digitate or subdigitate, with 5–7(–9) pinnae clustered at apex of stipe and no discernable rachis.

3a.       Margins of sterile pinnae entire ................................................................  3. P. stenophylla

3b.       Margins of sterile pinnae serrate.

4a.       Mature fronds monomorphic or nearly so.

5a.       Stipes castaneous, sometimes margins slightly straw-colored; sterile pinnae margins coarsely and acutely serrate ......................................................................................  18. P. plumbea

5b.       Stipes straw-colored, brownish at base; sterile pinnae margins finely serrate.

6a.       Rhizome long creeping or procumbent; plant 15–40 cm tall; stipes 15–30 cm; pinnae mostly 5–7, digitate ...............................................................................................  7. P. dactylina

6b.       Rhizome erect or ascending; plant 5–15, lateral pinnae often one pair and each forked.

7a.       Terminal pinna linear or linear-lanceolate, acuminate at apex; lateral pinnae each 2–3(–4)-forked into linear-lanceolate pinnules ..........................................................  9. P. gallinopes

7b.       Terminal pinna ovato-lanceolate, obtuse at apex; lateral pinnae oblong-obovate, each at base on basiscopic side with an almost freely divided lobe ........................  35. P. xiaoyingiae

4b.       Mature fronds distinctly dimorphic.

8a.       Lamina with many false veins between true veins (Taiwan) ..........................  22. P. kidoi

8b.       Lamina without false veins.

9a.       Fertile pinnules 3–5 mm wide (Taiwan) .........................................  21. P. ryukyuensis

9b.       Fertile pinnules 5–12 mm wide

10a.     Margins of rhizome scales sparsely ciliate (widespread but not recorded from Hainan)  19. P. cretica

10b.     Margins of rhizome scales entire (Hainan) ................................................  6. P. morii

2b.       Fronds simple or 2- or 3-pinnulate.

11a.     Base of lamina cordate ..............................................................................  32. P. liboensis

11b.     Base of lamina tapered, cuneate or subobtuse.

12a.     Fronds always simple (Hainan) ...................................................  34. P. changjiangensis

12b.     Sterile fronds 3-pinnulate, sometimes also simple.

13a.     Plants often with both simple and 3-forked fronds (Yunnan).

14a.     Fertile fronds simple or 3-forked; fronds (or pinnae) lanceolate or linear-lanceolate, up to 15 cm; sterile fronds simple or 2-forked; fronds (or pinnae) lanceolate, adaxially without pale green band along costa (SE Yunnan) ..........................................................  1. P. pseudopellucida

14b.     Fertile fronds simple, narrowly linear, up to 50 cm; sterile fronds 3-forked, pinnae linear, adaxially with distinct pale green band along costa (S Yunnan) ....................  2. P. undulatipinna

13b.     Fronds always 3-pinnulate, never simple.

15a.     Terminal pinna sessile or subsessile.

16a.     Pinna broadly lanceolate or lateral pair subovate, 1–2 cm wide.

17a.     Plants 10–25(–40) cm tall; pinnae often 5–8 cm; terminal pinna acuminate (some times shortly acuminate); lateral pinnae never lobed ............................................  12. P. deltodon

17b.     Plants 4–6.5(–7) cm tall; pinnae often 1.5–2.5(–3.1) cm; terminal pinna obtuse at apex; each lateral pinna at base with or without a basiscopic lobe ................  35. P. xiaoyingiae

16b.     Pinna linear, 0.3–1 cm wide.

18a.     Pinna margin entire or slightly undulate; (Xizang) ..........................  3. P. stenophylla

18b.     Pinna margin finely toothed or with undulate teeth.

19a.     Pinna margin finely undulate toothed, veinlets ca 28 per cm (Taiwan)  4. P. confertinervia

19b.     Pinna margin denticulate, veinlets ca. 10 per cm ............................  7. P. dactylina

15b.     Terminal pinna stalked, stalk 1–2(–3) cm.

20a.     Fronds conspicuously dimorphic; pinnae 2.5–3.5 cm wide.

21a.     Fronds obviously dimorphic; sterile lamina with lateral pinnae falcate-ovate to lanceolate, simple; fertile lamina with lateral pinnae simple or forked, linear in outline, caudate toward apex          P. sanduensis (see P. deltodon)

21b.     Fronds monomorphic or not obviously dimorphic, with ovate-oblong lateral pinnae, simple in both infertile and fertile fronds.

22a.     Sterile pinnae 2.5–3 cm wide, margins crenulate; terminal pinna of fertile frond subsessile; (Hainan) .................................................................................  5. P. crassiuscula

22b.     Sterile pinna ca. 3.5 cm wide, margins sparsely serrate; terminal pinna of fertile frond long stalked .................................................................................  33. P. nanlingensis

20b.     Fronds monomorphic; pinnae up to 2 cm wide.

23a.     Terminal pinna broadly lanceolate ..................................................  12. P. deltodon

23b.     Terminal pinna lanceolate.

24a.     Pinna margins coarsely and acutely serrate; lateral pinnae as long as terminal pinna; rhizome erect, ca. 6 mm thick1 .................................................................  13. P. olivacea

24b.     Pinna margins serrate only on upper portion, lower parts entire, lateral pinnae shorter than terminal pinna; rhizome prostrate, ca. 3 mm thick1 ...................  14. P. baksaensis

1b.       Fronds 1(–3)-pinnate, lateral pinnae 2 pairs or more, sometimes forked.

25a.     Fronds 3-pinnate; pinnules 20 or more, 0.5–1.3 cm .....................  30. P. cryptogrammoides

25b.     Fronds 1(–2)-pinnate, pinnules mostly much more than 5 cm, if less then fronds 1-pinnate, pinnules fewer than 10.

26a.     Lateral pinnae not forked (basal fertile pinnae occasionally with single basiscopic pinnule).

27a.     Lateral pinnae often 30–40 pairs, shorter gradually downward ....................  28. P. vittata

27b.     Lateral pinnae less than 16 pairs, not gradually reduced downward.

28a.     Margins of sterile pinnae entire, ± undulate.

29a.     Pinnae shiny and smooth adaxially, pinnae margins distinctly pleated .....  25. P. venusta

29b.     Pinnae not shiny or smooth adaxially, pinnae slightly undulate.

30a.     Pinnae glabrous .................................................................................  26. P. insignis

30b.     Pinnae basal parts and costa sparsely softly hairy ........................  27. P. menglaensis

28b.     Margins of sterile pinnae dentate or serrate.

31a.     Sterile pinnae ovate or narrowly rounded lanceolate, margins with rough and acute teeth              12. P. deltodon

31b.     Sterile pinnae narrowly linear or linear-lanceolate to lanceolate, margins finely dentate or serrate.

32a.     Lamina with many false veins between true veins (Taiwan) ......................  22. P. kidoi

32b.     Lamina without false veins.

33a.     Sterile pinnae narrowly linear, often 2–4 mm wide ..............................  10. P. henryi

33b.     Sterile pinnae, broadly linear-lanceolate to lanceolate, 9–35 mm wide.

34a.     Sterile pinnae linear-lanceolate, margins with fine undulate teeth; veinlets very closely spaced, ca. 28 per cm ......................................................................  4. P. confertinervia

34b.     Sterile pinnae lanceolate, margins finely dentate; veinlets fewer.

35a.     Median sterile pinnae 4–6 Χ ca. 1 cm .....................................  29. P. ensiformis

35b.     Median sterile pinnae 9–23 Χ 1.5–3.5 cm.

36a.     Stipe glabrous; margins obtusely serrate.

37a.     Fronds herbaceous (Hainan) .......................................................  6. P. morii

37b.     Fronds papery (Yunnan) .........................................................  19. P. cretica

36b.     Sterile stipe scaly; pinnule often with conspicuous pale central band, margins irregularly dentate (Taiwan)

38a.     Pinnule often with conspicuous pale central band; fertile pinnules ca. 15 mm wide          20. P. nipponica

38b.     Pinnule uniformly green; fertile pinnules 3–5 mm wide ......  21. P. ryukyuensis

26b.     Lateral pinnae forked forked or basiscopically branched.

39a.     Lateral pinnae often with 1–3 basiscopic branches, lowest pair (or basal pairs) often with 2 or 3 branches or subpinnate.

40a.     Lateral pinnae forked; ultimate pinnules of sterile fronds linear, apically long acuminate.

41a.     Stipe and rachis light straw-colored, glabrous or ± rough ........................  10. P. henryi

41b.     Stipe and rachis chestnut-brownish, rough, sometimes glabrous  11. P. actiniopteoides

40b.     Lateral pinnae (especially sterile frond) subpinnate; pinnules of sterile frond or segments broadly lanceolate or oblong, obtuse, or linear-lanceolate, apically acuminate.

42a.     Apical pinnae decurrent at base forming thin wings along rachises; sterile pinnules linear-lanceolate, apically acuminate .............................................................................  31. P. multifida

42b.     Apical pinnae not decurrent on rachises; sterile pinnules broadly lanceolate or oblong, apically obtuse, sometimes acute ..................................................................  29. P. ensiformis

39b.     Basal pinnae with 1 or 2 basiscopic branches, occasionally second and third pairs with basiscopic branches (rarely to fifth pair).

43a.     Fronds with 2 pairs of lateral pinnae.

44a.     Sterile pinnae 2–4(–8) mm wide, margins with narrow, acute teeth.

45a.     Fertile fronds longer than sterile fronds; lamina hard papery when dry  8. P. angustipinna

45b.     Fertile fronds as long as sterile fronds; lamina hard herbaceous when dry.

46a.     Stipe and rachis slightly straw-colored, glabrous or ± rough ...............  10. P. henryi

46b.     Stipe and rachis castaneous-brownish, sometimes glabrous or rough  11. P. actiniopteroides

44b.     Sterile pinnae 10–20 mm wide.

47a.     Margins of sterile pinnae slightly serrate (Hainan) .....................................  6. P. morii

47b.     Margins of sterile pinnae densely and sharply serrate.

48a.     Stipe and rachis castaneous, midvein brown near base; (E and S China)  18. P. plumbea

48b.     Stipe up to basal part of lamina straw-colored, basal part of midvein abaxially slightly brown-straw-colored or straw-colored.

49a.     Pinnae narrowly lanceolate, ca. 1 cm wide, shortly acuminate, basal pair with stalk ca. 1 cm ..........................................................................................  15. P. quinquefoliata

49b.     Pinna lanceolate to broadly lanceolate, acuminate, basal pair sessile or subsessile  16. P. guangdongensis

43b.     Fronds with more than 3 pairs of lateral pinnae, rarely only 3 pairs.

50a.     Pinnae margins entire.

51a.     Pinnae shiny and smooth, margins distinctly pleated ............................  25. P. venusta

51b.     Pinnae not shiny or smooth, margins slightly undulate.

52a.     Apical pinnae with stipes, not decurrent at base; lateral pinnae 6–14 pairs, pinnae fertile only on upper portion, lower pinnae sterile and stalked .................................  26. P. insignis

52b.     Apical pinnae decurrent along rachis; lateral pinnae 3 pairs, basal pair sessile, second and third pairs of pinnae with decurrent base along rachis ..........................  24. P. longipinna

50b.     Pinnae margins ± serrate.

53a.     Apical 2 or 3 pairs or more of lateral pinnae long decurrent at base, forming thin wings along rachises ..........................................................................................  31. P. multifida

53b.     Apical pinna 3-forked or sometimes adjacent pair of lateral pinnae ± decurrent, rarely not decurrent.

54a.     Fronds monomorphic, sometimes subdimorphic.

55a.     Pinnules linear, ca. 3 mm wide; fronds monomorphic, sometimes slightly dimorphic.

56a.     Stipe and rachis light straw-colored, smooth or ± rough ................  10. P. henryi

56b.     Stipe and rachis castaneous-brownish, rough or sometimes glabrous  11. P. actiniopteroides

55b.     Pinnules lanceolate to oblong-lanceolate, 1.5–2 cm wide; fronds monomorphic.

57a.     Pinnae lanceolate or broadly lanceolate, apically acuminate, basal pinnae pair sessile or nearly so; (Guangdong) ...........................................................  16. P. guangdongensis

57b.     Pinnae oblong-lanceolate, apically obtuse, basal pinnae pair stalked; (Guangxi)  17. P. hui

54b.     Fronds conspicuously dimorphic.

58a.     Lamina herbaceous (Guangdong and Hainan).

59a.     Median sterile pinnae 9–23 Χ 1.5–2.5 cm ........................................  6. P. morii

59b.     Median sterile pinnae 4–6 Χ ca. 1 cm .....................................  29. P. ensiformis

58b.     Lamina papery to slightly leathery (E to SW China, not in Hainan).

60a.     Plants 50–70 cm; sterile pinnae narrowly lanceolate or lanceolate, 1–2 cm wide, papery when dry; stipe always smooth ....................................................  19. P. cretica

60b.     Plants up to 1.5 m; sterile pinnae lanceolate to oblong-lanceolate, 2–5 cm wide, slightly leathery when dry; stipe sometimes with distal part verrucose-tuberculate  23. P. esquirolii

Key to Pteris sect. Quadriauricula

1a.       Frond 3-partite; lateral divisions same shape as terminal one but smaller (P. ser. Longipedes)  66. P. longipes

2a.       Lateral pinnae 12–20 per rachis ...................................................................  66. P. longipes

2b.       Lateral pinnae 4–6 per rachis ...........................................................  66bis. P. paucipinnula

1b.       Frond pinnate (P. ser. Quadriauritae).

3a.       Fronds dimorphic ...........................................................................................  44. P. cadieri

3b.       Fronds monomorphic.

4a.       Pinnae entire; frond with 2 or 3 pairs of lateral pinnae .....................  66ter. P. quadristipitis

4b.       Pinnae pectinately divided; frond usually with 4 or more pairs of lateral pinnae.

5a.       Lateral pinnae asymmetrical along costae, lobes shorter and/or fewer, sometimes almost absent, on one side.

6a.       Basiscopic side of costa with 0–3 segments, acroscopic side with 3–6 segments; segments 3.5–6 Χ ca. 0.6 cm, lowermost sometimes with secondary lobes so pinna 2-pinnatifid  43. P. formosana

6b.       Basiscopic side of costa regularly pectinately lobed, acroscopic side with fewer and smaller segments, sometimes almost entire.

7a.       Stipe straw-colored or dark straw-colored; costa with finely toothed protuberances or short spines along groove.

8a.       Plants often to 1.5(–2) m; lamina segments distinctly decurrent at base  39. P. malipoensis

8b.       Plants 0.8–1 m; lamina segments separate, basiscopic not distinctly decurrent at base  41. P. inaequalis

7b.       Stipe castaneous-red or dark castaneous; with erose margin along adaxial groove of costa.

9a.       Plants often 1–1.5 m; rhizome scales with paler margins .................  38. P. dissitifolia

9b.       Plants often 0.3–0.8 m; rhizome scales uniformly blackish brown.

10a.     Segments distant, almost entirely restricted to basiscopic side, acroscopic side entire or with very few shorter segments, distinctly decurrent at base; veins of sterile frond ending at base of shortly acuminate teeth .............................................................  37. P. semipinnata

10b.     Segments closely spaced, on both sides with acroscopic segments only slightly shorter than basiscopic segments, not distinctly decurrent at base; veins of sterile frond ending within long acuminate teeth .................................................................................  36. P. dispar

5b.       Lateral pinnae (at least of sterile fronds) symmetrical along costa with acroscopic side as wide as basiscopic side.

11a.     Basal pinnae similar to upper pinnae, without any basiscopic branches.

12a.     Abaxial side of costa and costule, and abaxial surface of segments glabrous  40. P. excelsa

12b.     Lower part of costa with coarse hairs; costules and abaxial surface of segments ± hairy.

13a.     Pinnae segments 8–12 pairs restricted to basal half of pinna, pinna apex linear, more than half as long as pinna ..........................................................................  46. P. heteromorpha

13b.     Pinnae segments 15–20 pairs distributed along most of length of pinna, pinna apex shorter than lower segments ............................................................................  47. P. decrescens

11b.     Basal pinnae with one or more basiscopic branches near their bases.

14a.     Lamina with narrow lines (false veins) between veins ......................  45. P. grevilleana

14b.     Lamina without lines.

15a.     Margins of sterile segments serrate ....................................................  42. P. amoena

15b.     Margins of sterile segments entire, sometimes slightly undulate.

16a.     Two veins from adjacent segments reaching base of incision of pinnae, forming an acute triangle, sometimes forming areoles; pinnae often pinnatifid ............................  65. P. linearis

16b.     Two veins from adjacent segments reaching incision of pinnae, forming a short obtuse triangle, pinnae pinnate or deeply pinnatisect.

17a.     Segment apex mucronate.

18a.     Costae with coarse hairs on basiscopic side; midvein, and basiscopic side of segments ± hairy ..................................................................................  48. P. longipinnula

18b.     Costa smooth or rough.

19a.     Lower vein of basal pair of veins of segment near to costa and subparallel to costa; basal pinnae with ca. 2 cm stalk .....................................................  51. P. splendida

19b.     Lower vein of basal pair of veins of segment oblique, not parallel to costa; basal pinnae sessile or subsessile.

20a.     Stipe, rachis and costae all green ....................................  48. P. longipinnula

20b.     Stipe, rachis and costae straw-colored or partly ± purple.

21a.     Stipe as long as lamina; lamina oblong-ovate; indusia purple or straw-colored             49. P. aspericaulis

21b.     Stipe 2 Χ as long as lamina; lamina ovate-triangular, 3-forked; indusia yellowish          50. P. caiyangheensis

17b.     Segment apex not mucronate.

22a.     Pinnae abaxially with short or flakelike hairs.

23a.     Pinnae abaxially with flakelike hairs (especially along costa); costa blue-green; segment costules with needlelike spines ............................................  60. P. viridissima

23b.     Pinnae abaxially sparsely shortly pubescent on basiscopic side; costa straw-colored, sometimes light purple; segment costules with flat spines.

24a.     Segments acute, adaxially sparsely shortly hairy; costules adaxially with thin acute spines                                                                                          61. P. hirsutissima

24b.     Segments obtuse, adaxially glabrous; costules adaxially without spines  59. P. puberula

22b.     Pinnae abaxially glabrous.

25a.     Lamina subleathery when dry; costae with soft flat spines adaxially  62. P. subquinata

25b.     Lamina papery or herbaceous when dry; costae with distinct, needlelike or oblate spines.

26a.     Fronds with (9–)11–16 pairs of pinnae, basal pinnae often each with (2 or)3 or 4 basiscopic branches ...............................................  52. P. setulosocostulata

26b.     Fronds with (3–)5–9 pairs of pinnae, basal pinnae each with 1 or 2 basiscopic branches.

27a.     Pinnae divided to costa; stipe reddish brown.

28a.     Stipe glossy, scales expanded at base ....................................  63. P. bella

28b.     Stipe not glossy, muricate, scales not expanded at base  64. P. scabristipes

27b.     Pinnae pinnatifid; stipe mainly straw colored.

29a.     Plants more than 1.3 m tall; lateral pinnae ca. 35 cm, median pinnule segments 35–55 Χ 7–9 mm .....................................................................  58. P. majestica

29b.     Plants 0.5–1 m tall, lateral pinnae 12–23 cm, median pinnule segments 10–16 Χ 4–5 mm.

30a.     Lateral pinnae oblique upward.

31a.     Lateral pinnae widest at base (triangular) or oblong ..  54. P. oshimensis

31b.     Lateral pinnae widest at their middle ...............................  55. P. fauriei

30b.     Lateral pinnae spreading or oblique.

32a.     Lateral pinnae triangular (widest at base) or oblong in outline  53. P. kiuschiuensis

32b.     Lateral pinnae often widest at their middle.

33a.     Median pinnae 12–16 Χ 2–2.5 cm; segments narrowed to apex, linear-lanceolate; 2–4 mm apart ................................................  56. P. angustipinnula

33b.     Median pinnae 17–23 Χ 4–6.5 cm; segments not narrowed to apex, linear, obtuse, ca.1 mm apart............................................ 57. P. obtusiloba

Key to Pteris sect. Campteria

1a.       Lamina pinnate; basal pair of pinnae each with 1 (sometimes 2) basiscopic pinnules.

2a.       Stipes straw-colored to pale green; lamina hard papery .................................  67. P. biaurita

2b.       Stipes chestnut; lamina herbaceous.

3a.       Costa wings to 5–6 mm wide on both sides; segments 6–9 mm wide .........  68. P. maclurei

3b.       Costa wings ca. 3 mm wide; segments 4–6 mm wide ..........................  69. P. maclurioides

1b.       Lamina 3-partite to pedate, divided into 3 or more branches, each branch pinnate; basal pair of pinnae not branched on basicopic side.

4a.       Areoles along costules of segments as well as along costa; stipes and rachis straw-colored or light brown.

5a.       Segments 2.5–7 cm, 7–12 mm apart .............................................................  74. P. finotii

5b.       Segments 1–3 cm, 2–5 mm apart .............................................................  75. P. tripartita

4b.       Areoles restricted to along main costa, absent from segments; veins above areoles all free (sometimes basal veins of segment forming areoles); stipes and rachis straw-colored or red or brown-red.

6a.       Stipes and rachis straw-colored; segments 30–40 Χ ca. 3 mm, 5–8 mm apart (Taiwan)  73. P. taiwanensis

6b.       Stipes and rachis red or brown-red, or rachis rarely straw-colored; segments 10–25 Χ ca. 4 mm, up to 4 mm apart.

7a.       Stipe and rachis with purple-brownish bristles .....................................  70. P. wallichiana

7b.       Stipe and base of rachis glabrous (sometimes stipes slightly hairy).

8a.       Lamina abaxially with red-brown bristles .........................................  71. P. austrosinica

8b.       Lamina abaxially subglabrous.

9a.       Lateral pinnules 20–25 Χ ca. 4.5 cm; segments 2–4 mm apart  72. P. occidentalisinica

9b.       Lateral pinnules 11–20(–25) Χ 2–2.5(–3.5) cm; segments 1–2 mm apart
.....................................................................................................  70. P. wallichiana

1. Pteris sect. Pteris

凤尾蕨组  feng wei jue zu

Plants small or medium sized (0.05–1.5 m tall). Fronds often dimorphic or subdimorphic, pedate, digitate or imparipinnate, seldom simple; pinnae or basal pinnae pair (sometimes few pairs) forked, never pectinate or pinnate; pinnae often narrowly linear or lanceolate, acuminate, with cartilaginous margins, sterile pinnae or sterile margins serrulate, seldom entire, costae or midveins adaxially erose; venation free. Sori linear, along segment margins, absent at apex and sinuses; indusia gray-brown or brown, linear, membranous, persistent.

35 species (20 endemic) in China.

Species in China are distributed in tropical and subtropical areas, from North to South (Qin Ling). Some species are calciphilous and can be found on limestone and walls.

1. Pteris pseudopellucida Ching, Lingnan Sci. J. 15: 395. 1936.

单叶凤尾蕨  dan ye feng wei jue

Pteris pellucida C. Presl f. simplex Hooker; P. subsimplex Ching.

Plants 30–40 cm tall. Rhizome erect, short, ca. 1.5 cm in diam., scaly at growing tips. Fronds dimorphic, simple or 2- or 3-forked; stipe straw-colored to light brown, slightly lustrous, 15–20 cm Χ ca. 2 mm, glabrous; sterile fronds: lamina (or pinnae) lanceolate, ca. 20 Χ 2.5–3 cm, base broadly cuneate or subobtuse, margins subcartilaginous, undulate, serrate upward, apically shortly serrulate, apex acuminate; fertile fronds (or pinnae) linear-lanceolate, ca. 20 Χ 1–2 cm, entire, apex sterile, slightly serrulate. Lamina pale green, papery when dried, glabrous; midvein straw-colored, convex abaxially; veins simple or 2-forked.

Common in ravines, bamboo forests; 100–200 m. SE Yunnan (Hekou) [N Vietnam (Lao Cai)].

The type of Pteris pseudopellucida is from Hekou, Yunnan.

2. Pteris undulatipinna Ching, Acta Bot. Austro Sin. 1: 3. 1983.

波叶凤尾蕨  bo ye feng wei jue

Plants 40–60 cm tall. Rhizome erect, short, ca. 1 cm in diam., scaly at growing tips. Fronds dimorphic; stipe light straw-colored, 15–20 cm Χ ca. 1.5 mm. Sterile fronds: lamina linear, up to 40 Χ ca. 1 cm, base sterile and slightly undulate, apex caudate and slightly serrulate. Fertile fronds: 3-forked, terminal pinnae linear, 25–30 Χ 1–1.5 cm, basally cuneate, sessile, separate, margin entire at base, revolute, spreading serrate upward, apex long acuminate; lateral pinnae as apical pinna but shorter, somewhat falcate; lamina pale green, thinly papery when dried, glabrous with black longitudinal striation ca. 4 mm wide on both sides along midvein; midvein light straw-colored, abaxially convex; veins simple or 2-forked.

* Bamboo forests; ca. 900 m. S Yunnan (Mengla).

3. Pteris stenophylla Wallich ex Hooker & Greville, Icon. Filic. t. 130. 1829.

狭羽凤尾蕨  xia yu feng wei jue

Pteris cretica Linnaeus var. stenophylla (Wallich ex Hooker & Greville) Baker; P. digitata Wallich, nom. nud.; P. pellucida C. Presl var. stenophylla (Wallich ex Hooker & Greville) C. B. Clarke.

Plants ca. 50 cm tall. Rhizome ascending, ca. 6 mm in diam. Fronds subdimorphic, clustered; stipe straw-colored, 30–35 cm Χ ca. 1.5 mm, glabrous; lamina digitate, pinnae 3–5, clustered at stipe apex, linear-lanceolate, up to 20 cm Χ 8–10 mm (sterile fronds wider than fertile fronds), base cuneate, margin entire and cartilaginous, apex long acuminate; lamina gray-green, papery when dried, glabrous; midvein straw-colored, convex adaxially, veins simple, rarely 2-forked.

Among dry rocks in open forests; 2500–3000 m. SE Xizang (Himalaya) [Bhutan, N India, Nepal].

The type of Pteris stenophylla is from Nepal.

4. Pteris confertinervia Ching, Acta Bot. Austro Sin. 1: 4. 1983.

密脉凤尾蕨  mi mai feng wei jue

Plants ca. 35 cm tall. Rhizome erect, short, apically scaly. Fronds submonomorphic, clustered; stipe light-brown, slender, ca. 1 mm in diam., glabrous. Sterile fronds: stipe up to 8 cm, lamina similar to those of fertile fronds in shape but shorter, or sometimes with more lateral pinnae pairs, apical pinna decurrent on lateral pinnae forming a wing. Fertile fronds: stipe up to 13 cm; lamina pedate, connate at base; terminal pinna linear-lanceolate, ca. 25 Χ 0.9 cm, margins densely undulate-denticulate at base of pinnae, apex long acuminate; lateral pinnae similar but shorter. Lamina light brown-green, slightly lustrous, herbaceous when dried; midvein convex on both sides; veins closely spaced, ca. 28 per cm.

* Taiwan (Taibei).

Fraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes, 101. 2008) regarded Pteris confertinervia as a synonym of P. cretica var. cretica.

5. Pteris crassiuscula Ching & C. H. Wang, Acta Phytotax. Sin. 8: 142. 1959.

厚叶凤尾蕨  hou ye feng wei jue

Plants ca. 50 cm tall. Rhizome erect, short, ca. 1 mm in diam., apex with brownish black scales. Fronds dimorphic. Sterile fronds: stipe straw-colored, light brown, or castaneous-brown, slightly lustrous, 10–25 cm Χ 1–2 mm, glabrous; lamina 3-pinnulate, terminal pinnae lanceolate, 20–25 Χ 2.5–3 cm, basally narrowly cuneate, somewhat decurrent, shortly stalked (1–5 mm), margins cartilaginous and slightly serrate, apex acuminate and serrate; lateral pinnae as terminal pinnae but shorter and sessile. Fertile fronds: stipe up to 45 cm; lamina oblong, 25–30 cm, 1-pinnate; pinnae 2 pairs per frond, subsessile, decumbent, ca. 5 cm apart, narrowly linear, 20–25 cm Χ ca. 5 mm. Lamina pale green or brown-green, leathery when dried, firm, both surfaces glabrous; midvein straw-colored or castaneous-brown, abaxially convex, veins conspicuous, simple or 2-forked from base.

* Forests along valleys; ca. 700 m. Hainan (Baoting, Diaoluoshan).

6. Pteris morii Masamune, Trans. Nat. Hist. Soc. Formosa 32: 342. 1942.

琼南凤尾蕨  qiong nan feng wei jue

Plants 50–70 cm tall. Rhizome erect, short, 1–1.5 cm in diam., apex with black-brown, entire scales. Fronds clustered, dimorphic; stipe pale straw-colored or castaneous, 30–45 cm Χ ca. 2 mm, firm, glabrous. Sterile fronds: lamina 1-pinnate, triangular-ovate in outline; pinnae 2–4 pairs, sometimes digitate, basal 1 or 2 pairs often forked, opposite, slightly decumbent, shortly stalked, remainder sessile, simple, narrowly lanceolate, 9–23 Χ 1.5–2.5 cm, base narrowly cuneate, margins cartilaginous and obtusely serrate, apex acuminate or acute and serrate. Fertile fronds: pinnae 2–5 pairs, basal pair often 2-forked, lower ones stalked (3–5 mm), upper ones sessile, linear, sometimes linear-lanceolate, 20–25 Χ 0.8–1 cm, sterile margins serrulate. Lamina pale green, thickly leathery when dried, glabrous, sterile pinnae sometimes with grayer lines along midvein adaxially, midvein abaxially convex, straw-colored, veins abaxially conspicuous, simple or forked.

* Dense forests along valleys; near sea level to 900 m. Hainan.

7. Pteris dactylina Hooker, Sp. Fil. 2: 160. 1858.

指叶凤尾蕨  zhi ye feng wei jue

Plants 20–40 cm tall. Rhizome prostrate, short, ca. 3 mm in diam., apex scaly. Fronds clustered, dimorphic; sterile fronds as long as fertile fronds; stipe straw-colored, brown at base, slightly lustrous, slender, 15–30 cm Χ ca. 1 mm, smooth or rarely scabrous; lamina digitate, pinnae (3–)5–7, basal pairs nearly 3-forked, rarely or apical pinnae 2- or 3 forked, clustered; terminal pinna narrowly linear, 8–10(–15) Χ (0.2–)0.3–0.4(–0.8) cm, sessile or shortly stalked, base cuneate, not decurrent, apex acuminate; lateral pinnae similar, shorter, somewhat falcate, sessile, margins pinnae denticulate, midvein straw-colored, glabrous, abaxially convex, adaxially grooved; veins conspicuous, separate, straight, parallel to each other, slightly oblique, simple or rarely forked from base; fertile pinnae similar but margins subentire, serrulate only at apex; lamina gray-green, firmly herbaceous when dried, both surfaces glabrous. Indusia linear, pallid, membranous, subentire.

Guizhou (Fanjingshan), Sichuan, SE Xizang, Yunnan [Bhutan, NE India, Nepal].

According to W. C. Shieh (Fl. Taiwan: 1: 224. 1994), Pteris dactylina is distributed in SE and SW Taiwan. The type is from Sikkim.

Pteris pseudodactylina Ching & S. K. Wu (Fl. Xizang. 1: 66–68. 1983) from SE Xizang is closely related, differing by the wider sterile pinnae, ca. 1 cm wide, with larger cartilaginous marginal teeth and fertile fronds with the 3 central pinnae decurrent at base.

Pteris gracillima Ching & S. K. Wu (Fl. Xizang. 1: 65–66. 1983) from SE Xizang is also closely related. It is described as a more delicate plant, 13–28 cm tall, with 3-pinnulate fronds, the pinnae 8–22 Χ ca. 0.4 cm, with the lateral pinnae much shorter than the terminal pinna, and the margins of the sterile pinnae undulate or sinuate, not toothed. Pteris stenophylla is also rather similar to P. gracillima but has pinnae more equal in length and ca. 1 cm wide.

8. Pteris angustipinna Tagawa, Acta Phytotax. Geobot. 4: 203. 1935.

细叶凤尾蕨  xi ye feng wei jue

Plants 20–40 cm tall. Rhizome and base of stipes scaly; scales castaneous-brown, linear or subulate-linear, ca. 3 mm, entire, basally sparsely glandular pubescent. Fronds somewhat dimorphic. Sterile fronds: stipe straw-colored, light brown at base, slightly scabrous, glabrous, adaxially grooved, 3–10 cm; lamina digitate or rarely pinnate (2 pairs of lateral pinnae); terminal pinna linear, 10–17 cm Χ 2–3(–5) mm, base shortly cuneate, shortly stalked or sessile, margins serrulate, apex acuminate; lateral pinnae narrower, 5–10 cm, sessile or shortly stalked; basal pinnae pair sometimes forked. Fertile fronds: stipe 5–17 cm; lamina pinnate (2 pairs of lateral pinnae); terminal pinna linear, 10–27 cm Χ 2–4 mm, stalked (3–15 mm); lateral pinnae shorter; basal pair sometimes forked, sterile margins serrulate. Lamina thinly papery when dried; veins conspicuous, simple or forked.

* On rocks in forests, very rare; 2000–2600 m. Taiwan.

9. Pteris gallinopes Ching, Acta Bot. Austro Sin. 1: 4. 1983.

鸡爪凤尾蕨  ji zhua feng wei jue

Plants small, 10–20 m tall. Rhizome erect, short, ca. 1 cm in diam., apex with black scales. Fronds clustered, monomorphic; stipe straw-colored, light brown at base, opaque, 6–10 cm Χ ca. 1 mm, glabrous, adaxially shallowly grooved; lamina digitate, 4–6 Χ 4–7 cm; pinnae 5(–9), clustered at apex of stipe, linear, 3–7 cm Χ 3–4 mm, middle ones slightly longer, stalked (2–4 mm), base cuneate, margins of sterile fronds slightly serrate, fertile fronds entire, apex acuminate and slightly serrulate; lateral pinnae similar but smaller and slightly falcate, apically acuminate or mucronate. Lamina gray-green, herbaceous when dried, glabrous; midvein prominent on both sides, straw-colored to pale green, glabrous; veins conspicuous, separate, parallel to each other, ca. 1.5 mm apart, oblique, simple or forked. Sori absent at segment base and apex; indusia gray-brown, linear, thinly membranous, entire.

* Limestone crevices within forests; 800–1700 m. Guizhou (Anshun, Douyun), Hubei (Hefeng), Sichuan (Baoxing, Emeishan, Leshan), Yunnan.

The type of Pteris gallinopes is from Yunnan.

10. Pteris henryi H. Christ, Bull. Herb. Boissier 6: 957. 1898.

狭叶凤尾蕨  xia ye feng wei jue

Plants 30–50 cm tall. Rhizome ascending, short, ca. 1 cm in diam., apex with brownish black scales. Fronds clustered, monomorphic or somewhat dimorphic; sterile fronds shorter than fertile fronds; stipe light straw-colored, 15–20 cm (stipes of sterile fronds shorter), 1–2 mm in diam., smooth or ± rough, glabrous, angulate; lamina 1-pinnate, oblong-ovate in outline, 20–30 Χ 10–15 cm; pinnae (2–)4–6 pairs, opposite, 5–7 cm apart near base of frond, very decumbent, basal pair shortly stalked, often 3- or 4-forked; upper pairs sessile, often 2–4-forked, seldom simple; terminal pinna 2- or 3-forked, rarely simple and shortly stalked; pinnules linear, 10–20 Χ (0.2–)0.3–0.4 cm (sterile segments broader), base broadly cuneate and slightly oblique, sterile margins slightly serrulate, fertile margins entire, apex long acuminate. Lamina gray-green, papery when dried, glabrous on both surfaces; midvein prominent on both sides, pale straw-colored; veins conspicuous, slightly inflexed, spreading, simple or forked. Sori narrowly linear; indusia brown, linear, membranous, entire.

* Limestone crevices; 400–2300 m. Guangxi (Lengyun, Leye), Guizhou, SW Henan (Xichuan), S Shaanxi, Sichuan (Chongqing, Leibo), Yunnan.

The type of Pteris henryi is from Yunnan (Mengzi).

11. Pteris actiniopteroides H. Christ, Bull. Herb. Boissier 7: 6. 1899.

猪鬣凤尾蕨  zhu zong feng wei jue

Pteris cretica Linnaeus var. melanocaulis Baker.

Plants 5–30(–60) cm tall. Rhizome erect, short, 1–1.5 cm in diam., apex with black-brown, entire scales. Fronds many, clustered, monomorphic or somewhat dimorphic, sterile fronds shorter than fertile fronds; stipe erect or spreading, castaneous-brown, 3–6(–20) cm (stipes of sterile fronds shorter), 0.5–1 mm in diam., scabrous or occasionally smooth; sterile lamina 1-pinnate (juvenile plants digitate), oblong-ovate or broadly triangular in outline; lateral pinnae 1 or 2 pairs, opposite, slightly decumbent, 2-forked or basal pair 3-forked, base of apical 3-forked, pinnae not or slightly decrescent; pinnules narrowly linear, often ca. 10 cm Χ 4–5 mm, base cuneate, margins serrate, apex long acuminate; fertile fronds: lateral pinnae often 2–4 pairs, opposite, 2–4 cm apart, slightly decumbent, basal pair 2–4-forked and shortly stalked, gradually simpler upward and sessile, base of 3-forked terminal pinna slightly or not decurrent; pinnules/segments narrowly linear, often 10–18 cm Χ 2–3 mm, base cuneate, margins serrulate toward apex only, apex long acuminate. Lamina pale green, thinly papery when dried, glabrous; midvein convex on both sides, light straw-colored, sometimes castaneous-brown at base; veins conspicuous on both sides, ca. 1 mm apart, slightly oblique, simple or forked. Sori absent at segment apices; indusia slightly broad, pallid, thinly membranous, entire.

* Limestone crevices; 600–2000 m. Chongqing, SE Gansu (Wenxian), N Guangxi (Luocheng, Yishan), Guizhou, N Henan (Huixian), Hubei (Yichang), S Shaanxi (Xixiang), Sichuan, Yunnan.

Pteris actiniopteroides is a xeric and calciphilous plant.

The type of Pteris actiniopteroides is from Yunnan (Mengzi).

12. Pteris deltodon Baker, J. Bot. 26: 226. 1888.

岩凤尾蕨  yan feng wei jue

Pteris nana H. Christ; P. trifoliata H. Christ (1899), non Fιe (1857).

Plants 15–25 cm tall. Rhizome erect, short, ca. 1 cm in diam., apex with black-brown scales. Fronds clustered, submonomorphic; stipe brown at base, upper part straw-colored, slightly lustrous, 10–20 cm Χ 1–2 mm; sterile lamina similar to fertile lamina in shape but shorter and wider, terminal pinnae oblong-lanceolate, lateral pinnae ovate, margins serrate except at base; fertile lamina pinnate, ovate to triangular-ovate, 10–20 Χ 4–7 cm; pinnae 3–5; terminal pinna broadly lanceolate, 5–8 Χ 1.2–2 cm wide, sessile or shortly stalked, base broadly cuneate, sterile margins triangular-serrate toward apex, remainder entire, apex acuminate; lateral pinnae smaller, opposite, decumbent, falcate, sessile, base obtuse and oblique, apex mucronate. Lamina light brown, papery when dried, glabrous on both surfaces; costae straw-colored, abaxially convex; veins conspicuous, simple or forked. n = 53, 2n = 106.

On dark and slightly dry limestone, common in limestone soils in SW China; 300–2500 m. SW Guangxi (Longsheng), Guizhou (Douyun, Pingtang, Anlong), Sichuan, Taiwan, Yunnan (Guangnan, Maguan, Pingbian), Zhejiang [Japan (Ryukyu Islands), Laos, Vietnam].

The type of Pteris deltodon is from Sichuan (Emeishan).

Pteris sanduensis X. Y. Wang & P. S. Wang (Pterid. Fl. Guizhou, 598–599. 2001) was compared in the protologue to P. deltodon and P. crassiuscula where it was separated by the dimorphic fronds, the sterile fronds with the terminal pinna 8–11 Χ 7–8 cm.

13. Pteris olivacea Ching, Acta Bot. Austro Sin. 1: 5. 1983.

长羽凤尾蕨  chang yu feng wei jue

Plants 25–30 cm tall. Rhizome erect, short, ca. 6 mm in diam., apex with dark brown scales. Fronds clustered, monomorphic; stipe 12–14 cm Χ 1.5–2 mm; lamina 3-pinnulate; lateral pinnae/pinnules 1 pair, opposite, decumbent, lanceolate, somewhat falcate, stalked (ca. 2 mm), ca. 12 Χ 1.5 cm, basally asymmetrical, margins mostly entire, upper 1/3 of sterile pinnules serrate, apex long acuminate; terminal pinna similar in shape but longer, long stalked (1.5–3.5 cm), base symmetrically cuneate; lamina brown-green, thinly papery when dried, both surfaces glabrous; costae straw-colored, glabrous, grooved adaxially; veins inconspicuous, oblique, parallel, ca. 1.5 mm apart, simple or forked.

* Under scrub by valleys; 1100–1300 m. SE Yunnan (Daweishan, Pingbian).

14. Pteris baksaensis Ching, Acta Phytotax. Sin. 9: 348. 1964.

白沙凤尾蕨  bai sha feng wei jue

Plants ca. 40 cm tall. Rhizome prostrate, ca. 3 mm in diam., apex with brown, linear-lanceolate scales. Fronds spaced, monomorphic; stipe brown at base, apically straw-colored, ca. 25 cm (stipes of sterile fronds shorter), glabrous, adaxially grooved; lamina digitate, pinnae 3, ca. 20 cm, middle part 6–10 cm; lateral pinnae sessile, opposite, very decumbent, lanceolate, subfalcate, ca. 1.2 cm wide, base slightly oblique, broadly cuneate, margins cartilaginous, sterile fronds entire at base, finely and acutely serrate toward apex, apex acuminate; terminal pinna similar, narrowly lanceolate, up to 22 Χ ca. 1.4 cm, base symmetrical, cuneate, stalked (1–1.5 cm); lamina gray-green, papery when dried, costae with pallid vertical stripes on both sides on adaxial surface, both surfaces glabrous; costae straw-colored, glabrous, adaxially grooved; veins inconspicuous, dense, parallel, oblique, 1- or 2-forked.

* Forests. Hainan (Baisha).

15. Pteris quinquefoliata (Copeland) Ching, Fl. Reipubl. Popularis Sin. 3(1): 24. 1990.

五叶凤尾蕨  wu ye feng wei jue

Pteris nana H. Christ var. quinquefoliata Copeland, Philipp. J. Sci., Sect. C. Bot. 3: 282. 1908.

Plants ca. 25 cm tall. Rhizome not seen. Fronds clustered, monomorphic; stipe straw-colored, slender, ca. 15 cm; lamina pinnate or digitate, ca. 10 Χ 10 cm; lateral pinnae often 2 pairs, opposite, oblique, basal pair shortly stalked, 2-forked, second lateral pinnae equal in shape to apical pinnae but shorter, basal pinnule oblong-lanceolate, ca. 3 cm, slightly reflexed; terminal pinnae longest, shortly stalked (ca. 1 cm), lanceolate, ca. 8 Χ 1 cm, base cuneate, margins entire below apex, apex mucronate and acute, serrate; lamina papery when dried, both surfaces glabrous; midvein adaxially brown-straw colored, convex; veins simple or 2-forked at base.

* On limestone. N Guangdong (Lianxian).

16. Pteris guangdongensis Ching, Acta Bot. Austro Sin. 1: 5. 1983.

广东凤尾蕨  guang dong feng wei jue

Plants 30–50 cm tall. Rhizome not seen. Fronds clustered, monomorphic; stipe brown at base, apex and rachis straw-colored, slightly lustrous, 20–35 cm Χ ca. 2 mm, apex compressed, glabrous; lamina 1-pinnate, ovate in outline, 20–25 Χ 10–15 cm; pinnae 2–6 pairs, opposite, sessile or basal pair subsessile, decumbent; lateral pinnae similar to terminal pinnae in shape, but shorter, sometimes broadly lanceolate to oblong-lanceolate, basal pair 2-forked, sterile pinnae tips acutely serrate, margins of sterile pinnae slightly irregularly and acutely serrate, fertile pinnae entire; terminal pinna largest, ± connate with nearest lateral pair of pinnae, lanceolate, 10–15 Χ 2–2.5 cm,  base not or only slightly decurrent, broadly cuneate, margin entire, apex acuminate; lamina brown-green, thinly leathery when dried, matte, glabrous on both surfaces; with scales on axil of pinnae, scales black-brownish, linear; midvein abaxially convex, straw-colored; veins conspicuous and convex on both sides, simple or forked.

* On rocks; ca. 800 m. Guangdong (Wengyuan, Yangchun, Yunfu).

The type of Pteris guangdongensis is from Guangdong (Yunfu).

17. Pteris hui Ching, Sinensia 1: 9. 1929.

胡氏凤尾蕨  hu shi feng wei jue

Plants ca. 25 cm tall. Rhizome erect, short, apex with black-brown scales. Fronds clustered, monomorphic; stipe straw-colored, 6–17 cm Χ ca. 2 mm, as long as rachis, glabrous; lamina 1(or 2)-pinnate, broadly ovate, 5–10 Χ 10–12 cm; lateral pinnae 1 or 2 pairs, opposite, stalked, basal pair largest, with 2 or 3 pinnules; middle pinnule largest, oblong-lanceolate to oblong-ovate, 3.5–6 Χ 1.5–2 cm, stalked (5–10 mm), base broadly cuneate, sterile margins serrate, apex obtuse; upper pair of lateral pinnae simple, similar to middle pinnule of basal pair; terminal pinna up to 8–9 cm; lamina pale green, subleathery when dried, midvein abaxially convex, light brown-straw-colored; veins 2-forked.

* Limestone crevices; ca. 700 m. NW Guangxi (Fengshan).

18. Pteris plumbea H. Christ, Notul. Syst. (Paris) 1: 49. 1909.

栗柄凤尾蕨  li bing feng wei jue

Pteris plumbea var. sintenensis Masamune.

Plants 25–35 cm tall. Rhizome erect or slightly oblique, apex with black or brown scales. Fronds clustered, submonomorphic; stipe castaneous, 4-angled, 10–20 cm Χ 1–2 mm; rachis similar (sometimes straw-colored when young), margin sometimes straw-colored, shiny, glabrous; lamina (mature) monomorphic, 1-pinnate, oblong or ovate-oblong in outline, 20–25 Χ 10–15 cm; pinnae often 2 pairs, opposite, decumbent; basal pinnae often with 2 or 3 pinnules; lateral pinnules much shorter, uppermost one often adnate to apical pinna, base ± decrescent; upper lateral pinnae usually simple, sessile, base not decurrent, margins cartilaginous, sterile parts serrate, fertile parts entire; terminal pinna linear-lanceolate, 10–15 Χ 0.8–1 cm, base broadly cuneate, slightly oblique, apex acuminate; lamina gray-green or adaxially brown, herbaceous when dried, glabrous; midvein biconvex, veins conspicuous, simple or forked.

Limestone crevices within open forests; 200–700 m. Fujian (Xianxialing), Guangdong, Guangxi, S Hunan (Yizhang), SW Jiangsu (Jiangning), W Jiangxi (Pingxiang), Taiwan, Zhejiang (Jinhua, Leqing) [Cambodia, India (Assam), Japan (Ryukyu Islands), Philippines, Thailand, N Vietnam].

The type of Pteris plumbea is from Guangdong (Shantou). Fl. Taiwan treats P. plumbea var. sintenensis as a synonym of P. cadieri.

19. Pteris cretica Linnaeus, Mant. Pl. 1: 130. 1767.

欧洲凤尾蕨  ou zhou feng wei jue

Rhizome creeping or ascending, ca. 1 cm in diam., apex with black-brown scales. Fronds clustered, dimorphic or subdimorphic; stipe straw-colored, sometimes brown, seldom castaneous, 10–45 cm; sterile fronds: stipe shorter, ca. 2 mm in diam., glabrous; lamina 1-pinnate or sometimes digitate, ovate in outline, 10–30[–40] Χ 6–20[–35] cm; pinnae (2 or)3–5(–7) pairs, often opposite, decumbent, basal pinnae each with 2(or 3) pinnules, shortly stalked, upper pinnae narrowly lanceolate or lanceolate, sometimes with basiscopic lobe, 10–18(–24) Χ 1–1.5(–3) cm, sessile, base broadly cuneate, margin white cartilaginous and serrate, apex acuminate. Fertile fronds longer: stipe 30–45 cm; pinnae 3–5(–8) pairs, opposite or upper alternate, decumbent, basal pinnae with (1 or)2(or 3) pinnules, shortly stalked (2–5 mm); upper pinnae linear, sometimes with lateral pinnule, 12–25 Χ 0.5–1.2 cm, sessile, base broadly cuneate, terminal pinna 3-lobed, base decurrent; midvein abaxially extremely convex, straw-colored, glabrous; veins conspicuous on both surfaces, distant, oblique, simple or forked at base; lamina green or gray-green, papery when dried, glabrous. 2n = 58*, 59, 87*.

Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, SW Henan, Hubei, Hunan, Jiangxi, S Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, W Zhejiang [Bhutan, Cambodia, India, Japan, Kashmir, Laos, Nepal, Philippines, Sri Lanka, Thailand, Vietnam; Africa, SW Asia, Europe, Pacific islands (Fiji, Hawaii)].

The type is from the Mediterranean coast. The species as a whole occurs widely throughout the tropics and subtropics. It is frequently cultivated and escapes, obscuring the natural distribution.

Two varieties are usually recognized. The correct placement of Pteris cretica var. sylvatica X. Y. Wang & P. S. Wang (Guizhou Sci. 12(2): 54. 1994) is uncertain.

1a. Stipe straw-colored, surface smooth; sterile pinna margins flat and straight, teeth small and regularly angled; plants usually on calcareous substrates ....................................................  19a. var. cretica

1b. Stipe often brown, surface scabrous; sterile pinna margins undulate, teeth longer and at rather irregular angles; plants sometimes (?always) on more acidic substrates .................................  19b. var. laeta

19a. Pteris cretica var. cretica

欧洲凤尾蕨(原变种)  ou zhou feng wei jue (yuan bian zhong)

Pteris cretica var. intermedia (H. Christ) C. Christensen; P. cretica var. nervosa (Thunberg) Ching & S. H. Wu; P. nervosa Thunberg; P. pentaphylla Willdenow; P. serrulata Forsskεl var. intermedia H. Christ; P. sichuanensis H. S. Kuang; P. xichouensis W. M. Chu & Z. R. He.

Stipe straw-colored, surface smooth. Pinna base broadly cuneate, decurrent or not, sterile margins serrate and often rough and acute, apex acuminate and acutely serrate.

Among shrubs on limestone terrain; 400–3200 m. Anhui, Chongqing, Fujian (Nanping), Guangdong (Lianxian), Guangxi, Guizhou, SW Henan (Jigongshan, Neixiang, Zhenping), Hubei, Hunan, Jiangxi (Jianggangshan, Lushan), S Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, W Zhejiang (Shouchang) [Cambodia, India, Japan, Laos, Nepal, Philippines, Sri Lanka, Vietnam; Pacific islands (Fiji, Hawaii)].

Fraser-Jenkins (Indian Subcontinental Pteridophytes, 101. 2008) is followed in including Chinese material previously named as Pteris cretica var. nervosa (more correctly var. intermedia) within var. cretica.

Pteris xichouensis and P. sichuanensis are probably best regarded as somewhat depauperate forms of this species with mostly strictly 1-pinnate fronds with entire, unbranched pinnae (the basal pinnae occasionally forked in P. sichuanensis).

Pteris dangiana X. Y. Wang & P. S. Wang (Pterid. Fl. Guizhou, 581–582. 2001) appears to be another member of the P. cretica complex. It was likened to P. guangdongensis in the protologue, but that species has monomorphic fronds.

19b. Pteris cretica var. laeta (Wallich ex Ettingshausen) C. Christensen & Tardieu, Notul. Syst. (Paris) 6: 137. 1937.

粗糙凤尾蕨  cu cao feng wei jue

Pteris laeta Wallich ex Ettingshausen, Farnkr. Jetzw. 96. 1864 [“1865”]; P. cretica var. cartilagidens H. Christ; P. cretica subsp. laeta (Wallich ex Ettingshausen) Fraser-Jenkins; P. cretica var. rosthornii Diels; P. plumbea Wu, Wong & Pong (1932), not H. Christ (1909).

Stipe often brown, surface scabrous. Pinna base narrowly cuneate, decurrent, margins undulate, apex long acute.

Acidic soils of valleys; 900–2600 m. Fujian, Guangdong (Ruyuan), Guangxi, Guizhou (Kaili), Jiangxi (Huanggangshan, Tonggushan), Sichuan (Emeishan, Yaan), SE Xizang, Yunnan [Bhutan, Cambodia, N India, Nepal, Vietnam].

Pteris cretica var. laeta is taller and stronger than var. intermedia. The type is from Nepal. It is an apomictic triploid.

Pteris pellucidifolia Hayata, described from Taiwan (Icon. Pl. Formosan. 6, Suppl. (Gen. Ind. Fl. Formos.): 114. 1917), is most closely related to P. cretica but differs by having 6–8 pairs of lateral pinnae to 2 cm wide and with the lower 1–3 pairs branched. W. C. Shieh (Fl. Taiwan, ed. 2, 1: 231. 1994) indicates that it also occurs in “Indo-China” and SW China. Fraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes, 100. 2008) regarded P. pellucidifolia and the following species, P. nipponica, as synonyms of P. cretica var. laeta.

20. Pteris nipponica W. C. Shieh, Bot. Mag. (Tokyo) 79: 285. 1966.

日本凤尾蕨   ri ben feng wei jue

Evergreen fern of medium size. Rhizome shortly creeping. Fronds closely spaced; scales linear, brown, margins entire. Fronds subdimorphic. Sterile fronds: stipe glabrous, straw-colored, brownish at base, 10–30 cm, scaly; laminae pinnate or subpedate, basal pinnae sometimes with a basiscopic lobe or pinnule; lateral pinnae 1–3 pairs, linear-oblong, 10–20 Χ 1.5–3.5 cm, glabrous, margin irregularly dentate, apex acute; terminal pinnae distinct. Fertile fronds: stipe 14–50 cm; lateral pinnae linear, somewhat falcate, 20–30 Χ ca. 1.5 cm, shortly stalked; lamina green, thickly papery to chartaceous, sterile pinnules often with broad adaxial central white stripe. 3n = 87.

Moderate slopes, forest margins. Taiwan [Japan, S Korea].

Plants of this species were originally treated as a cultivar of Pteris cretica by Japanese botanists.

21. Pteris ryukyuensis Tagawa, Acta Phytotax. Geobot. 4: 204. 1935.

琉球鳳尾蕨   liu qiu feng wei jue

Evergreen fern of small to medium size. Rhizome short, ascending; scales small, narrow, blackish brown. Fronds dimorphic. Sterile fronds: stipes straw-colored to tinted brown, lower portion ± polished brown, dark at base, scaly; lateral pinnae commonly one pair, sometimes forked, 1–1.5 cm wide; apical pinnae twice as long as lateral ones, margin irregularly dentate, apex moderately acute. Fertile fronds twice as tall as sterile ones, ca. 30 cm tall; stipes much longer than laminae; lateral pinnae 1 or 2 pairs, lowest one sometimes forked, 3–5 mm wide. n = 58.

Along trails or on stone walls in village areas or on forest floor in lowlands; near sea level to 600 m. Taiwan [Japan, Philippines].

22. Pteris kidoi Kurata, J. Geobot. 13: 8. 1964.

城戶氏凤尾蕨   cheng hu shi feng wei jue

Small evergreen fern. Rhizome short, ascending; scales small, dark brown. Fronds dimorphic. Sterile fronds: stipe straw-colored, brown at base, 2.5–8 cm, with linear dark brownish scales; lamina digitate to imparipinnate; lateral pinnae (0–)1–2(–3) pairs, (4–)7–17 Χ 0.1–1.4 cm, margin dentate; terminal pinna 7–20 Χ 0.7–1.4 cm; false veinlets distinct and rather dense. Fertile fronds taller; stipe 8–22 cm; lamina 7–20 cm; lateral pinnae 1–3 pairs; lowest pinnae often forked, to 15 Χ 0.4–0.7 cm; terminal pinna linear, up to 22 Χ 0.2–0.5 cm; lamina with many false veins in mesophyll between true veins. 2n = 58.

Crevices in limestone rock faces; 300–900 m. Taiwan, ?Zhejiang [Japan].

23. Pteris esquirolii H. Christ, Notul. Syst. (Paris) 1: 50. 1909.

阔叶凤尾蕨  kuo ye feng wei jue

Plants robust, 1–1.5 m tall. Rhizome erect, short, robust, 2–3 cm in diam., woody, with black-brown scales. Fronds clustered, dimorphic; stipe dark straw-colored, sometimes reddish brown, shiny, 60–90 cm (stipes of sterile fronds shorter), 4–6 mm in diam., firm, glabrous, surfaces smooth or ± rough. Sterile fronds: lamina 1-pinnate, 25–30 Χ ca. 20 cm; rachis straw-colored, adaxially grooved; pinnae 3 or 4(–6) pairs, decumbent, subopposite, basal pairs 4–5 cm apart, basal pair each shortly stalked and with a basiscopic lobe, upper pinnae sessile, simple, lanceolate to oblong-lanceolate, 15–28 Χ 2–5 cm, base broadly cuneate, margins cartilaginous, entire except for dentate to serrate apex, apex acuminate or tapering. Fertile fronds: lamina 1-pinnate 40–50 Χ ca. 35 cm, ; pinnae 5 or 6 pairs, opposite, decumbent, basal 2 or 3 pairs 5–6 cm apart, often with basiscopic lobes, shortly stalked (4–8 mm), upper pinnae sessile, linear, up to 30 Χ 1–1.3 cm, base cuneate, margin entire, apex acuminate; terminal pinna 3-lobed base decurrent; midvein extremely convex adaxially, straw-colored, sometimes brownish; lamina pale green, thinly leathery when dried, glabrous, veins conspicuous on both sides, straight, parallel, subspreading (sterile pinnae slightly decumbent), simple or forked.

Among rocks in dense forests; 800–1500 m. Fujian, Guangdong (Lechang, Ruyuan, Xinyi), Guangxi (Longjin), Guizhou, N Hunan, Sichuan (Emeishan), Yunnan [N Vietnam].

1a. Stipe of fertile frond smooth ............................................................................  23a. var. esquirolii

1b. Stipe of fertile frond rough, distal part sparsely verrucose-tuberculate ............  23b. var. muricatula

23a. Pteris esquirolii var. esquirolii

阔叶凤尾蕨(原变种)  kuo ye feng wei jue (yuan bian zhong)

Stipe of fertile frond smooth.

Among rocks in dense forests; 800–1500 m. Fujian, Guangdong (Lechang, Ruyuan, Xinyi), Guangxi (Longjin), Guizhou, Sichuan (Emeishan), Yunnan [N Vietnam].

The type of Pteris esquirolii is from Guizhou.

23b. Pteris esquirolii var. muricatula (Ching) Ching & S. H. Wu, Fl. Reipubl. Popularis. Sin. 3(1): 31. 1990.

刺柄凤尾蕨  ci bing feng wei jue

Pteris muricatula Ching, Bull. Fan Mem. Inst. Biol. Bot. 10: 173. 1940.

Sterile fronds not seen. Stipe of fertile frond rough, distal part sparsely verrucose-tuberculate.

* N Hunan.

This plant is perhaps a different species, but since we did not see the sterile fronds, we are treating it here as a variety.

24. Pteris longipinna Hayata, J. Coll. Sci. Imp. Univ. Tokyo 30: 444. 1911.

长叶凤尾蕨  chang ye feng wei jue

Plants 60–90 cm tall. Sterile fronds: stipe dark brown like costae, 30–50 cm Χ ca. 3 mm, robust, glabrous; lamina 1-pinnate, ovate in outline, ca. 40 Χ 15 cm; lateral pinnae 3 or 4 pairs, opposite, basal pairs ca. 4 cm apart, extremely oblique, narrowly linear, 24–27 Χ ca. 1 cm, sessile, often slightly decurrent at base forming thin wings on basiscopic side, margin entire or slightly undulate, apex long acuminate (apical fertile parts 6–8 cm); basal 1 or 2 pinnae pairs pinnate; terminal pinnae with 2 or 3 pinnules, decurrent to lower pair of lateral pinnae at base; midvein raised abaxially, dark straw-colored or dark brown; veins approximate, spreading, simple or 2-forked.

* Rock crevices; 700–1500 m. Taiwan.

25. Pteris venusta Kunze, Bot. Zeitung. 6: 195. 1848.

爪哇凤尾蕨  zhao wa feng wei jue

Pteris matsudai Masamune.

Plants 50–80 cm tall. Rhizome erect, short, robust, 1.5–2 cm in diam., apex with brown scales. Fronds clustered; stipe straw-colored, seldom brown or castaneous, slightly lustrous, 30–50 cm Χ 3–4 mm, glabrous, basal part scaly, scales caducous; lamina 1-pinnate, oblong-ovate, 35–45 Χ 25–30 cm; pinnae (3 or)4–6 pairs, opposite, decumbent, linear-lanceolate, 15–20 Χ 2–3.2 cm, basally broadly cuneate, margin entire, slightly undulate, obviously reflexed, and cartilaginous, apex long caudate; basal pair shortly stalked, middle pairs sessile, upper pairs often connate with rachis and ± decurrent to form narrow wings; terminal pinna with 3 pinnules, not decurrent or slightly decurrent at base; basal pinnae not shortened, lowest 1 or 2 pairs often with basiscopic pinnule; lamina gray-green, thinly herbaceous when dried, glabrous, shiny especially adaxially, smooth; midvein abaxially convex, light straw-colored; veins slender, contiguous, slightly raised on both surfaces, often simple or rarely forked at base.

Acidic soils beneath open forests; 800–1500 m. ?SW Taiwan (Gaoxiong), Yunnan [Bhutan, Cambodia, India, Indonesia, Laos, Malaysia, N Myanmar, Nepal, Thailand, Vietnam].

According to W. C. Shieh (1994. in T. C. Huang (ed.), Fl. Taiwan, ed. 2, 1: 232), it is also distributed in SW Taiwan (Gaoxiong).

The type of Pteris venusta is from Indonesia (Java). There is apparent confusion between this species and P. pellucida in Taiwan (e.g., https://picasaweb.google.com/116136418529949606360/PterisPellucidaZhuaWaFengWeiJue, accessed 06 Jan 2012).

26. Pteris insignis Mettenius ex Kuhn, J. Bot. 1868: 269. 1868.

全缘凤尾蕨  quan yuan feng wei jue

Pteris indochinensis H. Christ; P. platysora Baker.

Plants 1–1.5 m tall. Rhizome ascending, robust, ca. 3 cm in diam., woody, apex with black-brown scales. Fronds clustered; stipe dark straw-colored and slightly lustrous, castaneous-brown near base, firm, 60–90 Χ 0.5–0.7 cm, sparsely scaly; scales brownish black, caducous; lamina 1-pinnate, ovate-oblong in outline, 50–80 Χ 20–30 cm; pinnae 6–14 pairs, opposite or sometimes subopposite, decumbent, linear-lanceolate, 16–20 cm, base cuneate, margin entire, slightly undulate, and cartilaginous, apex acuminate; lower pairs of pinnae sterile, ca. 2.5 cm wide, stalk ca. 1 cm; pinnae 4–6 cm apart, basal pair sometimes with a short lateral pinna; upper pinnae fertile, 1–1.5 cm wide, shortly stalked; veins conspicuous, midvein abaxially convex, dark straw-colored, veins oblique, prominent on both surfaces, sparse, simple or forked at base; lamina gray-green to brown-green, without luster, thinly papery when dried, glabrous; rachises light brown; indusia pallid or gray-brown, linear, entire.

Dense forests along valleys or streams; 200–800 m. Fujian, Guangdong, Guangxi, Guizhou (Dushan, Jiangkou), Hainan (Lingshui), Hunan (Jiangyong, Qianyang, Yizhang), Jiangxi, Yunnan (Honghe), S Zhejiang (Taishun) [Malaysia, Vietnam].

The type of Pteris insignis is from Guangdong.

27. Pteris menglaensis Ching, Acta Bot. Austro Sin. 1: 6. 1983.

勐腊凤尾蕨  meng la feng wei jue

Plants ca. 60 cm tall. Rhizome erect, short, apex densely clothed with brown scales. Stipe firm, straw-colored, ca. 27 cm Χ 2 mm, glabrous; lamina 1-pinnate, oblong in outline, up to 30 Χ ca. 15 cm; pinnae 4 or 5 pairs, subalternate, decumbent, linear-lanceolate, basal pair up to 17 Χ 1–1.3 cm, stalked (ca. 1 cm), base rounded-cuneate, margins entire and cartilaginous, apex acuminate; upper two pairs of pinnae with base cuneate, connate with rachis, not decrescent, stipitate; midvein prominent abaxially, straw-colored; veins dense, simple or 2-forked; lamina blue-green, papery when dried, abaxially sparsely hairy near midvein.

* Open forests; ca. 700 m. S Yunnan (Mengla).

28. Pteris vittata Linnaeus, Sp. Pl. 2: 1074. 1753.

蜈蚣草  wu gong cao

Pteris vittata f. cristata Ching.

Plants (20–)30–100(–150) cm tall. Rhizome erect, short and robust, 2–2.5 cm in diam., woody, apex densely clothed with yellow-brown scales. Fronds clustered; stipe firm, dark straw-colored or light brown, 10–30 cm or larger, 3–4 mm in diam., densely scaly when young, scales like those of rhizome, sparse; rachis straw-colored, sparsely scaly; lamina 1-pinnate, oblanceolate-oblong in outline, 20–90 Χ 5–25 cm or larger; lateral pinnae up to 40 pairs, alternate or sometimes subopposite; lower pinnae 3–4 cm apart, decumbent, sessile, not connate with rachises, progressively shorter toward base, basal pair auriculiform, middle pinnae longest, narrowly linear, 6–15 Χ 0.5–1 cm, base slightly expanded and cordate, both sides slightly auriculiform, upper ones larger and overlapping rachis; pinnae 1–1.5 cm apart, sterile margin minutely and evenly serrate, not cartilaginous, apex acuminate; midvein prominent abaxially and light straw-colored; veins slender, contiguous, oblique, simple or forked; terminal pinna similar to lateral pinnae in shape; lamina pale green, opaque, thinly leathery, glabrous.

Calcareous soils, on limestone, also on stone and on walls; below ca. 2000 m. Anhui, Fujian, SE Gansu (Kangxian), Guangdong, Guangxi, Guizhou, SW Henan, Hubei, Hunan, Jiangxi, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [widely distributed in tropics and subtropics of the Old World].

Plants of Pteris vittata are very different in shape and size, varying according to their habitats.

The type of Pteris vittata is from Guangdong.

29. Pteris ensiformis N. L. Burman, Fl. Ind. 230. 1768.

剑叶凤尾蕨  jian ye feng wei jue

Plants 30–50 cm tall. Rhizome ascending or prostrate, slender, 4–5 mm in diam., apex with black-brown scales. Fronds dense, dimorphic; sterile fronds 1.5–2(–3) cm apart, shorter than fertile fronds; stipe straw-colored as well as rachises, slightly lustrous, 10–30 cm (stipes of sterile fronds shorter), 1.5–2 mm in diam., glabrescent; lamina oblong-ovate, 10–25 Χ 5–15 cm, pinnate to bipinnate; pinnae 2–6 pairs, opposite, slightly decumbent, upper ones sessile, lower pairs shortly stalked; sterile fronds often pinnate, triangular in outline, 2.5–3.5(–8) Χ 1.5–2.5(–8) cm, acuminate; pinnules (1 or)2 or 3 pairs, opposite, contiguous, sessile, decumbent, oblong-oblanceolate to broadly lanceolate, basally decurrent and entire, upward and apices with acute teeth, apex obtuse; pinnae of fertile fronds distant (basal pairs 5–7 cm apart), 1–3-forked, middle fork longest, apical pinnae not decurrent at base, basal two pairs sometimes pinnate; pinnules 2 or 3(or 4) pairs, decumbent, narrowly linear, basally decurrent, margins entire except at apices, sterile parts with dense teeth, apically acuminate; midvein straw-colored, adaxially prominent; veins dense, often forked; lamina gray-green to brown-green, sometimes with nearly white bands along each side of midvein, herbaceous when dried, glabrous. n = 58.

Wet acidic soils beneath forests, stream margins; 100–1000 m. Chongqing, Fujian, Guangdong, Guangxi, SW Guizhou (Anlong), Hainan, S Jiangxi, Sichuan (Emeishan, Yaan), Taiwan, S Yunnan, S Zhejiang (Pingyang) [Bhutan, Cambodia, N India, Japan (Ryukyu Islands), Laos, Malaysia, Myanmar, Nepal, Sri Lanka, Thailand, Vietnam; Australia, Pacific islands (Fiji, Polynesia)].

The type of Pteris ensiformis is from Sri Lanka.

1a. .................................................................... Pinnae with a ± white band along each side of midvein .......................................................................................................................  29b. var. victoriae

1b. Pinnae midveins lacking off-white bands.

2a. Basal pinnae each with 2 or 3 pinnules, second and third (to fifth) pairs with 2 pinnules; median sterile lateral pinnae narrowly lanceolate, 4–6 Χ ca. 1 cm; fronds often 1-pinnate, or ± bipinnate at base ...........................................................................................................  29c. var. merrillii

2b. Basal pinnae pair or lateral pinnae often with 2–4 pinnules, lowest pair (or basal pairs) often 3- or 4-forked or subpinnate.

3a. Fertile pinnae apically not forked, (or 2- or 3-forked); fronds bipinnate, pinnae 3–6 pairs, 2.5–3.5(–8) cm .......................................................................................  29a. var. ensiformis

3b. Fertile pinnae apically 1- or 2-forked, pinnae 2–3 cm, spreading ............  29d. var. furcans

29a. Pteris ensiformis var. ensiformis

剑叶凤尾蕨(原变种)  jian ye feng wei jue (yuan bian zhong)

Phorolobus chinensis Desvaux; Pteris crenata Swartz; P. serrulata Forsskεl var. obtusata H. Christ; P. striata Poiret.

Pinnae bipinnate, 3–6 pairs, fertile pinnae often 2- or 3-forked.

Wet acidic soils beneath forests, stream margins; 100–1000 m. Chongqing, Fujian, Guangdong, Guangxi, SW Guizhou (Anlong), Hainan (Nanshanling, Sanya), S Jiangxi, Sichuan (Emeishan, Yaan), Taiwan, S Yunnan, S Zhejiang (Pingyang) [Cambodia, N India, Japan (Ryukyu Islands), Laos, Malaysia, Myanmar, Sri Lanka, Vietnam; Australia, Pacific islands (Fiji, Polynesia)].

The plants are indicators of acidic soils, with a pH of 4.5–5.0.

29b. Pteris ensiformis var. victoriae Baker, Gard. Chron., ser. 3, 7: 756. 1890.

白羽凤尾蕨  bai yu feng wei jue

Middle pinnae with pallid off-white bands on both sides of midvein.

Forests; ca. 300 m. Hainan (Nanshanling, Sanya) [N India, Malaysia, Myanmar, Sri Lanka].

29c. Pteris ensiformis var. merrillii (C. Christensen ex Ching) S. H. Wu, Fl. Reipubl. Popularis Sin. 3(1): 39. 1990.

少羽凤尾蕨  shao yu feng wei jue

Pteris merrillii C. Christensen ex Ching, Acta Phytotax. Sin. 9: 348. 1964 [“merilli”].

Fronds simple, often 1-pinnate or more divided, or bipinnate at base; pinnae 2–4 pairs, basal pair simple or 2- or 3-forked, sometimes pinnate; sterile fronds: margins of pinnae or pinnules serrate, apex of pinnae obtuse, acute, or acuminate; middle pinnae narrowly lanceolate, 4–6 Χ ca. 1 cm.

* Forests. Guangdong (Zhujiang Estuary and adjacent islands), Guangxi (Guiping, Lingui, Tengxian), Hainan, Taiwan (Taibei).

The type of Pteris ensiformis var. merrillii is from Hainan.

29d. Pteris ensiformis var. furcans Ching, Acta Bot. Austro Sin. 1: 6. 1983.

叉羽凤尾蕨  cha yu feng wei jue

Fertile pinnae apically 1- or 2-forked, segments 2–3 cm, spreading.

* Among rocks on slopes; ca. 400 m. Chongqing.

30. Pteris cryptogrammoides Ching, Acta Bot. Austro Sin. 1: 7. 1983.

珠叶凤尾蕨  zhu ye feng wei jue

Plants short, 10–15 cm tall. Rhizome erect, short, 3–4 mm in diam., apex with light brown scales. Fronds dimorphic; stipe light straw-colored, slender, 6–8 cm (stipes of sterile fronds shorter), less than 1 mm in diam., glabrous; sterile fronds slightly shorter than fertile fronds, pinnules similar in shape but wider, margins with acute teeth; fertile fronds: lamina 2- or 3-pinnate, triangular in outline, 5–10 Χ 3–5 cm; pinnae ca. 7 pairs, alternate or subopposite, basal pairs ca. 2.5 cm apart, spreading, bipinnate, narrowly ovate, ca. 4 Χ 2 cm, stalked (ca. 6 mm); pinnules ca. 3 pairs, slightly ascending; ultimate pinnules oblong-lanceolate, 0.7–0.9 Χ 0.2–0.25 cm, base broadly cuneate, margin entire, apex obtuse; upper pinnae simpler, sessile; midvein conspicuous, pale green, veins not conspicuous, simple or forked; lamina pale green, herbaceous when dried, glabrous; rachises and costae pale green and with narrow wings.

* Fujian (Xiamen).

31. Pteris multifida Poiret in Lamarck, Encycl. 5: 714. 1804.

井栏边草  jing lan bian cao

Pteris serrulata Linnaeus f., Suppl. Pl. 445. 1782, nom. illeg.

Plants 30–45 cm tall. Rhizome erect, short, 1–1.5 cm in diam., apex with black-brown scales. Fronds many, clustered, distinctly dimorphic. Sterile fronds: stipe straw-colored or dark brown with straw-colored margins, slightly lustrous, 15–25 cm Χ 1.5–2 mm, glabrous; lamina 1-pinnate, ovate-oblong in outline, 20–40 Χ 15–20 cm; pinnae often 3 pairs, opposite, ascending, linear-lanceolate, 8–15 Χ 0.6–1 cm, sessile, base acuminate, margins cartilaginous, with irregular acute teeth, basal pinna often forked, with 1 or 2 basal lobes, sometimes subpinnate, upper pinnae decurrent at base to form wings along rachis 3–5 mm wide, attenuate at base of rachis; terminal pinna 3-forked. Fertile fronds: stipe long, pinnae 4–6 pairs, linear, 10–15 Χ 0.4–0.7 cm, sterile margins serrate, remainder entire, basal pair subpinnate, with stipe ca. 1 cm, others sessile, basal 2 or 3 pairs often 2- or 3-forked, upper pairs with base long decurrent along rachis to form wings 3–4 mm wide; midvein prominent on both sides, straw-colored; veins conspicuous, sparse, simple or forked, sometimes both surfaces with short raised false veins parallel to veins; lamina pale green, herbaceous when dried, glabrous throughout; rachises straw-colored, slightly lustrous. 2n = 116.

On walls or on limestone; below ca. 1000 m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hebei (Beidaihe), Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi (Qinling), Shandong (Laoshan, Lushan, Taishan), Sichuan, Taiwan, Zhejiang [Japan (including Ryukyu Islands), S Korea, Philippines, Thailand, Vietnam].

The type of Pteris multifida is from a cultivated plant at the Botanical Garden in Paris.

32. Pteris liboensis P. S. Wang, Acta Bot. Yunnan. 9: 398. 1987.

荔波凤尾蕨  li bo feng wei jue

Plants 30–40 cm tall. Rhizome erect, short, scaly at growing tips; scales thick, black, lanceolate-subulate. Fronds clustered, apparently monomorphic; stipe straw-colored, thin, 18–22 cm Χ 1–1.5 mm, glabrous; lamina simple, less often 3-lobed or pinnate, linear-lanceolate, ca. 15 Χ 3–4 cm, base cordate, margins basally and apically sterile, cartilaginous with acute teeth, median part fertile and entire, apex ± acuminate; veins forked, subspreading, conspicuous on both surfaces; lamina green-brownish, thinly papery, both surfaces glabrous. Sori continuous, gray, firm, entire; spores triangular, with irregular tubercles.

* Forests; ca. 1000 m. Guangxi (Napo), S Guizhou (Libo).

33. Pteris nanlingensis R. H. Miau, Acta Sci. Nat. Univ. Sunyatseni 36: 114. 1997.

南岭凤尾蕨  nan ling feng wei jue

Plants perennial, up to 76 cm tall. Rhizome short, thick, decumbent, ca. 1 cm in diam. Fronds dimorphic, clustered. Sterile fronds: stipe straw-colored, light-brown at base, 20–24 cm Χ ca. 1 mm, glabrous, adaxially grooved; lamina 3-lobed, or apical pinnae forked into 2 pairs of pinnules, 15–20 cm; lateral pinnules similar to terminal pinnule but shorter, basal stalk ca. 2 mm, margins cartilaginous, slightly reflexed, with small acute, sparse teeth, entire near base, apex acuminate; terminal pinnule lanceolate, ca. 18 Χ 3.2 cm, basally slightly cuneate, basal stalk 2.4–2.6 cm. Fertile fronds pinnate, lateral pinnae 2 pairs; terminal pinna linear, 19.5–22 cm Χ 8–10 mm, basal stalk ca. 2 cm; lateral pinnules slightly short, stalked (5–8 mm); upper pair sessile, sterile margins with acute teeth; midvein straw-colored or light brownish, thinly grooved adaxially, raised abaxially; veins conspicuous, simple or forked; lamina gray-green, subleathery or leathery, glabrous. Sori linear, along with margin, only apices sterile; indusia gray-green, linear, membranous, subentire.

* Guangdong (Ruyuan).

34. Pteris changjiangensis X. L. Zheng & F. W. Xing, Ann. Bot. Fenn. 47: 156. 2010.

昌江凤尾蕨  chang jiang feng wei jue

Plants 20–30 cm tall. Rhizome ascending, tenuous, ca. 3 cm Χ 2–4 mm, densely scaly; scales dark brown, glossy, linear-lanceolate or lanceolate, 1–2 mm, entire. Fronds monomorphic. Stipe rufous-straw-colored in lower part, light green upward, 5–15 cm, base distinctly scaly, nitid, adaxially grooved; lamina simple, lanceolate, 10–20 Χ 0.8–1.2 cm, base attenuate, margins narrow, cartilaginous, subentire and repand at base, serrulate above, apex acuminate; costa straw-colored, abaxially prominent, smooth; veins conspicuous, straight, parallel, simple and bifurcate; lamina grass green when dried, thinly papery, glabrous. Sori brown, on margin of upper lamina except for apex; indusium light gray, narrow, membranous, entire.

* On surface of ground and rocks; 400–800 m. Hainan (Wangxia Village, Changjiang).

35. Pteris xiaoyingiae H. He & L. B. Zhang, Syst. Bot. 35: 696. 2010.

筱英凤尾蕨  xiao ying feng wei jue

Plants 4–6.5(–7) cm tall. Rhizome ascending, ca. 1.2 cm Χ 3.1–4 mm, apex sparsely scaly; scales subulate, dark brown. Fronds dispersedly distichous; stipe 2.5–4(–5.5) cm, 0.4–0.6 mm in diam. at middle, adaxially grooved, smooth and glabrous above base, sparsely scaly; lamina 3–5-foliolate, most often ± pedate in outline, 2–2.5 Χ 2.5–4 cm; pinnae 3, digitately arranged; terminal pinna largest, ovate-lanceolate, 1.5–2.5(–3.1) Χ 1–1.4 cm, sessile or occasionally stalked (ca. 4 mm), base cuneate and sometimes decurrent, apex blunt; lateral pinnae opposite, oblong-obovate, 1–1.5 Χ 0.7–1 cm, sessile or stalked (1–1.5 mm), base cuneate or basiscopically broadly cuneate, apex rounded or blunt; usually each lateral pinna with an almost free basal basiscopic lobe, lobes mostly oblong-obovate, apex rounded or blunt; margin of pinnae and lobes entire on cuneate bases, with teeth upward on sterile pinnae and 3- or 4-toothed on each side above sori on fertile pinnae; teeth deltoid or dentate, with cartilaginous tip but not aristate; lateral veins free, spreading or slightly ascending, distinct abaxially and merely conspicuous adaxially, simple or 1(or 2)-times 2-forked, almost reaching cartilaginous margins; lamina thickly papery to leathery in texture, both surfaces glabrous. Sori marginal on both sides of fertile pinna from 3–5 mm above base to 4–5 mm below apex, linear; indusia gray, ca. 0.8 mm wide, membranous, entire on margin.

* Limestone walls at openings of dry karst caves. S Guizhou (Libo).

2. Pteris sect. Quadriauricula Ching, Acta Bot. Austro Sin. 1: 2. 1983.

篦形凤尾蕨组  bing xing feng wei jue zu

Plants often medium-sized (0.2–2 m tall). Rhizome often short, sometime long, erect or decumbent. Fronds monomorphic, seldom dimorphic, lamina 2-pinnatipartite (often 3-pinnatipartite at base); pinnae/pinnules regularly pectinately divided nearly to costae, lower pinnae often with 1 or more pinnules near base on basiscopic side, these similar in shape to main part of pinnae but smaller; segments lanceolate or ± oblong-falcate, apices obtuse or acute, margins entire, rarely minutely toothed, usually not cartilaginous; costae with spines along groove (often ± with spines along midrib) or ± praemorse; veins free, 2-forked, or rarely 2-pinnate. Sori linear; indusia gray-brown or brown, linear, membranous, entire, persistent.

Mainly distributed in the tropics and subtropics; 30 species (ten endemic) in China.

36. Pteris dispar Kunze, Bot. Zeitung. 6: 539. 1848.

刺齿半边旗  ci chi ban bian qi

Pteris semipinnata Linnaeus var. dispar (Kunze) Hooker & Baker; P. taiwaniana Masamune & Suzuki.

Plants 30–80 cm tall. Rhizome ascending, 7–10 mm in diam., apices of rhizome and bases of stipes with brownish black scales, scale apices ciliate and slightly curly. Fronds clustered (10–15), subdimorphic; stipe chestnut-colored, shiny, 15–40 cm Χ ca. 2 mm, rachis similar; lamina 2-pinnatipartite or at one side deeply bipinnate lobed, ovate-oblong in outline, 25–40 Χ 15–20 cm; lateral pinnae 5–8 pairs, similar to terminal pinna in shape, opposite or subopposite, decumbent, basal pair 6–12 Χ 2.5–4 cm at base, shortly stalked, base oblique, divided nearly to costae on both or basiscopic side, apex caudate-acuminate; segments as in terminal pinna but basiscopic lobes slightly longer than acroscopic lobes, and lobe nearest base on basiscopic side longest, inclined, sometimes basal 1 or 2 segments developing into pinnule pectinately lobed on basiscopic side; terminal pinna pectinately partite nearly to costa, lanceolate in outline, 12–18 Χ 2–3 cm, base rounded, apically acuminate; segments 12–15 pairs, opposite, spreading, contiguous, broadly lanceolate or linear-lanceolate, somewhat falcate, 10–20 Χ 3–5 mm, base basiscopically not decurrent or slightly decurrent, margins of sterile fronds with long acute spinelike teeth, apices obtuse or sometimes acute; costae abaxially convex, chestnut at base; upper straw-colored, grooved adaxially, grooves with thin raised edges (minutely and irregularly toothed); veins conspicuous, decumbent, 2-forked, veinlets reaching cartilaginous point of teeth; lamina green or pale green, herbaceous when dried, glabrous. 2n = 58, 116.

Open forests along valleys; 300–1000 m. S Anhui (Qimen), Chongqing, Fujian, Guangdong, Guangxi (Lingui, Tengxian), Guizhou, Henan, Hubei (Jiugongshan), Hunan, S Jiangsu (Yixing), Jiangxi, Sichuan (Emeishan, Luxian), Taiwan, Zhejiang [Japan, Korea (Cheju Island), Malaysia, Philippines, ?Thailand, Vietnam].

The type of Pteris dispar is from Japan.

DeVol (Ferns East Centr. China, Notes Bot. Chin. Mus. Heude No. 7, 109. 1945) included material of Pteris dispar within his concept of P. semipinnata.

37. Pteris semipinnata Linnaeus, Sp. Pl. 2: 1076. 1753.

半边旗  ban bian qi

Plants 35–80(–120) cm tall. Rhizome long creeping, 1–1.5 cm in diam., apex with blackish brown scales; scales also at base of stipes. Fronds clustered, submonomorphic; stipe 15–55 Χ 1.5–3 cm, stipe and rachis castaneous-reddish, shiny, glabrous; lamina pinnate, oblong-lanceolate in outline, 15–40(–60) Χ 6–15(–18) cm, at one side deeply bipinnate-lobed; lateral pinnae 4–7 pairs, opposite or subalternate, spreading, basal pairs shortly stalked; upper ones sessile; lamina half-triangular and slightly falcate, 5–10(–18) Χ 4–7 cm, basiscopic side pectinately lobed nearly to costa, acroscopic side reduced to subentire wing to 6 mm wide, base oblique, basiscopic side decurrent, apex caudate-acuminate; basiscopic segments 3–6 or more, falcate-lanceolate, basal part longest, 1.5–4(–8.5) Χ 0.3–0.6(–1.1) cm, basally margins of sterile segments acutely serrate, fertile segments entire except for with one spine or 2 or 3 acute teeth near apex, apex mucronate or obtuse; terminal pinna pectinately divided almost to rachis, broadly lanceolate to narrowly triangular in outline, 10–18 Χ 3–10 cm, apex caudate; segments 6–12 pairs, opposite, spreading, 3–5 mm apart, falcate-lanceolate, 2.5–5 Χ 0.6–1 cm, upper ones gradually reduced, basiscopic obtriangular wing decrescent to next pair of pinnae along costule, apices shortly acuminate; costae prominent abaxially, chestnut adaxially, upper straw-colored, grooved adaxially, grooves with thin, minutely and irregularly toothed, raised edges. Veins conspicuous, decumbent, 2-forked or bipinnate forked, veinlets reaching base of teeth; lamina gray-green, herbaceous when dried, glabrous. 2n = 116.

 Acidic soil beneath open forests, by streams or rocks; below ca. 900 m. Fujian, Guangdong, Guangxi, S Guizhou (Ceheng, Sandou), Hunan, S Jiangxi (Anyuan, Xunwu), Sichuan (Leshan), Taiwan, S Yunnan [Bhutan, N India, Indonesia (Borneo), Japan (Ryukyu Islands), Laos, Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].

The type of Pteris semipinnata is from Guangdong.

Pteris semipinnata is different from P. dispar in shape and distribution. The former is primarily distributed in tropical areas, while P. dispar is primarily distributed in subtropical and northern tropical areas. Both are in Fujian and Taiwan, but any intersecting types have not been seen.

38. Pteris dissitifolia Baker, J. Bot. 28: 263. 1890.

疏羽半边旗  shu yu ban bian qi

Pteris semipinnata Linnaeus var. dissitifolia (Baker) C. Christensen & Tardieu.

Plants 1–1.5 m tall. Rhizome ascending or erect, 1–1.5 cm in diam., apex and stipes basally densely clothed with scales; scales median brownish black, light brown to gray at margins, without luster, linear-lanceolate, ca. 2 mm in diam., subentire, apex long acuminate. Fronds clustered; stipe castaneous-brown, upper side sorrel-red, sometimes nearly brownish, shiny, 40–80 cm Χ 4–5 mm, glabrous, adaxially grooved; rachises castaneous-brown; lamina 2-pinnatipartite on both sides or either side, ovate-oblong in outline, 35–50 Χ 25–30 cm; lateral pinnae 5–8 pairs, opposite or subopposite, basal pairs 6–8 cm apart, stalked (1–3 mm), upper sessile, decumbent, lamina oblong-lanceolate, 15–25 Χ 4–6 cm, base oblique, apex tapering; basal 1 or 2 pairs of pinnae often pinnatipartite on both sides; basal pair of pinnae often with a pinnule near base on basiscopic side; median pinnae often pinnatipartite on basiscopic side, entire on acroscopic side or only pinnatipartite on upper parts; upper pinnae pinnatipartite on basiscopic side, entire on acroscopic side; segments lanceolate, 4–12 Χ 1–1.3 cm, gradually shorter upward, acuminate; basal segments on basiscopic longest, basally decrescent to a broad wing, sterile margins slightly serrate; terminal pinna pinnatipartite, divided nearly to costa, triangular-ovate in outline, ca. 15 Χ 10–12 cm, base broadly cuneate, somewhat decurrent, apex acuminate; segments 8–10 pairs, alternate or subopposite, ca. 1 cm apart, decumbent, falcate-lanceolate, 6–8 Χ 0.6–0.8 cm, gradually shorter upward, base conspicuously decrescent, with interrupted broad wing along costules, apex mucronate; costae prominent abaxially, straw-colored at apex, castaneous-brown downward, adaxially grooved, grooves with thin raised pale gray edges (minutely and irregularly toothed); veins conspicuous on both sides, sparse, decumbent, often 1- or 2-forked; lamina pale green, firmly herbaceous when dried, glabrous.

Sparse, shaded forests; 100–300 m. Guangdong, Hainan (Baoting), S Yunnan (Hekou, Xishuangbanna) [Laos, N Vietnam].

The type of Pteris dissitifolia is from N Vietnam.

39. Pteris malipoensis Ching, Acta Bot. Austro Sin. 1: 7. 1983.

大羽半边旗  da yu ban bian qi

Plants up to 2 m tall. Rhizome prostrate, firm, ca. 1 cm in diam., woody. Stipe castaneous-brown at base, opaque, ca. 100 cm Χ 4 mm, firm, glabrous; lamina 2-pinnatipartite on either side, oblong-ovate; pinnae alternate, decumbent, sessile, basal pairs ca. 11 cm apart, falcate, narrowly triangular, ca. 40 Χ 15 cm, base oblique, apex long caudate; pinnae entire on acroscopic margin and apical 1/4–1/3 of basiscopic margin, deeply lobed on rest of basiscopic margin, with 6–8 gradually shorter segments; segments linear-lanceolate, basal ones ca. 13 Χ 1.8 cm, up to 2 cm apart, base oblique, sterile margins serrate, apex acuminate; veins distinct on both surfaces, spreading, sparse, 1- or 2-forked; lamina pale green, herbaceous when dried, glabrous; costae and rachises straw-colored, grooved on adaxial side, with few spines on raised edges.

* Mixed forests; 1200–1500 m. Hunan (Tongdao), SE Yunnan (Malipo).

Pteris malipoensis is represented by larger plants, rare, and similar to P. semipinnata but the former has stipes robust, straw-colored, basal pinnae up to 40 cm, segments at base ca. 13 Χ 1.8 cm, pale green when dried.

40. Pteris excelsa Gaudichaud, Voy. Uranie 388. 1827.

溪边凤尾蕨  xi bian feng wei jue

Pteris excelsissima Hayata; P. inaequalis Baker var. aequata (Miquel) Tagawa; P. kleiniana H. Christ (1896), non Campteris kleiniana Presl (1836); P. longipinnula Franchet & Savatier (1876), not Wallich ex J. Agardh (1839); P. semipinnata Linnaeus var. aequata Miquel.

Plants up to 180 cm tall. Rhizome erect, short, robust, to 2 cm in diam., woody, apex with black-brown scales. Fronds clustered; stipe dark brown, upper side straw-colored, slightly lustrous, firm, robust, 70–90 cm Χ 6–10 mm, glabrous; rachis straw-colored, adaxially grooved; lamina 2-pinnatipartite, broadly triangular, 60–120 or more Χ 40–90 cm; lateral pinnae 5–10 pairs, alternate or subopposite, basal pairs 10–15 cm apart, shortly stalked, spreading, similar to terminal pinna, basal pair largest, more than 40 cm, sometimes forked at base, upper pinnae smaller, sessile; terminal pinna pectinately divided to nearly to costa, broadly lanceolate-oblong in outline, 20–30 cm or larger Χ 7–12 cm, attenuate toward apex, apex acuminate and caudate; segments 20–25 pairs, alternate, subspreading, falcate-lanceolate, 3.5–8(–10) Χ 0.6–1 cm, base slightly expanded, basiscopic side decurrent, sterile margins of apical fronds slightly serrate, apex acuminate; costae abaxially convex, straw-colored, glabrous, adaxially shallowly grooved, with thick spines along groove; veins abaxially conspicuous, sparse, decumbent, often 2-forked; lamina often pale green, herbaceous when dried, glabrous, base of pinnae rarely pubescent abaxially. 2n = 58, 87.

Open forests, scrub; 600–2700 m. Chongqing, Guangdong (Lechang, Yingde), Guangxi (Guilin, Lingui, Xiuren), Guizhou (Yinjiang, Zunyi), Hubei (Badong), Hunan (Qianyang, Yongshun), Jiangxi (Lushan, Xiushui), Sichuan, Taiwan, Xizang (Chayu, Cuona), Yunnan, Zhejiang [N India, Japan (Honshu, Kyushu, Shikoku), S Korea, Laos, Malaysia, Nepal, Pakistan, Philippines, Vietnam; Pacific islands (Fiji, Hawaii)].

The type of Pteris excelsa is from Hawaii.

41. Pteris inaequalis Baker, J. Bot. 13: 199. 1875.

中华凤尾蕨  zhong hua feng wei jue

Pteris excelsa Gaudichaud var. inaequalis (Baker) S. H. Wu; P. excelsa var. simplicior (Tagawa) W. C. Shieh; P. inaequalis var. simplicior Tagawa; P. sinensis Ching.

Plants 40–100 cm tall. Rhizome prostrate, apex with brown, lanceolate scales. Fronds monomorphic; stipe straw-colored or brownish straw-colored, 26–46 cm; lamina 2-pinnatipartite, ovate to oblong-lanceolate in outline, 34–42 Χ 10–32 cm, base rounded or rounded-cuneate, apex caudate; lateral pinnae 3–6 pairs, opposite, oblique, sessile or subsessile, basiscopically pectinately lobed, acroscopically entire or few lobed, narrowly triangular in outline, falcate and ascending, apex long caudate; basal pair of pinnae 10–22 Χ 5–11 cm, basiscopic 2–7 segments falcate-lanceolate, basal one largest, 3–10 Χ 0.7–1.5 cm, sterile margins dentate, apex obtuse or tapering; upper pinna sometimes entire, linear; terminal pinna larger, triangular-ovate, pectinately divided; lamina herbaceous, glabrous; grooves of rachis and costa with short spines; midvein abaxially protuberant; veins abaxially conspicuous, adaxially inconspicuous; lateral veins oblique, forked or single.

 Forests, stream margins, limestone caves; 400–1400 m. Fujian (Chongan), Guangdong (Lechang), Guangxi (Longsheng, Xiuren), Guizhou, Jiangxi (Jinggangshan, Tonggu), Sichuan (Dechang, Emeishan, Hechuan), Yunnan (Kunming, Mengzi), Zhejiang (Longquan) [India, Japan].

The type of Pteris inaequalis is from Yunnan (Mengzi). Pteris sinensis was based on Baker’s taxon P. inaequalis Fιe (1857), which seems to be a nomen nudum, though perhaps it was just a listing of P. inaequalis C. Presl (nom. nud. 1836), in which case P. inaequalis Baker is valid and the correct name for this taxon.

42. Pteris amoena Blume, Enum. Pl. Javae 210. 1828.

红秆凤尾蕨  hong gan feng wei jue

Pteris tokioi Masamune.

Plants 80–150 cm tall. Rhizome 1–1.5 cm in diam. Stipe basally castaneous-brown, upper part, rachis, and lower parts of costae sorrel-red, shiny, stipe 45–80 cm, 3–5 mm in diam., glabrous; lamina 2- or 3-pinnatipartite, ovate in outline, 35–65 Χ 30–40 cm; lateral pinnae 4–8 pairs, subopposite, decumbent, basal pair sometimes 2-pinnatipartite, shortly stalked; upper pinnae sessile, pectinately divided, lanceolate in outline, 25–30 Χ ca. 5 cm, base rounded-cuneate and slightly oblique, apex acute, caudate for 3–5 cm; terminal pinna similar but wider and stalked (3–4 cm), basal pair of pinnae often with a basiscopic pinnule near base, similar in shape to main part of pinna but smaller; segments 25–30 pairs, alternate, 3–4 mm apart, sinuses wide and rounded, slightly decumbent, subfalcate, lanceolate, 2.5–4 Χ 0.5–0.7 cm, base slightly expanded and decurrent, sterile margins densely serrulate, apex obtuse or mucronate, basal segments of pinnae slightly shorter and more widely spaced, basally decrescent to costules; costae prominent abaxially, adaxially straw-colored, shiny, glabrous, adaxially grooved, with spines along groove and bases of midribs; veins conspicuous, oblique, apical pairs of veins simple, remainder 2-forked, basal basiscopic ones from costae, others arising from midribs.

Hainan (Changjiang), Taiwan, SE Xizang (Mκdog), Yunnan (Guangnan, Xishuangbanna), Zhejiang (Cangnan) [India, Indonesia, Myanmar].

The type of Pteris amoena is from Indonesia (Java).

43. Pteris formosana Baker, J. Bot. 23: 103. 1885.

美丽凤尾蕨  mei li feng wei jue

Pteris decurrentipinnulata Bonaparte; P. takeoi Hayata.

Plants 70–100 cm tall. Rhizome prostrate, ca. 1 cm in diam.; scales yellowish brown to pale brown, linear to lanceolate, to 1.5 cm, entire. Fronds clustered; stipe reddish brown, shiny, 30–70 cm Χ 3–4 mm, glabrous; rachis castaneous, shiny, grooved adaxially; lamina 2-pinnatipartite, ovate-triangular in outline, 45–60(–100) Χ ca. 30 cm; lower lateral pinnae 5 or 6 pairs, opposite, 5–7 cm apart, pinnae 1(–2)-pinnatisect, decumbent, oblong-lanceolate in outline, 12–18 Χ 5–7 cm, shortly stalked, base decrescent, apex narrowly lanceolate-caudate; pinnae anisomerous: basiscopic side with 0–3 segments toward its apex, acroscopic side with 3–6 pectinately arranged segments; segments lanceolate, somewhat falcate, 35–60 Χ ca. 6 mm, base slightly narrowed, basiscopically decurrent, margins entire, apex long acute to acuminate, ca. 4 mm apart, decumbent; costae grooved adaxially, grooves with pallid thin raised edges; veins oblique, forked; distal lateral pinnae alternate, 3–5 each side, decumbent, lanceolate, 8–10 Χ 0.8–1 cm, sessile; terminal pinna pinnatifid, ca. 16 Χ 6 cm; lamina pale green, firmly herbaceous when dried, glabrous. 2n = 58.

Wet and shaded cliffs, often along streams or waterfalls; 600–2500 m. Taiwan [Japan (Ryukyu Islands)].

The type of Pteris formosana is from Taiwan.

44. Pteris cadieri H. Christ, J. Bot. 19: 72. 1905.

条纹凤尾蕨  tiao wen feng wei jue

Plants 30–40 cm tall. Rhizome erect, short, 0.5–1 cm in diam., apex with black-brown scales. Fronds clustered (10–20), distinctly dimorphic; stipe castaneous-brown, upward straw-colored, or sometimes dark straw-colored, shiny, 15–25 cm; rachis castaneous-brown, shiny, adaxially grooved with pale straw-colored, narrowly winged margins; sterile fronds: stipe shorter, ca. 1 mm in diam., glabrous, apex with thin narrow green wings; laminae 2-pinnatipartite, 3-forked, ovate-triangular in outline, 10–15 Χ 6–8 cm; lateral pinnae 1 pair, next to apical pinnae, falcate-triangular, 4–6 Χ 3–4 cm, both sides or basiscopically with 3–5 pectinate lobes (sometimes at base basiscopically only with one lobe), basal basiscopic one largest, simple or ± pectinate; terminal pinna largest, broadly lanceolate, 10–15 Χ 3–4 cm, sessile, base cuneate, middle sometimes with 4–8 pairs of pectinately arranged segments, margins acutely dentate, apex acuminate; segments oblong, somewhat falcate, 10–15 Χ 6–8 mm, contiguous, apex obtuse; fertile fronds 1–2-pinnate or sometimes subdigitate, pinnae linear, margins entire or slightly undulate, sterile margins serrate, apex acuminate; midvein abaxially convex, shallowly grooved and with spines, pale straw-colored; veins conspicuous on both surfaces, oblique, often forked, with irregular short raised false veins between veins. Lamina pale green, herbaceous when dried, glabrous. 3n = 87.

Forests, on wet rocks near creeks; 200–700 m. Fujian (Minhou, Nanjing), Guangdong, Guangxi, NE and SE Guizhou, Hainan, Jiangxi, Taiwan [Japan (Ryukyu Islands), N Vietnam].

The type of Pteris cadieri is from N Vietnam.

1a. Fertile fronds 1-pinnate or sometimes subdigitate, basally decrescent to costae bearing narrow wings; pinna margins entire or slightly undulate, rarely with a few lobes, lateral pinnae 1 or 2 pairs, opposite, spreading, sessile, same as apical pinnae in shape but shorter, basal basiscopically decurrent, basal pair often 2-forked ....................................................................................................  44a. var. cadieri

1b. Fertile fronds 1- or 2-pinnate; pinna margins rounded-undulate, or with lobes along rachis or pinnatipartite ..............................................................................................  44b. var. hainanensis

44a. Pteris cadieri var. cadieri

条纹凤尾蕨(原变种)  tiao wen feng wei jue (yuan bian zhong)

Pteris dimorpha Copeland; P. grevilleana Wallich var. diffusa Wu Wong & Pong.

Fertile fronds: lamina 1-pinnate or sometimes subdigitate, basally decrescent to costae bearing narrow wings; pinnae linear, 8–12 Χ 0.6–1 cm, margins entire or slightly undulate, rarely with a few lobes, apex acuminate; lateral pinnae 1 or 2 pairs, opposite, spreading, sessile, same as terminal pinna in shape but shorter, basally basiscopic side decurrent, basal pair often 2-forked.

Forests, near creeks, among wet rocks; 200–500 m. Fujian (Minhou, Nanjing), Guangdong, Guangxi, NE and SE Guizhou, Taiwan [Japan (Ryukyu Islands), N Vietnam].

44b. Pteris cadieri var. hainanensis (Ching) S. H. Wu, Fl. Reipubl. Popularis Sin. 3(1): 54. 1990.

海南凤尾蕨  hai nan feng wei jue

Pteris hainanensis Ching, Acta Phytotax. Sin. 8: 142–3. 1959.

Fertile fronds: lamina 1- or 2-pinnate; pinnae rounded-undulate at margin, or with lobes along rachis, or pinnatipartite.

* Forests; 300–700 m. Guangdong (Gaoyao, Wengyuan), Guangxi (Longjin, Yaoshan), Hainan (Baoting, Ledong).

The type of Pteris cadieri var. hainanensis is from Hainan.

Pteris grevilleana sensu Wu, Wang & Pong (Bull. Dept. Biol. Sun Yatsen Univ. 3: 234. 1932), not Wallich ex J. Agardh (1839).

45. Pteris grevilleana Wallich ex J. Agardh, Recens. Spec. Pter. 23. 1839.

林下凤尾蕨  lin xia feng wei jue

Plants 20–45 cm tall. Rhizome erect, short, 0.5–1 cm in diam., apex with black-brown scales. Fronds clustered (10–15 per plant), monomorphic; stipes of fertile fronds 2 Χ as long as those of sterile fronds, castaneous-brown, shiny, 20–30 cm Χ 1–1.5 mm, glabrous, with narrow wings apically; rachis similar, adaxially grooved and with narrow wings; lamina 2-pinnatipartite, broadly ovate-triangular, 10–15 Χ 8–12 cm; lateral pinnae 1 or 2 pairs, opposite, decumbent, sessile, in shape of apical lateral pinnae but shorter, base decrescent to narrow wings; basal pinnae 2-forked, basiscopic pinnule smaller; fertile pinnae similar to sterile pinnae, but segments shorter and fewer; terminal pinna broadly lanceolate, 8–12 Χ 2.5–3.5 cm, base decurrent, connate below pinnae, pectinately divided nearly to costae on both sides, apex caudate; segments decumbent, contiguous, falcate-linear to lanceolate, 15–20 Χ 3–4 mm, margins serrulate, apex obtuse; lower segments shorter; costae adaxially convex, basally castaneous-brown, upper straw-colored, shallowly grooved, with acute spines (ca. 0.8 mm) on both sides; veins inconspicuous, parallel, very decumbent, often 2-forked, occasionally 3-forked; lamina pale green, firmly herbaceous when dried, glabrous, both surfaces with short raised false veins between veins.

Among rocks within forests; 100–900 m. Guangdong (Foshan, Gaoyao, Xuwen), Guangxi, Hainan, Taiwan, Yunnan (Hekou, Xishuangbanna) [Bhutan, N India, Indonesia, Japan (Ryukyu Islands), Malaysia, Nepal, Philippines, Thailand, Vietnam].

The type of Pteris grevilleana is from NW India.

1a. Pinnae same color throughout ......................................................................  45a. var. grevilleana

1b. Pinnae with parallel pallid spots along both sides of costa, spots ca. 1 cm wide ....  45b. var. ornata

45a. Pteris grevilleana var. grevilleana

林下凤尾蕨(原变种)  lin xia feng wei jue (yuan bian zhong)

Pteris quadriaurita Retzius var. digitata Baker.

Pinnae same color throughout.

Among rocks within forests; 100–900 m. Guangdong (Foshan, Gaoyao, Xuwen), Guangxi, Hainan, Taiwan, Yunnan (Hekou, Xishuangbanna) [Bhutan, N India, Indonesia, Japan (Ryukyu Islands), Malaysia, Nepal, Philippines, Thailand, Vietnam].

45b. Pteris grevilleana var. ornata Alderwerelt, Malayan Ferns 364. 1908.

白斑凤尾蕨  bai ban feng wei jue

Pinnae with parallel pallid spots along both sides of costa adaxially, spots ca. 1 cm wide.

Forests. Guangdong (Gaoyao), Guangxi (Wuming). [N India, Indonesia, Malaysia, Myanmar, Philippines, Thailand, Vietnam].

Pteris grevilleana var. ornata is a beautiful ornamental plant.

46. Pteris heteromorpha Fιe, Gen. Fil. 127. 1852.

长尾凤尾蕨  chang wei feng wei jue

Pteris cretica Linnaeus var. heteromorpha Beddome.

Plants 60–80 cm tall. Rhizome ascending, ca. 1 cm in diam., apex with brown scales. Fronds clustered (8–10 fronds); stipe dark straw-colored to brown, slightly lustrous, 25–45 cm, 2–3 mm in diam., glabrous, ± rough; rachis similar, scabrous, with sparse short hairs; lamina 2-pinnatipartite, oblong-ovate in outline, 30–40 Χ 20–25 cm; terminal pinna with linear-caudate tails, 12–15 Χ 0.8–1 cm, slightly decurrent at base, pectinately divided nearly to costa on both sides, sometimes not divided, apex acuminate; segments 0–8 pairs, alternate or subopposite, contiguous and slightly imbricate, subspreading, linear-lanceolate, 2.5–3 cm, entire and with cartilaginous edges; lateral pinnae 4–6 pairs, opposite, decumbent, stalked (4–6 cm), similar to terminal pinna, distal 1 or 2 pairs of pinnae shorter, sessile, linear, long caudate, oblique, on both sides segments 1–15 pairs, or at basiscopic side only 1 or 2 segments and other side without segment; costae prominent abaxially, dark straw-colored or castaneous, surfaces ± rough, with shorter hairs, deeply grooved adaxially, with spines along grooves; veins conspicuous on both surfaces, oblique, 2-forked; lamina gray-green, firmly herbaceous when dried, abaxially with sparse multicellular gray short hairs.

Forests; 900–1200 m. S Yunnan (Jinghong, Simao) [Malaysia, Myanmar, Philippines, Vietnam].

The type of Pteris heteromorpha is from the Philippines (Luzon).

47. Pteris decrescens H. Christ, Bull. Acad. Int. Gιogr. Bot. 16: 244. 1906.

多羽凤尾蕨  duo yu feng wei jue

Plants 60–70 cm tall. Rhizome erect, short, 1–2 cm in diam., apex with black-brown scales. Fronds clustered (8–12 per plant); stipe dark straw-colored, slightly lustrous, 15–20 cm Χ ca. 2 mm, ± rough, with sparse gray bristles when young, bristles deciduous; rachis similar to stipe; lamina 2-pinnatipartite, oblong or broadly lanceolate in outline, 30–50 Χ 12–15 cm, attenuate toward base and apex; lateral pinnae 10–15 pairs, opposite, ca. 5 cm apart at base, oblique, slightly shortly stalked, upper sessile; terminal pinna lanceolate, 7–10 Χ 2–2.5 cm, wider at base, shortly stalked, base broadly cuneate, upward attenuate, apex narrowly linear-caudate, 1.5–4 cm; segments 15–20 pairs, alternate or subopposite, subspreading, 1–2 mm apart, linear, 5–12 Χ ca. 2.5 mm, slightly expanded at base, margins entire and with cartilaginous edge, apex obtuse and with 2–7(–12) small teeth; costae abaxially convex, straw-colored, with sparse multicellular gray bristles, adaxially deeply grooved, with narrow ridges on both sides along groove, with a spine at bifurcation of costa of segments; veins conspicuous and prominent on both surfaces, contiguous, oblique, 2-forked; lamina brown-green to pale green, herbaceous when dried.

Evergreen forests; 700–1200 m. Guangdong (Yangchun), Guangxi, Guizhou, Yunnan [Cambodia, Thailand, Vietnam].

The type of Pteris decrescens is from Guizhou.

1a. Lateral pinnae 10–15 pairs, caudate apex 1.5–2(–3) cm, basal pinnae gradually shorter, 5–7 cm, middle 8–10 Χ 2–3 cm, slightly shorter upward ........................................................  47a. var. decrescens

1b. Lateral pinnae 12–14 pairs, ca. 7 Χ 1.3 cm, caudate apex 3–4 cm, basal pair almost not shortened  47b. var. parviloba

47a. Pteris decrescens var. decrescens

多羽凤尾蕨(原变种)  duo yu feng wei jue (yuan bian zhong)

Lateral pinnae 10–15 pairs, oblique, slightly shortly stalked, upper ones sessile, rounded-truncate at base, caudate apex 1.5–2(–3) cm; basal pinnae gradually shorter, 5–7 cm, wide, middle 8–10 Χ 2–3 cm, slightly shorter upward.

Evergreen forests; 700–1200 m. Guangdong (Yangchun), Guangxi, Guizhou, Yunnan [Cambodia, Vietnam].

47b. Pteris decrescens var. parviloba (Christ) C. Christensen & Tardieu, Notul. Syst. (Paris) 6: 137. 1937.

大明凤尾蕨  da ming feng wei jue

Pteris parviloba Christ, Bull. Gιogr. Bot. 1907: 149. 1907; P. quadriaurita Retzius var. parviloba Christ.

Lateral pinnae 12–14 pairs, ca. 7 Χ 1.3 cm, narrowly linear, caudate apex 3–4 cm; basal pair almost not shortened.

Guangxi (Damingshan) [N Vietnam].

The type of Pteris decrescens var. parviloba is from N Vietnam.

48. Pteris longipinnula Wallich ex J. Agardh, Rec. Sp. Gen. Pteris. 19. 1839.

翠绿凤尾蕨  cui lό feng wei jue

Plants 0.5–1.2 m tall. Rhizome erect, short, 1–2 cm in diam., apex with black-brown scales. Fronds clustered; stipe brown, 30–70 cm, ca. 1 mm in diam., glabrous or with bristles, adaxially grooved; lamina 2-pinnatipartite, ovate, 30–60 Χ 20–25 cm; lateral pinnae 4 or 5 pairs, opposite, adjacent to each other, decumbent, basal pair shortly stalked, upper sessile, broadly lanceolate, 20–25 Χ ca. 5 cm, base narrower and shortly cuneate, deeply pectinately pinnatifid forming narrow wings, apex acuminate; terminal pinna similar but wider and with stalk 2–6 cm; segments 20–30 pairs, opposite or alternate, decumbent, incised acute, 2–3 mm apart, 20–40 Χ 5–8 mm, base slightly expanded, margin entire and with cartilaginous border, apex obtuse and with a sharp point; costae with short spines along groove; veins biconvex, oblique, forked at base; lamina pale green, firmly herbaceous when dried, glabrous.

Hainan (Diaoluoshan), Yunnan [Bhutan, NE India, Myanmar, Nepal].

Pteris longipinnula is slightly similar to P. decrescens, but differs because is slightly larger, with lateral pinnae fewer (only 4 or 5 pairs), and much longer (to 25 cm). The segments are longer, and not serrate. It is also similar to Pteris heteromorpha, but differs in having the stipe straw-colored, and costae abaxially hairy.

1a. Plants 1–1.2 m tall; stipe, rachis, costae, and costules glabrous ...................  48a. var. longipinnula

1b. Plants 0.5–0.8 m tall; stipe, rachis, costae, and costules with bristles ....................  48b. var. hirtula

48a. Pteris longipinnula var. longipinnula

翠绿凤尾蕨(原变种)  cui lό feng wei jue (yuan bian zhong)

Pteris umbraculifera Mettenius.

Plants 1–1.2 m tall. Stipe, rachis, costae, and costules glabrous. Stipe 60–70 cm, upper part of stipe and rachis, costae, and midvein blue-green; basal segments abruptly shortened, narrowly triangular, ca. 6 mm.

* Hainan (Diaoluoshan), S Yunnan (Hekou, Xishuangbanna).

48b. Pteris longipinnula var. hirtula C. Christensen, Contr. U.S. Natl. Herb. 26: 312. 1931.

毛叶凤尾蕨  mao ye feng wei jue

Pteris hekouensis Ching.

Plants 50–80 cm tall. Stipes 30–50 cm, adaxial groove with gray short bristles; costae adaxially prominent, costae and costules with gray sparse bristles. Rachis with gray multicellular bristles.

Evergreen rain forests; 100–200 m. SE Yunnan (Hekou) [Myanmar].

49. Pteris aspericaulis Wallich ex J. Agardh, Recens. Spec. Pter. 22. 1839.

紫轴凤尾蕨  zi zhou feng wei jue

Plants 0.3–1.5 m tall. Rhizome ascending, short, 1.5–2 cm in diam., apex with black-brown scales. Fronds clustered; stipe as long as lamina, glabrous, ± rough, with scattered pustules, scaly at base; lamina 2- or 3-pinnatipartite, oblong-ovate in outline, (18–)50–80 Χ (3–)20–25 cm; lateral pinnae (1–)6–14 pairs, opposite, decumbent, lanceolate, 2–18(–23) cm, sessile or basiscopically shortly stalked, base cuneate, slightly oblique, pectinate divided to near costae, apex caudate with linear pointed tail, basal pair of pinnae often with 1(or 2) basiscopic pinnules near base similar in shape to main part of pinnae but smaller; segments (17–)22–25(–32) pairs, alternate, contiguous, subspreading, oblong, somewhat falcate, 10–30 Χ 3.5–7 mm, base slightly expanded, entire or sterile margins occasionally slightly undulate, apex mucronate or obtuse, and with a sharp corner; terminal pinna similar to median lateral pinnae, stalked (1–1.5 cm); costae prominent abaxially, glabrous, grooved adaxially, with spines along groove; veins conspicuous on both sides, decumbent, 2-forked from base, lateral veins of segment arising from rachis, 2 opposite veins of base reaching incision of pinnae; lamina gray-green, often mauve at margin, subleathery when dried, glabrous (abaxially glabrous, adaxially subglabrous when young).

Forests; 800–2900 m. Guangxi (Baise), Sichuan (Shimian), Xizang (Chayu, Nielamu), Yunnan [S Himalaya: Bhutan, N India, Kashmir, Nepal].

1a. Stipe, rachis, costae, lamina margin, and indusia ± purple or mauve.

2a. Plants 1–1.5 m tall ................................................................................  49a. var. aspericaulis

2b. Plants 0.3–0.5 m tall ..............................................................................  49c. var. subindivisa

1b. Stipe and rachis brown or straw-colored, lamina margin and indusia green.

3a. Pinnae uniformly green .............................................................................  49b. var. cuspigera

3b. Pinnae with wide white or rose-colored band along sides of costa ................  49d. var. tricolor

49a. Pteris aspericaulis var. aspericaulis

紫轴凤尾蕨(原变种)  zi zhou feng wei jue (yuan bian zhong)

Pteris roseo-lilacina Hieronymus; P. tibetica Ching.

Plants 1–1.5 m tall. Stipe as long as lamina, 4–5 cm in diam.; stipe, rachis, costae, lamina margin, and indusia ± mauve, sometimes straw-colored; lamina oblong-ovate, 50–80 Χ 20–25 cm; lateral pinnae 6–14 pairs, opposite, decumbent, sessile or basiscopically shortly stalked, lanceolate, (6–)15–18(–23) Χ (1.5–)2.5–3(–5) cm.

Forests; 800–2900 m. Guangxi (Baise), Sichuan (Shimian), Xizang (Chayu, Nielamu), Yunnan [Bhutan, N India, Nepal].

49b. Pteris aspericaulis var. cuspigera Ching, Acta Bot. Austro Sin. 1: 8. 1983.

高原凤尾蕨  gao yuan feng wei jue

Pteris shimianensis H. S. Kuang.

Plants 1–1.5 m tall, not purple. Stipe base brown, upper part, rachis, and costae straw-colored; stipe ± rough, sometimes glabrous.

* Forests along valleys; 1900–2300 m. Sichuan (Shimian), Xizang (Chayu, Neilamu), Yunnan (Gongshan).

The type of Pteris aspericaulis var. cuspigera is from Yunnan (Gongshan).

49c. Pteris aspericaulis var. subindivisa (C. B. Clarke) Ching ex S. H. Wu, Fl. Reipubl. Popularis Sin. 3(1): 61. 1990.

高山凤尾蕨  gao shan feng wei jue

Pteris subindivisa C. B. Clarke, Trans. Linn. Soc., Bot., ser. 2, 1: 467. 1880; P. quadriaurita Retzius var. subindivisa (C. B. Clarke) Beddome.

Plants 0.3–0.5 m tall, precocious. Stipe ca. 8 cm; stipe, rachis, costae, lamina margin, and indusia ± purple; lamina to 18 Χ 3–3.5 cm, lateral pinnae 1 or 2 pairs, 2–6 cm.

W Yunnan (Gongshan) [common in S Himalaya].

The type of Pteris aspericaulis var. subindivisa is from the Himalaya.

49d. Pteris aspericaulis var. tricolor T. Moore ex Lowe, New Ferns 19, t. 9. 1861.

三色凤尾蕨  san se feng wei jue

Pteris tricolor Linden, Gard. Chron. 1860: 217. 1860.

Plants 0.3–0.5 m tall. Pinnae white or rosy, with pink costa with wide whitish or rose-colored bands along both sides, margins green.

SE Yunnan (Yingjiang) [Bhutan, India (Sikkim)].

Pteris aspericaulis var. tricolor is a beautiful ornamental plant.

50. Pteris caiyangheensis L. L. Deng, Bull. Bot. Res., Harbin 16: 423. 1996.

莱阳河凤尾蕨  lai yang he feng wei jue

Plants ca. 50 cm tall. Rhizome ascending, short, ca. 5 mm in diam., apex with yellow-brownish scales. Fronds monomorphic, clustered; stipe 2 Χ as long as lamina, 1–2 mm in diam., mauve as are rachis and frond margin, with yellow-brownish scales at base, glabrous, ± rough; lamina 2-pinnatipartite, 3-forked, ovate-triangular in outline, 15–20 Χ 10–15 cm; lateral pinnae 1 pair, with basiscopic pinnule similar to terminal pinna; segments 2–25 pairs, opposite or subopposite, contiguous, subspreading, somewhat falcate, 20–30 Χ ca. 5 mm, entire or sometimes slightly undulate, apex mucronate or obtuse, and with a protuberance; terminal pinna broadly lanceolate, basally broadly cuneate, stalked (1–2 cm), pectinate nearly to costae, apically tapering; costae prominent abaxially, glabrous, grooved adaxially, with 2–3 mm spine at intersecting point of segments and costa; veins conspicuous on both surfaces, 2-forked at base of segments, lateral veins of segment base arising from rachis; lamina green with mauve margins, subherbaceous when dried, glabrous; indusia flavescent.

* Evergreen broad-leaved forests, monsoon forests; 800–1000 m. Yunnan (Simao).

Pteris caiyangheensis is similar to P. aspericaulis var. aspericaulis, but differs in having lamina ovate-triangular, 2-pinnatipartite, 3-forked; stipe 2 Χ as long as lamina; indusia flavescent. It is treated as a synonym of P. aspericaulis var. tricolor by Zhang Xianchun in the Catalogue of Life, China (http://data.sp2000.cn/2010_cnnode_e/show_species_details.php?name_code=90ed8812-a375-40c9-9b84-c224c315ed89, accessed 09 Jan 2012)

51. Pteris splendida Ching, Acta Bot. Austro Sin. 1: 9. 1983.

隆林凤尾蕨  long lin feng wei jue

Plants up to 1 m tall. Rhizome not seen. Stipe brown at base, dark, ca. 80 cm Χ 5 mm, upper part straw-colored, adaxially with 2 angular grooves; rachis straw-colored, glabrous; lamina 2- or 3-pinnatipartite, narrowly ovate in outline, ca. 55 Χ 25 cm; lateral pinnae ca. 7 pairs, opposite, decumbent, approximate, basal pair of pinnae with stalks ca. 2 cm, upper pairs shortly stalked to gradually sessile; basiscopic pinnae broadly lanceolate, 25–30 Χ 4–8 cm, basally narrower, subtruncate, symmetrical, pectinately divided nearly to costae, apex linear-caudate (ca. 5 cm); basal pair 2-forked at base, with a deeply pectinately pinnatifid pinnule at base; terminal pinnae similar to lateral pinnae, but with longer stalk; segments ca. 30 pairs, alternate, contiguous, decumbent, linear-lanceolate, somewhat falcate, median ones 30–50 Χ 5–8 mm, upper ones gradually reduced, apex acute or obtuse, with a sharp point, entire; costae glabrous, adaxially grooved, with flat needlelike spines at both narrow sides of costa, and short spines on midvein; veins conspicuous, prominent abaxially, 2-forked at base, oblique; lamina papery when dried, glabrous.

* Densely shaded or open forests; 700–1000 m. NW Guangxi (Longlin), Guizhou, Hunan (Fenghuang, Tongdao).

1a. Lateral pinnae ca. 25 Χ 8 cm; median segments ca. 50 Χ 8 mm, adaxially pale green when dried  51a. var. splendida

1b. Lateral pinnae 26–30 Χ 4–5 cm; segments to 30 Χ 5–6 mm, adaxially brown-green when dried  51b. var. longlinensis

51a. Pteris splendida var. splendida

隆林凤尾蕨(原变种)  long lin feng wei jue (yuan bian zhong)

Lateral pinnae ca. 25 Χ 8 cm, apices linear, caudate; segments linear-lanceolate, somewhat falcate, median segments ca. 50 Χ 8 mm; lamina abaxially gray-green, adaxially pale green.

* Open forests in sand-shale areas; ca. 700 m. NW Guangxi (Longlin), Guizhou, Hunan (Fenghuang, Tongdao).

The type of Pteris splendida is from Guangxi.

The typical variety is similar to Pteris fauriei, but the former’s pinnae are deeply pinnatifid, with many long spines, and with a short acute spine at the apices of segments, a pair of veins at the segment base extends at near right angles, almost parallel to the rachis.

51b. Pteris splendida var. longlinensis Ching & S. H. Wu, Acta Bot. Austro Sin. 1: 9. 1983.

细羽凤尾蕨  xi yu feng wei jue

Lateral pinnae lanceolate, 26–30 Χ 4–5 cm; segments up to 30 Χ 5–6 mm; fronds brown-green when dried, slightly lustrous.

* Densely shaded forests; ca. 1000 m. NW Guangxi (Longlin), Hunan (Fenghuang).

52. Pteris setulosocostulata Hayata, Icon. Pl. Formosan. 4: 241. 1914 [“setuloso-costulata”].

有刺凤尾蕨  you ci feng wei jue

Plants 90–120 cm tall. Rhizome erect, robust, 2–3 cm in diam., woody, apex densely clothed with black-brown, subulate scales. Fronds clustered (6–10 per plant); stipe 50–60 cm, ca. 4 mm in diam., basally brown and sparsely scaly, upper part and rachis straw-colored, sometimes brown-green, glabrous; lamina 2(or 3)-pinnatipartite, oblong, 40–60 Χ 20–30 cm; lateral pinnae 9–16 pairs, opposite, decumbent, lowest pair slightly shortly stalked, 15–18 Χ 2–3 cm, with 2–4 pinnules on basiscopic side, pectinately divided; upper pinnae sessile, linear-lanceolate, base broadly cuneate, pectinately divided nearly to costa, apex acuminate; terminal pinna similar to median lateral pinnae but larger, with stipe ca. 1 cm; segments 28–32 pairs, alternate, slightly decumbent, 1–2 mm apart, oblong, sometimes falcate, 10–18 Χ 3–5 mm, entire; costae straw-colored, sometimes light brown or mauve, abaxially convex, glabrous, adaxially grooved, with needlelike spines along groove and midvein; veins conspicuous on both surfaces, oblique, 2-forked from base, veins of segment base oblique, reaching margin of incision; lamina pale green, firmly herbaceous when dried, glabrous.

Forests; 1000–2500 m. Guizhou (Pu’an), Sichuan (Emeishan), Taiwan, S and W Yunnan [Japan, Philippines]. 

The type of Pteris setulosocostulata is from Taiwan.

53. Pteris kiuschiuensis Hieronymus, Hedwigia 55: 341. 1914.

平羽凤尾蕨  ping yu feng wei jue

Plants 60–80 cm tall. Rhizome erect, short, ca. 1 cm in diam., apex with yellowish brown scales. Fronds clustered; stipe reddish brown at base, 25–55 cm, 2–3 mm in diam.; stipe and rachis straw-colored or brown-straw-colored, slightly lustrous, glabrous; lamina 2(or 3)-pinnatipartite, ovate in outline, 25–35(–40) Χ 20–25(–30) cm; lateral pinnae 4–9 pairs, opposite, basal pairs 3–4 cm apart, sessile, 12–16(–18) Χ 1.8–3.7 cm, base subtruncate or rounded-truncate, pectinate and forming narrow wings along costae, apex acuminate or long caudate (2–4 cm); upper pinnae slightly smaller than basal ones, spreading or oblique; apical pinnae similar to lateral pinnae but wider, sometimes narrower at base, stalked (1–2 cm); basal pair of pinnae often branched basiscopically near base, branch similar to main part of pinna but smaller; segments 23–28 pairs, alternate or subopposite, contiguous or at intervals of ca. 1 mm, slightly oblique, oblong, sometimes falcate, 1–1.5 cm Χ 4–5 mm, base slightly expanded, margin entire, apex obtuse; costae prominent abaxially, straw-colored, glabrous, adaxially grooved, with short and flat spines along groove, midvein sparsely spiny, spines sometimes poorly developed; veins conspicuous on both surfaces, oblique, 2-forked from base, veins at segment base oblique, reaching margin of incision. Lamina grass green, thinly herbaceous when dried, glabrous.

Open forests, near river margins; 300–1200 m. Chongqing, Fujian (Chongan, Jianyang, Wuyishan), Guangdong (Guangzhou, Luofushan, Renhua), Guangxi, Guizhou (Dushan, Jiangkou, Langdai), Hunan, Jiangxi, Sichuan (Emeishan), S Yunnan [Japan].

1a. Lateral pinnae spreading by right angles, linear-lanceolate, straight or somewhat falcate, 1.8–2.5(–2.8) cm wide ...................................................................................................  53a. var. kiuschiuensis

1b. Lateral pinnae often decumbent, up to 3.7 cm wide ...............................  53b. var. centrochinensis

53a. Pteris kiuschiuensis var. kiuschiuensis

平羽凤尾蕨(原变种)  ping yu feng wei jue (yuan bian zhong)

Lateral pinnae spreading, at right angles to rachis, linear-lanceolate, straight or somewhat falcate, 12–16(–18) Χ 1.8–2.5(–2.8) cm.

Open forests; 500–1200 m. Chongqing, Fujian (Chongan, Jianyang, Wuyishan), Guangdong (Guangzhou, Luofushan, Renhua), Guangxi, Guizhou (Dushan, Jiangkou, Langdai), Hunan, Jiangxi (Nanping, Yifeng), Sichuan (Emeishan), S Yunnan [Japan].

The type of Pteris kiuschiuensis is from S Japan.

53b. Pteris kiuschiuensis var. centrochinensis Ching & S. H. Wu, Acta Bot. Austro Sin. 1: 10. 1983 [“centro-chinensis”].

华中凤尾蕨  hua zhong feng wei jue

Lateral pinnae often oblique, up to 3.7 cm wide.

* Near river margins; 300–800 m. Chongqing, Fujian (Chongan, Jianyang), Guangdong, Guangxi (Pingnan, Rongshui, Tianyang), Guizhou (Dushan, Langdai), Hunan (Baojing, Dongan, Huitong), Jiangxi, Sichuan (Emeishan), S Yunnan.

The type of Pteris kiuschiuensis var. centrochinensis is from Guangdong.

54. Pteris oshimensis Hieronymus, Hedwigia 55: 367. 1914.

斜羽凤尾蕨  xie yu feng wei jue

Plants 50–80 cm tall. Rhizome erect, short, apex with brownish black scales. Fronds clustered; stipe basally brown, upper part and rachis and costae straw-colored, 25–50 cm, ca. 2 mm in diam., glabrous; lamina 2(or 3)-pinnatipartite, oblong in outline, 30–40 Χ 10–18 cm; lateral pinnae 7–9 pairs, opposite, decumbent, sessile, lanceolate, base broadly cuneate, symmetrical, lowest pair of pinnae cut obliquely from base, pectinately lobed nearly to costa, often with 1(or 2) basiscopic pinnules similar in shape to main part of pinna; terminal pinna same as lateral pinnae, with stipe ca. 1 cm; segments 22–30 pairs, alternate or opposite, ca. 1 mm apart, spreading or decumbent, lanceolate, somewhat falcate, 10–15 Χ 3–4 mm, slightly expanded, slightly narrower toward top, obtuse, entire; costae glabrous, abaxially grooved, with needlelike spines along groove, midvein with a few needlelike spines or spineless; veins conspicuous on both surfaces, 2-forked at base, oblique, basal pair of veins reaching margin above sinus; lamina pale green or brown-green when dried, herbaceous, glabrous.

Open forests; 300–900 m. Chongqing, Fujian (Nanping), Guangdong, Guangxi (Baise, Yangshuo), Guizhou (Pingtang, Zhenfeng), Hunan, Jiangxi (Dayu, Quannan, Yifeng), Sichuan, Zhejiang [Japan, N Vietnam].

1a. Basal lateral pinnae (8–)11–14(–17) cm, with short, acute tips 1–2 cm .........  54a. var. oshimensis

1b. Basal lateral pinnae 14–25 cm, with long tips (3–)4–9 cm .......................  54b. var. paraemeiensis

54a. Pteris oshimensis var. oshimensis

斜羽凤尾蕨(原变种)  xie yu feng wei jue (yuan bian zhong)

Pteris omeiensis Ching.

Basal lateral pinnae (8–)11–14(–17) Χ (1.7–)2–2.8 cm, apically attenuate to form linear, acute tip 1–2 cm, not or slightly narrower at base. 3n = 87.

Open forests; 300–900 m. Chongqing, Fujian (Nanping), Guangdong, Guangxi (Baise, Yangshuo), Guizhou (Pingtang, Zhenfeng), Hunan, Jiangxi (Dayu, Quannan, Yifeng), Sichuan (Emeishan) [Japan, N Vietnam].

The type of Pteris oshimensis is from S Japan. Christ (Bull. Herb. Boissier 2: 1016. 1901) treated material of this species as P. asperula J. Smith.

54b. Pteris oshimensis var. paraemeiensis Ching, Acta Bot. Austro Sin. 1: 10. 1983.

尾头凤尾蕨  wei tou feng wei jue

Lower lateral pinnae 14–25 Χ 1.7–3.5 cm, apically attenuate into long tips, (3–)4–9 cm.

* Forests; 500–600 m. Chongqing, Guangxi (Baise), Hunan (Shimen), Sichuan.

The type of Pteris oshimensis var. paraemeiensis is from Sichuan (Emeishan). Material was originally included with P. omeiensis Ching (Fang 17956).

55. Pteris fauriei Hieronymus, Hedwigia 55: 345. 1914.

傅氏凤尾蕨  fu shi feng wei jue

Plants 50–90 cm tall. Rhizome ascending, short, ca. 1 cm in diam., apex densely scaly; scales dark brown, brown at margin, linear-lanceolate, ca. 3 mm. Fronds clustered; stipe straw-colored, 30–50 cm, 2–4 mm in diam., with dark brownish scales, glabrous, adaxially grooved; rachis similar; lamina 2(or 3)-pinnatipartite, ovate to ovate-triangular in outline, 25–45 Χ 17–24(–30) cm; lateral pinnae 3–6(–9) pairs, basal pairs opposite, 4–8 cm apart, decumbent, rarely slightly decumbent, basal pair sessile or shortly stalked, often with a basiscopic pinnule near base similar to main part of pinna but smaller; upper pairs sessile, base attenuate, broadly cuneate, pectinate and forming narrow wings along costae, apex linear-caudate, 2–3(–4.5) cm, acuminate; terminal pinnae similar but wider and stalked (2–4 cm); segments 20–30 pairs, alternate or opposite, conjoined or ca. 1 mm apart (fertile segments up to 2 mm apart), decumbent, falcate, broadly lanceolate, usually basiscopic segments longer than acroscopic segments, basal pair or several pairs of segments shortened at base, entire, apex obtuse; costae abaxially prominent, straw-colored, glabrous, grooved adaxially, with needlelike flat spines on both sides of groove, and few spines on costa; veins conspicuous on both surfaces, oblique, 2-forked at base, basiscopic vein of segment arising from rachis, and acroscopic vein arising from base of costa, opposite veins at base oblique up to margin of incision; lamina light green to pale green, papery when dried, glabrous (rarely glabrescent when young); indusia gray-brown, linear, membranous, persistent. n = 29, 2n = 58, 87, 89

 Acidic soils of forests along valleys; sea level to 800 m. Anhui, Fujian (Chongan, Shaowu), Guangdong, Guangxi (Douyun), Hainan (Lizhishan), S Hunan (Tongdao, Yizhang, Yongshun), Jiangxi, Taiwan, Xizang, SE Yunnan (Hekou), Zhejiang (Nanhui, Tiantaishan) [Japan, N Vietnam].

Fraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes, 109. 2008) includes Pteris fauriei within P. khasiana (C. B. Clarke) Hieronymus, which would extend the distribution to Bangladesh, NE India, Myanmar, Nepal, Philippines, and Vietnam.

Pteris medogensis Ching & S. K Wu (Fl. Xizang. 1: 71–72, t. 16,  fig. 1–4. 1983) and P. bomiensis Ching & S. K. Wu (loc. cit.: 72, tab. 19) appear to fall within the range of variation accepted for P. fauriei. Pteris wulaiensis C. M. Kuo (Bot. Bull Acad. Sin., n.s., 30: 143) is an apomictic diploid intermediate between P. fauriei and P. bella and is presumed to be of hybrid origin.

1a. Lateral pinnae falcate-lanceolate, 3–4 cm wide; median segments of pinnae 15–22 Χ 4–6 mm  55a. var. fauriei

1b. Lateral pinnae broadly lanceolate, 4–6 cm wide; median segments of pinnae 20–35 Χ (5–)6–8 mm  55b. var. chinensis

55a. Pteris fauriei var. fauriei

傅氏凤尾蕨(原变种)  fu shi feng wei jue (yuan bian zhong)

Pteris fauriei var. minor Hieronymus; P. guizhouensis Ching; P. linearis Poiret var. fauriei (Hieronymus) C. Christensen & Tardieu; P. longipinnula H. Christ (1906), not Wallich ex J. Agardh (1839); P. quadriaurita Retzius var. abbreviata Rosenstock.

Lateral pinnae falcate-lanceolate, 13–23 Χ 3–4 cm; median segments of pinnae 15–22 Χ 4–6 mm.

Acidic soils of forests along valleys; sea level to 800 m. Fujian (Chongan, Shaowu), Guangdong, Guangxi (Douyun), Hainan (Lizhishan), S Hunan (Tongdao, Yizhang, Yongshun), Jiangxi, Taiwan, SE Yunnan (Hekou), Zhejiang (Nanhui, Tiantaishan) [Japan, N Vietnam].

The type of Pteris fauriei is from S Japan. W. J. Hooker (Sp. Fil. 2: 179. 1858) and others included material of this taxon within P. quadriaurita, while C. Christensen (Index Filic. 593. 1906) included it within P. biaurita.

55b. Pteris fauriei var. chinensis Ching & S. H. Wu, Acta Bot. Austro Sin. 1: 10. 1983.

百越凤尾蕨  bai yue feng wei jue

Pteris bifurcata Ching.

Lateral pinnae broadly lanceolate, 16–22 Χ 4–6 cm; median segments of pinnae 20–35 Χ (5–)6–8 mm.

* Forests along valleys; 300–700 m. Fujian (Chongan), Guangdong (Gaoyao, Qujiang, Yunfu), Guangxi, S Guizhou (Dushan), Hainan (Chengmai, Danxian, Lingao), Taiwan.

The type of Pteris fauriei var. chinensis is from Guangdong. Material of this taxon was identified as P. longipinnula by Merrill (Lingnan Sci. J. 5: 16. 1927) and as P. fauriei s.s. by Ching (Icon. Fil. Sin. 3: t. 140. 1935).

56. Pteris angustipinnula Ching & S. H. Wu, Acta Bot. Austro Sin. 1: 11. 1983.

线裂凤尾蕨  xian lie feng wei jue

Plants ca. 1 m tall. Rhizome not seen. Stipe 43–65 cm, ca. 3 mm in diam., sparsely scaly; stipe apically straw-colored, or slightly dark chestnut, slightly lustrous, glabrous, grooved on upper side; scales brown, lanceolate, ca. 2 mm, gradually deciduous; rachis straw-colored, glabrous, adaxially grooved; lamina 2- or 3-pinnatipartite, oblong-ovate in outline, 35–45 Χ 16–24 cm; lateral pinnae 5–7 pairs, opposite, slightly decumbent, basal pairs 5.5–7.5 cm apart; basal pair slightly shortly stalked; distal pairs sessile, median pinnae broadly lanceolate, 17–22 Χ 4–6.5 cm, base rounded-cuneate, pectinately divided with narrowly winged costa, apex caudate-acuminate (4–5.5 mm); pinnae asymmetrical; basal pairs slightly wider, with 1 or 2 basiscopic pinnules similar in shape to main part of pinna but smaller; segments 21–25 pairs, opposite or alternate, decumbent, 2–4 mm apart, linear-lanceolate, somewhat falcate, basal segments slightly shorter; middle segment 25–40 Χ 4–5 mm, apical segment shorter and becoming narrow, obtuse, slightly expansive at base, entire; terminal pinna similar to lateral pinnae, but wider, and shortly stalked (5–15 cm); costae and midvein prominent abaxially, light straw-colored, glabrous, adaxially grooved, with needlelike flat spines along groove of costae, and with a few needlelike spines along midvein; veins conspicuous on both sides, 2-forked, decumbent, oblique, basiscopic vein at segment base arising from rachis, and acroscopic vein together extending up to margin of incision; lamina gray-green, papery when dried, glabrous, or rarely minutely puberulent; indusia gray-brown, linear, membranous, entire, persistent.

* Forests in valleys. SW Guangxi (Fusui, Longjin).

Pteris angustipinnula is similar to P. fauriei, but differs in having linear-lanceolate segments that are distant from each other.

57. Pteris obtusiloba Ching & S. H. Wu, Acta Bot. Austro Sin. 1: 11. 1983.

江西凤尾蕨  jiang xi feng wei jue

Plants ca. 70 cm tall. Rhizome not seen. Stipe light straw-colored or chestnut abaxially, shiny, ca. 40 cm, up to 2 mm in diam., shallowly grooved adaxially, basally scaly, distally glabrous; scales brown with brown margins, lanceolate, ca. 1 mm; lamina 2- or 3-pinnatipartite, oblong-ovate in outline, ca. 30 Χ 20 cm; lateral pinnae 6 or 7 pairs, opposite, oblique, 3.5–4.5(–7) cm apart; basal pair of pinnae shortly stalked, other pairs sessile, lanceolate, 12–16 Χ 2–2.5 cm, base rounded-cuneate, apex caudate-acuminate, pinnae anisomerous, pectinately deeply divided leaving narrowly winged rachis, basiscopic side wider, basal pair of pinnae often with a basiscopic pinnule near base similar to main part of pinna but smaller; segments 19–23 pairs, alternate or subopposite, oblique, ca. 1 mm apart, linear, somewhat falcate, basal lobes shorter, median 13–16 Χ ca. 4 mm, parallel on both sides, base widened, margin entire, apex obtuse; terminal pinnae same shape and size as lateral pinnae, but symmetrical on both sides, shortly stalked (ca. 1 cm); costae and midvein prominent abaxially, pale straw-colored, glabrous, costae shallowly grooved adaxially, with spines on both sides, and a few spiculate thorns in pinna costae; veins conspicuous on both surfaces, 2-forked, oblique, lateral veins of segment base arising abaxially from rachis, other lateral vein adaxially extended to margin of incision; lamina green, herbaceous when dried, glabrous; rachises same as stipes in color, glabrous, slightly grooved adaxially; indusia gray-brown, linear, membranous, entire, persistent.

* Forests along valleys; ca. 500 m. Hunan (Lingxiang, Yongshun), SW Jiangxi (Chongyi), Zhejiang.

Pteris obtusiloba is similar to P. fauriei, but differs by the herbaceous lamina, narrow pinnae, median 2–2.5 cm wide, with shorter segments, 13–16 Χ ca. 4 mm, linear, slightly falcate, parallel to both margins, apex rounded, with spiculate spines on both sides of groove adaxially.

58. Pteris majestica Ching, Acta Bot. Austro Sin. 1: 12. 1983.

硕大凤尾蕨  shuo da feng wei jue

Plants more than 1.3 m tall. Rhizome not seen. Stipe straw-colored, ca. 80 cm, ca. 5 mm in diam. at middle, glabrous, grooved adaxially; rachis similar, slightly lustrous, glabrous, grooved adaxially; lamina 2- or 3-pinnatipartite, ovate-oblong in outline, ca. 65 Χ 30–50 cm; lateral pinnae ca. 7 pairs, opposite or subopposite, slightly decumbent, stalked (5–10 mm), distal ones sessile, slightly asymmetrically broadly lanceolate, ca. 35 Χ 8 cm, base broadly cuneate, pectinately divided to narrowly winged costae, apex caudate-acuminate; basal pair of pinnae often with a basiscopic pinnule near base similar to main part of pinna but smaller; segments 30–35 pairs, alternate, 1.5–3 mm apart, decumbent, lanceolate, somewhat falcate, median ones 35–55 Χ 7–9 mm, basally wider, margins entire, apex obtuse; basal segments conspicuously shorter; terminal pinna similar to middle lateral pinnae, but symmetrical and with basal stalk ca. 1 cm; costae and midvein prominent abaxially, straw-colored, glabrous, costae adaxially shallowly grooved, with flat rigid spine on both sides, midvein with a few very short spines on upper surface; veins conspicuous on both sides, decumbent, 2-forked at base, lateral veins of segment base abaxially arising from rachis, and other lateral vein adaxially extended at margin of incision; lamina grass green, thickly papery when dried, glabrous; indusia gray-brown, linear, membranous, entire, persistent.

* Dense forests, wet valleys; ca. 2700 m. Guangdong, Sichuan (Emeishan, Ya’an), SE Yunnan (Jingdong, Wuliangshan).

Pteris majestica is similar to P. fauriei, but much taller, more than 1.3 m, lateral pinnae ca. 35 Χ 8 cm, middle segments 3.5–5.5 cm Χ 7–9 mm.

59. Pteris puberula Ching, Bull. Fan Mem. Inst. Biol. Bot. 11: 52. 1941.

柔毛凤尾蕨  rou mao feng wei jue

Plants 50–75 cm tall. Rhizome erect, short, ca. 1 cm in diam., apex densely clothed with brown scales. Fronds clustered; stipe brown straw-colored, rarely pale straw-colored, shiny, 17–40 cm, 2–3 mm in diam., glabrous; rachis similar in color, slightly lustrous, sparsely gray puberulent or glabrescent; lamina 2- or 3-pinnatipartite, broadly ovate in outline, 30–44 Χ (14–)22–27 cm; lateral pinnae 5–7 pairs, opposite, oblique or decumbent, basal pair shortly stalked, remainder sessile, broadly lanceolate, 14–17 Χ 3.2–4.5 cm, base rounded-truncate, pectinately divided to narrowly winged costae, apex acuminate, lobate or shortly caudate; basal pair of pinnae often with 1(or 2) basiscopic pinnules similar to main part of pinna but smaller; segments 25–30 pairs, alternate or subopposite, slightly decumbent, 1–2 mm apart, falcate, broadly linear, 20–25 Χ 5–6 mm, base conspicuously expanded, margin entire, apex obtuse; terminal pinna similar to median lateral pinnae, stalk 1–1.5 cm; costae prominent abaxially, light straw-colored, slightly lustrous, sparsely gray strigose, with soft flat spines along grooves, and sparsely gray puberulent abaxially on midvein; veins conspicuous on both sides, decumbent, 2-forked at base, pair of opposite veins at segment base extending up to margin of incision; lamina light green when dried, thinly herbaceous, abaxially sparsely gray strigose, adaxially glabrous.

* Forests; 2500–2800 m. Xizang (Neilalmu), Yunnan (Dangbi, Dayao).

The type of Pteris puberula is from Yunnan (Dangbi).

60. Pteris viridissima Ching, Acta Bot. Austro Sin. 1: 13. 1983.

绿轴凤尾蕨  lό zhou feng wei jue

Plants ca. 70 cm tall. Rhizome erect, short, ca. 1.5 cm in diam., apex with brown scales. Fronds clustered; stipe dark brownish on lower parts, slightly lustrous, ca. 30 cm Χ 2 mm, slightly flat, upper part of stipes and rachises and costae blue-green, slightly lustrous, glabrous; lamina 2(or 3)-pinnatipartite, oblong-ovate in outline, ca. 45 Χ 20 cm; lateral pinnae ca. 9 pairs, decumbent, lower subopposite, shortly stalked, upper subopposite or alternate, sessile, broadly lanceolate, 13–15 Χ 4–5 cm, basally rounded-cuneate, pectinately divided to narrowly winged costae, apex acuminate and lobate; basal pair of pinnae basiscopically branched near base, branch similar to main part of pinna but smaller; segments ca. 25 pairs, alternate, slightly decumbent, contiguous or narrowly separated, broadly linear and straight, 20–30 Χ 4–5 mm, basally slightly expanded, margins entire, apex mucronate or subobtuse; terminal pinnae similar to lateral pinnae but with stipe ca. 1 cm; costae grooved with flat spines on both sides, and spiculate spinules on midvein; veins conspicuous on both surfaces, decumbent, 2-forked at base, pair of opposite veins at segment base extending up to margin of incision; lamina blue-green, thinly herbaceous when dried, pinnae abaxially glandular along both sides of rachis.

* Limestone soils; 600–2000 m. Guizhou (Pingtang), Hunan (Shimen), Yunnan.

The type of Pteris viridissima is from Yunnan.

61. Pteris hirsutissima Ching, Acta Bot. Austro Sin. 1: 13. 1983.

微毛凤尾蕨  wei mao feng wei jue

Plants 75–90 cm tall. Rhizome ascending, short and thick. Fronds clustered; stipe brown at base, apical portion straw-colored, 30–45 cm Χ 2–3 mm, with brown scales at base; lamina 2(or 3)-pinnatipartite, oblong, 40–45 Χ 20–25 cm; lateral pinnae 4–8 pairs, subopposite or opposite, decumbent, sessile or basal pair stalked, stalk lanceolate, ca. 1 cm; basal pairs 20–23 Χ 4.5–5.5 cm, basally rounded-cuneate, pinnate to costa, apex acuminate, caudate; basal pair of pinnae each with basiscopic pinnule similar in shape to main part of pinna but smaller; segments 24–30 pairs, oblique, ca. 2 mm apart, lanceolate, 2–3.5 cm, basally enlarged (6–7 mm wide), margin entire, apex attenuate, acuminate; basiscopic segments slightly longer than acroscopic segments; terminal pinna similar to median lateral pinnae, stalked (2.5–3.5 cm); costae prominent abaxially, pale brown-straw-colored, sometimes mauve, with flat short spines on both sides of groove, and 3–5 thin spines on midvein; costae, midvein, and abaxial surface of segments sparsely strigose; rachis straw-colored, glabrous; veins conspicuous on both surfaces, oblique, 2-forked from base, and opposite veins at segment base oblique up to margin of incision; lamina brown-green, herbaceous when dried.

* Forests; ca. 1400 m. Sichuan (Mianning).

62. Pteris subquinata Wallich ex J. Agardh, Recens. Spec. Pter. 21. 1839.

勐海凤尾蕨  meng hai feng wei jue

Pteris monghaiensis Ching; P. quadriaurita Retzius var. subquinata (Wallich ex J. Agardh) Beddome.

Plants 80–90 cm tall. Rhizome not seen. Stipe straw-colored, 50–60 cm Χ ca. 3 mm, glabrous, grooved adaxially; lamina 2- or 3-pinnatipartite, triangular-ovate in outline, ca. 30 Χ 20 cm; lateral pinnae 3 pairs, opposite, decumbent, sessile, lanceolate, 15–17 Χ ca. 3.5 cm, base rounded-cuneate, pectinately pinnate, apex acuminate and lobate; basal pair of pinnae often with a basiscopic pinnule similar to main part of pinna but smaller; segments 30–35 pairs, alternate or subopposite, spreading, 2–3 mm apart, linear and substraight, 15–20 Χ ca. 3 mm, base enlarged, margin entire, apex obtuse; terminal pinna similar to median lateral pinnae, stalked (ca. 1 cm); costae prominent abaxially, straw-colored, glabrous, grooved adaxially, with soft flat spines along grooves; veins conspicuous on both surfaces, oblique, thick and dense, 2-forked at base, acroscopic vein on segment base, and basiscopic vein arising from rachis, and opposite veins at base oblique up to margin of incision; lamina gray-brown, subleathery when dried, glabrous.

* Forests along valleys; ca. 1000 m. S Yunnan (Menghai, Xishuangbanna).

63. Pteris bella Tagawa, Acta Phytotax. Geobot. 8: 166. 1939.

栗轴凤尾蕨  li zhou feng wei jue

Pteris wangiana Ching.

Plants 50–70 cm tall. Rhizome long ascending, ca. 1 cm in diam., woody, apex with light brown scales. Fronds clustered; stipe with basal part castaneous-brown, upper part and rachis castaneous-red, shiny, 30–40 cm, ca. 2 mm in diam., glabrous, scales with expanded bases; lamina 2- or 3-pinnatipartite, broadly ovate to oblong in outline, 30–35 Χ 15–20 cm; lateral pinnae 5 or 6, opposite, decumbent; basal pair shortly stalked, each often with 1(or 2) basiscopic pinnules similar to main part of pinna but smaller, middle pinnae sessile, lanceolate, 10–15 Χ 2–3 cm, pectinately divided to near costa, apex long acuminate; segments 20–25 pairs, alternate, approximate, slightly decumbent, narrowly oblong, 10–15 Χ 3–4 mm, base rounded-cuneate, basally enlarged with basiscopic side slightly decurrent, margin entire, apex obtuse; terminal pinna similar to median lateral pinnae, but with stipe ca. 1 cm; costae abaxially light straw-colored, sometimes chestnut abaxially, shiny, glabrous, slightly grooved adaxially, with aciculate spines on both sides of groove; veins conspicuous, sparse, oblique, basal part forked but simple upward; lamina grass green, herbaceous when dried, glabrous.

Shaded forests; 600–1600 m. Hainan (Wuzhishan), Hunan (Yizhang), N and S Taiwan, Yunnan [S Vietnam].

The type of Pteris bella is from SE Yunnan.

64. Pteris scabristipes Tagawa, Acta Phytotax. Geobot. 5: 103. 1936.

红柄凤尾蕨  hong bing feng wei jue

Rhizome short, ascending, scaly. Stipe reddish, 25–45 cm, muricate, scaly at base, scales not expanded at base; lamina 2-pinnatifid, oblong-ovate in outline, 30–50 cm; lateral pinnae 4–9 pairs, opposite; basal pinnae each with a basiscopic pinnule similar to main part of pinna but smaller; middle pinnae sessile, narrowly lanceolate, pectinately divided; segments 20–26 pairs, often opposite, narrowly oblong-falcate, to ca. 15 Χ 5 mm, margin entire, apex subacute.

* Taiwan.

65. Pteris linearis Poiret in Lamarck, Encycl. 5: 723. 1804.

线羽凤尾蕨  xian yu feng wei jue

Pteris arisanensis Tagawa; P. biaurita Linnaeus var. intermittens C. Christensen.

Plants 1–1.5 m tall. Rhizome erect, short, 1.5–2 cm in diam., apex with black-brown scales. Fronds clustered (6–8 per plant); stipe basally brown, upper part straw-colored, slightly lustrous, as long as fronds, 3–4 mm in diam., glabrous; lamina 2- or 3-pinnatipartite, oblong-ovate in outline, 50–70 Χ 20–30 cm; lateral pinnae 5–15 pairs, opposite, slightly decumbent, sessile or basal pairs shortly stalked, lanceolate, 15–25(–33) Χ 3–4(–5.5) cm, base rounded-cuneate and slightly oblique, deeply pectinately divided to winged costa, apex long caudate; basal pair of pinnae often each with basiscopic pinnule similar to main part of pinnae but smaller; segments 25–35 pairs, alternate, subspreading or oblique, falcate-oblong, 20–30 Χ 5–8 mm, base slightly enlarged, margins entire, apex obtuse or mucronate; terminal pinna similar to lateral pinnae; costae with 6–10 mm wide wings, prominent abaxially, straw-colored, glabrous, grooved adaxially, with spines; veins conspicuous and convex on both sides, decumbent, 2-forked at base, basiscopic vein of segment base arising from rachis, and acroscopic vein from base of costa, 2 opposite veins of pinnae base arriving at margin of incision and forming a fork or triangle, or sometimes interlinked into a continuous triangular mesh, and other veins outward from mesh separate and extending to base of incision; lamina green, yellowish green, or brown-green, subleathery when dried, glabrous.

Dense forests, wet areas near stream margins; 100–1800 m. Guangdong, Guangxi, SW Guizhou (Ceheng), Hainan, Sichuan (Dechang, Yanbian), Taiwan, C, S, and W Yunnan [widely distributed in tropical Africa, Asia, and Madagascar].

The type of Pteris linearis is from E India.

66. Pteris longipes D. Don, Prod. Fl. Nepal. 15. 1825.

三轴凤尾蕨  san zhou feng wei jue

Hypolepis pteridioides Hooker; Pteris pellucens J. Agardh; P. brevisora Baker; P. zollingeri Mettenius ex Miquel.

Plants 1.2–1.5 m tall. Rhizome erect, short, 1.5–2 cm in diam., woody, apex with dark brown scales. Fronds clustered; stipe dark straw-colored to light brown, slightly lustrous, 70–80 cm, 6–8 mm in diam., firm, glabrous; lamina 3-pinnatipartite, triangular-ovate in outline, 60–70 Χ 35–45 cm; lateral pinnae 12–20 pairs, subopposite or alternate, oblique, sessile, lanceolate, 10–12 Χ 1.5–2 cm, base truncate, pectinately divided nearly to costae, apex caudate (2–3 cm), with linear lobes; segments 25–28 pairs, alternate, interlinked and slightly decumbent, oblong, ca. 10 Χ 3–4 mm, base slightly enlarged, apex obtuse and crenate; apical pinnule same as middle lateral ones, stalked (ca. 1 cm); costae prominent abaxially, straw-colored, glabrous, slightly grooved adaxially, with needlelike spines on both sides; veins conspicuous on both surfaces, oblique, 2-forked at base, and opposite two veins oblique up to margin of incision; lamina pale green to green-brown, herbaceous when dried, glabrous.

Forests; 600–2400 m. W Guangxi (Tianlin), Hunan (Tongdao), C and SW Taiwan, Yunnan (Jingdong, Malipo, Simao) [widely distributed in Bhutan, India, Indonesia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].

The type of Pteris longipes is from Nepal.

66bis. Pteris paucipinnula X. Y. Wang & P. S. Wang, Pterid. Fl. Guizhou, 596. 2001.

稀羽凤尾蕨  xi yu feng wei jue

Plants ca. 1 m tall. Rhizome not seen. Stipe puce at base, ca. 55 cm, ca. 3 mm in diam., stipe and rachis brown-straw-colored, slightly lustrous, glabrous; lamina 3-pinnatipartite, with 3 pinnatisect divisions, lateral branches same in shape with central division, oblong-ovate, 38–44 Χ 20–26 cm; lateral pinnules 4–6 pairs, opposite or nearly opposite, oblique, sessile, median pinnules linear-lanceolate, 16–20 Χ 2.5–3.2 cm, base broadly cuneate, pectinately divided nearly to costae, apex linear, apical pinnules same as lateral ones in shape and size, but with stalks 1–1.5 cm; segments 22–28 pairs, alternate, slightly oblique, oblong to linear-oblong, 1–2 cm Χ 3–4 mm, slightly enlarged at base, margin entire, apex blunt; lamina yellow-green, herbaceous when dried, glabrous; costae abaxially prominent, adaxially grooved, with spines along groove, midvein sparsely spiny, spines sometimes poorly developed; veins conspicuous on both surfaces, oblique, 2-forked from base, veins at segment base oblique, reaching margin of incision.

* Forests. N Guizhou (Chishui).

Pteris paucipinnula is most similar to P. longipes but has only 4–6 pairs of lateral pinnae with entire sterile margins.

66ter. Pteris quadristipitis X. Y. Wang & P. S. Wang, Pterid. Fl. Guizhou, 598. 2001.

方柄凤尾蕨  fang bing feng wei jue

Plants to 63 cm tall. Rhizome erect. Fronds clustered, monomorphic; stipe chestnut brown, 30–38 cm, cross sections nearly quadrate, glabrous; lamina odd-pinnate, ovate, 18–25 Χ 14–20 cm; lateral pinnae 2 or 3 pairs, opposite, oblique, basal pair shortly stalked, 2-forked, upper pairs sessile, linear-lanceolate, 10–13 Χ 1.3–1.6 cm, base cuneate or rounded-cuneate, margin undulate, often slightly lobed or with 1 or 2 lobes, apex acuminate; apical pinnae same in shape as lateral ones but longer, stalked, apex sterile, and sterile margin with short teeth; lamina gray-green when dried, papery, glabrous; veins conspicuous, midvein abaxially prominent, adaxially grooved, with spines along groove, lateral veins 2-forked. Sori linear, indusia gray-white, membranous, entire.

* Forests by streams; ca. 400 m. S Guizhou (Libo).

Pteris quadristipitis looks superficially like P. cretica because of the rather few undivided pinnae; however, the entire pinnae margins and spines along the costae place it in P. sect. Quadriauricula.

3. Pteris sect. Campteria (Presl) Ching, Acta Bot. Austro Sin. 1: 2. 1983.

网眼凤尾蕨组  wang yan feng wei jue zu

Campteria Presl, Tent. Pterid. 146. 1836.

Plants often large. Fronds monomorphic, division patterns and lobe shape same as for sect. Quadriauricula but often divided and 3-divaricate (lateral pinnae usually again divided into 2 forks); lobes without cartilaginous margin, serrate; costae spiny along both sides of groove on adaxial side; venation anastomosing to form a series of narrow areoles along costae, veinlets free beyond areolae and extending to margins. Sori linear, extending continuously along lobe margins, often sterile at apex; indusia brown or light brown, linear, membranous, entire, persistent.

Mainly distributed in the tropics; nine species (three endemic) in China.

67. Pteris biaurita Linnaeus, Sp. Pl. 2: 1076. 1753.

狭眼凤尾蕨  xia yan feng wei jue

Campteris biaurita (Linnaeus) Hooker; Pteris flavicaulis Hayata.

Plants 70–110 cm tall. Rhizome erect, robust, 2–2.5 cm in diam., woody, apex densely clothed with brown scales. Fronds clustered; stipe light brown, apically straw-colored to pale green, slightly lustrous, 40–60 cm, 3–5 mm in diam., glabrous, scaly, rarely with a few scales, adaxially narrowly grooved; rachis straw-colored, glabrous, narrowly grooved adaxially; lamina 2- or 3-pinnatipartite, oblong-ovate in outline, 40–55 Χ 20–30 cm; lateral pinnae 8–10 pairs, decumbent, opposite, lower ones shortly stalked, 3–5 cm apart; upper ones sessile, broadly lanceolate, 15–20 Χ 3–5.5 cm, base broadly cuneate, pectinately divided to broadly winged costa (to 8 mm wide), apex narrowly lanceolate and caudate (2–3 cm); basal pair of pinnae often each with 1 or sometimes 2 basiscopic pinnules similar to main part of pinna but smaller; segments 20–25 pairs, alternate, subspreading, incised obtuse-rounded, 2–5 mm apart, falcate, broadly lanceolate to falcate-oblong, 18–35 Χ 5–7 mm, base slightly enlarged, margins entire, apex slightly narrow and obtuse; terminal pinna similar to middle lateral pinnae, stalked (ca 1.5 cm); costae prominent abaxially, straw-colored, glabrous, slightly grooved adaxially, with short spines on both sides; veins slightly raised, conspicuous on both surfaces, acroscopic veinlet of lobe base and a basiscopic veinlet of upper lobe base combining to form an arcuate vein, anastomosing to form a series of narrow areoles along costules, with 5 or 6 free veinlets extending to margin at arcuate vein, and a majority of veinlets outward from areole usually 2-forked; lamina gray-green, thickly papery when dried, glabrous; indusia light brown, membranous, entire, persistent.

Dry sloping fields; 200–1500 m. Guangdong, Guangxi (Baise), Hainan, C and S Taiwan, Yunnan [Bhutan, India, Indonesia, Laos, Malaysia, Nepal, Philippines, Sri Lanka, Thailand; S Africa, Americas (tropical Brazil, West Indies (Jamaica)), Madagascar, Oceania].

68. Pteris maclurei Ching, Bull. Dept. Biol. Sun Yatsen Univ. No. 6. 28. 1933.

两广凤尾蕨  liang guang feng wei jue

Pteris biaurita Wu, Wong & Pong (1932), non Linnaeus (1753); P. nakasimae Tagawa.

Plants 80–90 cm tall. Rhizome ascending, ca. 1 cm in diam., apex densely clothed with brown scales. Fronds subclustered; stipe dark chestnut, apically light chestnut, shiny, 40–50 cm, 3–4 mm in diam., glabrous, scaly, narrowly grooved adaxially; lamina 2- or 3-pinnatipartite, broadly ovate in outline, 40–45 Χ 25–30 cm; lateral pinnae 5–7 pairs, decumbent, opposite, 7–10 cm apart, shortly stalked with a narrow wing, or sessile toward apex, broadly lanceolate, 15–20(–32) Χ 4–6(–8) cm, base cuneate, pectinately divided to with 5–6 mm of costa, apex lobate or long caudate (2–7 cm), basal pinnae with 1–3 pairs of pinnae often with a basiscopic pinnule similar to main part of pinna but smaller; segments 14–20 pairs, alternate or subopposite, sinuses obtuse, 4–8 mm wide, slightly oblique, falcate-lanceolate, 20–65 Χ 6–9 mm, base enlarged, margins serrate or crenate, entire, apex acuminate or obtuse; terminal pinna similar to median lateral pinnae; costae abaxially prominent, light brown, upper 1/3 part straw-colored, straw-colored and slightly grooved adaxially, with needlelike spines, winged, wing 5–6 mm wide on both sides of costule; veins anastomosing to form a series of narrow areoles along costules, areoles 1/3–2/3 as wide as costule wing, acroscopic arcuate veinlet of areole arising from rachis of segment below base of costa, veinlets outward from areole free, and veinlet of segment 2-forked besides 2 or 3 apex pairs; lamina brown-green, thinly herbaceous when dried, glabrous.

Dense wet forests; 600–700 m. Fujian (Jianou), Guangdong (Lechang, Meixian, Yingde), Guangxi (Xingde, Xiangxian, Yaoshan), S Hunan (Jianghua, Jiangyong, Tongdao), S Jiangxi, ??Zhejiang [S Japan, N Vietnam].

The type of Pteris maclurei is from Guangdong (Meixian).

69. Pteris maclurioides Ching, Acta Bot. Austro Sin. 1: 15. 1983.

岭南凤尾蕨  ling nan feng wei jue

Plants 60–75 cm tall. Rhizome ascending, apex densely clothed with dark brown scales. Fronds clustered; stipe dark chestnut at base, upper part paler, shiny, ca. 30 cm Χ 2 mm, adaxially narrowly grooved; lamina 2- or 3-pinnatipartite, ovate in outline, ca. 45 Χ 25 cm; lateral pinnae 7–9 pairs, subopposite, decumbent, 14–18 Χ 3.5–4 cm, base slightly attenuate-cuneate, deeply pectinately divided to within ca. 3 mm of costa, apex acutely lanceolate caudate (3–5 cm); basal pinnae pair largest, each shortly stalked and with a basiscopic pinnule near base similar to main part of pinna but smaller; segments 15–18 pairs, alternate, oblique, narrowly lanceolate, 13–25 Χ ca. 4 mm, margin not densely serrate, interval equal to width of segment; apical pinnae same as middle lateral pinnae in shape and size, but cuneate and decurrent at base; costae prominent abaxially, below middle part maroon, upper 1/3 part straw-colored, straw-colored and slightly grooved adaxially; veins anastomosing to form a series of narrow areoles along costules, width of areole equal to 1/2 wing width to almost same, occasionally incised, veinlet in segment 2-forked; lamina brown-green, herbaceous when dried, glabrous.

* Open forests; ca. 400 m. Guangdong (Shixing), ?Guangxi [?N Vietnam].

Pteris maclurioides is closely related to P. maclurei but is smaller with pinnae 14–18 Χ 3.5–4 cm, segments 13–25 Χ ca. 4 mm, 3–5 mm apart, costae with ca. 3 mm wide wings, and areoles from 1/2 as wide as wing to as wide.

Ching and S. H. Wu (Fl. Reipubl. Popularis Sin. 3(1): 81. 1990) gave a brief note on “Pteris maclurioides var. tonkinensis” (中越凤尾蕨 zhong yue feng wei jue) from Guangxi and N Vietnam, but as it lacked a Latin description or diagnosis and no type was designated the name was not validly published (Vienna Code, Art. 36.1, 37.1). It differed by the more oblique pinnae, longer basal pinnae, to 30 cm, and the larger segments, 35–45 Χ 5–6 mm, with tapering apices.

70. Pteris wallichiana J. Agardh, Recens. Spec. Pter. 69. 1839.

西南凤尾蕨  xi nan feng wei jue

Rhizome erect, short, thick, 1.5–2 cm in diam., woody, apex with brown scales. Fronds clustered; stipe 60–80 cm, slightly extended at base, 1–2 cm in diam., firm, scabrous, glabrous or with purple-brownish bristles, broadly grooved adaxially; lamina usually 3-partite, lateral branches usually pedately divided, broadly ovate-pentagonal in outline, 70–85 Χ ca. 60 cm; central division 1- or 2-pinnatipartite, 50–70 Χ 20–25 cm, basal stalk 7–10 cm, thicker; lateral pinnae more than 20 pairs, alternate, oblique or decumbent, basal ones shortly stalked, 3–4 cm apart, apical ones sessile, lanceolate, 11–15(–20) Χ 2–2.5(–3.5) cm, base subtruncate to broadly cuneate, deeply pectinately divided to leave narrowly winged costule, margins flat serrate, apex linear-caudate (1–2 cm); basal pinnules slightly shorter; segments 23–30 pairs, alternate, contiguous or acutely incised, 1–2 mm apart, oblique, oblong or broadly lanceolate, 10–13(–18) Χ 3.5–4.5 mm, margins obtusely flat serrate, apex acuminate or obtuse; terminal pinna similar to lateral pinnae, base cuneate with short stalk; costules prominent abaxially, glabrous, slightly grooved adaxially, with short spines; veins conspicuous on both surfaces, oblique, acroscopic vein of segment base anastomosing with basiscopic vein of adjacent segment into an arcuate vein, to form a series of narrow areoles along costules, and with several free simple veinlets arriving at incision in outer edge of arcuate vein, apical 2 or 3-pairs simple, other 2-forked at base, decumbent; lamina pale green or gray-green, firmly herbaceous when dried, subglabrous; glabrous, slightly grooved adaxially.

Chongqing, Guangdong (Xinyi), Guangxi (Huaping), Guizhou (Qingzhen), Hainan (Sanya), Hunan, Jiangxi (Wugongshan), Sichuan, Taiwan, Xizang (Cuona), Yunnan [Bhutan, India, Indonesia, Japan, Kashmir, Laos, Malaysia, Nepal, Philippines, Thailand, Vietnam].

1a. Stipes and rachis with dense purple-brownish hispid hairs ...........................  70b. var. yunnanensis

1b. Stipes and base of rachis glabrous (sometimes stipes slightly hairy).

2a. Segments acuminate ..............................................................................  70a. var. wallichiana

2b. Segments obtuse ..........................................................................................  70c. var. obtusa

70a. Pteris wallichiana var. wallichiana

西南凤尾蕨(原变种)  xi nan feng wei jue (yuan bian zhong)

Campteris wallichiana T. Moore; Litobrochia wallichiana Fιe (1850), not C. Presl (1836); Pteris morrisonicola Hayata.

Plants ca. 1.5 m tall; stipes castaneous-reddish, costa and costules straw-colored; segment apices usually acuminate; segments 3.5–4.5 mm wide.

Forests within valleys; 800–2600 m. Chongqing, Guangdong (Xinyi), Guangxi (Huaping), Guizhou (Qingzhen), Hainan (Sanya), Hunan, Sichuan (Emeishan, Hongxi, Hanyuan), Taiwan, Xizang (Cuona), Yunnan (Jingdong, Yangbi, Yingjiang) [Bhutan, India, Indonesia, Japan, Laos, Malaysia, Nepal, Philippines, Thailand, Vietnam].

The type of Pteris wallichiana is from NW India (Kumaon).

70b. Pteris wallichiana var. yunnanensis (H. Christ) Ching & S. H. Wu, Fl. Reipubl. Popularis Sin. 3(1): 83. 1990.

云南凤尾蕨  yun nan feng wei jue

Pteris yunnanensis H. Christ, Bull. Herb. Boissier 6: 957. 1898; P. tomentella Handel-Mazzetti.

Plants up to 2 m tall. Stipes and rachises with dense purple-brown multicellular hispid hairs; costules abaxially slightly purple-brown and with multicellular hispid hairs.

* Mountain forests, along valleys; 500–2300 m. S and W Yunnan.

70c. Pteris wallichiana var. obtusa S. H. Wu, Acta Bot. Austro Sin. 1: 15. 1983.

圆头凤尾蕨  yuan tou feng wei jue

Plants ca. 1 m tall. Stipes castaneous-reddish, slightly scabrous; segments obtuse, 1.5–1.8 cm wide.

* Forests. Jiangxi (Wugongshan), Sichuan (Emeishan), S Yunnan.

The type of Pteris wallichiana var. obtusa is from Jiangxi.

71. Pteris austrosinica (Ching) Ching, Acta Phytotax. Sin. 10: 302. 1965.

华南凤尾蕨  hua nan feng wei jue

Pteris wallichiana J. Agardh var. austrosinica Ching, Bull. Dept. Biol. Sun Yatsen Univ. 6. 27. 1933.

Plants ca. 1.5 m tall. Rhizome erect, short, thick, ca. 2 cm in diam., woody, apex with brown scales. Fronds clustered; stipe pale castaneous, up to 1 m, ca. 1.5 cm in diam., firm, glabrous, broadly grooved adaxially; rachis pale chestnut or reddish brown, sometimes brown straw-colored, with sparse reddish brown multicellular hairs, narrowly grooved adaxially; lamina usually 3-pinnatipartite, pentagonal-broadly ovate in outline, 80–100 Χ ca. 70 cm, central branch a long ovate column, 60–70 cm, middle ones ca. 25 cm wide, stalked (8–10 cm), lateral branches smaller, usually again divided; lateral pinnules 14–20 pairs, alternate, decumbent, sessile or slightly shortly stalked, basal several pairs slightly shorter, ca. 1.5 cm apart, middle pinnules lanceolate, 15–20 Χ 3–4 cm, base broadly cuneate, subsymmetric, deeply pectinately divided leaving broadly winged costule, apex shortly linear-caudate; segments 17–25 pairs, alternate, sinuses obtuse-acute, 3–5 mm wide, slightly decumbent, falcate-lanceolate, 20–25 Χ ca. 3 mm, basally enlarged, apex shortly acuminate, sterile apex obtusely dentate; terminal pinnules similar to median lateral pinnules, stalked (ca. 1 cm); costules straw-colored, glabrous, with short spines on both sides of adaxial groove; veins conspicuous, oblique, anastomosing to form a series of narrow areoles along costules, several simple veinlets reaching incision in outer edge of arcuate vein, and veinlet free outward from areole, and basal veinlet of segment 2-forked at base; lamina brown-green, papery when dried, below with brown slender multicellular hairs.

* Dense forests within wet valleys; 400–1000 m. Guangdong, Guangxi, Jiangxi.

The type of Pteris austrosinica is from Guangxi (Yaoshan).

72. Pteris occidentalisinica Ching, Acta Bot. Austro Sin. 1: 15. 1983.

华西凤尾蕨  hua xi feng wei jue

Plants to 2 m tall. Rhizome not seen. Stipe basally dark brown, ca. 5 mm in diam.; lamina usually 3-branched from apex of stipe, lobes deeply 3-pinnatifid; pinnae oblong-lanceolate, ca. 80 cm, middle ones ca. 40 cm; pinnules more than 20 pairs, alternate or subopposite, oblique, contiguous with each other; middle ones longest, lanceolate, 20–25 Χ ca. 4.5 cm, base broadly cuneate, apex long acuminate (ca. 2 mm); segments opposite or alternate, 2–4 mm apart, obliquely patent, 20–25 Χ ca. 5 mm, apex obtuse or bluntly acute, flat serrate; costae straw-colored, slightly brown; costules abaxially straw-colored, with short spines along both sides of adaxial groove; veins conspicuous on both surfaces, anastomosing to form a series of narrow areoles along costules; lamina pale green when dried, thinly papery, subglabrous.

* Forests; ca. 2400 m. S Sichuan (Laibo).

73. Pteris taiwanensis Ching, Acta Bot. Austro Sin. 1: 16. 1983.

台湾凤尾蕨  tai wan feng wei jue

Plants ca. 2 m tall. Rhizome not seen. Stipe light straw-colored, glabrous, adaxially grooved; lamina 3-pinnatipartite, broadly ovate in outline, 80–100 Χ 60–70 cm; segments of central branch oblong, up to 100 Χ 30–35 cm, lateral branches 2, shorter than central one; pinnules more than 20 pairs, opposite or subopposite, lower ones stipitate, upper sessile, lanceolate, 20–30 Χ 5–7 cm, base broadly cuneate, pectinately divided to leave narrowly winged costule, apex long acuminate, serrulate; base of pinnules with several pairs of free segments; segments 30–35 pairs, alternate, subspreading, 5–8 mm apart, linear, 25–35 Χ ca. 3 mm, base slightly enlarged, margins flat serrate, apex acuminate; costules abaxially convex, straw-colored, sparsely multicellular brown strigose or glabrous, adaxially flat grooved, with short spines on both sides; veinlets conspicuous abaxially, flared, anastomosing to form a series of narrow areoles along costules; lamina brown-green, herbaceous when dried, brown multicellular strigose on both surfaces; rachises straw-colored, grooved adaxially.

* About 800 m. Taiwan (Wulai).

74. Pteris finotii H. Christ, J. Bot. (Morot) 19: 72–73. 1905.

疏裂凤尾蕨  shu lie feng wei jue

Pteris finotii var. obtusa Tagawa.

Plants to 2.5 m tall. Rhizome erect, short and thick, ca. 2 cm in diam., apex with bright brown scales. Fronds clustered; stipe dark straw-colored to dark brown, ca. 100 cm, 7–12 mm in diam., glabrous; rachis straw-colored, glabrous; lamina with 3 pinnatisect divisions, ovate-triangular in outline, to more than 100 Χ 60 cm, central division ovate, column, to more than 80 Χ 35–40 cm, stalk 6–10 cm; two lateral branches smaller; lateral pinnules 6–8 pairs, opposite or subalternate, oblique, stalked (ca. 1 cm), upper ones sessile, broadly lanceolate, 20–27 Χ 6–10 cm, base rounded-truncate, apex caudate (4–8 cm); segments 13–20 pairs, alternate or basal pairs subopposite, linear-lanceolate, 25–70 Χ 4–7 mm, basally slightly enlarged, sterile margins slightly serrate, apex obtuse or acuminate; sterile pinnules with wider segments; terminal pinnule triangular, 20–25 Χ 15–18 cm, stalked (1.5–2 cm), base rounded-cuneate, along costule narrowly cuneate, pectinately divided with broadly winged costa, margins flat, obtusely serrate, apex linear-lanceolate (4–5 cm); costules prominent abaxially, straw-colored, glabrous, grooved adaxially, with inconspicuous spines or without spines; veinlets slender, conspicuous only abaxially, anastomosing to form a series of narrow areoles along costules and midvein, areoles along costules long and narrow, areoles along midvein irregularly polygonal, veins of segments free; lamina gray-green, herbaceous when dried, glabrous.

Forests, by small streams; 100–500 m. Guangdong, Hainan, S Yunnan (Hekou, Xishuangbanna) [N Vietnam].

The type of Pteris finotii is from N Vietnam.

75. Pteris tripartita Swartz in Schrader, J. Bot. 1800[2]: 67. 1801.

三叉凤尾蕨  san cha feng wei jue

Litobrochia marginata (Bory) C. Presl; L. tripartita (Swartz) C. Presl; Pteris marginata Bory; P. tripartita Swartz.

Plants more than 2 m tall. Rhizome erect, short, ca. 2 cm in diam., apex with gray-brown scales. Fronds clustered; stipe dark brown or straw-colored, 100–150 cm, ca. 1 cm in diam., glabrous; lamina with 3 1- or 2-pinnatipartite divisions, broadly ovate in outline, 80–100 Χ ca. 70 cm, central division oblong in outline, 80–100 Χ 25–30 cm, stalked (10–12 cm), lateral branches smaller, and basal ones usually again divided into 2 or 3 forks; pinnules 20–30 pairs, alternate or subopposite, oblique, shortly stalked, upper ones sessile; pinnules at base and apex slightly shorter, middle pinnules lanceolate, 15–21 Χ 4–5 cm, base rounded-truncate, pectinately divided and forming two broad wings of costule, apex acute and lanceolate-caudate (3–4 cm); segments 14–25 pairs, alternate, slightly oblique, 2–5 mm apart, falcate-lanceolate, 10–30 Χ 4–6 mm, base slightly enlarged, sterile margins serrate, apex mucronate or obtuse; terminal pinnule similar to middle lateral pinnae but stalked; costules abaxially convex, straw-colored, glabrous, shallowly grooved adaxially, with short spines on both sides; veinlets slender, conspicuous only abaxially, veinlets from adjacent segment anastomosing to form a series of narrow areoles along costa, veinlets further anastomosing to form a series of narrow polygonal areoles along costules; lamina brown-green, thinly papery when dried, glabrescent.

Guangdong, Guangxi (Baise, Napo), Hainan (Lingshui), Hunan (Shimen), Taiwan [India, Indonesia, Malaysia, Philippines, Sri Lanka, Thailand, Vietnam; Africa, Australia, Madagascar, Pacific islands (Polynesia), South America (Brazil)].

Hainan and Taiwan are the northernmost limits for the distribution of this pantropical species.

6. ANOGRAMMA Link, Fil. Spec. 137. 1841.

翠蕨属  cui jue shu

Zhang Gangmin (张钢民); Tom A. Ranker

Ferns annual, small, terrestrial or on rocks. Rhizomes short, poorly developed, sparsely scaly; scales small, fibrous. Fronds many, monomorphic, tufted; stipe chestnut-colored or chestnut brown, slender, rounded abaxially, grooved adaxially, glabrous; lamina ovate, ovate-deltoid to ovate-lanceolate, or lanceolate, 1–3-pinnate, thinly herbaceous or submembranous, both surfaces usually glabrous. Ultimate pinnules or segments small, ovate-elliptic, spatulate, or obovate, margin entire, or shallowly lobed at apex. Veins free, forked, 1 veinlet per segment, far from frond margins. Indusia absent. Sori borne along veinlets, paraphyses absent. Spores tetrahedral, somewhat ridged on surface. Gametophytes still alive for a long time after sporophytes are produced. x = 29, 58.

About six species: worldwide tropics, subtropics, and European region along Atlantic Ocean; two species in China.

1a.       Ultimate pinnules or segments obovate or obdeltoid (pinnae fan-shaped when lamina pinnate), apex shallowly lobed ............................................................................................  1. A. leptophylla

1b.       Ultimate pinnules or segments ovate-elliptic or spatulate, apex obtuse or mucronulate  2. A. microphylla

1. Anogramma leptophylla (Linnaeus) Link, Fil. Spec. 137. 1841.

薄叶翠蕨  bo ye cui jue

Polypodium leptophyllum Linnaeus, Sp. Pl. 2: 1092. 1753; Acrostichum leptophyllum (Linnaeus) Lamarck & Candolle; Grammitis leptophylla (Linnaeus) Swartz; Gymnogramma leptophylla (Linnaeus) Desvaux.

Rhizomes small, short; scales few, pellucid. Fronds tufted; stipe chestnut brown, 2–6 cm Χ 0.5–1 mm, glabrous or occasionally with 1 or 2 pellucid and jointed long hairs; lamina yellowish green, ovate-deltoid to ovate-lanceolate, 4–8 Χ 1.5–3 cm, submembranous when dry, glabrous, 2-pinnate to 2-pinnate-pinnatifid. Basal pinnae usually larger than adjacent ones; distal pinnae gradually reduced. Ultimate pinnules or segments obovate or obdeltoid, base cuneate and decurrent, apex shallowly lobed. Veins visible on both surfaces. 2n = 58.

Evergreen broad-leaved forests, stream banks; below 2900 m. Taiwan, Yunnan [India, Vietnam; Africa, Australia, Europe, Pacific islands, South America].

Anogramma leptophylla is variable in frond shape. The laminae of young plants can be orbicular-fan-shaped, bifurcate, or tripartite, or in 1-pinnate individuals narrowly ovate, with all the pinnae fan-shaped and shallowly lobed at apex.

2. Anogramma microphylla (Hooker) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 259. 1899.

翠蕨  cui jue

Gymnogramma microphylla Hooker, Icon. Pl. 10: t. 916. 1854; Cerosora microphylla (Hooker) R. M. Tryon; Grammitis microphylla (Hooker) Beddome.

Rhizomes short, together with stipe base sparsely scaly; scales blackish, less than 0.8 mm. Fronds many, clustered; stipe chestnut brown, 2–10 cm Χ ca. 0.6 mm, glabrous; lamina green, ovate-deltoid or ovate-lanceolate, 2–5 Χ 2–4 cm, thinly herbaceous when dry, both surfaces glabrous, 2-pinnate-pinnatifid to 3-pinnate. Pinnae 5–7 pairs, basal pair largest, ovate-deltoid, 1–2 Χ 0.7–1.2 cm, inequilateral, shortly stalked. Pinnules anatropous, with winged short stalks. Ultimate pinnules or segments elliptic or spatulate, base long cuneate and decurrent to costules, margin entire, apex obtuse or mucronulate, rarely emarginate. Veins visible on both surfaces. 2n = 116.

Montane gorges, on rocks, in rock crevices; 1100–2900 m. E Guangxi, Guizhou, Taiwan, Yunnan [Bhutan, India, Myanmar, Nepal].

7. TAENITIS Willdenow ex Sprengel, Anleit. Kennt. Gew. 3: 374. 1804.

竹叶蕨属  zhu ye jue shu

Dong Shiyong (董仕勇); Masahiro Kato

Plants terrestrial. Rhizome creeping, apex covered with dark, rigid bristles; bristles with a single row of cells at apex and a few rows of cells at base. Fronds monomorphic or dimorphic; stipes with 1, 2, or 4 vascular bundles near base, adaxially grooved; lamina simple or 1-pinnate, with terminal pinnae similar to lateral pinnae; pinnae simple, entire, thickly papery to leathery, glabrous; veins reticulate, areoles without included free veinlets. Sori linear and forming a narrow longitudinal band between midrib and margin, or borne irregularly along veins, or converging to orbicular or oblong sori, or in long inframarginal grooves, or acrostichoid on abaxial surface of contracted fertile pinnae, exindusiate; paraphyses abundant, multicellular. Spores trilete, tetrahedral-globose, with tubercles or rodlets. n = 44, 108, 110, 114.

About 15 species: from Sri Lanka and S India to S China, through Malaysia and Indonesia to N Queensland and Fiji; one species in China.

The taxonomic state and position of this group has been disputed. Ching (1978) treated it as a monogeneric family and thought it related to Lindsaeaceae. In other later systems, it was treated as a genus related to Syngramma J. Smith within the Pteridaceae, a view accepted by Smith et al. (2006) and Christenhusz et al. (2010).

1. Taenitis blechnoides (Willdenow) Swartz, Syn. Fil. 24, 220. 1806.

竹叶蕨  zhu ye jue

Pteris blechnoides Willdenow, Phytographia, 13. 1794.

Rhizome long creeping, 1–2 mm in diam., with bristles at apex; bristles dark brown, stiff, 2–3 mm. Fronds distant, 2–4 cm apart; stipe castaneous at base, straw-colored or brown distally, 40–70 cm, adaxially grooved, glabrous; lamina 1-pinnate, slightly dimorphic, 20–40 Χ 20–30 cm, with 3–8 pairs of lateral pinnae, papery to leathery, glabrous; sterile pinnae 15–25 Χ 2.5–4 cm, lanceolate or linear-lanceolate, shortly stalked, base narrowly cuneate, margin entire, apex acuminate; costae abaxially distinctly raised, adaxially flat or slightly sunken; fertile pinnae narrower, 13–25 Χ 1.2–3 cm. Veins fully reticulate without included veinlets. Sori linear and forming a narrow longitudinal band between midrib and margin, rarely interrupted, 1–1.5 mm wide.

Terrestrial in forests; 400–1000 m. Hainan [Cambodia, India, Indonesia, Laos, Malaysia, Sri Lanka, Thailand, Vietnam; Pacific islands].

8. PITYROGRAMMA Link, Handbuch 3: 19. 1833.

粉叶蕨属  fen ye jue shu

Zhang Gangmin (张钢民); Tom A. Ranker

Plants terrestrial. Rhizomes short, erect or ascending, dictyostelic; scales reddish brown, thin, lanceolate, margins entire. Fronds monomorphic, clustered; stipe purplish black, lustrous, rounded proximally, upward with single longitudinal groove adaxially, base scaly, glabrous distally; lamina 2- or 3-pinnate, ovate to oblong in outline, apex acuminate. Pinnae many, oblique, lanceolate, somewhat stalked, 2- or 3-pinnate, herbaceous to somewhat leathery, abaxially white or yellow farinose, adaxially glabrous, base subequilateral, apex acuminate. Pinnules many, anatropous, base inequilateral, usually adnate to costae, margins serrate. Veins of ultimate segments free, obscure, pinnately branched or not. Indusia absent. Sori along veins but not to vein tips, without paraphyses. Spores dark colored, globose-tetrahedral, perispore reticulate with irregular ridges. x = 29 (30).

About 20 species: tropical Africa, America, and Asia; one species in China.

1. Pityrogramma calomelanos (Linnaeus) Link, Handbuch 3: 20. 1833 [“calomelas”].

粉叶蕨  fen ye jue

Acrostichum calomelanos Linnaeus, Sp. Pl. 2: 1072. 1753; Ceratopteris calomelanos (Linnaeus) Underwood; Gymnogramma calomelanos (Linnaeus) Kaulfuss; Neurogramma calomelanos (Linnaeus) Diels.

Stipe 40–50 cm; lamina 2-pinnate, long oblong or oblong-lanceolate, 15–40 Χ 10–20 cm, thickly papery when dry, abaxially densely white farinose, adaxially grayish green, glabrous, base broadly cuneate, apex acuminate; rachis and costae purple-black, lustrous, glabrous, grooved adaxially; pinnae 16–20 pairs, oblique, with short, winged stalks; basal pair of pinnae not shortened, lanceolate, 10–15 Χ 2–5 cm, base slightly inequilateral; pinnules (or segments) 16–18 pairs, deltoid, ovate-lanceolate, or lanceolate, 1.1–1.4 Χ ca. 0.5 cm, base inequilateral, somewhat decurrent to costae, apex acute or acuminate; proximal pinnule margins serrate or lobed; lobes serrate or entire, with only 1 or 2 teeth at apex. Sori nearly confluent throughout abaxial pinnule surface when mature. 2n = 240.

Forest margins, stream banks; below 600 m. Hainan, Taiwan, S Yunnan (Luchun) [Cambodia, Laos, Vietnam; Africa, South America].

Pityrogramma calomelanos is a cultivated ornamental plant of New World origins, which has become widely naturalized in the Old World.

9. ONYCHIUM Kaulfuss, Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 45. 1820.

金粉蕨属  jin fen jue shu

Zhang Gangmin (张钢民); George A. Yatskievych

Plants terrestrial or less commonly on rocks. Rhizomes long creeping or rarely short and decumbent, siphonostelic, scaly, scales concolorous, light brown to reddish brown, opaque to somewhat translucent, nonclathrate, linear-lanceolate to ovate-lanceolate, margins entire. Fronds monomorphic to somewhat dimorphic, widely or closely spaced. Stipe straw-colored above a usually darker proximal portion, occasionally uniformly straw-colored or reddish brown to dark brown (especially abaxially), glabrous above scaly base, with (1 or)2 vascular bundles at base, adaxially grooved. Lamina ovate-deltate or ovate-lanceolate, less often elongate-lanceolate, herbaceous or papery, finely 2–5-pinnate-pinnatifid, rarely 2-pinnate, glabrous or nearly so or fertile segments yellow-farinose abaxially; rachis and costae usually grooved adaxially. Pinnae alternate. Ultimate segments or lobes narrow and small, lanceolate to oblong-lanceolate or narrowly oblong, base cuneate and often decurrent, apex acute or acuminate. Veins simple or pinnately branched, free, slightly thickened, sometimes raised abaxially, in fertile segments veinlets connected by an inframarginal commisural vein. Sori borne along commisural veins, linear. False indusia usually well developed, membranous, continuous but interrupted at segment apex and base, reaching to midvein or nearly so (narrow in Onychium tenuifrons), margins entire, slightly undulate, or less commonly erose. Spores trilete, globose-tetrahedral, perispore coarsely tuberculate-reticulate on distal face, proximal face tuberculate and with prominent, coarse ridges parallel to equatorial flange. x = 29.

About ten species: tropical and subtropical Africa (one species), Asia; eight species in China.

The taxonomy of Onychium is poorly understood and is complicated by polyploidy and apomixis in several of the taxa. The descriptions below contrast sterile and fertile fronds, but it should be noted that plants of some species often also produce intermediate fronds that are fertile distally and sterile proximally. Small specimens of Onychium sometimes are confused with dissected-leaved Asplenium species; collectors should examine rhizome scales (clathrate in Asplenium), sori (true indusia lateral on veins in Asplenium), and spores (monolete, mostly ellipsoidal, and lacking an equatorial flange in Asplenium).

1a.       Longest sori 1–2 cm or more; fertile lamina with yellow farina abaxially; rhizome scales linear-lancolate    1. O. siliculosum

1b.       Sori usually all less than 1 cm; fertile lamina not farinose; rhizome scales lanceolate to ovate-lanceolate.

2a.       Rhizomes short, decumbent; fronds closely spaced, ± clustered; scales of stipe base pale brown; margins of ultimate segments ± serrulate toward tip; margins of false indusia erose ....  2. O. tenuifrons

2b.       Rhizomes long creeping (sometimes shortly creeping in O. angustifrons); fronds mostly widely spaced; scales of stipe base reddish brown or dark brown; margins of ultimate segments entire; margins of false indusia entire or slightly undulate.

3a.       Lamina less than 10 cm wide, narrowly deltate-lanceolate, 2–3-pinnate.

4a.       Lamina of fertile frond 4–10 cm wide, 3-pinnate ..................................  7. O. moupinense

4b.       Lamina of fertile frond 1–3 cm wide, 2-pinnate ...................................  8. O. angustifrons

3b.       Lamina 12–30 cm or more wide, ovate to ovate-deltate, 3–5-pinnate-pinnatifid.

5a.       Lamina thinly herbaceous; ultimate segments crowded, sometimes overlapping  6. O. plumosum

5b.       Lamina ± papery; ultimate segments mostly well separated, not overlapping.

6a.       Lamina ovate-deltate to ovate-lanceolate; ultimate segments crowded, sometimes overlapping       5. O. japonicum

6b.       Lamina usually broadly ovate or elliptic; ultimate segments not crowded.

7a.       Stipe bases black; pinna apices acuminate; fertile segment apices acute to slightly acuminate; lamina green to grayish green ..........................................................  3. O. cryptogrammoides

7b.       Stipe bases dark brown; pinna apices caudate; fertile segment apices acuminate; lamina dark green ...........................................................................................................  4. O. tibeticum

1. Onychium siliculosum (Desvaux) C. Christensen, Ind. Filic., fasc. 8: 469. 1906.

金粉蕨  jin fen jue

Pteris siliculosa Desvaux, Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 5: 324. 1811; Allosorus auratus (Kaulfuss) C. Presl; Acrostichum viviparum Cavanilles (1802, non Linnaeus f., 1782); Cryptogramma aurata (Kaulfuss) Prantl; Onychium auratum Kaulfuss; O. chrysocarpum (Hooker & Greville) C.Christensen; O. tenue H. Christ; Pteris aurata (Kaulfuss) Mettenius; P. chrysocarpa Hooker & Greville.

Rhizomes short, erect or ascending; scales dark brown, linear-lanceolate. Fronds somewhat dimorphic (fertile fronds with longer, broader segments), clustered. Stipe (5–)12–30 cm Χ 1–5 mm, straw-colored above a brown base, brown color sometimes extending abaxially to rachis. Lamina of sterile fronds ovate-deltate to narrowly ovate-deltate, 2- or 3-pinnate-pinnatifid, apex acuminate, papery when dry, green to grayish green, not farinose. Lamina of fertile fronds ovate-lanceolate to oblong-ovate, (5–)15–35 Χ 9–18 cm, 3- or 4-pinnate or rarely 2-pinnate-pinnatifid., with long, linear, simple, terminal segments; lateral pinnae (4–)10–15 pairs, basal pair slightly largest, oblong-lanceolate or deltate, with stalks 3–6 mm. Ultimate segments of sterile lamina sometimes closely spaced but not overlapping, sessile, less than 1 mm wide at base, slightly wedge-shaped, apex wider and with 1–3 small teeth or narrowly notched, slightly thickened or recurved. Ultimate pinnules of fertile lamina not overlapping, linear when young, 5–15 Χ 1.5–2 mm, wider at maturity (2–3 mm), stalks 2–3 mm, base cuneate, apex acuminate or acute. Longest sori 10–20 mm or more, intermixed with bright yellow farina. False indusia linear, extending nearly to midvein, with entire margins. 2n = 58.

Rock crevices of dry valleys; 100–1500 m. E Hainan, Taiwan, Xizang, Yunnan [Bangladesh, Bhutan, Cambodia, India, Laos, Malaysia, Myanmar, Nepal, Thailand, Vietnam; Oceania].

2. Onychium tenuifrons Ching, Lingnan Sci. J. 8: 500. 1934.

蚀盖金粉蕨  shi gai jin fen jue

Onychium japonicum (Thunberg) Kunze var. delavayi H. Christ.

Rhizomes short, decumbent; scales light brown, narrowly lanceolate. Fronds slightly dimorphic (fertile fronds with narrower segments than sterile ones), closely spaced, often appearing clustered. Stipe straw-colored, sometimes slightly darkened at very base, 7–20 cm Χ ca. 2 mm (longer on fertile than sterile fronds). Lamina of sterile fronds ovate to narrowly ovate or elliptic-lanceolate, 3-pinnate-pinnatifid, thinly herbaceous. Lamina of fertile fronds elliptic-lanceolate, 15–25 Χ 5–10 cm, 3- or 4-pinnate, apex acuminate, firmly papery when dry, green, not farinose; lateral pinnae 8–10 pairs; longest pinnae lanceolate, stalks 4–10 mm. Ultimate pinnules (segments) of sterile lamina crowded but mostly not overlapping, obovate or oblong, with sharp teeth at apex, 1 veinlet per tooth, margins often slightly thickened or somewhat cartilaginous. Ultimate pinnules (segments) of fertile lamina not overlapping, oblong-linear or lanceolate, 1–6 Χ 0.4–0.9 mm, sessile or decurrent on costule, apex acuminate or acute. Sori 3–5 mm, glabrous. False indusia linear to narrowly oblong, not approaching mdvein, with erose margins. 2n = 87.

Forest margins, under shrubs; 100–2700 m. W Guizhou, Sichuan, Yunnan [Bhutan, India, N Myanmar, Nepal, Philippines].

Onychium tenuifrons is similar to O. japonicum, but differs in having clustered fronds, lighter and thinner rhizome scales, and false indusia with erose margins.

3. Onychium cryptogrammoides H. Christ, Notul. Syst. (Paris) 1: 52–53. 1909.

黑足金粉蕨  hei zu jin fen jue

Onychium japonicum (Thunberg) Kunze var. intermedia C.B. Clarke; O. japonicum var. multisecta C.B. Clarke.

Rhizomes long creeping; scales dark brown, lanceolate. Fronds monomorphic, closely or widely spaced. Stipe (15–)20–50 cm Χ 2–3 mm, straw-colored above a black base. Lamina ovate to ovate deltate or often broadly ovate, (12–)20–38 Χ 10–26 cm, apex acuminate, finely 4-pinnate-pinnatifid, thinly papery when dry, green to grayish green, not farinose. Lateral pinnae 10–14 pairs, basal pair largest, ovate-deltate, apex acuminate, 10–25 Χ 5–14 cm, stalked to ca. 1 cm. Ultimate pinnules (segments) not overlapping, linear to narrowly oblanceolate, narrowly oblong, or narrowly elliptic, 2–5 Χ0.5–1.5 mm wide, margins often cartilaginous, enitire or with 1–2 sharp teeth, base cuneate, decurrent, apex acute to slightly-acuminate. Sori 1–3 mm. False indusia linear to oblong, extending nearly to midvein, entire.. 2n = 87, 116.

Open woods, stream banks, valleys, often forming large, dense, clonal colonies; 1200–3500 m. S Gansu, Guizhou, Sichuan, Taiwan, S Xizang, NW Yunnan [Bhutan, Cambodia, India, Laos, Myanmar, Nepal, Thailand, Vietnam].

The Onychium cryptogrammoides group is a polyploid complex that requires further study. Onychium cryptogrammoides resembles O. japonicum, but differs in its black stipe bases, finely divided lamina, and more slender costae and costules. The two species have different distributions. The name O. contiguum Wall. ex C. Hope has been applied to this taxon in much of the literature on Asian pteridophytes, but that epithet is a nom. superfl. for the taxon here treated as O. japonicum var. lucidum, as the basionym Leptostegia lucida D. Don was cited in synonymy in the protologue.

4. Onychium tibeticum Ching & S.K. Wu in C.Y. Wu, Fl. Xizang. 1: 77–78. 1983.

西藏金粉蕨  xi zang jin fen jue

Rhizomes long creeping; scales brown or reddish brown, lanceolate. Fronds monomorphic, widely spaced. Stipe 10–28 cm, straw-colored above a dark brown base. Lamina elliptic, 24–28 Χ ca. 15 cm, base cuneate, apex acuminate, 3-pinnate-pinnatifid to 4-pinnate, thinly papery to papery when dry, dark green, not farinose. Lateral pinnae 8–10 pairs, alternate, crowded, basal pair similar to but larger than adjacent pinnae, elliptic, 13–16 Χ 3–5 cm, stalked to ca. 1 cm, base cuneate, apex caudate. Ultimate sterile pinnules (segments) similar to but shorter than fertile ones. Ultimate fertile pinnules (segments) not overlapping, linear to elliptic-lanceolate, 5–7(–10) Χ ca. 1 mm, apex acuminate. Sori restricted to median portions of ultimate fertile pinnules (segments), 2–5 mm. False indusia oblong to narrowly oblong, extending nearly to midvein, entire.

* Banks of ditches under trees; 2000–2300 m. Xizang.

Onychium tibeticum is similar to O. cryptogrammoides, but differs in having a brown stipe base, caudate pinna apices, and longer ultimatesegments. However, some botanists consider it to represent a mere variant of O. japonicum that was described based on a specimen with the frond intermediate between sterile and fertile morphologies.

5. Onychium japonicum (Thunberg) Kunze, Bot Zeitung (Berlin) 6:. 507. 1848.

野雉尾金粉蕨  ye zhi wei jin fen jue

Rhizomes long creeping; scales brown or reddish brown, lanceolate. Fronds slightly dimorphic (fertile fronds slightly more coarsely divided than sterile ones), mostly widely spaced. Stipe 7–45(–60) cm, 2–4 mm in diam., straw-colored throughout or with a, brown, to dark brown base, dark color sometimes extending to rachis abaxially. Lamina ovate-deltate or narrowly ovate, (9–)20–48 Χ 10–20 cm, apex acuminate, finely 2-pinnate-pinnatifid to 4-pinnate, firmly papery when dry, green to dark green or grayish green, not farinose. Lateral pinnae 10–15 pairs, deltate-lanceolate or oblong-lanceolate; basal pair largest, 9–20 Χ 5–10 cm, stalked to ca. 1–2 cm, apex acuminate. Ultimate fertile pinnules (segments) crowded, sometimes overlapping, linear to linear-lanceolate, 4–10 Χ 1–2 mm, apex acute. Ultimate sterile pinnules (segments) similar to fertile ones but mostly somewhat shorter, apex shortly acuminate. Sori (2–)3–6 mm. False indusia extending nearly to midvein, entire. 2n = 58, 87, 116, 174.

Streambanks, roadsides, forest margins, mountain slopes; 200–2400 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, W Hebei, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Shandong, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Indonesia, Japan, Korea, Myanmar, Nepal, Pakistan, Philippines, Vietnam; Pacific islands].

The O. japonicum/O. moupinense group is a polyploid complex whose taxonomy requires further study. Although the two varieties reported for China have been treated as separate species by some botanists, they exhibit a great deal of morphological overlap and perhaps should not be formally recognized.

1a. Plants (16–)40–60 cm tall; stipe straw-colored above a sometimes brown to dark brown base; ultimate pinnules or segments usually 5–7 mm ..............................................................  5a. var. japonicum

1b. Plants (39–)45–80 cm tall; stipe brown to dark brown above base, at least abaxially, dark color sometimes extending to rachis; ultimate pinnules or segments mostly 7–10 mm .....  5b. var. lucidum

5a. Onychium japonicum var. japonicum

野雉尾金粉蕨(原变种)  ye zhi wei jin fen jue (yuan bian zhong)

Trichomanes japonicum Thunberg, Fl. Jap. 340. 1784; Caenopteris japonica (Thunberg) Thunberg; Cryptogramma japonica (Thunberg) Prantl; Darea japonica (Thunberg) Willdenow; Onychium chinense (Desv.) Fιe; Phorolobus chinensis Desvaux; Pteris japonica (Thunberg) Mettenius.

Plants (16–)40–60 cm tall. Rhizomes ca. 3 mm in diam. Stipe straw-colored, at most brown proximally. Ultimate pinnules or segments usually 5–7 mm.

By streams, roadsides, forest margins; 200–2200 m. Anhui, Fujian, S Gansu, Guangxi, Guizhou, W Hebei (Xinle), Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Shandong, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Thailand; Oceania].

5b. Onychium japonicum var. lucidum (D. Don) H. Christ, Bull. Soc. Bot. France Mem. 1: 60. 1905.

栗柄金粉蕨  li bing jin fen jue

Leptostegia lucida D. Don, Prod. Fl. Nepal. 14–15. 1825; Onychium lucidum (D. Don) Sprengel; Onychium contiguum Wallich ex C. Hope (nom. superfl.).

Plants40–80 cm tall or more. Rhizomes ca. 4 mm in diam. Stipe usually brown to chestnut-brown. Ultimate pinnules or segments mostly 7–10 mm.

Roadsides, forest margins, mountain slopes; 200–2400 m. Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hubei, Jiangxi, Shaanxi, Sichuan, Xizang, Yunnan, Zhejiang [Bhutan, India, Myanmar, Nepal, Pakistan, Vietnam].

6. Onychium plumosum Ching, Lingnan Sci. J. 13: 499–500. 1934

繁羽金粉蕨  fan yu jin fen jue

Onychium japonicum var. parvisorum R. Bonaparte

Rhizomes stout, decumbent; scales dark brown, shiny, subulate-lanceolate. Fronds monomorphic or nearly so (fully fertile fronds slightly more coarsely divided than entirely sterile ones, but many fronds intermediate), closely spaced, often appearing clustered. Stipe straw-colored above a brown base, (12–)25–30 cm Χ ca. 4 mm. Lamina deltate-oblong or deltate-ovate, 35–60 Χ 20–30 cm, apex acuminate, 4–5-pinnate-pinnatifid, thinly  herbaceous when dry, bright green to grayish green, not farinose. Lateral pinnae 9–15 pairs, basal pair largest, (8–)15–22 Χ 6–16 cm, stalked to ca. 1.5 cm, apex acuminate. Pinnules.deltate or elongate-ovate, stalked;, stalks with narrow wings. Ultimate segments crowded, sometimes overlapping, linear to narrowly oblanceolate, 2–3(–7) Χ ca. 1 mm, apex acuminate. Sori 1–3 mm. False indusia extending nearly to midvein, margins entire. 2n = 116.

Forests and ditches; 1200–2800 m. SW Sichuan, NW Yunnan [Bhutan, India, Nepal, Thailand].

7. Onychium moupinense Ching, Lingnan Sci. J. 8: 500. 1934.

木坪金粉蕨  mu ping jin fen jue

Rhizomes long creeping; scales dark brown, subulate-lanceolate to lanceolate. Fronds monomorphic or slightly dimorphic (fertile laminae somewhat wider and more finely divided than sterile ones), closely to widely spaced. Stipe straw-colored above a sometimes brown base, 5–32 cm Χ ca. 1 mm. Lamina of sterile fronds narrowly lanceolate to lanceolate, 10–15 Χ 2.5–3.5 cm, 2-pinnate or 2-pinnate-pinnatifid. Lamina of fertile fronds larger than sterile lamina, lanceolate to ovate-lanceolate, 14–32 Χ 6–10 cm, apex long attenuate to somewhat caudate, 3-pinnate, papery when dry, green to grayish green, not farinose. Lateral pinnae 8–15 pairs, obliquely patent or curved upward, basal pair largest, lanceolate or ovate-lanceolate, 3–8 Χ 2–3 cm, stalked to 5–10 mm, apex elongate-acuminate or caudate. Ultimate pinnules (segments) of sterile fronds obliquely ovate, apex sharply serrate. Ultimate (segments) of fertile fronds linear, 3–8 Χ ca. 1.5 mm, base narrowly winged and decurrent on costae, apex acuminate. Sori 2–7 mm. False indusia oblong to linear, entire. 2n = 87, 174.

Rock crevices under shrubs or in broad-leaved forests, shaded moist places; 500–1900 m. Hubei, Shaanxi, Sichuan, Yunnan [?Japan, Nepal].

As noted under O. japonicum, the O. japonicum/O. moupinense group is a polyploid complex that requires further study. Two varieties are tentatively accepted here, both in China.

1a. Fronds subdimorphic; sterile lamina lanceolate; fertile lamina ovate-lanceolate; ultimate segments 6–8 mm; sori 5–7 mm .........................................................................................  7a. var. moupinense

1b. Fronds monomorphic or nearly so; sterile and fertile laminae lanceolate; ultimate segments 3–5 mm; sori 2–3 mm .....................................................................................................................  7b. var. ipii

7a. Onychium moupinense var. moupinense

木坪金粉蕨(原变种)  mu ping jin fen jue (yuan bian zhong)

Onychium japonicum Thunberg subsp. moupinense (Ching) Fraser-Jenkins

Plants to 25–70 cm tall. Fronds subdimorphic. Sterile lamina lanceolate; fertile lamina ovate-lanceolate. Ultimate segments linear, 6–8 mm. Sori 5–7 mm.

Crevices in shrubs or broad-leaved forests; 500–1900 m. Hubei, Shaanxi, Sichuan, Yunnan [?Japan, Nepal].

7b. Onychium moupinense var. ipii (Ching) K.H. Shing, Fl. Reipubl. Popularis Sin. 3(1): 112. 1990.

湖北金粉蕨  hu bei jin fen jue

Onychium ipii Ching, Lingnan Sci. J. 15: 282. 1936.

Plants mostly 20–40 cm tall. Fronds monomorphic. Sterile and fertile laminae lanceolate. Ultimate segments short, 3–5 mm. Sori 2–3 mm.

* Shaded, moist places; below 1100 m. W Hubei (Gucheng).

Some botanists continue to treat this taxon as a separate species and O. angustifrons is sometimes treated as a synonym.

8. Onychium angustifrons Ching, Fl. Reipubl. Popularis Sin. 3(1): 112. 1990.

狭叶金粉蕨  xia ye jin fen jue

Onychium angustifolium Ching, Acta Phytotax. Sin. 20: 233. 1982, not O. angustifolium (Kunth) Kunze (1848).

Rhizomes long creeping or occasionally shortly creeping; scales dark brown, subulate-lanceolate. Fronds slightly dimorphic, mostly relatively closely spaced. Stipe straw-colored above a ± black base, 4–11 cm Χ ca. 1 mm. Lamina of sterile fronds narrowly lanceolate, 5–7 cm, pinnate-pinnatifid. Lamina of fertile fronds larger than sterile lamina, lanceolate or narrowly lanceolate, 7–12 Χ 1–3 cm, apex pinnatifid and elongate-acuminate, 2-pinnate, thinly papery when dry, green to grayish green, not farinose. lateral pinnae 7–15 pairs, strongly ascending obliquely, basal pair largest, deltate-lanceolate or oblong-lanceolate, 1.5–2 Χ 0.4–0.5 cm. Pinnules (segments) of sterile fronds oblong, 2 Χ 1 mm, apex acute or with 2 or 3 sharp teeth, 1 tooth per veinlet. Ultimate pinnules (segments) of fertile fronds linear, 4–8 Χ ca. 0.7 mm, base narrowly cuneate and narrowly winged-decurrent on costae, apex acuminate. Sori 3–6 mm. False indusia oblong to linear, entire.

* Exposed rock crevices. C Sichuan.

Onychium angustifrons has lamina less divided than most other Onychium species and is small in stature, mostly less than 20 cm tall. It resembles O. moupinense in habit. Further studies are needed to show whether it is indeed a distinct species or merely a reduced form of O. moupinense in an extreme habitat.

4. Subfam. CHEILANTHOIDEAE

碎米蕨亚科  sui mi jue ya ke

Zhang Gangmin (张钢民); George A. Yatskievych, Tom A. Ranker

Plants terrestrial or on rocks, mostly medium or small in stature. Rhizomes erect or ascending to prostrate, caespitose to long creeping, siphonostelic, infrequently dictyostelic, bearing scales. Fronds monomorphic or rarely dimorphic, clustered to widely scattered. Stipe brown to black, with 1 or 2 vascular bundles near base, terete or adaxially grooved, glabrous, hairy, or scaly. Lamina variously shaped, 2–4-pinnate, sometimes also simple or pinnatifid, herbaceous to papery or leathery, glabrous or hairy, scaly, and/or farinose. Veins free, occasionally anastomosing (without included free veinlets). Sori submarginal at vein tips or along veins, variously shaped, discrete to confluent, sometimes along commissural vein connecting vein tips (Doryopteris). Indusia absent (Parahemionitis and Paragymnopteris) or more commonly present, then formed by reflexed and somewhat modified lamina margin (false indusium). Spores mostly brown, trilete, globose-tetrahedral to globose, perispore variously ornamented.

About 19 genera and ??? species: worldwide, mainly in subtropical areas; seven genera and 46 species in China.

Molecular data has shown that the cheilanthoid ferns form a well-defined monophyletic group, Cheilandthoideae, within Pteridaceae. In FRPS, this fern group was separated into two different families, Sinopteridaceae and Hemionitidaceae.

1a.       Sori borne along veins, indusia absent.

2a.       Fronds somewhat dimorphic, simple, ovate or oblong, base cordate or hastate; veins anastomosing       15. Parahemionitis

2b.       Fronds monomorphic, 1–3-pinnate; veins free or connected only near frond margin .........  16. Paragymnopteris

1b.       Sori submarginal at vein tips, usually covered by reflexed and somewhat modified lamina margins (false indusia).

3a.       Lamina with white or yellow farina on abaxial surface ...............................  14. Aleuritopteris

3b.       Lamina lacking white or yellow farina on abaxial surface at maturity (sometimes farinose when young).

4a.       Lamina pentagonal (i.e. proximal basiscopic pinnules of basal pinnae enlarged).

5a.       Sori borne at vein tips, rounded, separate (confluent when mature); rhizome scales brown or dark brown, concolorous or with narrow lighter brownish margins ...............  14. Aleuritopteris

5b.       Sori borne along marginal commissural vein, linear; rhizome scales bicolored, with black and thick central band, broad brownish and membranous margins.

6a.       Rhizomes long creeping; fronds mostly widely spaced ....................  10. Calciphilopteris

6b.       Rhizomes short, erect or ascending; fronds clustered ............................  11. Doryopteris

4b.       Lamina lanceolate, oblong-lanceolate, or oblong-triangular to deltate-lanceolate or deltate-ovate.

7a.       Lamina margins not or only slightly reflexed, indusia absent; lamina with dense, yellow to brownish, long hairs abaxially .................................................................................  13. Cheilanthes

7b.       Lamina margins reflexed and modified, forming scarious false indusia; lamina glabrous or relatively sparsely hairy (in a few species also glandular and/or scaly) abaxially.

8a.       Lamina thick, leathery, pinnules mostly hastate to ovate-hastate ...................  12. Pellaea

8b.       Lamina herbaceous or papery, pinnules or segments entire or pinnately lobed.

9a.       Scales of rhizome lanceolate or linear-lanceolate, usually black or dark brown, with an entire margin .........................................................................................................  13. Cheilanthes

9b.       Scales of rhizome ovate-lanceolate, brown, semi-hyaline, with a sparsely serrate margin ...........  14. Aleuritopteris

10. CALCIPHILOPTERIS Yesilyurt & H. Schneider, Phytotaxa 7: 53. 2010.

戟叶黑心蕨属  ji ye hei xin jue shu

Zhang Gangmin (张钢民); George Yatskievych

Plants terrestrial or on rocks. Rhizomes shortly to long creeping; scales lanceolate to narrowly ovate, mostly bicolorous, with a dark brown, ± clathrate, central band and lighter-brown margins. Fronds scattered to ± clustered, dimorphic, fertile fronds with longer stipes and more finely divided laminae than sterile ones. Stipe longer than lamina (except sometimes in sterile fronds), shiny, dark brown to black, rounded or distally somewhat flattened to slightly grooved adaxially, with 1 vascular bundle at base, scaly at base, glabrescent or with sparse, short, hairs and/or scales distally. Lamina pedate-pinnatifid to pedate-bipinnatifid, pedately lobed, or trilobate (smaller fronds sometimes unlobed), pentagonal to broadly cordate, hastate, sagittate, or ovate in outline, margins entire and often white-cartilaginous, papery to leathery, glabrous on both surfaces or rarely sparsely glandular-hairy and/or scaly abaxially at base, midvein and costae blackish abaxially. Veins faint, anastomosing, those of fertile fronds connected to a marginal commissural vein, those of sterile fronds free near margins, ending in submarginal, adaxial hydathodes. Sori with numerous sporangia continuous along commissural vein, margin of false indusium entire to slightly erose. False indusia formed by reflexed lamina margins, continuous except at segment tips. Spores globose to globose-tetrahedral, dark brown, echinate to cristate. x = 29.

Four species: SE Asia to New Guinea and Australia; one species in China.

Calciphilopteris was segregated from Doryopteris recently based mainly on molecular data. The plants tend to grow in limestone areas and have creeping rhizomes and semiclathrate rhizome scales.

1. Calciphilopteris ludens (Wallich ex Hooker) Yesilyurt & H. Schneider, Phytotaxa 7: 56. 2010.

戟叶黑心蕨  ji ye hei xin jue

Pteris ludens Wallich ex Hooker, Sp. Fil. 2: 210. 1858; Doryopteris ludens (Wallich ex Hooker) J. Smith; Litobrochia ludens (Wallich ex Hooker) Beddome; Pellaea ludens (Wallich ex Hooker) Prantl.

Rhizomes long creeping, ca. 4 mm in diam., scales 2–3 mm, lanceolate to narrowly ovate, margins entire. Fronds mostly widely spaced. Stipe rounded, sparsely scaly proximally, with sparse, scales and hairs 1–2 mm distally, glabrescent with age; 10–35 cm (sterile) or 20–40 cm (fertile). Lamina 10–25 cm, deeply cordate at base, medium green adaxially, yellowish green to pale green abaxially; sterile lamina coarsely divided, sagittate or hastate to pedately lobed (rarely smallest fronds unlobed); fertile lamina more finely divided, pedately lobed to pedate-pinnatifid or pedate-bipinnatifid with 3 or 4 pairs of primary lobes, basal pair largest, strongly produced basiscopically, ultimate lobes caudate. 2n = 232.

Limestone rocks by streams in forests; 400–1000 m. S Yunnan [Bangladesh, Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Philippines, Thailand, Vietnam; Australia].

11. DORYOPTERIS J. Smith, J. Bot. 3: 404. 1841.

黑心蕨属  hei xin jue shu

Zhang Gangmin (张钢民); George Yatskievych

Plants terrestrial or on rocks. Rhizomes shortly creeping to ascending or erect, compact; scales lanceolate to narrowly ovate, mostly bicolorous, with a dark brown, nonclathrate, central band and lighter-brown margins. Fronds ± clustered, monomorphic to somewhat dimorphic, fertile fronds sometimes with longer stipes and more finely divided laminae than sterile ones, Stipe longer than to ± as long as lamina, shiny, reddish brown to black, rounded or grooved adaxially, with 1 or 2 vascular bundles at base, scaly at base, glabrescent or with sparse, short, hairs. Lamina mostly pedately 1–3 times lobed or pinnatifid, in a few species pinnate, trilobate, or entire, pentagonal to broadly cordate, hastate, sagittate, or ovate in outline, margins entire to weakly crenulate, often with a white, brown, or black cartilaginous border, papery to leathery, glabrous on both surfaces or rarely sparsely glandular-hairy and/or scaly abaxially at base, in some species with a proliferous bud at base, midvein and costae reddish brown to blackish abaxially. Veins faint, free or anastomosing, those of fertile fronds connected to a marginal commissural vein, those of sterile fronds free near margins, slightly to noticeably enlarged, then ending in submarginal, adaxial hydathodes. Sori with numerous sporangia continuous along commissural vein, margin of false indusium entire to slightly erose. False indusia formed by reflexed lamina margins, usually continuous except at segment tips. Spores globose to globose-tetrahedral, yellow to brown, rugulate or echinate to cristate. x = 29, 30.

About 35 species: Africa, Australia, N America, S America, Asia; one species in China.

1. Doryopteris concolor (Langsdorff & Fischer) Kuhn in Decken, Reisen Ost-Afrika 3(3): 19. 1879.

黑心蕨  hei xin jue

Pteris concolor Langsdorff & Fischer, Pl. Voy. Russes Monde 19, t. 21. 1810; Allosorus concolor (Langsdorff & Fischer) Kuntze; Cheilanthes concolor (Langsdorff & Fischer) R. A. Tryon & A. F. Tryon; Pellaea concolor (Langsdorff & Fischer) Baker; Pteris cheilanthoides Hayata.

Rhizomes short, erect or ascending. Fronds monomorphic, clustered. Stipe 6–24 cm Χ 1–2 mm, shallowly grooved adaxially, rounded abaxially, reddish brown, shiny, with 1 U-shaped vascular bundle proximally. Lamina pentagonal to elongate-pentagonal, as long as wide, 4–8 cm, pedate-bipinnatisect to pedate quadripinnatisect, lacking proliferous buds, base deeply cordate, apex of segments acute, bluntly to sharply pointed, thin to relatively firm in texture, green to yellowish green, glabrous. Middle primary segment broadly rhombic, 3.5–5 Χ 2–4 cm, base broadly cuneate, decurrent along rachis. Lateral pair of pinnae deltate, 3–4.5 Χ 2.5–3.5 cm, base decurrent and connected to middle pinna with broad wings; basiscopic pinnules much longer than acroscopic ones, proximal basiscopic pinnule longest, pinnatipartite margins entire or slightly crenate, with a white to brown border. Veins free, obscure, free vein ends forming adaxial hydathodes. Sori along segment margins, interrupted at sinuses and segment tips. Spores yellow, rugulate. 2n = 60.

Terrestrial or on rocks, along streams, in forests or open areas; 200–800 m. Guangdong, Guangxi, Hainan, Taiwan [India, Malaysia, Sri Lanka, Vietnam; Africa, Australia, Caribbean Islands, Central and South America, Madagascar, Mexico].

12. PELLAEA Link, Fil. Sp. Cult. t. 49, 59. 1841.

旱蕨属  han jue shu

Zhang Gangmin (张钢民); George Yatskievych

Plants usually on rocks. Rhizomes erect to decumbent or long creeping, siphonostelic; scales concolorous or bicolorous, brown to nearly black, sometimes with narrow pale margins, narrowly lanceolate or subulate-lanceolate to lanceolate, margins entire or toothed. Fronds monomorphic or slightly dimorphic, clustered or more widely spaced. Stipe usually brown to black, usually lustrous, terete or adaxially grooved, with 1 vascular bundle at base. Lamina oblong-lanceolate to deltate-lanceolate, usually leathery, thick, 1–4-pinnate, with a discrete, terminal pinnule or segment, papery or leathery, glabrous on both surfaces or occasionally hairy or scaly. Ultimate pinnules or segments stalked, or if sessile then narrowly attached, variously shaped, sometimes with 1 or 2 blunt basal lobes, margins entire, apex rounded to acute. Veins free or rarely anastomosing, pinnately branched. Sori small, orbicular, at vein tips, usually confluent at maturity. False indusia linear, continuous, usually narrow, margins entire or erose. Spores globose-tetrahedral, perispore granulate, cristate, rugose or rarely tuberculate to verrucate. x = 29, 30.

About 30 species: S Africa, S America, N America, Asia, New Zealand, Australia; two species in China.

Traditionally, the genus Pellaea was circumscribed broadly to include 45–50 species in 4 sections distributed on most continents. However, this version of Pellaea is polyphyletic and includes several distantly related groups. Of the two Chinese species treated here, P. calomelanos is clearly a member of sect. Holchochlaena Hooker & Baker, which includes 10 or more species with a center of diversity in Africa. In the future, it is likely that this group will be raised to generic status, as it is only distantly related to Pellaea sect. Pellaea. The systematic position of the rare P. connectens has not yet been evaluated because of the lack of recently collected material, but likely it is related to P. calomelanos. Other species included in Pellaea in some of the literature on Asian ferns are here treated in Cheilanthes, including those sometimes segregated under the generic name Mildella.

1a.       Pinnules ovate to ovate-hastate, base cordate, terminal pinnules almost always with a pair of blunt, spreading lobes .........................................................................................  1. P. calomelanos

1b.       Pinnules oblong to ovate, unlobed, base rounded to truncate .........................  2. P. connectens

1. Pellaea calomelanos (Swartz) Link, Fil. Spec. 61. 1841.

三角羽旱蕨  san jiao yu han jue

Pteris calomelanos Swartz, J. Bot. (Schrader) 1800(2): 70. 1801; Pellaea hastata (Thunberg) Prantl (1882), not Link (1841); Pteris hastata Thunberg.

Rhizomes short and stout, 4–8 mm in diam., decumbent or ascending; Scales bicolorous, black, with narrow pale brown margins, lustrous, lanceolate to narrowly lanceolate. Fronds clustered. Stipe (3–)5–12(–30) cm, dark brownish black, terete, with sparse narrow scales at base, glabrous above. Lamina ovate to oblong-deltate, (5–)8–15(–25) Χ 4–6(–15) cm, 2-pinnate with discrete terminal pinnule, leathery, grayish green, pale abaxially, glabrous on both surfaces; rachis, costae, and pinnule stalks dark brownish black, terete, glabrous or rarely sparsely shortly hairy or -glandular. Pinnae 11–21, oblique, basal or second pairs of pinnae largest, medial and distal pinnae gradually reduced, undivided or imparipinnate with 3–11 pinnules, stalks 3–10 mm. Pinnules 3–7(–11), 6–25 Χ 4–22 mm, discrete, ovate to ovate-hastate, base cordate, apex obtuse, with stalks to ca. 3 mm. Veins obscure. Sori confluent. False indusia slender, brown, continuous, margins entire.

Rock crevices of dry valleys; 900–1800 m. S Sichuan, Yunnan [India, Nepal, Pakistan; Africa, S Europe, Madagascar, Indian Ocean islands (Mascarene Islands)].

This widespread species sometimes has been divided into two or more varieties in Africa. The Chinese plants correspond to var. calomelanos.

2. Pellaea connectens C. Christensen, Acta Hort. Gothob. 1: 84. t. 18, f. d; t. 19, f. a. 1924.

四川旱蕨  si chuan han jue

Rhizomes short, erect; scales concolorous, brown, subulate-lanceolate, thin and twisted. Fronds densely clustered. Stipe 3–8 cm, terete, dark brown, with sparse narrow scales at base, occasionally with sparse, brown, hairlike scales distally, brittle. Lamina lanceolate to somewhat oblong-triangular, 5–10 Χ 2–4 cm, base not reduced, apex shortly acuminate, 2-pinnate or occasionally 3-pinnate with a discrete terminal pinnule or segment, papery, brownish green when dry, glabrous on both surfaces; rachis straight; rachis, costae, and costules chestnut-colored, lustrous. Pinnae 7–21, oblique, basal or second pair of pinnae largest, medial and distal pinnae gradually reduced, undivided or imparipinnate with 3–9 pinnules, stalks 1–3 mm. Ultimate pinnules or segments discrete, oblong to ovate, unlobed, 3–6 Χ 2–3 mm, shortly stalked, base rounded to truncate, apex rounded or obtuse. Sori ± confluent, continuous or sometimes interrupted at pinnule or segment apex. False indusia poorly developed, narrow, pale green, margins undulate or irregularly crenate.

* Rare; limestone crevices in hot, dry valleys; 1900–2800 m. SW and W Sichuan.

13. CHEILANTHES Swartz, Syn. Fil. 5, 126. 1806, nom. cons.

碎米蕨属  sui mi jue shu

Zhang Gangmin (张钢民); George Yatskievych

Cheilosoria Trevisan, misapplied; Mildella Trevisan; Notholaena R. Brown, misapplied.

Plants terrestrial or on rock. Rhizomes short and erect, less commonly decumbent or creeping, siphonostelic; scales concolorous or bicolorous, brown to chestnut-black, sometimes with a pale border, subulate to lanceolate. Fronds monomorphic, tufted, clustered, or relatively closely spaced along rhizome. Stipe chestnut-colored to black, sometimes lustrous, terete or grooved adaxially, with 1 vascular bundle at base, sparsely to densely scaly and hairy, at least proximally or when young, sometimes glabrescent at maturity. Lamina lanceolate, oblong-lanceolate, oblong, or ovate-pentagonal, 1–3-pinnate-pinnatifid, herbaceous or papery to leathery, glabrous or hairy on both surfaces. Ultimate segments sessile to shortly stalked, broadly or narrowly attached, variously shaped. Veins of ultimate segments free (but sometimes obscure), unbranched or forked distally. Sori orbicular, at vein tips, often confluent at maturity. False indusia absent or formed by reflexed margin, this interrupted to continuous, sometimes somewhat modified, margins entire, erose, serrulate, or ciliate. Spores globose-tetrahedral, perispore granulate, pseudo-reticulate, cristate, or rarely rugulate. x = 28, 29, 30.

More than 100 species: Asia, Africa, Australia, Europe, Oceania, South America; 17 species in China.

As treated here, the genus Cheilanthes is an unnatural assemblage that includes members of two groups. The genera Cheilosoria Trevisan and Notholaena R. Brown are typified by New World species and these names have been misapplied to the Asian taxa. The Asian species that have been treated under these names are part of a lineage that includes the species of Aleuritopteris, according to several independent studies of molecular phylogeny. These taxa are included in Cheilanthes for convenience, both because the necessary combinations in Aleuritopteris have not been published and because taxonomic relationships within the Aleuritopteris lineage are still poorly understood. The species formerly included in Mildella, which sometimes have been misclassified in Pellaea, also are members of this lineage. On other hand, C. tenuifolia is part of Cheilathes s.s., a group of ca. 35 species with centers of diversity in Australia, S Africa, and South America.

1a.       Lamina margins not or only slightly reflexed, differentiated false indusia absent; lamina densely covered by yellow to brownish long hairs abaxially.

2a.       Rhizomes long creeping; fronds scattered; lamina pinnate-pinnatifid; pinnae subsessile  2. C. chinensis

2b.       Rhizomes short, erect or decumbent; fronds clustered; lamina 2-pinnate-pinnatifid; pinnae distinctly stalked ......................................................................................................  3. C. nudiuscula

1b.       Lamina margins strongly reflexed and modified forming scarious false indusia; lamina frequently glabrous abaxially.

3a.       Lamina thickly papery to leathery, coarsely divided, often caudate at apex; false indusia continuous.

4a.       Stipes straw-colored, sometimes tan proximally.

5a.       Fronds subdimorphic; lamina pinnate-pinnatifid, to 15 Χ 10 cm; segments 2–4 cm  12. C. christii

5b.       Fronds monomorphic; lamina 2-pinnate-pinnatifid, to 10 Χ 3 cm; pinnules 1–2 cm.

6a.       Stipes longer (usually much longer) than laminae; false indusia broad, reaching nearly to midvein, margins sparsely fimbriate ........................................................................  13. C. smithii

6b.       Stipes ± as long as laminae; false indusia narrow, not approaching midvein, margins entire to somewhat erose .....................................................................................  14. C. tibetica

4b.       Stipes brown to black.

7a.       Pinnules deltate, ovate, or oblong, at least those of basal pinnae stalked, apex obtuse; rachis flexuous .........................................................................................................  11. C. trichophylla

7b.       Pinnules lanceolate or linear, base adnate to costa, apex acuminate; rachis straight or nearly so.

8a.       Stipes and rachises glabrous above scaly base; laminae ovate-pentagonal, ± as long as wide          17. C. bhutanica

8b.       Stipes and rachises shortly hairy and/or with broader scales; laminae oblong to oblong-deltate or ovate-deltate, nearly 2 times as long as wide.

9a.       Stipes sparsely scaly above base, but densely shortly hairy; apices of lamina and pinnae obtuse or shortly caudate, caudate portion less than 1 cm; pinnules (segments) closely spaced  15. C. nitidula

9b.       Stipes moderately to densely scaly throughout, also shortly hairy; apices of lamina and pinnae long caudate; caudate portion to ca. 2 cm; pinnules (segments) widely spaced  16. C. brausei

3b.       Lamina herbaceous to somewhat papery, finely divided, apex pinnatifid, acuminate; false indusia usually interrupted.

10a.     Lamina ovate-pentagonal or elongate-deltate, 3-pinnate; basal pair of pinnae larger than adjacent pinnae and proximal basiscopic pinnules conspicuously enlarged.

11a.     Stipe terete or slightly flattened ..............................................................  10. C. hancockii

11b.     Stipe noticeably flattened or grooved abaxially.

12a.     Scales of rhizome and stipe base brownish, soft; stipe slightly grooved adaxially, without two sharp margins ................................................................................................  1. C. tenuifolia

12b.     Scales of rhizome and stipe base black, stiff; stipe and rachis flattened to shallowly grooved abaxially with two sharp margins ..............................................................................  8. C. patula

10b.     Lamina lanceolate, pinnate-pinnatifid to 3-pinnate; basal pair of pinnae not larger than adjacent pinnae; proximal basiscopic pinnules not conspicuously enlarged.

13a.     Lamina 3-pinnate; costae sparsely clothed with glandular hairs adaxially ......  9. C. insignis

13b.     Lamina pinnate-pinnatifid to 2-pinnate; costae glabrous or with nonglandular hairs and/or scales adaxially.

14a.     Pinnae obtuse at apex, sparsely hairy on both surfaces; margins of false indusia fimbriate  4. C. fragilis

14b.     Pinnae acute or acuminate at apex, glabrous on both surfaces; margins of false indusia entire to slightly irregular or undulate.

15a.     Pinnae widely separated, elongate-acuminate at apex; basal pinnae of lamina same size as adjacent pinnae; basal-basiscopic pinnules larger than acroscopic ones, proximal basiscopic pinnule largest; false indusia continuous, rarely somewhat interrupted ...........................  7. C. belangeri

15b.     Pinnae closely spaced, acute at apex, basal pinnae of lamina somewhat smaller than adjacent pinnae; basal-basiscopic pinnules same size as acroscopic ones; false indusia interrupted.

16a.     Winglike ridges along margins of stipe and rachis groove with short hairs and narrow scales       5. C. chusana

16b.     Winglike ridges along margins of stipe and rachis groove glabrous
..........................................................................................................  6. C. opposita

1. Cheilanthes tenuifolia (N. L. Burman) Swartz, Syn. Fil. 129, 332. 1806.

薄叶碎米蕨  bo ye sui mi jue

Trichomanes tenuifolium N. L. Burman, Fl. Ind. 237. 1768; Cheilosoria tenuifolia (N. L. Burman) Trevisan.

Rhizomes short, erect; scales yellowish brown, subulate, not sclerified. Fronds clustered. Stipe 6–25 cm, chestnut brown, grooved adaxially, margins of groove blunt, rounded abaxially, sparsely scaly proximally, glabrous distally. Lamina deltate, pentagonal-ovate, or broadly ovate-lanceolate, shorter than stipe, 4–18 Χ 4–12 cm, tripinnate or pinnate-bipinnatifid, herbaceous, thin, green to yellowish green, with sparse, short hairs adaxially, glabrous abaxially; rachis and costae grooved adaxially, margins of grooves narrowly winged on distal rachis and costae. Pinnae 6–8 pairs; basal pair largest, ovate-deltate or ovate-lanceolate, 3–9 Χ 2.5–4.5 cm, stalked (stalks 0.3–1 cm), apex acuminate, proximal pinnae inequilateral, basal basiscopic pinnules largest. Ultimate segments connected by very narrow wings, lobed, lobes elliptic. Sori confined to distal portion of lobes, interrupted at segment bases. False indusia continuous or not, margin entire to somewhat undulate or erose. 2n = 108, 112, 120.

On rocks in forests and along stream banks, roadsides, rice fields; 100–1000 m. Fujian, Guangdong, Guangxi, Hainan, Hong Kong, S Hunan, Jiangxi, Taiwan, Yunnan [Cambodia, India, Laos, Malaysia, Nepal, Sri Lanka, Thailand, Vietnam; Australia, Oceania, New Zealand].

2. Cheilanthes chinensis (Baker) Domin, Biblioth. Bot. 85: 133. 1913.

中华隐囊蕨  zhong hua yin nang jue

Notholaena chinensis Baker, Gard. Chron. n.s. 14: 494. 1880.

Rhizomes creeping, 2–3 mm in diam.; scales linear to linear-lanceolate, bicolorous with dark brown central stripe and lighter margins, subulate-lanceolate. Fronds ± closely spaced. Stipe dark brown, lustrous, terete, (2–)3–14 cm, scaly proximally. Lamina narrowly deltate to oblong-triangular, 7–18 Χ 2–8 cm, pinnate-pinnatifid, papery, adaxially sparsely to moderately villous with fine, brown hairs, abaxially moderately to densely tomentose with fine, long, yellowish brown to orangish brown hairs. Pinnae 10–20 pairs; basal pair largest, 2–4 Χ 1.5–3 cm, sessile, apex rounded to angled, proximal 2–4 pairs strongly inequilateral, basiscopic segments greatly enlarged, much longer than adjacent acroscopic ones; distal pinnae gradually reduced, ± equilateral, pinnatifid or not, apex obtuse. Ultimate segments sessile, broadly adnate to costae, margins slightly recurved but not covering sori. Sori ± continuous or sometimes interrupted at segment sinuses, containing few sporangia, immersed in pubescence. False indusium absent.

* Limestone crevices; 400–800 m. Chongqing, N Guangxi (Lingui), NE Guizhou, W Hubei, Sichuan.

3. Cheilanthes nudiuscula (R. Brown) T. Moore, Index Fil. 249. 1861.

隐囊蕨  yin nang jue

Pteris nudiuscula R. Brown, Prodr. 155. 1810. Pteris hirsuta Poiret; Cheilanthes hirsuta (Poiret) Mettenius (1859), not Link (1833); Cheilanthes densa Fιe; Notholaena hirsuta (Poiret) Desvaux; N. nudiuscula (R. Brown) Desvaux; N. sulcata Link; Pellaea nudiuscula (R. Brown) Hooker.

Rhizomes short, erect; scales concolorous, reddish brown, subulate-lanceolate. Fronds clustered. Stipe chestnut-colored, slightly lustrous, 8–12 cm, shallowly grooved adaxially, margins of groove blunt, villous and with sparse linear scales, sometimes glabrescent with age. Lamina oblong to oblong-lanceolate, 10–16 Χ 2.5–8 cm, 2-pinnate-pinnatifid, papery to somewhat leathery, adaxially sparsely to moderately villous with multicellular gray hairs, abaxially moderately to more commonly densely villous to subtomentose with dense, long, multicellular brown hairs; rachis, costae chestnut brown, sparsely to densely villous. Pinnae 8–10 pairs, proximal pairs ovate-lanceolate or oblong-lanceolate, 2–4.5 Χ 1–3 cm, stalked (stalks 3–5 mm), apex rounded or angled, proximal 2–4 pairs inequilateral, basal basiscopic pinnules somewhat larger than acroscopic ones. Ultimate segments sessile, margins shallowly lobed to entire, slightly recurved but not covering sori. Sori ± continuous or sometimes interrupted at segment sinuses, containing several sporangia, embedded in pubescence. False indusium absent. 2n = 168.

On rocks along rivers or rice fields; 200–400 m. Fujian, Guangdong, Guangxi (Nanning), Hong Kong, Taiwan (Taizhong) [Australia, Malesia, Oceania].

4. Cheilanthes fragilis Hooker, Fil. Exot. t. 96. 1859.

脆叶碎米蕨  cui ye sui mi jue

Cheilosoria fragilis (Hooker) Ching & Shing.

Rhizomes erect; scales concolorous, grayish brown, subulate-lanceolate. Fronds clustered. Stipe chestnut brown to nearly black, 5–6 cm, with dense glandular hairs and a few lanceolate scales. Lamina narrowly lanceolate, 20–25 Χ 4–6 cm, slightly narrowed proximally, apex acuminate, 2-pinnate, herbaceous, thin when dry, green to dark green, sparsely with glandular hairs on both surfaces; rachis chestnut-black, sparsely pubescent and with glandular hairs on both surfaces. Pinnae 15–20 pairs, almost patent; medial pinnae largest, 3–4 Χ ca. 1 cm, lanceolate, shortly stalked, slightly narrowed proximally, apex somewhat obtuse, basal pinnae equilateral, basiscopic pinnules not enlarged. Pinnules narrowly oblong, medial ones larger, to 7 Χ 3 mm, somewhat adnate to costa, margins irregularly crenate, somewhat ciliate, apex obtuse-rounded. Veins pinnate, lateral veins forked. Sori interrupted. False indusia well developed, discontinuous, reniform to nearly semicircular, margin fimbriate.

Limestone rock; ca. 1000 m. S Yunnan (Yiwu, Mengla) [Malaysia, Myanmar].

5. Cheilanthes chusana Hooker, Sp. Fil. 2: 95. t. 106 B. 1852.

毛轴碎米蕨  mao zhou sui mi jue

Cheilosoria chusana (Hooker) Ching & Shing; Adiantopsis fordii (Baker) C. Christensen; Cheilanthes bockii Diels; C. boltoni Copeland; C. fordii Baker; C. mysurensis var. chusana (Hooker) Christ; C. mysurensis var. giraldii Christ.

Rhizomes short, erect; scales concolorous and reddish brown or bicolorous and dark brown to black with narrow orangish brown margin, lanceolate. Fronds tufted. Stipe 2–7 cm, dark brown, lustrous, densely scaly and with sparse, short hairs, scales occasionally bicolorous at base, but then grading abruptly to concolorous, reddish brown, lanceolate or subulate-lanceolate, adaxially grooved, groove with a pair of narrow winglike ridges along margins, these bearing brown, short hairs and/or narrow scales, some of hairs and scales often minutely gland-tipped. Lamina elliptic-lanceolate to lanceolate, 8–28 Χ 2–6 cm, somewhat narrowed at base, shortly tapered at apex, pinnate-pinnatisect, green, herbaceous, glabrous on both surfaces. Pinnae 10–20 pairs, relatively closely spaced, oblique, subsessile; medial pinnae largest, 1–3.5 Χ 0.8–1.5 cm, lanceolate-triangular, acute or obtuse at apex; pinnae gradually reduced proximally and distally, widely separated; basal pair of pinnae oblong to triangular-oblong, equilateral, basiscopic pinnules not enlarged. Ultimate segments oblong to narrowly oblong, often decurrent at base, margins crenate, apex rounded to obtuse. Sori discrete. False indusia well developed, discontinuous, elliptic-reniform to reniform, margins otherwise entire or slightly irregular.

Rock crevices on roadsides, streambanks, or in forests; 100–1500 m. Anhui, Chongqing, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Sichuan, Taiwan, Zhejiang [Japan, Philippines, Vietnam].

6. Cheilanthes opposita Kaulfuss, Enum. 211. 1824.

碎米蕨  sui mi jue

Cheilanthes mysurensis Wallich ex Hooker; Cheilosoria mysurensis (Wallich ex Hooker) Ching & K. H. Shing.

Rhizomes short, erect; scales mostly bicolorous and dark brown to black with narrow orangish brown margin, narrowly lanceolate. Fronds tufted. Stipe 2–7 cm, dark brown to black, densely scaly proximally, sparsely scaly distally, scales concolorous, reddish brown, lanceolate or subulate-lanceolate, adaxially grooved, groove with a pair of narrow winglike ridges along margins, these glabrous, occasionally with a few minute teeth. Lamina elongate-lanceolate to narrowly oblong-elliptic, 8–18(–23) Χ 1–2 cm, base gradually tapered, apex acuminate, bipinnate, rarely bipinnate-pinnatifid, herbaceous, brown when dry, glabrous on both surfaces. Pinnae 10–20 pairs, relatively closely spaced; medial pinnae deltate or deltate-lanceolate, 1–1.5 Χ 0.5–0.8 cm, subsessile; lower pinnae widely separated and gradually shortened proximally, oblong to narrowly deltate; basal pair of pinnae often reduced to small auricles, equilateral or nearly so. Pinnules of medial pinnae usually divided with 3 or 4 pairs of orbicular lobes. Sori 1 or 2 per lobe, discrete. False indusia discontinuous, reniform or somewhat elliptic-reniform, margins otherwise entire or slightly irregular. 2n = 60.

On rocks on streambanks or under shrubs; 100–300 m. Fujian, Guangdong (Dapu), Hainan, Taiwan (Wulei) [India, Myanmar, Sri Lanka, Vietnam].

Cheilanthes opposita is easily confused with the closely related C. chusana.

7. Cheilanthes belangeri (Bory) C. Christensen, Index Filic., fasc. 3: 172. 1905.

疏羽碎米蕨  shu yu sui mi jue

Pteris belangeri Bory in Belanger, Voy. Bot. 2: 44. 1883; Cheilosoria belangeri (Bory) Ching & K. H. Shing; Pteridella belangeri (Bory) Mettenius; Cheilanthes varians Wallich ex Hooker; Allosorus cambodiensis (Baker) Kuntze.

Rhizomes short, erect; scales concolorous, dark brown, subulate-lanceolate. Fronds tufted. Stipe 8–23 cm, dark brown to nearly black, adaxially shallowly grooved, groove rounded or blunt along margins, scaly toward base, scales concolorous, dark brown, subulate lanceolate to subulate. Lamina narrowly triangular-lanceolate, 9–26 Χ 3–5 cm, base not or only slightly narrowed, apex elongate-acuminate, pinnate-pinnatifid to 2-pinnate, papery, brown when dry, glabrous on both surfaces; rachis and costae dark brown, adaxially grooved, usually with minute stalked glands, especially adaxially, sometimes glabrous abaxially. Pinnae 12–18 pairs, widely separated; basal pinnae slightly shorter to ± as long as second pair, elongate-deltate, 1.5–3.5 Χ 0.6–1.5 cm, stalked (stalks 1–4 mm), base broadly cuneate, apex acuminate and often somewhat caudate. Pinnules (segments) of proximal pinnae 4–6 pairs, adnate to costa, inequilateral, basiscopic pinnules somewhat longer than adjacent acroscopic ones; basal basiscopic pinnule longest, oblong-lanceolate, somewhat lobed. Sori interrupted at segment sinuses. False indusia discontinuous, margins entire or slightly undulate. 2n = 120.

On rocks or occasionally in or moist soil; 100–800 m. Hainan [Bangladesh, Cambodia, India, Indonesia, Myanmar, Nepal, Philippines, Thailand, Vietnam].

8. Cheilanthes patula Baker, J. Bot. 26(7): 225–226. 1888.

平羽碎米蕨  ping yu sui mi jue

Cheilosoria patula (Baker ) P. S. Wang; Pellaea patula (Baker) Ching.

Rhizomes short, erect; scales concolorous, black to blackish brown, subulate-lanceolate. Fronds tufted. Stipe 8–15 cm Χ ca. 2 mm, dark brown, adaxially flattened to shallowly grooved, groove with a pair of narrow winglike ridges along margins, these glabrous, scaly toward base, scales dark brown, subulate. Lamina elongate-deltate, 12–20 Χ 6–11 cm, broadest at base, apex acuminate, 2- or 3-pinnate, papery, brownish green when dry, glabrous on both sides; rachis somewhat flexuous; rachis and costae dark brown, adaxially grooved, groove with a pair of narrow winglike ridges along margins. Pinnae 8–10 pairs, stalked, stalks 2–4 mm; basal pair of pinnae largest, oblong-deltate, 3–6 Χ 1.6–2 cm; pinnules deltate, 8–12 Χ 4–6 mm, shortly stalked, inequilateral, basiscopic pinnules longer than adjacent acroscopic ones; basal basiscopic pinnule longest. Ultimate segments oblong, margins entire, apex obtuse. Sori usually not interrupted on segments. False indusia continuous or rarely somewhat discontinuous, margins entire.

* Rock crevices; 400–900 m. Chongqing, Guangxi, Guizhou, W Hubei.

9. Cheilanthes insignis Ching, Fl. Tsinling. 2: 208, t. 18, f. 7–9. 1974.

厚叶碎米蕨  hou ye sui mi jue

Cheilosoria insignis (Ching) Ching & K. H. Shing.

Rhizomes short, creeping to ascending; scales concolorous or bicolorous, black, sometimes with brown margins, shiny, lanceolate, stiff. Fronds closely spaced. Stipe 10–15 cm Χ ca. 2 mm, reddish brown to brown, lustrous, grooved adaxially, groove rounded along margins, scaly toward base or on smaller fronds scales sometimes distributed nearly to lamina, scales dark brown, mostly concolorous, lanceolate-subulate to hairlike. Lamina broadly lanceolate, 15–17 Χ 5–6 cm, 3-pinnate, papery, brownish green when dry, glabrous on both surfaces or with minute stalked glands, mostly adaxially; rachis and costae dark brown, adaxially grooved, with sparse, minute, stalked glands adaxially. Pinnae 9–14 pairs, oblique; second or third pinnae largest, ovate-lanceolate or broadly lanceolate, 4–5 Χ 1.5–2.5 cm, shortly stalked, acroscopic basal pinnule parallel to rachis, basiscopic base cuneate, apex acuminate. Pinnules of basal pinnae; slightly inequilateral, basiscopic pinnules slightly longer than adjacent acroscopic ones; basal basiscopic pinnule shorter than adjacent basiscopic ones, oblong to oblong-ovate, lobed, axes narrowly winged. Ultimate segments ovate or oblong, margins shallowly lobed. Sori interrupted at segment sinuses. False indusia discontinuous, margins slightly erose.

* On rocky slopes or under shrubs in dry-hot valleys; 1700–3300 m. N Sichuan, E Xizang.

10. Cheilanthes hancockii Baker, Bull. Misc. Inform. Kew 1895(99): 54. 1895.

大理碎米蕨  da li sui mi jue

Cheilosoria hancockii (Baker) Ching & K. H. Shing Cheilanthes bonatiana Brause; Cheilanthes henryi Christ; Cheilanthes taliensis Christ; Cheilanthes wilsonii Christ.

Rhizomes short, erect; scales bicolorous, dark brown to black with lighter. margins, subulate-lanceolate. Fronds clustered. Stipe 6–20 cm Χ ca. 1 mm, dark brown, lustrous, terete, scaly toward base, scales dark brown, concolorous to slightly bicolorous, lanceolate-subulate. Lamina pentagonal-ovate to broadly deltate, 5–15 Χ 5–9 cm, apex acuminate to elongate-acuminate, 3-pinnate, herbaceous, brownish green when dry, glabrous on both surfaces; rachis and costae flattened to shallowly grooved adaxially, glabrous. Pinnae 5–7 pairs; basal pair longest, elongate-deltate, (3–)5–9 Χ 2–4 cm, shortly stalked, acroscopic basal pinnule parallel to rachis, basiscopic base cuneate, basal basiscopic pinnule largest, much longer than adjacent acroscopic ones, apex acuminate. Ultimate segments oblong, ca. 1 Χ 0.2–0.3 cm, base narrowly winged, margins undulate or crenate, apex obtuse or rounded. Sori discrete at vein tips, False indusia discontinuous, reniform, semi-orbicular, or oblong, sometimes adjacent ones fused and indusium then 2-lobed, margins otherwise entire or somewhat erose. 2n = ca. 120.

On rock in woods or under shrubs by road; 1400–3000 m. Gansu (Wenxian), W Guizhou, Sichuan, Xizang (Chayu), Yunnan [Bhutan].

11. Cheilanthes trichophylla Baker, Ann. Bot. 5:211. 1891.

毛旱蕨  mao han jue

Pellaea trichophylla (Baker) Ching; C. delavayi Baker; C. undulata Hope & C. H. Wright.

Rhizomes short, decumbent to erect; scales concolorous or somewhat bicolorous, dark brown to black, sometimes with very narrow, lighter brown margins, lustrous, subulate-lanceolate, stiff. Fronds tufted. Stipe 10–30 cm Χ 1–2 mm, dark brown to black, terete, with fine, orangish brown to brown, multicellular hairs of 2 kind: moderate to dense, short, ascending hairs and scattered, longer, spreading hairs. Lamina deltate-lanceolate, at least slightly longer than stipe, 5–15 cm wide, apex shortly acuminate, 2-pinnate-pinnatifid, papery, dull brownish green when dry, strigose on both surfaces; rachis flexuous; rachis, costae, and pinnule stalks dark brown, terete, densely pubescent with short, multicellular hairs. Pinnae 6–10 pairs, basal pair longest, deltate or deltate-lanceolate, 5–11 Χ 2.5–5 cm, stalks 5–12 mm, apex acute, equilateral. Pinnules 3–5 pairs, ovate-deltate, 1.5–2.5 Χ 1–2 cm, stalks 1–2 mm, base rounded to truncate or somewhat cordate, pinnatifid, costules winged; segments oblong, margins usually entire. Sori confluent, sometimes interrupted at segment tip. False indusia continuous, narrow, margins undulate to crenulate. 2n = 60.

Rock crevices in hot-dry valley or under woods; 800–2500 m. W Sichuan, E Xizang, W Yunnan [possibly India (Sikkim)].

12. Cheilanthes christii Fraser-Jenkins & Yatskievych in Fraser-Jenkins, Pangtey & Khullar, Indian Fern J. 27: 213. 2010 [Aug 2011].

凤尾旱蕨  feng wei han jue

Pteris paupercula Christ, Bull. Acad. Int. Gιogr. Bot. 16: 131. 1906, not Cheilanthes paupercula (Kunze) Mettenius (1856); Cheilanthes paupercula (Christ) Fraser-Jenkins (2008, nom. illeg.); Pellaea paupercula (Christ) Ching; Mildella paupercula C. C. Hall & Lellinger.

Rhizomes short, erect; scales bicolorous, dark brown to black with light brown margins, narrowly lanceolate, shiny, thick and stiff. Fronds clustered, subdimorphic (sterile fronds somewhat shorter and with slightly broader segments). Stipe 5–22 cm, straw-colored, sometimes tan proximally, terete or flattened to grooved adaxially, with, concolorous, reddish brown, soft, narrowly lanceolate scales extending onto rachis, also pubescent toward tip with moderate to dense, short and sparse, longer, multicellular hairs. Rachis and costae straw-colored, densely pubescent with short, viscid hairs and sparse hairlike scales mostly adaxially. Sterile lamina shorter than fertile lamina, herbaceous, pale brownish green when dry, glabrous on both surfaces. Fertile lamina oblong, 5–15 Χ 3–10 cm, apex caudate, 2-pinnate; pinnae 3–7 pairs, proximal pinnae ovate-deltate, 3–7 Χ 2–4 cm, shortly stalked, apex caudate. Ultimate sterile segments oblong or broadly linear, 5–6 mm wide, margins entire or occasionally obscurely crenulate. Ultimate fertile segments linear, 2–3 mm wide, base decurrent on costa, apex acuminate. Sori confluent, not interrupted at sinuses or segment tips. False indusia continuous, margins erose and sparsely ciliate.

* Rock crevices of dry valleys; 1200–2900 m. W Sichuan.

Cheilanthes christii superficially resembles some of the smaller species of Pteris, but these Pteris species have more strongly dimorphic fronds, usually with the margins of the sterile fronds finely dentate-serrulate, and stipes that are grooved and glabrous above the base.

13. Cheilanthes smithii (C. Christensen) R. M. Tryon, Amer. Fern J. 76: 185–186. 1966.

西南旱蕨  xi nan han jue

Pellaea smithii C. Christensen, Acta Hort. Gothob. 1: 84, t. 18, f. a–c. 1924. Mildella smithii (C. Christensen) C. C. Hall & Lellinger.

Rhizomes short, erect or ascending; scales bicolorous, dark brown to black with light brown margins, lanceolate. Fronds clustered, monomorphic or nearly so. Stipe 6–20 cm Χ ca. 1 mm, straw-colored, sometimes tan proximally, terete or flattened to slightly grooved adaxially, with concolorous, reddish brown, soft, narrowly lanceolate scales mostly at base, also pubescent adaxially with shorter and longer, lax, unicellular and multicellular, viscid hairs, short hairs mostly glandular distally. Lamina oblong, much shorter than stipe, 4–10 Χ 2–3 cm, 2-pinnate, leathery, pale brownish green when dry, glabrous on both surfaces; rachis straw-colored. sometimes light brownish tinged, grooved, with sparse viscid hairs adaxially. Pinnae 2–7 pairs, ovate-deltate, 1–2.3 Χ 1–2 cm, sessile. Pinnules or segments 3–5 pairs, linear, 0.5–1.5 cm Χ 1.2–1.6 mm, base decurrent on costa, apex obtuse or mucronate, margins entire or obscurely crenulate, sometimes with 1–3 small lobes. Sori confluent, not interrupted at sinuses or segment tips. False indusia continuous, broad, nearly to midvein, margins fimbriate.

* Rock crevices under shrubs in dry valleys; 1300–2600 m. W Sichuan, NW Yunnan.

14. Cheilanthes tibetica Fraser-Jenkins & Wangdi in Fraser-Jenkins & Dulawat, Fern Gaz. 18: 218. 2009.

禾杆旱蕨  he gan han jue

Pellaea straminea Ching, Bull. Fan Mem. Inst. Biol. 2: 203, t. 17. 1931, non Cheilanthes straminea Brause (1913). Mildella straminea (Ching) Hall & Lellinger; Pellaea straminea var. tibetica Ching.

Rhizomes short, erect or ascending; scales somewhat bicolorous, with dark brown to black central stripe grading to lighter brown margins, shiny, lanceolate. Fronds clustered, monomorphic. Stipe to 7 cm Χ ca. 1 mm, straw-colored, grooved adaxially, at least toward tip, brittle scaly, scales concolorous, brown proximally, grading to light brown below midpoint, also pubescent with sparse to dense, short, mostly bicellular, viscid hairs adaxially. Lamina ovate-oblong to ovate-deltate, (3–)5–8 Χ 1.5–3 cm, pinnate-pinnatifid, acute or acuminate at apex, herbaceous to thinly papery, grayish green when dry, glabrous on both surfaces; rachis and costae straw-colored, grooved adaxially, with dense, short, viscid hairs adaxially. Pinnae 3–5(–7) pairs, subsessile, basal pair longest, elongate-ovate or ovate-deltate, 1.5–2 Χ 1–1.4 cm, apex caudate. Segments linear-lanceolate, 6–15 Χ 1–2 mm, bases adnate to costae, margins entire or somewhat undulate. Sori ± confluent, not interrupted at sinuses or segment tips, occasionally interrupted along lateral margins. False indusia narrow, continuous or occasionally somewhat interrupted, membranous, margins entire.

On rocks; 3000–4500 m. Gansu (Linxia), S Qinghai (Nangqian), Xinjiang, Xizang [Bhutan].

Pellaea straminea var. tibetica was described based on a specimen from Xizang. It was said to differ from var. straminea only in its larger stature (to 22 cm tall) and likely represents plants collected in moister or shadier than usual habitats. The name Cheilanthes tibetica is based on type of Pellaea straminea and was published without reference to the existence of the heterotypic infrataxon, P. straminea var. tibetica.

15. Cheilanthes nitidula Wallich ex Hooker, Sp. Fil. 2: 112. 1852.

旱蕨  han jue

Pellaea nitidula (Wallich ex Hooker) Baker; M. nitidula (Wallich ex Hooker) Hall & Lellinger; Pellaea henryi Christ; Mildella henryi (Christ) Hall & Lellinger; Cheilanthes nitidula subsp. henryi (Christ) Fraser-Jenkins.

Rhizomes ascending to erect, sometimes with creeping branches; scales bicolorous, dark brown with very narrow, light brown margins, subulate-lanceolate. Fronds numerous, clustered or closely spaced. Stipe 6–20 cm Χ 1–1.5 mm, dark brown to nearly black, terete, with ± bicolorous, subulate-lanceolate to linear scales near base, these grading abruptly to sparse, unicellular scales, also with sparse to occasionally dense, short, reddish brown, unicellular hairs, these sometimes mostly adaxial. Lamina oblong to oblong-deltate, 4–12 Χ 3–6 cm, 2-pinnate-pinnatifid, blunt or shortly caudate at apex, leathery, brownish green when dried, glabrous on both surfaces; rachis and costae dark brown, densely shortly hairy. Pinnae 3–5 pairs, sessile or subsessile, basal pair longest, deltate, 2.5–3.5 Χ 2–2.5 cm; pinnules relatively closely spaced, those of basal pinnae 4–6 pairs, adnate to costae; basiscopic pinnules enlarged, basal basiscopic pinnules longer than adjacent acroscopic ones, oblong, 1.5–2 Χ 0.8–1.5 cm, pinnatifid, with 5–7 pairs of lanceolate to deltate segments, other pinnules lanceolate or linear, few lobed to entire. Sori confluent, not interrupted at sinuses or segment tips. False indusia continuous, brown, membranous, margins irregularly dentate to erose, sparsely ciliate. 2n = 58, 116.

On rocks in forests and dry valleys; 200–2400 m. Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, Kashmir, Nepal, Pakistan, Vietnam].

Traditionally, two taxa have been accepted in this complex. Subspecies henryi occurs in the eastern portion of the species range and differs from subsp. nitidula mostly in its denser rachis and stipe pubescence with the hairs often tending to extend around the stipe (vs. restricted to the adaxial side). The ranges of the two subspecies overlap extensively and there are too many intermediates to allow formal recognition of subspecies.

16. Cheilanthes brausei Fraser-Jenkins, Taxon. Revis. Indian Subcontinental Pteridophytes, 122. 2008.

滇西旱蕨  dian xi han jue

Pellaea mairei Brause, Hedwigia 54: 201. 1914, not Cheilanthes mairei Brause (1913); Mildella mairei (Brause) C. C. Hall & Lellinger.

Rhizomes ascending to erect, sometimes with creeping branches; scales bicolorous, dark brown to black with narrow, light brown to reddish brown margins, narrowly lanceolate. Fronds clustered or closely spaced. Stipe 8–16 cm Χ 1.5–2 mm, dark brown, lustrous, ± terete, densely scaly at base, scales often slightly bicolorous, narrowly lanceolate, grading abruptly to moderate to dense, concolorous, linear-subulate (above a slightly expanded base) scales, also moderately to densely pubescent with short, unicellular hairs, mostly adaxially. Lamina ovate-deltate or oblong-deltate, 6–12 Χ 4–7 cm, pinnate-pinnatifid, apex caudate, papery, brownish green when dry, glabrous on both surfaces; rachis dark brown, slightly grooved adaxially, densely shortly hairy and with few small scales. Pinnae 3–5 pairs, basal pair longest, ovate-deltate, 2.5–5 Χ ca. 2 cm, subsessile, apex long caudate (caudate portion ca. 2 cm); segments widely spaced, basiscopic segments enlarged, basal basiscopic segments longer than adjacent acroscopic ones, linear-lanceolate to narrowly oblong, 1.5–2 Χ 0.2–0.3 cm, margins entire, crenulate, or with 1 or 2 small lobes. Sori confluent, not interrupted at lobe sinuses or segment tips. False indusia continuous, appearing somewhat inflated, margins entire to erose, irregularly ciliate. 2n = 120.

* Rock crevices and stone walls; 1000–3200 m. Guizhou, Hunan, Shaanxi (Ningshan), SW Sichuan, Yunnan.

Cheilanthes brausei is similar to C. nitidula, but has long-caudate pinnae, more widely separated segments, and scales extending up the stipe.

17. Cheilanthes bhutanica Fraser-Jenkins & Wangdi in Fraser-Jenkins & Dulawat, Fern Gaz. 18: 220. 2009.

云南旱蕨  yun nan han jue

Pellaea yunnanensis Ching, Acta Phytotax. Sin. 20(2): 235. 1982, not Cheilanthes yunnanense Brause (1913).

Rhizomes ascending to erect, sometimes with creeping branches; scales bicolorous, black with narrow, light brown margin, lanceolate to narrowly lanceolate. Fronds clustered or closely spaced. Stipe to 19 cm Χ 1.5–2 mm, shiny, terete or slightly flattened adaxially, dark brown to nearly black, scaly toward base, scales mostly bicolorous, narrowly lanceolate, glabrous above. Lamina ovate-pentagonal, 9–12 Χ ca. 9 cm, 2-pinnate-pinnatifid, acute to shortly acuminate at tip, papery, brownish green when dry, glabrous on both surfaces; rachis grooved adaxially, glabrous. Pinnae ca. 6 pairs, basal pair largest, deltate, 4–6 Χ ca. 5 cm, shortly stalked (2–3 mm), apex acuminate; pinnules 4 or 5 pairs, basiscopic pinnules enlarged, basal basiscopic pinnules longer than adjacent acroscopic ones, oblong-lanceolate to narrowly deltate, 2–4 Χ 0.8–2 cm, sessile, base unequally cuneate, apex acuminate. Ultimate segments 3 or 4 pairs, oblong, margins mostly entire, 6–15 Χ 3–5 mm. Sori confluent, but mostly interrupted at sinuses. False indusia continuous, margins somewhat erose.

Rock crevices in forests; 1500–1900 m. Sichuan (Kangding), Yunnan (Kunming) [Bhutan].

14. ALEURITOPTERIS Fιe., Mem. Foug., Gen. Filic. 153–154. 1852.

粉背蕨属  fen bei jue shu

Zhang Gangmin (张钢民); George Yatskievych

Leptolepidium K. H. Shing & S. K. Wu; Sinopteris C. Christensen & Ching.

Plants small, usually on rocks. Rhizomes short, erect or ascending; scales brown to black, concolorous or bicolorous with dark, central stripe and lighter margins, mostly narrowly to broadly lanceolate. Fronds numerous, clustered. Stipe and rachis black, dark brown, or reddish brown, shiny, terete or rarely sulcate, scaly proximally or less commonly throughout, occasionally also hairy. Lamina variously shaped, pinnate-pinnatifid to pinnate-tripinnatifid, adaxially glabrous or rarely with hairs or scales, abaxially farinose or less commonly lacking farina, farina white to milky yellow or yellow, otherwise glabrous or occasionally hairy and/or scaly abaxially; rachis grooved adaxially. Pinnae sessile or subsessile, at least some of them often somewhat falcate, basal pinnae usually largest; basal basiscopic segments enlarged, longer than adjacent acroscopic ones. Veins free but often obscure, pinnate, branched or unbranched. Sori consisting of 1–10 sporangia, orbicular, at vein tips, separate when young, often confluent at maturity. False indusia membranous or herbaceous, brown, grayish brown, or rarely light green, often drying brown, continuous, or interrupted, margins entire, erose, lacerate, or fimbriate. Spores globose to globose-tetrahedral or tetrahedral, trilete, perispore reticulate, cristate, echinate, rugulate, or granulate. x = 29, 30.

About 40 species; 29 species in China.

The genus Aleuritopteris is here circumscribed relatively broadly to include Leptolepidium and Sinopteris, which were segregated based on morphological characters that represent specializations within Aleuritopteris. Additionally, most of the species assigned by previous authors to Cheilosoria Trevisan and Notholaena R. Brown are more closely related to Aleuritopteris based on molecular data, which would raise the total number of species in the genus to ca. 70. See Cheilanthes for further discussion of these groups. Circumscription of Aleuritopteris remains controversial, and a stable resolution must await future publication of a comprehensive generic classification for the cheilanthoid ferns that incorporates the full range of morphological, cytological, and molecular data.

1a.       Veins dark brown to black (sometimes partially obscured by farina), stout, prominently raised abaxially; sori with 1(or 2) large sporangia with broad annuli.

2a.       Lamina 7–10 cm; medial pinnae pinnatifid; false indusia interrupted ..........  1. A. grevilleoides

2b.       Lamina 3.5–8 cm; medial pinnae bipinnatifid; false indusia continuous ............  2. A. albofusca

1b.       Veins green (sometimes obscured by farina), slender, not or only slightly raised abaxially; sori consisting of (1 or)2 to several small sporangia with narrow annuli.

3a.       Lamina mostly pentagonal, nearly as wide as long; pinnae lobed to pinnatifid or only proximal 1–3 pairs of pinnae separated by wingless rachis; false indusia continuous, with entire or undulate margins.

4a.       Lamina moderately to densely scaly on tissue and veins abaxially ................  3. A. squamosa

4b.       Lamina lacking scales abaxially or scaly only on rachis and costae.

5a.       Lamina glabrous, lacking farina abaxially.

6a.       Pinnae curved upward; basal pinnae strongly inequilateral, acroscopic pinnules strongly reduced or lacking; lamina coarsely divided, ultimate segments falcate ....................  12. A. duclouxii

6b.       Pinnae patent or obliquely angled upward, moderately inequilateral, basal acroscopic pinnules developed, but shorter than basiscopic ones; lamina finely divided, ultimate segments relatively straight.

7a.       Second pair of pinnae (from lamina base) shorter than basal and third ones  9. A. tamburii

7b.       Second pair of pinnae (from lamina base) shorter than basal pair but longer than third ones          11. A. argentea

5b.       Lamina with white or yellow farina abaxially (note that herbarium specimens dried with heat or treated with chemicals sometimes appear to lack farina and also rare hybrids involving farinose parental taxa sometimes do not produce farina).

8a.       Lamina small, usually less than 5 cm, with snow-white farina abaxially; sori consisting of (1 or)2 or 3 sporangia, discrete.

9a.       Laminae longer than stipes; stipes with scales and twisted hairs, these extending onto rachis and costae .................................................................................................  4. A. pygmaea

9b.       Laminae shorter than stipes; stipes scaly only at base, not hairy; rachis and costae not hairy or scaly.

10a.     Lamina with basal pair of pinnae nearly equilateral, basal acroscopic segments lobed or pinnatifid, nearly as long as opposing basal basiscopic segments .........................  5. A. speciosa

10b.     Lamina with basal pair of pinnae inequilateral, basal acroscopic segments simple, entire, much shorter than opposing basal basiscopic segments ..............................  6. A. niphobola

8b.       Lamina larger, usually more than 5 cm, with milky white or yellow farina abaxially; sori consisting of 3 to several sporangia, often confluent at maturity.

11a.     Scales of rhizome concolorous, dark brown to black.

12a.     Lamina with white farina abaxially and with relatively dense red glands (actually farina-producing glands with a red apical cell) .........................................................  7. A. likiangensis

12b.     Lamina with milky yellow farina abaxially, lacking red glands or some of farina-producing glands with apical cell light brown, rarely somewhat reddish tinged ..................  8. A. veitchii

11b.     Scales of rhizome bicolorous, with a brown to black central stripe and lighter margins.

13a.     Lamina with light yellow to lemon yellow farina abaxially ..................  12. A. duclouxii

13b.     Lamina with white farina abaxially.

14a.     Second pair of pinnae (from lamina base) shorter than basal pair but longer than third ones      11. A. argentea

14b.     Second pair of pinnae (from lamina base) shorter than basal and third ones.

15a.     Medial pinnae rounded or bluntly pointed at tips; distal lobes of pinnae oblong, oblong-triangular, or nearly semicircular, with narrow V-shaped sinuses between lobes  9. A. tamburii

15b.     Medial pinnae attenuate at tips; distal lobes of pinnae linear to narrowly oblong, with broad U-shaped sinuses between lobes
..............................................................................................  10. A. yalungensis

3b.       Lamina triangular-ovate, oblong-lanceolate, or oblong; pinnae usually 5–10 pairs or more, separated from each other by wingless rachis; false indusia usually interrupted, rarely continuous.

16a.     Lamina lacking farina at maturity (young fronds sometimes farinose).

17a.     False indusia continuous, margin undulate; lamina hairy abaxially ............  28. A. subvillosa

17b.     False indusia interrupted, margin laciniate; lamina glandular abaxially or glabrous except for inconspicuous hairs at pinna bases.

18a.     Fronds (15–)25–40 cm; stipe terete; lamina glabrous abaxially or occasionally sparsely and inconspicuously hairy at pinna bases ....................................................  26. A. leptolepis

18b.     Fronds 7.5–15 cm; stipe grooved distally; lamina with brown glands along costae and veins abaxially                                                                                                                  27. A. duthiei

16b.     Lamina with white or yellow farina abaxially.

19a.     Stipe with short, dark brown glands ....................................................  13. A. sichouensis

19b.     Stipe not glandular, but often scaly, occasionally also hairy.

20a.     Stipe, rachis, and costae scaly.

21a.     Scales on stipe broadly lanceolate, distinctly bicolorous, with conspicuous pale margins  23. A. albomarginata

21b.     Scales on stipe narrowly lanceolate, reddish brown, concolorous or mixed with occasional scales having narrow, lighter margins.

22a.     Lamina glabrous or rarely sparsely hairy adaxially; lamina widest at base  24. A. dubia

22b.     Lamina sparsely hairy and with sparse slender scales adaxially; lamina usually widest above base ...............................................................................................................  25. A. rufa

20b.     Stipe scaly at base, sometimes up to rachis, but scales not extending to costae.

23a.     Lamina with milky yellow or golden yellow farina abaxially.

24a.     Stipe black, basalmost stipe scales black, slightly bicolorous, with very narrow, brown margins ........................................................................................................  14. A. ebenipes

24b.     Stipe usually reddish brown to dark brown; basalmost stipe scales concolorous, reddish brown.

25a.     Lamina with golden yellow farina abaxially; stipe scales linear-lanceolate  15. A. chrysophylla

25b.     Lamina with milky yellow farina abaxially; stipe scales broadly lanceolate  16. A. krameri

23b.     Lamina with white farina abaxially.

26a.     Stipe and rachis densely scaly .......................................................  17. A. formosana

26b.     Stipe scaly proximally, rachis not scaly (occasionally sparsely scaly at base in A. anceps).

27a.     Rhizome scales brown, ovate-lanceolate, thin and translucent.

28a.     Lamina hairy along costae and veins ..........................................  28. A. subvillosa

28b.     Lamina lacking hairs .......................................................................  29. A. kuhnii

27b.     Rhizome scales black, lanceolate, relatively thick and opaque.

29a.     Rhizome scales concolorous ..............................................  18. A. gongshanensis

29b.     Rhizome scales bicolorous.

30a.     False indusia poorly developed to nearly absent ........................  19. A. dealbata

30b.     False indusia well developed.

31a.     False indusia interrupted, with laciniate margin ...........................  20. A. anceps

31b.     False indusia continuous or sometimes interrupted, margins entire or undulate.

32a.     Lamina 2–6 cm, thinly herbaceous; false indusia poorly developed, narrow; stipe scales linear-lanceolate ...................................................................  21. A. rosulata

32b.     Lamina (3–)10–20 cm, leathery; false indusia well developed, broad; stipe scales broadly lanceolate ................................................................................  22. A. grisea

1. Aleuritopteris grevilleoides (Christ) G. M. Zhang, comb. nov.

中国蕨  zhong guo jue

Basionym: Cheilanthes grevilleoides Christ in Lecomte, Notul. Syst. 1: 51, f. 1A, B. 1909; Sinopteris grevilleoides (Christ) C. Christensen & Ching.

Rhizomes short, erect; scales bicolorous, black with brown margin, lanceolate. Fronds clustered. Stipe 10–18 cm. Lamina pentagonal, as long as broad, 7–10 cm, almost equally tripartite, leathery, brownish green adaxially when dry, with white farina abaxially. Middle pinna largest, oblong-lanceolate, 6–9 Χ 3–4.5 cm, base abruptly reduced, cuneate and decurrent, connected to lateral pinnae by a narrow wing, apex shortly acuminate, pinnatifid; segments ca. 15 pairs, obliquely patent, linear-lanceolate, medial segments larger than proximal and distal ones, 2–3 Χ 0.3–0.5 cm, margins entire or with few gross teeth. Lateral pinnae deltate, 3.5–6 cm, inequilateral, 2-pinnatifid; proximal basiscopic pinnules especially enlarged, 3–4.5 Χ 1–1.5 cm, pinnatifid, connected to costules by narrow wings. Veins free, pinnate, close together and prominently raised abaxially as rows of tiles on roof. Sori consisting of 1(or 2) large sporangium, sporangia with broad annulus. False indusia narrow, margins divided into deltoid gross teeth, concealing sori.

* Rock crevices along riverbanks or under shrubs; 1000–1800 m. N Sichuan (Qingchuan), W Yunnan (Binchuan, Dayao, Qiaojia).

2. Aleuritopteris albofusca (Baker) Pichi-Sermolli, Nuov. Giorn. Bot. Ital. n.s., 53: 154. 1946.

小叶中国蕨  xiao ye zhong guo jue

Cheilanthes albofusca Baker, Kew Bull. 1895: 54. 1895; C. mairei Brause; Sinopteris albofusca (Baker) Ching; S. hopeiensis C. Christensen & Ching.

Rhizomes short, erect; scales bicolorous, chestnut-black with narrow, brown margin, lanceolate. Fronds clustered. Stipe 4–12 cm. Lamina as long as broad, 3.5–6 cm, tripartite, leathery, dull green adaxially when dry, abaxially farinose, farina white. Middle pinna somewhat rhombic, as long as wide, 3–5 cm, base cuneate and decurrent, usually connected with lateral pinnae by a narrow wing, rarely free, apex acuminate, 2-pinnatifid; pinnules 4 or 5 pairs, proximal pair largest, 2–3 Χ 0.4–0.8 cm, linear-lanceolate, apex obtuse or acute; ultimate segments 6–9 pairs, oblong or triangular, margins entire, apex obtuse or obtusely acute. Lateral pinnae triangular, 2–4 Χ 2–3 cm, inequilateral, 2-pinnatifid; proximal basiscopic segment especially enlarged, 1–2 Χ 0.3–0.6 cm, lanceolate or oblong, apex shortly acuminate or acute, pinnatifid, connected to costule with narrow wings. Veins free, prominently raised abaxially. Sori consisting of 1(or 2) large sporangium, sporangia with broad annulus. False indusia usually wide, continuous, margins irregularly undulate-crenulate.

* Rock crevices under trees and shrubs; 500–3200 m. Gansu (Wenxian), Guizhou, Hebei, S Hunan, Sichuan, Xizang, Yunnan.

3. Aleuritopteris squamosa (C. Hope & C. H. Wright) Ching, Hongk. Nat. 10: 199. 1941.

毛叶粉背蕨  mao ye fen bei jue

Pellaea squamosa C. Hope & C. H. Wright in S. T. Dunn, J. Linn. Soc., Bot. 35: 518. 1903; Cheilanthes hopeana C. Christensen; Doryopteris squamosa C. Christensen.

Rhizomes short, erect; scales bicolorous, black, with light brown margin, lanceolate. Fronds clustered. Stipe 5–15 cm, chestnut-colored, densely covered with light brown, broadly lanceolate, often slightly bicolorous scales, becoming sparsely scaly with age. Lamina pentagonal, nearly as long as broad, 5–10 cm, apex shortly acuminate, 3-pinnatipartite, thick papery when dry, brownish green, adaxially glabrous, abaxially with snow-white farina and membranous, translucent, light brown, broadly lanceolate scales, scale margins erose-serrulate. Pinnae 5–7 pairs, connected by a narrow wing; basal pair largest, subtriangular, 4–5 cm, obliquely patent upward, 2-pinnatipartite; pinnules 4–6 pairs, proximal basiscopic pinnule largest, 3–3.5 Χ 1–1.5 cm, obliquely patent, ultimate segments 3–4 pairs, 3–8 Χ ca. 2 mm; second and upper pairs of pinnae gradually shortened distally, lanceolate or oblong-lanceolate, apex shortly acute, base decurrent and connected to rachis with broad wings, pinnatipartite. Sori consisting of few sporangia. False indusia continuous, narrow, membranous, margins entire.

* Rock crevices or under shrubs in dry-hot valleys; 400–1000 m. Hainan, S Yunnan.

This species is quite distinct and is distinguished from other members of Aleuritopteris with pentagonal fronds by the presence of scales on abaxial lamina, which also has dense, snow-white farina.

4. Aleuritopteris pygmaea Ching in S. K. Wu, Acta Phytotax. Sin. 19(1): 61. 1981.

矮粉背蕨  ai fen bei jue

Rhizomes short, ascending; scales bicolorous, dark brown with light brown margin, lanceolate. Fronds clustered. Stipe 1–2 cm, reddish brown, densely covered with concolorous brown scales and fibrous, twisted hairs, these extending onto rachis and costae. Lamina ovate, 1.3–1.5 Χ ca. 1 cm, bipinnatifid, papery when dry, adaxially glabrous, abaxially with snow-white farina and sparse hairs. Pinnae 3 or 4 pairs, opposite, slightly oblique upward; basal pair of pinnae widely separated from adjacent one, ca. 0.6 cm, pinnatipartite, ultimate segments 3 or 4 pairs, obliquely patent, basal basiscopic segment larger, entire; second pair of pinnae subequal to or slightly longer than basal one, base decurrent or not. Veins obscure. Sori sparse, discontinuous, consisting of 2 or 3 sporangia. False indusia continuous, narrow, margins entire.

* Rock crevices in valleys; 3800–3900 m. Xizang.

This species is the smallest in Aleuritopteris. It is distinguished from other members by its reddish brown stipe, which is densely covered with scales and twisted, long hairs.

5. Aleuritopteris speciosa Ching & S. K. Wu, Acta Phytotax. Sin. 19(1): 61. 1981.

美丽粉背蕨  mei li fen bei jue

Rhizomes short, erect; scales concolorous, reddish brown, narrowly lanceolate. Fronds usually clustered. Stipe 10–18 cm, reddish brown, lustrous. Lamina pentagonal, nearly as long as broad, ca. 4 cm, apex shortly acuminate, pinnate-bipinnatifid, leathery to thin leathery when dry, dark green, adaxially glabrous or with sparse white farina, abaxially with dense snow-white farina; rachis, costae, and costules dark brown, lustrous. Pinnae 3 or 4 pairs, opposite, oblique, separated from each other; basal pair largest, deltate, ca. 2 Χ 2 cm, apex acuminate, 2-pinnate-pinnatifid, acroscopic pinnules pinnatifid, proximal basiscopic pinnule larger, ca. 1.5 Χ ca. 1 cm, elliptic, 1-pinnate-pinnatifid; second and upper pairs of pinnae gradually shortened distally, pinnatifid. Veins pinnate, obscure. Sori consisting of 1 or 2 sporangia, discrete. False indusia thick membranous, pale green, broad.

* Rock crevices; ca. 3100 m. E Xizang (Zhag’yab).

A rare small fern, known thus far only from the type (PE).

6. Aleuritopteris niphobola (C. Christensen) Ching, Hongk. Nat. 10: 197. 1941.

雪白粉背蕨  xue bai fen bei jue

Cheilanthes niphobola C. Christensen, Acta Hort. Gothob. 1: 88. 1924; Aleuritopteris niphobola var. pekingensis Ching & Hsu; A. niphobola var. concolor Ching.

Rhizomes short, erect; scales concolorous or often bicolorous, uniformly reddish brown or with dark central stripe and lighter margin, lanceolate. Fronds clustered. Stipe 4–13 cm, reddish brown or ebony, glabrous or occasionally with sparse scales as those of rhizome. Lamina pentagonal, nearly as long as broad, 1.5–4 cm, tripinnatifid, papery or herbaceous when dry, dark green, adaxially glabrous, abaxially with snow-white farina; rachis and costae of same color as stipe. Pinnae 2 or 3 pairs, connected to rachis by narrow wings; basal pair largest, triangular, inequilateral, bipinnatifid; basal basiscopic pinnule largest, pinnatifid, ultimate segments narrowly oblong, margins entire; second pinnae pinnatifid or entire, oblong-lanceolate or lanceolate. Veins pinnate, obscure. Sori consisting of 1–3 sporangia, discrete. False indusia continuous, broad, pale green, margins entire.

* Rock crevices; 300–3400 m. Gansu, Hebei, Nei Mongol, Ningxia, Shaanxi, Shanxi, Sichuan, Xizang.

Aleuritopteris niphobola var. concolor was based on a specimen from Gansu apparently lacking abaxial frond farina. Examination of the type specimen at PE reveals that it possesses dense farinose glands abaxially, but that the farinose exudate was eliminated during the drying process.

7. Aleuritopteris likiangensis Ching in S. K. Wu, Acta Phytotax. Sin. 19(1): 64. 1981.

丽江粉背蕨  li jiang fen bei jue

Rhizomes short, erect; scales concolorous, dark brown to black, narrowly lanceolate. Fronds clustered. Stipe 5–20 cm, ebony, lustrous, base covered with reddish brown, concolorous, lanceolate scales. Lamina ovate-pentagonal, (4–)6–15 Χ (4–)6–12 cm, 3-pinnatipartite, apex elongate-acuminate or caudate, thin leathery when dry, dark green, adaxially glabrous, abaxially with white farina, also appearing to have contrasting red glands (these actually red terminal cells of farinose glands); rachis and costae of same color as stipe. Pinnae 2–4 pairs, sessile, basal pair largest, subtriangular, 2-pinnatipartite; pinnules 4 or 5 pairs, basal basiscopic pinnule largest, 4–6.5 cm Χ 3–4 mm 1-pinnatipartite; second and third pinnae gradually shortened distally, 1-pinnatipartite, narrowly triangular. Veins pinnate, obscure. Sori consisting of 3 to several sporangia, confluent at maturity False indusia continuous, margins entire.

* Rock crevices by dry, hot valleys; 1500–2900 m. SW Sichuan, NW Yunnan.

Specimens of A. likiangensis and A. veitchii are often difficult to separate. Many specimens determined as A. veitchii have at least some colored apical cells on farinose glands, especially along costae, but they are tan rather than red. The species are retained here provisionally, but in the future may prove to be merely forms of a single species.

8. Aleuritopteris veitchii (Christ) Ching, Hongk. Nat. 10: 200. 1941.

金爪粉背蕨  jin zhua fen bei jue

Doryopteris veitchii Christ, Bull. Acad. Int. Gιogr. Bot. 16: 134. 1906; Aleuritopteris cremea Ching; Cheilanthes veitchii (Christ) Ching.

Rhizomes short, ascending; scales concolorous, dark brown to black, narrowly lanceolate. Fronds clustered. Stipe 8–15 cm, chestnut red to dark brown, lustrous, with reddish brown, linear scales at base. Lamina ovate-pentagonal, 7–15 Χ 5–10 cm, pinnate-bipinnatifid, papery when dry, brownish green, adaxially glabrous, abaxially with white to milky yellow farina, often also with at least some of farinose glands having apical cell tan to pale red; rachis and costae of same color as stipe. Pinnae 4 or 5 pairs, sessile, separated from each other, basal pair largest, subtriangular, 1-pinnate-pinnatifid; pinnules 4–7 pairs, basal basiscopic pinnule largest, 3–4 Χ 1–2 cm, 1-pinnatifid; ultimate segments oblong or ovate. Veins pinnate, obscure. Sori consisting of 3 to several sporangia, borne at vein tips, False indusia continuous, membranous, margins entire.

* Rock crevices; 1300–2000 m. Guangxi, S and SW Sichuan.

In earlier references, this species sometimes was confused with Aleuritopteris duclouxii (Christ) Ching var. sulphurea Ching, which has bicolorous rhizome scales.

9. Aleuritopteris tamburii (Hooker) Ching, Hongk. Nat. 10: 198. 1941.

阔羽粉背蕨  kuo yu fen bei jue

Pellaea tamburii Hooker, Sp. Fil. 2: 134. 1858; Allosorus tamburii (Hooker) Kuntze; Cheilanthes argentea (S. G. Gmelin) Kunze var. tamburii (Hooker) Beddome; C. tamburii (Hooker) Moore; Doryopteris tamburii (Hooker) C. Christensen.

Rhizomes short, erect; scales bicolorous, brown or reddish brown with lighter margin, lanceolate. Fronds clustered. Stipe 10–35 cm, reddish brown, lustrous, scaly at base, scales mostly bicolorous, narrowly lanceolate. Lamina pentagonal, 8–15(–24) Χ 6–12(–18) cm, tripinnatifid to pinnate-bipinnatifid, herbaceous or papery when dry, adaxially glabrous, abaxially with snow-white farina or rarely glabrous; rachis and costae chestnut-brown. Pinnae 3–5 pairs, sessile, connected by broad wings, distal lobes oblong, oblong-triangular, or nearly semicircular, with narrow V-shaped sinuses between lobes; basal pair largest, 7–8 Χ 4–5 cm, triangular, obliquely patent upward, 2-pinnatifid, medial pinnae rounded or bluntly pointed at tips; pinnules 4 or 5 pairs, basal basiscopic pinnule largest; second pair of pinnae shorter than basal and third ones, simple, triangular; other pairs gradually shortened distally; ultimate segments deltate or ovate, broad, apex obtuse, closely approaching. Veins pinnate, distinct. Sori consisting of 3 to several sporangia, borne at vein tips. False indusia continuous, narrow, margins entire.

Thickets, rock crevices in dry hot valleys; 1900–2700 m. S Sichuan, NW Yunnan [India, Nepal].

This species is easily distinguished from related taxa in its large fronds that are coarsely dissected and with broad, somewhat rounded ultimate segments.

1a. Fronds with white farina abaxially .......................................................................  9a. var. tamburii

1b. Fronds glabrous, lacking farina abaxially ................................................................  9b. var. viridis

9a. Aleuritopteris tamburii var. tamburii

阔羽粉背蕨(原变种)  kuo yu fen bei jue (yuan bian zhong)

Fronds with white farina abaxially.

Thickets, rock crevices in dry hot valleys; 1900–2700 m. S Sichuan, NW Yunnan [India, Nepal].

9b. Aleuritopteris tamburii var. viridis H. S. Kung, Acta Bot. Yunnan. 5(4): 356–358. 1983.

Chinese  pinyin

Fronds glabrous abaxially.

* Rare; rock crevices; ca. 2000 m. SW Sichuan (Yan-Bian).

10. Aleuritopteris yalungensis H. S. Kung, Acta Bot. Yunnan. 5(4): 356. 1983.

雅砻粉背蕨  ya long fen bei jue

Rhizomes short, ascending; scales bicolorous, dark brown with lighter margin, linear-lanceolate. Fronds clustered. Stipe 18–25 cm, brown or reddish brown, lustrous. Lamina ovate-pentagonal, 15–24 Χ 12–18 cm, 3-pinnatifid, papery when dry, adaxially glabrous, abaxially with white farina; rachis and costae brown. Pinnae 5 or 6(–8) pairs, angled obliquely upward, decurrent at base, distal lobes linear to narrowly oblong, with broad U-shaped sinuses between lobes; basal pair largest, 8–12 Χ 4–9 cm, triangular, inequilateral, 2-pinnatifid, medial pinnae attenuate at tips; pinnules ca. 5 pairs, alternate, basal basiscopic pinnule largest, 5–7 Χ 1.5–2 cm, lanceolate, pinnatifid; second pinnae short than basal one and third one, 2–4 cm Χ 5–8 mm, other pairs gradually shortened distally, 1-pinnatifid or simple, lanceolate; ultimate segments lanceolate, narrow, distantly apart. Veins pinnate, obscure. Sori consisting of 3 to several sporangia, borne at vein tips. False indusia continuous, gray, membranous, margins entire.

* Rare; ca. 2000 m. SW Sichuan (Dechang).

The species resembles A. tamburii, but differs in its finely dissected fronds and lanceolate ultimate segments.

11. Aleuritopteris argentea (S. G. Gmelin) Fιe, Gen Fil. 154. 1852.

银粉背蕨  yin fen bei jue

Pteris argentea S. G. Gmelin, Novi Comment. Acad. Sci. Imp. Petrop. 12: 519. 1768; Allosorus argentea (S. G. Gmelin) C. Presl.; Cheilanthes argentea (S. G. Gmelin) Kunze; Doryopteris argentea (S. G. Gmelin) Christ.

Rhizomes erect or ascending; scales bicolorous, brown with lighter margin, lanceolate. Fronds clustered. Stipe 10–22 cm, chestnut-colored to black, lustrous, scaly at base. Lamina pentagonal or ovate-pentagonal, 5–13 Χ 5–10 cm, 3-pinnatifid, papery or somewhat leathery when dry, adaxially glabrous; rachis and costae of same color as stipe. Pinnae 3–5 pairs, opposite; basal pair largest, 3–5 Χ 2–4 cm, right-triangular; pinnules 3 or 4 pairs, proximal basiscopic pinnule largest, 2–2.5 cm Χ 5–10 mm, oblong-lanceolate, pinnatifid or simple; other pairs gradually shortened distally, 1-pinnatifid or simple, lanceolate; ultimate segments deltate or falcate. Veins pinnate, obscure. Sori consisting of 3 to several sporangia, confluent at maturity. False indusia continuous, yellowish green, membranous, margins entire. 2n = 116.

Limestone crevices, cliffs, rock walls; below 3900 m. Widely distributed throughout China [Bhutan, Japan, Korea, Mongolia, Nepal, Russia].

1a. Fronds with white or pale-yellow farina abaxially .............................................  11a. var. argentea

1b. Fronds glabrous, lacking farina abaxially ...........................................................  11b. var. obscura

11a. Aleuritopteris argentea var. argentea

银粉背蕨(原变种)  yin fen bei jue (yuan bian zhong)

Aleuritopteris argentea var. flava Ching & S. K. Wu; A. argentea var. geraniifolia Ching & S. K. Wu; A. argentea var. major Ching; A. michelii (Christ) Ching; A. subargentea Ching; A. qianguiensis W. M. Chu & H. G. Zhou; Cheilanthes subargentea (Ching) C. M. Kuo; Doryopteris michelli Christ.

Fronds with white or pale yellow farina abaxially.

Limestone crevices or rocky wall; below 3900 m. Widely distributed throughout China [Bhutan, Japan, Korea, Mongolia, Nepal, Russia].

Aleuritopteris argentea var. qianguiensis refers to specimens in which the laminae appear glandular but not farinose abaxially. This likely resulted during the drying process, as application of heat during pressing can cause the farina to melt and soak into the pressing papers.

11b. Aleuritopteris argentea var. obscura (Christ) Ching, Hong Kong Naturalist 10: 198. 1941.

陕西粉背蕨  shan xi fen bei jue

Cheilanthes argentea Kunze var. obscura Christ, Nuov. Giorn. Bot. Ital. N. s. 4: 88. 1897; A. shensiensis Ching; A. nuda Ching.

Fronds finely dissected, green, lacking farina abaxially.

* Rock crevices; below 2600 m. Gansu, Guizhou, Hebei, Henan, Liaoning, Qinghai, Shandong, Shaanxi, Shanxi, Sichuan, Yunnan.

12. Aleuritopteris duclouxii (Christ) Ching in Hongk. Nat. 10: 199. 1941.

裸叶粉背蕨  luo ye fen bei jue

Doryopteris duclouxii Christ, Bull. Acad. Int. Gιogr. Bot. 11: 231. 1902; D. mairei Brause; D. muralis Christ; Cheilanthes duclouxii (Christ) Ching.

Rhizomes short, ascending; scales bicolorous, dark brown to black with lighter margin, lanceolate. Fronds clustered. Stipe 6–25 cm, reddish brown to black, lustrous, scaly at base; scales mostly bicolorous, narrowly lanceolate. Lamina pentagonal or ovate-pentagonal, nearly as long as broad, 5–13 Χ 5–10 cm, apex acuminate, pinnate-pinnatifid to tripinnatifid, thinly leathery when dry; rachis and costae of same color as stipe. Pinnae 2 or 3 pairs, basal pair largest, 3–5 Χ 2–4 cm, triangular; pinnules 3 or 4 pairs, basal basiscopic pinnule largest, 2–2.5 cm Χ 5–10 mm, oblong-lanceolate, pinnatifid or simple; other pairs gradually shortened distally, 1-pinnatifid or simple, lanceolate; ultimate segments deltate or falcate. Veins pinnate, obscure. Sori consisting of several sporangia, confluent at maturity. False indusia continuous, yellowish green, membranous, margins entire.

* Rock crevices or rock outcrops in forests; 800–2300 m. Hunan, Guangxi, Guizhou, Sichuan, Yunnan.

1a. Lamina glabrous on both surfaces ...................................................................  12a. var. duclouxii

1b. Lamina glabrous adaxially, with light yellow farina abaxially .............................  12b. var. sulphurea

12a. Aleuritopteris duclouxii var. duclouxii

裸叶粉背蕨(原变种)  luo ye fen bei jue (yuan bian zhong)

Lamina relatively coarsely dissected, glabrous on both surfaces; pinnae strongly inequilateral, basal basiscopic pinnules much longer than acroscopic ones.

* Rock crevices; 800–2300 m. Guangxi, Guizhou, Hunan, Sichuan, Yunnan.

12b. Aleuritopteris duclouxii var. sulphurea (Ching) Ching, Hongk. Nat. 10: 199. 1941.

硫磺粉背蕨  liu huang fen bei jue

Cheilanthes duclouxii Christ var. sulphurea Ching, Icon. Filic. Sin. 3: 133. 1935.

Lamina finely dissected, glabrous adaxially, with light yellow farina abaxially, basal basiscopic pinnules longer to much longer than acroscopic ones.

* Rock crevices or in woods; 1300–2200 m. Sichuan, Yunnan.

13. Aleuritopteris sichouensis Ching & S. K. Wu, Acta Phytotax. Sin. 19(1): 69. 1981.

西畴粉背蕨  xi chou fen bei jue

Rhizomes short, erect; scales bicolorous, brown with lighter margin, subulate. Fronds clustered. Stipe 15–25 cm Χ 1–1.5 mm, black, lustrous, scaly only at base, densely covered with short, dark brown glands. Lamina ovate-deltate or ovate-lanceolate, 12–15 Χ 8–10 cm, pinnate-bipinnatifid, leathery when dry, adaxially glabrous, abaxially with white farina; rachis and costae with same glands as stipe; pinnae 8–10 pairs, opposite, basal ones largest, ovate-lanceolate, bipinnatifid; pinnules 6–8 pairs, alternate; proximal basiscopic pinnules largest, linear-lanceolate, 2–2.5 cm Χ 3–4 mm, pinnatifid; ultimate segments 4 or 5 pairs, deltate; second and upper pairs of pinnae gradually shortened distally, bipinnatifid, lanceolate. Veins obscure. Sori consisting of 3 or 4 sporangia. False indusia continuous, poorly developed, narrow, margins entire.

* Limestone cliffs; 1400–1500 m. S Yunnan (Xichou).

The species is distinguished from other members by its glandular stipe and rachis, subulate rhizome scales, and very narrow false indusia.

14. Aleuritopteris ebenipes X. C. Zhang, Acta Phytotax. Sin. 32: 94. 1994.

黑柄粉背蕨  hei bing fen bei jue

Rhizomes short, ascending; scales inconspicuously bicolorous, black with very narrow brown margin, lanceolate. Fronds clustered. Stipe 5–12 cm, black, lustrous, scaly at base; basalmost scales black, with narrow brown margin, grading to concolorous, lanceolate. Lamina deltate, 5–10 Χ 4–8 cm, papery, adaxially glabrous, abaxially with milky yellow farina, pinnate-bipinnatifid; rachises similar to stipes; pinnae 5 or 6 pairs, sessile, basal pair largest, deltate, strongly inequilateral, 2–4 Χ 1.5–3 cm, bipinnatifid; pinnules 6 or 7 pairs, proximal basiscopic pinnules largest, oblong, up to 3 cm, pinnatifid; other pairs gradually shortened distally, 1-pinnatifid or simple, lanceolate; ultimate segments deltate to narrowly oblong or lanceolate. Veins pinnate, obscure. Sori consisting of several sporangia, confluent at maturity. False indusia continuous, narrow, membranous, with a subentire margin.

* Rock crevices; 500–1300 m. Guizhou (Danzhai, Guiding, Pingfa).

This species is similar to A. veitchii in its lamina being covered with milky yellow farina abaxially, but differs in its bicolorous scales with a brown, narrow margin, which in some specimens can be difficult to detect. The difference noted in couplet 3 of the key of the triangular-ovate lamina in A. ebenipes (vs. pentagonal in A. veitchii) also may provide a clue for distinguishing these two species. However, some specimens can be difficult to assign to one or the other of these taxa. Further research is needed on separation of A. ebenipes from A. veitchii.

15. Aleuritopteris chrysophylla (Hooker) Ching, Hong Kong Naturalist 10: 201. 1941.

金粉背蕨  jin fen bei jue

Cheilanthes chrysophylla Hooker, Sp. Fil. 2: 113–114. 1852; Aleuritopteris flavopygmaea S. R. Ghosh; A. humatifolia X. C. Zhang & L. Shi; Cheilanthes argentea Kunze var. chrysophylla (Hooker) Hooker; C. farinosa Kaulfuss var. chrysophylla (Hooker) C. B. Clarke.

Rhizomes short, erect; scales concolorous, dark brown, linear-lanceolate to lanceolate. Fronds clustered. Stipe 2–10 cm Χ ca. 1 mm, reddish brown or ebony, lustrous, sparsely scaly above base; scales concolorous, reddish brown, linear-lanceolate scales. Lamina ovate-deltate, 3–5 Χ 2–3 cm, pinnate-pinnatifid to bipinnatifid, papery or thinly leathery when dry, adaxially glabrous, abaxially with golden yellow farina; rachis and costae same color as stipe; pinnae 3 or 4 pairs, subopposite, sessile, basal pair largest triangular, pinnatifid; pinnules 3 or 4 pairs, alternate; basal basiscopic pinnules largest, lanceolate; second and upper pairs of pinnae gradually shortened distally, pinnatifid, lanceolate. Veins obscure. Sori consisting of several sporangia, confluent at maturity. False indusia continuous, membranous, margin undulate. 2n = 60.

Rock crevices; 1000–2600 m. Guangxi, Hainan, Yunnan [Bhutan, India, Nepal].

16. Aleuritopteris krameri (Franchet & Savatier) Ching, Hong Kong Naturalist 10: 202. 1841.

克氏粉背蕨  ke shi fen bei jue

Cheilanthes krameri Franchet & Savatier, Enum. Pl. Jap. 2: 212, 619–620. 1879. Aleuritopteris agetae Saiki. Cheilanthes agetae (Saiki) C. M. Kuo.

Rhizomes short, erect, scales bicolorous, black with narrow brown margin, lanceolate. Fronds clustered. Stipe 10–25 cm Χ 1–1.5 mm, dark brown, lustrous, scaly from base to ca. midpoint; basalmost scales similar to rhizome scales, grading abruptly to concolorous, orangish brown, and broadly lanceolate. Lamina triangular or ovate-triangular, 6–20 Χ 4–12 cm, pinnate-bipinnatifid, papery or herbaceous when dry, adaxially glabrous, abaxially with milky yellow to white farina; pinnae 5–9 pairs, separated from each other by wingless rachis, basal pair triangular or triangular-lanceolate, 2–5 Χ 1–3 cm, bipinnatifid; basal basiscopic pinnules largest, pinnatifid; second and third pairs of pinnae with same shape as basal pair, but shorter and narrower; distal pinnae lanceolate. Veins obscure. Sori consisting of 5–7 sporangia, confluent at maturity. False indusia usually interrupted, broad, margin undulate to irregularly crenate, somewhat corrugated, usually farinose externally.

Rock crevices on slopes; 1800–2500 m. Taiwan [Japan, Thailand].

The status of the Taiwanese endemic A. agetae remains unresolved. This taxon is said to differ from A. krameri s.s. in its milky yellow (vs. white) farina and stipe scales that are sometimes slightly bicolorous. Ralf Knapp (pers. comm.), the author of a recent authoritative guide to the pteridophytes of Taiwan, has stated that where the two taxa grow together they are readily distinguishable in the field. However, based on examination of herbarium material collected in Taiwan, the present authors could not consistently discern two discrete morphological entities. Some of the specimens from mainland China also appear to have milky-colored farina. The situation requires further study.

17. Aleuritopteris formosana (Hayata) Tagawa, Acta Phytotax. Geobot. 14: 191. 1952.

台湾粉背蕨  tai wan fen bei jue

Cheilanthes formosana Hayata Enum. Pl. Form. 612. 1906; Cheilanthes anceps Blanford var. brevifrons Khullar; C. brevifrons (Khullar) Khullar.

Rhizomes short, erect; scales bicolorous, black with pale brown margin, narrowly lanceolate. Fronds clustered. Stipe 5–23 cm Χ 1–2 mm, dark brown, lustrous, scaly up to rachis; scales bicolorous, grading to concolorous above stipe midpoint, lanceolate. Lamina oblong-lanceolate, 4–17 Χ 2–7 cm, pinnate-bipinnatifid, papery when dry, adaxially glabrous, abaxially with white farina, rachis and costae same color as stipe; pinnae 4–6 pairs, discrete along rachis, basal pair triangular, bipinnatifid; acroscopic ones smaller, basal basiscopic pinnules largest, 1.5–2.5 Χ 0.5–1 cm, lanceolate, pinnatifid; second pair of pinnae oblong-lanceolate, shorter and narrower than basal one; third and upper pairs of pinnae lanceolate. Veins obscure. Sori consisting of several sporangia, confluent at maturity. False indusia membranous, relatively broad, interrupted, margin laciniate. 2n = 60.

Rock crevices on slopes; 600–2000 m. Fujian, Guangdong, Guangxi, Guizhou, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, India, Nepal, Pakistan, Philippines, Thailand].

18. Aleuritopteris gongshanensis G. M. Zhang, Novon 14: 513–515. 2004.

贡山粉背蕨  gong shan fen bei jue

Plants 20–30 cm. Rhizomes short, erect; scales concolorous, black or dark brown, concolorous, lanceolate, ca. 0.5 Χ 6 mm. Fronds clustered. Stipe 10–18 cm Χ 1–1.5 mm, ebeneous, lustrous, scaly at base (scales similar to those of rhizome). Lamina ovate-deltate, 8–13 Χ 4.5–8.5 cm, apex caudate, papery or somewhat leathery, adaxially glabrous, abaxially with white farina, pinnate-bipinnatifid; rachises similar to stipes, grooved adaxially; pinnae 5–8 pairs, usually opposite or sometimes alternate, oblique, sessile, basal pinnae largest, deltate, strongly inequilateral, 4–7 Χ 4–5 cm, bipinnatifid; pinnules 4–7 pairs, alternate; acroscopic pinnules much smaller, usually simple, ca. 8 Χ 3 mm, apex obtuse, broadly adnate to costa; basal basiscopic pinnules largest, oblong, ca. 4 Χ 1.5 cm, pinnatifid; ultimate segments 4–5 pairs, oblong to ovate, middle ones to 1.5 Χ 0.3 cm; second pinnae 1.5–4 cm, apart from basal ones, similar but smaller; upper pairs gradually shortened distally. Sori consisting of several sporangia, confluent at maturity. False indusia narrow, membranous, continuous, with a denticulate margin.

* Under thickets in dry, hot valleys; 1000–1700 m. C and NW Yunnan.

This species is similar to Aleuritopteris veitchii in its concolorous rhizome scales, but differs in its ovate-deltate fronds with white farina abaxially, false indusia margin distinctly lacerate, and spores with a reticulate-cristate perine.

19. Aleuritopteris dealbata Fιe, Mem. Foug., Gen. Filic. 154. 1852.

无盖粉背蕨  wu gai fen bei jue

Cheilanthes dealbata Don (1825, non Pursh, 1813); Cheilanthes doniana Fraser-Jenkins & Khullar; Aleuritopteris doniana S. K. Wu; A. interrupta Saiki; A. sikkimensis Ghosh.

Rhizomes short, erect; scales bicolorous, black with pale margin, lanceolate. Fronds clustered. Stipe (15–)20–35 cm Χ 1.5–2 mm, dark brown to black, sparsely scaly at base, scales lanceolate, those at stipe base bicolorous, dark brown with lighter margins, often grading to concolorous above. Lamina lanceolate to somewhat oblong-lanceolate, 20–40 Χ 7–16 cm, apex acuminate, pinnate-bipinnatifid, papery or thinly leathery when dry, adaxially glabrous, abaxially with white farina; rachis and costae same color as stipe; pinnae 5–20 pairs, sessile, distinct along rachis, basal pair of pinnae largest, inequilateral, triangular-lanceolate, bipinnatifid; pinnules ca. 10 pairs, acroscopic ones smaller, basal basiscopic pinnules largest, 3–4 cm Χ 1–1.5 mm, pinnatifid; second and third pairs of pinnae of same shape as basal one, but slightly shorter; fourth and upper pairs of pinnae lanceolate. Sori consisting of several sporangia, confluent at maturity. False indusia poorly developed, very narrow, indistinct.

On rocks; 600–1500 m. Guizhou (Xingyi), Yunnan [Bhutan, India, Nepal].

This beautiful species is distinguished from its relatives by its large fronds and poorly developed false indusia. It was called A. doniana in much of the literature on Asian ferns, but the valid publication of A. dealbata predates it by more than 150 years.

20. Aleuritopteris anceps (Blanford) Panigrahi, Bull. Bot. Surv. India 2: 321. 1961.

粉背蕨  fen bei jue

Cheilanthes anceps Blanford, J. Simla Naturalists’ Soc. 1(2): 21. 1886; Aleuritopteris farinosa Fιe var. anceps (Blanford) Ching; A. pseudofarinosa Ching & S. K. Wu; A. pseudofarinosa var. glandulosa H. G. Zhou; A. wuyishanensis Ching; Cheilanthes farinosa Kaulfuss var. anceps (Blanford) Blanford; C. pseudofarinosa (Ching & S. K. Wu) K. Iwatsuki.

Rhizomes short, erect; scales bicolorous, black with pale brown margin, lanceolate. Fronds clustered. Stipe 8–30 cm Χ 1–2 mm, dark brown, lustrous, scaly toward base, scales sometimes extending up stipe to rachis, those at stipe base bicolorous, dark brown to black with lighter margins, some of distal scales concolorous or darker-tipped. Lamina ovate-deltate, oblong, or ovate-lanceolate, 10–25 Χ 5–10 cm, pinnate-bipinnatifid, papery or thinly leathery when dry, adaxially glabrous, abaxially with white farina; rachis sometimes with a few scales similar to those of stipe; pinnae 5–10 pairs, distinct along rachis, basal pair inequilateral, triangular, bipinnatifid; pinnules 5–8 pairs, basal basiscopic pinnules largest much longer than adjacent acroscopic ones, lanceolate, pinnatifid; second and upper pairs of pinnae lanceolate. Veins obscure. Sori consisting of several sporangia, confluent at maturity. False indusia membranous, interrupted, with a laciniate margin. 2n = 58, 60, 116.

Rock crevices on slopes; 300–2600 m. Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Yunnan, Zhejiang [Bhutan, India, Nepal, Pakistan].

21. Aleuritopteris rosulata (C. Christensen) Ching, Hong Kong Naturalist 10: 197. 1941.

莲座粉背蕨  lian zuo fen bei jue

Cheilanthes rosulata C. Christensen, Acta Hort. Gothob. 1: 89. 1924.

Rhizomes short, erect; scales bicolorous, dark brown with light brown margins, narrowly lanceolate. Fronds clustered. Stipe 3–8 cm, reddish brown, lustrous, sparsely scaly, scales concolorous to slightly bicolorous, thin, light brown, linear-lanceolate. Lamina ovate-pentagonal, slightly longer than broad, 2–5 cm, 3-pinnatipartite, thinly herbaceous when dry, adaxially glabrous or sparsely farinose, abaxially with white farina, rachis reddish brown. Pinnae 3–5 pairs; basal pair of pinnae ovate-triangular, apex obtusely acute, basal basiscopic pinnule largest, longer than adjacent acroscopic ones, ca. 1 cm, pinnatipartite, ultimate segments 4 or 5 pairs; second pair of pinnae ovate-lanceolate, ca. 2 Χ 1 cm, pinnatipartite; other pinnae gradually shortened distally. Sori sparse, consisting of mostly 2 or 3 sporangia, not fully confluent at maturity. False indusia narrow, membranous, continuous, margins entire to slightly undulate.

* Rock crevices; ca. 1500–2700 m. W Sichuan.

A rare fern with laminae thin and herbaceous. The species was originally described as having dimorphic fronds, but subsequent collections have shown that this is incorrect.

22. Aleuritopteris grisea (Blanford) Panigrahi, Bull. Bot. Surv. Ind. 2: 321. 1961.

阔盖粉背蕨  kuo gai fen bei jue

Cheilanthes grisea Blanford, J. Simla Naturalists’ Soc. 1(2): 21. 1886; Aleuritopteris farinosa (Kaulfuss) Fιe var. grisea (Blanford) Ching; A. grisea var. alpina (Ching ex S. K. Wu) S. K. Wu; A. platychlamys Ching; A. platychlamys Ching var. alpina Ching; A. stenochlamys Ching; Cheilanthes farinosa Kaulfuss f. minor C. B. Clarke & Baker; C. farinosa var. grisea (Blanford) Blanford; C. farinosa var. tenera C. B. Clarke & Baker; C. platychlamys (Ching) Fraser-Jenkins.

Rhizomes short, erect; scales slightly bicolorous, black with brown margin, narrowly lanceolate. Fronds clustered. Stipe 5–30 cm Χ 0.8–2 mm, dark brown, lustrous, scaly at base or to ca. midpoint, scales concolorous, reddish brown, broadly lanceolate. Lamina oblong-lanceolate or ovate-lanceolate, apex acuminate, (3–)10–20 Χ 2–14 cm, tripinnatifid, papery or thinly leathery when dry, adaxially glabrous, abaxially with white farina; rachis and costae same color as stipe; pinnae 6–12 pairs, distinct along rachis, basal pair of pinnae subtriangular, (1–)2–7 Χ (0.8–)2–3 cm, pinnate-pinnatifid, inequilateral, basal basiscopic pinnules larger than adjacent acroscopic ones, pinnatifid; second and third pairs of pinnae with same shape as pinnae, but slightly shorter and narrower; distal pairs lanceolate, equilateral. Veins obscure. Sori consisting of several sporangia, confluent at maturity. False indusia broad, sometimes nearly to midvein, continuous or somewhat interrupted, margins entire or undulate. 2n = 60.

Rock crevices on slopes; 1000–4600 m. Guangxi, Guizhou, Hebei, Sichuan, Xizang, Yunnan [Bhutan, India, Nepal, Pakistan, Thailand].

The name A. stenochlamys refers to populations occurring from N India and C Nepal to SW China (Sichuan, Yunnan, Xizang) at elevations below ca. 3000 m. These plants have relatively large fronds and seem to differ from typical A. grisea in their spores (cristate-reticulate vs. echinate-rugulate). Likely A. grisea s.l. represents a species complex that requires more detailed study to tease apart its cryptic component taxa.

23. Aleuritopteris albomarginata (C. B. Clarke) Ching, Hong Kong Naturalist 10: 199. 1941.

白边粉背蕨  bai bian fen bei jue

Cheilanthes albomarginata C. B. Clarke, Trans. Linn. Soc., Bot. 1: 456. pl. 52. 1880; Cheilanthes farinosa Kaulfuss var. albomarginata (C. B. Clarke) Beddome.

Rhizomes short, erect; scales bicolorous, black with broad, pale margin, lanceolate. Fronds clustered. Stipe 6–12 cm Χ 1–2 mm, dark brown, lustrous, scaly, scales black or dark brown, with conspicuous lighter margins, broadly lanceolate. Lamina oblong-deltate to deltate, 9–19 Χ 4–10 cm, pinnate-bipinnatifid at base, papery when dry, glabrous adaxially, with white or pale yellow farina abaxially, also with scales along costae and midveins abaxially; pinnae 3–5 pairs, sessile, distinct along rachis, basal pair of pinnae ovate-deltate, inequilateral, bipinnatifid; pinnules 6–8 pairs, basal basiscopic pinnules larger than adjacent acroscopic ones, 2–3 cm Χ 0.5–1 mm, lanceolate, pinnatifid; second pair of pinnae with same shape as basal ones; third and upper pairs of pinnae lanceolate, pinnatifid. Sori consisting of several sporangia, confluent at maturity. False indusia broad, often nearly to midvein, interrupted, margin fimbriate. 2n = 58, 60, 116.

Rock crevices on slopes; 1300–2700 m. Guangxi, Guizhou, Sichuan, S Xizang, Yunnan [Bhutan, India, Nepal, Pakistan].

24. Aleuritopteris dubia (C. Hope) Ching, Hong Kong Naturalist 10: 200. 1941.

中间粉背蕨  zhong jian fen bei jue

Cheilanthes dubia C. Hope, J. Bombay Nat. Hist. Soc. 12: 528–529, t. 2. 1899; Aleuritopteris subrufa (Baker) Ching; Cheilanthes leveillei Christ; C. subrufa Baker; C. wusukungii Miyamoto & H. Ohba.

Rhizomes short, erect; scales bicolorous, black with narrow brown margin, lanceolate. Fronds clustered. Stipe usually noticeably shorter than lamina, 5–15 cm Χ 1–2 mm, dark brown or black, lustrous, densely scaly and often with multicellular hairs, scales concolorous or somewhat bicolorous with lighter margins, narrowly lanceolate to hairlike. Lamina oblong to deltate, 6–25 Χ 4–10 cm, pinnate-bipinnatifid, papery when dry, glabrous or rarely sparsely hairy adaxially, with white or pale yellow farina abaxially, also with hairlike to narrowly lanceolate scales along costae and midveins abaxially; pinnae 4–12 pairs, distinct along rachis, basal pair of pinnae ovate-deltate, inequilateral, bipinnatifid; basal basiscopic segments larger than adjacent acroscopic ones; second and upper pairs of pinnae lanceolate, pinnatifid. Sori consisting of several sporangia, confluent at maturity. False indusia interrupted, with a fimbriate margin.

Rock crevices on slopes, rock walls; 1300–2700 m. Guizhou, Sichuan, Yunnan [Bhutan, India, Nepal, Thailand].

Some specimens appear to be intermediate between A. dubia and A. albomarginata or A. rufa. The complex requires further study. Some of the intermediates may represent interspecific hybrids.

25. Aleuritopteris rufa (D. Don) Ching, Hong Kong Naturalist 10: 200–201. 1941.

棕毛粉背蕨  zong mao fen bei jue

Cheilanthes rufa D. Don, Prod. Fl. Nepal. 16. 1825.

Rhizomes short, erect; scales bicolorous, black with narrow brown margin, narrowly lanceolate. Fronds clustered. Stipe usually noticeably shorter than lamina, 3–9 cm Χ 1–2 mm, dark brown or black, lustrous, densely scaly and hairy, scales light orangish brown, narrowly lanceolate to hairlike. Lamina oblong, 4–12 Χ 2–7 cm, pinnate-pinnatifid, herbaceous or papery when dry, with sparse linear-lanceolate scales and multicellular hairs adaxially, with milky yellow farina and also dense hairs and scales abaxially; rachis and costae scales similar to those of stipe; pinnae 4–10 pairs, opposite, sessile, distinct along rachis, often relatively straight and spreading, basal pair of pinnae angular or oblong-lanceolate, 2–4 Χ 1–2 cm, inequilateral, pinnatifid; segments 5–8 pairs, basal basiscopic pinnules longer than adjacent acroscopic ones, 1–1.5 cm Χ 2–4 mm, oblong; second and upper pairs of pinnae oblong-lanceolate, pinnatifid. Sori consisting of several sporangia, confluent at maturity. False indusia interrupted, with a fimbriate margin. 2n = 58.

Rock crevices on slopes, rock walls; 1000–3000 m. Guangxi, Guizhou, Yunnan [India, Myanmar, Nepal, Philippines, Thailand].

26. Aleuritopteris leptolepis (Fraser-Jenkins) Fraser-Jenkins, Taxon. Revis. Indian Subcontinental Pteridophytes, 127. 2008.

薄叶粉背蕨  bo ye fen bei jue

Cheilanthes leptolepis Fraser-Jenkins, Bot. Helv. 102: 144. 1992. Cheilanthes dalhousiae Hooker, nom. rej.; C. farinosa Kaulfuss var. dalhousiae (Hooker) C. B. Clarke, nom. inval.; Aleuritopteris dalhousiae (Hooker) Ching, nom. inval.; Leptolepidium dalhousiae (Hooker) K. H. Shing & S. K. Wu, nom. inval.; L. leptolepis (Fraser-Jenkins) Kholia & Punetha.

Rhizomes short, erect; scales bicolorous, black with narrow brown margin, ovate to broadly lanceolate. Fronds clustered. Stipe shorter than lamina, 10–15 cm, dark brown, lustrous, scaly at base or to ca. midpoint, basalmost scales bicolorous, dark brown to black with lighter margin, grading to concolorous, light brown, thin and translucent, ovate to broadly lanceolate. Lamina ovate-triangular, 15–22 Χ 8–12 cm, pinnate-bipinnatifid, thinly herbaceous when dry, usually farinose during development but becoming nonfarinose or nearly so at maturity, glabrous adaxially, glabrous except for inconspicuous hairs at pinna bases abaxially; pinnae 10–12 pairs, basal pair of pinnae triangular-lanceolate, 6–10 Χ 4–6 cm, bipinnatifid; pinnules 8–10 pairs, basal basiscopic segments longer than adjacent acroscopic ones, to 3 cm, lanceolate, pinnatifid; second and upper pairs of pinnae of same shape as basal one, but slightly shorter and narrower. Veins pinnate, relatively conspicuous. Sori consisting of several sporangia, confluent at maturity. False indusia membranous, interrupted, with a laciniate margin. 2n = 60.

Rock crevices in forests; 1900–3500 m. SW Sichuan, SE Xizang, NW Yunnan [Bhutan, India, Myanmar, Nepal, Pakistan, Philippines].

This species was long known to Asian botanists as A. dalhousiae, but the basionym Cheilanthes dalhousiae was typified by a mixed collection and was thus of ambiguous application; the epithet has been officially rejected from further use.

27. Aleuritopteris duthiei (Baker) Ching, Bull. Fan. Mem. Inst. Biol., n.s.: 1: 271. 1949.

杜氏粉背蕨  du shi fen bei jue

Cheilanthes duthiei Baker, Ann. Bot. (Oxford) 5: 210. 1891; Leptolepidium duthiei (Baker) X. C. Zhang & G. M. Zhang.

Rhizomes short, erect; scales concolorous, dark brown, broadly lanceolate. Fronds clustered. Stipe 2.5–9 cm, grooved, dark brown to black, scaly, at least a few scales extending to rachis, scales concolorous, dark brown to reddish brown, broadly lanceolate at stipe base, grading abruptly to narrowly ovate above base. Lamina oblong-deltate to ovate-deltate or subpentagonal, 5–7 Χ 3–5 cm, pinnate-bipinnatifid to pinnate-tripinnatifid, thinly herbaceous to membranous when dry, glabrous adaxially, nonfarinose abaxially but with short, brown to dark brown, capitate glands along costae, costules, and veins; pinnae 4 or 5 pairs, distinct along rachis, basal pair of pinnae oblong-deltate, ca. 1.5 Χ ca. 1 cm, bipinnatifid to tripinnatifid; pinnules 3or 4 pairs, proximal basiscopic ones largest, pinnatifid to pinnate; second and upper pairs of pinnae same shape as basal one, but slightly shorter and narrower. Veins pinnate, relatively conspicuous. Sori consisting of 2–4 sporangia, confluent at maturity. False indusia interrupted, margin irregularly crenate or dentate.

Rock crevices; 3500–4400 m. SW Sichuan, Xizang [Bhutan, India, Nepal].

This relatively small-fronded species is readily distinguished from other species of Aleuritopteris by its lamina not farinose or villous; the grooved stipes and rachises, and with the distinctive, dense, short, brown to dark brown capitate glands on costae, costules and veins abaxially. It is apparently a rare species, occasionally collected at high elevations in the Himalayan region.

28. Aleuritopteris subvillosa (Hooker) Ching, Hong Kong Naturalist 10: 203. 1941.

绒毛粉背蕨  rong mao fen bei jue

Cheilanthes subvillosa Hooker, Sp. Fil. 2: 87–88, t. 98B. 1852; Cheilanthes yunnanensis Brause; C. yunnanensis var. dilatata Brause; Leptolepidium subvillosum K. H. Shing & S. K. Wu.

Rhizomes short, erect; scales concolorous or slightly bicolorous, brown, occasionally with lighter margin, sometimes slightly translucent, lanceolate. Fronds clustered. Stipe 10–15 cm, dark brown to black, lustrous, scaly at base or to ca. stipe midpoint, scales concolorous or bicolorous, light brown or brown with lighter margin, broadly ovate to ovate-lanceolate with filiform tips. Lamina oblong-lanceolate, 15–30 Χ 3–10 cm, pinnate-bipinnatifid, thinly herbaceous when dry, adaxially glabrous, abaxially hairy along costae and costules, sometimes also with white farina abaxially; pinnae 7–10 pairs, widely separated along rachis, basal pair of pinnae elliptic-triangular, 3–4 Χ 2–3 cm, sessile, bipinnatifid; pinnules 4 or 5 pairs, basal basiscopic segments longer than adjacent acroscopic ones, 1.5–2 cm Χ 7–10 mm, oblong, pinnatifid; second pair of pinnae triangular, slightly longer than basal pair; third and upper pairs gradually shortened distally. Veins pinnate, relatively conspicuous. Sori consisting of several sporangia, confluent at maturity. False indusia herbaceous, continuous or rarely discontinuous, margins undulate. 2n = 60, ca. 120.

Under shrubs or in rock crevices, usually on dry slopes; 2000–3900 m. Guizhou, Sichuan, Xizang, Yunnan [Bhutan, India, Myanmar, Nepal].

1a. Lamina without farina abaxially, except sometimes on developing fronds .........  28a. var. subvillosa

1b. Lamina with moderate to dense farina abaxially at maturity .................................  28b. var. tibetica

28a. Aleuritopteris subvillosa var. subvillosa

绒毛粉背蕨(原变种)  rong mao fen bei jue (yuan bian zhong)

Aleuritopteris subvillosa var. dilatata (Brause) H. S. Kung; L. subvillosum var. dilatatum (Brause) K. H. Shing & S. K. Wu.

Lamina hairy but lacking farina abaxially at maturity, sometimes somewhat white-farinose when young.

Under shrubs and in rock crevices; 2000–3900 m. Guizhou, Sichuan, Xizang, Yunnan [Bhutan, India, Myanmar, Nepal].

Although plants typical of this variety are easily distinguished from the relatively densely farinose var. tibetica, there is a lot of variation among populations. In most cases, plants with nonfarinose adult fronds produce at least some farina on croziers. Additionally, some plants have sparse abaxial farina persistent on mature fronds.

28b. Aleuritopteris subvillosa var. tibetica (Ching & S. K. Wu) H. S. Kung, Fl. Sichuan. 6: 258. 1988.

西藏粉背蕨  xi zang fen bei jue

Leptolepidium subvillosum K. H. Shing & S. K. Wu var. tibeticum Ching & S. K. Wu, Acta Bot. Yunnan. 1: 116. 1979.

Lamina moderately to densely white-farinose abaxially, also hairy.

* Rock crevices on dry slopes; usually above 3500 m. S and SW Sichuan, Xizang, NW Yunnan.

29. Aleuritopteris kuhnii (Milde) Ching, Hong Kong Naturalist 10: 202. 1941.

华北粉背蕨  hua bei fen bei jue

Cheilanthes kuhnii Milde, Bot. Zeitung (Berlin) 1867: 149. 1867; Aleuritopteris caesia (Christ) Ching; A. kuhnii var. brandtii (Franchet & Savatier) Tagawa; A. kuhnii f. efarinosa (Makino) Tagawa; A. tenella Saiki; Cheilanthes brandtii Franchet & Savatier; C. brandtii var. efarinosa Makino; C. caesia Christ; C. farinosa Kaulfuss var. brandtii (Franchet & Savatier ) C. Christensen; C. kuhnii f. gracilis Kitagawa; C. kuhnii var. brandtii Tagawa; C. kuhnii var. caesia (Christ) C. Christensen; C. lanceolata C. Christensen; Leptolepidium caesium (Christ) K. H. Shing & S. K. Wu; L. kuhnii (Milde) K. H. Shing & S. K. Wu; L. kuhnii K. H. Shing & S. K. Wu var. brandtii K. H. Shing & S. K. Wu; L. tenellum Ching & S. K. Wu.

Rhizomes short, erect; scales bicolorous, black with reddish brown margin, broadly lanceolate. Fronds clustered. Stipe 10–20 cm, reddish brown, lustrous, scaly at base or to ca. midpoint, scales bicolorous, brown with lighter margin, translucent, narrowly ovate to ovate-lanceolate. Lamina oblong-lanceolate, 15–30 Χ 5–13 cm, pinnate-bipinnatifid, herbaceous when dry, adaxially glabrous, abaxially with white farina or rarely nonfarinose and usually glabrous; pinnae 8–12 pairs, sessile or subsessile, distinct along rachis, basal pair of pinnae slightly shorter than or equal to second pair, ovate-triangular to oblong-lanceolate, 2.5–4(–7) cm, bipinnatifid; pinnules 4–6 pairs, basal basiscopic pinnules longer than adjacent acroscopic ones, 1–1.5 cm Χ 5–7 mm, ovate-oblong, pinnatifid; second and upper pairs gradually shortened distally. Veins pinnate, obscure or more conspicuous on adaxial surface. Sori consisting of several sporangia, confluent at maturity. False indusia herbaceous, continuous, margins undulate.

Rock crevices on dry slopes or in forests; 1000–3500 m. Gansu, Hebei, Henan, Jilin, Liaoning, Nei Mongol, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan [Japan, Korea, Russia].

This species is widely distributed and exhibits great variation in lamina form. Aleuritopteris caesia Ching was based on large-frond plants of A. kuhnii.

15. PARAHEMIONITIS Panigrahi, Indian Fern J. 9: 244. 1993.

泽泻蕨属  ze xie jue shu

Zhang Gangmin (张钢民); Tom A. Ranker

Plants terrestrial. Rhizomes erect, short, bearing small, fluffy scales and long, jointed hairs, dictyostelic. Fronds somewhat dimorphic, clustered; stipe chestnut-colored or purple-black, densely scaly and hairy as on rhizomes; stipe of fertile frond usually 1–3 Χ length of stipe of sterile frond; lamina simple, ovate, oblong-ovate, or hastate, herbaceous, brown and sparsely covered with small subulate scales abaxially, glabrous adaxially, base strongly cordate, apex obtuse or rounded. Veins anastomosing, areoles numerous, densely arranged, elongated hexagonal. Indusia absent. Sori following course of veins, confluent throughout abaxial surface when mature. Spores globose-tetrahedral, cristate. x = 30.

One species: tropical Asia.

Parahemionitis cordata was usually included in the genus Hemionitis, but it has subulate scales instead of hairs abaxially, differing from Hemionitis species in the Americas. Panigrahi thought it reasonable to put the Asian species into another genus Parahemionitis.

1. Parahemionitis cordata (Roxburgh ex Hooker & Greville) Fraser-Jenkins, New Sp. Syndr. Indian Pteridol. 187. 1997.

泽泻蕨  ze xie jue

Hemionitis cordata Roxburgh ex Hooker & Greville, Icon. Filic. 1: t. 64. 1828; Gymnogramma sagittata (Fιe) Ettingshausen; H. sagittata Fιe; H. toxotis Trevisan.

Rhizomes erect, short; scales brownish, narrowly lanceolate. Stipe of fertile frond much longer than that of sterile frond (more than 2–3 Χ in dense forests, ca. 1 Χ in dry shrublands), 6–18 cm Χ ca. 1 mm; lamina abaxially brown, adaxially brownish green, ovate, narrowly ovate, or hastate, 3–6(–10) Χ 2–4(–6) cm, forming an oblique angle with stipe, herbaceous when dry, abaxially with sparse, small subulate scales (scales rather dense along main vein), adaxially glabrous, base strongly cordate, margin with sparse, reddish brown, jointed hairs, apex obtuse or rounded. Sori brown, following course of veins, confluent throughout abaxial surface when mature. 2n = 120.

Wet soil and rock crevices of stream valleys in dense forests, or in shrublands; below 1000 m. Hainan, SW Taiwan, S Yunnan (Menghai) [Cambodia, India, Laos, Malaysia, Philippines, Sri Lanka, Thailand, Vietnam].

16. PARAGYMNOPTERIS K. H. Shing, Indian Fern J. 10: 227. 1994.

金毛裸蕨属  jin mao luo jue shu

Zhang Gangmin (张钢民); Tom A. Ranker

Plants usually growing on rocks, xeric. Rhizomes erect or decumbent, short, dictyostelic, scaly and villous; scales tan, linear or subulate, margin entire. Fronds monomorphic, clustered; stipe chestnut brown, lustrous, terete, densely villous distally; lamina oblong-lanceolate, 1- or 2-pinnate, papery or leathery, soft, densely tan (gray when old), fine sericeous throughout (especially abaxially), or with pellucid, lanceolate, entire, and imbricately arranged scales abaxially. Ultimate pinnules or segments ovate, oblong, or oblong-lanceolate, base rounded or cordate, margin entire, apex obtuse or rounded. Veins free, pinnately branched and divergent distally, or occasionally connected near frond margin. Indusia absent. Sori linear, following entire course or distal portion of veins, covered by hairs or scales, somewhat revealed when mature. Spores globose-tetrahedral, perispore cristate. x = 30.

About five species: Bhutan, China, India, Nepal, Russia, Thailand; Europe; five species (three endemic) in China.

1a.       Lamina densely covered with imbricately arranged ovate-lanceolate scales abaxially.

2a.       Lamina 1-pinnate; stipe covered with jointed long hairs or subglabrous; rachis glabrous adaxially; plants usually growing on bare limestone rocks .........................................................  1. P. delavayi

2b.       Lamina 1-pinnate-pinnatifid; stipe covered with fibrous scales; rachis sparsely scaly adaxially; plants growing in crevices of noncalcareous rocks ..................................................  2. P. marantae

1b.       Lamina densely sericeous abaxially.

3a.       Lamina 1-pinnate; pinnae rounded at base (occasionally proximal pinnae somewhat cordate at base); veins connected near frond margin ....................................................................  3. P. vestita

3b.       Lamina 1- or 2-pinnate; pinnae or pinnules strongly cordate or with 1 or 2 segments at base; veins free, not connected near frond margin.

4a.       Lamina 1- or 2-pinnate; pinnules ovate, 7–14 mm ......................................  4. P. bipinnata

4b.       Lamina 2-pinnate; pinnules ovate-deltoid or hastate, 4–5 mm .......................  5. P. sargentii

1. Paragymnopteris delavayi (Baker) K. H. Shing, Indian Fern J. 10: 229. 1994.

滇西金毛裸蕨  dian xi jin mao luo jue

Gymnogramma delavayi Baker, Ann. Bot. (Oxford) 5: 484. 1891; Gymnopteris delavayi (Baker) Underwood; G. delavayi var. intermedia Ching; G. marantae (Linnaeus) Ching var. intermedia Ching; Neurogramma delavayi (Baker) Diels; Notholaena bureaui Christ; N. delavayi (Baker) C. Christensen; N. marantae (Linnaeus) R. Brown var. delavayi (Baker) Tagawa; Paraceterach delavayi (Baker) R. M. Tryon; Paragymnopteris marantae (Linnaeus) Ching var. intermedia (Ching) K. H. Shing.

Rhizomes ascending or decumbent, short, stout; scales tan, narrowly subulate. Fronds clustered; stipe chestnut black, 8–12 cm Χ 1–2 mm, base slightly scaly and hairy, distally hairy or subglabrous; lamina adaxially pale green, oblong-lanceolate or broadly linear-lanceolate, 5–14 Χ 2–4 cm, leathery when dry, abaxially densely covered with brown, pellucid, ovate-lanceolate scales, adaxially glabrous, apex shortly acuminate or caudate, imparipinnate; rachis with sparse, narrow subulate scales. Pinnae (5–)10–15 pairs, falcate-lanceolate or lanceolate, 1.5–2.5 Χ ca. 0.5 cm, shortly stalked (distal pinnae sessile), base rounded or with a macroscopic auricle (sometimes a small oblong segment). Veins obscure, pinna midrib convex abaxially, slightly concave adaxially. Sori usually covered by scales, with 32 spores per sporangium.

* Limestone crevices in open woods; 2200–4600 m. Gansu, W Guizhou, Qinghai, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan.

Paragymnopteris delavayi looks very much like P. marantae except for its undivided pinnae and jointed long hairs (rarely with hairy scales with 2 or 3 rows of cells) on stipes. In addition, both are distributed in NW Yunnan, even in the same locality, only the former grows on calcareous substrates and the latter mostly on noncalcareous rocks.

2. Paragymnopteris marantae (Linnaeus) K. H. Shing, Indian Fern J. 10: 229. 1994.

欧洲金毛裸蕨  ou zhou jin mao luo jue

Acrostichum marantae Linnaeus, Sp. Pl. 2: 1071. 1753; A. canariense Willdenow; A. subcordatum Cavanilles; Ceterach marantae (Linnaeus) Candolle; Cheilanthes marantae (Linnaeus) Domin; Cincinalis marantae (Linnaeus) Desvaux [“maranthae”]; Gymnogramma marantae (Linnaeus) Mettenius; Gymnopteris marantae (Linnaeus) Ching; Notholaena marantae (Linnaeus) R. Brown; Paraceterach marantae (Linnaeus) R. M. Tryon.

Rhizomes decumbent or ascending, short, stout; scales tan, narrowly subulate. Fronds clustered or closely spaced; stipe 6–17 cm Χ 1.5–2.5 mm, covered with fibrous scales; lamina adaxially brownish green, broadly lanceolate, (8–)12–20 Χ 2.5–5 cm, thinly papery, abaxially densely covered with reddish brown, pellucid, ovate-lanceolate scales, adaxially glabrous, pinnate-pinnatifid, gradually tapered to pinnatifid apex; rachis and costae with sparse, small subulate-lanceolate scales. Pinnae (6–)10–16 pairs, obliquely patent, ovate-deltoid or somewhat deltoid, 2–3 Χ 1–1.5 cm, pinnatifid nearly to costae; proximal pairs of pinnae shortly stalked. Segments 2–5 pairs, oblong-lanceolate or oblong, 5–8 Χ ca. 3 mm, apex rounded. Veins obscure. Sori spreading along lateral veins, covered by scales. 2n = 58.

Rock crevices in understories of woods in dry regions; 1800–4200 m. Sichuan, Xizang, Yunnan [India, Nepal; Europe].

3. Paragymnopteris vestita (Hooker) G. M. Zhang & Ranker, comb. nov.

金毛裸蕨  jin mao luo jue

Basionym: Gymnogramma vestita Hooker, Icon. Pl. 2: t. 115. 1837; Gymnopteris vestita (Hooker) Underwood; Hemionitis vestita (Hooker) J. Smith; Neurogramma vestita (Hooker) Diels; Notholaena himalaica Fraser-Jenkins; Paraceterach vestita (Hooker) R. M. Tryon; Syngramma vestita (Hooker) T. Moore.

Rhizomes decumbent or ascending, short, stout; scales tan, narrowly subulate. Fronds clustered or closely spaced; stipe (6–)10–20 cm Χ 1–2.5 mm, densely brownish sericeous; lamina brown when dry, lanceolate, 10–15 Χ 2.5–5(–7) cm, imparipinnate, soft leathery, abaxially densely golden sericeous, adaxially sparsely pale brown sericeous; rachis and costae sericeous. Pinnae (7–)10–17 pairs, ovate or narrowly ovate, 1.5–4 Χ 1–2 cm, alternate, shortly stalked, base rounded or sometimes slightly cordate, rarely macroscopically auriculate, margin entire, apex obtuse. Veins obscure, pinnate and divergent, connecting near pinna margins. Sori covered by hairs. 2n = 60.

Among shrubs in rock crevices; 800–3900 m. W Guizhou, Hebei, Shanxi, Sichuan, Taiwan, Xizang, Yunnan [India, Nepal, Pakistan, N Thailand].

“Paragymnopteris vestita (Wallich ex C. Presl) K. H. Shing” (Indian Fern J. 10: 230. 1994) is an invalid combination because it was based on the nomen nudum “Gymnogramma vestita Wallich ex C. Presl” (Tent. Pterid. 218. 1836).

4. Paragymnopteris bipinnata (Christ) K. H. Shing, Indian Fern J. 10: 230. 1994.

川西金毛裸蕨  chuan xi jin mao luo jue

Rhizomes decumbent, short, stout, densely scaly; scales brown or dark brown, shiny, narrowly subulate. Fronds closely spaced; stipe densely pale brown sericeous when young; lamina adaxially brownish green, lanceolate or broadly lanceolate, soft leathery when dry, abaxially densely sericeous, adaxially sparsely sericeous, 1- or 2-imparipinnate; rachis and costae sericeous. Pinnae long stalked. Ultimate pinnules ovate or narrowly ovate, base cordate, margin entire or with 1 or 2 small segments, apex obtuse. Veins obscure. Sori covered by hairs.

* Rock cliffs, crevices by ditches, on rocks beneath shrubs or woods; 800–3600 m. Gansu, Hebei, Henan, W Hubei, Nei Mongol, Shaanxi, Sichuan, Xizang, Yunnan.

1a. Lamina 2-pinnate; inframedial pinnae lanceolate or deltoid-lanceolate ................  4a. var. bipinnata

1b. Lamina 1-pinnate; pinnae ovate or narrowly ovate, margin entire, or at most with 1 or 2 free segments in proximal pinnae ..............................................................................................  4b. var. auriculata

4a. Paragymnopteris bipinnata var. bipinnata

川西金毛裸蕨(原变种)  chuan xi jin mao luo jue (yuan bian zhong)

Gymnopteris bipinnata Christ, Notul. Syst. (Paris) 1(2): 55. 1909; Hemionitis bipinnata (Christ) Mickel; Paraceterach bipinnata (Christ) R. M. Tryon.

Stipe 10–22 cm Χ 1–3 mm; lamina 15–25 Χ 3–7 cm, 2-pinnate (apical portion 1-pinnate). Pinnae 10–17 pairs, lanceolate or deltoid-lanceolate, imparipinnate. Lateral pinnules 1–6 pairs, ovate or narrowly ovate, shortly stalked or sessile, base somewhat cordate, margins entire or rarely with 1 or 2 small lobes, apex obtuse; terminal pinnule as lateral ones but larger and with longer stalk.

* Rock cliffs, crevices by ditches; 1100–3200 m. Gansu, Shaanxi, Sichuan, Xizang, Yunnan.

4b. Paragymnopteris bipinnata var. auriculata (Franchet) K. H. Shing, Indian Fern J. 10: 230. 1994.

耳羽金毛裸蕨  er yu jin mao luo jue

Gymnogramma vestita Hooker var. auriculata Franchet, Nouv. Arch. Mus. Hist. Nat., sιr. 2, 10: 123. 1887; Gymnopteris bipinnata var. auriculata (Franchet) Ching; G. borealisinensis Kitagawa.

Lamina 1-pinnate. Pinnae ovate or narrowly ovate, stalked (stalks to 1 cm), base strongly cordate, margin entire or often enlarged forming auricles at both sides near base, sometimes proximal pinnae with 1 or 2 free segments, apex obtuse.

* On rocks below shrubs or woods; 800–3600 m. Gansu, Hebei, Henan, W Hubei, Nei Mongol, Sichuan, Xizang, Yunnan.

Paragymnopteris bipinnata var. auriculata is similar to P. vestita. It differs in its long pinna stalks (to 1 cm), strongly cordate pinna base, and free veins.

5. Paragymnopteris sargentii (Christ) K. H. Shing, Indian Fern J. 10: 230. 1994.

三角金毛裸蕨  san jiao jin mao luo jue

Gymnopteris sargentii Christ, Bot. Gaz. 51: 355. 1911; Notholaena sargentii (Christ) Fraser-Jenkins; Paraceterach sargentii (Christ) R. M. Tryon.

Plants small, less than 40 cm tall. Rhizomes decumbent or ascending, short, stout; scales shiny, brown, narrowly lanceolate. Fronds clustered; stipe light chestnut-colored, 5–12 cm Χ 1.5–2 mm, densely light brown sericeous proximally, becoming sparsely so distally; lamina adaxially brownish green, deltoid-lanceolate or oblong-lanceolate, 10–25 Χ 4–10 cm, 2-pinnate (apical portion 1-pinnate), leathery when dry, abaxially densely sericeous, adaxially sparsely sericeous; rachis and costae with same hairs as on lamina surface. Pinnae 8–14 pairs, inframedial pinnae alternate, lanceolate, 3.5–7 Χ 1–1.5 cm, stalked (stalks 3–7 mm), imparipinnate; pinnules 3–8 pairs, anatropous, ovate-deltoid or hastate, 4–5 Χ 3–4 mm, shortly stalked, base cordate, apex obtuse; terminal pinnule as lateral ones but with longer stalk and often inequilateral. Veins obscure. Sori covered by hairs.

* Rocky slopes, among shrubs in dry valleys; 1900–3300 m. W Sichuan, Xizang, NW Yunnan.

5. Subfam. VITTARIOIDEAE

书带蕨亚科  shu dai jue ya ke

[check all authors cited; order OK?]
Zhang Xianchun (张宪春), Lin Youxing (林尤兴); Michael G. Gilbert

Epiphytic or epilithic plants. Rhizome creeping, bearing roots with very numerous water-absorbing root hairs, and clathrate scales with iridescent lamina. Fronds simple and entire, glabrous, epidermis with large elongate siliceous cells. Sori exindusiate, usually immersed in grooves, elongate along fertile veins, with abundant paraphyses. Spores transparent, trilete or monolete, mostly ellipsoidal, sometimes fusiform, or tetrahedral-globose.

Eleven genera and ca. ?300 species: pantropical; four genera and 58 species (17 endemic) in China.

Molecular data has shown that Vittariaceae and Adiantaceae form a well-defined monophyletic group, Vittarioideae, within the Pteridaceae.

rbcL gene analysis by Crane (1997) indicated that the traditional Antrophyum and Vittaria are both polyphyletic or paraphyletic. Ten monophyletic genera are recognized in Vittariaceae, i.e., Anathacorus, Anetium, Antrophyum, Haplopteris, Hecistopteris, Monogramma, Polytaenium, Radiovittaria, Scoliosorus, and Vittaria. Only Antrophyum, Haplopteris, and Monogramma are distributed in China. All the Chinese species formerly classified in Vittaria are now treated in Haplopteris.

1a.       Fronds compound with slender glossy black or reddish brown stipe and rachis and well-defined, often articulate, petiolulate pinnules (rarely reduced to single orbicular to reniform pinnule); sporangia in discrete marginal sori, protected by false indusium formed from reflexed lamina margin .....  1. Adiantum

1b.       Fronds simple, without glossy stipe or distinct pinnules; sporangia distributed along veins, often in grooves, without any indusium.

2a.       Fronds obovate to oblanceolate, widest above middle, rarely linear, costa none or only at lower part; sori multiseriate along veins, in grooves along veins on abaxial surface of lamina, rarely one row on each sides; spores monolete
....................................................................................................................  2. Antrophyum

2b.       Fronds filiform to linear or ribbonlike, rarely lanceolate, costa distinct up to apex or upper part; sori biseriate or uniseriate, in marginal or submarginal grooves, rarely superficial, or along costa; spores trilete or monolete.

3a.       Sori biseriate in marginal or submarginal grooves, rarely superficial; spores monolete  3. Haplopteris

3b.       Sori uniseriate, 1–3 along costa or on lateral veins, immersed; spores trilete
                                                                                                                 4. Monogramma

17. ADIANTUM Linnaeus, Sp. Pl. 2: 1094. 1753.

铁线蕨属  tie xian jue shu

Lin Youxing (林尤兴); Michael G. Gilbert

Ferns, terrestrial or epilithic (growing on rocks), medium-sized or small; outlines highly variable. Rhizomes short and erect, or long and creeping, with siphonostele, covered with scales; scales brown or black, lanceolate, thick in texture, usually entire. Fronds monomorphic, whorled and clustered, scattered in 2 rows or compact, not articulate; stipe black or reddish brown, glossy, fine and rounded, hard, with 1 or 2 vascular bundles and combined into one upward; lamina mostly 1–3 or more pinnate or 1–3 dichotomous with pedate branches, rarely reduced to solitary entire and orbicular-flabellate pinnule, herbaceous or papery, less often leathery or membranous; rachises, costae, and stalks same color as stipe; pinnules sometimes with articulate petiolules, ovate, flabellate, orbicular-flabellate, or dimidiate, margins serrate, lobed, or entire, usually deciduous when dried; veins free, reaching margins, simple or dichotomously forked, often radiate [rarely anastomosing (Hewardia)], usually visible on both surfaces of lamina. Sori borne on veins of recurved, membranous margins (false indusium); false indusium orbicular, reniform, lunate, elongated-square, or oblong, free or continuous, upper margins forming deep sinus, retuse or truncate. Sporangia globose, long stalked, annuli erect; mostly consisting of 18–28 incrassate cells. Spores yellowish, tetrahedral, trilete, transparent, smooth and experine. x = 15(30), 29.

More than 200 species: from cold temperate zone to tropics, most in South America; 34 species (17 endemic) in China.

1a.       Fronds each with a single suborbicular or orbicular-reniform pinnule ................  1. A. reniforme

1b.       Fronds each with several to very many pinnules.

2a.       Fronds 1–3-palmate or dichotomously branched.

3a.       Fronds 2- or 3-dichotomously or nearly dichotomously branched, primary branches with 1-pinnate pinnae on both sides.

4a.       Pinnules including indusia hairy ............................................................  18. A. hispidulum

4b.       Pinnules glabrous ............................................................................  19. A. flabellulatum

3b.       Fronds pedately branched, with 2–6(or 8) 1-pinnate pinnae per branch.

5a.       Sori 1 or 2 per pinnule; lamina ca. 15 cm; pinnules ca. 10 Χ 3–4 mm
.........................................................................................................  17. A. subpedatum

5b.       Sori 4–6 per pinnule; pinnules ca. 20 Χ 6 mm.

6a.       Pinnules with upper margin divided for 1/3–1/2 of length, apex with blunt serrations, abaxially green; indusia retuse at upper margin ...............................................................  15. A. pedatum

6b.       Pinnules with upper margin more shallowly lobed, apex with acute triangular serrations, abaxially glaucous; indusia deeply sinuate at upper margin ..............................  16. A. myriosorum

2b.       Fronds 1–4-pinnate.

7a.       Fronds 1-pinnate, lanceolate or linear-lanceolate in outline.

8a.       Pinnules entire, petiolule apex ± articulate and pinnule easily lost when dry.

9a.       Plants small, creeping, 1.5–3 cm tall, lamina with 3–5 small orbicular pinnules; indusia orbicular, upper margin truncate, one per pinna .......................................................  2. A. mariesii

9b.       Plants erect, above 3 cm tall, lamina usually with 5–7 or more flabellate pinnules; indusia reniform or elongated, upper margins retuse or truncate, one to many per pinna.

10a.     Plants soft and weak; lamina narrowly flabellate, thin in texture, stipe as slender as a hair, 1/3–1/2 as long as pinnae (N Guangdong) ...................................................  3. A. lianxianense

10b.     Plants strong; lamina broadly flabellate, stipe robust, less than 1/5 as long as lamina.

11a.     Pinnules obtriangular, indusia depressed at upper margins, scale margins denticulate  4. A. gravesii

11b.     Pinnules orbicular-flabellate or obtriangular, indusia truncate at upper margins, scale margins entire.

12a.     Pinnules orbicular-flabellate, usually with 3 or 4 sori ..................  5. A. juxtapositum

12b.     Pinnules obtriangular, with 1(or 2) sori ................................................  6. A. chienii

8b.       Pinnules ± divided, petiolule not articulate and pinnule persistent when dry.

13a.     Apex of frond not prolonged into whiplike structure ..........................  19. A. diaphanum

13b.     Apex of frond usually elongated and whiplike, often forming plantlet at end.

14a.     Stipes, rachises, and both sides of lamina multicellular brown hirsute.

15a.     Pinnule upper margins subentire or crisped or lobed for ca. 1/3 of breadth of pinnule or with 3 or 4 short, broad lobes.

16a.     Stipe 4–12 cm, densely hairy; lamina 1.2–2.4 cm wide; sori 2 or 3 per pinnule (Yunnan, rare)                                                                                                            9. A. sinicum

16b.     Stipe 10–20 cm, very sparsely hairy or glabrous; lamina 3–6 cm wide; sori 5–16 per pinnule (Taiwan, rare) ......................................................................  10. A. meishanianum

15b.     Pinnule upper margins with many long lobes (widely distributed S of Chang Jiang, common).

17a.     Basal pair of pinnules smaller than immediately distal pinnules; abaxial surface of rachis and lamina sparsely hairy, hairs facing all directions; rhizome scale margins entire  7. A. caudatum

17b.     Basal pair of pinnules larger than more distal pinnules; abaxial surface of rachis and lamina densely hairy, hairs appressed and facing laminal front; rhizomes scale margins serrate             8. A. malesianum

14b.     Stipes, rachises, and both surfaces of lamina glabrous or occasionally with 1 or 2 stiff hairs.

18a.     Stipes, rachises, and stalks with membranous, brown wings .........  11. A. soboliferum

18b.     Stipes, rachises, and stalks terete.

19a.     Pinnules semidimidiate, subsessile .............................................  14. A. edgeworthii

19b.     Pinnules semilunar, orbicular-flabellate, or orbicular, with longer stalks.

20a.     Pinnules lunate, base asymmetrical, petiolule ca. 10 mm ...........  12. A. philippense

20b.     Pinnules orbicular-flabellate or orbicular, base symmetrical, stalks petiolule 2–3 mm             13. A. capillus-junonis

7b.       Fronds 2–4-pinnate.

21a.     Pinnules with sparse deep brown and appressed long needlelike setae
...........................................................................................................  21. A. diaphanum

21b.     Pinnules glabrous.

22a.     Rachises, and petiolules brown hispidulous.

23a.     Rhizome short and erect, fronds clustered; pinnules 2 pairs per pinna; sori 4–6 per pinnule (rarely 1 or 2 on ultimate pinnules) ................................................................  20. A. induratum

23b.     Rhizome long creeping, fronds spaced; pinnules 4 or 5 pairs per pinna, stiffly papery; sori 1 (rarely 2) per pinnule .......................................................................................  24. A. davidii

22b.     Rachises, petiolules and pinnules glabrous.

24a.     Pinnules with upper margin entire, bluntly toothed, or undulate-crenate.

25a.     Rhizome long creeping; upper margin of pinnules distinctly bluntly triangular serrate  29. A. monochlamys

25b.     Rhizome shortly creeping or ascending; upper margin of pinnules entire or undulate-crenate.

26a.     Upper margin of pinnules undulate-crenate; sori mostly 1 per pinnule
                                                                                             30. A. erythrochlamys

26b.     Upper margin of pinnules entire; sori 2–4 per pinnule.

27a.     Pinnules 8–14 mm wide; ...........................................................  33. A. refractum

27b.     Pinnules 4–7 mm wide.

28a.     Pinnules triangular, leathery ..................................................  31. A. roborowskii

28b.     Pinnules fan-shaped, herbaceous ........................................  32. A. formosanum

24b.     Pinnules with upper margin densely finely sharply toothed to erose-dentate.

29a.     Plants to 10 cm tall; fronds 2-pinnate; indusia rectangular, truncate at upper margins; pinnules nearly as long as wide ...................................................................  22. A. fengianum

29b.     Plants more than 15 cm tall; fronds usually 3-pinnate; indusia orbicular, orbicular-reniform, or oblong.

30a.     Stipe base densely brownish, multicellular setose (setae deciduous but leaving marks and rough feel) .........................................................................................  23. A. bonatianum

30b.     Stipe base glabrous.

31a.     Pinnule 12–20 Χ 10–15 mm, upper margin 2–4-lobed; sori 3–10 per pinnule  34. A. capillus-veneris

31b.     Pinnule 3–11(–13) Χ 3–13 mm, upper margin truncate, curved, or emarginate; sori 1–3(–5) per pinnule.

32a.     Indusia orbicular or reniform, upper margins emarginate or curved.

33a.     Indusia orbicular, upper margins sinuate, pinnule abaxially green, veins distinctly raised     25. A. venustum

33b.     Indusia reniform, upper margins curved, pinnule abaxially glaucous, veins indistinct           26. A. tibeticum

32b.     Indusia elongated or orbicular-reniform, upper margins truncate or slightly curved.

34a.     Upper margins of pinnules with long and sharp serrations
..........................................................................................  27. A. fimbriatum

34b.     Upper margins of pinnules with short and obtuse serrations
.....................................................................................  28. A. breviserratum

1. Adiantum reniforme Linnaeus var. sinense Y. X. Lin, Acta Phytotax. Sin. 18: 102. 1980.

荷叶铁线蕨  he ye tie xian jue

Plants 5–20 cm tall. Rhizomes erect, short, scales brown, lanceolate, also multicellular villous. Fronds clustered; stipe dark castaneous, 3–14 cm, 0.5–1.5 mm in diam., base densely scaly, distally villous, hairs easily rubbed off when dried; lamina a single terminal pinnule, orbicular or orbicular-reniform, 2–6 cm in diam., papery or hard papery, greenish when dried, dark brown upon natural drying, adaxially laxly brown multicellular villous, base symmetrical, cordate, sinus shallow to deep with overlapping lobes forming 1–3 concentric rings around apex of stipe, margins bluntly crenate but crenation indistinct on fertile lamina; veins radiate from bases to all sides, multidichotomous, visible on both surfaces. Sori many per pinnule; false indusia deep brown, orbicular or subrectangular, membranous, flat and straight at upper margins and spreading along margins, approximate or wider spaced, persistent. 2n = 120*.

* Gregarious on rocks or in rock crevices; ca. 300 m. Sichuan (Shizhu).

This plant is endangered in its native habitat by road building and collection for medicinal use (L. K. Fu & J. M Jin, China Red Data Book 1: 2–3. 1992). It is now cultivated (e.g., in the Wuhan Botanical Garden) as an ornamental.

Adiantum reniforme var. sinense has been used in Chinese medicine for more than 100 years, known as “荷叶金钱草 he ye jing quan cao” in Sichuan. The whole plant is used in traditional Chinese medicine to clear away heat and toxic materials, promote diuresis, and to treat hepatitis of a type infecting urinary organs, urinary stones, and middle ear infections.

The typical variety occurs in the Atlantic Ocean islands while further varieties are recorded from scattered localities throughout Africa, Madagascar, Mauritius, and Rιunion.

2. Adiantum mariesii Baker, Gard. Chron., n.s., 16: 494. 1880.

小铁线蕨  xiao tie xian jue

Adiantum acrocarpon Christ; A. nanum Ching.

Plants epilithic, small, 2–3 cm tall. Rhizomes erect, short, scales black-brown lanceolate, margins entire. Fronds clustered; stipe dark castaneous, ca. 1 cm; as slender as silk; lamina 1-pinnate, ovate in outline, ca. 2 Χ 1 cm; costae and stalks same color as stipe, pinnules 1–3 pairs, subopposite, easily deciduous when dry; petiolule 1–2 mm, articulate; blade obliquely spreading, orbicular or subovate-orbicular, 2.5–4 Χ 2.5–4 mm, papery, abaxially slightly pale blue-grayish, glabrous, adaxially brown-green and glossy, base rounded or rounded-cuneate, lateral margins entire or slightly undulate, upper margin rounded and with indistinct sinuses in middle; veins undivided, 4 forked veinlets running from base to margin, visible on both surfaces. Sori 1 per pinnule, false indusia brown, orbicular, upper margins flat and straight, slightly depressed, persistent.

* Usually gregarious on wet limestone cliffs; ca. 200 m. Guangxi, Guizhou, Hubei, Sichuan.

The type specimen was collected from Hubei (Badong).

3. Adiantum lianxianense Ching & Y. X. Lin, Acta Phytotax. Sin. 18: 102. 1980.

粤铁线蕨  yue tie xian jue

Plants 5–7 cm tall. Rhizomes erect, short, scales black lanceolate. Fronds clustered; stipe castaneous-black, slightly glossy, 2–3 cm, as slender as a hair, glabrous; lamina oblong in outline, 3–5 Χ 1–1.5 cm, 1-pinnate, costae and stalks same color as stipe, pinnules 3 or 4 pairs, 6–9 mm apart, opposite, obliquely spreading, petiolule 1.5–4 mm, 1/3–1/2 as long as pinnule, articulate at end, persistent after pinnule fall; pinnules narrowly ovate or narrowly flabellate, 4–6 Χ 2.5–4 mm, papery, brownish green, both surfaces glabrous, base cuneate, both sides entire and undulate, upper margins subtruncate, with one sinus at middle; terminal pinnae same shape as and slightly larger than lateral pinnae; veins undivided, 4 forked veinlets arising from bases, visible on both surfaces. Sori 1 per pinnule; false indusia brown, reniform or elliptic, leathery, upper margins depressed, persistent.

* Gregarious on wet calcareous rocks and calcitic soils. Guangdong (Lianxian).

Ching initially misidentified this species as Adiantum gravesii Hance. Adiantum lianxianense is believed to be extinct due to habitat loss (China Plant Specialist Group 2004. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.4. <www.iucnredlist.org>. Downloaded on 27 Apr 2011).

4. Adiantum gravesii Hance, J. Bot. 13: 197. 1875.

白垩铁线蕨  bai e tie xian jue

Adiantum greenii Ching; A. leveillei Christ.

Plants terrestrial, 4–14 cm tall. Rhizomes erect, short, scales black subulate-lanceolate, margins denticulate. Fronds clustered; stipe castaneous-black, glossy, 1–6 cm, slender, glabrous; lamina 1-pinnate, oblong or ovate-lanceolate in outline, 3–6 Χ 2–2.5 cm; costae and stalks same color as stipes; pinnules 2–4 pairs, 1–2 cm apart; alternate, obliquely upward, petiolule up to 3 mm (ca. 1/5 as long as pinna or shorter), articulate, persistent after pinnule fall; broadly obovate or broadly ovate-triangular, ca. 1 Χ 1 cm, papery, abaxially glaucous, adaxially grayish green, both surfaces glabrous, base rounded-cuneate or rounded, both sides slightly undulate, apex rounded, with one (rarely 2) shallow sinus(es), entire; veins dichotomously branching, and reaching cartilaginous margins, visible on both surfaces. Sori 1 (rarely 2) per pinna; false indusia brown, reniform or lunate (rarely orbicular), leathery, upper margin depressed, persistent.

Gregarious on wet cliffs, in rock crevices, on chalky soils in caves; 600–1500 m. Guangdong, Guangxi, Guizhou, Hubei, Hunan, Sichuan, Yunnan, Zhejiang [N Vietnam].

The type specimen was collected from Guizhou.

Adiantum gravesii is very similar to Adiantum lianxianense Ching & Y. X. Lin, but differs in the plants being taller and stronger, having larger, broadly ovate or broadly obovate-triangular pinnules that are abaxially glaucous, and with shorter petiolules.

5. Adiantum juxtapositum Ching, Acta Phytotax. Sin. 6: 312. 1957.

仙霞铁线蕨  xian xia tie xian jue

Plants 8–20 cm tall. Rhizomes erect, short, scales black lanceolate. Fronds clustered; stipe castaneous, glossy, 2–9 cm, ca. 1 mm in diam., glabrous; lamina 1-pinnate, lanceolate in outline, 4–11 Χ 1.5–3 cm; costae and stalks same color as stipe, smooth; pinnules 5–9 pairs, opposite, nearly alternate upward, horizontally spreading; petiolule 2–3 mm, articulate, persistent after pinnule fall; pinnules orbicular or broadly orbicular-flabellate (rarely obtriangular), ca. 1 Χ 1.2–1.5 cm, subleathery, adaxially greenish, abaxially glaucous and glabrous, base rounded-cuneate (rarely cuneate), upper margin rounded, slightly undulate; distal pinnules slightly smaller, obtriangular or flabellate; veins multidichotomously branching and reaching cartilaginous margins, visible on both surfaces. Sori (1–)3 or 4 (or 6) per pinnule; false indusia black, orbicular or orbicular-reniform (rarely elongated), leathery, upper margins flat and straight or slightly depressed, entire or slightly undulate, persistent. Perispore indistinctly granulate.

* Limestone crevices. N Fujian (Xian Xia Shan).

6. Adiantum chienii Ching, Sinensia 1: 50. 1930.

北江铁线蕨  bei jiang tie xian jue

Plants usually 20–30 cm tall. Rhizomes erect, short, scales castaneous-brown, linear. Fronds clustered, horizontally spreading; stipe castaneous-black, glossy, 5–10 cm, ca. 1 mm in diam., glabrous; lamina 1-pinnate, linear-lanceolate in outline, 13–18 Χ 3–4 cm, ca. 2 cm spaced; costae and petiolules same color as stipe, pinnules opposite or subopposite, 6–8 pairs, petiolule 2–3 mm, articulate at apex, persistent after pinnule fall; blade obovate-triangular, ca. 1.5 Χ 1.5 cm, papery, brownish, abaxially glaucous and glabrous, adaxially glossy, base rounded or rounded-cuneate, margin entire, upper margin truncate; distal pinnules similar but smaller; veins multidichotomously branching, reaching cartilaginous margins, visible on both surfaces. Sori 1(or 2) per pinnule; false indusia brown, rectangular, up to 1 cm, leathery, upper margins flat and straight, entire, persistent.

* Guangdong (Beijiang).

The outline of Adiantum chienii is similar to A. gravesii Hance; the plant differs by being much taller and larger, and the indusia narrowly linear, horizontally attached at upper margin.

7. Adiantum caudatum Linnaeus, Mant. Pl. 308. 1771.

鞭叶铁线蕨  bian ye tie xian jue

Adiantum caudatum var. angustilobatum Bonaparte; A. lyratum Blanco.

Plants terrestrial or epilithic, 10–40 cm tall. Rhizomes erect, short, scales deeply castaneous, lanceolate, margins entire, Fronds clustered; stipe castaneous, 1–10 cm, densely dark brown or brownish multicellular hirsute; lamina 1-pinnate, lanceolate in outline, 15–30 Χ 2–4 cm, base slightly narrow; rachis same color as stipe and similarly sparsely hirsute, glabrescent when old, apex usually prolonged into a whiplike stolon rooting at tip to form new plantlet, pinnules 20–44 pairs, 5–8 mm apart; alternate, or lower ones subopposite, horizontally spreading or slightly obliquely spreading, lower pinnules gradually reduced, middle ones ± dimidiate, suboblong, 0.7–2 Χ 0.6–1 cm, papery, brownish green, both surfaces sparsely multicellular hirsute and densely pubescent, base asymmetrical, lower margins nearly straight and entire, upper and outer margins deeply divided into many narrow lobes, upper side truncate; lobes linear, margins entire, upper part again lobed into fine linear segments, apex truncate, fine segments truncate or few dentate at apex; veins multidichotomously branching, visible on both surfaces. Sori 5–12 per pinna; false indusia dark brown, orbicular or oblong, hairy, upper margins flat and straight, entire, persistent, Perispore granulate, 2n = 90.

On rocks or in rock crevices of forest understories or mountain valleys; 100–1200 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [Bhutan, Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Nepal, Philippines, Thailand, Vietnam; tropical and subtropical regions of Africa and Asia].

8. Adiantum malesianum J. Ghatak, Bull. Bot. Surv. India 5: 73. 1963.

假鞭叶铁线蕨  jia bian ye tie xian jue

Adiantum caudatum Linnaeus var. latilobatum Bonaparte.

Plants terrestrial or epilithic, 10–40 cm tall. Rhizomes erect, short, densely covered with scales; scales brown, lanceolate, scale margins dentate. Fronds clustered; stipe brown when young, castaneous-black and partly fallen when old, 3–20 cm, base covered with same brown scales as rhizome and multicellular articulate hairs on whole stipe; lamina linear-lanceolate, 12–20 Χ 1.5–3 cm, attenuate upward, base not narrow, 1-pinnate; pinnae 18–30 pairs, sessile to shortly stalked, rachises same color as stipes and densely covered with same stiff hairs; apex of rachis usually prolonged into a whiplike stolon and rooting to form new plantlets. petiolules ca. 1 mm, horizontally spreading, alternate or subopposite, ca. 1 cm spaced; proximal pair of pinnules suborbicular flabellate, ± reflexed downward; middle lateral pinnules dimidiate, 0.7–2 Χ 0.6–1 cm, papery, brown-green, abaxially densely covered with brown multicellular stiff hairs, with appressed short hairs facing outer margins, sparsely covered with short hairs adaxially, upper and outer margins parted; segments 5 or 6 pairs per pinna, rectangular, lower and inner margins flat and straight, apex depressed; terminal pinnae subtriangular, upper margins rounded and parted; veins multidichotomously forked, distinctly raised adaxially and less visible abaxially; Sori 5–12 per pinna; indusia brown, orbicular-reniform, papery, upper margins flat and straight, densely covered with hairs adaxially, entire, persistent.

On rocks or in rock crevices under thickets on slopes; below 100–1400 m. Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan [India, Indonesia, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, Vietnam; Pacific islands (Polynesia)].

Material of this species was originally misidentified as Adiantum caudatum Linnaeus by Beddome (Ferns S. India t. 2. 1863).

9. Adiantum sinicum Ching, Bull. Fan Mem. Inst. Biol., n.s., 1: 268. 1949.

苍山铁线蕨  cang shan tie xian jue

Plants 15–30 cm tall. Rhizomes erect, short, scales lanceolate. Fronds clustered; stipe castaneous-black, 4–12 cm, densely rufous-brown multicellular villous; lamina 1-pinnate, lanceolate in outline, 12–15 Χ 1.2–2.4 cm; costae same color as stipe, also with stiff hairs, apex acuminate, occasionally prolonged into a whiplike stolon and rooting to form new plantlets. Pinnules 20–25 pairs, alternate, mostly horizontally spreading, basal pair obliquely reflexed, semidimidiate, 1–1.4 Χ 0.4–0.6 cm, progressively smaller distally, papery, greenish, abaxially covered with stiff brown multicellular hairs, adaxially with lax short hairs, base asymmetrical, lower margin straight and entire, inner margins nearly straight, upper margin truncate, entire or undulate-crenate; terminal pinnules small, obtriangular, base cuneate, outer margins rounded and 2–5-lobed; veins multidichotomously forked, visible on both surfaces. Sori 2 or 3 per pinnule; false indusia dark brown, rectangular, papery, hairy on upper sides, upper margins flat and straight, entire, persistent.

* On soil by sides of limestone rocks; ca. 1600 m. Yunnan.

Adiantum sinicum is listed as endangered in the IUCN Red Book (<A2c http://www.iucnredlist.org/apps/redlist/details/46604/0>, accessed 12/07/11.

10. Adiantum meishanianum F. S. Hsu ex Yea C. Liu & W. L. Chiou, Novon 19: 59(–61; fig.1). 2009.

梅山铁线蕨  mei shan tie xian jue

Rhizome erect to suberect, short, scales brown with black center, linear, to 4 mm, margins entire. Stipe castaneous to blackish purple, lustrous, 10–20 cm, scales like those on rhizome but uniformly brown; stipe and rachis glabrous or with sparse multicellular hairs; lamina 1-pinnate, linear to lanceolate in outline, 10–50 Χ 3–6 cm; apex usually prolonged into a whiplike stolon and rooting to form new plantlets, occasionally secondarily prolonged. Pinnules 5–20 pairs, basal pinnules largest, reflexed; petiolule distinct, usually 3–5 mm, at 60–90 angle to rachis, mostly dimidiate, usually 20–30 Χ 10–15 mm, softly herbaceous, both surfaces glabrous, lower margin straight, rounded at apex, upper margin subentire, crisped or lobed for ca. 1/3 of breadth of pinnule, sinuses narrow, lobes rounded to quadrangular, rounded to truncate at subentire or toothed apex. distal pinnules gradually smaller toward apex. Sori 5–16 per pinnule, false indusia 2–3 mm wide, glabrous.

* Margin of secondary forests; ca. 1000 m. Taiwan.

Adiantum meishanianum is known only from a single very small population with less than 100 individuals and was rated in the protologue as critically endangered.

11. Adiantum soboliferum Wallich ex Hooker, Sp. Fil. 2: 13. 1851.

翅柄铁线蕨  chi bing tie xian jue

Adiantum alatum Copeland; A. balansae Baker; A. caudatum Linnaeus var. soboliferum (Wallich ex Hooker) Beddome; A. dolabriforme Hooker; A. mettenii Kuhn; A. lunulatum N. L. Burman var. mettenii Beddome.

Plants ?terrestrial, 25–30 cm tall. Rhizomes erect, short, scales dark brown, linear-lanceolate, margins entire. Fronds clustered; stipe castaneous-black, glossy, 9–16 cm, each side with a narrow brown membranous wing, base with same scales as rhizome, distally glabrous; lamina 1-pinnate, lanceolate in outline, 9–20 Χ 3.5–5.5 cm; rachis and stalks similar to stipe, apex of usually sterile rachis prolonged into whiplike stolon and rooting to form new plantlets; pinnules 5–10 pairs, alternate; petiolule 2–4 mm, articulate, persistent after pinnule fall; basal 1 or 2 pairs of pinnules slightly reflexed pinnules below middle uniform in size, horizontally spreading, dimidiate, suboblong, 2–3.3 Χ 0.9–1.4 cm, papery, brownish green, both surfaces glabrous, base asymmetrically broadly cuneate, lower and inner margins nearly flat and straight, upper and outer margins obtuse and 4–6-lobed; distal pinnules similar but slightly smaller; terminal pinnules flabellate, outer margin with 3–6 sinuses; veins multidichotomously forked, reaching cartilaginous margins, visible on both surfaces. Sori 3–7 per pinnule; false indusia dark brown, elliptic or reniform, papery, upper margins flat and straight, entire, persistent. Perispore granulate. 2n = 120.

Wet forest soils. Guangdong, Guangxi, Hainan, Taiwan, N Yunnan [India, Indonesia, Nepal, Philippines, Vietnam; tropics and subtropics of N Africa and Asia].

12. Adiantum philippense Linnaeus, Sp. Pl. 2: 1094. 1753.

半月形铁线蕨  ban yue xing tie xian jue

Adiantum arcuatum (Poiret) Swartz; A. lunatum Cavanilles; A. lunulatum N. L. Burman; A. lunulatum var. limbatum Christ; A. lunulatum var. subjunonicum Christ; Polypodium arcuatum Poiret; Pteris lunulata (N. L. Burman) Retzius.

Plants terrestrial or epilithic, 10–50 cm tall. Rhizomes erect, short, scales dark brown, lanceolate, margins denticulate. Fronds clustered; stipe castaneous, glossy, 6–25 cm, terete, base with same scales as rhizome, distally glabrous; lamina 1-pinnate, lanceolate in outline, 12–25 Χ 2–6.5 cm; rachis, costae and petiolules same color as stipe, apex usually prolonged into a whiplike stolon and rooting to form new plantlets. Pinnules 6–12 pairs, alternate, obliquely spreading, petiolule 10–20 mm, articulate, persistent after pinnule fall; pinnules below middle subequal in size, dimidiate-lunate or semiorbicular-reniform, 1–4 Χ 1–2.3 cm, herbaceous, green or brown-green, both surfaces glabrous, upper margin rounded, apex obtuse or bent downward, sterile pinnules stalked, both sides asymmetrical; margins undulate-lobed, segments obtuse and serrulate at apex, lower margins entire, truncate or slightly bent downward, rarely broadly cuneate, fertile pinnules subentire or with 2–4 shallow sinuses, or slightly undulate; distal pinnules slightly smaller; terminal pinnules flabellate, slightly larger than basal pinnules; veins multidichotomously forked and reaching margins, visible on both surfaces. Sori 2–6 per pinna; false indusia dark brown or brown-green, linear-oblong, membranous, upper margins flat and straight or slightly depressed, entire, persistent. Perispore finely granulate.

Gregarious on shaded and wet places or on acid soil of forest understories, sometimes rupicolous; 100–2000 m. Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, Yunnan [Bhutan, India, Indonesia, Kashmir, Malaysia, Myanmar, Nepal, Philippines, Thailand, Vietnam; tropics and subtropics of Africa, Oceania].

There has been controversy as to the correct name of this taxon. Pichi Sermolli (Webbia 12 : 665. 1957 lectotypified Adiantum philippense with the illustration “Adiantum Philippense, folio rotundo laciniato” in Petiver, Gazophyl. Nat., 8, t. 4, f. 4, 1702–1709. Some (e.g., Morton in Contr. U. S. Natl. Herb. 38: 370. 1974, Verdcourt, Fl. Trop. East Africa: Adiantaceae: 58. 2002) have argued that this illustration is unidentifiable and might not even be a fern. Fraser-Jenkins (Taxon. Revis. Indian Subcontinental Pteridophytes, 144. 2008) claims that Petiver’s drawing was copied from a drawing sent to him by Kamel and that the original drawing and herbarium material upon which it was based are available in the Sloane Herbarium (BM) and prove that A. philippense is indeed this species. Verma and Fraser-Jenkins (in S. C. Verma, S. P. Khullar & Cheema, Perspec. Pterido. 82–83. 2009 [“2008”], fidι Fraser-Jenkins, loc. cit.) recognized two further subspecies: the diploid sexual subsp. teestae S. C. Verma & Fraser-Jenkins in which they included Chinese material, the diploid and tetraploid apomictic subsp. intermedium S. C. Verma & Fraser-Jenkins, and epitypified subsp. philippense with material of a known triploid apomict from India.

Adiantum philippense is an indicator of acidic conditions, as it grows in soils of pH 4.5–5.

13. Adiantum capillus-junonis Ruprecht, Beitr. Pflanzenk. Russ. Reiches 3: 49. 1845.

团羽铁线蕨  tuan yu tie xian jue

Adiantum cantoniense Hance [??“cantonense”].

Plants epilithic, 8–20 cm tall. Rhizomes erect, short, scales dark brown, lanceolate, margins entire. Fronds clustered; stipe dark castaneous, glossy, 2–6 cm, as slender as iron wire, base covered with same scales as rhizome, distally glabrous; lamina 1-pinnate, lanceolate in outline, 8–15 Χ 2.5–3.5 cm, costae and stalks castaneous, apex usually prolonged into a whiplike stolon and rooting to form new plantlets. Pinnules 4–8 pairs, lower pinnules opposite, upper ones subopposite, spreading obliquely upward; petiolule 2–3 mm, articulate, persistent after pinnule fall; lower several pairs of pinnules subequal in size, orbicular-flabellate or orbicular, 1.1–1.6 Χ 1.5–2 cm, membranous, green, both surfaces glabrous, base symmetrically rounded-cuneate or rounded, lateral sides entire, upper margins rounded, upper margin of sterile pinnules denticulate; fertile pinnules with 2–5 shallow sinuses, denticulate on sterile parts; distal and terminal pinnules similar to lower pinnules but smaller; veins multidichotomously forked, reaching margins, visible on both surfaces. Sori 1–5 per pinna; false indusia brown, orbicular or reniform, papery, upper margins flat and straight, persistent. Perispore granulate.

Gregarious on wet limestone, in basal crevices of walls, shaded and wet chalky soil; 300–2500 m. Gansu, Guangdong, Guangxi, Guizhou, Hebei, Henan, Shandong, Sichuan, Taiwan, Yunnan [Japan, Korea].

The type specimen was collected in Beijing.

14. Adiantum edgeworthii Hooker, Sp. Fil. 2: 14. 1851.

普通铁线蕨  pu tong tie xian jue

Adiantum caudatum Linnaeus var. edgeworthii (Hooker) Beddome; A. caudatum var. rhizophorum Wallich ex C. B. Clarke; A. edgeworthii var. spencerianum (Copeland) Tagawa; A. guilelmi Hance; A. spencerianum Copeland.

Plants terrestrial, 10–30 cm tall. Rhizomes erect, short, scales black-brown lanceolate, margins denticulate. Fronds clustered; stipe dark purplish, glossy, 4–16 cm, base covered with scales, distally smooth; lamina 1-pinnate, linear-lanceolate in outline, 6–23 Χ 2–3 cm, base subattenuate, apex acuminate; rachis castaneous, with sparse reddish brown linear scales, smooth, apex often prolonged into a whiplike stolon and rooting to form new plantlets. pinnules 10–30 each side, opposite or alternate, if rachis prolonged into a whiplike stolon then upper pinnules gradually further apart from each other; petiolule ca. 1 mm, several basal pairs of pinnules shorter and relatively broader than middle pinnules and slightly reflexed, middle pinnules horizontally spreading, dimidiate, 1–1.5 Χ 0.5–0.8 cm, papery, dark brownish or brownish green, both surfaces glabrous, base asymmetrical with acroscopic side truncate, lower and inner margins straight and entire, upper margin shallowly 2–5-lobed, apex acute or obtuse; segments suborbicular, entire or slightly undulate; upper pinnules progressively smaller distally, terminal pinnules subflabellate, base cuneate, upper margins divided; veins multidichotomously forked, visible on both surfaces. Sori 2–5 per pinna, borne at apex of segments; false indusia orbicular or oblong, membranous, upper margins flat and straight, entire, persistent. Perispore granulate.

Shaded and wet places, on rocks; 700–2500 m. Gansu, Hebei, Henan, Liaoning, Shandong, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, N India, Japan, Malaysia, Myanmar, Nepal, Philippines, N Thailand, Vietnam].

15. Adiantum pedatum Linnaeus, Sp. Pl. 2: 1095. 1753.

掌叶铁线蕨  zhang ye tie xian jue

Adiantum boreale C. Presl; A. pedatum var. aleuticum Ruprecht; A. pedatum var. glaucinum C. Christensen (1927), non Christ (1898); A. pedatum var. kamtschaticum Ruprecht.

Plants terrestrial, 40–60 cm tall. Rhizomes erect or decumbent, scales dark brown, broadly lanceolate, margins entire. Fronds clustered or approximate; stipe castaneous or brown, 20–40 cm, covered with same scales as rhizome, distally glabrous; lamina pedately dichotomous, broadly flabellate in outline, up to 30 Χ 40 cm; pinnae 4–6 per branch, 1-imparipinnate, linear-lanceolate in outline, rachises and stalks castaneous-red, glabrous. inner pinnae up to 28 Χ 2.5–3.5 cm, outer pinnae slightly shorter; pinnules 20–30 pairs per pinna, alternate, obliquely spreading, petiolule 1–2.5 cm; basal pinnules slightly smaller, flabellate or semiorbicular, with longer petiolules; middle pinnules dimidiate, narrowly triangular, ca. 2 Χ 0.6 cm, herbaceous, green, both surfaces glabrous, base asymmetrical, cuneate, inner and lower margins straight and entire, apex undulate or with blunt teeth, upper margin divided to halfway, apex obtuse; segments ± square, entire or depressed at middle or undulate-crenate; distal pinnules similar but gradually smaller toward apices, terminal pinnules flabellate, divided at middle, bilateral sides lobed, equal in size or slightly larger than middle pinnules; veins multidichotomously forked, reaching margin, visible on both surfaces. Sori 4–6 per pinnule, horizontally attached in shallow sinuses; false indusia grayish green or dark brown, orbicular or reniform, membranous, entire, persistent. Perispore granulate.

Near stream margins of forest understories; 300–3500 m. Gansu, Hebei, Heilongjiang, Henan, Jilin, Liaoning, Qinghai, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan [Bhutan, NE India, Japan, Korea, Nepal; North America].

The whole plant is used to promote diuresis and to stop bleeding for dysuria, stanguria, toothaches, menorrhagia, and is also used to clear away lung-heat and relieve coughs.

16. Adiantum myriosorum Baker, Bull. Misc. Inform. Kew. 1898: 230. 1898.

灰背铁线蕨  hui bei tie xian jue

Adiantum myriosorum var. recurvatum Ching & Y. X. Lin; A. pedatum Linnaeus var. glaucinum Christ; A. pedatum var. myriosorum (Baker) Christ; A. pedatum var. protrusum Christ.

Plants terrestrial, 40–60 cm tall. Rhizomes erect or ascending, scales dark brown, broadly lanceolate, margins entire. Fronds clustered or approximate; stipe nearly black, 12–25 cm, covered with same scales as rhizome, distally glabrous; lamina pedately dichotomous, broadly flabellate in outline, 25–35 cm; pinnae 3–7 per branch, 1-imparipinnate, linear-lanceolate in outline, outer pinnae progressively shorter, rachises and stalks glabrous; pinnules 20–30 pairs per pinna, alternate, obliquely spreading, petiolulate; basal pinnules slightly smaller, flabellate or semiorbicular, with longer petiolules; middle pinnules dimidiate, triangular-elliptic, ca. 2 Χ 0.6 cm, herbaceous, abaxially glaucous, both surfaces glabrous, base asymmetrical, cuneate, inner and lower margins straight and entire, upper margin lobed, apex obtuse; segments ± square, with acute triangular teeth; distal pinnules similar to middle pinnules but gradually smaller toward apices, terminal pinnules flabellate, divided at middle, bilateral sides lobed, equal in size or slightly larger than middle pinnules; veins multidichotomously forked, reaching margin, visible on both surfaces. Sori 4–6 per pinnule, horizontally attached in shallow sinuses; false indusia grayish green or dark brown, orbicular-reniform, membranous, entire, persistent. Perispore reticulate.

Understories of dense forests; 900–2600 m. Anhui, Gansu, Guizhou, Henan, Hubei, Hunan, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, NE India, Kashmir, N Myanmar, Nepal].

Adiantum myriosorum is a beautiful plant that is cultivated as an ornamental.

17. Adiantum subpedatum Ching, Bull. Bot. Res., Harbin 3(3): 2(–3, f. 2.2). 1983.

昌化铁线蕨  chang hua tie xian jue

Plants terrestrial, 24–28 cm tall. Rhizomes erect, short scales brown, broadly lanceolate, Fronds clustered; stipe blackish, 10–15 cm, ca. 1 mm thick, glabrous; lamina pedately dichotomous, ca. 15 Χ 11–20 cm; pinnae 2–4(–5) per branch, 1-imparipinnate, linear-lanceolate in outline, inner pinnae up to 13 Χ 2 cm, outer pinnae slightly shorter; pinnules ca. 24 pairs per pinna, alternate, patent, petiolulate; basal pinnules slightly smaller, flabellate, with longer petiolules; middle pinnules oblong, ca. 1 Χ 0.3–0.4 cm, submembranous, drying green, base cuneate, inner and lower margins straight and entire, upper margin minutely acutely dentate, Veins extending into marginal teeth, visible abaxially. Sori 1 or 2 per pinnule, horizontally attached in shallow sinuses; false indusia grayish green, orbicular or reniform, membranous, persistent.

* On calcareous rocks; ca. 1000 m. Zhejiang.

The possibility that Adiantum subpedatum is a depauperate form of the preceding species, A. myriosorum, merits further investigation.

18. Adiantum hispidulum Swartz in Schrader, J. Bot. 1800: 82. 1802.

毛叶铁线蕨  mao ye tie xian jue

Adiantum pubescens Schkuhr; A. pedatum [sensu?] G. Forster (1786), non Linnaeus (1753).

Plants ca. 40 cm tall. Rhizomes erect, short, scales purple-black, lanceolate, margins subentire. Fronds clustered; stipe dark castaneous, glossy, ca. 20 cm, ca. 2 mm in diam., densely brownish villous, glabrescent; lamina dichotomously 2- or 3-pinnate, broadly ovate-triangular in outline, ca. 18 Χ 10 cm, middle pinnae usually longest, 1-imparipinnate; linear-lanceolate in outline, ca. 16 cm; all costae and stalks castaneous, densely brown pubescent; pinnules ca. 30 pairs per pinna, alternate, obliquely spreading, petiolule short; lower 1 or 2 pairs of pinnules slightly reduced, flabellate or flabellate-rectangular, middle pinnules nearly uniform in size, orbicular, ca. 8 Χ 4–5 mm, papery, dark brown or deep olive-green, both surfaces sparsely brown pubescent, base asymmetrically cuneate, inner and lower margins straight and entire, with narrow shallow sinuses, outer margins rounded, upper margins rounded-truncate, sterile parts undulate and dentate. all lateral pinnae same shape as middle pinnae and gradually becoming shorter toward bilateral sides; fertile segments truncate and entire on upper margins, distal pinnules gradually smaller toward apex; terminal pinnules obtriangular, slightly larger than lower lateral ones; veins multidichotomously forked, visible on both surfaces. Sori 4–12 at sinuses of pinnules; false indusia dark brown, orbicular, leathery, pubescent adaxially, upper margins deeply depressed or sinuate, entire, persistent. 2n = 360.

E Guangdong, E Taiwan, SE Yunnan [India, Indonesia (Java), Malaysia, Philippines; tropical and subtropical regions: Africa, Asia, Pacific islands].

19. Adiantum flabellulatum Linnaeus, Sp. Pl. 2: 1095. 1753.

扇叶铁线蕨  shan ye tie xian jue

Adiantum amoenum Wallich ex Hooker & Greville; A. fuscum Retzius.

Plants terrestrial, 20–45 cm tall. Rhizomes erect, short, scales dense, yellowish to brown, glossy, linear-lanceolate, margins entire. Fronds clustered; stipe black-purple, glossy, 10–30 cm, adaxially grooved with short stiff brown hairs inside, base covered with same scales as rhizome, distally glabrous; lamina pedately 2- or 3-dichotomously branched, flabellate in outline, 10–25 cm, middle pinnae usually longer, 1-imparipinnate; costae and stalks purple-black, abaxially glabrous, adaxially with dense, short, brown-red, hairs; middle pinnae linear-lanceolate, 6–15 Χ 1.5–2 cm, 1-pinnate; outer pinnae similar but slightly shorter, up to 5 cm; pinnules 8–15 pairs per pinna, alternate, horizontally spreading, petiolule 1–2 mm; pinnules below middle ± uniform in size, dimidiate-semiorbicular when fertile, rhomboid when sterile, 6–15 Χ 5–10 mm, thinly leathery, green or dark brown, both surfaces glabrous, base broadly cuneate or flabellately cuneate, inner and lower margins straight and entire, outer and upper margins subrounded or rounded-truncate, fertile parts shallowly sinuate, segments entire, sterile parts denticulate; distal pinnules similar but slightly smaller; terminal pinnules obovate or flabellate, equal to or slightly larger than lower pinnules; veins multidichotomously forked and reaching margins, visible on both surfaces. Sori 2–5 per pinnule, horizontal, at upper and outer margins of segments; false indusia dark brown, semiorbicular or oblong, glabrous, upper margins flat and straight, entire, persistent. Perispore indistinctly granulate. 2n = 116.

Open acidic red and yellow soils; 100–1100 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [India, Indonesia, Japan (including Ryukyu Islands), Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, Vietnam].

The whole plant is used in traditional Chinese medicine to clear away heat and toxic materials, relax tendons and activate blood, promote diuresis, dissipate phlegm, subsidence of swelling, stop bleeding and alleviate pain from trauma; it is applied to burns and scalds, bites from poisonous snakes, centipedes, and scabies; it is also used to treat inflammatory diseases of pigs with redness of skin and rinderpests of ruminants.

The plant is an indicator of acidity, growing on red and yellow soils with a pH of 4.5–5.

20. Adiantum induratum Christ, J. Bot. (Morot) 21: 265. 1908.

圆柄铁线蕨  yuan bing tie xian jue

Plants 15–40 cm tall. Rhizomes erect, short, scales dense, dark brown, Fronds clustered; stipe brown when young, black when old, glossy, 4–20 cm, less than 1.5 mm in diam., adaxially grooved with stiff brown hairs inside, terete when old, glabrescent; lamina 2–4-pinnate, broadly ovate in outline, 10–20 Χ 6–10 cm; pinnae 2–4 each side, obliquely spreading, stalked; rachis, costae and stalks dark brown and glossy, abaxially glabrous, adaxially densely brown-yellow pubescent, basal pair of pinnae largest, narrowly ovate in outline, 4–9 Χ 2–4 cm, stalk 1–1.2 cm, 2- or 3-paripinnate; outer pinnae progressively shorter; pinnules 2 pairs per pinna, alternate, oblique upward, petiolule ca. 3 mm; articulate at end, persistent after pinnule fall; pinnules of fertile fronds subrounded, 5–9 Χ 8–11 mm, subleathery, dark brown-yellow or dark brown-green, abaxially sometimes slightly glaucous, both surfaces glabrous, base broadly cuneate or truncate, inner and lower margins straight and entire, outer and upper margins rounded or subrounded, entire or few sinuate; pinnules of sterile fronds slightly larger, oblong, 9–12 Χ 11–16 mm, bases broadly cuneate, entire, outer and upper margins undulately lobed, denticulate; distal pinnules similar but slightly smaller; terminal pinnules: petiolule 2–4 mm, blade flabellate, slightly larger than lower pinnules. Veins multidichotomously forked, reaching margins, visible on both surfaces. Sori 4–6 (rarely 1 or 2) on ultimate pinnules, horizontally attached at upper margins; false indusia dark brown, reniform, broadly linear, or oblong, leathery, upper margins truncate or slightly depressed, entire, persistent.

On acidic soil of forest understories by roadsides or forest margins; 100–800 m. Guangdong, Guangxi, Hainan, Yunnan [Vietnam].

21. Adiantum diaphanum Blume, Enum. Pl. Javae 2: 215. 1828.

长尾铁线蕨  chang wei tie xian jue

Adiantum affine Willdenow; A. diaphanum var. affine (Willdenow) Alderwerelt; A. heteromorphum Colenso ex Field, nom. nud.; A. setulosum J. Smith.

Plants epilithic, 15–30 cm tall. Rhizomes erect, short, scales dark brown, lanceolate, margins denticulate. Fronds clustered; stipe castaneous, glossy, 4–20 cm, slender, adaxially grooved, base sparsely covered with same scales as rhizome, distally glabrous; lamina 1-pinnate or 2-pinnate with a few lateral pinnae nearly as large as terminal pinna, deltoid-lanceolate in outline, 6–11 Χ 2–3 cm; rachis, costae, and petiolules castaneous, glabrous; pinnules 8–16 pairs per pinna, alternate, obliquely spreading or lower ones subhorizontally spreading, petiolule less than 2 mm; pinnules uniform in size, dimidiate, rhombic or quadrangular, 1–1.8 Χ 0.5–0.9 cm, membranous, abaxially slightly glaucous and with lax, appressed, brown, unicellular needlelike setae; adaxially glossy, dark brown or deep olive-green, base asymmetrical, broadly cuneate, margin with rounded sinuses, outer margin obtuse or truncate, upper margin truncate or rounded, distal pinnules similar but slightly smaller; terminal pinnules rhombic, slightly larger; veins flabellately forked, reaching margins, visible on both surfaces. Sori 2–10 per pinnule, inserted in sinuses between lobes except those attached along veinlet but also on mesophyll between veinlets; false indusia dark brown, orbicular, leathery, upper margins deeply sinuate and with unicellular brown needlelike setae, persistent. Perispore indistinctly granulate. 2n = 116, 232.

Wet soils or on rocks near stream margins within forests; 400–2200 m. Fujian, Guangdong, Hainan, Jiangxi, Taiwan [Indonesia, Malaysia, Vietnam; Australia, Pacific islands (New Zealand, Polynesia)].

22. Adiantum fengianum Ching, Bull. Fan Mem. Inst. Biol. Bot., n.s., 1: 267. 1949.

冯氏铁线蕨  feng shi tie xian jue

Plants ca. 10 cm tall. Rhizomes decumbent, slender, scales dense, dark brown, Fronds approximate; stipe dark castaneous, glossy, 3–6 cm, as slender as silk, smooth above base; lamina 2-pinnate, narrowly ovate to lanceolate in outline, 3–5 Χ 1–2 cm, apex obtuse; pinnae 4 or 5 each side, horizontally or obliquely spreading, rachises, costae, and petiolules same as stipe; pinnae below middle only, ± uniform in size, imparipinnate, narrowly ovate in outline, 1–1.5 Χ ca. 0.8 cm; pinnules alternate, petiolule short, slender; pinnules spreading obliquely upward, uniformly obtriangular, ca. 4 mm, progressively slightly smaller distally, herbaceous, green, both surfaces glabrous, sterile pinnules: base symmetrically cuneate, bilateral margins straight and entire, upper margins divided and sharply serrate, apex rounded, fertile pinnules apex truncate or with few sharp serrations on bilateral acroscopic corner; veins multidichotomously forked and reaching margins, visible on both surfaces. Sori 1 per pinnule, horizontally attached at upper margin; false indusia glaucous or dark brown, elongate, nearly as wide as upper margin of pinnule, membranous, upper margins truncate or slightly undulate, persistent. Perispore indistinctly granulate.

* Wet rocks of forest understories; ca. 3400 m. N Yunnan (Lijiang, Yulong Shan).

Adiantum fengianum is listed as endangered in the IUCN Red Book (A2c; B1ab(iii) <http://www.iucnredlist.org/apps/redlist/details/46602/0>, accessed 12/07/11.

23. Adiantum bonatianum Brause, Hedwigia 54: 206. 1914.

毛足铁线蕨  mao zu tie xian jue

Plants 25–60 cm tall. Rhizomes creeping, slender, scales black lanceolate, also brown multicellular villous. Fronds approximate; stipe black-purple, glossy, 10–20 cm, base densely scaly and villous, hairs easily rubbed off when dried to leave rough feeling surface, distally smooth; lamina 3- or 4-pinnate, broadly ovate to triangular in outline, 20–40 Χ 19–25 cm, apex acuminate; primary pinnae 5–7 each side, 2- or 3-pinnate, obliquely spreading, stalk ca. 1 cm; rachis and costae same color as stipe, glabrous; basal 1 or 2 pairs of pinnae largest, triangular, 8–18 Χ 4–9 cm; pinnules 2–4, alternate; petiolule castaneous-red, ca. 1 mm, hairlike; blade flabellate, 5–9 Χ 4–11 mm, thinly herbaceous, green, both surfaces glabrous, base symmetrical, cuneate, bilateral margins entire, apex rounded, with dense short and broad triangular serrations sometimes with cartilaginous, often crooked, tips extended into long awns; veins multidichotomously forked, reaching tips of serration, visible on both surfaces. Sori 1–4 per pinnule; false indusia dark brown, orbicular or orbicular-reniform, membranous, upper margins deeply sinuate, entire, persistent. Perispore indistinctly granulate.

* Slightly acidic, damp soil in forest understories, in rock crevices at forest margins; 1400–2500 m. Guizhou, Sichuan, Yunnan.

1a. Apical serrations of ultimate pinnules with cartilaginous, usually crooked long awns at tips  23a. var. bonatianum

1b. Apical serrations of ultimate pinnules acute and without cartilaginous awn at tip  23b. var. subaristatum

23a. Adiantum bonatianum var. bonatianum

毛足铁线蕨(原变种)  mao zu tie xian jue (yuan bian zhong)

Apical serrations of ultimate pinnules extended into cartilaginous, usually crooked, long awn points.

* Slightly acidic damp soil in forest understories, in rock crevices at forest margins; 1400–2500 m. Guizhou, Sichuan, Yunnan.

23b. Adiantum bonatianum var. subaristatum Ching, Acta Phytotax. Sin. 6: 338. 1957.

无芒铁线蕨  wu mang tie xian jue

Adiantum pseudobonatianum Ching, nom. nud.

Apical serrations of ultimate pinnulets acute, not extended into cartilaginous awn points.

* On rocks of forest understories; ca. 1400 m. Sichuan.

24. Adiantum davidii Franchet, Nouv. Arch. Mus. Hist. Nat., ser. 2, 10: 112. 1887.

白背铁线蕨  bai bei tie xian jue

Adiantum aristatum Christ; A. davidii var. aristatum (Christ) C. Christensen; A. davidii var. latedeltoideum (Christ) Ching; A. davidii var. longispinum Ching; A. monochlamys Eaton var. latedeltoideum Christ; A. prattii Baker.

Plants epilithic, 20–30 cm tall; rhizomes long creeping, slender, scales dark brown, glossy, lanceolate, margins entire. Fronds spaced apart; stipe dark castaneous, 10–21 cm, with same scales as rhizome, distally glabrous; lamina 3-pinnate, triangular-ovate in outline, 10–15 Χ 6–10 cm, apex acuminate; pinnae 3–5 each side, obliquely spreading, rachises, costae, and stalks same color as stipes, with brown multicellular, articulate hairs at base of pinnae and pinnules. basal pair of pinnae largest, 2-pinnate, narrowly triangular in outline, 5–7 Χ 3–4 cm; pinnules 4 or 5 pairs per pinna, slightly overlapping, obliquely spreading, apex obtuse; petiolule castaneous 1–4 mm, slender; pinnules uniform in shape, flabellate, 4–7 Χ 4–7 mm, progressively smaller distally, hard herbaceous, abaxially gray-green, adaxially green or glaucous, both surfaces glabrous, base cuneate, bilateral margins entire, apex rounded, with dense short and broad serrations with short cartilaginous awn points; Veins multidichotomously forked, reaching apices of serrations, visible on both surfaces. Sori 1 (rarely 2) per pinnule, horizontally attached at apical sinus of pinnule; false indusia dark brown, reniform or orbicular-reniform, papery, upper margins shallowly depressed, entire, persistent. Perispore thickly granulate. 2n = 240*.

* On rocks by streamsides; 1100–3400 m. Gansu, Guizhou, Hebei, Henan, Shaanxi, Shanxi, Sichuan, Yunnan.

The type specimen was collected in Shaanxi.

25. Adiantum venustum D. Don, Prodr. Fl. Nepal. 17. 1825.

细叶铁线蕨  xi ye tie xian jue

Adiantum venustum var. wuliangense Ching & Y. X. Lin.

Plants terrestrial, 25–50 cm tall. Rhizomes decumbent, thick, scales dense, dark brown, ovate-lanceolate, margins entire. Fronds remote; stipe castaneous-brown, 10–20 cm, base covered with same scales as rhizome, distally glabrous; lamina 3-pinnate, broadly ovate in outline, 10–25 Χ 5–12 cm, apex acuminate and 1-pinnate, pinnae ca. 6 each side, obliquely spreading, stalk 2–7 mm; rachises, costae, and stalks ± zigzag, same color as stipe, glabrous, basal pair of pinnae largest, ovate-elliptic, 5–13 Χ 4–7 cm, 2-pinnate; pinnules 4 or 5 pairs per lateral pinna, alternate, upwardly oblique; petiolule short; pinnules flabellate, 7–10 Χ 7–10 mm, green or brown-green, herbaceous, both surfaces glabrous, base shortly cuneate, with short stalk, apex rounded and denticulate. Veins multidichotomously forked, reaching teeth at pinnule margin, visible on both surfaces; Sori 1 or 2 (rarely 3) per pinnule, attached at rounded sinus; false indusia brownish, orbicular, membranous, upper margin deep sinuslike, entire, persistent. 2n = 120.

In rock crevices, mountain slopes; 2000–2900 m. Xizang, Yunnan (Jingdong, Wuliangshan) [Bhutan, India, Kashmir, N Myanmar, Nepal].

26. Adiantum tibeticum Ching, Acta Phytotax. Sin. 18: 104. 1980.

西藏铁线蕨  xi zang tie xian jue

Plants ca. 30 cm tall. Rhizomes creeping, ca. 1.5 mm in diam., scales dense, brown, lanceolate. Fronds remote, erect; stipe castaneous-brown, glossy, ca. 15 cm, terete, adaxially grooved, distally glabrous; lamina 3-pinnate, ovate in outline, 10–16 Χ 5–7 cm, distally 1-pinnate; pinnae 4 or 5 each side, oblique upward, with short stalks; rachises, costae, and stalks zigzag, same color as stipes, smooth; basal pinnae 2-pinnate, narrowly ovate in outline, ca. 6 Χ 2.8 cm; pinnules 3 or 4 pairs per pinna; petiolule short, silklike[?very slender]; fertile pinnules flabellate, usually 7–11(–13) Χ 7–13 mm, thinly herbaceous, abaxially glaucous, adaxially greenish, both surfaces glabrous, base cuneate, oblique to irregularly shaped, apex rounded, with rounded sharp triangular teeth; terminal pinnules slightly larger. Veins multidichotomously forked, obscure. Sori 1 or 2(or 4) on ultimate pinnule; false indusia brownish, orbicular-reniform or narrowly reniform, membranous, upper margin with a large depression, entire, persistent.

* Forest understories by roadsides; 2800–3200 m. SE Xizang (Jilong).

27. Adiantum fimbriatum Christ, Bull. Soc. Bot. France: Mem. 1: 62. 1905.

长盖铁线蕨  chang gai tie xian jue

Adiantum fimbriatum var. shensiense (Ching) Ching & Y. X. Lin; A. smithianum (C. Christensen) Ching; A. smithianum (C. Christensen) Ching var. shensiense Ching; A. venustum D. Don var. smithanum C. Christensen.

Plants terrestrial or epilithic, (10–)25–35 cm tall. Rhizomes creeping, slender, scales dense, brown, glossy, ovate-lanceolate. Fronds spaced; stipe reddish castaneous, glossy, 10–20 cm, base covered with same scales as rhizome, distally glabrous; lamina ovate-triangular in outline, 7–25 Χ 10–20 cm, 3- or 4-pinnate, apex obtuse; pinnae 3–7 each side, oblique upward, stalk ca. 10 mm; rachises, costae, and stalks zigzag, same color as stipes, glossy, basal pair of pinnae largest, 2-pinnate, ovate-triangular in outline, 6–10 Χ 4–8 cm, apex obtuse; distal pinnae 1-pinnate. pinnules 3–5 pairs per ultimate pinna, petiolulate, pinnule obovate or flabellate, 6–7 Χ 6–7 mm or longer than wide, progressively smaller distally, thinly herbaceous, greenish or gray-green, both surfaces glabrous, base symmetrical or slightly oblique, cuneate, bilateral sides entire, apex rounded or ± oblique, with dense small sharp narrowly triangular teeth without awned tips. Veins flabellately forked, ending in marginal teeth, visible on both surfaces. Sori (1 or)2 or 3(–5) per pinnule; false indusia brownish, rectangular, reniform, orbicular, or orbicular-reniform, membranous, less often slightly depressed, upper margins more often flat and straight, entire, persistent. Perispore indistinctly granular.

* On rocks or in rock crevices of forest understories by streamsides; 2700–3600 m. Gansu, Hebei, Qinghai, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan.

28. Adiantum breviserratum (Ching) Ching & Y. X. Lin, Acta Phytotax. Sin. 18: 104. 1980.

圆齿铁线蕨  yuan chi tie xian jue

Adiantum venustum D. Don var. breviserratum Ching, Acta Phytotax. Sin. 6. 335. 1957.

Plants 20–30 cm tall. Rhizomes creeping, scales dense, brown, lanceolate. Fronds tufted or approximate; stipe castaneous-brown, glossy, 10–20 cm, base with same scales as rhizome, distally smooth; lamina 3- or 4-pinnate, subovate-triangular in outline, ca. 15 Χ 10 cm, apex acuminate; primary pinnae 5 or 6 each side, spreading obliquely upward, stalks 2–6 mm; basal pair of pinnae largest, 2-pinnate, ovate or subovate-triangular in outline, 7–9 Χ 2.5–4 cm, apex obtuse; secondary pinnae 4 or 5 pairs, basal pair of pinnae largest, 1-pinnate, broadly ovate in outline, 2.5–3.5 Χ ca. 1.5 cm, apex obtuse; pinnules 2–4 pairs per ultimate pinna, petiolule silklike; pinnule obovate or narrowly flabellate, 3–6 Χ 3–6 mm, progressively smaller distally, thinly herbaceous, grayish green, both surfaces glabrous, base cuneate, bilateral sides entire, upper margins rounded and crenate, Veins multidichotomously forked, reaching teeth of upper margin of pinnule, visible on both surfaces. Sori 1 per pinnule, horizontally attached at upper margin; false indusia brownish, suborbicular or rectangular, membranous, upper margins flat and straight, upper margins flat and straight, entire, persistent.

* On limestone; 2400–3500 m. Xizang, N Yunnan.

The type specimen was collected in Yunnan (Zhenkang).

29. Adiantum monochlamys D. C. Eaton, Proc. Amer. Acad. Arts 4: 110. 1858.

单盖铁线蕨  dan gai tie xian jue

Adiantum monochlamys var. simozawai Masamune; A. veitchii Hance; A. venustum D. Don var. monochlamys (Eaton) Luerssen.

Plants ?terrestrial, 15–55 cm tall. Rhizomes long creeping, scales dense, castaneous-black, glossy, narrowly lanceolate, margins entire. Fronds close together or spaced; stipe castaneous-black or castaneous, glossy, 15–28 cm, base covered with same scales as rhizome, distally glabrous; lamina, 2(–3)-pinnate, narrowly ovate-triangular in outline, 15–30 Χ 3–10 cm, base broadly cuneate, apex acuminate and 1-pinnate, pinnae 6–8 each side, spreading obliquely upward, stalk 1–1.5 cm; rachises, costae, and stalks zigzag, same color as stipes, glabrous; basal pair of pinnae largest, triangular-ovate, 4–8 Χ 3–4.5 cm; 2-pinnate; secondary pinnae 2 or 3 pairs per pinna; pinnules 3–5 pairs per pinna, obliquely ascending, petiolule castaneous, short, slender; pinnules obtriangular, 6–10 Χ 5–8 mm, progressively slightly smaller distally, herbaceous, abaxially grayish green, both surfaces glabrous, base cuneate, apex rounded-truncate; fertile pinnules deeply depressed at middle, both sides with sharp triangular teeth, bilateral margins straight and entire; terminal pinnules with longer petiolule. Veins multidichotomously forked, reaching ends of teeth of pinnules, visible on both surfaces. Sori 1 (or 2) per pinnule, in sinus of upper margin; false indusia dark red-brown, reniform, thinly papery, upper margin deeply sinuate, entire or slightly undulate, persistent. 2n = 116.

Forest understories on mountain lands, shaded rocks and cliffs in forests; 700–2500 m. Guizhou, Sichuan, Taiwan, Zhejiang [Japan, S Korea].

Adiantum monochlamys var. plurisorum Christ is an invalid nomen nudum.

30. Adiantum erythrochlamys Diels, Bot. Jahrb. Syst. 29: 201. 1900.

肾盖铁线蕨  shen gai tie xian jue

Adiantum roborowskii Maximowicz var. robustum Christ.

Plants epilithic, 16–35 cm tall. Rhizomes shortly creeping or ascending, scales dense, castaneous-black, glossy, narrowly lanceolate, margins entire. Fronds clustered or closely spaced; stipe castaneous, glossy, 5–22 cm, base densely covered with same scales as rhizome, distally glabrous; lamina 3-pinnate, lanceolate-triangular in outline, 4–22 Χ 4–8 cm, base cuneate, apex acuminate; pinnae 4–7 each side, stalk 5–10 mm; costae and stalks similar to stipe, glabrous, obliquely ascending; basal pinnae slightly larger, 2-pinnate below middle, narrowly ovate, 2.5–4 Χ ca. 2 cm, and with 2 pairs of pinnules; distal pinnae 1-pinnate; pinnules 3 or 4 pairs per pinna, alternate, obliquely ascending; petiolule ca. 1 mm, slender, pinnules narrowly flabellate or obovate, 5–14 Χ 4–10 mm, progressively slightly smaller distally, papery, yellow-green or dark brown-green, both surfaces glabrous, base narrowly cuneate, bilateral sides symmetrical, undulate-crenate, outer and inner margins entire; sterile pinnules with upper margins distinctly bluntly triangular serrate, upper margin of fertile pinnules with wide and deep sinus at middle. Veins multidichotomously forked, reaching margins, visible on both surfaces; Sori 1 (or 2) per pinnule, in sinus of upper margin; false indusia dark brown, orbicular or orbicular-reniform, subleathery, upper margins deeply sinuate, entire, persistent. Perispore granulate.

* On rocks or in rock crevices of forest understories by streamsides; 600–3500 m. Guizhou, Henan, Hubei, Sichuan, Taiwan, Xizang.

31. Adiantum roborowskii Maximowicz, Mιlanges Biol. Bull. Phys.-Math. Acad. Imp. Sci. Saint-Pιtersbourg 11: 867. 1883.

陇南铁线蕨  long nan tie xian jue

Plants epilithic, 9–25(–35+) cm tall. Rhizomes erect or ascending, short, scales dense, dark brown, margins entire. Fronds clustered or approximate; stipe reddish castaneous, glossy, 2–20 cm, terete, base covered with same scales as rhizome, distally glabrous; lamina 3-pinnate on lower part, 1-pinnate on upper part, lanceolate or ovate-elliptic in outline, 2–18 Χ 2–6 cm, apex acuminate, pinnae 3–6 each side, all parts obliquely ascending, stalk 3–6 mm; rachises, costae, and stalks reddish castaneous, glabrous. basal pair of pinnae slightly larger, 2-pinnate, ovate-triangular, 1.5–3 Χ 1.2–1.4 cm, base rounded-cuneate, apex obtuse; distal pinnae progressively smaller, 1-pinnate; pinnules 1 or 2 pairs per pinna, alternate; petiolule reddish castaneous; short, slender, pinnules subtriangular, narrowly flabellate, triangular, broadly ovate or suborbicular-flabellate, 5–9 Χ 4–7 mm, papery or nearly hard papery, gray-green, both surfaces glabrous, base broadly or rounded cuneate, sterile pinnules with upper margins rounded and with slightly undulate projection; upper margins of fertile pinnules entire and with 1 or 2 deep sinuses at middle. Veins multidichotomously forked, nearly reaching margins, visible on both surfaces; Sori 1 or 2 per pinnule, in deep sinuses of upper margin; false indusia dark brown, orbicular or orbicular-reniform, subleathery, upper margins deeply sinuate, entire, persistent.

* In rock crevices of wet forest understories, on mossy or damp rocks and cliffs, on rocks or cliffs by ditch sides; 1000–3500 m. Gansu, Guizhou, Hubei, Qinghai, Shaanxi, Sichuan, Xizang.

1a. Ultimate pinnules suborbicular-flabellate, base rounded-cuneate ............................  31c. var. faberi

1b. Ultimate pinnules triangular, obovate, or broadly ovate, base broadly cuneate.

2a. Stipe 4–20 cm, lamina 5–18 cm, pinnules narrowly flabellate .................  31a. var. roborowskii

2b. Stipe 2–8 cm, lamina 2–7 cm, pinnules triangular or broadly ovate
..........................................................................................................  31b. var. taiwanianum

31a. Adiantum roborowskii var. roborowskii

陇南铁线厥(原变种)  long nan tie xian jue (yuan bian zhong)

Stipe 4–20 cm; lamina 4–18 cm, ultimate pinnules narrowly flabellate, base broadly cuneate.

* In rock crevices of wet forest understories, on cliffs or rocks by ditch sides; 1000–2000 m. Gansu, Guizhou, Qinghai, Shaanxi, Sichuan.

The type specimen was collected in Qinghai.

31b. Adiantum roborowskii var. taiwanianum (Tagawa) W. C. Shieh, J. Chinese Forestry 2: 167. 1968.

台湾高山铁线蕨  tai wan gao shan tie xian jue

Adiantum taiwanianum Tagawa, Acta Phytotax. Geobot. 4: 93. 1935.

Stipe 2–8 cm; lamina 2–7 cm, ultimate pinnules triangular or broadly ovate, base broadly cuneate.

* On mossy rocks and cliffs; 2200–3500 m. C and S Taiwan.

31c. Adiantum roborowskii var. faberi (Baker) Y. X. Lin & Prado, comb. nov.

峨眉铁线蕨  e mei tie xian jue

Basionym: Adiantum faberi Baker, J. Bot. 26: 225. 1888; A. roborowskii f. faberi (Baker) Y. X. Lin.

Ultimate pinnules suborbicular-flabellate, base rounded-cuneate.

* On damp rocks; 1200–3000 m. Hubei, Sichuan.

32. Adiantum formosanum Tagawa, J. Jap. Bot. 14: 315. 1938.

深山铁线蕨  shen shan tie xian jue

Rhizomes creeping, short, scaly. Stipe dark castaneous, glossy, 5–12 cm. lamina 2- or 3-pinnate, narrowly ovate-triangular in outline, 4–10 Χ 3–4 cm; pinnae 3 or 4 each side, stalked; rachises and stalks zigzag, smooth, basal pair of pinnae slightly larger, 1- or 2-pinnate; pinnules 3 or 4 pairs per pinna, alternate, petiolule short, pinnules fan-shaped, 5–10 Χ 4–6 mm, herbaceous, both surfaces glabrous, base cuneate, upper margin rounded, entire or shallowly 2-lobed. Sori (1–)2(–4) per pinnule, on lobe apices of upper margin; oblong to linear; indusium reniform.

* On wet mossy rocks and cliffs; 2300–3500 m. Taiwan.

33. Adiantum refractum Christ, Bull. Acad. Int. Gιogr. Bot. 11: 224. 1902.

月芽铁线蕨  yue ya tie xian jue

Adiantum delavayi Christ; A. edentulum Christ; A. edentulum f. muticum (Ching) Y. X. Lin; A. edentulum f. refractum (Christ) Y. X. Lin; A. muticum Ching; A. subemarginatum Christ, p.p.; A. veneris Linnaeus var. sinuatum Christ.

Plants epilithic, 15–30(–50) cm tall. Rhizomes erect or ascending, short, scales dense, brown, lanceolate, margins entire. Fronds clustered; stipe castaneous-black, glossy, 5–17 cm, base covered with same scales as rhizome, distally smooth; lamina usually 2- or 3-pinnate, ovate or ovate-lanceolate in outline, 10–15 Χ 4.5–12 cm, base cuneate, apex acute; pinnae 4–6 each side, all parts obliquely ascending, stalked, rachises and stalks similar to stipe, zigzag, smooth, basal pair of pinnae largest, 1- or 2-pinnate, narrowly ovate or narrowly triangular-ovate in outline, 2.5–12 Χ 2–3 cm; pinnae from second pair upward all similar but progressively smaller; pinnules 4 or 5 pairs per pinna, alternate; petiolule 0.5–4 mm, slender as silk; pinnules asymmetrical, flabellate, 5–15 Χ 8–14 mm, papery, abaxially gray-green, both surfaces glabrous, base shortly cuneate, bilateral sides entire, upper margin undulate-rounded, 1–4-lobate or semilobate (divided to middle), sterile pinnules entire or slightly undulate, fertile segments shallowly 2 or 4-lobed, terminal pinnules similar but slightly larger. Veins multidichotomously forked, reaching margins, visible on both surfaces. Sori 2–4(or 5) per pinnule, on lobe apices of upper margin; false indusia brown, narrowly reniform or orbicular-reniform, membranous, upper margins flat and straight or depressed, entire, persistent. Perispore reticulate.

* On rocks of forest understories, wet rocks covered with bryophytes in ditches, shaded and wet cliffs, ditch sides of mixed forest understories; 1000–3600 m. Guizhou, Hubei, Hunan, Shaanxi, Sichuan, Xizang, Yunnan, Zhejiang.

Adiantum refractum is very similar to the Himalayan species A. wattii Baker, and perhaps should be included within that species.

34. Adiantum capillus-veneris Linnaeus, Sp. Pl. 2: 1096. 1753.

铁线蕨  tie xian jue

Adiantum capillus-veneris f. dissectum (M. Martens & Galeotti) Ching; A. capillus-veneris f. fissum (Christ) Ching; A. capillus-veneris var. fissum Christ; A. capillus-veneris var. trifidum Christ; ?A. lingii Ching; A. michelii Christ; A. subemarginatum Christ, p.p.; A. tenerum Swartz var. dissectum M. Martens & Galeotti.

Plants terrestrial or epilithic, 10–40 cm tall. Rhizomes creeping, slender, scales dense, brown, lanceolate, margins entire. Fronds remote or closely spaced; stipe castaneous-black, glossy, 3–20 cm, slender, base covered with same scales as rhizome, distally glabrous; lamina mostly 2-pinnate below middle, 1-pinnate above middle, ovate-triangular in outline, 6–25 Χ 8–16 cm, base cuneate, apex acute; pinnae 3–5 each side, obliquely ascending, stalk up to 15 mm, rachises, costae, and stalks same color as stipes, slightly zigzag; color passing into lamina base; basal pair of pinnae larger, 1(or 2)-pinnate, narrowly ovate in outline, 3–9 Χ 2.5–4 cm, apex obtuse, pinnae from second pair upward all similar but progressively smaller; pinnules 2–4 pairs per ultimate pinna, alternate, obliquely ascending, petiolule castaneous-black, 1–2 mm, slender, pinnules subequal in size or basal pair slightly larger, 12–20 Χ 10–15 mm, thinly herbaceous, green or dark brown-green, both surfaces glabrous, base cuneate, sides entire, upper margin rounded, 2–4-lobed or divided into twiglike segments; sterile pinnules with apex obtuse with marginal teeth broadly triangular or erose; fertile segments with apex truncate, straight or slightly depressed, entire or with erose teeth on both sides; terminal pinnules flabellate, usually larger, base narrowly cuneate, petiolule up to 1 cm. Veins multidichotomously forked, reaching margins, visible on both surfaces. Sori 3–10 per pinnule, on apices of lobes of upper margin; indusium yellowish green, brown when old, narrowly reniform or orbicular-reniform, membranous, upper margins flat and straight, entire, persistent. Perispore thickly granulate.

On limestone near running streams, bottom of limestone caves, cliffs of dripping water; 100–2800 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Shanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Japan, Vietnam; widely distributed in temperate and tropical regions in Africa, America, Asia, Europe, Oceania].

Adiantum capillus-veneris is an indicator of calcareous soil.

Plants with ultimate pinnules shallowly divided on upper margins have been treated as f. capillus-veneris, while plants with ultimate pinnules deeply divided into several lobes have been treated as f. dissectum (M. Martens & Galeotti) Ching.

The whole plant is used in traditional medicine to clear away heat, urinary problems, lymph tuberculosis, and to stop coughs.

Material of Adiantum lingii Ching (Acta Phytotax. Sin. 6: 341. 1957, from Sichuan) has not been seen and is placed here on the basis of the protologue. It was compared with A. capillus-veneris but differed by the smaller pinnules that dry olive-green, each with 4–6 sori with orbicular to orbicular-reniform false indusia.

18. ANTROPHYUM Kaulf., Enum. Fil. 197, 282. 1824.

车前蕨属  che qian jue shu

Zhang Xianchun (张宪春); Michael G. Gilbert

Small to medium-sized epiphytic or epilithic ferns. Rhizome short erect or creeping, densely covered with clathrate, iridescent scales, and roots with numerous water-absorbing root hairs. Fronds simple; laminae fleshy, or leathery, shrunk when dry, broadly lanceolate or oblanceolate, sometimes linear, spatulate, obovate, or suborbicular, rarely forked at apex; mostly gradually narrowed into a stipelike base; costa usually only present in basal part; lateral veins abundantly reticulate, without free included veinlets. Sori forming coenosorus, soral lines superficial or immersed, on lateral veins, netted or branched, less often simple; paraphyses abundant, club-shaped with a capitate head, or taeniform, or filiform. Spores trilete, tetrahedral-globose, surface low-papillate, often with scattered spherules and rodlets.

About 40 ill-defined species: Old World tropics; nine species in China.

The species, which badly need taxonomic revision, are either epiphytic or epilithic. In dry periods the fronds shrivel and curl up to some extent. Like most tropical plants, the species of Antrophyum reach their northernmost limits of distribution in mainland Asia, south of the Chang Jiang in C China, while along the Pacific islands up to C Japan.

Ching (1940, 1978) treats this genus in a rather strict sense, which does not include the tropical American Anetium, Polytaenium and Scoliosorus. This is also supported by the molecular studies of Crane (1995, 1997). Kramer (1990) believed that “Although the genus has been divided into the Old World Antrophyum s.s., with the costa evanescing above the base and paraphyses present among the sporangia, and the New World genus Polytaenium, with percurrent costa and without paraphyses, the overall characters are basically not much different, yet another monotypic American genus, namely Anetium might be included in it.”

The Asian species could be divided into three main groups according to the types of paraphyses, i.e., A. obovate group with capitate paraphyses; A. henryi group with taeniform paraphyses; and A. callifolium group with filamentous paraphyses. A. obovate group might be the most primitive group, while A. callifolium group be the most advanced one in this genus.

1a.       Paraphyses club-shaped, capitate (A. obovatum group).

2a.       Fronds 2–8 cm, 0.5–1 cm wide; lamina oblanceolate, spatulate, or elliptic; stipe indistinct  3. A. parvulum

2b.       Fronds 10–25 cm, broadest above middle, 2–8 cm wide.

3a.       Lamina obovate, as long as stipe; margin of scales toothed 1 .....................  1. A. obovatum

3b.       Lamina oblanceolate; margin of scales entire .............................................  2. A. castaneum

1b.       Paraphyses taeniform, or filiform, spirally twisted.

4a.       Paraphyses taeniform (A. henryi group).

5a.       Lamina oblong-lanceolate, 1.5–3 cm wide; stipe short or indistinct .........  4. A. formosanum

5b.       Lamina linear-lanceolate, 0.5–1.8 cm wide; stipe indistinct ...............................  5. A. henryi

4b.       Paraphyses filiform (A. callifolium group).

6a.       Lamina linear, 0.7–1 cm wide; sori usually in one row near margin on both sides of lamina  9. A. vittarioides

6b.       Lamina broad; soral lines many.

7a.       Lamina oblanceolate, 1–1.5 cm wide; stipe indistinct; soral lines simple, paralleled  8. A. coriaceum

7b.       Lamina oblong-lanceolate to broadly oblanceolate, 1.5–10 cm wide; soral lines usually netted.

8a.       Stipe 3–10 cm ....................................................................................  6. A. callifolium

8b.       Stipe indistinct ..................................................................................  7. A. sessilifolium

1. Antrophyum obovatum Baker, Kew Bull. 233. 1898.

长柄车前蕨  chang bing che qian jue

Rhizome short erect; scales dark brown, lanceolate, 6–10 mm, apex long acuminate, bristlelike, margin minutely denticulate, clathrate; fronds clustered; stipes 2–15 cm, appressed, base covered with scales like those on rhizome, upward with sparse different sized scales, gradually glabrous; laminae thin coriaceous, obovate, 2–10 Χ 2–8 cm, widest at middle or above, apex long acuminate or caudate, sometimes upper part irregular lacerate at margin, base attenuate along stipe; costa none; veins abundantly reticulate. Soral lines superficial or slightly immersed, on lateral veins, netted, fertile in middle part of lamina; paraphyses club-shaped with a capitate head. Spores trilete, tetrahedral-globose, surface papillate.

In evergreen broad-leaved forests, epiphytic on lower part of tree trunks or on rocks; 200–2400 m. Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, India, Japan, Myanmar, Nepal, N Thailand, Vietnam].

This species is very similar to A. plantagineum (Cav.) Kaulf., and need more studies.

2. Antrophyum castaneum H. Ito, J. Jap. Bot. 12: 473, f. 7: 1. 1936.

栗色车前蕨  li se che qian jue

Rhizome short erect or short creeping; scales dark brown, linear-lanceolate, 3–8 mm, apex long acuminate, bristlelike, margin entire, clathrate; fronds clustered; stipes 5–10 cm, base covered with scales like those on rhizome, upward with scales, gradually glabrous; laminae coriaceous, oblanceolate, 10–20 Χ 1–3 cm, widest above middle, apex acute, base attenuate along stipe; costa none; veins abundantly reticulate. Soral lines slightly immersed, netted or parallel; paraphyses club-shaped with a capitate head. Spores trilete, tetrahedral-globose, surface papillate.

* 1500–1900 m. Taiwan.

3. Antrophyum parvulum Blume, Enum. Pl. Javae 2: 110. 1828, & Fl. Jav. 78, t. 34, f. 3. 1828.

小车前蕨  xiao che qian jue

Rhizome short erect; scales brown, lanceolate, margin minutely denticulate, clathrate; fronds clustered; stipes indistinct, appressed, base covered with small scales, gradually glabrous; laminae coriaceous, oblanceolate, spatulate, or oblong, 1–6 Χ 0.5–1 cm, apex acuminate or rounded, often lacerate or bifurcate, base attenuate; costa indistinct or only visible at lower 1/3; veins abundantly reticulate, invisible on both surfaces. Soral lines slightly immersed, forked but not netted; paraphyses club-shaped with a capitate head. Spores trilete, tetrahedral-globose, surface papillate.

Epiphytic on tree trunks or on limestone; 400–1600 m. Hainan, Taiwan [India, Indonesia, Malaysia, Philippines, Thailand, Vietnam].

Antrophyum parvulum might be conspecific to A. immersum (Bory ex Willd.) Mett., which needs further studies. It is quite different from A. obovatum Baker by its small size and indistinct stipe.

4. Antrophyum formosanum Hieron., Hedwigia 57: 210. 1916.

台湾车前蕨  tai wan che qian jue

Rhizome slender, short creeping or ascending; scales dark brown, lanceolate, 3–5 mm, ca. 0.5 mm wide at base, apex long acuminate, margin minutely denticulate, clathrate; fronds clustered; stipes with wings; laminae coriaceous, oblong-lanceolate, 10–20 Χ 1.5–3 cm, widest at middle or above, apex acute, base long attenuate; costa indistinct; veins abundantly reticulate. Soral lines slightly immersed, netted; paraphyses taeniform. Spores trilete, tetrahedral-globose, surface papillate.

On moist rocks by streams in forests; below 1300 m. Taiwan [Japan (Ryukyu Islands)].

Antrophyum formosanum and its relative A. henryi Hieron. both have taeniform paraphyses, but the latter is much smaller with soral lines not anastomosing.

5. Antrophyum henryi Hieron., Hedwigia 57: 208. 1916.

车前蕨  che qian jue

Rhizome slender, short creeping or erect; scales pale-brown, linear-lanceolate, 1.5–3.5 Χ 0.1–0.3 mm, margin obviously denticulate, clathrate; fronds clustered; stipes indistinct; laminae subcoriaceous, linear-lanceolate, 5–15 Χ 0.8–1.5 cm, widest at middle or above, apex narrowly acute, base long attenuate; costa indistinct; veins abundantly reticulate, raised above, invisible below. Soral lines 3–5, zigzag, subparallel, continuous or interrupted, or forming nets, lower 1/3 not fertile; paraphyses taeniform. Spores trilete, tetrahedral-globose, surface papillate.

On moist mossy rocks or epiphytic on tree trunks in valley; 300–1600 m. Guangdong, Guangxi, Guizhou, ?Taiwan, Yunnan [India (Assam, Sikkim), N Thailand].

6. Antrophyum callifolium Blume, Enum. Pl. Javae 2: 111. 1828, & Fl. Jav. 83, t. 35. 1829.

美叶车前蕨  mei ye che qian jue

Antrophyum annamense Tardieu & C. Christensen.

Rhizome short erect; scales dark brown, linear-lanceolate, ca. 6 Χ 1 mm, apex long acuminate, bristlelike, margin sparsely denticulate, clathrate; fronds clustered; stipes 1–3 cm, appressed, glabrous; laminae coriaceous, obovate-lanceolate or oblong-lanceolate, 15–40 Χ 2–10 cm, widest above middle, apex acuminate or caudate, base attenuate along stipe; costa only visible at base; veins abundantly reticulate, raised above, invisible below. Soral lines continuous or interrupted, partly netted; paraphyses filiform, longer than sporangia. Spores trilete, tetrahedral-globose, surface papillate.

Epiphytic on tree trunks or epilithic on rocks; 100–1600 m. Guangxi, Hainan, Yunnan [Cambodia, India, Indonesia, Laos, Malaysia, Philippines, Sri Lanka, Thailand, Vietnam; Australia].

A very variable species, particularly in the width of the lamina. Antrophyum annamense Tardieu & C. Chr. represents an extreme form with large, broad lamina, which are found in Hainan and N Vietnam, but intermediate forms link them together. It might be conspecific to A. reticulatum (Forster) Kaulf.

7. Antrophyum sessilifolium Spring, Syst. Veg. 4: 67. 1827.

无柄车前蕨  wu bing che qian jue

Antrophyum cumingii Fιe, Mem. Foug. 4: 72, t. 4, f. 7. 1851–1852.

Rhizome short erect; scales linear-lanceolate, 5–6 mm, margin minutely denticulate, clathrate; fronds clustered; stipes indistinct; laminae coriaceous, obovate-lanceolate or oblong-lanceolate, 15–30 Χ 1.5–2.5 cm, widest at middle or above, base attenuate; costa visible at base. Soral lines immersed; paraphyses filiform. Spores trilete, tetrahedral-globose, surface papillate.

On moist mossy rocks in dense forests, also epiphytic and close to streams; 100–300 m. Taiwan (Lan Yu) [Philippines].

No specimens of this species from Taiwan are available for comparison. Some specimens from Luzon in the Kew herbarium determined as this species are quite like A. coriaceum (D. Don) Wallich ex T. Moore.

8. Antrophyum coriaceum Wallich ex T. Moore, Index Fil. 5: 80. 1858.

革叶车前蕨  ge ye che qian jue

Rhizome short creeping; scales dark brown, subulate-lanceolate, 6–9 mm, ca. 0.5 mm wide at base, apex acuminate, bristlelike, margin denticulate, clathrate; fronds clustered; stipes indistinct; laminae coriaceous, oblanceolate, 15–30 cm, ca. 2 cm wide at middle, widest above middle, up to 3 cm wide, apex rounded, base long attenuate to very base; costa only visible at base; veins abundantly reticulate, raised above, forming several parallel lines, invisible below. Soral lines continuous, rarely interrupted, parallel, not netted, only lower 1/3 fertile; paraphyses filiform, longer than sporangia. Spores trilete, tetrahedral-globose, surface papillate.

Usually epiphytic on tree trunks in evergreen forests; 1300–1500 m. Xizang, Yunnan [Bhutan, NE India, N Myanmar, Nepal].

Antrophyum coriaceum sometimes approaches the narrow-leaved forms of A. callifolium Blume, except for the obvious stipes of the latter. Dried fronds are olive-green.

9. Antrophyum vittarioides Baker, J. Bot. 28: 267. 1890.

书带车前蕨  shu dai che qian jue

Antrophyum stenophyllum Baker, Kew Bull. 233, 1898, not Rovirosa 1909.

Rhizome short creeping, apex ascending or erect; scales pale-brown, subulate-lanceolate, 4–5 mm, 0.5–1 mm wide at base, margin dentate, base lacerate, clathrate; fronds clustered; stipes ca. 1 cm or shorter, appressed and winged, base slightly swollen, covered with scales like those on rhizome, verrucas after deciduous; laminae coriaceous, linear, 15–20 Χ 0.7–1 cm; costa obvious, infra-medial visible, evanescent upward; veins reticulate, forming 2 or 3 linear areolas. Soral lines linear, submarginal, one row on each side of costa, parallel to costa, or interrupted, slightly immersed in grooves, middle part fertile; paraphyses filiform, longer than sporangia. Spores trilete, tetrahedral-globose, surface ornamentation obscure.

Epiphytic by streams in dense forests; 300–1000 m. Guizhou, Yunnan [Vietnam].

Antrophyum vittarioides is close to A. brookei Hooker and A. subfalcatum Brack. from tropical Asian islands in the soral lines usually in one row near the margin on both sides of fronds. Superficially like Haplopteris fudzinoi (Makino) E. H. Crane.

19. HAPLOPTERIS C. Presl, Tent. Pterid. 141.1836.

书带蕨属  shu dai jue shu

Zhang Xianchun (张宪春); Michael G. Gilbert

Epiphytic or epilithic grasslike plants. Fronds simple, linear, entire, glabrous. Venation consisting of a costa and oblique unbranched veins, free except for a submarginal fertile connecting vein. Sori in one elongate marginal or submarginal groove on each side of lamina, immersed, rarely superficial; paraphyses long with a dark obconic head. Spores monolete, most ellipsoidal, sometimes fusiform, smooth, transparent.

About 40 unclear species: tropical and subtropical Asia; 13 species in China.

Most of the Old World species formally classified in Vittaria belong to Haplopteris as reflected in modern rbcL phylogenetic system.

1a.       Soral line marginal, immersed in groove, open outward.

2a.       Lamina up to 100 cm or longer, ca. 10 mm wide; stipe long, slender; scales dark brown  12. H. elongata

2b.       Lamina 8–30 cm, 2–4 mm wide; stipe short; scales yellow-brown
.....................................................................................................  13. H. anguste-elongata

1b.       Soral line submarginal.

3a.       Soral line superficial; lamina margins straight or revolute.

4a.       Scales dark brown, small, straight, areola wall thick; soral line submarginal, covered by revolute margin of lamina ...............................................................................................  1. H. amboinensis

4b.       Scales yellow-brown, large, soft, twisted, areola wall thin; soral line with a broad area between margins of lamina.

5a.       Lamina 2–3 mm wide; scales 3–7 Χ ca. 0.2 mm; soral line close to costa
..............................................................................................................  5. H. mediosora

5b.       Lamina over 6 mm wide; scales 10–20 Χ 0.5–2.5 mm; soral line with a broad area between costa.

6a.       Stipe short, thick; lamina 10–30 mm wide; scales 10–20 mm ...................  2. H. doniana

6b.       Stipe none, or slender; lamina 5–12 mm wide; scales ca. 10 mm.

7a.       Lamina 6–12 mm wide; soral line 1–1.5 mm away from margins of lamina  3. H. taeniophylla

7b.       Lamina 5–6 mm wide; soral line ca. 1 mm away from margins of lamina  4. H. himalayensis

3b.       Soral line immersed in groove between costa and margins of lamina; lamina margins revolute.

8a.       Lamina 5–10 mm wide, or more.

9a.       Costa not raised adaxially, lamina with a narrow concave line adaxially .  6. H. plurisulcata

9b.       Costa raised adaxially, lamina with a broad concave line on each side of costa adaxially  7. H. fudzinoi

8b.       Lamina (0.5–)2–4(–11) mm wide.

10a.     ............................................................................................................  8. H. linearifolia

10b.    

11a.     Scales dark brown, linear-lanceolate, 5–10 mm; lamina with a convex line close to margins            9. H. hainanensis

11b.     Scales yellow-brown, subulate-lanceolate or lanceolate, 2–3 mm; lamina without a convex line close to margins, only with a narrow concave line adaxially.

12a.     Costa flattened abaxially, wide; soral line covered by costa ............  10. H. sikkimensis

12b.     Costa raised abaxially, narrow; soral line with a broad area between costa
..........................................................................................................  11. H. flexuosa

1. Haplopteris amboinensis (Fιe) X. C. Zhang, Ann. Bot. Fennici 40: 460. 2003.

剑叶书带蕨  jian ye shu dai jue

Vittaria amboinensis Fιe, Mιm. Foug. 3: 14. 1852; V. ensata Christ, J. Bot. 21: 240. 1908; V. lauana Ching; V. chingii B. S. Wang.

Rhizome thick, long creeping, bearing roots with very numerous water-absorbing root hairs; scales dark brown, obscure iridescent, subulate-lanceolate, 3–5 mm, 0.5 mm wide at base, apex acuminate, margin prominent denticulate, apical areola not transparent, wall thick, obviously verrucate, dark colored; fronds clustered; stipes 2–4 mm distant, 4–10 cm, slender, appressed, base covered with scales; laminae hard papyraceous, or subcoriaceous, brown when dry, lanceolate, 20–40 cm Χ 10–25 mm, gradually narrowed to both ends, apex long acuminate, base long attenuated along stipe, margin slightly revolute when dry; costa ± invisible and narrowly concave adaxially, raised abaxially, flattened, veins evident. Soral line submarginal, superficial, or slightly immersed, infra-medial and apical part sterile; paraphyses many, long, with an obconic head, width ± half of length of head. Spores monolete, oblong in outline, surface papillate.

Epiphytic or epilithic in evergreen forests. Guangdong, Guangxi, Hainan, Yunnan [Cambodia, N India, Indonesia Laos, Malaysia, Myanmar, Thailand, Vietnam].

V. latifolia Ching, non Benedit.

2. Haplopteris doniana (Mett. ex Hieron.) E. H. Crane, Syst. Bot. 22: 514. 1997.

带状书带蕨  dai zhuang shu dai jue

Vittaria doniana Mett. ex Hieron., Hedwigia 57: 204. 1916; V. forrestiana Ching.

Rhizome thick, short creeping; scales yellow-brown, bright iridescent, soft, twisted, linear-lanceolate, 10–20 Χ 1–2.5 mm, lower margin subentire, upper part minutely denticulate, apex long acuminate, areola wall thin; fronds clustered; stipes thick, short, 1–2 cm; laminae thick coriaceous, shrink when dry, ribbonlike, 15–35 cm or longer, 10–30 mm wide, widest at middle or above, apex long attenuate or caudate, base long attenuate and winged on stipe, margin cartilaginous; costa thick, slightly raised adaxially, carinate abaxially. Soral line 1–2 mm to margin, superficial; paraphyses long, with an obconic head, length longer than width of head. Spores monolete, oblong in outline, surface rugate.

Epiphytic or epilithic; 1600–3300 m. Guangxi, Guizhou, Xizang, Yunnan [N India, N Myanmar].

Vittaria forrestiana Ching perhaps represents a thin fronded form of this species, which was found mainly from NW Yunnan. The report of it from Thailand, Indochina, and Japan are all misidentifications of Haplopteris amboinensis (Fιe ) X. C. Zhang (V. amboinensis Fιe).

3. Haplopteris taeniophylla (Copel.) E. H. Crane, Syst. Bot. 22: 514.1997.

广叶书带蕨  guang ye shu dai jue

Vittaria taeniophylla Copel., Philip. J. Sci. Suppl. 1: 157. 1906.

Rhizome short creeping; scales pale-brown, bright iridescent, lanceolate, ca. 10 Χ 0.5 mm; fronds clustered; stipes none; laminae subcoriaceous, ribbonlike, 30–60 cm or longer, 6–12 mm wide, widest at middle or above, apex acute, base long attenuate; costa distinct. Soral line 1–1.3 mm to margin, superficial or slightly immersed.

Epiphytic on base of tree trunk; 2000–2600 m. Taiwan [Philippines].

4. Haplopteris himalayensis (Ching) E. H. Crane, Syst. Bot. 22: 514. 1997.

喜马拉雅书带蕨  xi ma la ya shu dai jue

Vittaria himalayaensis Ching, Sinensia 1(12): 190, f. 5B: 1–2. 1931.

Rhizome short creeping, 3–5 mm thick; scales dark brown, bright iridescent, linear-lanceolate, ca. 10 Χ 1 mm, apex bristlelike, margin obviously denticulate; fronds clustered; stipes slender, short or long; laminae thin herbaceous, linear, 30–60 cm Χ 5–6 mm, wider usually near middle, narrowed gradually to both ends, base attenuate to very base; costa invisible adaxially, raised abaxially, flattened. Soral line ca. 1 mm to margin, superficial; paraphyses long, with an obconic head, length longer than width of head. Spores monolete, oblong in outline, surface papillate.

Epiphytic or epilithic; 1700–2900 m. S Xizang, NW and W Yunnan [Bhutan, N India, Nepal].

5. Haplopteris mediosora (Hayata) X. C. Zhang, Ann. Bot. Fennici 40: 460. 2003.

中囊书带蕨  zhong nang shu dai jue

Vittaria mediosora Hayata, Icon. Pl. Formosan. 5: 346, f. 149: g-i. 1915; V. stenophylla Copel.; V. tibetica Ching & S. K. Wu.

Rhizome short creeping or ascending; scales brown, bright iridescent, linear-lanceolate, 3–7 Χ ca. 0.2 mm, margin minutely denticulate, apex bristlelike; fronds clustered; stipes slender, ca. 1 mm thick, 4–5 cm; laminae herbaceous, linear, 10–25 cm Χ 2–3 mm, margin straight or slightly revolute, apex gradually narrowed, base attenuate; costa invisible adaxially, raised abaxially, flattened, slender. Soral line superficial, between costa and margin of lamina; paraphyses long, with an obconic head, length and width equal of head. Spores monolete, elliptic in outline, surface ornamentation obscure.

Epiphytic or epilithic; 2300–3500 m. W Sichuan, C Taiwan, Xizang, C Yunnan [E Himalaya, Philippines].

6. Haplopteris plurisulcata (Ching) X. C. Zhang, Ann. Bot. Fennici 40: 461. 2003.

曲鳞书带蕨  qu lin shu dai jue

Vittaria plurisulcata Ching, Sinensia 1: 186, pl. 4, f. 1–3. 1931.

Rhizome short creeping; scales light brown, lanceolate, 5–8 mm, ca. 1 mm wide at base, areola wall thin, soft, twisted, margin entire, apex bristlelike; fronds clustered; stipes short; laminae herbaceous, lanceolate, 30–40 cm Χ 5–8 mm, gradually narrowed to both ends, margin slightly revolute; costa invisible adaxially, slightly raised abaxially, slender. Soral line immersed in groove, close to revolute margin, ca. 2 mm to costa, lower 1/3 not fertile; paraphyses long, with an obconic head, width ± half of length of head. Spores monolete, elliptic in outline, surface ornamentation obscure.

Epiphytic or epilithic; 1800–2900 m. W Sichuan, C Taiwan, Xizang, C Yunnan [N Vietnam].

7. Haplopteris fudzinoi (Makino) E. H. Crane, Syst. Bot. 22: 514. 1997.

平肋书带蕨  pin le shu dai jue

Vittaria fudzinoi Makino, Bot. Mag. Tokyo 12: 28. 1898; V. centrochinensis Ching ex J. F. Cheng.

Rhizome short creeping, or ascending; scales yellow-brown, bright iridescent, soft, twisted, small ones ca. 5 Χ 1 mm, subulate-triangular, margin denticulate, large ones ca. 8 mm, 0.1–0.2 mm wide, linear-lanceolate, apex bristlelike, margin sub-entire; fronds clustered; stipes dark, 1–6 cm, or nearly absent; laminae thick coriaceous, linear or narrow ribbonlike, 15–55 cm Χ ca. 5 mm, sometimes up to 8–10 mm wide, narrowed gradually to both ends, base long attenuate; costa raised adaxially, adaxial surface with 2 parallel long grooves beside costa, also raised abaxially, broadened, flattened. Soral line close to margin, immersed in groove or ± superficial; paraphyses long, with an obconic head, length longer than width of head. Spores monolete, oblong in outline, surface with obscure papillate.

Epiphytic or epilithic; 1300–2800 m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Yunnan [Japan].

The broad frond form, with soral line ± superficial, scales more brown in color are found mainly from Zhejiang, Jiangxi, Hunan, Guangdong and Guangxi, which has been published as Vittaria centrochinensis J. F. Chen. It approaches the Philippine Vittaria taeniophylla Copel.

8. Haplopteris linearifolia (Ching) X. C. Zhang, Ann. Bot. Fennici 40: 460. 2003.

线叶书带蕨  xian ye shu dai jue

Vittaria linearifolia Ching, Sinensia 1(12): 183, pl. 1, f. 1–3. 1931.

Rhizome short creeping; scales light brown, bright iridescent, lanceolate, ca. 10 mm, ca. 1.5 mm wide at base, twisted, margin denticulate; fronds clustered; stipes shorter, slender, glabrous, light brown at base; laminae thick coriaceous, linear, 20–50 cm Χ 2–4 mm, margin strongly revolute; costa invisible adaxially, raised abaxially, flattened, very broad, up to half of width of lamina. Soral line deeply immersed in groove, very close to costa; paraphyses long, with an obconic head, length slightly shorter than width or equal of head. Spores monolete, oblong in outline, surface ornamentation obscure.

Epiphytic or epilithic; 1700–3400 m. SE Xizang, NW and W Yunnan [India (Assam), N Myanmar].

9. Haplopteris hainanensis (C. Chr. ex Ching) E. H. Crane, Syst. Bot. 22: 514. 1997.

海南书带蕨  hai nan shu dai jue

Vittaria hainanensis C. Chr. ex Ching, Sinensia 1(12): 182, pl. 1, f. 1–4. 1931.

Rhizome short creeping, ca. 2 mm thick; scales dark brown, bright iridescent, linear-lanceolate, 5–10 mm, ca. 0.5 mm wide at base, apex bristlelike, marginal one row areola thin walled, middle areola wall thick, dark colored; fronds clustered; stipes none; laminae very coriaceous, linear or lanceolate, 10–30 cm Χ 2–4 mm, rarely 5–11 mm wide, broad usually near middle, narrowed gradually to both ends, flattened to very base, margin strongly revolute; costa visible abaxially, sometimes adaxially, not raised; veins usually not evident. Soral line submarginal, immersed in deep groove, thus raised above; paraphyses long, with an obconic head, width and length of head ± equal. Spores monolete, oval in outline, surface obscure papillate.

On palm trees; 100–1000 m. Hainan, S Yunnan [N Vietnam].

Whether Haplopteris hainanensis is conspecific with H. ensiformis (Swartz) E. H. Crane or not is still a problem.

10. Haplopteris sikkimensis (Kuhn) E. H. Crane, Syst. Bot. 22: 514. 1997.

锡金书带蕨  xi jin shu dai jue

Vittaria sikkimensis Kuhn, Linnaea 36: 66. 1869.

Plants very small. Rhizome very slender, short creeping or ascending; scales pale-brown, bright iridescent, subulate-lanceolate, 2–4 Χ 0.25–0.5 mm, marginal one row areola thin walled, middle areola wall thick, dark colored; margin minutely denticulate, apex long bristlelike; fronds clustered; stipes short, slender, appressed; laminae thin herbaceous, linear, 1.5–12 cm Χ 0.5–1.5 mm, normally 4–6 cm Χ ca. 1 mm, margin slightly revolute, apex acute or rounded, base long attenuate; costa invisible adaxially, raised abaxially, wide, flattened. Soral line deeply immersed in groove, covered by costa, occupying area between costa and margin when mature; paraphyses long, with an obconic head, length longer than width of head. Spores monolete, elliptic in outline, surface sparsely papillate.

Epiphytic or epilithic in evergreen broad-leaved forests, mixed with Leucobryum; 1400–2200 m. SE Xizang, S, SE, and W Yunnan [N India, Myanmar, N Thailand, N Vietnam].

11. Haplopteris flexuosa (Fιe) E. H. Crane, Syst. Bot. 22: 514. 1997.

书带蕨  shu dai jue

Vittaria flexuosa Fιe, Mιm. Foug. 3: 16. 1852; V. caricina Christ; V. filipes Christ; V. modesta Hand.-Mazz.; V. nana Ching; V. ophiopogonoides Ching; V. costularis Ching.

Rhizome short creeping, fronds very close, scales deciduous; rhizome scales yellow-brown, bright, iridescent, subulate-lanceolate, (2–)5–6 mm, 0.2–0.5 mm wide at base, apex bristlelike, margin denticulate, areola wall thick, dark colored; fronds clustered; stipes slender, short, base pale-brown, with small fibrous scales; laminae herbaceous, linear, 15–40 cm or longer, 4–6 mm wide, small ones 6–12 cm Χ 1–2.5 mm, narrowed gradually to both ends, base long attenuate; costa narrowly concave adaxially, raised abaxially, slender, veins invisible, margin of lamina revolute and partly cover sori. Soral line submarginal, immersed in groove, with a broad area between costa, but in narrow-frond plants line occupying whole area when mature, fertile medially; paraphyses many, long, with an obconic head, length and width ± equal of head. Spores monolete, oblong in outline, surface with obscure papillate.

Epiphytic or epilithic; 100–3200 m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, Cambodia, India (Sikkim), Japan, Korea, Laos, Myanmar, Nepal, Thailand, Vietnam].

The narrow-frond form, with soral line occupying area between costa, growing in crevices of rocks in dry conditions or on wet cliffs of caves, are forms called Vittaria caricina Christ, V. nana Ching, and V. modesta Hand.-Mazz.; the epiphytic long-frond form from humid evergreen monsoon forests in E Himalaya is called V. ophiopogonoides Ching; while intermediate forms named as V. filipes Christ or V. costularia Ching before. This species is widespread in E Asia very variable in size but constant in scales and paraphyses characters, margin of lamina always revolute to cover partly the soral line.

12. Haplopteris elongata (Sw.) E. H. Crane, Syst. Bot. 22: 514. 1997.

唇边书带蕨  chun bian shu dai jue

Vittaria elongata Sw., Syn. Fil. 109, 302. 1806; V. pauciariolata Ching.

Rhizome rather long, creeping, much branched, bearing roots with very numerous water-absorbing root hairs; scales dark brown, bright iridescent, subulate-lanceolate, 4–5 mm, 0.5–1 mm wide at base, apex bristlelike, margin prominent denticulate, areola wall thick, dark colored; fronds clustered, drooping; stipes ca. 0.5 mm distant; laminae subcoriaceous, linear or ribbon like, up to 100 cm Χ 5–20 mm, apex rounded or obtuse, base gradually narrowed; costa slender, not prominent, veins evident. Soral line marginal, immersed in deep groove, open outward, fertile throughout; paraphyses many, long, with an obconic head, length longer than width of head. Spores monolete, narrow oblong in outline, surface ornamentation obscure.

Epiphytic or epilithic; 100–1400 m. Fujian, Guangdong, Guangxi, Hainan, SE Xizang, S Yunnan [Indochina, Indonesia, S Japan, Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand; Australia, Madagascar].

Haplopteris elongata always grows with other epiphytic ferns, notably with Pseudodrynaria coronans (Wallich ex Mettenius) Ching, and species of the Asplenium nidus and Asplenium laserpitifolium groups. It is a polymorphic species, with different size, and color of scales changes in different habitat. All Chinese specimens identified as Vittaria zosterifolia (Borry) Willdenow and V. merrillii Christ belong to this species.

13. Haplopteris anguste-elongata (Hayata) E. H. Crane, Syst. Bot. 22: 514. 1997.

姬书带蕨  ji shu dai jue

Vittaria anguste-elongata Hayata, Icon. Pl. Formosan. 6: 161. 1916.

Rhizome slender, long creeping, ascending; scales yellow-brown, bright iridescent, linear-lanceolate, 5–7 mm, 0.2–0.3 mm wide at base, apex bristlelike, often with a glandular like head, margin subentire, margin areola wall thin, dark brown; fronds clustered; stipes indistinct, slender, appressed, glabrous; laminae thin herbaceous, brown when dry, linear, 8–30 cm Χ 2–4 mm, gradually narrowed to both ends, apex short acuminate or caudate; costa slender, slightly raised above, indistinct below, veins not evident. Soral line marginal, immersed in deep groove, open outward, fertile throughout; paraphyses many, long, with an obconic head, width ± half of length of head. Spores monolete, oblong in outline, surface ornamentation obscure.

Epiphytic or epilithic. Fujian, Hainan, Taiwan [Japan (Ryukyu Islands), Philippines].

20. MONOGRAMMA Commerson ex Schkuhr, Krypt. Gewδchse 1: 82. 1809.

一条线蕨属  yi tiao xian jue shu

Zhang Xianchun (张宪春); Michael G. Gilbert

Vaginularia Fιe, Congr. Sci. France, Dix. Sess. 1: 178. 1843.

Fronds very small, simple, linear, entire, glabrous. Venation consisting of a costa and at most a few almost parallel fertile veins. Sori born in a single groove along costa or on fertile veins, continuous or interrupted; paraphyses without a head, short. Spores trilete, smooth, transparent.

About nine species: tropical Asia to Australia, and the Pacific islands; two species in China (Hainan and Taiwan).

Vaginularia and Monogramma are both diminutive derived plants that resemble each other and can not be distinguished properly.

1a.       Sorus 1 per frond, on groove of costa ............................................................  1. M. paradoxa

1b.       Sori 1 or 2 per frond, on lateral veins ............................................................  2. M. trichoidea

1. Monogramma paradoxa (Fιe) Bedd., Ferns Brit. India Suppl. 24. 1876.

连孢一条线蕨  lian bao yi tiao xian jue

Pleurogramma ? paradoxa Fιe, Mιm. Foug. 3: 38. 1852.

Small grass like plants, fronds filiform. Rhizome less than 1 mm thick, creeping, stipes ca. 1 mm apart. Scales dark brown, clathrate, lanceolate, 0.8–1.25 Χ 0.25–0.5 mm, margin denticulate; fronds clustered; laminae 3–12 cm Χ 0.5–1 mm, filiform, apex acute, base long attenuate, without lateral vein. Sorus only one in groove of costa, elongate, not interrupt, covered by raised margins of costa, fertile medially, swollen when mature; paraphyses filiform, without a head. Spores trilete, surface ornamentation obscure.

Very rare, epiphytic or epilithic; 700–1400 m. Taiwan [Indonesia, Philippines, Sri Lanka, Thailand; Pacific islands (Micronesia, Polynesia)].

2. Monogramma trichoidea (Fιe) Hook., Sp. Fil. 5: 123. 1864.

针叶蕨  zhen ye jue

Vaginularia trichoidea Fιe, Mιm. Foug. 3: 34. 1852.

Small grass like plants, fronds filiform. Rhizome less than 1 mm thick, creeping, stipes ca. 1 mm apart. Scales brown, subulate-lanceolate, ca. 0.5 mm, clathrate; fronds clustered; laminae filiform, 5–12 cm, ca. 0.5 mm wide at sterile part; costa throughout, with 1 or 2 lateral veins. Sorus 1 or 2 per frond, on lateral veins, covered by costa and outer side of raised lateral vein; paraphyses without a head, many. Spores trilete, surface ornamentation obscure.

Epilithic on shaded wet rocks in dense forests in valley; 700–1400 m. Hainan, Taiwan [Indonesia (Sumatra), Malaysia, Philippines, S Thailand].


[1] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

[2] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

[3] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

[4] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

[5] The Nature Conservancy, Eugene, Oregon 97402, U.S.A.

[6] Missouri Botanical Garden, c/o Herbarium, Library, Art and Archives, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom.

[7] Department of Botany, National Museum of Nature and Science, Tsukuba 305-0005, Japan.

[8] Herbarium (ALA), University of Alaska Museum of the North, University of Alaska Fairbanks, Fairbanks, Alaska 99775, U.S.A.

[9] Herbario “Maria Eneyda P.K. Fidalgo,” Instituto de Botanica, Caixa Postal 4005, Sao Paulo, 01000 Sao Paulo, Brazil.

[10] Herbarium, University of Colorado Museum, Campus Box 218, Boulder, CO 80302, U.S.A.[University of Hawaii per IH!]