GLEICHENIACEAE [Draft: 10 Jul 2012]

里白科 li bai ke

Ding Bingyang (丁炳扬), Jin Xiaofeng (金孝锋); Kunio Iwatsuki

Ferns, terrestrial, often forming thickets. Rhizomes long creeping, protostelic, with scales or multicellular hairs. Fronds monomorphic, evergreen, erect, scrambling, or climbing, vernation circinate; stipe not articulate, distant, cylindrical, apex forked, with dormant bud; apical bud covered with hairs or scales; rachis simple to several times pseudodichotomously branched [or pinnate], with stellate hairs and/or ciliate-like scales when young, these persistent or glabrescent; pinnules pectinately pinnatisect; lobes elliptic or lanceolate, apex obtuse or acute; veins free, forked; lamina papery or subleathery, frequently glaucous abaxially. Sori in 1(–3) lines both sides of lobe costules, orbicular, exindusiate; sporangia 2–10[–16], sessile, turbinate, with a mid-transverse annulus; spores tetrahedral or reniform, without perispore, smooth.

Three to five genera depending on concepts of authors and over 150 species: tropical and subtropical areas; three genera and 16 species (seven endemic) in China.

Most species in the Gleicheniaceae are characteristic of very mineral poor soils.

Ching Ren-chang, Fu Shu-hsia, Wang Chu-hao & Shing Gung-hsia. 1959. Gleicheniaceae. In: Ching Ren-chang, ed., Fl. Reipubl. Popularis Sin. 2: 116–132, 346–350.

1a.       Veins 2- or 3-forked with (3 or )4–6 veinlets in each group; rhizome with hairs only; fronds dichotomously divided with a pair of lateral pinnules at most dichotomies; sporangia 5–18 per sorus  2. Dicranopteris

1b.       Veins 1-forked with 2 veinlets in each group; rhizome with scales; fronds without lateral pinnules at dichotomies; sporangia 2–5(–6) per sorus.

2a.       Rachis undivided, producing successive pairs of 2- or 3-pinnatifid pinnae; penultimate axes of frond without lamina; spores tetrahedral. ..........................................................  1. Diplopterygium

2b.       Rachis pseudodichotomous; penultimate (and often lower) axes with well-developed lamina; spores ellipsoid ............................................................................................................  3. Sticherus

1. Diplopterygium (Diels) Nakai, Bull. Natl. Sci. Mus., Tokyo 29: 47. 1950.

里白属 li bai shu

Jin Xiaofeng (金孝锋), Ding Bingyang (丁炳扬); Kunio Iwatsuki

Gleichenia sect. Diplopterygium Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 353. 1900; G. subg. Diplopterygium (Diels) Holttum.

Rhizomes long creeping, branched, with protosteles, covered with scales; scales brown, lanceolate, margin entire or ciliate. Fronds distant; rachis undivided, producing 1 to several successive pairs of pinnae; stipe and rachis with lanceolate scales and stellate hairs when young, these glabrescent or persistent; dormant apical bud with dense brown scales, outer bracts pinnatifid, leaflike; primary pinnae opposite, 2(–3)-pinnatifid, spreading or drooping; ultimate pinnules many, obliquely or horizontally spreading, deeply pectinately pinnatifid to costa, lanceolate in outline, sessile or shortly petiolate at base, apex acuminate; lobes linear to narrowly oblong-lanceolate, margin entire and slightly reflexed, apex emarginate; veins once forked with a pair in each group, spreading to lobe margins. Sori in single lines either side of costule; sporangia 2–4; spore tetrahedral, without perispore.

About 20 species: tropical and subtropical areas, mainly distributed in tropical Asia; nine species (four endemic) in China.

The FRPS account of the Gleicheniaceae treated all species of Diplopterygium as members of the genus Hicriopteris, but Holttum (Reinwardtia 4: 261. 1957) showed that the type of Hicriopteris belonged in Dicranopteris.

One of us (Iwatsuki) considers that Diplopterygium, as treated here, is split into too many species.

1a.       Pinnules with a short stipe at base, stipe 3–4 mm ...........................................  1. D. blotianum

1b.       Pinnules sessile.

2a.       Costae and costules with dense scales.

3a.       Lamina papery; stipes, costae, and costules with brown scales and stellate hairs; pinnules 22–24 mm wide; sori and veins bare .............................................................................  2. D. chinense

3b.       Lamina thinly leathery; stipes, costae, and costules with dense rufous scales and stellate hairs; pinnules 10–12 mm wide; sori and veins ± densely covered with rufous hairs .................  3. D. rufum

2b.       Costae and costules without scales, sometimes with sparse stellate hairs.

4a.       Lobes obliquely spreading.

5a.       Fronds 3-pinnate; pinnules irregular, entire, pinnatifid, or pinnatisect ........  4. D. irregulare

5b.       Fronds 2-pinnate; pinnules regular, pinnatifid or pinnatisect.

6a.       Pinnules pinnatisect, 15–25 mm wide, glossy; lobes 25–40 pairs
                                                                                                          5. D. laevissimum

6b.       Pinnules pinnatifid, 30–40 mm wide; lobes 60–70 pairs ......................  6. D. cantonense

4b.       Lobes horizontally or subhorizontally spreading.

7a.       Pinnae and pinnules smaller; pinnules 11–17 Χ 1.5–2.5 cm; veins 10 or 11 pairs on abaxial lobe surface .....................................................................................................  7. D. glaucum

7b.       Pinnae and pinnules larger; pinnules 15–22 Χ 2.4–4 cm; veins more than 15 pairs on abaxial lobe surface.

8a.       Lobes thinly leathery; pinnules glaucous abaxially .................................  8. D. giganteum

8b.       Lobes thickly membranous; pinnules green abaxially .............................  9. D. maximum

1. Diplopterygium blotianum (C. Christensen) Nakai, Bull. Natl. Sci. Mus., Tokyo 29: 49. 1950.

阔片里白 kuo pian li bai

Gleichenia blotiana C. Christensen, Bull. Mus. Natl. Hist. Nat., sιr. 2, 6: 103. 1934; Hicriopteris blotiana (C. Christensen) Ching; H. tamdaoensis Ching & P. S. Chiu.

Plants 2–3 m tall. Fronds 2-pinnate; pinna rachises stramineous, 2–4 mm wide; pinnae 2-pinnatifid, lamina 60–140 Χ 20–30 cm; pinnules many, alternate, deeply pectinately pinnatifid, lanceolate or narrowly lanceolate, 10–25 Χ 2–4 cm, base attenuate and with a 3–4 mm stipe, apex acuminate; lobes 20–40 pairs, linear-lanceolate to broadly lanceolate, 12–22 Χ 3–4 mm, apex emarginate; lamina thickly papery, glabrous adaxially, costae, costules, and veins abaxially with sparse brown stellate hairs; costae flat adaxially; veins prominent or slightly prominent on both surfaces. Sori brown, with 4 or 5 sporangia.

600–1400 m (in Taiwan). S Guangdong (Yangshan), S Guangxi, Hainan, C Taiwan, SE Yunnan (Pingbian) [Cambodia, Laos, Vietnam].

2. Diplopterygium chinense (Rosenstock) De Vol, Fl. Taiwan 1: 92. 1975 [“chinensis”].

中华里白 zhong hua li bai

Gleichenia chinensis Rosenstock, Repert. Spec. Nov. Regni Veg. 13: 120. 1913; Hicriopteris chinensis (Rosenstock) Ching.

Plants ca. 3 m tall. Rhizomes long creeping, ca. 5 mm in diam., densely covered with brown scales. Fronds 2-pinnate; stipe dark brown, 5–6 mm wide, with dense rufous scales, glabrescent, pinna rachises stramineous, 2–4 mm wide; pinnae 2-pinnatifid, lamina oblong, 80–120 Χ 20–30 cm; pinnules many, alternate, deeply pectinately pinnatifid, lanceolate, 12–18 Χ 2.5–3 cm, base sessile, apex acuminate; lobes alternate, 40–60 pairs, narrowly lanceolate to lanceolate, 10–15 Χ 2.5–3.5 mm, apex obtuse or sometimes emarginate; lamina papery, costae, costules, and veins with dense brown stellate hairs abaxially; costae plane adaxially; veins prominent on both surfaces. Sori with 3 or 4 sporangia.

Dense forests, near forests or valleys; 100–2800 m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Taiwan, Xizang (Chayu), Yunnan, Zhejiang [N Vietnam].

Two collections (Qinghai-Tibet Exped. 73-805), which have pinna costae with sparse brown scales, were provisionally labeled as “Diplopterygium tsayuensis Ching & S. K. Wu.” Here they are regarded as an ecotype of D. chinense.

3. Diplopterygium rufum (Ching) Ching ex X. C. Zhang, Novon 14: 150. 2004.

厚毛里白 hou mao li bai

Hicriopteris rufa Ching, Fl. Reipubl. Popularis Sin. 2: 347. 1959; Diplopterygium rufopilosum (Ching & P. S. Chiu) Ching ex X. C. Zhang; H. rufopilosa Ching & P. S. Chiu.

Plants 0.5–2 m tall. Fronds 2-pinnate; pinna rachises with dense rufous stellate hairs and ciliate scales; pinnae 2-pinnatifid, lamina oblong, 30–40 Χ 14–17 cm, base attenuate, sessile, apex acuminate; pinnules 30–35 on each side, alternate, deeply pectinately pinnatifid to costa, narrowly lanceolate, ca. 10 Χ 1–1.3 cm, apex acuminate; lobes alternate, 20–35 on each side, broadly lanceolate, 5–6 Χ 2–2.2 mm, apex obtuse; lamina thinly leathery, costae, costules, and veins with dense rufous stellate hairs abaxially; costae depressed adaxially; veins obscure or slightly prominent abaxially. Sori covered with rufous hairs, obscure or slightly prominent abaxially; sporangia 3 or 4.

* Thickets, mountaintops; 2000–2400 m. Yunnan.

4. Diplopterygium irregulare W. M. Chu & Z. R. He, Acta Bot. Yunnan. 22: 255. 2000.

参差里白 can cha li bai

Plants 1–1.8 m tall. Fronds 3-pinnate; primary pinnae opposite, oblong, 60–100 Χ 50–70 cm; secondary pinnae alternate, 30–40 on each side, pinnatisect, narrowly oblong, 10–40 Χ 1.5–15 cm, base attenuate, apex caudate-acuminate; ultimate pinnules irregularly entire, pinnatifid, or pinnatisect, linear or narrowly lanceolate, 10–65 Χ 2–3 cm; lobes alternate, 20–35 on each side, obliquely spreading, lanceolate, 2–7 Χ ca. 1 mm; lamina thinly leathery, greenish gray abaxially, glabrous; costae flat adaxially; veins prominent on both surfaces. Sori with 2–4 sporangia.

* Margins of evergreen broad-leaved forests; ca. 1600 m. S. Yunnan (Xichou).

Diplopterygium irregulare is a distinctive species, which is close to D. laevissimum but differs by having tripinnate fronds with irregular pinnules.

5. Diplopterygium laevissimum (Christ) Nakai, Bull. Natl. Sci. Mus., Tokyo 29: 52. 1950.

光里白 guang li bai

Gleichenia laevissima Christ, Bull. Acad. Int. Gιogr. Bot. 11: 268. 1902; G. kiusiana Makino; Hicriopteris laevissima (Christ) Ching.

Plants 1–1.5 m tall. Rhizomes creeping, dark brown, cylindric, covered with brown scales. Fronds 2-pinnate; pinnae opposite, ovate-oblong, 30–50 Χ 20–26 cm; pinnules alternate, 20–30 on each side, pinnatisect, narrowly lanceolate, 12–18 Χ 1.5–2.5 cm, base attenuate, apex caudate-acuminate; lobes alternate, 25–40 on each side, obliquely spreading, lanceolate, 7–13 Χ ca. 2 mm, apex acute; lamina thinly leathery, greenish gray abaxially, glabrous; costae flat adaxially; veins prominent on both surfaces. Sori orbicular, exindusiate, with 4 or 5 sporangia.

Forests, mountain slopes, forest margins; 500–2500 m. Anhui, Fujian, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Philippines, Vietnam].

6. Diplopterygium cantonense (Ching) Nakai, Bull. Natl. Sci. Mus., Tokyo 29: 49. 1950.

广东里白 guang dong li bai

Gleichenia cantonensis Ching, Lingnan Sci. J. 15: 391. 1936; Hicriopteris cantonensis (Ching) Ching.

Plants ca. 3 m tall. Rhizomes creeping. Fronds 2-pinnate; pinnae opposite, oblong, 40–70 Χ 30–40 cm; pinnules alternate or subopposite, 30–40 on each side, deeply pinnatisect, linear-lanceolate, 25–30 Χ 3–4 cm, base broadly cuneate, apex long acuminate; lobes alternate, 60–70 on each side, obliquely spreading, linear, 15–20 Χ ca. 2 mm, apex obtuse; lamina thickly papery, glaucous abaxially, glabrous; costae flat adaxially; veins obscure on both surfaces. Sori with 2–4 sporangia.

* Guangdong (Huangpu, Shenzhen, Xinyi), Hainan, S Jiangxi (Longnan).

7. Diplopterygium glaucum (Thunberg ex Houttuyn) Nakai, Bull. Natl. Sci. Mus., Tokyo 29: 51. 1950.

里白 li bai

Polypodium glaucum Thunberg ex Houttuyn, Nat. Hist. 14: 177. 1783; Gleichenia glauca (Thunberg ex Houttuyn) Hooker[(1844), not Swartz (1806)??]; G. japonica Sprengel; Hicriopteris glauca (Thunberg ex Houttuyn) Ching; H. remota Ching.

Plants ca. 1.5 m tall. Rhizomes creeping, ca. 3 mm in diam., covered with brown scales. Fronds 2-pinnate; pinnae opposite, oblong, 60–90 Χ 18–28 cm, base slightly attenuate, apex acuminate; pinnules alternate or subopposite, 25–35 on each side, pinnatifid, linear-lanceolate to narrowly lanceolate, 11–16 Χ 1.2–2.4 cm, base truncate, apex acuminate; lobes alternate, 20–35 on each side, plane[horizontally spreading?], lanceolate, 7–12 Χ 2–3 mm, rarely lowest pairs reflexed to rachises, apex obtuse; lamina glaucous abaxially, green adaxially, papery, glabrous, costae and costules with sparse stellate hairs, glabrescent; costae flat adaxially; veins prominent on both surfaces. Sori with 3 or 4 sporangia.

Thickets, mountain slopes, forests, forest margins; 300–2300 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [India, Japan].

8. Diplopterygium giganteum (Wallich ex Hooker & Bauer) Nakai, Bull. Natl. Sci. Mus., Tokyo 29: 50. 1950.

大里白 da li bai

Gleichenia gigantea Wallich ex Hooker & Bauer, Gen. Fil. t. 39. 1840; Diplopterygium criticum (Ching & P. S. Chiu) Ching ex X. C. Zhang; D. simulans (Ching) Ching ex X. C. Zhang; D. yunnanense (Ching) Ching ex X. C. Zhang; Hicriopteris critica Ching & P. S. Chiu; H. gigantea (Wallich ex Hooker & Bauer) Ching; H. glaucoides Ching; H. omeiensis Ching & P. S. Chiu; H. reflexa Ching & P. S. Chiu; H. simulans Ching; H. yunnanensis Ching.

Plants 2–3 m tall. Rhizomes creeping, cylindrical, ca. 4 mm in diam., covered with rufous lanceolate scales. Fronds 2-pinnate; pinnae opposite, oblong, 90–120 Χ 27–40 cm, base slightly attenuate, apex acuminate; pinnules many, alternate or subopposite, pinnatifid, linear-lanceolate to lanceolate, 12–21 Χ 1.5–3.5 cm, base slightly attenuate, apex acuminate; lobes alternate, 30–50(–70) on each side, plane[horizontally spreading?], narrowly lanceolate to lanceolate, rarely linear, 10–15(–20) Χ 2.5–3.5 mm, apex obtuse; lamina papery, glaucous or gray-green abaxially, glabrous, costae and veins with sparse stellate hairs; costae plane or depressed adaxially; veins prominent on both surfaces. Sori with 3–5 sporangia.

Forest margins, mountain slopes, moist places; 800–2800 m. Hainan (Anding), Hubei (Lichuan), Sichuan (Emei Shan), Xizang (Chayu), Yunnan [Bhutan, India, Myanmar, Nepal].

9. Diplopterygium maximum (Ching) Ching & H. S. Kung, Fl. Sichuan. 6: 134. 1988.

绿里白 lό li bai

Hicriopteris maxima Ching, Fl. Reipubl. Popularis Sin. 2: 348. 1959.

Plants ca. 3 m tall. Fronds 2-pinnate; stipe brown, ca. 1 m Χ 5–7 mm; pinnae elliptic, 90–100 Χ 45–50 cm, base slightly attenuate, apex acuminate; pinnules many, plane, alternate or subopposite, deeply pinnatifid, narrowly lanceolate, 20–25 Χ 2.5–4 cm, apex long acuminate; lobes alternate, 50–70 on each side, plane[horizontally spreading?], linear or lanceolate, 15–20 Χ 3–3.5 mm, apex obtuse or sometimes emarginate; lamina submembranous, costae and veins abaxially with sparse stellate hairs; costae depressed adaxially; veins prominent on both surfaces. Sori with 3 sporangia.

* Under rocky cliffs; 1200–1400 m. Sichuan (Emei Shan).

2. Dicranopteris Bernhardi, Neues J. Bot. 1(2): 38. 1805.

芒萁属 mang qi shu

Ding Bingyang (丁炳扬), Jin Xiaofeng (金孝锋); Kunio Iwatsuki

Rhizomes slender, long creeping, covered with multicellular hairs. Fronds distant, erect or somewhat scandent; rachis often several times dichotomously branched with dormant buds in dichotomies, with normal pinnules only at apex, each dichotomy usually with a pair of lateral pinnae, these shallowly to more usually deeply pectinately pinnatifid; dormant apical buds covered with stiff hairs, usually subtended by stipulelike bracts; ultimate pinnae deeply pectinately pinnatifid, lanceolate or broadly lanceolate, sessile; lobes plane[horizontally spreading?], margin entire or rarely crenate, often narrowly revolute; most veins 2- or 3-forked with (3 or)4–6 veinlets. Sori orbicular, exindusiate, in 1(–3) line(s) on each side of costule; sporangia 6–18, sessile; spores tetrahedral [or ellipsoid], without perispore, smooth.

About ten species: mainly in tropical and subtropical areas; six species (three endemic) in China.

1a.       Rachis with 1(or 2) dichotomy, without lateral pinnae; sporangia 12–18 per sorus ....................              1. D. splendida

1b.       Rachis 1–8 Χ dichotomous[“times dichotomously branched” (and “Χ dichotomously branched”) was used throughout but has been mostly changed under this genus only to “Χ dichotomous.” OK to stet original?], most dichotomies with a pair of lateral pinnae; sporangia 5–8(–15) per sorus.

2a.       Sori in 2 or 3 irregular lines on each side of costule; ultimate pinna lobes 8–10 mm wide  2. D. ampla

2b.       Sori in 1 line on each side of costule; ultimate pinna lobes 2–6 mm wide.

3a.       Ultimate pinna lobes broader, 4–6 mm wide; apical buds with subtending bracts fugacious, so buds often apparently ebracteate ....................................................................  3. D. taiwanensis

3b.       Ultimate pinna lobes narrower, 2–4.5 mm wide; apical buds with subtending bracts more persistent.

4a.       Frond rachis 5–8 Χ dichotomously branched, without lateral pinnules at first dichotomy; ultimate pinnule lobes 15–40 on each side, lanceolate or linear-lanceolate, 10–20 Χ 2–3 mm  6. D. linearis

4b.       Frond rachises 1–3 Χ dichotomously branched, with lateral pinnules at every dichotomy; ultimate pinnule lobes 35–80 on each side, linear-lanceolate, 15–80 Χ 3–4.5 mm.

5a.       Plants up to 4 m tall; ultimate pinnae 35–40 Χ 9–15 cm; lobes 3–8 cm
                                                                                                                4. D. gigantea

5b.       Plants ca. 1 m tall; ultimate pinnae 15–25(–35) Χ 4–8(–10) cm; lobes 1.5–5 cm  5. D. pedata

1. Dicranopteris splendida (Handel-Mazzetti) Tagawa, Acta Phytotax. Geobot. 8: 164. 1939.

大羽芒萁 da yu mang qi

Gleichenia splendida Handel-Mazzetti, Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 61: 81. 1924.

Plants 0.7–1 m tall. Rhizomes creeping, ca. 2 mm in diam., covered with brown hairs. Stipe brown, cylindrical, 20–30 cm, ca. 2 mm wide, glabrous; rachis 1(or 2) Χ dichotomous (apical buds degenerate); apical bud ovate, covered with dense brown hairs; bracts ovate, margin crenate; rachis without lateral pinnae at dichotomy. Ultimate pinnae oblong-lanceolate, 15–25 Χ 5–7 cm, apex attenuate; lobes 35–50 on each side, lanceolate, 30–40 Χ 6–8 mm, margin entire, apex obtuse or emarginate; lamina papery, glaucous abaxially, dark green adaxially, glabrous; costae prominent on both surfaces; veins 4 or 5 in each group. Sori in 1 line on each side of costule; sporangia 12–18.

Forests, forest margins; 2000–2800 m. NW Yunnan (Qiujiang) [Myanmar].

Images of Dicranopteris splendida from Thailand (http://rbg-web2.rbge.org.uk/thaiferns/factsheets/index.php?q=Dicranopteris_splendida.xml, accessed 4 Jul. 2012) do not match this species.

2. Dicranopteris ampla Ching & P. S. Chiu, Acta Phytotax. Sin. 8: 161. 1959.

大芒萁 da mang qi

Plants 1–2 m tall. Rhizomes creeping, 2–4 mm in diam., woody, covered with brown multicellular hairs. Stipe dark brown, ca. 80 cm, 3.5–5 mm wide, glabrous; rachis 3 or 4 Χ dichotomous; apical buds ovate; bracts ovate, 17–20 mm, margin irregularly toothed, apex obtuse; rachis with a pair of lateral pinnae at each dichotomy; lateral pinnae deeply pectinately pinnatifid, oblong-lanceolate, 15–22 Χ 4.5–13 cm; ultimate pinnae lanceolate or oblong, 20–40 Χ 8–15 cm, base asymmetrically attenuate, apex acuminate, not subtended by lateral pinnae; lobes lanceolate or linear-lanceolate, 5–10 Χ 0.8–1 cm, with a pair of lateral stipulelike pinnules at base, margin entire or crenate and sometimes pinnatifid, apex obtuse or emarginate; pinnules linear, 6–10 Χ 0.7–1 cm, margin undulate or crenate; lamina thinly leathery, gray-green abaxially, dark green adaxially, glabrous; costae prominent adaxially; veins 5–7 in each group. Sori in 1–3 irregular lines on each side of costule; sporangia 7–15.

* Thickets, forest understories; 700–1900 m. SW Guangdong, W Guangxi, S Hainan, S Xizang, S Yunnan.

Collections of Dicranopteris ampla with the lobes crenate, sometimes pinnatifid, were recognized as a form (D. ampla f. irregularis B. Y. Ding & X. F. Jin, Acta Bot. Yunnan. 30: 9. 2008). Ching (Icon. Filic. Sin. 4: t. 153. 1937) initially identified material of D. ampla as D. splendida Handel Mazzetti.

3. Dicranopteris taiwanensis Ching & P. S. Chiu, Fl. Reipubl. Popularis Sin. 2: 346. 1959.

台湾芒萁 tai wan mang qi

Dicranopteris linearis (N. L. Burman) Underwood var. montana Holttum.

Plants ca. 1 m tall. Rhizomes not seen. Stipe dark brown, ca. 35 cm, ca. 3 mm wide, glabrous; rachis 2 or more Χ dichotomous; apical buds ovate, small, covered with brown hairs, subtending bracts small and fugacious, so buds often apparently ebracteate; rachis with a pair of lateral pinnae at each dichotomy; first pair of lateral pinnae obliquely spreading, broadly lanceolate, 7–9 Χ 3–4 cm, base and apex attenuate; second pair smaller, ca. 4 Χ 1.5 cm; ultimate pinnae lanceolate or oblong-lanceolate, 15–25 Χ 4–7 cm, base attenuate, apex acuminate, subtended by lateral pinnae; lobes 25–40 on each side, lanceolate or linear-lanceolate, 3–3.5 Χ 0.4–0.6 cm, margin entire, apex obtuse or emarginate; lamina papery, glaucous abaxially, green adaxially, glabrous; costae prominent abaxially; veins 4 or 5 in each group. Sori in 1 line on each side of costule; sporangia 6–8.

Broad-leaved forests, forest margins; 1000–1700 m. Taiwan [India, Indonesia, Malaysia, Sri Lanka].

Dicranopteris taiwanensis was reported as the only species in Dicranopteris that has apical buds covered only by hairs, not by bracts. Further field studies (R. Knapp, pers. comm.) have shown that there are bracts though these are small and fugacious. This removes the main distinction from D. linearis var. montana and extends the distribution outside Taiwan.

4. Dicranopteris gigantea Ching, Fl. Reipubl. Popularis Sin. 2: 346. 1959.

乔芒萁 qiao mang qi

Plants 3–4 m tall. Rhizomes not seen. Stipes dark brown, cylindrical, 1–2 m, ca. 5 mm wide, glabrous; rachis 2 or 3 Χ dichotomous; apical buds ovate, ca. 2 mm, covered with dense brown hairs; bracts ovate-oblong, 6–10 mm, margin entire or rarely crenate, apex obtuse; rachis with a pair of lateral pinnules at each dichotomy; lateral pinnules oblong, 17–25 Χ 6–7 cm; lobes 30–40 on each side; ultimate pinnae lanceolate, 35–40 Χ 9–15 cm, base slightly attenuate, apex long caudate; lobes 50–60 on each side, linear, 3–8 Χ 0.4–0.45 cm, margin entire, apex obtuse, bilobate, or emarginate; lamina papery, bluish green abaxially, green adaxially, glabrous; costae prominent adaxially; veins inconspicuous, 4 or 5 in each group. Sori in 1 line on each side of costule; sporangia 5–8.

* Forests; 100–300 m. Hainan, Yunnan (Hekou, Mengzi).

5. Dicranopteris pedata (Houttuyn) Nakaike, Enum. Pterid. Jap. 114. 1975.

芒萁 mang qi

Polypodium pedatum Houttuyn, Nat. Hist. 14: 174. 1783; Dicranopteris dichotoma (Thunberg) Bernhardi; Gleichenia dichotoma (Thunberg) Hooker; Mertensia dichotoma (Thunberg) Willdenow; P. dichotomum Thunberg (1784), not Houttuyn (1783).

Plants 0.5–2(–3) m tall. Rhizomes creeping, ca. 2 mm in diam., covered with dense dark brown hairs. Stipe stramineous, 0.3–1(–2) m, 2–5 mm wide, glabrous; rachis 1–3 Χ dichotomous, basal internode ca. 10 cm, covered with dark brown hairs, glabrescent, second internode 3–5 cm; apical buds ca. 2 mm, covered with dense brown hairs; bracts ovate to ovate-oblong, 5–10 mm, margin irregularly crenate, rarely entire; rachises with a pair of lateral stipulelike pinnae at each dichotomy; lower lateral pinnules broadly lanceolate, 10–25 Χ 4–7 cm, upper ones smaller, 4–10 Χ 2–4 cm; ultimate pinnae lanceolate or broadly lanceolate, 15–25(–35) Χ 4–8(–10) cm, base attenuate, apex caudate; lobes 35–50 on each side, linear-lanceolate, 15–50 Χ 3–4 mm, margin entire, apex emarginate or obtuse; lamina papery, glaucous abaxially, yellowish green adaxially, with sparse brown hairs on costae and veins abaxially; costae prominent on both surfaces; veins 3–5 in each group. Sori in 1 line on each side of costule; sporangia 5–8.

Thickets, forests, hillsides; below 100–2200 m. Anhui, Fujian, Gansu (Wenxian), Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangsu, Jiangxi, Shanxi, Sichuan, ?Taiwan, Yunnan, Zhejiang [India, Japan, Vietnam].

One of us (Iwatsuki) cannot distinguish Dicranopteris pedata from D. linearis with any clear discriminative characters. The Japanese form is easily included in the range of variable forms of so-called D. linearis, and no infraspecific taxa are recognized under that variable species in the present treatment.

6. Dicranopteris linearis (N. L. Burman) Underwood, Bull. Torrey Bot. Club 34: 250. 1907.

铁芒萁 tie mang qi

Polypodium lineare N. L. Burman, Fl. Indica, 235. 1768; Dicranopteris linearis var. subpectinata (Christ) Holttum; D. linearis var. tetraphylla (Rosenstock) Nakai; D. subpectinata (Christ) C. M. Kuo; D. tetraphylla (Rosenstock) C. M. Kuo; Gleichenia linearis (N. L. Burman) C. B. Clarke; G. linearis var. tetraphylla Rosenstock; G. subpectinata Christ.

Plants 1–1.5 m tall. Rhizomes creeping, ca. 3 mm in diam., covered with brown multicellular hairs. Stipe dark brown, ca. 60 cm, ca. 6 mm wide, when young with brown hairs at base, glabrescent; rachis 5–8 Χ dichotomous, basal internode 13–18 cm, ca. 3.5 mm wide, others shorter, ultimate 3.5–6 Χ ca. 0.1 cm, fluted; apical buds ovate, covered with dense brown hairs; bracts ovate, with triangular lobes at margin; rachises with a pair of lateral pinnae at each dichotomy except first; lateral pinnae spreading obliquely downward; ultimate pinnae 5.5–15 Χ 2.5–4 cm, subtended or not by lateral pinnae; lobes 15–40 pairs, lanceolate or linear-lanceolate, 10–20 Χ 2–3 mm, margin entire, apex obtuse; lamina papery, glaucous abaxially, green adaxially, glabrous or rarely with sparse stellate hairs abaxially; costae prominent abaxially; veins 3 in each group. Sori in 1 line on each side of costule; sporangia 5–7.

Valleys, by rivers, hillsides; below 700–1900 m. S Guangdong, S Guizhou, Hainan (Hongmao Shan), Taiwan, SE Yunnan [India, Indonesia, Malaysia, Nepal, Singapore, Sri Lanka, Thailand, Vietnam; Australia].

Dicranopteris linearis is widely distributed in tropical areas of S Asia and is sometimes an invasive weed. It is very variable, and Holttum (Fl. Males., ser. 2, 1: 33–36. 1959) recognized thirteen varieties, four of which have been reported in Taiwan (D. linearis var. linearis, D. linearis var. montana (= D. taiwanensis), D. linearis var. subpectinata, and D. linearis var. tetraphylla). These were separated by the presence or absence of hairs on the pinnules, rachis dichotomies equal or unequal, presence or absence of lateral pinnae subtending the ultimate pinnae, and the dimensions of the pinnae (Fl. Taiwan 1: 92–94. 1994); however, we believe that these vary too continuously to justify the recognition of distinct taxa.

3. Sticherus C. Presl, Tent. Pterid. 51. 1836.

假芒萁属 jia mang qi shu

Ding Bingyang (丁炳扬), Jin Xiaofeng (金孝锋); Kunio Iwatsuki

Gleichenia subg. Mertensia (Willdenow) Hooker; Mertensia Willdenow (1804), not Roth (1797).

Rhizomes creeping, covered with scales. Fronds distant, erect; lamina often repeatedly pseudo­dichotomously divided, without lateral pinnules at dichotomies, upper axes pinnulelike with rows of lobes along each side; lamina gray-green or glaucous abaxially; veins once forked. Sori orbicular, exindusiate, in 1 line on each side of costule; sporangia 4 or 5(or 6), sessile; spores 2-sided or reniform, monolete.

About 100 species: tropical areas, mainly distributed in South America; one species in China.

Some authorities (e.g., http://www.botanyvn.com/cnt.asp?param=edir&v=Gleichenia%20truncata&list=species&lg=en) follow Holttum (Fl. Males., Ser. II, Pterid. 1: 20, f. 7a–b & 9. 1959, accessed 4 Jul. 2012) who included Sticherus (and Diplopterygium) within Gleichenia and Sticherus laevigatus within S. truncatus.

1. Sticherus truncatus (Willdenow) Nakai, Bull. Natl. Sci. Mus. 29: 20. 1950.

假芒萁 jia mang qi

Mertensia truncata Willdenow, Kongl. Vetensk. Acad. Nya Handl. 25: 169, t. 5, f. A. 1804; Gleichenia laevigata (Willdenow) Hooker; G. truncata (Willdenow) Sprengel; M. laevigata Willdenow; Sticherus laevigatus (Willdenow) C. Presl.

Plants 1–2 m tall. Rhizomes long creeping, covered with peltate scales. Stipe sprawling or scandent, 0.8–1.6 m; lamina several times dichotomous, all but basal axes pinnulelike with 2 rows of lobes; apical buds covered with brown ciliate scales and 2-pinnate bracts; ultimate pinnae deeply pectinately pinnatifid, broadly lanceolate, 12–18 Χ 2.5–5 cm; lobes plane[horizontally spreading?], opposite, 10–30 Χ 2–3 mm, base expanded, margin entire, apex obtuse or emarginate; lamina papery, with sparse brown stellate hairs and scales when young, glabrescent; costa prominent on both surfaces. Sori with 4 or 5 sporangia.

Hainan, SE Yunnan (Hekou, Jinping) [Cambodia, Malaysia, Sri Lanka, Thailand, Vietnam].