BLECHNACEAE [Draft]

乌毛蕨科  wu mao jue ke

Wang Faguo (王发国)[1], Xing Fuwu (邢福武)1; Masahiro Kato[2]

Ferns, mostly terrestrial, sometimes tree ferns (like small trees). Rhizome mostly erect, or creeping or scandent, dictyostelic, with brown, entire scales. Fronds uniform or dimorphic, mostly long-stipitate, stipe with 2 to several bundles, scaly at base; lamina pinnate, pinnatifid, or bipinnatifid, rarely simple, thickly papery to leathery, glabrous or usually small-scaly; veins free or anastomosing with one or more series of areoles, without included free veinlets. Sori elongate or continuous along vascular network or commissure on either side of midrib; indusium facing toward costa or midrib, rarely exindusiate, annulus longitudinal, interrupted; spores elliptic, bilateral, monolete.

A family of 13 genera worldwide, with ca. 240 species; seven genera and 12 species in China [??#endemic].

A natural generic classification of the family should be based on molecular phylogenetic relationships and comparative morphology because published and unpublished molecular data do not support the conventional classifications (e.g., Fl. Reipubl. Popularis Sin. 4(2). 1999; the system adopted here).

1a.       Tree ferns with trunklike stem, up to ca. 1 m tall.

2a.       Stem 1–2 cm in diam.; lamina bipinnatifid; sori indusiate, discrete
................................................................................................................  7. Diploblechnum

2b.       Stem 10–15 cm in diam.; lamina pinnate; sori exindusiate, covering pinna surface ..  3. Brainea

1b.       Rhizome creeping to suberect; plants not like tree ferns.

3a.       Sori discrete.

4a.       Rhizome creeping; stipes distant; pinnae accrete, conjoined by narrow wing along rachis ..  5. Chieniopteris

4b.       Rhizome short and erect; stipes tufted; pinnae separate ..............................  4. Woodwardia

3b.       Sori continuous.

5a.       Plants epiphytic or lithophytic; veins anastomosing ......................................  2. Blechnidium

5b.       Plants terrestrial; veins free.

6a.       Lamina monomorphic, lamina marginal sori prominent ..................................  1. Blechnum

6b.       Lamina dimorphic, lamina marginal sori much reduced ...........................  6. Struthiopteris

1. BLECHNUM Linnaeus, Sp. Pl. 2: 1077. 1753.

乌毛蕨属  wu mao jue shu

Plants terrestrial, moderate-sized or large. Rhizome stout, usually erect or ascending, dictyostelic, densely scaly; scales dark brown, glossy, lanceolate, entire, basally attached. Fronds clustered, long-stipitate; stipe robust, scaly at base, above naked; lamina pinnate, with many reduced auricular pinnae, leathery; pinnae linear, margin entire or serrulate; veins free, parallel, simple or forked. Sori long, linear, adjacent and parallel to costa, coenosori; indusium attached to commissure, facing toward costa; annulus longitudinal, interrupted, of 14–28 cells. Spores elliptic, usually smooth, with perispore.

About 35 species: pantropical; one species in China.

In an unpublished molecular phylogeny, Blechnum orientale is not included in the genus Struthiopteris s. str. represented by B. occidentale (type) and relatives.

1. Blechnum orientale Linnaeus, Sp. Pl. 2: 1077. 1753.

乌毛蕨  wu mao jue

Rhizome erect, dark brown, short, densely covered with scales; scales dark brown at center, brown near margin, narrowly linear, ca. 1 cm, entire. Stipes tufted, 10–60 cm, 3–10 mm in diam., base dark brown and covered with scales as rhizome; lamina imparipinnate, monomorphic, ovate-lanceolate, 55–100 Χ 20–60 cm, subleathery; pinnae numerous, close, alternate; lower ones contracted to small round auricles, 0.3–1 cm; upper ones oblique, distant, linear or linear-lanceolate, 10–30 Χ 0.8–1.8 cm, base sessile, rounded or subtruncate, or adnate, decurrent to rachis on basiscopic side, gradually narrowed to acuminate apex, terminal pinna similar to middle pinnae; veins free, parallel, simple or forked near costa, close. 2n = 66.

Exposed shrubby or low hillsides; 200–1000 m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Japan; tropical Asia, Australia, Pacific islands].

1a. Apex of lateral pinnae entire or slightly undulate .................................................  1a. var. orientale

1b. Apex of lateral pinnae cristate ..........................................................................  1b. var. cristatum

1a. Blechnum orientale var. orientale

乌毛蕨(原变种)  wu mao jue (yuan bian zhong)

Blechnopsis orientalis (Linnaeus) C. Presl.

Apex of lateral pinnae entire or slightly undulate.

Exposed shrubby or low hillsides; 200–1000 m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Japan; tropical Asia, Australia, Pacific islands].

1b. Blechnum orientale var. cristatum[??cristata] J. Smith in Seemann, Bot. Beechey Voy. 1: 427. 1857. [??Cult. Ferns]

冠羽乌毛蕨  guan yu wu mao jue

A monstrous form with cristate apex of lateral pinnae.

* Guangdong.

2. BLECHNIDIUM T. Moore, Ferns Great Brit. Nat. Printed oct. ed. 2. 210. 1860. [??Octavo Nat.-Print. Brit. Ferns]

乌木蕨属  wu mu jue shu

Plants epiphytic, small to moderate-sized. Rhizome creeping, dark brown, densely covered with lanceolate or ovate-lanceolate, rufous scales. Fronds distant, long-stipitate; stipe dark brown, grooved above, sparsely scaly at base; lamina brownish green when dry, lanceolate, subleathery, both surfaces glabrous, pectinate; pinnae adnate, oblong, margin entire, apex subacute or obtuse; lower few pairs of pinnae shortened; veins anastomosing with 1–3 rows of areoles. Sori linear, adjacent and parallel to costa; indusium attached to commissure, facing toward costa. Spores elliptic, with perispore.

One species in India, Myanmar, and China.

This monotypic genus is the most closely related to Struthiopteris and isolated from the rest.

1. Blechnidium melanopus (Hooker) T. Moore, Brit. Ferns Nat. Pr. octavo ed., 2. 210. 1860.

乌木蕨  wu mu jue

Blechnum melanopus Hooker, Sp. Fil. 3: 64. 1860[??1859]; Blechnidium plagiogyriifrons Hayata; Blechnum plagiogyriifrons Hayata.

Rhizome dark brown, long-creeping, ca. 3 mm in diam., densely scaly; scales rufous, lanceolate to ovate-lanceolate, ca. 5 mm, membranous, subentire, acuminate or fibriform. Fronds distant, long-stipitate; stipe dark brown, 12–25 cm, ca. 1 mm in diam., grooved above, sparsely scaly at base; lamina brownish green when dry, lanceolate, 15–25 Χ 3–7 cm, subleathery, both surfaces glabrous, gradually narrowed to base, pectinate, acuminate; pinnae 15–25 pairs, adnate; middle ones oblong, usually subfalcate, 1.5–5 Χ 0.5–1 cm, entire, subacute or obtuse; lower pinnae shortened, 0.4–1.3 cm; veins inconspicuous, anastomosing with 1–3 rows of hexagonal areoles. Sori linear, adjacent and parallel to costa; indusium attached to commissure, linear, facing toward costa. Spores elliptic, with perispore.

Epiphytic on tree trunks or rocks in forests; 1600–2800 m. Taiwan, Yunnan [India, Myanmar].

3. BRAINEA J. Smith, Cat. Ferns Gard. Kew 5. 1856.

苏铁蕨属  su tie jue shu

Tree ferns, terrestrial. Rhizome[??Stem] erect, blackish brown, stout, up to ca. 1 m tall, 10–15 cm in diam., woody, apex scaly; scales rufous, linear, up to 3 cm, membranous, acuminate. Fronds clustered in terminal crown, slightly dimorphic; stipe brownish stramineous, 10–30 cm, basal part scaly; lamina pinnate, elliptic-lanceolate, 50–100 cm, slightly narrowed downward, leathery, abaxially with some small scales along costa and veins; pinnae 30–55 pairs, opposite or alternate, subsessile, linear to narrowly oblong; middle pinnae 10–15 Χ 0.7–1.2 cm, base asymmetrical, subauriculate, margin serrulate; basal pinnae slightly shortened; veins free, simple or 1- or 2-forked, except for costal row of areolae nearly triangular; fertile pinnae somewhat shorter, margin sometimes irregularly lobed; rachis stramineous, grooved above. Sori borne on costal veins, exindusiate, abundant, covering whole abaxial surface of pinnae when mature. 2n = 66.

One species: widely distributed in tropical Asia.

The monotypic genus Brainea is isolated from the rest.

1. Brainea insignis (Hooker) J. Smith, Cat. Kew Ferns 5. 1856.

苏铁蕨  su tie jue

Browningia insignis Hooker, J. Bot. 5: 237. 1853; Brainea formosana Hayata; B. insignis var. formosana (Hayata) Tagawa.

Characters for this species are the same as in the genus.

Moist[??wet or damp] and exposed hillsides; 300–1700 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan, Yunnan [tropical regions of Asia].

4. WOODWARDIA J. E. Smith, Mιm. Acad. Sci. Turin 5: 411. 1793.

狗脊属  gou ji shu

Plants terrestrial, of moderate to large size. Rhizome erect or ascending, sometimes creeping, stout, dictyostelic, densely scaly; scales brown, non-clathrate, basally attached. Stipes tufted; lamina deeply bipinnatifid, elliptic, papery or subleathery, long-stipitate; pinnae narrowly oblong, gradually narrowed to narrowly deltoid-ovate, sometimes stalked, firm, glabrous, margin entire or serrulate; veins anastomosing with a series of areoles along costae and costules, free to margin, simple or forked. Sori occupying costular areoles, discrete, linear or elliptic, attached to outer veins forming areoles, superficial or sunken; indusia facing toward costa, dark brown, thickly papery; sporangia with long stalk, annulus of 17–24 thickened cells. Spores elliptic, perispore rugose.

About 10 species: distributed in temperate to tropical areas in Asia, Europe, and America [??North and South]; five species in China.

1a.       Rachis with large subterminal bulbils on rachis ............................................  1. W. unigemmata

1b.       Rachis without such bulbils.

2a.       Base of lower pinnae asymmetrical, 1–3 basiscopic lobes lacking; sori sunken, oblong.

3a.       Lamina 35–45 cm; 1 basiscopic basal pinna-lobe lacking; lobes acute or acuminate  2. W. orientalis

3b.       Lamina 30–120 cm; 1–3 basiscopic basal pinna-lobes lacking; lobes acuminate or caudate  3. W. prolifera

2b.       Base of lower pinnae symmetrical, basal pairs of lobes contracted or not; sori not sunken, linear.

4a.       Basal basiscopic lobe of lower pinnae contracted, apex obtuse ...................  4. W. japonica

4b.       Basal basiscopic lobe of lower pinnae slightly contracted, lanceolate, apex acute  5. W. magnifica

1. Woodwardia unigemmata (Makino) Nakai, Bot. Mag. (Tokyo) 39: 103. 1925.

顶芽狗脊  ding ya gou ji

Woodwardia radicans (Linnaeus) Smith var. unigemmata Makino, J. Jap. Bot. 2(2): 7. 1918; W. himalaica Ching & S. K. Wu; W. latiloba Ching & P. S. Chiu; W. maxima Ching; W. yunnanensis Ching & P. S. Chiu.

Rhizome decumbent, dark brown, stout, up to 3 cm in diam., densely scaly; scales brown, lanceolate, 2.2–2.7 cm, membranous, entire, long acuminate. Stipes tufted, 30–100 cm, 5–8 mm in diam., base densely scaly; upper part of stipe and rachis with fewer brown, fibriform scales; lamina deeply bipinnatifid, elliptic or ovate-lanceolate, 30–100 Χ 20–60 cm, leathery; pinnae 7–15 pairs, subsessile or shortly stalked, broadly lanceolate; middle pinnae 15–35 Χ 4–12 cm, base rounded-truncate, unequal, caudate, cut 3/4 way to costa; lobes 14–20 pairs, oblique, close, oblong-lanceolate, sometimes falcate, lower pairs of lobes slightly shortened, middle lobes 1–6 cm Χ 8–12 mm at base, margin serrulate with sharp, long teeth. Veins anastomosing with 2 or 3 rows of areoles along costae and costules, marginally free, simple or forked. Bulbils single or multiple, large, scaly, subterminal below rachis tip at base of upper pinna. Sori occupying costular areoles, oblong, sunken in rimmed depressions; indusia attached to outer vein, dark brown, thickly membranous. 2n = 68.

In shrubs by roadsides, forests; 400–3000 m. Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Shanxi, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, India, Japan, Myanmar, Nepal, Philippines, Vietnam].

2. Woodwardia orientalis Swartz in Schrader, J. Bot. 1800(2): 76. 1801.

东方狗脊  dong fang gou ji

Rhizome decumbent, dark brown, stout, densely scaly; scales dark brown, lanceolate, 1–4 cm, entire, membranous, apex fibriform. Stipes close, 20–55 cm, 3–6 mm in diam., base densely scaly; upper part of stipe and rachis sparsely covered with brown, broadly lanceolate scales; lamina deeply bipinnatifid, brown or slightly green when dry, ovate, 35–45(–70) Χ 15–45 cm, leathery, glabrous, base rounded-truncate, apex acuminate; pinnae 6–8 pairs, shortly stalked, lanceolate; lower and middle pinnae 10–30 Χ 4–9 cm, base asymmetrical with 1 basiscopic lobe lacking, acuminate, deeply pinnatifid to 2–3(–4) mm from costa, basal pinnae shortened; lobes 10–18 pairs, close, oblique, oblong, 3–5.5(–7) cm Χ 8–10(–13) mm, usually broadest at base, margin cuspidately serrulate, apex acute or acuminate; veins obvious, anastomosing to form one row of areoles along costae and 2 or 3 rows of discrete, polygonal areoles, remainder free to margin, simple or forked. Small bulblets borne on adaxial surfaces of pinna-lobes or not. Sori occupying costular areoles, crescent-shaped or elliptic, sunken in rimmed depressions; indusia dark brown, thickly papery. 2n = 136.

Roadsides, mountain slopes; ca. 500 m. Anhui, Fujian, Guangdong, Guangxi, Hunan, Jiangxi, Taiwan, Zhejiang [Japan, Philippines].

3. Woodwardia prolifera Hooker & Arnott, Bot. Beech. Voy. 275. 1841 [1838].

珠芽狗脊  zhu ya gou ji

Woodwardia angustiloba Hance; W. exaltata Nakai; W. orientalis Swartz var. formosana Rosenstock; W. orientalis var. prolifera (Hooker & Arnott) Ching; W. radicans (Linnaeus) Sm.[??author name] var. prolifera [??basionym] C. Christensen.

Rhizome decumbent, dark brown, stout, densely scaly; scales red-brown, lanceolate, 2–4 cm, membranous, entire or with a few teeth, fibriform. Stipes close, 30–110 cm, 3–15 mm in diam., base densely scaly; upper part of stipe and rachis sparsely covered with brown, broadly lanceolate scales, surface rough with scale scars; lamina deeply bipinnatifid, brown or slightly green when dry, oblong-ovate or elliptic, 35–120 Χ 30–40 cm, leathery, glabrous, acuminate; pinnae 5–9(–13) pairs, shortly stalked, lanceolate; lower and middle pinnae 10–30 Χ 4–9 cm, base asymmetrical with 1–3 basiscopic lobes lacking, deeply pinnatifid to 1–2 mm from costa, long-acuminate or caudate; lobes 10–14(–24) pairs, close or separate, oblique, oblong-lanceolate, 3–7(–9) cm Χ 5–9 mm, slightly narrowed to base, margin cuspidately serrulate, occasionally lobed, long acuminate or caudate; veins obvious, anastomosing with one row of areoles along costae and 2 or 3 rows of discrete, polygonal areoles, free distally, simple or forked. Leaf-bearing bulblets small and usually abundant on adaxial surfaces of pinna-lobes. Sori occupying costular areoles, crescent-shaped or elliptic, sunken in rimmed depressions; indusia dark brown, thickly papery. 2n = 68.

Mountain slopes, light[??sunny/open] and wet places in sparse forests, near streams; 100–1100 m. Anhui, Fujian, Guangdong, Guangxi, Hunan, Jiangxi, Taiwan, Zhejiang [Japan].

Herbarium and field observations of Woodwardia prolifera show that there is ecological variation in the frequency of bulblets on the lamina; some mature fronds are not bulbiferous.

4. Woodwardia japonica (Linnaeus f.) Sm.[??author name], Mιm. Acad. Sci. Turin 5: 411. 1793.

狗脊  gou ji

Blechnum japonicum Linnaeus f., Supp. Syst. Veg. 447[??445]. 1781 [1782]; Woodwardia affinis Ching & P. S. Chiu; W. intermedia H. Christ; W. japonica var. contigua Ching & P. S. Chiu; W. omeiensis Ching & P. S. Chiu[??only Ching]; W. rubricaulis Ching & H. S. Kung [??not in Tropicos and IPNI].

Rhizome decumbent, dark brown, stout, 3–5 cm in diam., densely scaly; scales dark brown, lanceolate or linear-lanceolate, ca. 1.5 cm, membranous, entire, acuminate, sometimes fibriform. Stipes close, 15–70 cm, 5–8 mm in diam., base densely scaly; upper part of stipe and rachis sparsely covered with brown, fibriform scales; lamina bipinnatifid, elliptic or lanceolate, 25–85 Χ 18–45 cm, leathery, acuminate; pinnae 7–15 pairs, subsessile or shortly stalked, broadly lanceolate; middle pinnae 12–25 Χ 2–4 cm, base acroscopically truncate, basiscopically rounded, cut 1/2 way to costa, apex acuminate; lobes 11–16 pairs, close, oblique, hemielliptic or hemiovate, lowest pair shortened, basiscopic lobe rounded, ovate or auriculate, 5–10 mm, apex rounded; middle lobes 1.3–2.2 Χ 0.7–1 cm, serrulate, acute; veins anastomosing with 2 or 3 rows of areoles along costae and costules, distally free, simple or forked. Sori occupying costular areoles, discrete, linear, sunken; indusia brown. 2n = 68.

Ridges, exposed slopes, shaded forests; 300–1500 m. Widely distributed south of the Yangtze River [Japan, Korea, Vietnam].

Woodwardia japonica is variable in the length, number, and arrangement of the pinnae, and the shape of the pinna lobes.

5. Woodwardia magnifica Ching & P. S. Chiu, Acta Phytotax. Sin. 12: 247. 1974.

滇南狗脊  dian nan gou ji

Rhizome decumbent, dark brown, stout, densely scaly; scales brown or dark brown, lanceolate, ca. 1.5 cm, membranous, entire, apex fibriform. Stipes close, brown-stramineous, 35–76 cm, 6–10 mm in diam., base densely scaly, sparsely scaly upward; lamina bipinnatifid, brown or brown-green when dry, oblong-ovate, 60–95 Χ 30–65 cm, leathery, glabrous, long-acuminate; pinnae 13–15 pairs, alternate or lower ones opposite, oblique, subsessile or shortly stalked, basal pair slightly shortened; middle ones linear-lanceolate, 30–35 Χ 4–8 cm, base rounded-truncate or rounded-cuneate, pinnatifid, caudate-acuminate; lobes 20–22 pairs, alternate, lower ones slightly shortened; middle one longest, lanceolate or narrowly triangular-lanceolate, 2.6–5 cm, base 1–1.5 cm wide, serrulate, acute; veins obvious, anastomosing with one row of areoles along rachis, costae, and costules, distally simple or forked, ending in submarginal fusiform hydathodes; rachis or costae densely covered with brown, fibriform, deciduous scales. Sori occupying costular areoles, linear, discrete; indusia dark brown.

Near forests, roadsides; 1400–1600 m. Yunnan [Vietnam].

5. CHIENIOPTERIS Ching, Acta Phytotax. Sinica 9: 37. 1964.

祟澍蕨属  chong shu jue shu

Plants terrestrial, of small to moderate size. Rhizome long-creeping, dark brown, apex covered with brown, lanceolate scales. Fronds distant, long-stipitate; lamina simple, trifid, or deeply pinnatifid, thickly papery or subleathery, glabrous; pinnae (or lobes) 1–5 pairs, lanceolate, acuminate, base slightly narrowed, terminal pinnae, if any, similar to but longer than lateral ones; veins anastomosing with 3 or 4 rows of areoles, marginal veins free. Sori oblong or linear, borne along costal and costular areolae; indusia facing toward costa, dark brown, linear-oblong, papery. Spores elliptic, perispore rugose, granulate.

Two species: distributed in China, south to Vietnam and east to Japan.

Chieniopteris is treated as a section of the genus Woodwardia in Cranfill and Kato (2003, in S. Chandra and M. Srivastava (eds.), Pteridology in the New Millennium, pp. 25–48, Kluwer).

1a.       Lamina simple or imparipinnate with 1–4 pairs of lateral pinnae and longer terminal pinna  1. C. harlandii

1b.       Lamina pinnatifid, with pinnatifid apex, lateral pinnae 5–7 pairs ± lobed ................  2. C. kempii

1. Chieniopteris harlandii (Hooker) Ching, Acta Phytotax. Sin. 9: 39. 1964.

祟澍蕨  chong shu jue

Woodwardia harlandii Hooker, Fil. Exot. t. 7. 1857; Lorinseria harlandii (Hooker) J. Smith.

Rhizome long-creeping, dark brown, 4–6 mm in diam., densely scaly; scales brown, lanceolate, 4–6 mm, membranous, margin entire or with few hairlike teeth. Fronds distant, subdimorphic with fertile lobes narrower than sterile lobes, 30–120 cm; stipe 12–80 cm, base dark brown, scaly; lamina simple, ternate, or deeply pinnatifid, gray-green or brown when dry, thickly papery or subleathery, glabrous; pinnae (or lobes) 1–4 pairs, opposite, 4–5 cm apart, lanceolate, acuminate, base adnate to rachis and decurrent, margin entire or undulate, usually reflexed when dry, basal pinnae 20–29 Χ 2–3 cm, upper pinnae shortened, terminal pinna similar to but longer and broader than lateral ones, rachis wing often very narrow or lacking between basal pinnae; veins anastomosing with 1 row of costal areoles, 2 or 3 rows of hexagonal areoles, marginal veins free. Sori linear, interrupted, 10–22 mm, borne along costal and costular areoles; indusia rufous when mature, papery.

In valleys, moist[??damp or wet] forests; 400–1300 m. Fujian, Guangdong, Guangxi, Hainan, S Hunan, Taiwan [Japan, Vietnam].

2. Chieniopteris kempii (Copeland) Ching, Acta Phytotax. Sin. 9: 39. 1964.

裂羽祟澍蕨  lie yu chong shu jue

Woodwardia kempii Copeland, Philipp. J. Sci. 3: 280. 1908; W. harlandii Hooker var. takeoi (Hayata) Masamune; W. heteropinnata B. S. Wang; W. takeoi Hayata.

Rhizome long-creeping, dark brown, 4–7 mm in diam., densely scaly; scales brown, lanceolate, 4–5 mm, membranous, entire, apex acuminate. Fronds distant, subdimorphic, fertile fronds larger and more deeply dissected than sterile fronds; stipe 30–70 cm, base dark brown, densely scaly, upward sparsely scaly and glabrous, brown-stramineous; lamina pinnatifid, brown when dry, deltoid-ovate, 13–26 Χ 11–28 cm, nearly leathery, glabrous, base rounded-truncate, apex acuminate; sterile lamina pinnatifid, basal lobes largest, lanceolate, 6.5–10 Χ 1–2.5 cm, base slightly narrowed, united with broad wing, margin serrulate, apex acuminate; fertile lamina bipinnatifid; pinnae 5–7 pairs, opposite, oblique or basal pair nearly spreading, pinnatifid; basal pair largest, elliptic-lanceolate, 11–20 Χ 4.5–10 cm, base narrowed, sessile or shortly stalked, ca. 3 cm apart from above, pinnatipartite to broad wing of costa, apex acuminate; middle lobes lanceolate or linear, 8–9 Χ 1.2–1.5 cm, margin irregularly undulate or lobed, entire or with irregular and distant sera, basal lobes much shortened or rounded-auriculate; veins invisible but midvein prominent on both surfaces, anastomosing with 1 row of long, narrow areoles along costa, secondary veins anastomosing with 2 rows of areoles, free near margin. Sori linear, 5–17 mm, borne in costal, rachis, or midvein areoles; indusium dark brown when mature, papery.

Forests, moist[??damp or wet] forests. Fujian, Guangdong, Guangxi, Taiwan [Japan].

6. STRUTHIOPTERIS Scopoli, Meth. Pl. 25. 1754.

荚囊蕨属  jia lang jue shu

Lithophytes. Rhizome erect or ascending, short, scaly; scales brown, linear or lanceolate, entire. Fronds tufted, monomorphic; lamina pectinate-pinnatisect, oblanceolate, narrowed downward, leathery; pinnae spreading, oblong-subdeltoid, base decurrent to rachis wing; veins invisible, free, furcate, basal ones always trifurcate, not reaching laminar margin. Sori linear, one on either side of costa; indusium facing toward costa, thinly papery, inrolling young sporangia, opening when mature; spores elliptic, perispore corrugated. 2n = 62, 68.

About 10 species: mainly distributed in temperate areas of the northern hemisphere and Australia; two species in China (one endemic).

1a.       Sterile lamina 2–5 cm wide; stipe 3–24 cm ........................................................  1. S. eburnea

1b.       Sterile lamina 5–8 cm wide; stipe almost absent ...............................................  2. S. hancockii

1. Struthiopteris eburnea (H. Christ) Ching, Sunyatsenia 5: 243. 1940.

荚囊蕨  jia lang jue

Rhizome erect and short, or ascending and long, densely scaly; scales brown, lanceolate, 5–6 mm, thickly membranous, margin entire or with few teeth, apex fibriform. Fronds tufted, dimorphic; stipe stramineous, almost absent or 3–24 cm, 1–2 mm in diam., base scaly, glabrous upward; lamina pectinate-pinnatisect, dark green or brown when dry, linear-oblanceolate, 15–45 Χ 2–5(–8) cm above middle, thickly leathery, glabrous; pinnae 20–40 pairs, spreading, lower ones gradually shortened, basal pair small auriculate; upper pinnae oblong, 1.5–3 Χ ca. 0.5 cm, base adnate to rachis, entire, slight narrowed to apex, slightly reflexed, spreading, close or distant; veins invisible, pinnate, furcate, not reaching laminar margin; rachis stramineous, glabrous, shallowly grooved on upper side. Fertile lamina as long as but narrower than sterile lamina. Sori linear, from near costa to near laminar margin, one on either side of costa. 2n = 66*.

* On rocks, limestone; 500–1800 m. Anhui, Fujian, Guangxi, Guizhou, Hubei, Hunan, Sichuan, Taiwan.

1a. Lamina to 5 cm wide; pinnae acute .....................................................................  1a. var. eburnea

1b. Lamina to 3 cm wide; pinnae obtuse or rounded ...................................................  1b. var. obtusa

1a. Struthiopteris eburnea var. eburnea

荚囊蕨(原变种)  jia lang jue (yuan bian zhong)

Blechnum eburneum H. Christ, Bull. Acad. Int. Gιogr. Bot. 11: 233. 1902; Lomaria eburnea (H. Christ) Ching; Spicanthopsis eburnea (H. Christ) Tagawa.

Lamina to 5 cm wide; pinnae acute.

* On rock or limestone; 500–1800 m. Anhui, Fujian, Guangxi, Guizhou, Hubei, Hunan, Sichuan.

1b. Struthiopteris eburnea var. obtusa (Tagawa) Tagawa, Acta Phytotax. Geobot. 14: 192. 1952. [??not in Tropicos or IPNI]

天長羅蔓蕨  pinyin

Spicantopsis eburnea var. obtusa Tagawa, Acta Phytotax. Geobot. 9: 88. 1940.

Lamina to 3 cm wide; pinnae obtuse or rounded.

* On limestone at middle elevations of central mountains. Taiwan.

2. Struthiopteris hancockii (Hance) Tagawa, Acta Phytotax. Geobot. 14: 192. 1952.

宽叶荚囊蕨  kuan ye jia lang jue

Blechnum hancockii Hance, J. Bot. 21: 267. 1883; Lomaria apodophylla Baker; Spicantopsis hancockii Masamune[??not in Tropicos or IPNI]; S. nipponica var. hancockii Nakai[??not in Tropicos or IPNI].

Rhizome erect, short, densely scaly; scales dark brown, linear, ca. 1.5 cm, membranous, entire, apex subulate. Fronds tufted, subdimorphic with fertile lamina narrower than sterile lamina, subsessile or shortly stipitate; lamina pectinate-pinnatisect, brown when dry, elliptic or oblanceolate, broadest at or above middle, 30–40 Χ 5–8 cm, thickly leathery, glabrous, base gradually narrowed with small auricles, apex acuminate or acute; pinnae 20–40 pairs, spreading, close or distant, linear-oblong, slightly falcate, largest 2.5–4 Χ ca. 0.5 cm, in fertile fronds 1.5–3 Χ ca. 0.3 cm, base adnate to rachis, entire, apex acute, slightly reflexed when dry; veins invisible, pinnate, furcate, ending in submarginal fusiform hydathodes; rachis brown or stramineous, shallowly grooved on upper side, sparsely scaly; scales brown, lanceolate. Sori linear, one on either side of costa, occupying most [??of] pinna-lamina.

Forests. Taiwan [Japan].

7. DIPLOBLECHNUM Hayata, Bot. Mag. (Tokyo) 41: 702. 1928[??1927].

扫把蕨属  sao ba jue shu

Tree ferns, terrestrial. Stem black, erect, up to ca. 0.8 m tall, 1–2 cm in diam., covered with stubs of persistent frond bases, apex densely scaly; scales dark brown, narrowly deltoid-lanceolate, ca. 1 cm, entire, subulate. Fronds tufted, shortly stipitate; stipe ca. 3 cm, scaly; lamina bipinnatifid, elliptic-lanceolate, narrowed downward, up to 40 Χ 10–14 cm, papery, both surfaces glabrous, acuminate; pinnae to 30 pairs, pinnatifid, oblong-lanceolate, 6.5–8.5 Χ 1.4–1.8 cm, base decurrent to narrow rachis wing with triangular lobes between pinnae, apex acuminate; lower pinnae much shorter, basal ones triangular auriculate; pinna-lobes 13–15 pairs, oblong or triangular-oblong, up to 9 Χ 3 mm, entire, acute; veins free, pinnate, secondary veins 2- or 3-furcate, not reaching laminar margin; rachis and costae grooved adaxially. Fertile lamina as large as sterile lamina, slightly contracted, pinnae caudate. Sori linear, costal, single on either side of costa, borne on straight unbranched fertile veins parallel to costa; indusia facing toward costa, brown, thin, inrolling when young, opening to costa when mature; spore reniform.

One species: distributed in Taiwan and the Pacific islands.

1. Diploblechnum fraseri (A. Cunningham) deVol in H. L. Li et al., Fl. Taiwan 1: 153. 1975.

扫把蕨  sao ba jue

Lomaria fraseri A. Cunningham, Companion Bot. Mag. 2: 264[??364]. 1836; Blechnum fraseri (A. Cunningham) Luerssen; B. integripinnulum Hayata; Diploblechnum integripinnulum (Hayata) Hayata; L. fraseri var. philippinensis H. Christ.

Characters for this species are the same as for the genus.

Moist[??damp or wet] and somewhat exposed path sides. Taiwan [Malaysia, New Zealand, Philippines].

The genus Diploblechnum is an ingroup of Blechnum (R. Cranfill, Y. Nakahira, and M. Kato, unpubl. data), although the two are distinct morphologically.



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

[2] Department of Botany, National Museum of Nature and Science, Tsukuba 305-0005, Japan.