DRYOPTERIDACEAE: part I [DRAFT]

Zhang Xianchun (张宪春); Dave Barrington

#. CYCLOPELTIS J. Smith, Bot. Mag. 72: 36. 1846.

[Good genus but allied to Tectariaceae (Zhang) or Lomariopsidaceae (Barrington).]

拟贯众属  ni guan zhong shu

Zhang Xianchun (张宪春); Dave Barrington

[Hemicardion Fιe.]

Rhizome ascending, short, thick, bearing numerous scales on rhizome apex and stipe, smaller upward on rachis and lower side of costae; scales chestnut brown, linear, margin toothed. Fronds tufted, bright bluish green above when fresh, narrowly elliptic-lanceolate, 1-pinnate; stipe and rachis not or only slightly channeled, not winged; pinnae alternate, mostly sessile, lowest pinnae shortly stalked, upper pinnae falcate, glabrescent adaxially, subentire, usually articulate to rachis; costa prominent above, not grooved; veins free, forked several times. Sori round, dorsal on veinlets, in 1–4 rows on each side of costa; indusia peltate; spores bilateral, ellipsoid, with perispore folds, granulate. n = 41.

Species five or six: tropical Asia and America, one in China.

The genus Cyclopeltis should be classified within the tectaroid ferns, not the polystichoid ferns as Ching (1957 [citation]) suggested. The multicellular hairs on fronds are not found in Dryopteridaceae but in Tectariaceae.

This genus is possibly in Lomariopsidaceae according to Smith [??citation].

1. Cyclopeltis crenata (Fιe) C. Christensen, Index Filic., Suppl. 3, 64. 1934.

拟贯众  ni guan zhong

Hemicardion crenatum Fιe, Mιm. Foug., 5: 283. 1852 [??1850].

Rhizome ascending, stiff, short, thick, apex and base of stipes with dense scales; scales brown, linear, up to ca. 1.5 cm, margin with numerous regular sharply pointed teeth. Fronds 70–90 cm, stipe 25–55 cm, 4–5 mm in diam. at base, brown [??the fronds and/or stipe]; lamina bright bluish green above when fresh, narrowly elliptic-lanceolate, 50–65 Χ 12–25 cm, 12–25 cm wide at middle, base attenuate, 1-pinnate, apex long acuminate; pinnae 10–15 pairs, alternate, articulate to rachis, lanceolate, middle pinnae 8–14 Χ 1.5–2.8 cm, base truncate on acroscopic side, cordate on basiscopic side and auriculate over rachis, margin undulate-serrate, apex subacuminate or caudate; veins free, 4-forked, reaching margin; texture papery; with short erect multicellular hairs at pinnae base near rachis, along costa beneath, and sparsely at pinna margins and on surfaces. Sori round, dorsal on veinlets, in 1–4 rows on each side of costa; indusia peltate, with a few minute marginal hairs; spores monolete, with perispore folds, granulate.

Clay soil of forest understories, beside rocks along streams; 400–1300 m. [??add Guizhou], Hainan, [??add Yunnan] [Myanmar, Vietnam to the Philippines [list countries]].

1. CYRTOGONELLUM Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 327. 1938.

[Maintain or place in Polystichum? Barrington: Probably best left as is, though this leaves an artificial Polystichum.]

柳叶蕨属  liu ye jue shu

Zhang Xianchun (张宪春); Dave Barrington

Plants small to medium sized. Rhizome erect, short. Fronds tufted, glossy green, sparsely scaly above base of stipe; scales uniformly brown, ovate-lanceolate, thickly textured, margin fimbriate, acuminate; stipes stramineous, sulcate adaxially; lamina yellow or green when dry, 1-pinnate, ovate-oblong to lanceolate, apex pinnatifid or with an apical pinna conform[??]; pinnae 4–34 pairs, alternate or opposite basally, stalked, ovate-oblong to lanceolate, bases equal, subequal, or unequal, margin entire to crenate-serrate; lateral main veins absent, veinlets 3–5 in subpinnate groups, free or anastomosing toward margin and forming 1 or 2 rows of oblique elongate areoles each with one simple soriferous included veinlet; lamina texture papery to leathery, with small fibrillose scales along veins abaxially; rachises and costae adaxially sulcate, interrupted at junction. Sori round, terminal on acroscopic basal veinlet of each group, or on included veinlets, uniseriate and medial between costa and margin, not parallel; indusia brown, round, scalelike, peltate, membranous, entire, fugacious; spores 32 per sporangium, bilateral, ellipsoid, opaque, coarsely granulate.

About four closely related apomictic species: confined to the limestone rocky areas in SW China and Vietnam, with the center of distribution in Guizhou and adjacent regions, inhabiting niches in calcareous rocks; all four species in China (two endemic).

The genus Cyrtogonellum is an intermediate between Cyrtomium and Polystichum.

1a.     Pinnae rarely longer than 2 cm, ovate, shortly acuminate, bases unequal .........  4. C. inaequale

1b.     Pinnae (3–)5–11 cm, lanceolate or ovate-lanceolate, acuminate, bases equal or subequal.

2a.    Fronds generally with conform[??] apical pinna; pinnae usually 3–6(–10) pairs; veins normally regularly anastomosing .......................................................................................  3. C. fraxinellum

2b.    Fronds with pinnatifid apex; pinnae usually above 10 pairs; [veins?].

3a.     Pinnae 10–20 pairs, lanceolate or linear-lanceolate, bases subequal .........  1. C. caducum

3b.     Pinnae 20–30 pairs, falcate or lanceolate-falcate, bases unequal, ± auriculate on acroscopic side     2. C. xichouense

1. Cyrtogonellum caducum Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 330. 1938.

离脉柳叶蕨  li mai liu ye jue

Cyrtogonellum falcilobum Ching ex Y. T. Hsieh; C. salicifolium Ching ex Y. T. Hsieh; C. tenuium Ching.

Rhizome erect or ascending. Stipe 12–25(–46) cm, scaly at base, smooth upward; scales brown, ovate-lanceolate, margin fimbriate; lamina green when dry, 1-pinnate, lanceolate, 15–30(–54) Χ 6–12 cm, apex pinnatifid; pinnae 13–25 pairs, lanceolate, 3.5–10 Χ 0.8–2 cm, bases subequal, margin subentire to regularly crenate-serrate above base, apex acuminate, pinnatifid; texture papery, with small fibrillose scales along veins abaxially; veins 3- or 4-forked, occasionally joining toward margin, large sori terminal on acroscopic basal veinlets of each group, sori round; indusia peltate, round, entire, fugacious.

Crevices of limestone rocks in forests; 300–1800 m. Chongqing, Guangxi, Guizhou, Hunan, Sichuan, Yunnan [Vietnam].

2. Cyrtogonellum xichouense S. K. Wu & S. Mitsuta, Acta Phytotax. Geobot. 36: 25. 1985.

西畴柳叶蕨  xi chou liu ye jue

Cyrtogonellum simile Ching ex Y. T. Hsieh.

Rhizome short, erect. Stipe dark brown, lanceolate, 18–30 cm, scaly at base, smooth upward, margin denticulate, apex fimbriate; lamina green when dry, 1-pinnate, lanceolate, 30–35 Χ 5–6(–9) cm, apex pinnatifid; pinnae 16–30 pairs, falcate, 2.5–3(–5) Χ 0.8–1 cm, bases unequal, acroscopically slightly auriculate, margin regularly crenate, apex acute; texture papery, with small fibrillose scales along veins abaxially; veins 2- or 3-forked, free, large sori terminal on acroscopic basal veinlets of each group, round; indusia peltate, round, entire, fugacious.

* Crevices of limestone rocks in forests; 900–1700 m. Guizhou, Yunnan.

3. Cyrtogonellum fraxinellum (H. Christ) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 329. 1938.

柳叶蕨  liu ye jue

Aspidium fraxinellum H. Christ, Bull. Herb. Boissier 7: 15. 1899; Cyrtogonellum omeiense Ching ex Y. T. Hsieh; C. Χ rupicola P. S. Wang & X. Y. Wang; Polystichum fraxinellum (H. Christ) Diels.

[Rhizome unknown?] Stipe to 23 cm; lamina pale brownish abaxially, ovate, normally conform[??], rarely apex slightly pinnatifid; pinnae 3–6 pairs, ovate-lanceolate, 7–11 Χ 2–3 cm, bases equal and cuneate, rather long stalked, acuminate; uppermost pinnae barely smaller, entire except obscurely crenate toward apex; texture rather thickly leathery, wrinkled abaxially, with small fibrillose scales along veins abaxially; veinlets 4 or 5 per group, immersed, quite regularly anastomosing toward margin, forming a row of elongate oblique areoles each with one included acroscopic basal veinlet terminated by a large sorus; large sori terminal on acroscopic basal veinlets of each group, round; indusia peltate, round, entire, fugacious.

Crevices of limestone rocks in forests; 500–1800 m. Guangxi, Guizhou, Hunan, Sichuan, Taiwan, Yunnan [N Vietnam].

4. Cyrtogonellum inaequale (H. Christ) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8: 331. 1938.

斜基柳叶蕨  xie ji liu ye jue

Cyrtomium fraxinellum (H. Christ) Ching var. inaequale H. Christ, Bull. Acad. Int. Gιogr. Bot. 11: 264. 1902; C. minium Y. T. Hsieh[??okay].

Rhizome erect or ascending. Stipe 6–20 cm, densely scaly at base, smooth upward; scales brown, lanceolate, margin fimbriate; lamina pale brownish abaxially, imparipinnate, linear-lanceolate, 20–30 Χ 3–4 cm; apical pinna reduced; pinnae 7–34 pairs, alternate, close, obliquely ovate, 1.5–2.5 Χ 0.6–1.4 cm, base unequal, hardly cuneate, stalked, margin remotely but regularly incised-serrate above entire base, apex shortly acuminate or acute, seldom obtuse; texture rather thickly leathery, wrinkled abaxially, costa distinct on both sides, with small fibrillose scales along veins abaxially; veins free, slightly immersed, 3-forked; large sori terminal on acroscopic basal veinlets of each group, round, forming one row between costa and margin of pinna, submarginal, 8–10 on acroscopic side of costa, and about half as many on basiscopic side; indusia peltate, round, thick, entire, fugacious.

* Crevices of limestone rocks in forests; 500–1300 m. Chongqing, Guizhou.

Cyrtogonellum inaequale resembles C. fraxinellum only in texture, color, and the free apical pinna. It differs in more numerous pairs of small ovate pinnae with unequal bases, free venation (rarely the lower larger pinnae with few veinlets anastomosing, forming few areoles), and the regularly incised-serrate margin above the entire base.

2. CYRTOMIDICTYUM Ching, Bull. Fan Mem. Inst. Biol. Bot. 10: 182. 1940.

[Reduce to synonymy?, place species in Polystichum? or Cyrtogonellum? Barrington: Belongs with lime-loving species of Polystichum in section Haplopolystichum and with Cyrtogonellum, would prefer to sink into Polystichum.]

鞭叶蕨属  bian ye jue shu

Zhang Xianchun (张宪春); Dave Barrington

Rhizome erect or ascending, short. Fronds tufted, long stipitate, drying brown, lanceolate to broadly oblong-lanceolate, 1-pinnate or pinnatifid in lower part, uniform or subdimorphic, some sterile fronds bearing a scaly bulbil on an extended, whiplike rachis, bulbil retained and rooting to yield a new plant upon reaching the ground; stipe and rachis abaxially covered copiously with broad, ovate, acuminate, long fimbriate, concolorous, spreading scales; pinnae ovate to lanceolate, entire or toothed at margin, acute or acuminate, ± falcate; texture thickly papery to leathery, abaxially copiously covered with broad, amorphous, ciliate, thin, appressed small scales, glabrous adaxailly; veins obscure, in pinnate groups, obliquely ascending, 2- or 3-forked, free or casually anastomosing. Sori round, small, dorsal on veinlets, in 1 or 2 rows on each side of costa of pinnae, exindusiate; spores # per sporangium, bilateral, ellipsoid, perispores verrucose or with broad winglike folds.

Two species: E and SE China (including Taiwan) to C and S Japan, and S Korea. A genus with many essential characters similar to the 1-pinnate and tripartite species of Polystichum. Both species in China (one endemic).

1a.     Lamina 1-pinnate; pinnae acroscopically auriculate; sori usualy in two rows on each side of the costa      1. C. lepidocaulon

1b.     Lamina simple or usually pinnatifid; pinnae adnate and not auriculate at acroscopic base; sori in one row on each side of costa ..........................................................................................  2. C. basipinnatum

1. Cyrtomidictyum lepidocaulon (Hooker) Ching, Bull. Fan Mem. Inst. Biol. Bot. 10: 162. 1940.[??p. 182]

鞭叶蕨  bian ye jue

Aspidium lepidocaulon Hooker, Sp. Fil. 4: 12. 1862; Cyrtomidictyum conjunctum Ching; C. faberi (Baker) Ching; C. lepidocaulon var. incisa Ching; Dryopteris lepidocaulis (Hooker) Kuntze; Nephrodium faberi Baker; Polystichum lepidocaulon (Hooker) J. Smith.

Rhizome erect or ascending, short, densely covered at growing tip with dark brown lanceolate scales. Stipe 10–30 cm, densely covered with scales; scales thin, brown to light brown, dimorphic, smaller scales ovate-lanceolate, larger scales ovate, margins of scales prominently fimbriate; lamina 1-pinnate, linear-lanceolate, 15–35 Χ 5–15 cm, apex acuminate, pinnatifid; free pinnae 5–15 pairs, shortly stalked, or subsessile upward, spreading, alternate, basal pinnae as long as next ones above or rarely longer, lanceolate-falcate, 4–10 Χ 0.8–1.8 cm, abaxially with scattered small appressed, thin, fimbriate scales, glabrous adaxially, base unequal, oblique below, with a triangular auricle above, entire throughout, acuminate; texture thickly papery or subleathery; veins obscure, 3–6 in pinnate groups, reaching margin of pinnae except for basal acroscopic fertile veinlet, free or occasionally basal ones of adjacent groups or same group on same side joining. Sori round, medial, small, abaxial on lower 2 or 3 veinlets of each group, exindusiate, normally in 2 rows (sometimes 1 row along each side of costa, sometimes 3 rows on auricle). Sterile rooting fronds with more slender and longer scaly stipes, lamina much narrower, usually consisting of fewer pairs of smaller remote pinnae, rachis elongating into a long whiplike stolon, ending in a scaly bud.

Wet forests; 300–2400 m. Anhui, Fujian, Guangxi, Hunan, Jiangsu, Jiangxi, Taiwan, Zhejiang [C and S Japan, S Korea].

Cyrtomidictyum conjunctum and C. faberi are smaller forms of C. lepidocaulon. There is no important difference among them. Whether the basal veinlets in each group reach the margin of the lamina is not useful in defining species, since the two venations, which were used to distinguish C. lepidocaulon and C. faberi, co-occur on individual plants. In sterile fronds, the acroscopic basal veinlets always reach the margin, but not in fertile fronds.

2. Cyrtomidictyum basipinnatum (Baker) Ching, Acta Phytotax. Sin. 6: 262. 1957.

单叶鞭叶蕨  dan ye bian ye jue

Aspidium basipinnatum Baker, J. Bot. (London) 27: 176. 1889; Polystichum basipinnatum (Baker) Diels.

Rhizome erect, short, densely covered at growing tip with dark brown, lanceolate scales. Stipe 8–14 cm, densely covered with scales; scales thin, brown to light brown, dimorphic, smaller scales ovate-lanceolate, larger scales ovate, margins of scales prominently fimbriate; lamina drying brown, linear-lanceolate, 15–20 Χ 2.5–3.5 cm, apex acuminate, pinnatisect below, apex pinnatifid, shallowly pinnatifid to entire; pinnae [numerous(give number)], contiguous, lower ones oblong-ovate, 1.2–1.4 Χ ca. 0.7 cm, entire, acute, base adnate to rachis, upper side not auriculate; texture subleathery, glabrous adaxially, scaly on surface and rachis abaxially; veins obscure, in pinnate groups, all 2-forked, or rarely 3-forked, ascending, reaching margin of pinnae. Sori round, small, medial and uniseriate on each side of costa. Sterile lamina simple, or similar to fertile ones, usually less deeply pinnatifid, sometimes rachis elongate into a long whiplike nodding stolon and viviparous at apex.

* Very rare, in forests, by streams. Guangdong, Guangxi.

Although Cyrtomidictyum basipinnatum appears to be a young stage of Cyrtomidictyum lepidocaulon, it sometimes has a simple sterile or fertile fronds not found in that species. [Perhaps too little evidence to recognize as separate species?]

#. SOROLEPIDIUM H. Christ, Bot. Gaz. 51: 350. 1911.

玉龙蕨属  yu long jue shu

[Place species in Polystichum. Agreed by Zhang and Barrington.]

According to our new molecular analysis, the species of Sorolepidium are nested within Polystichum, in the section of high alpine Asian small species, i.e., sect. Sorolepidium (Christ) Tagawa, or sect. Lasiopolystichum Daigobo. Two species are recognized in Flora Reipublicae Popularis Sinicae 5(2). An examination of more materials and field observations confirm that they pertain to one polymorphic species.

1. Sorolepidium glaciale (H. Christ) H. Christ, Bot. Gaz. 51: 350. 1911.

玉龙蕨  yu long jue

Polystichum glaciale H. Christ, Bull. Soc. Bot. France. 52, Mιm. 1: 28. 1905; Sorolepidium ovale Y. T. Hsieh.

Rhizome erect, thick, densely rooted. Fronds tufted, 4–7; stipes 1–5(–8) cm, densely covered with pale brown or ochreous-yellow scales of two types: short, fibrillose, and long, broadly ovate, to 6 mm, acuminate; lamina linear-lanceolate, 10–15 Χ 1.5–2.5 cm, broad at middle, tapering to both ends, 1-pinnate; rachis thickly covered with lanceolate subulate scales; pinnae [confert], 15–25 pairs, alternate, subsessile, 0.5–1.2 Χ 0.3–0.6 cm, oblong, margin strongly reflexed, crenate, truncate or cordate with whitish appressed setaceous scales adaxially, abaxially densely covered with large lanceolate, brown scales on costa and smaller ones on veins; scales linear-lanceolate, ca. 2 mm, scarious; veins entirely hidden, free, forked or subpinnate near base. Sori 3 or 4 on each side of costa, almost round, hidden, exindusiate; surface of spores with echinate fenestrate folds.

Calcareous rock crevices up to snow line or by glaciers on cold alpine mountains; 2600–4700 m. Gansu, Qinghai, Sichuan, Taiwan, Xizang, Yunnan [India (Himalayas)].