ONOCLEACEAE [Draft]

球子蕨科  qiu zi jue ke

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

Ferns, terrestrial. Rhizome robust and short, erect or creeping, dictyostelic, covered with ovate-lanceolate to lanceolate scales. Fronds tufted or distant, dimorphic. Sterile lamina pinnatifid to bipinnatifid, green, deltoid-ovate, elliptic-lanceolate, or oblanceolate, papery or thinly papery; pinnae alternate, sessile or adnate, linear-lanceolate to narrowly elliptic, undulate or pinnatifid; pinnules hemielliptic-oblong, entire or subserrate with few teeth; veins free, pinnate or anastomosing, without included veinlets. Fertile lamina pinnate or bipinnate, elliptic to linear; pinnae or pinnules much inrolled, dull purple to dark brown, cylindrical or globose; veins free, pinnate or furcate. Sori round, indusiate or exindusiate; sporangia long-stalked, annulus a longitudinal row consisting of 36–40 thickly walled cells; spores monolete, with perispore transparent, membranous, with a few folds.

Three (or four) genera: distributed in the temperate zone of the northern hemisphere and Mexico; two (or three) genera in China.

1a.       Rhizome long-creeping; veins in sterile lamina anastomosing; fertile pinnae moniliform  1. Onoclea

1b.       Rhizome short, erect or creeping; veins in sterile lamina free; fertile pinnae cylindrical  2. Matteuccia

1. ONOCLEA Linnaeus, Sp. Pl. 2: 1062. 1753.

球子蕨属  qiu zi jue shu

Plants medium-sized. Rhizome long-creeping, dark brown, covered with brown scales. Fronds distant. Sterile lamina pinnate, green, ovate-triangular, papery, both surfaces glabrous, with white long filiform hairs only on rachis and costae; pinnae narrowly elliptic, margin shallowly lobed, basal 1 or 2 pairs shortly stalked, adnate to rachis; rachis always with narrow wings; veins obvious, anastomosing with hexagonal areoles, without included veinlets. Fertile lamina bipinnate, much contracted; pinnae linear, shortly stalked, much oblique; pinnules much inrolled into small spheres, subopposite, separate from each other. Sori round, with raised receptacles, terminal at ends of veinlets, indusiate. Sporangia globose, slender-stalked; spores narrowly elliptic, perispore transparent, with folds.

One species: distributed in East Asia (var. interrupta) and North America (var. sensibilis).

1. Onoclea sensibilis Linnaeus var. interrupta Maximowicz, Prim. Fl. Amur. 337. 1859.

球子蕨  qiu zi jue

Onoclea interrupta (Maximowicz) Ching & P. C. Chiu.

Rhizome long-creeping, dark brown, sparsely scaly; scales brown, broadly ovate, ca. 5 mm, membranous, margin entire or slightly undulate, apex acuminate. Stipe of sterile frond 20–50 cm, stramineous except at chestnut-brown base, sparsely brown scaly; sterile lamina basally pinnatifid-pinnate, apically pinnatifid, dark green when dry, ovate-triangular or ovate, 14–30 Χ 13–25 cm, papery, slightly scaly, glabrous when mature; pinnae 5–8 pairs, 1.5–3 cm apart, narrowly elliptic, basal 1 or 2 pairs largest, 8–12 Χ 1.5–3 cm, shortly stalked, margin undulate or lobed, sessile or adnate upward, connected by rachis wing; veins reticulate. Stipe of fertile frond 20–45 cm; fertile lamina bipinnate, much contracted, 15–25 Χ 2–4 cm; pinnae narrowly linear; pinnules globose, hardened, separate from each other. Indusium membranous, fixed at posterior base, free distally.

Wet meadows and wetlands in valleys; 200–900 m. Hebei, Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol [Japan, Korea, Russia].

[The typical variety occurs in ???]

2. MATTEUCCIA Todaro, Giorn. Sci. Nat. Econ. Palermo 1: 235. 1866, nom. cons.

荚果蕨属  jia guo jue shu

Rhizome robust, erect or oblique, covered with brown lanceolate scales. Fronds tufted or approximate. Sterile lamina bipinnatifid and apex pinnatifid, ovate-lanceolate, elliptic-lanceolate, or oblanceolate, papery or thinly papery, nearly glabrous or with few hairs or scales on rachis and costae; pinnae alternate, sessile, narrowly lanceolate, pinnatifid to 1/2; lobes oblique, oblong to elliptic, subentire or subserrate; veins free, pinnate, veinlets reaching lamina margin. Fertile lamina smaller than sterile lamina; lamina pinnate, elliptic to broadly oblanceolate; pinnae linear, alternate, subsessile, much inrolled into brown cylinders, concealing sori. Sori globose, with raised receptacles, borne at ends of veinlets, indusiate or exindusiate; sporangia spheroidal, slender-stalked, annulus longitudinal, consisting of ca. 40 thickly walled cells; spores elliptic, perispore transparent, slightly folded. n = 40.

Three species: distributed in the temperate zone of northern hemisphere; three species in China.

Because Matteuccia is paraphyletic, Pentarhizidium is better segregated from it.

1a.       Basal pinnae of sterile lamina gradually contracted to small auricles ...........  1. M. struthiopteris

1b.       Basal pinnae of sterile lamina shortened or slightly shortened.

2a.       Pinnae 2–3.5 cm wide; sori indusiate ...........................................................  2. M. orientalis

2b.       Pinnae 1.5–1.8 cm wide; sori exindusiate ...................................................  3. M. intermedia

1. Matteuccia struthiopteris (Linnaeus) Todaro, Giorn. Sci. Nat. Econ. Palermo 1: 235. 1866.

荚果蕨  jia guo jue

Rhizome short and robust, erect, dark brown, stoloniferous, densely covered with brown scales; scales lanceolate, ca. 5 mm, membranous, entire, apex fibriform. Fronds tufted; stipe brown, 6–10 Χ 0.5–1 cm, deeply grooved adaxially, base swollen, triangular and densely scaly, sparsely scaly upward. Sterile lamina bipinnatifid, green or brown-green when dry, elliptic-oblanceolate to oblanceolate, 50–100 Χ 18–25 cm, markedly narrowed downward, papery, rachis and costae hairy or scaly; pinnae 40–60 pairs, alternate or subopposite, sessile, oblique, basal pairs gradually contracted to small auricles; pinnae above middle largest, lanceolate or linear-lanceolate, 10–15 Χ 1–1.5 cm, apex acuminate, sessile, pinnatifid, pectinate; lobes 20–25 pairs, close, oblong or subtriangular; veins obvious, 1.5–2 cm apart, pinnate, veinlets simple. Fertile lamina shorter than sterile lamina, pinnate, oblanceolate, 20–40 Χ 1–8 cm, gradually narrowed downward, lower pinnae much reduced; pinnae dark brown, linear-moniliform, hardened, much inrolled, concealing sori. Indusium membranous, fixed at posterior end, free distally.

Forest understories, valley wetlands; 100–3800 m, also cultivated as ornamentals. Gansu, Hebei, Heilongjiang, Henan, Hubei, Jilin, Liaoning, Shanxi, Sichuan, Xinjiang, Xizang [Japan, Korea, Russia; Europe, North America].

1a. Pinna lobes hemielliptic or nearly oblong, apex obtuse .................................  1a. var. struthiopteris

1b. Pinna lobes triangular-elliptic, apex acute ..........................................................  1b. var. acutiloba

1a. Matteuccia struthiopteris var. struthiopteris

荚果蕨(原变种)  jia guo jue (yuan bian zhong)

Osmunda struthiopteris Linnaeus, Sp. Pl. 2: 1066. 1753; Onoclea germanica Willdenow ex Hooker[??not in Tropicos or IPNI]; O. struthiopteris (Linnaeus) Hoffmann; Pteris sinuata Thunberg; Struthiopteris filicastrum Allioni; S. germanica Willdenow.

Pinna lobes hemielliptic or nearly oblong, obtuse at apex.

Forest understories, valley wetlands; 100–3000 m, also cultivated as ornamentals. Gansu, Hebei, Heilongjiang, Henan, Hubei, Jilin, Liaoning, Shanxi, Sichuan, Xinjiang, Xizang [Japan, Korea, Russia; Europe, North America].

1b. Matteuccia struthiopteris var. acutiloba Ching, Fl. Tsinling. 2: 221. 1974.

尖裂荚果蕨  jian lie jia guo jue

Pinna lobes triangular-elliptic, acute at apex.

* Forest understories, valley wetlands; 1500–3800 m. Henan, Hubei, Shanxi, Sichuan.

2. Matteuccia orientalis (Hooker) Trevisan, Atti Ist. Veneto Sci. 3(14)[??vol. 14]: 586. 1869.

东方荚果蕨  dong fang jia guo jue

Struthiopteris orientalis Hooker, Sec. Cent. Ferns 4, t. 4. 1861; Matteuccia japonica (Hayata) C. Christensen; Onoclea orientalis (Hooker) Hooker; Pentarhizidium japonicum Hayata; P. orientale (Hooker) Hayata; Pterinodes orientalis (Hooker) O. Kuntze.

Rhizome short-creeping, apex ascending, dark brown, apical part densely covered with brown scales; scales lanceolate, ca. 2 cm, membranous, entire, apex fibriform. Fronds approximate. Stipe of sterile frond brown, 30–70 Χ 0.3–1 cm, base brown, swollen, triangular, bearing short pneumatophores, densely scaly, upward sparsely scaly; sterile lamina bipinnatifid, ovate-deltoid, narrowed upward, slightly narrowed downward, 40–82 Χ 18–40 cm, thinly papery, with sparse fibriform scales on rachis and costae; pinnae 15–20 pairs, alternate, oblique, 2.5–3 cm apart, basal pinnae as long as above, base slightly narrowed, sessile, apex acuminate; lobes narrowly hemielliptic, oblique, entire or slightly dentate, apex obtuse or acute; veins obvious, pinnate, veinlets simple or occasionally furcate, reaching lamina margin. Stipe of fertile frond 20–45 cm; lamina as long as or shorter than sterile lamina, pinnate, elliptic, 12–40 Χ 5–10 cm; pinnae oblique, close, dark purple, linear, 10 Χ ca. 0.5 cm, hardened. Sori fused into linear coenosori when mature; indusium membranous, fixed at posterior end, free distally. 2n = 80, 82.

Streams of dense forest understories; 1000–2700 m. Anhui, Chongqing, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangxi, Shanxi, Sichuan, Taiwan, Xizang, Zhejiang [India, Japan, Korea, Russia].

3. Matteuccia intermedia C. Christensen, Bot. Gaz. 56: 337. 1913.

中华荚果蕨  zhong hua jia guo jue

Onoclea intermedia (C. Christensen) M. Kato, T. Suzuki & Nakato; Pentarhizidium intermedium (C. Christensen) Hayata.

Rhizome short and robust, erect, dark brown, apical part densely covered with dark brown scales; scales broadly lanceolate, ca. 1.5 cm Χ 4 mm, membranous, entire, apex acuminate. Fronds tufted. Stipe of sterile frond brown, 20–30 Χ ca. 0.5 cm, base dark brown, firm, bearing prominent pneumatophores, with sparse lanceolate scales; sterile lamina bipinnatifid, elliptic, 40–60 Χ 15–25 cm, slightly narrowed downward, chartaceous[thinly papery], glabrous, with brown linear-lanceolate scales on rachis; pinnae 20–25 pairs, alternate, oblique, close, midveins ca. 1.5 cm apart, basal 2 or 3 pairs of pinnae shortened, slightly reflexed; middle pinnae largest, linear or oblong-lanceolate, 13–15 Χ 1.5–2 cm, base truncate, sessile, pinnatifid, apex acuminate; lobes 20–30 pairs, close, oblong, entire, apex rounded or truncate with points; veins obvious, pinnate, veinlets simple or sometimes furcate, reaching lamina margin. Fertile frond shorter than sterile frond; stipe 20–25 cm, 5–8 mm in diam.; fertile lamina shorter than sterile lamina, pinnate, elliptic, 30–45 Χ 8–15 cm; pinnae close, dark purple, linear, 3.5–5.5 Χ 0.2–0.3 cm. Sori exindusiate, fused into linear coenosori when mature. 2n = 80.

Forest understories; 1500–3200 m. Hebei, Hubei, Shanxi, Sichuan, Yunnan [India (Sikkim)].



[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.