裸子蕨科 luo zi jue ke
Zhang Gangmin
Plants terrestrial. Rhizomes creeping, ascending, or erect, bearing scales and/or hairs, dictyostelic or siphonostelic. Leaves monomorphic or somewhat dimorphic, distant, closely spaced, or tufted. Petiole straw-colored or chestnut-colored, with single U-shaped or rounded vascular bundle in cross section. Blade 13-pinnate (rarely simple, base cordate or hastate), herbaceous (less often soft leathery), green, somewhat hairy or scaly (rarely glabrous, or with white farina abaxially as in Pityrogramma). Veins free, rarely anstomosing, without including free veinlets in areoles. Indusia absent. Sori following course of veins. Spores tetrahedral or globose-tetrahedral, verrucose, echinate, or reticulate, rarely plain.
About 17 genera: mostly in tropical and subtropical regions; 5 genera in China.
1a. Leaves somewhat dimorphic, simple, ovate or oblong, base cordate or hastate; veins
anastomosing....................................................................................................1. Parahemionitis
1b. Leaves monomorphic, 13-pinnate (young leaves of Coniogramme simple, lanceolate); veins
free or connected only near leaf margin, if anastomosing, only forming few or 13 rows of
areoles at both sides of main vein.
2a. Blade with white farina abaxially..........................................................2. Pityrogramma
2b. Blade without white farina abaxially.
3a. Blade 12-pinnate, soft leathery, densely covered with imbricate-arranged scales or
long sericeous hairs abaxially...........................................................3. Paragymnopteris
3b. Blade 13-pinnate, herbaceous or papery, glabrous or sparsely pubescent abaxially.
4a. Small plants, usually less than 10 cm tall; blade finely pinnate (round-fan-
shaped when young).............................................................................4. Anogramma
4b. Large plants, to 1 m tall or so; blade not finely pinnate; simple pinnae or
pinnules often over 10 X 1 cm, lanceolate........................................5. Coniogramme
1. PARAHEMIONITIS Panigrahi, Amer. Fern J. 83(3): 90. 1993.
泽泻蕨属 ze xie jue shu
Plants terrestrial. Rhizomes short, erect, bearing small, fluffy scales and long, jointed hairs,
dictyostelic. Leaves somewhat dimorphic, clustered. Petiole chestnut-colored or purple-black,
densely scaly and hairy as on rhizomes; petiole of fertile leaf usually 13 times length of that of
sterile leaf. Blade ovate, oblong-ovate, or hastate, simple, base strongly cordate, apex obtuse or rounded, herbaceous, glabrous adaxially, brown and sparsely covered with small subulate scales abaxially. Veins anastomosing, areoles numerous, densely arranged, elongated hexagonal.
Indusia absent. Sori following course of veins, confluent all over abaxial surface when mature.
Spores globose-tetrahedral, cristate. x = 30.
One species: tropical Asia; 1 species in China. This species was usally put into the genus Hemionitis, but it has strongly dimorphic leaves with subulate scales intead of hairs abaxially, differing from Hemionitis species in America. Panigrahi thought that is reasonable to put the Asian species into another genus Parahemionitis.
1. Parahemionitis cordata (Hooker & Greville) Fraser-Jenkins. New Sp. Syndrome Indian Pterid. & Ferns Nepal, 187. 1997.
泽泻蕨 ze xie jue
Hemionitis cordata Hooker & Greville, Ic. Fil. t. 64. 1828; Gymnogramme sagittata Ettingshausen; Hemionitis sagittata Fιe; H. toxotis Trevisan.
Rhizomes short, erect; scales brownish, narrowly lanceolate. Petiole of fertile leaf much longer than that of sterile leaf (over 23 times in dense forests, ca. 1 time in dry shrubs), 618 cm X ca. 1 mm. Blade ovate, narrowly ovate, or hastate, 36(--10) X 24(--6), forming an oblique angle with petiole, base strongly cordate, margins with sparse, reddish brown, and jointed hairs, apex obtuse or rounded, herbaceous when dry, brownish green and glabrous adaxially, brown and sparsely covered with small subulate scales abaxially (scales rather dense along main vein). Sori following course of veins, confluent all over abaxial surface when mature, brown. 2n = 120.
Moist soil and rock crevices of stream valley in dense forests, or in shrubs; below 1000 m; Hainan, S Taiwan, S Yunnan (Menghai) [Cambodia, India, Laos, Malaysia, Philippines, Sri Lanka, Thailand, Vietnam].
2. PITYROGRAMMA Link, Handb. Gew. 3: 19. 1833.
粉叶蕨属 fen ye jue shu
Plants terrestrial. Rhizomes short, erect or ascending, dictyostelic; scales reddish brown, thin,
lanceolate, margins entire. Leaves monomorphic, clustered. Petiole purplish black, lustrous,
rounded proximally, upwards with single longitudinal groove adaxially, base scaly, glabrous
distally. Blade ovate to oblong, apex acuminate, 23-pinnate, herbaceous to somewhat
leathery, adaxially glabrous, abaxially farinose, farina white or yellow. Pinnae numerous, oblique,
lanceolate, somewhat stalked, base subequilateral, apex acuminate. Pinnules numerous,
anadromous, 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 globose-tetrahedral, dark colored, perispore reticulate with irregular ridges. x = 29 (30).
About 20 species: tropical Africa, America, and Asia; 1 species in China.
1. Pityrogramma calomelanos (Linnaeus) Link, Handb. Gewachse 3: 20. 1833.
粉叶蕨 fen ye jue
Acrostichum calomelanos Linnaeus, Sp. Pl. 2: 1072. 1753; Ceratopteris calomelanos Underwood; Gymnogramme calomelanos Link; Neurogramme calomelanos Diels.
Petiole 4050 cm X ca. 3 mm. Blade elongate-oblong or oblong-lanceolate, 1540 X 1020 cm, base broadly cuneate, apex acuminate, 2-pinnate, thick papery when dry, adaxially grayish green, glabrous, abaxially densely farinose, farina white; rachis and costae purple-black, lustrous, glabrous, grooved adaxially. Pinnae 1620 pairs, oblique, with short, winged stalks; basal pair of pinnae not shortened, lanceolate, 1015 X 25 cm, base slightly inequilateral. pinnules (or segments) 1618 pairs, deltate, ovate-lanceolate or lanceolate, 1.11.4 X ca. 0.5 cm, base inequilatral, somewhat decurrent to costae, apex acute or acuminate; proximal pinnule margins serrate or lobed, lobes serrate or entire, only with 12 teeth at apex. Sori nearly confluent all over abaxial pinnule surface when mature. 2n = 240.
Forest margin or stream bank; below 600 m; Hainan, Taiwan, S Yunnan (Luchun) [Cambodia, Laos, Vietnam; Africa, S America].
Like many species of Pityrogramma, P. calomelanos is an ornamental plant, which is also cultivated in China.
3. PARAGYMNOPTERIS Shing in Indian Fern J. 10:227. 1993.
金毛裸蕨属 jin mao luo jue shu
Plants usually on rock, xeric. Rhizomes short, erect or decumbent, dictyostelic; scales tan, linear or subulate, margins entire, intermixed with villose. Leaves monomorphic, clustered. Petiole
chestnut-brown, lustrous, terete, densely villosous distally. Blade oblong-lanceolate, 12-pinnate, papery or leathery, soft, densely covered with tan (gray when old), fine sericeous hairs throughout
(especially abaxially), or with pellucid, lanceolate, entire, and imbricate-arranged scales abaxially. Ultimate pinnules or segments ovate, oblong, or oblong-lanceolate, base rounded or cordate, margins entire, apex obtuse or rounded. Veins free, pinnately branched and divergent distally, or occasionally connected near leaf 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.
About 5 species: Bhutan, India, Nepal, Russia, Thailand; Europe; 5 species in China.
1a. Blade densely covered with imbricate-arranged ovate-lanceolate scales abaxially.
2a. Blade 1-pinnate; petiole clothed with jointed long hairs or nearly glabrous; rachis glabrous
adaxially; usually growing on bare limestone rocks.........................................1. P. delavayi
2b. Blade 1-pinnate-pinnatifid; petiole clothed with fibrous scales; rachis sparsely scaly
adaxially; growing in crevices of noncalcareous rocks...................................2. P. marantae
1b. Blade densely covered with sericeous hairs abaxially.
3a. Blade 1-pinnate; pinnae rounded at base (occasionally proximal pinnae somewhat cordate
at base); veins connected near leaf margin .................................................3. P. vestita
3b. Blade 1- or 2-pinnate; pinnae or pinnules strongly cordate or with 12 segments at base; veins free, not connected near leaf margin.
4a. Blade 12-pinnate; pinnules ovate, 714 mm ...........4. P. bipinnata
4b. Blade 2-pinnate; pinnules ovate-deltate or hastate, 45 mm....................5. P. sargentii
1. Paragymnopteris delavayi (Baker) K. H. Shing, Indian Fern J. 10: 229. 1993.
滇西金毛裸蕨 dian xi jin mao luo jue
Gymnogramme delavayi Baker, Ann. Bot. 5: 484. 1891; Gymnopteris delavayi Underwood; Gymnopteris delavayi Underwood var. intermedia Ching;
Neurogramme delavayi Diels; Notholaena bureaui Christ; N. delavayi C. Christensen; N. marantae (Linnaeus) Desvaux var. delavayi Tagawa; Paraceterach delavayi R. M. Tryon.
Rhizomes short, stout, ascending or decumbent; scales tan, narrowly subulate. Leaves clustered. Petiole 812 cm X 12 mm, chestnut-black, base slightly scaly and hairy, distally hairy or nearly glabrous. Blade
oblong-lanceolate or broadly linear-lanceolate, 514 X 24 cm, apex shortly acuminate or caudate, imparipinnate, leathery when dry, adaxially pale green, glabrous, abaxially densely covered with brown, pellucid, ovate-lanceolate scales; rachis sparsely with narrow subulate scales. Pinnae (5--)1015 pairs, falcate-lanceolate or lanceolate, 1.52.5 X ca. 0.5 cm, shortly stalked (distal pinnae sessile), base rounded or with an auricle acroscopicly (sometimes a small oblong segment). Veins obscure, pinna midrib slightly concave adaxially, convex abaxially. Sori usually covered by scales. 32 spores per sporangium.
*Limestone crevices in open woods; 22004600 m; Gansu, W Guizhou, Qinghai, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan.
Paragymnopteris delavayi looks very much like P. marantae except its undivided pinnae and jointed long hairs (rarely some hairy scales with 23 rows of cells) on petioles. In addition, both of them distribute in NW Yunnan, even in same locality, only the former grows on calcareous substrates and the later, mostly on noncalcareous rocks. Perhaps P. delavayi is the substitute species of P. marantae.
2. Paragymnopteris marantae (Linnaeus) K. H. Shing, Indian Fern J. 10: 229. 1993.
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 D. Candolle; Cheilanthes marantae Domin; Cinlinalis marantae Desvaux; Gymnogramma marantae Mettenius; Gymnopteris marantae Ching; Notholaena marantae Desvaux; Paraceterach marantae R. M. Tryon.
Rhizomes short, stout, decumbent or ascending; scales tan, elongate-subulate. Leaves clustered or closely spaced. Petiole 617 cm X 1.52.5 mm, covered with fibrous scales. Blade broadly lanceolate,
(8--)1220 X 2.55 cm, pinnate-pinnatifid, gradually tapered to pinnatifid apex, thin papery, adaxially brownish green, glabrous, abaxially densely covered with reddish brown, pellucid, ovate-lanceolate scales; rachis and costae sparsely clothed with small subulate-lanceolate scales. Pinnae (6--)1016 pairs, obliquely patent, ovate-deltate or somewhat deltate, 23 X 11.5 cm, pinnatifid nearly to costae; proximal pairs of pinnae shortly stalked. Segments 25 pairs, oblong-lanceolate or oblong, 58 X ca. 3 mm, apex rounded. Veins obscure. Sori spreading along lateral veins, covered by scales. 2n = 58.
Rock crevices under woods in dry regions; 18004200 m; Sichuan, Xizang, Yunnan [India, Nepal; Europe].
3. Paragymnopteris vestita (Wallich ex Presl) K. H. Shing, Indian Fern J. 10: 230. 1993.
金毛裸蕨 jin mao luo jue
Gymnogramme vestita Wallich ex Presl, Tent. Pterid. 218. 1836; Gymnopteris vestita Underwood; Hemionitis vestita J. Smith; Neurogramma vestita Diels; Notholaena himalaica Fraser-Jenkins; Syngramma vestita Moore; Paraceterach vestita R. M. Tryon.
Rhizomes short, stout, decumbent or ascending; scales tan, elongate-subulate. Leaves clustered or closely spaced. Petiole (6--)1020 cm X 12.5 mm, densely clothed with brownish sericeous hairs. Blade lanceolate, 1015 X 2.55(--7) cm, imparipinnate, soft leathery, brown when dry, adaxially sparsely with pale brown sericeous hairs, abaxially densely with golden sericeous hairs; rachis and costae also sericeous. Pinnae
(7--)1017 pairs, ovate or elongate-ovate, 1.54 X 12 cm, alternate, shortly stalked, base rounded or sometimes slightly cordate, rarely auriculate acroscopicly, margins entire, apex obtuse. Veins obscure, pinnate and divergent, connecting near pinna margins. Sori covered by hairs. 2n = 60.
Rock crevices in shrubs; 8003900 m; W Guizhou, Hebei, Shanxi, Sichuan, Taiwan, Xizang, Yunnan [India, Nepal, Pakistan, Tailand].
4. Paragymnopteris bipinnata (Christ) K. H. Shing, Indian Fern J. 10: 230. 1993.
川西金毛裸蕨 chuan xi jin mao luo jue
Gymnopteris bipinnata Christ in Lecomte, Not. Syst. 1: 55. 1909; G. vestita var. bipinnata Christ; Hemionitis bipinnata Michel; Paraceterach bipinnata R. M. Tryon.
Rhizomes short, stout, decumbent, densely scaly; scales brown, or dark brown, shiny, elongate-subulate. Leaves closely spaced. Petiole densely clothed with pale brown, sericeous hairs when young. Blade lanceolate or broadly lanceolate, 12-imparipinnate, soft leathery when dry, adaxially brownish green, sparsely sericeous, abaxially densely sericeous; rachis and costae clothed with same hairs. Pinnae long stalked. Ultimate pinnules ovate or elongate-ovate, base cordate, margins entire or with 12 small segments, apex obtuse. veins obscure. Sori covered by hairs.
Two varieties; endemic to China.
1a. Blade 2-pinnate; inframeidal pinnae
lanceolate or deltate-lanceolate..................
....................4a. P. bipinnata var. bipinnata
1b. Blade 1-pinnate; pinnae ovate or
elongate-ovate, margins entire, or at most
with 12 free segments in proximal
pinnae........................................................
..................4b. P. bipinnata var. auriculata
4a. Paragymnopteris bipinnata Christ var. bipinnata
川西金毛裸蕨 （原变种）chuan xi jin mao luo jue (yuan bian zhong)
Petiole 1022 cm X 13 mm. Blade 1525 X 37 cm, 2-pinnate (apical portion 1-pinnate). Pinnae 1017 pairs, lanceolate or deltate-lanceolate, imparipinnate. Lateral pinnules 16 pairs, ovate or elongate-ovate, shortly stalked or sessile, base somewhat cordate, margins entire or rarely with 12 small lobes, apex obtuse; terminal pinnule as lateral ones but larger and with longer stalk.
*Rock cliff or crevices by ditches; 11003200 m; Gansu, Shaanxi, Sichuan, Xizang, Yunnan.
4b. Paragymnopteris bipinnata Christ var. auriculata (Franchet) K. H. Shing, Indian Fern J. 10: 230. 1993.
耳羽金毛裸蕨 （变种）er yu jin mao luo jue (bian zhong)
Gymnopteris bipinnata Christ var. auriculata (Franchet) Ching, Lingnan Sci. J. 15: 398. 1936; Gymnogramma vestita Hooker var. auriculata Franchet, Arch. Mns. Paris ser. 2. 10: 123. 1877; Gymnopteris borealisanensis Kitagawa.
Blade 1-pinnate. Pinnae ovate or elongate-ovate, stalked (stalks to 1 cm long), base strongly cordate, margins entire or often enlarged forming auricles at both sides near base, sometimes with 12 free segments, apex obtuse.
*On rock under shrubs or woods; 8003600 m; Gansu, Hebei, Henan, W Hubei, Nei Mongol, Sichuan, Xizang, Yunnan.
Paragymnopteris bipinnata var. auriculata is much similar to the species 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. 1993.
三角金毛裸蕨 san jiao jin mao luo jue
Gymnopteris sargentii Christ, Bot. Gaz. 51: 355. 1911; Notholaena sargentii Fraser-Jenkins; Paraceterach sargentii R. M. Tryon.
Plants small, less than 40 cm. Rhizomes short, stout, decumbent or ascending; scales shiny, brown, narrowly lanceolate. Leaves clustered. Petiole light chestnut-colored, 512 cm X 1.52 mm, densely covered with light brown sericeous hairs proximally, becoming sparse distally. Blade deltate-lanceolate or oblong-lanceolate, 1025 X 410 cm, 2-pinnate (apical portion 1-pinnate), leathery when dry, adaxially brownish green, with few sericeous hairs, abaxially densely sericeous; rachis and costae with same hairs as on blade surface. Pinnae 814 pairs, inframedial pinnae lanceolate, 3.57 X 11.5 cm, alternate, stalked (stalks 37 mm), imparipinnate. pinnules 38 pairs, anadromous, ovate-deltate or hastate, 45 X 34 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.
*Rock slopes, dry valley in shrubs; 19003300 m; W Sichuan, Xizang, NW Yunnan.
4. ANOGRAMMA Link, Fil. Sp. Cult. 137. 1841.
翠蕨属 cui jue shu
Plants small, annual, terrestrial or on rock. Rhizomes short, poorly developed, sparsely scaly; scales small, fibrous. Leaves numerous, monomorphic, tufted. Petiole chestnut-colored or chestnut-brown, slender, grooved adaxially, rounded abaxially, glabrous. Blade ovate, ovate-deltate to ovate-lanceolate, or lanceolate, 13-pinnate, thin herbaceous or nearly membranous, usually glabrous on both surfaces. Ultimate pinnules or segments small, ovate-elliptic, spathulate, or obovate, margins entire, or shallowly lobed at apex. Veins free, forked, 1 veinlet for each segment, far from leaf margins. Indusia absent. Sori borne along veinlets, paraphyses absent. Spores tetrahedral, somewhat ridged on surface. Gametophytes still alive for a long time after sporophytes growing up. x = 29, 58.
About 6 species: world tropics, subtropics, and Europe area along Atlantic; 2 species in
China.
1a. Ultimate pinnules or segments obovate or obdeltate (pinnae fan-shaped when blade
1- pinnate), apex shallowly lobed.......................................................................1. A. leptophylla
1b. Ultimate pinnules or segments ovate-elliptic or spathulate, apex obtuse or mucronulate...........
..........................................................................................................................2. A. microphylla
1. Anogramma leptophylla (Linnaeus) Link, Fil. Sp. 137. 1841.
薄叶翠蕨 bo ye cui jue
Polypodium leptophyllum Linnaeus, Sp. Pl. 2: 1092. 1753; Acrostichum leptophyllum D. Candolle; Grammitis leptophylla Swartz; Gymnogramma leptophylla Desvaux.
Rhizomes small, short; scales few, pellucid. Leaves tufted. Petiole 26 cm X 0.51 mm,
Chestnut-brown, glabrous or occasionally with 12 pellucid and jointed long hairs. Blade ovate-deltate to ovate-lanceolate, 48 X 1.53 cm, 2-pinnate to 2-pinnate-pinnatifid, nearly membranous when dry, yellowish green, glabrous. Basal pinnae usually larger than adjacent ones. distal pinnae gradually reduced. Ultimate pinnules or segments obovate or obdeltate, base cuneate and decurrent, apex shallowly lobed. Veins visible on both surfaces. 2n = 58.
Evergreen broad-leaf forest along stream bank; below 2900 m; Taiwan, Yunnan [India,
Vietnam; Africa, Australia, Europe, Pacific Islands, and S America].
Anogramma leptophylla is various in its leaf shape. The blades of its young plants can be round-fan-shaped, bifurcate, or tripartite, or in 1-pinnate individuals elongate-ovate, and all the pinnae are 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, Ic. Pl. t. 916. 1854; Grammitis microphlla Beddome.
Rhizomes short, together with petiole base sparsely scaly; scales blackish, less than 0.8 mm. Leaves numerous, clustered. Petiole chestnut-brown, 210 cm X ca. 0.6 mm, glabrous. Blade ovate-deltate or ovate-lanceolate, 25 X 24 cm, 2-pinnate-pinnatifid to 3-pinnate, thin herbaceous when dry, green, glabrous on both surfaces. Pinnae 57 pairs, basal pair largest, ovate-deltate, 12 X 0.71.2 cm, inequilateral, shortly stalked. Pinnules anadromous, with winged short stalks. Ultimate pinnules or segments elliptic or spathulate, base elongate-cuneate and decurrent to costules, margins entire, apex obtuse or mucronulate, rarely emarginate. Veins visible on both surfaces. 2n = 116.
On rock or in crevices in gorges; 11002900 m; E Guangxi, Guizhou, Taiwan, Yunnan [Bhutan, India, Myanmar, Nepal, Sikkim].