DRYNARIACEAE [FIRST DRAFT]
槲蕨科 hu jue k e
Zhang Xianchun (张宪春)1;
Key to genera
1a. Frond internally dimorphic, separate sterile base frond absent.
2a. Frond pinnatifid, sessile with a dilated base. .. 1. Aglaomorpha
2b. Frond pinnate; stipe not winged. .. 2. Photinopteris
1a. Frond dimorphic, separate sterile base frond present (except Drynaria parishii). .. .. 3. Drynaria
1. AGLAOMORPHA Schott, Gen. Fil. ad t. 19. 1836.
连珠蕨属 liang zhu jue shu
Zhang Xianchun (张宪春)1;
Pseudodrynaria (C. Chr.) C. Chr. in Verdoon, Manual. Pterid. 548. 1938.
Epiphytic, epilithic or terrestrial. Rhizome thick, short to long creeping. Anatomy: vascular bundles 20 to many (50-100), in cross section arranged in 1 or 2 flattened circles with conspicuous dorsal invaginations or protrusions; sclerenchyma strands absent. Rhizome scales appressed or spreading, pseudopeltate or rarely peltate, margin toothed or ciliate with 1- or 2-celled glandular projections. Fronds non-articulate, monomorphic, usually internally dimorphic, sessile with a dilated base, fronds bases imbricated or separate, forming individual nests, rachises not persistent; laminae deeply pinnatifid or subpinnate, with conspicuous nectaries situated below the junctions of rachis and costae, or of costae and veins. Pinnae abscissile from costa and from each other by a line of abscission betweeb costa and base of sinus; gradually smaller towards the frond-apex, entire, apical pinna present. Venation highly complex, with main areoles delimited by the veins and connecting veins, filled with numerous small areoles containing excurrent and recurrent free veinlets, each veinlet terminating in a hydathode. Fertile parts similar to sterile or usually narrowed. Sori small, in rows along connecting veins or veinlets, or distinctly enlarged to soral patches, in one row between midrib and margin. Sporangia glabrous or sometimes with 1-3 acicular hairs. Spores with spines or small globules. Chromsome number n = 36, 37.
About 31 species restrict to tropical Asia, most abundantly in Malesia, from Himalays to Taiwan. Mainly in tropical forests, forming large nests on tree trunks or on rocks. Like Drynaria, but fronds internally dimorphic. Photinopteris possessing several unique characters is not included here otherwise the delimitation is according to Roos (1985).
Key to species
1a. Fertile pinnae similar to sterile pinnae .. 1. Aglaomorpha coronans
1b. Fertile pinnae distinctly contracted 2. Aglaomorpha meyeniana
1. Aglaomorpha coronans (Wall. ex Mett.) Copel., Univ. Calif. Publ. Bot. 16: 117. 1929.
崖姜 ya jiang
Polypodium coronans Wall., List no. 288. 1828. nom. nud.; Mett., Farngatt. 1. Polyp. 121. t. 3, f.40-41. 1857; Pseudodrynaria coronans (Wall. ex Mett.) Ching
Rhizome 2--3 cm thick or more; dilated fron dbases imbricated, rachises not persistent. Rhizome scales spreading, pseudopeltate, 5--20 x 0.5--1 mm, basal auricles short, the margin toothed, apex narrowly acuminate to short-filiform, midrib absent. Fronds monomorphic or with the fertile parts slightly narrowed, sessile, the dilated base lobed, upwards pinnatifid up to 0.2 cm from the rachis, 70--170 x 20--45 cm, glabrous, apical pinna present. Pinnae without basal contriction, 15--35 x 1.5--5 cm, margin entire, apex acute to acuminate. Sori usually present on all pinnae, in several rows between midrib and margin, one in each main areole, elongated, 1--3 mm in largest diameter, slightly sunken. Sporangia glabrous. Spores with spines and verrucate.
Epiphytic, forming a ring-shaped basket around tree trunks, or epilithic, primary and secondary forest. Alt. 100--1900 m. Fujian, Guangdong, Guangxi, Hainan, Hong Kong, Guizhou, Taiwan, S. Xizang, S. Yunnan [Indochina, Myanmar, India, Nepal, Peninsular Malaysia, Ryukyu Islands].
2. Aglaomorpha meyeniana Schott, Gen. Fil. t. 19. 1835 (1836).
连珠蕨 liang zhu jue
Rhizome 2--3 cm thick or more, short-creeping; dilated frond bases imbricated, rachises not persistent. Rhizome scales spreading, pseudopeltate, 6--15 x 0.4--1(--1.3) mm, basal auricles short, apex narrowly acuminate to long-filiform, strongly dentate, midrib absent. Fronds internally dimorphic, sessile, the dilated base lobed, upwords pinnatifis, 35--90 x 15--30 cm, glabrous or abscission vein with tufts of very short acicular hairs, apical pinna present. Sterile pinnae without basal constriction, 7.5--15 x 1.5--3.5 cm, apex rounded, acute to acuminate. Fully fertile pinnae in upper 2/3 of the frond, distinctly narrowed, up to 5--20 x 0.4--0.8 cm, contracted between the sori. Sorial patches in one row along the pinna-midrib, protruding bead-like, round, 1--3 mm wide. Sporangia glabrous. Spores with verrucate.
Epiphytic, forming a ring-shaped basket around tree trunk.Alt. 450--1600 m, or epilithic, or terrestrial, usually in exposed areas in primary forest. Taiwan [Philippines].
2. PHOTINOPTERIS J. Sm., J. Sm. in Hook. J. Bot. 3: 403. 1841.
顶育蕨属 ding yu jue shu
Zhang Xianchun (张宪春)1;
Aglaomorpha, M.C. Roos, Phylog. Syst. Drynar. Polyp. [Utrecht] 227. 1985, pro parte;
Like Drynaria but differs from it in: fronds stipitate, without basal fronds or expanded lamina at lower part, fertile pinnae very narrow, elongate, sori acrostichia. Several monotypic genera were segregated principally by the difference in the form of fertile pinnae which were all included in a single genus by the monographer Roos (1985). Photinopteris is maintained here pending futher researches especially by molecular evidences.
1. Photinopteris acuminata (Willd.) C.V. Morton in Contr. U. S. Nat. Herb. 38: 31. 1967.
顶育蕨 ding yu jue
Acrostichum acuminatum Willd., Sp. Pl. ed. 5, 116. 1810; Aglaomorpha acuminata (Willd.) Hovenkamp; Photinopteris speciosa (Blume) C. Presl; Aglaomorpha speciosa (Blume) M.C. Roos
Rhizome glaucous, long-creeping, diam. 7--20 mm; scales brown, linear-lanceolate, 3--10 x 0.2--1.2 mm, apex narrowly acuminate to filiform-sublate, margin dentate to ciliate, base auriculate; fronds internally dimorphic, stalked, stipes 10--35 cm long, with two rows of rudimentary pinnae; laminae pinnata, 30(--) 45--90 x 13--30 cm, coriaceous, glabrous to sparsely hairy with hairs up to 3 mm long, apical pinna present, nectaries on small extensions of the pinna-base at the basiscopic, sometimes also at the acroscopic side; sterile pinnae ovate, 7--30 x 3.5--10 cm, apex acuminate to caudate, base cuneate, pedicles to 1 cm long; fertile pinnae in upper 2/3 of the frond, a few to several pairs, strongly narrowed, linear, 10--27 cm long, 4--7 mm broad. Sori (coenosori) linear, covering the whole under surface of the fertile pinnae except on costa and marginal portion. Spores with small globules.
Epilithic, climbing on limestone cliffs in exposed or shaded places in tropical rain forests, alt. 1300--1400 m, forming large populations, but uncommon locally. S. Yunnan [Indochina,Thailand, Sumatra, Malaya, Borneo, Philippines, Sulawesi].
3. DRYNARIA (Bory) J. Sm., Hook. J. Bot. 3: 397. 1841 [nom. cons.: Pic. Serm., Taxon 21: 707. 1972].
槲蕨属 hu jue shu
Zhang Xianchun (张宪春)1;
Epiphytic, epilithic or terrestrial. Rhizome short to long-creeping; rhizome scales appressed or, often squarrosely, spreading, basifixed or peltate, margin toothed; fronds ususally dimorphic, with base and foliage frond, rarely monomorphic (base frond absent); base fronds sesile, rounded to ovate-elliptical, entire to lobed up to 2/3; foliage fronds internally monomorphic or apical fertile part slightly narrowed, stalked, pinnatifid, apex often aborted, with a lateral pinna taking its place, rarely pinnate (then pinnae articulted to the rachis, deciduous), hairs sometimes present, spread over the lamina; in pinnatifid species, pinnae abscissile from costa and from each other by a line of abscission between costa and base of sinus; basal pinnae reduced; costae, costules and main veins prominent beneath, cross-veins and minor venation evident, much anastomosing with occasional free-included-veinlets, veins forming glandular plexuses along costa, usually just above costules; mature fronds without hydathodes on upper surface, margins cartilaginous, laminae of mature fronds scaly at first, glabrrescent. Sori small, in rows along veins or connecting veins, very shallowly impressed. Sporangia glabrous or sometimes with glandular hairs. Spores with spines or small glabules. Chromsome number n = 36, 37.
Paleotropical, 16 species. 9 in China, mainly in subtropical and tropical region, epiphytic, epilithic, or terrestrial, one species (D. sinica) extends to North China.
Key to species
1a. Foliage fronds pinnate; pinnae articulate to rachis 1. D. rigidula
1b. Foliage fronds pinnatifid; pinnae not articulate to rachis.
2a. Base fronds thick, not transparent; sori in tow or more rows between costa and margin.
3a. Base fronds rounded, margin emtire or shallowly undulate; sori small, irregular between lateral veins. 2. D. bonii
3b. Base fronds ovate or cordate, margin distinctly lobed; sori large, 1--2 rows between lateral veins.
4a. Plants 30--40 cm high; base fronds 3--5 x 2--4 cm; sori in one row between lateral veins. 3. D. roosii
4b. Plants up to 100 cm high; base fronds 20--40 x 16--32 cm; sori in two rows between lateral veins. 4. D. quercifolia
2b. Base fronds thin, transparent; sori in one row between costa and margin.
5a. Scales imbricate, peltate, rigid; foliage fronds glabrous.
6a. Base fronds absent; foliage fronds with 5--9 pairs of pinnae. 5. D. parishii
6b. Base fronds exist; foliage fronds with 8--12 pairs of pinnae. 6. D. propinqua
5b. Scales basifixed, sorft and tuft; foliage fornds often with haris.
7a. Apex of foliage fronds not aborted; margin of lamina entire, long ciliate. 7. D. mollis
7b. Apex of foliage fronds aborted; margin of lamina serrute or sinuate, not ciliate, or with short hairs at the sinus.
8a. Base fronds oblong-lanceolate, 5--15 x 3--6 cm; foliage fronds 7--12 cm broad; pinnae 0.5--1.2cm broad, apex of lower pinnae rounded, of upper pinnae acute. 8. D. sinica
8b. Base fronds ovate, 6--14 x 4--9 cm; foliage fronds 12--20 cm broad; pinnae 1.2--2 cm broad, apex of pinnae acuminate. 9. D. delavayi
1．Drynaria rigidula (Sw.) Bedd., Ferns Brit. Ind. t. 314. 1869.
硬叶槲蕨 ying ye hu jue
Polypodium rigidulum Sw., J. Bot. 1800(2): 26. 1801.
Rhizome 1--2 cm thick, short-creeping; scales brown to dark brown, with a lighter margin, spreading, peltate, 5--13 x 0.5--1.5 mm, apex acute to acuminate, margin ciliate; fronds dimorphic, rachises persistent; base fronds 10--30 x 5--15 cm, lobed from 1/3 to 4/5, margin irregularly and finely denticulate; foliage fronds stalked, stipes to 40 cm long, not winged, with two rows of nectarines; laminae pinnate, 25--100(--200) x 12--50 cm, apex aborted; pinnae articulate to rachis, all equally long or smaller towards the apex, 8--25(--30) x 0.5--1.5 cm, base narrowly cuneate, margin crenate to serrate, apex obtuse to acuminate, free veinlets simple or absent, hydathodes absent. Sori in one row between costa and margin, costal, singly between the veins, round, sunken. Spores with short, blunt spines and globules.
Epiphytic, encircling the trunk many times, rarely epilithic. Alt. 0--2000(--2400) m. Hainan and SW Yunnan [Indochina, Myanmar, Thailand, Malesia, Polynesia and tropical Australia].
This species is different from the other species of Drynaria by pinnate fronds. It is classified as the sole member of Sect. Poronema to show its isolated position appropriately.
2. Drynaria bonii Christ in Not. Syst. [Paris] 1: 186. 1909.
团叶槲蕨 tuan ye hu jue
Rhizome terete to dorsiventrally flattened, 0.3 cm thick, 1--2 cm wide, short-creeping; scales squarrose or spreading form an appressed base, peltate, 2--12 x 1.5--3 mm, acumen often subulate, suddenly contracted from a wide base, often with a distinct, brown midrib, margin irregularly and shortly dentate at base, nearly entire in the acumen; fronds dimorphic, glabrous; base fronds contiguous or overlapping, often completely covering and surrounding the rhizome, sessile, (4--) 10--15 x (3.5--) 8--12 cm, naerly entire to shallowly lobed; foliage fronds stalked, stipes up to 20 cm long, conspicuously winged with a sinuose wing; laminae pinnatifid to ca. 1 mm from costa, 30--70 x 20--30 cm, apex aborted; pinnae 3--7 pairs, slighyly ascending, 1.5--3 cm distant, broadly lanceolate, 7--20 x 2.5--5 cm, apex obtuse to long acuminate, base slightly narrowed and decurent, margin subentire to shallowly sinuate. Sori in 2 irregular rows between costa and margin, 2--4 rows between lateral veins. Spores with spines.
Epiphytic or epilithic on dry or muddy rocks in deciduous forest at low to medium altitudes, alt. 100--1300 (--1700) m, fairly common on limestone rocks. Guangdong, Guangxi, Guizhou, Hainan, Yunan [India, Indochina, Thailand, and Malaysia].
Some times this species was wrongly identified as D. sparsisora (Desv.) T. Moore, a species in Southeast Aisa and Australia, similar, differs in base fronds lobed up to 1/3, rhizome scales with a very long acumen.
3. Drynaria roosii Nakaike, New Fl. Jap. Pterid. revised et enlarged 841, f.882. 1992.
槲蕨 hu jue
Polypodium fortunei Kunze ex Mett., Farngatt. 1, Polyp. 121, t. 3: 42-45. 1857; Drynaria fortunei (Kunze ex Mett.) J. Sm. in Seem., Bot. Voy. Herald [Fl. Hongkong] 425. 1857. non T. Moore in Gard. Chr. 708. 1855.
Rhizome 1--2 cm thick, short-creeping; scales peltate, 7--12 x 0.8--1.5 mm, margin dentate; fronds dimorphic, glabrous; base fronds overlapping, sessile, (2--) 5--9 x (2--) 3--7 cm, base cordate, shallowly lobed to 1/3, sparsely hairy benath; foliage fronds stalked, stipes 4--7(--13) cm long, conspicuously winged; laminae pinnatifid to 2--5 mm from costa, 20--45 x 10--15(--20) cm, apex aborted or not; pinnae 7--13 pairs, slighyly ascending, lanceolate, 6--10 x (1.5--) 2--3 cm, apex obtuse or acute, margin obscurely dentate. Sori in 2--4 rows between costa and margin, one row between lateral veins, paraphyses many, glandular. Spores with globules.
Epiphytic or epilithic, or on buildings, alt. 100--1800 m. Often on limestone rocks. Anhui, Chongqing, Fujian, Jiangsu, Jiangxi, Hong Kong, Hubei, Hunan, Guizhou, Guangdong, Guangxi, Sichuan and Yunnan [Indochina, N. Thailand, India (Assam)].
Rhizome is used as medicine herb, also known as a kind of Gu sui bu, the Davallia plants.
4. Drynaria quercifolia (L.) J. Sm. in J. Bot. (Hook.) 3: 398. 1841.
栎叶槲蕨 le ye hu jue
Polypodium quercifolium L., Sp. Pl. 2: 1087. 1753.
Rhizome 2--3 cm thick or more, short-creeping; scales brown-blackish, spreading, linear, 6--20 x 0.5--1 mm, pseudopeltate or peltate, towards the apex strongly dentate, apex long, narrow, acute; fronds dimorphic; base fronds sessile, 15--50 x 10--30 cm, shallowly lobed; foliage fronds stalked, stipes up to 30 cm, not or inconspicuously winged; laminae pinnatifid to 2--5 mm from costa, 40--100 x ca. 40 cm, apex aborted; pinnae broadly lanceolate, 15--25 x 2--3.5 cm, apex acute or acuminate. Sori in 2 rows between costa and margin, regular or irregular in two rows between lateral veins, slightly sunken. Spores with acuminate spines.
Epiphytic, spirally climbing, occasionaly epilithic or terrestrial, in various typies of primary and secondary forest, plantations. Altitude from sea level to about 1000 m. Hainan [Widely ranges from Sri Lanka, East Himalayas, Indochina to South East Asia and Australia].
5. Drynaria parishii (Bedd.) Bedd., Ferns S. Ind. Suppl. 24. 1876.
小槲蕨 xiao hu jue
Pleopeltis parishii Bedd., Ferns Brit. Ind. t. 125. 1866.
Rhizome terete 1 cm thick, rather long-creeping; scales appressed, peltate, 4--7 x 1--1.8 mm, grey-brown, dark in the middle, margin grey color, long-ciliate; fronds monomorphic, glabrous; base fronds absent; foliage fronds stalked, stipes5--10 cm long, conspicuously winged to the very base; laminae pinnatifid to ca. 2mm from costa, ca. 25 x 20 cm, apex aborted; pinnae 5--9(--11) pairs, spreading or slightly ascending, oblanceolate, 4--10 x 1--1.5(--2) cm, upper 1/3 the broadest, apex acute or acuminate, base towards base gradually narrowed, margin subentire; venation prominent, 2-5 rows areolae between to lateral veins, with free included veinlets. Sori in one row between costa and margin, close to costa, only one sorus between lateral veins, obviosuly sunken and prominent on upper surface.
Epiphytic or epilithic, often on trunks in moss forest, alt. 500--1600m. S. and SE. Yunnan [Indochina, Myanmar, and Thailand].
6. Drynaria propinqua (Wall. ex Mett.) J. Sm. ex Bedd., Ferns Brit. Ind. 160. 1866.
石莲姜槲蕨 shi liang jiang hu jue
Polypodium propinquum Wall., List 293. 1828. nom. nud.; Mett., Farngatt. 1. Polyp. 120. t. 3, f.50. 1857.
Rhizome terete 1--2 cm thick, rather long-creeping; scales appressed, peltate, 3--6 x 1--1.5 mm, brown, margin dentate; fronds dimorphic, glabrous; base fronds rounded or ovate, 10--20 x 7--18 cm, pinnatifid to 2/3 or more, margine irregularly dentate; foliage fronds stalked, stipes 8--20(--25) cm long, slightly winged; laminae pinnatifid to ca. 2mm from costa, (20--)30--50(--60) x (12--)20-30 cm, apex not aborted; pinnae 7--15 pairs, apex acute, margin dentate. Sori in one regular row between costa and margin, close to costa. Spores with verrucate ans spines.
Epiphytic or epilithic, alt. 500--1900(--2800) m. Guizhou, Guangxia, Sichuan, Xizang and Yunnan [Indochina, Myanmar, Thailand, N. Inida, Sikkim, Nepal, and Bhutan].
7. Drynaria mollis Bedd., Ferns Brit. Ind. t. 216. 1866.
毛槲蕨 mao hu jue
Drynaria costulisora Ching et S.K. Wu; Drynaria tibetica Ching et S.K. Wu
Rhizome 0.5--1(--2) cm thick, creeping; scales soft, tuft, basifixed, 5--11 x 0.5--1.2 mm, margin biserrate; fronds dimorphic, hairy; base fronds sessile, 7--15 x 3--7 cm, base cordate, auriculed, pinnatilobed to 2/3 or more, lobes 8--13 pairs, margin entire, ciliate at young stage, glabrous on both sides, lower side of costa slightly hairy and with small scales at base; foliage fronds stalked, stipes 1--3(--13) cm long, conspicuously winged; laminae pinnatifid to 0--2 mm from costa, 20--40(--50) x 7--12(--15) cm, apex not aborted; pinnae 15--18 pairs, spreading, lanceolate, 3--8 x 1--1.5(--2) cm, apex obtuse, margin entire, densely ciliate; under side along costa with samll scales. Sori in one stright row between costa and margin, one row between lateral veins. Spores with spines.
Epilithic on limstone rocks in Quercus forest, or on tree trunks in mixed fores, alt. 2700-3400 m. Xizang and NW. Yunnan [N. Inida, Sikkim, Nepal, and Bhutan].
8. Drynaria sinica Diels in Engl., Bot. Jahrb. 29(1): 208. 1900.
秦岭槲蕨 qin ling hu jue
Drynaria baronii (Christ) Diels; Drynaria sinica var. intermedia Ching et S.K. Wu
Rhizome 1--2 cm thick, with many old persist stipes, creeping; scales soft, tuft, basifixed, 4--11 x 0.5--1.5 mm, base auriculate, margin biserrate; fronds dimorphic, or sometimes base fronds absent; base fronds sessile or with very short stipe, oblong-lanceolate, 5--15 x 3--6 cm, base narrowed, not cordate, not auriculed, pinnatilobed to 2/3 or more, lobes 10--12(--20) pairs, margin slightly dentate; foliage fronds stalked, stipes 2--10 cm long, conspicuously winged; laminae pinnatifid, 22--50 x 7--12 cm, apex often aborted; pinnae 16--25(--30) pairs, spreading, lanceolate, 4--7 x 0.5--1.2 cm, apex acute, margin dentate, glabrous or sparesely ciliate; laminae sparsely hairy on both surfaces, especially along costa and veins. Sori in one stright row between costa and margin, close to costa, one row between lateral veins. Spores with verrucate and spines.
Often terrestrial, or epilithic, rarely epiphytic on tree trunks, alt. 1280--3800 m. Gansu, Qinghai, Shaanxi, Shanxi, Sichuan, E. Xizang and NW. Yunnan. Endemic to China.
9. Drynaria delavayi Christ, Bull. Soc. Bot. Fr. [ser 4(5)] 52. Mem. 1: 22. 1905.
川滇槲蕨 chuan dian hu jue
Rhizome 1--2 cm thick, with many old persist stipes, creeping; scales soft, tuft, basifixed, 4--10 x 0.5--1 mm, base auriculate, margin biserrate; fronds dimorphic; base fronds sessile, ovate to oblong, 613(--17) x 4--10 cm, base auriculate, pinnatilobed to 2/3 or more, lobes 5--7 pairs, margin slightly dentate; foliage fronds stalked, stipes 3--9 cm long, winged; laminae pinnatifid, 25--45 x 12--18 cm, apex aborted or not; pinnae 7--13(--17) pairs, spreading, lanceolate, (5--)7.5--12(--14) x 1.5--2(--3.5) cm, apex acute, margin shallowly dentate, glabrous or with few hairs; laminae sparsely hairy on both surfaces, especially along costa and veins. Sori in one stright row between costa and margin, close to costa, one row between lateral veins. Sporangia often with glandulars. Spores with verrucate.
Epiphytic or epilithic, alt. 1000--1900(--2800--3800--4200) m. S. Gansu, Qinghai, Shaanxi, Sichuan, E. Xizang and NW. Yunnan [Bhutan, Myanmar].