LOXOGRAMMACEAE [FIRST DRAFT]
剑蕨科 jian jue ke
Zhang Xianchun (张宪春)[1];
1. LOXOGRAMME (Blume) C. Presl, Tent. Pterid. 214, pl. 9, f. 8. 1836.
剑蕨属 jian jue shu
Rhizome very short to long-creeping, occasionally branching, roots forming a spongy mass; scales clathrate, orange-brown to blackish, concolorous, entire, elongate, basifixed. Articulation between phyllopodium and frond none, or evident but not functional, or functional. Entire plant, except for the roots, lacking sclerenchyma. Laminae simple, monomorphic to dimorphic, linear, narrowly elliptic, oblanceolate, spathulate to orbicular, entire, thin- to thick-papyraceous; margin not cartilaginous, drying revolute or involute. Laminar surface glabrous except for minute 2-celled clavate glandular hairs. Veins regularly anastomosing with numerous, few, or without free included veinlets; hydathodes lacking. Sori exindusiate, elongate oblique or nearly parallel to costa, discrete, paraphyses muticellular, hair-like. Sporangial stalk 1- or 2-celled at base, annulus with 12-16 indurated cells. Spores greenish (at time of dispersal), globose-trilete, or ellipsoida-monolete, surface finely verrucose. Chromosome number n = 35, 36.
About 33 species, pantropical, mainly in tropical Asia, one in Central America, one in Pacific Islands, 4 in Africa. 12 in China, south of the Tsinling ranges. Epiphytic or epilithic in wet forests, Holttum (1955) suggests the lack of articulation causes the fleshy nature of fronds, the fronds curled up in dry weather.
Key to species
1a. Plants less than 1 cm long; fronds dimorphic, sterile fronds suborbicular or obovate, 0.4-1 cm long .. 1. L. lankokiensis
1b. Plants more than 1 cm long; fronds monomorphic, or subdimorphic, sometimes dimorphic.
2a. Fronds 3-10 cm long, rarely over 15 cm; spores globose-trilete.
3a. Laminae spatulate, oblanceolate, or linear-lanceolate, 5-8 cm long. 2. L. grammitoides
3b. Laminae lanceolate or oblanceolate.
4a. Rhizome slender, long creeping; scales brown.
5a. Laminae 5-10(-20) x 0.5-1.0 cm. . 3. L. chinensis
5b. Laminae 3-6 x 0.3-0.4 cm. .. 4. L. acroscopa
4b. Rhizome short, thick; scales dark brown or black. 5. L. assimilis
2b. Fronds 20-35 cm long; spores ellipsoidal-monolete.
6a. Rhizome short, thick, erect or creeping; fronds close or in tuft, subsessile.
7a. Scales thick, somewhat opaque, cells small, dense, cell wall thin; without paraphyses. 6. L. formosana
7b. Scales thin, transparent, cells bigger; paraphyses many, dense.
8a. Costae distinctly raised on upper surface; scales dark brown, 5-10 x 0.5-1.2 mm. 7. L. avenia
8b. Costae not so distinctly raised on upper surface; scales greyish brown. ca. 7 x 4 mm. 8. L. involute
6b. Rhizome long creeping, slender; fronds distant.
9a. Base of stipe not shining purplish dark brown or black.
10a. Stipes short or not distinct.
11a. Paraphyses few, sparse. 9. L. cuspidate
11b. Paraphyses many, dense. .. 10. L. porcata
10b. Stipes long. . 11. L. salicifolia
9b. Base of stipe purplish dark brown or black, shining. .. 12. L. duclouxii
1. Loxogramme lankokiensis (Rosenst.) C. Chr., Ind. Fil. Suppl. 3: 125. 1934.
老街剑蕨 lao jie jian jue
Polypodium lankokiense Rosenst. in Med. Rijks Herb. 5: 31. 1917.
Rhizome long-creeping, slender, diam. ca. 0.5 mm, densely scaly throughout; scales dark brown, thin, distinctly clathrate, lanceolate, ca. 2 x 0.5 mm, margin entire. Fronds distant, dimorphic; laminae of sterile fronds suborbicular or obovate, length and width about same, 0.4 -- 1.0 cm long and wide, with a short ca. 2 mm long stipe, carnosa, entire at margin; laminae of fertile fronds oblanceolate, 1--3 cm long, widest at top, ca. 5 mm wide, apex obtusely acute, narrowed below, base decurrent to 2 mm from base of stipe; costae distinct on lower half; veins quit invisible; margin of lamina a little involved. Sori oblong, oblique, slightly sunken into lamina, 2-3 pairs on upper part of lamina, without paraphyses. Spores globose-trilete,.
Epilithic on mossy rocks in valley of forests, alt. 900-1400m. Guangdong, Yunnan and SE. Xizang [Indochina, N. Thailand].
This is one of the smallest plants in the genus, which has dimorphic fronds. A similar species, L. conferta Copel. distributed in the Philippines and Borneo, but differentent in scale characters.
2. Loxogramme grammitoides (Baker) C. Chr., Ind. Fil. Suppl. 2: 21. 1917.
匙叶剑蕨 chi ye jian jue
Gymnogramma grammitoides Baker in J. Bot. [London] 27: 178. 1889.
Loxogramme yigongensis Ching et S.K.Wu in C.Y. Wu, Fl. Xizangica 1: 351, f.94: 4-6. 1983.
Rhizome long-creeping, slender, densely scaly throughout; scales dark brown, clathrate, lanceolate, margin slightly dentate. Fronds close or distant; stipes short or indistinct, green; laminae spatulate, oblanceolate, or linear-lanceolate, 5--8 x 0.5-1 cm, apex acute or obtuse, base decurrent; costae raised on both sides, veins hidden, areolae narrow, oblique, without free veinlets; laminae papyraceous, glabrous. Sori oblong, oblique, parallel to costa in narrow frond form, slightly sunken into lamina, 2-5 paris on upper part of lamina, without paraphyses. Spores globose-trilete,.
Epilithic or epiphytic in evergreen broad-leaved forests. Anhui, Chongqing, Fujian, Gansu, Guizhou, Henan, Hubei, Hunan, Jiangxi, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan and Zhejiang [Japan].
3. Loxogramme chinensis Ching in Sinensia 1: 13. 1929.
中华剑蕨 zhong hua jian jue
Loxogramme confertifolia Tagawa in Acta Phytotax. Geobot. 8: 232. 1939; W.C. Shieh et al., Fl. Taiwan 1: 496. 1994.
Loxogramme fujiansis Ching in Fl. Fujianica 1: 254, f. 241, 602. 1982.
Rhizome long-creeping, slender, diam. 1-1.5 mm, densely scaly throughout; scales pale brown to dark brown, clathrate, lanceolate, (2-) 3-5 mm long, ca. 1 mm wide at base, margin subentire. Fronds close or distant; stipes short, densely scaly at base, pale green, narrowly winged to the very base; laminae spatulate, oblanceolate, or linear-lanceolate, 5-12 (-15-20) x 0.5-1.2 cm, apex acute, base decurrent, margin slightly undulate and involute when dry; thick papyraceous, glabrous, yellow green; costae raised on both sides, veins hidden. Sori oblong, very oblique, nearly parallel to costa, superficial, 5-8 pairs from middle to upper part of lamina, without paraphyses. Spores globose-trilete.
Epilithic on mossy rocks or epiphytic in evergreen broad-leaved forests. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan and Zhejiang [Bhutan, India, Nepal, Myanmar, Thailand, Vietnam].
4．Loxogramme acroscopa C. Chr. in Dansk Bot. Ark. 6: 48. 1929.
顶生剑蕨 ding sheng jian jue
Polypodium acroscopum Christ in J. de Bot. 9: 75. 1905.
Rhizome creeping, slender; scales pale brown, lanceolate, cells length longer than width, entire. Fronds distant, monomorphic; stipes slender, 2-3 cm long; laminae linear-lanceolate, 4-6 x 0.3-0.4 cm, apex obtusely acute, base decurrent to a winged stipe; costae raised on upper surface, veins hidden; texture thin. Sori linear, 3-5 pairs, onlique, nearly parallel to costa, without paraphyses. Spores globose-trilete.
On wet rocks by streams, may be a seasonal rheophyte, altitude 250-500 m. Similar to L. chinensis in characters of scales and spores, might be an ecological form of the latter. SE. Yunnan, S. Guizhou [N. Vietnam].
5. Loxogramme assimilis Ching in Bull. Dept. Biol. Coll. Sci. Sun Yatsen Univ. 6: 31. 1933.
黑鳞剑蕨 hei lin jian jue
Rhizome short to moderately creeping, densely scaly throughout; scales dark brown or black, linear-lanceolate, ca. 5 x 2 mm, entire. Fronds close or distant; stipes short or subsessile, densely scaly at base, pale green, narrowly winged to the very base; laminae linear-lanceolate, 10-15(-25) x 1.0-2.5 cm, widest at middle, apex acuminate or acute, base decurrent; thick papyraceous, glabrous on both surfaces, upper surface deep green, lower surface yellow green; costae not distinctly raised on both sides; veins hidden. Sori oblong, oblique, close to margin of frond than to costa, superficial, without paraphyses. Spores globose-trilete.
Epilithic on mossy rocks or epiphytic in evergreen broad-leaved forests, alt. 600-2200 m. Chongqing, Guangxi, Guizhou, Sichuan, and SE Yunnan [N. ietnam].
6. Loxogramme formosana Nakai in Bot. Mag. Tokyo 43: 8. 1929.
台湾剑蕨 taiwan jian jue
Loxogramme ensiformis Ching, Icon. Fil. Sin. 4: t. 191. 1937.
Loxogramme grandis Ching et Z.Y. Liu in Bull. Bot. Res. [Harbin] 4(4): 221, f. 51. 1984. non Copel. 1908.
Rhizome short, erect, densely scaly; scales pale brown, late-ovate, ca. 5 x 2.5 mm, acuminate, entire, cells small, dense. Fronds in tuft; stipes short, thick, 1-3 cm long, flattened, base shining brown; laminae oblanceolate, 20-35 x 3-3.5 cm, widest at upper 2/3, apex acuminate, base decurrent on stipe, coriaceous, thick and fleshy, deep green on upper surface, paler beneath, glabrous; costae slightly raised on both surfaces; veins invisible, anastomosing to form elongate areoles with included veinlets. Sori confined to the upper half of the fronds, oblique, rather close to the costa, more or less sunken into lamina, without paraphyses. Spores ellipsoidal-monolete.
Epilithic on mossy rocks or epiphytic in evergreen broad-leaved forests. Chongqing, Guizhou, Sichuan, Taiwan, and S. Yunnan. Endemic to China.
7. Loxogramme avenia (Blume) C. Presl, Tent. Pterid. 215. 1836.
剑蕨 jian jue
Grammitis avenia Blume, Enum. Pl. Jav. 117. 1828.
Rhizome short creeping, diam. more than 3 mm, densely covered with scales near apex and at base of stipes; scales dark brown, lanceolate, ca. 5-10 x 0.5-1.2 mm, entire. Fronds in tuft; stipes indistinct, dark castaneous when dry; laminae lanceolate, 15-25 x 1-2.5 cm or more bigger, acuminate at apex, broadest at middle to upper portion, narrowed below, base decurrent to 2-5 mm from base of stipe, margin recurved, green on upper surface, paler beneath; costae distinctly raised on upper surface, hardly beneath, pale green to brown; veins hardly visible on both surfaces, forming copious anastomosis; texture thick, leathery, fleshy; glabrous on upper surface, lower surface with sparse brown, 2-celles glandular hairs. Sori linear, oblique, 8-20 pairs, 1-2 cm long, continuous in middle between costa and the margin of frond, with about 3-4 mm between each, slightly sunken into lamina; paraphyses many, dense, longer than sporangia, completely covered the sori at early satge. Spores ellipsoidal-monolete.
Epilithic on mossy rocks or epiphytic on tree trunks in dense evergreen forests by streams at low altitude. Yunnan (Tengchong) [Myanmar, Thailand, Indochina, Java, W. Malesia].
Very similar to L. involuta and L. cuspidata, different in the scales smaller, dark brown; different from the letter in paraphyses abundant.
8. Loxogramme involuta (D. Don) C. Presl, Tent. Pterid. 215. 1836.
内卷剑蕨 nei juan jian jue
Grammitis involuta D. Don, Prodr. Fl. Nepal 14. 1825.
Rhizome short, ascending to creeping, diam. ca. 1.5 mm, densely covered with scales; scales subdeltoid, apex acuminate, up to 7 x 4 mm, thin, entire, grayish brown, cells dense, length and width almost same. Fronds in a tuft near apex of rhizome; stipes indistinct, or very short with wings; laminae lanceolate, up to 35 x 4 cm, apex caudately long acuminate, attenuate at base and decurrent to narrow wings of stipe nealy to the base, deep green on upper surface, paler beneath; costae more or less raised beneath, usually flat on upper surface, stramineous or pale green; veins all hidden, anastomosing with free included veinlets; lamianae curled up when dry, texture thick and fleshy, glabrous on both surfaces. Sori linear, very oblique, well spaced, 3-4 cm long, continuous from near costa to margin of frond, superficial; paraphyses many, dense, longer than sporangia. Spores ellipsoidal-monolete.
Epilithic on mossy rocks or epiphytic on tree trunks in evergreen broad-leaved forests, alt. 2000-2500 m. W. Yunnan, and S. Xizang [Nepal, Sikkim, NE. India, N. Thailand].
9. Loxogramme cuspidata (Zenker) M.G. Price in Amer. Fern J. 74(2): 61. 1984.
西藏剑蕨 xizang jian jue
Grammitis cuspidata Zenker, Plantae Indicae 1: t. 2. 1835.
Loxogramme tibetica Ching et S.K. Wu in C.Y. Wu, Fl. Xizangica 1: 353, f. 95: 4-5, pl. 12, f. 5-6. 1983.
Rhizome short to moderately creeping, densely covered with scales; scales lanceolate, dark brown, apex long acuminate, entire, cells dense, length longer than width. Fronds close; stipes winged; laminae lanceolate, 35-40 x 3 cm, widest at upper 1/3, apex acuminate or caudate, narrowed from middle downwards, attenuate at base and decurrent to narrow wings of stipe nearly to the base, green on upper surface, paler beneath; costae raised on upper surfaces, distinct but not raised beneath, stramineous or pale green; veins hidden, anastomosing with free included veinlets; lamianae curled up when dry, texture thick and fleshy, glabrous on both surfaces. Sori linear, oblique, up to 2.5 cm long, continuous from near costa to margin of frond, superficial; paraphyses few, sparse. Spores ellipsoidal-monolete.
Epilithic on mossy rocks or epiphytic on tree trunks in evergreen broad-leaved forests, alt. 2000-3500 m. Sichuan, Yunnan, and Xizang [India].
Very similar to L. involuta, scales smaller, colour, texture and cell size of scales are also different.
10. Loxogramme porcata M.G. Price in Amer. Fern J. 80(1): 6，f. 3. 1990.
拟内卷剑蕨 ni nei juan jian jue
Rhizome short creeping, densely covered with scales; scales linear-lanceolate, apex acuminate, 5-10 x 0.5-1.2 mm, thin, entire, grayish brown to dark brown, cells narrow, length longer than width. Fronds close; stipes indistinct, or very short with wings; laminae narrow elliptic to oblanceolate, 10-60 x 0.8-6 cm, apex long acuminate, attenuate at base and decurrent to narrow wings of stipe nealy to the base, deep green on upper surface, paler beneath; costae raised beneath, flat on upper surface, stramineous or pale green; veins hidden, anastomosing with free included veinlets; lamianae curled up when dry, texture thick and fleshy, glabrous on both surfaces. Sori linear, very oblique, well spaced, up to 6 cm long, continuous from near costa to margin of frond, superficial; paraphyses many, dense, soft, longer than sporangia, completely covered the sori at early satge. Spores ellipsoidal-monolete.
Epiphytic on tree trunks in evergreen broad-leaved forests, alt. 1200-1750 m. S. & W. Yunnan, and SE. Xizang [Myanmar, N. Thailand, Nepal, Sikkim, India].
Very similar to the sympatric L. involute, both develop abundant paraphyses, but different in scales smaller, the cells of scales narrow and long.
11. Loxogramme salicifolia (Makino) Makino in Bot. Mag. Tokyo 19: 138. 1905.
柳叶剑蕨 liu ye jian jue
Gymnogramme salicifolia Makino, Phan. et Pterid. Jap. Icon. Ill. pl. 34. 1899.
Loxogramme biformis Tagawa in Acta Phytotax. Geobot. 3: 93. 1934.
Rhizome long creeping, diam. ca. 2 mm, rather densely scaly; scales ovate-lanceolate, apex acuminate, entire, brown or reddish brown. Fronds subdimorphic or obviously dimorphic, distant, 15-35 cm long; stipes greenish yellow to paler, 2-5 (-10) cm long, or subsessile; laminae narrowly oblanceolate to linear, 15-20 (-32) x 1-1.5 (-3) cm, acuminate at apex, decurrent at base to form wings of upper portion of stipes; laminae texture relatively thin, fleshy; costae raised on upper surface, plane below; veins hidden, anastomosis without or with few included veinlets. Sori on upper portion of fronds, very oblique to nearly parallel to the costa, up to 10 or more pairs, 1-3 cm long, in middle between costa and margin of frond, slightly sunken into lamina, without paraphyses. Spores ellipsoidal-monolete.
Epilithic on mossy rocks or epiphytic on tree trunks in forests, alt. 200-1200 m. Anhui, Chongqing, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, and Zhejiang [Korea (Cheju Island), Japan].
Similar to the sympatric species, L. duclouxii, different in the stipe not shining black, and sori line not very close to costa.
This species is almost sympatric with L. duclouxii, but not found in Yunnan. The report by Christensen (1931) from Yunnan should be L. cuspidata; and the recent report by Chen (2005) from Yunnan might be confused with L. duclouxii or L. cuspidata. L. salicifolia is a low elevation species not found in the high plateau of Yunnan and Xizang.
12. Loxogramme duclouxii Chirst in Bull. Acad. Geogr. Bot. 16: 140. 1907.
褐柄剑蕨 he bing jian jue
Loxogramme remote-frondigera Hayata, Icon. Pl. Formosa 5: 323. 1915.
Loxogramme saziran Tagawa ex M.G. Price in Amer. Fern J. 80(1): 7, f. 4. 1990.
Rhizome long-creeping, diam. 1-1.6 mm, dark brown to black, sometime smooth when scales caducous; scales on rhizome dark brown to black, distinctly clathrate, subulate or ovate-lanceolate, 1-1.8 x 0.5-0.7 mm, entire; scales at base of stipe 3-4 x 0.9-1.6 mm, ovate, acute. Fronds subdimorphic, or sometimes obvious dimorphic, close or distant, stipes on obvious phyllopodia; phyllopodia 1-2 mm long; stipes up to 7 cm long, shining purplish dark brown or black when dry; laminae linear-oblanceolate, 10-35 x 1.5-2.5 (-3.5) cm, apex acuminate or caudate, base decurrent on stipe, coriaceous, thick and fleshy, deep green on upper surface, paler beneath, glabrous, margin involute when dry; costae raised on upper surface, a little distinct and flattened beneath, pale green; veins invisible, anastomosing to form elongate areoles usually with included veinlets. Sori confined to the upper half of the fronds, 10 or more pairs, usually close together, oblique, rather close to the costa, more or less sunken into lamina, without paraphyses, or with few short paraphyses, shorter than sporangia. Spores ellipsoidal-monolete.
Epilithic on mossy rocks or epiphytic in evergreen broad-leaved forests, alt. 800-2500 m. Anhui, Chongqing, Gansu, Shaanxi, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan and Zhejiang [NE. India, Thailand, N. Vietnam, Korea (Cheju Island), Japan].
Similar to the sympatric species, L. salicifolia, different in the peculiar shining black stipe and sori line rather close to costa.
[1] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, Peoples Republic of China.