阴地蕨科 yin di jue ke
Zhang Xian-Chun; N. Sahashi
Plants perennial, terrestrial or in rocky crevices, evergreen or winter green. Rhizomes short, erect, forming caudex up to 5 mm in diam., dictyostelic; gemmae absent or minute, spherical, apex hairy; rhizome [glabrous (subgenus Botrychium) or] with hairs light brown, long, uniseriate, non-septate. Roots occasionally laterally branching, yellowish to black, 0.52 mm diam., smooth or with corky ridges, not proliferous. Fronds 1(or 2) per stem, with common stipe divided into sterile, laminate, photosynthetic portion (sterile blade) and fertile, spore-bearing portion (fertile blade). Common stipe thick, fleshy, glabrous or hairy. Sterile blades ascending or perpendicular to common stipe, sessile or stalked; blade simple to 5-pinnate, linear, oblong, or deltoid, 425 Χ 135 cm. Pinnae (reduced to segments in many species) spreading to ascending, fan-shaped to lanceolate or linear, margins entire to dentate or lacerate, apex rounded or acute; veins free, arranged like ribs of fan or pinnate. Fertile blades normally 1 per frond, 13-pinnate, long-stalked, borne at ground level to high on common stipe. Sporangia sessile to shortly stalked, almost completely exposed, large, globose, glabrous, dehiscing by two valves, borne in 2 rows on pinnate (except in very small plants) fertile blade branches. Spores tetrahedral, trilete, thickly walled, surface rugate, tuberculate, baculate, sometimes joined in delicate network, with a large number of spores per sporangium. Gametophytes not green, broadly ovate, unbranched, subterranean, mycorrhizal. x = 44, 45, 92.
Fifty to sixty species: nearly worldwide; 12 species in China.
Once separated into four genera: Botrychium, Botrypus, Sceptridium and Japanobotrychium; here, all the species are treated in one single genus.
[Don Farrars review: Hauk et al. (2003), using rbcL markers found genetic differences between the Genera of Kato (1987) and subgenera of Clausen (1938) to be supportive of recognizing these taxa as separate genera. In North America I have accepted Hauks conclusion in recognizing Botrychium, Botrypus and Sceptridium as distinct genera. It is the prerogative of the authors to choose to recognize these entities as a single genus, but in the Flora, there is something to be said for treating this group similarly to those of Lycopodium, Equisetum, etc.]
阴地蕨属yin di jue shu
Description and distribution as for the family.
Classification mainly follows Zou and Wagner (1988) [??full citation]. [Don Farrars review: Many additional species in subgenus Botrychium have been described since the Zou and Wagner 1988 treatment, and also since Wagners treatment in Flora of North America. In subgenus Botrychium we are now recognizing 14 diploid species (n = 45), 16 tetraploid species (n = 90) and 1 hexaploid species (n = 135) in North America plus 1 additional diploid and 1 additional tetraploid in Europe for a total of 32 species worldwide. With the number of species in subgenus Botrychium, the total number of species in the broad genus as recognized by the authors needs to be recalculated.]
[Don Farrars review: Given the large number of species of moonworts (subgenus Botrychium) in North America, it is surprising that only 3 species have been reported for China. The authors may want to point this out and encourage field workers to seek additional species of moonworts. I would be pleased to look at photographs, scans or specimens for comparison to North American taxa.]
[FRPS and Fl. Liaoning: ??Botrychium multifidum (Gmelin) Ruprecht; Osmunda multifida Gmelin; Botrychium matricarae Sprengel; Osmunda matricarae Schrank]
[FRPS: ??Botrychium longipedunculatum Ching]
[Fl. Xizang: ??Botrypus tibeticus Ching]
1a. Plants small, 315 cm; frond primordium glabrous; sterile blade pinnate or ternate, 1- or 2-pinnate.
2a. Pinnae fan-shaped, pinna midrib absent ......................................................... 1. B. lunaria
2b. Pinnae elongate, pinna midrib present.
3a. Pinnae rounded at apex, often overlapping; primordial blade erect .............. 2. B. boreale
3b. Pinnae pointed at apex, usually well separated; primordial blade pendent 3. B. lanceolatum
1b. Plants medium to large, mostly more than 10 cm; frond primordium usually hairy; sterile blade ternately 24-pinnate.
4a. Fertile blade stalk joined to sterile blade stalk at or above base of sterile blade.
5a. Fertile blade stalk joined to rachis above base of sterile blade; frond very hairy to essentially glabrous 4. B. lanuginosum
5b. Fertile blade stalk joined to base of sterile blade; frond nearly to entirely glabrous.
6a. Ultimate segments contracted at base, deeply divided; sporangia on 2- or 3-pinnate fertile blade; sporangial cluster deltoid .............................................................. 5. B. virginianum
6b. Ultimate segments not contracted, shallowly lobed; sporangia on 1-pinnate fertile blade; sporangial cluster narrowly linear ........................................................................ 6. B. strictum
4b. Fertile blade stalk joined to sterile blade stalk below base of sterile blade.
7a. Plants sparsely hairy; frond primordium with many fine hairs; blade mostly large, 1025 cm.
8a. Fertile blade stalk joined to sterile blade stalk near base of the frond .. 7. B. japonicum
8b. Fertile blade stalk joined to sterile blade stalk 1/32/3 from base to sterile blade.
9a. Sterile blade up to 3-pinnate, papery; fertile blade stalk arising ½2/3 from base to sterile blade 8. B. daucifolium
9b. Sterile blade up to 4-pinnate, herbaceous; fertile blade stalk arising 1/32/3 from base to sterile blade .............................................................................................. 9. B. formosanum
7b. Plants mostly glabrous; frond primordium glabrous or only upper part hairy; blade mostly smaller, 515 cm.
10a. Terminal pinnules large, differentiated from lateral pinnules, 815 Χ 510 mm, elongate, acute at apex, margins finely serrate; fronds mostly turning reddish in winter ....... 10. B. nipponicum
10b. Terminal pinnules smaller, ± conforming with lateral pinnules, 310 Χ 29 mm, rounded to truncate at apex, margins nearly entire to finely dentate; fronds mostly green in winter.
11a. Segments blunt, rounded, or subacute; margins entire to shallowly and coarsely crenate or dentate; sterile blade leathery .................................................................. 11. B. robustum
11b. Segments mostly acute; margins shallowly to deeply denticulate; sterile blade herbaceous 12. B. ternatum
扇羽阴地蕨 shan yu yin di jue
Osmunda lunaria Linnaeus, Sp. Pl. 2: 1064. 1753; Botrypus lunaria Richard.
Rhizomes short, erect, annually producing one frond 515(25) cm tall. Common stipe greenish, cylindrical, 412 cm, 23 mm in diam., hollow, succulent. Sterile blade pinnate, sessile, broadly lanceolate or oblong, 38 Χ 1.52.5 cm, fleshy, leathery, glabrous, apex rounded or blunt; pinnae 46 pairs, 11.5(2) cm long and wide, shortly stalked to almost sessile, approximate, often overlapping, opposite or almost opposite, lowest 1 or 2 pairs 12 cm apart, flabellate (fan-shaped), lunate, or reniform, margin entire, crenate, or incised; veins free, flabellately forked, glabrous. Fertile blade with stalk 47 cm, glabrous; panicle 2- or 3-pinnate, racemose, 36 Χ 1.52 cm, glabrous; sporangia sessile, large, 0.50.1 mm[??reverse] diam. Spores yellowish, surface verrucose. 2n = 90.
Grasslands, meadows, forest understories; 13004000 m. Northeastern to southwestern China [??list provinces: Hebei, Henan, Nei Mongol, Qinghai, Shanxi, Sichuan, Taiwan, Xinjiang, Xizang, Yunnan], also in Taiwan, and Xinjiang [Japan, Korea, Sri Lanka; Asia, Australia, N Europe, Himalaya, North America, Pacific islands].
Throughout the range of Botrychium lunaria, sporadic plants have been found with incised sterile pinnae and which have been named as forms or varieties, such as B. lunaria f. subincisum Roeper, and B. lunaria var. onondagense (Underwood) House.
[Don Farrars reviewIn North America we now recognize 5 taxa in this complex: B. lunaria var. lunaria, B. lunaria var. crenulatum, B. neolunaria, B. tunux and B. yaaxudakeit. All of these except var. crenulatum occur in the Aleutian Islands and thus may also be present in Asia. B. yaaxudakeit is an allotetraploid derived from hybridization between B. lunaria and B. neolunaria. It is very difficult to distinguish morphologically from B. lunaria except by its spore size (46-57 microns) which is considerably larger that the average we obtained for B. lunaria (33-39 microns). I am very curious about the authors report of spore sizes reaching 48 in B. lunaria and wonder if plants having spores of that size might be B. yaaxudakeit.]
北方阴地蕨 bei fang yin di jue
Rhizomessuberect, shortly cylindrical, annually producing one frond 510 cm tall. Common stipe 38 cm. Sterile blade stalk sessile or nearly so; blade 2-pinnate at base, pinnate above, shiny, green, ovate-deltoid, 14 Χ 13 cm, fleshy, base cordate. Pinnae 35 pairs, ascending, mostly overlapping; basal pinnae largest, ovate, up to 2 cm wide, base subtruncate, apex obtuse; upper pinnae and segments of basal pinnae elliptic, up to 5 mm wide; veins free. Fertile blade arising at or near top of common stipe, 2-pinnate, with stalk 1.55 cm. Spore surface verrucose. 2n = 180.
Forests; ca. 1000 m. Nei Mongol [Japan, Korea; N Europe, Greenland].
[Don Farrars review: This species name was initially applied circumboreally to allotetraploid plants having originated through hybridization between B. lunaria and B. lanceolatum. With knowledge of the greater diversity within B. lunaria and B. lanceolatum than previously known, we now recognize three taxa from within B. boreale: B. boreale in Europe, and B. pinnatum and B. alaskense in North America. The American species are quite distinct from B. boreale both morphologically and genetically. Asian plants have not yet been tested genetically but all plants of this complex in Alaska, including the Aleutian Islands, have proven to be either B. alaskense or B. pinnatum. It would be good to genetically analyze the B. boreale plants in China to confirm that they are of that species and not one of the American species.]
长白山阴地蕨 chang bai shan yin di jue
Osmunda lanceolata S. G. Gmelin, Nov. Comm. Acad. Petrop. 12: 516. 1768; Botrychium manshuricum Ching; B. ramosum Wang Wei et al., [nom. invalid, not (Roth) Ascherson (1864)].
Rhizomes shortly cylindrical, annually producing one frond 520 cm tall. Common stipe green, 415 cm. Sterile blade 2-pinnatifid, subternate, medium green to yellow-green, somewhat shiny, broadly ovate-deltoid, 14 Χ 13 cm; pinnae lanceolate, up to 1 cm wide, deeply pinnatifid, apex acute; upper pinnae and segments of basal pinnae oblong, up to 2 cm wide; veins pinnate, free. Fertile blade arising at top of common stipe, narrowly deltoid, 16 cm, 2-pinnate; sporangia exposed and not immersed, mostly approximate or slightly separated and covering midrib of pinnae. Spore surface tuberculose to irregularly verrucose. 2n = 90.
Rocky places; high elevations. Jilin (Changbai Shan), [??Nei Mongol] [Japan; Eurasia, North America].
[Don Farrars review: This species has long been recognized as containing two subspecies: angustisegmentum and lanceolatum. More recently we have recognized a third taxon in this complex, subspecies viride, which is equally as distinct as the other two subspecies are from one another. A publication justifying recognizing all three subspecies at the level of species is in preparation.]
绒毛阴地蕨 rong mao yin di jue
[??Botrychium arisanense Masamune]; Botrychium decurrens Ching; B. modestum Ching; B. parvum Ching; [??B. virginianum Clarke (1880), not Swartz (1801)]; B. virginianum var. lanuginosum Moore; B. yunnanense Ching; Botrypus lanuginosus (Wallich ex Hooker & Greville) Holub; Japanobotrychium lanuginosum (Wallich ex Hooker & Greville) Nishida ex Tagawa [??not in IPNI or Tropicos]; Osmundopteris lanuginosa (Wallich ex Hooker & Greville) Nishida.
Rhizomes erect, short, apex hairy; hairs long, light brown. Common stipes stramineous to light brown, 1230 cm, 36 mm in diam., fleshy, woolly-hairy; hairs sparse, whitish, long. Sterile blade 3- or 4-pinnate, deltoid to subpentagonal, 1445 Χ 1038 cm, thin and herbaceous, degree of pubescence variable; pinnae 58 pairs, alternate, long stalked, 26 cm apart, lowest pair largest, 720 Χ 510 cm; pinnules 610 pairs, alternate, stalked, basiscopic pinnules larger than acroscopic ones, basal pinnule largest; ultimate lobes acute or obtuse; veins free, simple or forked. Fertile blade with stalk 210 cm, hairy, arising at various points between lowest two pairs of pinnae of sterile blade, usually close to base of second lowest pinnae. Spikes 2- or 3-pinnate, paniculate, 712 Χ 57 cm, hairy. Spores yellowish, surface reticulate, finely striate on ridges. 2n = 180, 270, 360.
Evergreen broad-leaved forests; 10003000 m. Guangxi, Guizhou, Hunan, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, India, Indonesia (Java, Sumatra), Malaysia, Myanmar, Nepal, New Guinea, Philippines, Sri Lanka, Thailand, Vietnam].
蕨萁 jue ji
Osmunda virginiana Linnaeus, Sp. Pl. 2: 1064. 1753; Botrypus virginianus (Linnaeus) Holub; Japanobotrychium virginianum (Linnaeus) M. Nishida ex Tagawa.
Rhizomes erect, short, cylindrical, fleshy, having many fleshy roots and annually producing one frond, 2570 cm tall. Sterile blade: stalk 1535 cm, subglabrous; blade subternate, tri- or quadripinnatifid, pale green, broadly pentagonal or subdeltoid, 528 Χ 730 cm, thinly herbaceous, soft, abaxially lanuginose on midrib, base subtruncate or slightly cordate, apex acute or obtuse; pinnae broadly ovate, with basal pinnules small and middle ones large, oblong or ovate-lanceolate: ultimate segments elliptic or oblong, margin deeply lobed or dentate, apex acuminate; veins pinnate, free. Fertile blade arising at top of common stipe; fertile blade stalk 1030 cm; lamina 3- or 4-pinnate, ovate-deltoid, 1020 cm; pinnae stalked; sporangia globose or elliptic, each on tip of very short, sometimes obscure axes. Spore surface coarse and distantly verrucose. 2n = 184.
Forests; 16003200 m. [??Anhui], Chongqing, Gansu, Guizhou, Henan, Hubei, Hunan, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan, Zhejiang [temperate regions within the northern hemisphere; Central and South America; temperate Asia: Himalaya, Japan, Korea, Russia].
劲直阴地蕨 jing zhi yin di jue
Botrypus strictus (Underwood) Holub; Japanobotrychium strictum (Underwood) M. Nishida ex Tagawa [??not in IPNI or Tropicos]; Osmundopteris stricta (Underwood) Nishida.
Plants summer-green. Rhizomes erect, short, cylindrical. Fronds 3070 cm tall. Sterile blade: stalk 1525 cm, up to 1 cm in diam., fleshy; blade subternate, bi- or tripinnatifid, dull green, broadly pentagonal; pinnules sessile, narrowly oblong, base decurrent to costal wing; ultimate segments elliptic, dentate, apex rounded; costae and costules beneath sparsely lanuginose. Fertile blade arising at top of common stipe, as long as sterile blade; lamina bipinnate, spikelike; pinnae short; sporangia ellipsoid-globular, each on tip of short but distinct ultimate axes or veins. Spore surface coarsely verrucose. 2n = 88, ca. 90.
Forests. Chongqing, Gansu, Heilongjiang, Henan, Hubei, Jilin, Liaoning, Nei Mongol, Shaanxi, Sichuan [Japan, Korea].
华东阴地蕨 hua dong yin di jue
Botrychium daucifolium Wallich var. japonicum Prantl, Jahrb. Kφnigl. Bot. Gart. Berlin 3: 340. 1884; Sceptridium japonicum (Prantl) Lyon.
Rhizomes erect, shortly cylindrical, having many fleshy roots and annually producing one frond (15)3050(70) cm tall. Sterile blade: stalk 1226(36) cm, 34 mm in diam., glabrous; blade subternate, tripinnatifid, subpentagonal, 1025 Χ 1525 cm, apex acute; basal pinnae much larger than upper ones, broadly deltoid-elliptic, 815 Χ 58 cm, with stalk 1.53 cm; pinnules broadly lanceolate, gradually narrowed toward acute apex; pinnule segments narrowly elliptic, margin sharply serrate or crenulate, apex acute or acuminate; basal segments fan-shaped, deeply lobed; stalks and costae sparsely lanuginose; veins pinnate, free. Fertile blade arising 26(10) cm above base of common stipe (or much below middle of common stipe), taller than sterile blade, subdeltoid, bipinnate, 2550 cm (stalk 2040 cm, lamina 510 cm); sporangia globular, attached on both sides of axes. Spore surface with dense spinules or papillae forming reticulate pattern with fine granules. 2n = 270.
Forests by streams; ca. 1200 m. [??Anhui], Fujian, Guangdong, Guizhou, Hunan, Jiangsu, Jiangxi, Taiwan, Zhejiang [Japan, Korea].
薄叶阴地蕨 bo ye yin di jue
Sceptridium daucifolium (Wallich ex Hooker & Greville) Lyon; Botrychium officinale Ching; [??B. subcarnosum Wallich ex Beddome]; S. officinale (Ching) Ching & H. S. Kung.
Rhizomes erect, shortly cylindrical, bearing many fleshy roots. Frond 3040 cm tall with stalk 1012 cm. Sterile blade: stalk 78 cm, glabrous; blade bipinnate to tripinnatifid, subpentagonal, 1520 Χ 1624 cm; pinnae 6 or 7 pairs, alternate, shortly stalked; basal pinnae largest, triangular, 1214 Χ 610 cm, bipinnatifid; pinnules 4 or 5 pairs, narrowly ovate to broadly lanceolate, lower basal pinnule largest, up to 8 Χ 3 cm, pinnatifid, ultimate segments sharply serrate, apex acute or acuminate; lamina herbaceous, veins obvious, rachis and costae with sparse, white, long hairs. Fertile blade arising from above middle of common stipe, as long as sterile blade, with stalk 1416 cm, bi-tripinnate, 1012 cm, with long soft hairs. Spore surface with dense papillae often forming reticulate pattern with minute granules. 2n = 180.
Forests, shaded, wet places. Chongqing, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [NE and S India, Indonesia (Sumatra), Japan (Ryukyus), Myanmar, Philippines, Sri Lanka, Vietnam].
华东阴地蕨 hua dong yin di jue
Sceptridium formosanum (Tagawa) Holub.
Rhizomes erect, cylindrical, ca. 2 cm, annually producing 1 or 2 fronds 1560 cm tall. Sterile blade: stalk (4)1335 cm, sparsely lanuginose; blade subternate, tripinnate, deltoid-pentagonal, 835 Χ 840 cm, thickly herbaceous; pinnules broadly lanceolate, apex acute; ultimate segments elliptic, irregularly roughly serrate, apex obtuse or subacute; veins pinnate, free. Fertile blade arising (2)615 cm below top of common stipe (or near middle of common stipe): stalk 720 cm; lamina deltoid, bipinnate, 518 cm; sporangia globular, attached on both lateral sides of axes. Spore surface with dense papillae often uniting in reticulate pattern, with minute granules. 2n = 180.
Guangdong, Guangxi, Guizhou, Jiangxi, Taiwan, Yunnan [Himalaya].
[??vernacular name]
Sceptridium nipponicum (Makino) Holub.
Plants small, winter-green. Rhizomes erect, short, cylindrical, annually producing one frond 2050 cm tall. Sterile blade: stalk 4.519 cm, glabrous; blade subternate, tripinnate, ca. 10 cm long and broad, herbaceous or membranous, reddish in winter; lower pinnae broadly ovate, long stalked, upper ones much narrower; ultimate segments elongate or lanceolate, serrate; veins pinnate, free. Fertile blade arising 1.56 cm above base of common stipe (or distinctly below middle of common stipe), much taller than sterile blade, deciduous after spore dispersal; stalk ca. 40 cm; lamina smaller than sterile blade; sporangia globular, attached on both lateral sides of axes. Spore surface irregularly reticulate with fine granules. 2n = 90.
Forests. Guangxi [Japan, Korea].
粗壮阴地蕨 cu zhuang yin di jue
Botrychium rutifolium A. Braun var. robustum Ruprecht ex Milde, Nova Acta Regiae Soc. Sci. Upsal., ser. 3, 26: 763. 1858; B. multifidum var. robustum C. Christensen ex Hultιn; [??B. obliquum H. Christ (1905), not Muhl. (1810);] B. sutchuanense Ching; [??B. ternatum Komarov (1901), not (Thunberg) Swartz (1801)].
Plants small, hardy. Rhizomes erect, short, cylindrical. Fronds 3540 cm tall. Sterile blade: stalk 27 cm or more, up to 1 cm in diam., fleshy; blade bi- or tripinnatifid, dull green, pentagonal, 611 Χ 711 cm; pinnae 46 pairs, subopposite, lower ones stalked; basal pinnae largest, ovate to triangular, 46 Χ 2.54.5 cm, bipinnate to tripinnatifid; pinnules 3 or 4 pairs, narrowly oblong or lanceolate, lower basal pinnule longest, 2.54 Χ 1.52 cm, ultimate segments ovate or oblong, margin dentate; costae and costules beneath sparsely lanuginose, lateral veins obscure. Fertile blade arising from middle or lower part of common stipe, with longer stalk, up to 1025 cm, lamina 411 cm, bipinnate to tripinnate, spikelike; pinnae short; sporangia ellipsoid-globular, each on tip of short but distinct ultimate axes or veins. Spore reticulate-like on surface, up to 2 μm in diam. 2n = 90.
Forests, grasslands; 10002000 m, sometimes up to 4000 m. Chongqing, Heilongjiang, Jilin, Liaoning, Sichuan, Yunnan [Japan, Korea].
阴地蕨 (pinyin[??])
Osmunda ternata Thunberg, Syst. Veg., ed. 14 (J. A. Murray), 927. 1784; Sceptridium ternatum (Thunberg) Lyon.
Plants small, winter-green. Rhizomes erect, short, annually producing one frond (10)1555 cm tall. Sterile blade: stalk 512 cm or longer, ca. 2 mm in diam.; blade ternate, tri- or quadripinnatifid, subpentagonal, dull green, 510 Χ 812 cm, somewhat thickly herbaceous, glabrous, apex acute; pinnae subdeltoid, basal ones stalked; segments broadly elliptic or broadly ovate, sparsely crenate, apex rounded; veins pinnate, free. Fertile blade arising 24 cm above base of common stipe (or below middle of common stipe), much taller than sterile blade: stalk 1225 cm; lamina bi-to tripinnate, subdeltoid, 410 cm; sporangia globular, attached on both lateral sides of axes. Spore surface irregularly reticulate with minute granules. 2n = 90.
In shaded shrubs; 4001000 m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Liaoning, Shaanxi, Shandong, Sichuan, Taiwan, Zhejiang [temperate and warm regions of E Asia including within Himalaya, India, Japan, Korea, Vietnam].