BOLBITIDACEAE [Draft]
实蕨科 shi jue ke
Xing Fuwu 1*, Wang Fa-guo1, Kunio Iwatsuki2
Plants small to medium-sized, terrestrial or creeping on rocks or river-banks or small trees. Rhizome creeping or short erect or scandent, dorsiventral, covered with ovate or lanceolate, brown scales. Leaves distant or tufted, dimorphic, simple or mostly pinnate. Sterile leaves: petioles dark brown or blackish near the base, bearing scales ±similar to those on the rhizome, usually on the basal part only; lamina herbaceous or papyraceous, apex usually with bulbils, margins entire, crenate or lobed, with or without teeth or spines, veins free or variously anastomosing and forming areolae, with or without included veinlets. Fertile leaves of similar shape as the sterile ones, usually with longer stipes and narrower lamina, sori acrostichoid, sporangia spreading all over the lower surface.
2 genera of about 90 species, pantropical in distribution, mainly in Asia and Pacific islands; one genus in China.
Bolbitis Schott, Gen, Fil. ad pl. 13. 1835.
实蕨属 shi jue shu
Egenolfia Schott, Gen. Fil. ad pl. 16. 1835.
Moderate size or small ferns, terrestrial, sometimes creeping on rocks or river-banks or small trees. Rhizome creeping, short erect or scandent, dorsiventral, covered with lanceolate, brown or blackish scales. Leaves dimorphic, evergreen, simple, pinnate, or rarely bipinnate. Sterile leaves: petioles bearing ovate or lanceolate, brown scales at base; lamina mostly more or less herbaceous, apex usually with bulbils, margins crenate to deeply lobed, with or without teeth or spines, venation pattern: veins either free or variously anastomosing, with or without included free veinlets. Fertile leaves of similar shape as the sterile ones, usually with longer stipes and narrower lamina. Sori acrostichoid, without indusia. Spores globose or nearly so, with thick epispores.
1a. Veins free.
2a. Sterile leaves pinnate, base asymmetrical (except B. hookeriana), apex obtuse or acute.
3a. Bulbil terminal on the lamina····················································· 1. B. rhizophylla
3b. Bulbil subterminal on the lamina.
4a. Stipe and rachis subglabrous; base of pinnae asymmetrical; sporangia spreading all over the lower surface, the arrangement fully acrostichoid.
5a. Sterile pinnae usually obtuse or rounded at apex; fertile pinnae 8-10 × 2-3 mm 2. B. appendiculata
5b. Sterile pinnae acute at apex; fertile pinnae 4-6 × ca. 0.8 cm··························· 3. B. medogensis
4b. Stipe and rachis densely scaly; base of pinnae symmetrical; sporangia on all sides of the costa and costules, usually arranged moniliform················································································ 4. B. hookeriana
2b. Sterile leaves bipinnatifid, base symmetrical, apex acuminate.
6a. Apex of lobes rounded, lobes 0-1 mm apart.
7a. Rhachis with a narrow wing except for the lowest part; base of lowermost pinnae symmetrical 5. B. sinensis
7b. Rhachis without wing; base of lowermost pinnae asymmetrical·························· 6. B. longiaurita
6b. Apex of lobes obtusely acute, lobes 2-3 mm apart.
8a. Stipe and rachis densely scaly; lobes of pinnae falcate, 1-1.5 cm long················ 7. B. tonkinensis
8b. Stipe and rachis subglabrous; lobes of pinnae triangular, 0.5-0.6 cm long 8. B. fengiana
1b. Veins more or less anastomosing.
9a. Venation pattern more or less irregular, veins along costae usually anastomosing and others free or rarely uniting; rachis narrowly winged or not.
10a. Pinnae pinnatifid; rachis wingless···································· H1. B. x yunnanensis
10b. Pinnae subnetire or crenate; rachis narrowly winged.
11a. Pinnae crenate; veins usually free except those along costae··························· H2. B.×laxireticulata
11b. Pinnae subnetire; veins anastomosing including those along costae H3. B.×nanjenensis
9b. Venation pattern regular, veins variously anastomosing; rachis wingless.
12a. 1-4 veinlets arising from either side of lateral veins uniting in sterile pinnae, leaves green or rarely purplish when dry.
13a. Venation without included free veinlets.
14a. Fertile pinnae linear; lateral pinnae 15-24 pairs········································· 9.B. angustipinna
14b. Fertile pinnae lanceolate; lateral pinnae 6-15(-17) pairs.
··································· 15a. Bulbilis terninal; lateral pinnae about 15 pairs; sterile pinnae truncate at base 11. B. tibetica
15b. Bulbils subterminal; lateral pinnae 6~12 paris.
16a. Sterila pinnae truncate to subcordate at base····················· 10. B. chirstensenii
16b. Sterila pinnae rounded at base········ 12. B. hekouensis.
13b. Venation with included free veinlets.
17a. Lateral pinnae 6-15 pairs; terminal segment triangular································· 13. B. subcordata
17b. Lateral pinnae 2-7 pairs; terminal segment conform to the central pinnae.
18a. Fertile pinnae lanceolate, 3-8 times longer than wide.
19a. Sterile pinnae 2-3 pairs; acrostichoid················································· 14. B. hainanensis
19b. Sterile pinnae 5-7 pairs; fertile pinnae non-acrostichoid.
20a. Leaves purplish when dry; 2-3 veinlets arising from lateral veins, free veinlets without thickened ends············································· 15. B. scalpturata
20b. Leaves stramineous when dry; 4-5 veinlets arising from lateral veins; free veinlets with thickened ends········································· 16. B. deltigera
18b. Fertile pinnae linear, 14-45 times longer than wide.
21a. Leaves purplish when dry; 5-8 veinlets arising from either side of lateral veins uniting in sterile pinnae ···························································· 17. B. costata
21b. Leaves greenish when dry; 4 veinlets arising from either side of lateral veins uniting in sterile pinnae ····························································· 18. B. virens
12b. 5-8 veinlets arising from either side of lateral veins uniting in sterile pinnae, leaves often dark-brownish when dry.
22a. Rhizome with 4 rows of leaves; scales on base of stipe with obvious brown margin 19. B. confertifolia
22b. Rhizome with 2 rows of leaves; scales on base of stipe without obvious brown margin.
23a. Leaves thickly chartaceous; rhizome 1-1.5 cm thick, high climbing 20. B. scandens
23b. Leaves thinly herbaceous; rhizome 3-6 mm thick, terrestrial or climbing on bases of tree-trunks 21. B. heteroclita
1. Bolbitis rhizophylla (Kaulf.) Hennipman, Blumea 18: 148. 1970 et Monogr. Gen. Bolbitis 199, fig. 52: d-r, fig. 53. 1977.
根叶刺蕨 gen ye ci jue
Gymnogramma rhizophylla Kaulf., Enum. Fil. 78. 1824. --- Egenolfia rhizophylla (Kaulf.) Fée, Gener. Polyp. 48. 1852.
Rhizome short-creeping, 4-6 mm thick, with 2 or 3 rows of leaves, densely covered with ovate-lanceolate, subclathrate, brown. Leaves close together; sterile leaves pinnate, 20-90 cm long; petiole 10-20 cm long, base 2-3 mm thick, lamina lanceolate, pinnate, 40-80 × 5-13 cm, herbaceous, green to blackish, dark green when dry, glabrous on both surfaces, the petiole and lower side of the basal part of the rhachis usually with small, blackish, spreading scales, the terminal segment 1-6 cm long; rhachis with a narrow wing except for the lowest part; pinnae 20-35 pairs, alternate to oppostie, elliptic-lanceolate, sessile, up to 3.5 cm apart, 2.5-4 × 1-1.5 cm, base rounded-truncate, ±symmetrical, margin finely serrate, or serrately lobed to 1/3 towards the costa, apex orbicular or acuminate, the two lowermost pinnae deflexed or not; veinlets 2-4 pairs, pinnate, simple. Fertile leaves 16-30 cm long; lamina lanceolate, 16-20 × 1.5-2.5 cm, widest usually about the middle; pinnae 20-25 pairs, elliptic, ca. 1.5 cm apart, 6-12 × 3-4 mm, margin entire or irregular undulate, apex obtuse. Sporangia inserted all over the lower surface. 2n=82.
On rocks in dense forests; 400-1000. Endemic toTaiwan (Tainan).
2. Bolbitis appendiculata (Willd.) K. Iwats., Acta Phytotax. Geobot. 18: 48. 1959.
刺蕨 ci jue
Acrostichum appenndiculatum Willd., Spec. Pl. 5: 114. 1810.
Egenolfia crenata Ching & P. S. Chiu ex Ching & C.H.Wang Acta Phytotax. Sin. 21: 212. 1983.
Rhizome short-creeping, up to 3 mm thick, with 2 or 3 rows of leaves, densely covered with scales, scales lanceolate, 2-3 × 1-1.5 mm, opaque or subclathrate, blackish, margin with irregular small tooth. Leaves close together; sterile leaves pinnate,15-40 cm long; petiole 5-15 cm long, ca. 1 mm thick, sparsely covered scales near the base; lamina lanceolate, 12-25 × 3.5-6 cm, herbaceous, thickly green when dry, the terminal segment, triangular, apex acuminate, sometimes somewhat prolonged; rhachis usually with a narrow wing; pinnae 15-30 pairs, opposite or alternate, elliptic-linear, up to 3 cm apart, 2-3.5 × 5-8 cm, base asymmetrical, usually oblique, margin ±entire, finely serrate or lobed to 1/3 towards the costa, apex acute or acuminate, the tow lowermost pinnae 1-2 mm stalked; secondary veins on either side with 2 or 3 tertiary veins. Fertile leaves pinnate, 25-40 cm long; petiole 20-25 cm long; pinnae 10-30, 8-10 × 2-3 mm; two lowermost pinnae 1-9 mm stalked. Sporangia inserted all over the lower surface, the arrangement fully acrostichoid.
Creeping on rocks or in soil in evergreen forests; 100-1300. Hainan, Guangdong, Guangxi, Hong Kong, Taiwan, Yunnan [Japan, Bangladesh, Bhutan, Sikkim, India, Laos, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, India, Indonesia, vietnam].
3. Bolbitis medogensis (Ching & S. K. Wu) S. Y. Dong, Acta Phytotax. Sin. 43(2): 105. 2005.
墨脱刺蕨 mo tuo ci jue
Egenolfia medogensis Ching & S. K. Wu in C. Y. Wu, Fl. Zizang. 1: 278. 1983.
Rhizome short creeping, densely covered with scales, scales linear-lanceolate, brown, entire. Leaves close together or tufted; sterile leaves pinnate, ca. 75 cm long; petiole 20-35 cm long, sparsely covered with scales; lamina broadly lanceolate, ca. 48 × 20-25 cm, green when dry, glabrous on both surfaces, apex caudately acuminate, subterminal with a bulbil; pinnae ca. 12 pairs, alternate, with short stalk, 3-5 cm apart, 10-13 × ca. 3.5 cm, base nearly truncate, margin crenate-serrate, with a fleshy spine in sinus; secondary veins slightly obvious, dichotomous. Fertile leaves pinnate; petiole ca. 50 cm long; lamina narrowly lanceolate, ca. 46 × 10-12 cm; pinnae 10-12 pairs, linear, 4-6 × ca. 0.8 cm; entire, subsessile. Sporangia inserted all over the lower surface.
On slope in evergreen forests; 700~900. Endemic to Xizang (Mêdog).
4. Bolbitis hookeriana K. Iwats., Acta Phytotax. Geobot. 18: 49. 1959.
虎克实蕨 hu ke shi jue
Polybotrya vivipara Buch.-Ham. ex Hook., Exot. Fl. 2: 107, pl. 107. 1825; Bolbitis appendiculata ssp. vivipara (Hook.) Hennipman, Blumea 18: 147. 1970.
Rhizome long-creeping, 5-12 cm thick, with usually 3 rows of leaves, without or with but few scleraenchyma strands, sterile leaves pinnate, 20-100 cm long, the petiole and rhachis usually throughout with spreading scales or sometimes glabrescent; petiole 5-25 cm long; lamina 20-85 × 4-20 mm, the terminal segment 3-10 cm long; pinnae 33-60 pairs, 3.5-10 ×1.2-2.5 cm, base ±symmetrical, margin usually entire, sometimes lobed to 1/5 towards the costa. 3-5 pairs, alternate, linear-lanceolate, usually widest about (to below) the middle, 15-25 2-3 cm, base ±symmetrical, broadly attenuate or truncate, margin slightly serrate-crenate, with a short spine in each sinus, apex acuminate, the two lowermost pinnae with short stalk; secondary veins on either side with 2 or 3 tertiary veins. Fertile leaves 40-90 cm long; petiole 16-45 cm long; lamina 14-40 × 4-10 cm; pinnae 35-50 pairs; central pinnae 2-8.5 cm long, the lowermost two pinnae 0-4 mm stalked. Sporangia inserted on all sides of the costa and costules, usually arranged moniliform.
On rocks in dense forests. Alt. 500 m. Yunnan (Yingjiang). [Bangladesh, India, Myanmar, Indo-China Peninsula].
5. Bolbitis sinensis (Baker) K. Iwats., Acta Phytotax. Geobot. 18: 49. 1959.
中华刺蕨 zhong hua ci jue
Acrostichum sinense Baker, Kew Bull. 1906: 14; Egenolfia sinensis (Baker) Maxon, Proc. Biol. Sc. Wash. 36: 173. 1923.
Egenolfia crassifolia Ching, Acta Phytotax. Sin. 21: 216. 1983.
Rhizome short creeping, ca. 1 cm thick, with 2 or 3 rows of leaves; scales ovate-lanceolate, subclathrate, brownish or blackish, ca. 3 mm long. Leaves close together; sterile leaves 20-110 cm long; petiole 10-35 cm long, base ca. 3 mm thick, grooved adaxially, lamina ovate-lanceolate, pinnate, 40-110 × 10-35 cm, herbaceous, green to blackish, dark green when dry, glabrous on both surfaces, the terminal segment 8-30 cm long; rhachis with a narrow wing except for the lowest part, sparsely covered with dark brown, ovate scales; pinnae 14-30 pairs, alternate to opposite, elliptic, sessile, 2-3 cm apart, 10-14 × 2-2.5 cm, base truncate, ±symmetrical, margin lobed to 1/3-2/3 towards the costa, apex orbicular or acuminate; lobes 16-20 pairs, ca. 1 mm spaced, 8-12 × 4-5 mm, margin usually entire or finely crenate-serrate, apex orbicular, with a spine in sinus; veinlets 6-7 pairs, pinnate, simple or biforked. Fertile leaves 16-80 cm long; petiole as long as the sterile ones; lamina lanceolate, 22-30 × 4-5 cm, apex caudate; pinnae 10-14 pairs, narrowly lanceolate, 2-3 cm apart, 4-5 × ca. 0.5 cm, basal pairs 3-4 mm stalked, base truncate to slightly cordate, apex acuminate. Sporangia inserted all over the lower surface. 2n=82.
In soil or on rocks in dense forests; 650~1900. Guizhou, Hong Kong, Yunnan [Bangladesh, Cambodia, India, Indonesia, Java, Myanmar, Thailand, Vietnam, Lesser Sunda Islands].
6. Bolbitis longiaurita F. G. Wang & F. W. Xing, Novon 16(3): 434. 2006.
长耳刺蕨 chang er ci jue
Bolbitis bipinnatifida K. Iwats., Acta Phytotax. Geobot. 18: 49. 1959, non (Kuhn) Ching, 1934; Egenolfia bipinnatifida J. Sm., Hist. Fil. 132. 1875.
Acrostichum appendiculatum Willd. var. costulatum Hook., Spec. Fil. 5: 252 1864; Bolbitis sinensis K. Iwats. var. costulata Tagawa & K.Iwats., Acta Phytotax. Geobot. 22: 102. 1967.
Rhizome short creeping, ca. 1 cm thick, densely covered with scales; scales ovate-lanceolate, dark brownish, ca. 3 mm long. Leaves close together; sterile leaves 40-70 cm long; petiole 11-15 cm long, base ca. 3 mm thick, grooved adaxially, sparsely covered with scales; lamina elliptic, bipinnate, 50-60 × 10-35 cm, papery, dark green when dry, glabrous on both surfaces, widest at base, the terminal segment long attenuate to flagelliform; rhachis shallowly grooved, glabrous; pinnae ca. 20 pairs, 14-16 × 2.5-3.5 cm, lanceolate, basal pairs nearly opposite, with short stalk, the two lowermost pinnae 3-6 mm stalked, apex caudately acuminate, base asymmetrical with its basiscopic 2-3 lobes undeveloped, other lobes longer than the acroscopic ones, other pairs of pinnae lobed to 1/2 to the costa, base truncate and symmetrical; lobes 18-20, elliptic, 1-1.5 × ca. 0.6 cm, margin sparsely rounded dentate, with a spine in sinus; veinlets 6-7 pairs, free, biforked or simple. Fertile leaves: petiole 20-40 cm long; lamina lanceolate, pinnate, 18-20 × 4-5 cm; pinnae 12-14 pairs, elliptic, 2-3 × ca. 0.5 cm, basal pairs with short stalk, base rounded truncate, apex obtuse. Sporangia inserted all over the lower surface.
On rocks in dense forests; 1200. Endemic to Yunnan.
This species is similar to B. sinensis, but differs in its rhachis without wing, the base of the two lowermost pinnae asymmetrical with its basiscopic 2-3 lobes undeveloped, other lobes longer than the acroscopic ones.
7. Bolbitis tonkinensis (C. Chr.) K. Iwats., Acta Phytotax. Geobot. 18: 49. 1959.
镰裂刺蕨 lian lie ci jue
Egenolfia tonkinensis C. Chr. ex Ching, Bull. Fan Mem. Inst. Biol. 2: 306. 1931.
Rhizome short-creeping, ca. 1 cm thick, densely covered with scales, scales ovate, ca. 3 mm long, fuscous, margin erose. Leaves close together; sterile leaves ca. 80 cm long; petiole 20-30 cm long, ca. 3 mm thick at the base, grooved adaxially, covered with ovate-lanceolate fuscous scales; lamina bipinnate, triangular-ovate, 50-60 × 20-30 cm, dark green when dry, glabrous on both surfaces, apex caudately acuminate, subterminal with a bulbil; pinnae 12-16 pairs, lanceolate, alternate, with short stalk 2-4 mm long, 13-15 × 3-3.5 cm, base circularly truncate and slightly asymmetrical, margin lobed to 2/3 to the costa, apex caudately acuminate, subterminal with a small bulbil; the terminal segment prolonged to flagelliform and pinnatifid, subterminal with a big bulbil; lobes 16-18 pairs, 2-3 mm apart, elliptic-lanceolate, falcate, 10-15 × 5-6 mm, with a long spine in sinus. veinlets 6-7 pairs, pinnate, simple or biforked. Fertile leaves pinnate; petiole 40-45 cm long; lamina narrowly lanceolate, ca. 25 × 6 cm; pinnae ca. 10 pairs, elliptic-lanceolate, 2.5-3 × 0.5 cm; entire. Sporangia inserted all over the lower surface.
On moist rocks in forests; 200~1200 m. S Yunnan [Vietnam (Tonkin), Thailand].
8. Bolbitis fengiana (Ching) S. Y. Dong, Acta Phytotax. Sin. 43(2): 107. 2005.
疏裂刺蕨 shu lie ci jue
Egenolfia fengiana Ching, Acta Phytotax. Sin. 21: 215. 1983.
Rhizome long-creeping, 4-6 mm thick, densely covered with scales, scales lanceolate, ca. 2 mm long, blackish brown. Leaves close together; sterile leaves 40-60 cm long; petiole 15-25 cm long, base ca. 2-3 mm thick, grooved adaxially, sparsely covered with scales; lamina broadly lanceolate, bipinnate, 22-30 × 10-15 cm, herbaceous, dark green when dry, glabrous on both surfaces, base rounded-truncate, apex long-caudate, subterminal with a small bulbil; rhachis with a narrow wing; pinnae 10-15 pairs, alternate, linear-lanceolate, ca. 1 cm apart, 5-9 × 1-1.5 cm, basal pairs with short stalk 2-3 mm long, base truncate, lobed to 1/3 towards to costa; lobes 8-12 pairs, triangular, 5-6 × 5-6 mm, margin with sparsely short tooth, with long acuate spine in sinus; veinlets 3 pairs, pinnate, simple. Fertile leaves pinnate; petiole ca. 25-40 cm long; lamina lanceolate, ca. 16 × 4-5 cm; pinnae 10-12 pairs, elliptic to elliptic-lanceolate, 1.5-2 cm apart, 2-2.5 × ca. 0.6 cm, basal pairs nearly opposite and 3-4 mm stalked, base broadly cuneate, margin subentire or slightly undulate, apex obtuse. Sporangia inserted all over the lower surface.
In dense forests; 1300-1800. Endemic to Yunnan.
9. Bolbitis angustipinna (Hayata) H. Ito, J. Jpn. Bot. 14: 443. 1938.
多羽实蕨duo yu shi jue
Bolbitis contaminans (Wall.) Ching in C.Chr. Ind. Fil. Suppl. 3: 47. 1934.
Leptochilus cuspidatus var. crenatus Ros., Hedwigia 56: 281. 1915.
Rhizome robust and long-creeping, 1-2 cm thick and 10-16 cm long, densely covered with scales, with 2-4 rows of leaves; scales lanceolate, 4-6 mm long, subclathrate, dark brown or blackish. Leaves close together; sterile leaves pinnate, 55-145 cm long; petiole 40-60 cm long, near the base ca. 8 mm thick, sparsely covered with scales, with 7-16 vascular bundles; lamina elliptic, 60-85 ×30-45 cm, widest somewhat below the middle, hard herbaceous, grayly green when dry; pinnae 12-20 pairs, alternate, linear-lanceolate, usually widest about (to below) the middle, 15-25 ×2-3 cm, base ±symmetrical, broadly attenuate or truncate, margin slightly serrate-crenate, with a short spine in each sinus, apex acuminate, the two lowermost pinnae with short stalk; the terminal segment joined to the rhachis, usually narrowly triangular, ±conform to the central pinnae, 10-20 cm long, subterminal with bulbil; rhachis not winged; secondary veins prominent; veins forming a costal areole and 3 rows distal areoles, without included free veinlets. Fertile leaves 50-150 cm long; petiole 30-85 cm long; lamina 70-90 × 15-25 cm; pinnae 80-120 × 5-8 mm. Sporangia inserted usually all over the lower surface, the arrangement fully acrostichoid, sometimes either with a sterile strip along the costa.
On rocks or near stream in dense forests. 250-1400 m. Hainan, Taiwan, Yunnan [Nepal, Bhutan, Sikkim, India, Sri Lanka, Myanmar, Thailand].
10. Bolbitis christensenii (Ching) Ching in C. Chr. Ind. Fil. Suppl. 3: 47. 1934.
贵州实蕨 gui zhou shi jue
Campium chistensenii Ching, Bull. Fan Mem. Inst. Biol. 2:214, pl. 31. 1931.
Rhizome robust and short-creeping, ca. 1 cm thick, densely covered with scales; scales lanceolate, ca. 9 mm long, blackish brown, margin maple, entire. Leaves tufted; sterile leaves pinnate; petiole 30-40 cm long, base ca. 4 mm thick, base densely covered with lanceolate scales; lamina elliptic, 40-60 ×20-25 cm, thinly herbaceous, grayish brown when dry, apical part pinnatipartite to lanceolate lobes, subterminal with bulbil; rhachis not winged, densely covered with lanceolate brown scales especially when young; pinnae 6-10 pairs, the basal ones nearly opposite, with short stalk 2-3 mm long, lanceolate, 6.5-12.0 ×1.2-2.0 cm, base circular, margin crenate-serrate, sometimes with a obtuse tooth in sinus, apex long-acuminate; secondary veins unconspicuous; veins forming 3 rows, long and narrow areoles, without included free veinlets. Fertile leaves as long as sterile leaves; lamina obviously narrower, linear-lanceolate; pinnae 3-4 × ca. 5 mm, base circularly cuneate, subsessile, entire, apex acuminate. Sporangia inserted all over the lower surface.
Near stream in forests. 400-1100 m. Guangxi, Guizhou. [Vietnam (Tonkin)].
11. Bolbitis tibetica Ching & S. K. Wu, C. Y. Wu, Fl. Xizang. 1: 276. 1983.
西藏实蕨 xi zang shi jue
Rhizome short and robust, ca. 1.5 thick, densely covered with scales; scales 4-5 mm long, narrowly lanceolate, nut-brown, entire. Leaves tufted, 80-110 cm long; sterile leaves pinnate; petiole ca. 25 cm long, stramineous, sparsely covered with scales, base 4-5 mm thick, grooved adaxially; lamina broadly lanceolate, 50-65 ×25-30 cm, herbaceous, sap green when dry, glabrous on both surfaces, apex pinnatilobate and elongate to flagelliform, terminal with bulbil; rhachis not winged, sparsely covered with nut-brown scales; pinnae ca. 15 pairs, alternate, 1-1.5 cm apart, lanceolate, 16-18 × 2.5-3 cm, lower pairs pinnae with short stalk, upper pairs base connate with rhachis, base truncate, margin with obtuse tooth, apex caudately acuminate; veins prominent on both surfaces; veins forming a costal areole and 3-4 rows tetragonal to hexagonal areoles, without included free veinlets. Fertile leaves 95-115 cm long; petiole 50-55 cm long; lamina 50-60 × 10-12 cm, apex pinnatilobate; pinnae narrowly lanceolate, 6-8 × 1.5-1.8 cm, base circularly truncate, apex acuminate. Sporangia inserted all over the lower surface. Chromosomes.
In broad-leaved forests. 800 m. endemic to Xizang (mêdog).
12. Bolbitis hekouensis Ching, Acta Phytotax. Sin. 21: 212. 1983.
河口实蕨 he kou shi jue
Rhizome robust and short creeping, 1-1.5 cm thick, densely covered with scales; scales lanceolate, ca. 4 mm long, dark brown, entire. Leaves close together, 28-51 cm long; sterile leaves pinnate; petiole 8-17 cm long, base ca. 3 mm thick, pale brown, grooved adaxially, densely covered with lanceolate, brown scales; lamina elliptic, 18-32 ×9.5-15.0 cm, thickly papyraceous, grayish brown when dry, apical part pinnatipartite to lanceolate lobes, subterminal with bulbil; rhachis not winged, densely covered with lanceolate, brown scales especially when young; pinnae 6-10 pairs, the basal ones nearly opposite, with short stalk 2-3 mm long, lanceolate, 6.5-12.0 ×1.2-2.0 cm, base circular, margin crenate-serrate, sometimes with an obtuse tooth in each sinus, apex long-acuminate; secondary veins unconspicuous; veins forming 3 rows long and narrow areoles, without included free veinlets. Fertile leaves as long as sterile leaves; lamina obviously narrower, linear-lanceolate; pinnae 3-4 × ca. 5 mm, base circularly cuneate, subsessile, entire, apex acuminate. Sporangia inserted all over the lower surface.
Usually in rocky crevices of limestone area in rain forests. 400-1100 m. Endemic to Hainan and Yunnan.
13. Bolbitis subcordata (Copel.) Ching in C.Chr., Ind. Fil. Suppl. 3: 50. 1934.
华南实蕨 hua nan shi jue
Campium subcordatum Copel., Phil. J. Sc. 37: 369, f. 23, pl. 16. 1928.
Bolbitis media Ching & C. H. Wang, Acta Phytotax. Sin. 21: 212. 1983.
Rhizome short-creeping, 8-15 mm thick, with lanceolate, clathrate, subentire, brown or blackish scales. Leaves ± close together, 60-120 cm long; sterile leaves pinnate, 20-110 ×15-28 cm; petiole 8-45 cm long, densely scaly near base, bearing small scales at apex, with 5-9 vascular bundles; lamina 12-65 cm long, widest below the middle, herbaceous, dark green or brownish, the terminal segment narrowly triangular, longer to shorter than the central pinnae, base trifid; rhachis not winged; pinnae 6-14 pairs, opposite below the middle, base broadly cuneate to subcordate, margin usually lobed to 1/3 towards to costa, crenate-serrate, with a prominent tooth in each sinus between the lobes, apex acuminate; secondary veins slightly prominent; veins forming a costal areole and distal areoles, areoles in part with usually one simple, excurrent free vein. Fertile leaves 30-125 cm long; petiole 15-70 cm long; pinnae 6-25, 2-6 cm apart, 4-8 × ca. 1 cm; two lowermost pinnae 1-7 mm stalked. Sporangia inserted all over the lower surface, the arrangement acrostichoid. Chromosomes, 2n = 82.
In forests or near streams. Alt. 100-600m. Hong Kong, Jiangxi, Macau, Taiwan, Fujian, Guangdong, Guangxi, Hainan, Yunnan, Zhejiang [Japan, Vietnam].
14. Bolbitis hainanensis Ching & C. H. Wang, Acta Phytotax. Sin. 21: 214. 1983.
厚叶实蕨 hou ye shi jue
Bolbitis yunnanensis Ching, Acta Phytotax. Sin. 21: 214. 1983.
Rhizome short-creeping, 1-1.5 cm thick and 5-9 cm long, densely covered with scales; scales lanceolate, ca. 3 mm long, dark brown, entire. Leaves close together, 4-7 mm apart; sterile leaves pinnate; petiole 20-40 cm long, near the base ca. 3.5 mm thick, densely covered with scales, glabrous upwards; lamina ovate, 25-35 × 20-25 cm, hard papyraceous, grayish brown when dry, glabrous on both surfaces; pinnae 3-5 pairs, alternate, elliptic-lanceolate, 16-25 ×4.5-6 cm, base narrowly cuneate, margin wrinkled undulate, apex caudately acuminate, the two lower pairs of pinnae with stalk ca. 1 cm long; the terminal segment ±conform to the central pinnae; rhachis not winged; secondary veins tenuous, prominent abaxially; veins unconspicuous, forming a costal big areole and 3 rows small areoles, sometimes included free short veinlets. Fertile leaves 35-75 cm long; petiole 30-55 cm long; lamina long ovate, 12-25 × 10-14 cm; pinnae 2-5 pairs, lanceolate, 6-10 × 1-1.5 c, base narrowly cuneate, apex acuminate. Sporangia inserted usually all over the lower surface, the arrangement acrostichoid.
In dense forests. Alt. 700-1100 m. Endemic to Hainan and Yunnan.
15. Bolbitis scalpturata (Fée) Ching in C. Chr., Ind. Fil. Suppl. 3: 50. 1934
红柄实蕨 hong bing shi jue
Heteroneuron scalpturatum Fée, Hist. Acrost. 95, pl. 56, p.p. 1845.
Rhizome long-creeping, up to 10 mm thick and ca. 10 cm long, with 2 rows of leaves, densely covered with lanceolate, subclathrate, blackish scales. Leaves close together; sterile leaves pinnate; petiole 15-24 cm long, sparsely scaly near the base, with 5-9 vascular bundles; lamina 20-55 × 7-30 cm, widest below the middle, herbaceous to subcoriaceous, purplesh or purplish-brown, the terminal segment usually subarticulate, 9-20 cm long, subterminal with a bulbil; rhachis not winged; pinnae 2-12 pairs, opposite or alternate, 1.5-4.5 cm apart, 10－12 × 2－2.5 cm, base ±symmetrical, narrowly to broadly cuneate, margin ±entire or serrate-crenate, without spines, apex acute to acuminate; veins forming a costal areole and one to few transverse rows of distal areoles, part of the areloles with few to several excurrent included free vein. Fertile leaves 30-60 cm long; petiole 15-40 cm long; pinnae 5-20, 2-6 cm apart, 4-8 × ca. 1 cm; two lowermost pinnae 0-6 mm stalked. Sporangia inserted all over the lower surface, or along the margin only, the arrangement pteridoid.
On rocks in forests. 0-1200 m. Hainan, Taiwan. [Myanmar, Thailand, Vietnam, Malaysia, Indonesia, Philippines].
16. Bolbitis deltigera (Bedd.) C. Chr., Ind. Fil. Suppl. 3: 48. 1934.
间断实蕨 jian duan shi jue
Poecilopteris costata (C.Presl) Moore var. deltigera Bedd., Ferns Br. Ind. 114, pl. 114. 1865; Bolbitis virens (Hook. & Grev.) Schott var. deltigera (Bedd.) Hennipman, Monogr. Gen. Bolbitis 184, f. 47, 48 k~m. 1977.
Rhizome up to 14 mm thick and 10-15 cm long, with 2 or 3 rows of leaves, covered with lanceolate (5-7 × ca. 1.5 mm), subclathrate, blackish scales. Sterile leaves pinnate, 40-115 cm long; petiole 20-60 cm long, 2-7 mm thick near the base, with 7-14 vascular bundles; lamina 20-75 ×15-50 cm, widest about or below the middle, herbaceous to coriaceous, purplesh or purplesh-brown, the terminal segment 14-40 cm long, subarticulate or joined to the rhachis, with a bulbil subterminally on the terminal segment; rhachis not winged; pinnae 6-20 pairs, opposite or alternate, up to 6 cm apart, 10－30 × 2.5-6.0 cm, base ±symmetrical, usually narrowly, sometimes broadly attenuate or cuneate, margin usually ±entire or serrate-crenate, apex acuminate, the tow lowermost pinnae 1-10 mm stalked; veins forming a costal areole and several equally large distal areoles, 4-5 veins arising from each side of lateral veins anastomosing. Fertile leaves 40-120 cm long; petiole 25-60 cm long; pinnae 7-18, up to 6 cm apart, central pinnae 5-17 × 1.5-4.5 cm; two lowermost pinnae 1-9 mm stalked. Sporangia inserted on the veins and near the margin only, the arrangement non-acrostichoid.
On slope or in valley under rain forests. 300-700 m. Hainan, Yunnan. [Bangladesh, Nepal, Bhutan, Sikkim, India, Myanmar, Thailand].
17. Bolbitis costata (C. Presl) Ching in C. Chr., Ind. Fil. Suppl. 3: 47. 1934.
紫轴实蕨 zi zhou shi jue
Campium costatum C.Presl, Tent. Pterid. 239, pl. 10. 1836.
Rhizome up to 15 mm thick and 10 cm long, with 2-4 rows of leaves, covered with lanceolate (ca. 5 × 1 mm), subclathrate, light brown scales. Sterile leaves pinnate, 55-120 cm long; petiole 20-60 cm long, 2-7 mm thick near the base, with 5-20 vascular bundles; lamina 35-75 × 20-40 cm, widest about or below the middle, subcoriaceous, purplish when dried, the terminal segment 20-40 cm long, with a bulbil subterminal on terminal segment or pinnae; rhachis not winged; pinnae 3-19 pairs, opposite or alternate, up to 8 cm apart, 16-45 × 4-9 cm, base symmetrical, acute to subcordate, margin entire or crenulate, sometimes crisped, apex rather abruptly narrowing into the acute or acuminate tip, the two lowermost pinnae 1-6 mm stalked; veins for the greater part forming 4 or more regular rows of areoles, with in part of the areloles one t three excurrent included free vein whick end in a thickening. Fertile leaves 40-170 cm long; petiole 20-100 cm long; lamina 15-85 × 7-30 cm, the terinal segment 8-20 cm long; pinnae 5-18, up to 11 cm apart, ca. 22 × 0.6-2 cm; two lowermost pinnae 1-6 mm stalked. Sporangia inserted mainly on the veins, arranged ususlly acrostichoid, rarely gymnogrammoid.
In forests near streams; 0-1200 m. Yunnan [Bangladesh, Nepal, Sikkim, India, Myanmar, Thailand].
Bolbitis costata is similar to B. scalpturata for their fronds usually turn purplesh when dry; it differs by having wider sterile pinnae (to 4-9 cm), 5-8 veinlets arising from each side of lateral veins, and fertile pinnae linear (14-16 times as long as wide).
18. Bolbitis virens (Hook. & Grev.) Schott, Gen. Fil. pl.13. 1835.
宽羽实蕨 kuan yu shi jue
Acrosticum virens Wall. ex Hook. & Grev., Ic. Fil. pl. 221. 1831.
Bolbitis latipinna Ching, Acta Phytotax. Sin. 21: 213. 1983.
Rhizome up to 13 mm thick and 10 cm long, with 2 or 3 rows of leaves, densely covered with scales, scales narrowly lanceolate, ca. 4 × 1.5 mm, subclathrate, brownish. Sterile leaves pinnate, 40-110 cm long; petiole 20-60 cm long, 2-7 mm thick near the base, with 7-14 vascular bundles; lamina 25-75 × 20-50 cm, widest about or below the middle, herbaceous to coriaceous, the terminal segment 15-40 cm long, subarticulate or joined to the rhachis, somewhat prolonged, with a bulbil subterminal on terminal segment or pinnae; rhachis not winged; pinnae 6-22 pairs, opposite or alternate, up to 6.5 cm apart, 10-30 × 2.5-6 cm, base ±symmetrical, sometimes broadly attenuate or cuneate, margin entire or serrate or serrate-crenate, apex acuminate, the two lowermost pinnae 1-10 mm stalked; veins forming a costal areole and several to many equally large distal ones. Fertile leaves 40-120 cm long; petiole 25-70 cm long; lamina 15-65 × 7-30 cm, the terinal segment 4-19 cm long; pinnae 7-18, up to 6 cm apart, 4-20 × 0.2-4.5 cm; two lowermost pinnae 1-6 mm stalked. Sporangia inserted all over the lower surfaces, arranged variously.
On
Rrocks in forests; 850 m. Yunnan [Bangladesh, Myanmar, Thailand].
19. Bolbitis confertifolia Ching, Acta Phytotax. Sin. 21: 211. 1983.
密叶实蕨 mi ye shi jue
Rhizome robust and long-creeping, ca. 1.5 cm thick and 12 cm long, densely covered with scales; scales ovate-lanceolate, ca. 3 cm long, median brunneus, margin brown, subentire. Leaves densely clustered on the upper side of rhizome; sterile leaves elliptic, pinnate; petiole 20-25 cm long, densely covered with scales as the rhizome, grooved adaxially; lamina 35-45 × ca. 15 cm, herbaceous, dark green when dry, the terminal segment elongate to long flagelliform, terminal with bulbil; pinnae 4-5 pairs, subsessile, lanceolate, 12-15 × 2.5-3 cm, base cuneate, margin repand, apex long acuminate; secondary veins prominent on abaxial surfaces; veins forming nearly hexagonal areoles, without included free veinlets. Fertile leaves unseen.
In rain forests. Endemic to Yunnan (Jinghong).
20. Bolbitis scandens W. M. Chu ex Ching & C. H. Wang, Acta Phytotax. Sin. 21: 213. 1983.
附着实蕨 fu zhe shi jue
Rhizome climbing on tree-trunks over 2-3 meters in height, 1-1.5 cm thick, densely covered with scales; scales lanceolate, 3-4 mm long, nut-brown, margin maple, entire. Leaves close together; sterile leaves pinnate; petiole 16-20 cm long, near the base ca. 4 mm thick, grooved adaxially, sparsely covered with scales below the middle; lamina ovate, 35-40 ×20-25 cm, widest somewhat below the middle, thickly chartaceous, grayly brown when dry; pinnae 2-3 pairs, subsessile, the two lowermost pinnae nearly opposite, elliptic-lanceolate, 15-18 ×4-5 cm, base circularly cuneate, margin slightly undulated, apex caudately acuminate; the terminal segment bigger than the central pinnae; secondary veins prominent on the lower surface, parallel and 6-7 mm apart; veins unconspicuous on both surfaces, forming a narrow costal areole and 3-4 rows distal areoles, sometimes with included free veinlets. Fertile leaves 40-60 cm long; lamina narrow-ovate, 25-30 × 10-12 cm; pinnae 3-4 pairs, broadly lanceolate,10-12 × 2-2.5 cm, apex acuminate, subsessile. Sporangia inserted all over the lower surface, the arrangement fully acrostichoid.
Climbing on tree-trunks in forests; 700 m. S Yunnan.
Bolbitis scandens is similar to B. heteroclita; it differs by having rhizome climbing on tree-trunks, and lateral pinnae of sterile fronds usually three-paired, terminal pinnae never obviously prolonged and leaf texture thickly papyraceous.
21. Bolbitis heteroclita (C. Presl) Ching in C. Chr. Ind. Fil. Suppl. 3: 48. 1934.
长叶实蕨 chang ye shi jue
Acrostichum heteroclitum C.Presl, Rel. Haenk.15, pl. 2, f. 2. 1825.
Bolbitis annamensis Tard.–Blot & C. Chr., Not. Syst. 7: 100. 1938 & in Lecomte, Fl. Gen. I.-C. 7:436, f. 50: 3-4. 1941.
Rhizome robust and long-creeping, 3-6 mm thick, densely covered with scales; scales ovate-lanceolate, 3-9 × 0.5-1.5 cm, usually subclathrate, brownish or blackish, subentire. Leaves ± spaced, 1-3 cm apart; sterile leaves simple or pinnate, 15- 80 cm long; petiole 10- 30 (-40) cm long, sparsely covered with scales, grooved adaxially; lamina 10-60 × 5-25 cm, thinly herbaceous, blackish when dry, apex elongate to long flagelliform, terminal with bulbil; rhachis not winged, sparsely covered with nut-brown scales; pinnae 1-6 pairs, alternate to opposite, broadly lanceolate, 10-19 × 3-6 cm, subsessile, base circularly cuneate, margin subentire or crenate-serrate, apex caudately acuminate; secondary veins prominent on both surfaces; veins forming 3 rows regularly tetragonal to hexagonal areoles, without included free veinlets. Fertile leaves 15-70 cm long; petiole 10-45 cm long; lamina 10-26 × 5-20 cm. Sporangia inserted all over the lower surface. Chromosomes. 2n = 82.
On rocks or at base of trees near streams in broad-leaved forests. 50~1500 m. Chongqing, Guangxi, Guangdong, Guizhou, Hainan, Sichuan, Yunnan, Taiwan. Asian tropics.
The following three forms are the putative hybrids recorded from China.
H1. Bolbitis ×yunnanensis (Ching & C. H. Wang) F. G. Wang, K. Iwats. & F. W. Xing comb. nov.
云南刺蕨 yun nan ci jue
Egenolfia yunnanensis Ching & P. S. Chiu ex Ching & C. H. Wang, Acta Phytotax. Sin. 1: 216. 1983.
Bolbitis angustipinna x sinensis Hennipman, Monogr. Gen. Bolbitis 284, fig. 83: a-c. 1977.
Rhizome creeping, 1-1.5 cm thick, densely covered with scales; scales ovate-lanceolate, dark brownish, 2-3 mm long. Leaves close together; sterile leaves 70-90 cm long; petiole 10-35 cm long, base covered with scales, ca. 3 mm thick, grooved adaxially, lamina ovate-lanceolate, bipinnate, 50-65 × 15-20 cm, papery, dark green when dry, glabrous, the terminal segment prolonged to flagelliform, subterminal with a big bulbil; rhachis with a narrow wing on the upper part; pinnae ca. 20 pairs, alternate, lanceolate, , 1-1.5 cm apart, 20-22 × ca. 3 cm, basal pairs with short stalk 3-4 mm, base broadly cuneate, margin lobed to 1/2 towards the costa, apex long-caudate; lobes 22-26 pairs, elliptic, ca. 1 mm spaced, 8-12 × ca. 5 mm, margin usually entire or slightly undulate, apex orbicular, with a long spine in sinus; veinlets 5-6 pairs, pinnate, simple, sometimes veins forming a costal areole. Fertile leaves: petiole ca. 35 cm long; lamina ca. 20 cm long; pinnae lanceolate, 4-6 × 0.6-0.9 cm, with short stalk, apex obtuse. Sporangia inserted all over the lower surface.
In soil in dense rain forests; 850. Guangxi, Yunnan.
H2. Bolbitis ×laxireticulata K. Iwats., Acta Phytotax. Geobot. 18: 50. fig. 7 & 8. 1959.
网脉实蕨 wang mai shi jue
Egenolfia laxireticulata (K. Iwats.) C. M. Kuo, Fl. Taiwan 1: 352. 1975.
B. appendiculata x B. subcordata
Rhizome short-creeping, ca. 4 mm thick, with scleraenchyma strands, sparsely covered with scales; scales ovate-lanceolate to lanceolate, subclathrate, 3-4 × 1.5-2 mm, brown-blackis. Leaves dimorphic; sterile leaves pinnate, 30-75 cm long; petiole 14-30 cm long, densely covered with scales; lamina oblong-elliptic, widest below or about the middle, 20-30 × 10-13 cm, papyraceous, grayish green when dry, with a spherical subterminal bulbil; rhachis sparsely covered with scales, with a narrow wing; pinnae alternate or opposite, 12-20 pairs, glabrous on both surfaces, 4.5－7 ×1.5－2, base sometimes symmetrical, variously cuneate, usually asymmetrical, margin ±entire to lobed to 2/5 towards the costa, lobes finely crenate-serrate, with a distinct spine in the sinuses, apex acute to acuminate, the two lowermost pinnae up to 1.5 mm stalked, terminal segment triangular; veins very irregular, forming a costal areole at least in part of the pinnae of one plant. Fertile leaves: petiole 25-40 cm long; lamina 14-28 cm long; pinnae 10-12 pairs, oblong-lanceolate, entire or undulate, ca. 2 × 0.4 cm. Sporangia inserted mainly on the veins.
On rocks along stream in rain forests, 500-650 m. Hainan, Guangdong, Hong Kong, Taiwan. [Japan (Ryukyu Isls.)].
H3. Bolbitis ×nanjenensis C. M. Kuo, Bot. Bull. Acad. Sin. 31: 308, fig. 1. 1990.
南仁实蕨 nan ren shi jue
B. appendiculata x B. heteroclita
Rhizome creeping, ca. 4 mm thick, densely covered with scales, scales ovate-lanceolate, dark brown, ca. 3 × ca. 1 mm, margin subentire, apex long acuminate. Sterile leaves 30-35 cm long, stramineous, sparsely covered scales; Lamina pinnate; rhachis usually with a narrow wing; pinnae 11-13 pairs, sessiles, oblong-lanceolate, ca. 5 × 1-1.2 cm, glabrous adaxially, sparsely scaly abaxially, base asymmetrical, usually oblique and broadly cuneate, margin ±entire, apex acute; the superior pairs of pinnae connate, terminal with bulbil; veins reticulate and forming a costal areole. Fertile leaves unseen.
In a ravine in semi-original dwarf forest, on the outermost mountain slope from the seaside exposed to wind. Taiwan (Pingtung).
B. ×nanjenensis is very close to B. ×laxireticulata, both having on their rachis narrow wings, but the latter have bigger pinnae 4.5－7 ×1.5－2 cm, margin ±entire to lobed to 2/5 towards the costa, with a distinct spine in each sinus.
1. South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China; 2.Department of Botany, Faculty of Science, University of Kyoto, Kyoto, Japan.
Correspondence to: XING Fu-Wu. E-mail: xinfw@scib.ac.cn