BOLBITIDACEAE [DRAFT]

实蕨科  　shi jue ke

Xing Fuwu 1*, Wang Fa-guo1, Kunio Iwatsuki2

Ferns, small to medium-sized, terrestrial or creeping on rocks or riverbanks or small trees. Rhizome creeping or shortly erect or scandent, dorsiventral, covered with ovate or lanceolate brown or blackish scales. Fronds distant or tufted, dimorphic, simple or mostly pinnate. Sterile fronds: stipe dark brown or blackish near base, with scales ± similar to those on rhizome, usually on basal part only; lamina herbaceous or papery, apex usually with bulbil, margin entire, crenate, or lobed, with or without teeth or spines, veins free or variously anastomosing and forming areolae, with or without included veinlets. Fertile fronds of similar shape as sterile ones, usually with longer stipes and narrower lamina; sori acrostichoid, sporangia spreading throughout abaxial surface.

Two genera of about 90 species; pantropical, mainly in Asia and Pacific Islands; one genus in China; 21 species (nine endemic) plus three putative hybrids (two endemic).

1. BOLBITIS Schott, Gen, Fil. t. 14. 1835.

实蕨属  shi jue shu

Egenolfia Schott; [??add]Poecilopteris C. Presl.

Fronds dimorphic, evergreen, simple, pinnate, or rarely bipinnate. Sterile fronds: stipe with ovate or lanceolate brown scales at base; lamina mostly ± herbaceous, apex usually with a bulbil, 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 fronds of similar shape as sterile fronds, usually with longer stipes and narrower lamina. Sori acrostichoid, without indusia. Spores globose or nearly so, with thick epispores.

1a.     Veins free.

2a.    Sterile fronds pinnate, base asymmetrical (except B. hookeriana), apex obtuse or acute.

3a.     Bulbil terminal on lamina ...................................................................  1. B. rhizophylla

3b.     Bulbil subterminal on lamina.

4a.     Stipe and rachis subglabrous; base of pinnae asymmetrical; sporangia spreading throughout abaxial surface, arrangement fully acrostichoid.

5a.    Sterile pinnae usually obtuse or rounded at apex; fertile pinnae 8–10 Χ 2–3 mm  2. B. appendiculata

5b.    Sterile pinnae acute at apex; fertile pinnae 4–6 Χ ca. 0.8 cm ......  3. B. medogensis

4b.     Stipe and rachis densely scaly; base of pinnae symmetrical; sporangia on all sides of costa and costules, usually arranged moniliform .........................................................  4. B. hookeriana

2b.    Sterile fronds bipinnatifid, base symmetrical, apex acuminate.

6a.     Apex of lobes rounded, lobes 0–1 mm apart.

7a.     Rachis with a narrow wing except for lowest part; base of lowermost pinnae symmetrical  5. B. sinensis

7b.     Rachis without wing; base of lowermost pinnae asymmetrical .........  6. B. longiaurita

6b.     Apex of lobes obtuse-acute, lobes 2–3 mm apart.

8a.     Stipe and rachis densely scaly; lobes of pinnae falcate, 1–1.5 cm ....  7. B. tonkinensis

8b.     Stipe and rachis subglabrous; lobes of pinnae triangular, 0.5–0.6 cm ....  8. B. fengiana

1b.     Veins ± anastomosing.

9a.    Venation pattern ± irregular, veins along costae usually anastomosing and others free or rarely uniting; rachis narrowly winged or not.

10a.   Pinnae pinnatifid; rachis wingless ...............................................  H1. B. Χyunnanensis

10b.   Pinnae subentire or crenate; rachis narrowly winged.

11a.   Pinnae crenate; veins usually free except those along costae ..  H2. B. Χlaxireticulata

11b.   Pinnae subentire; veins anastomosing including those along costae  H3. B. Χnanjenensis

9b.    Venation pattern regular, veins variously anastomosing; rachis wingless.

12a.   Veinlets 5–8 arising from either side of lateral veins uniting in sterile pinnae; fronds often dark brownish when dry.

13a.   Rhizome with 4 rows of fronds; scales on base of stipe with obvious brown margin  19. B. confertifolia

13b.   Rhizome with 2 rows of fronds; scales on base of stipe without obvious brown margin.

14a.   Lamina thickly papery; rhizome 1–1.5 cm in diam., high climbing .  20. B. scandens

14b.   Lamina thinly herbaceous; rhizome 3–6 mm in diam., terrestrial or climbing on bases of tree trunks ...............................................................................................  21. B. heteroclita

12b.   Veinlets 1–4, arising from either side of lateral veins uniting in sterile pinnae; fronds green or rarely purplish when dry.

15a.   Venation without included free veinlets.

16a.   Fertile pinnae linear; lateral pinnae 15–24 pairs .......................  9. B. angustipinna

16b.   Fertile pinnae lanceolate; lateral pinnae 6–15(–17) pairs.

17a.   Bulbils terminal; lateral pinnae ca. 15 pairs; sterile pinnae truncate at base  11. B. tibetica

17b.   Bulbils subterminal; lateral pinnae 6–12 pairs.

18a.   Sterile pinnae truncate to subcordate at base ................  10. B. chirstensenii

18b.   Sterile pinnae rounded at base .......................................  12. B. hekouensis

15b.   Venation with included free veinlets.

19a.   Lateral pinnae 6–15 pairs; terminal segment triangular .............  13. B. subcordata

19b.   Lateral pinnae 2–7 pairs; terminal segment conform[??] to central pinnae.

20a.   Fertile pinnae linear, 14–45 Χ longer than wide.

21a.   Fronds purplish when dry; veinlets 5–8 arising from either side of lateral veins uniting in sterile pinnae ................................................................................  17. B. costata

21b.   Fronds greenish when dry; veinlets 4 arising from either side of lateral veins uniting in sterile pinnae ..................................................................................  18. B. virens

20b.   Fertile pinnae lanceolate, 3–8 Χ longer than wide.

22a.   Sterile pinnae 2 or 3 pairs; acrostichoid .........................  14. B. hainanensis

22b.   Sterile pinnae 5–7 pairs; fertile pinnae not acrostichoid.

23a.   Fronds purplish when dry; veinlets 2 or 3 arising from lateral veins, free veinlets without thickened ends .........................................................  15. B. scalpturata

23b.   Fronds stramineous when dry; veinlets 4 or 5 arising from lateral veins; free veinlets with thickened ends .............................................................  16. B. deltigera

1. Bolbitis rhizophylla (Kaulfuss) Hennipman, Blumea 18: 148. 1970.

根叶刺蕨  gen ye ci jue

Gymnogramma rhizophylla Kaulfuss, Enum. Filic. 78. 1824; Egenolfia rhizophylla (Kaulfuss) Fιe.

Rhizome shortly creeping, 4–6 mm in diam., with 2 or 3 rows of fronds, densely scaly; scales brown, ovate-lanceolate, subclathrate. Fronds close together; sterile fronds pinnate, 20–90 cm long; stipe 10–20 cm, base 2–3 mm in diam.; lamina green to blackish, dark green when dry, lanceolate, pinnate, 40–80 Χ 5–13 cm, herbaceous, both surfaces glabrous; stipe and lower side of basal part of rachis usually with small, blackish, spreading scales; terminal segment 1–6 cm; rachis with a narrow wing except for lowest part; pinnae 20–35 pairs, alternate to opposite, sessile, up to 3.5 cm apart, elliptic-lanceolate, 2.5–4 Χ 1–1.5 cm, base rounded-truncate, ± symmetrical, margin finely serrate, or serrately lobed to 1/3 toward costa, apex rounded or acuminate, two lowermost pinnae deflexed or not; veinlets 2–4 pairs, simple pinnate. Fertile fronds 16–30 cm; lamina lanceolate, 16–20 Χ 1.5–2.5 cm, widest usually near middle; pinnae 20–25 pairs, elliptic, ca. 1.5 cm apart, 6–12 Χ 3–4 mm, margin entire or irregularly undulate, apex obtuse. Sporangia inserted throughout abaxial surface. 2n = 82*.

* On rocks in dense forests; 400–1000 m. Taiwan (Tainan).

2. Bolbitis appendiculata (Willdenow) K. Iwatsuki, Acta Phytotax. Geobot. 18: 48. 1959.

刺蕨  ci jue

Acrostichum appendiculatum Willdenow, Sp. Pl., ed. 4, 5: 114. 1810; Egenolfia crenata Ching & P. S. Chiu.

Rhizome shortly creeping, up to 3 mm in diam., with 2 or 3 rows of fronds, densely scaly; scales blackish, lanceolate, 2–3 Χ 1–1.5 mm, opaque or subclathrate, margin with irregular small teeth. Fronds close together; sterile fronds pinnate, 15–40 cm; stipe 5–15 cm, ca. 1 mm in diam., sparsely scaly near base; lamina dark green when dry, lanceolate, 12–25 Χ 3.5–6 cm, herbaceous; terminal segment triangular, apex acuminate, sometimes somewhat prolonged; rachis usually with a narrow wing; pinnae 15–30 pairs, opposite or alternate, elliptic-linear, up to 3 cm apart, 2–3.5 Χ 5–8 cm, base asymmetrical, usually oblique, margin ± entire, finely serrate, or lobed to 1/3 toward costa, apex acute or acuminate, two lowermost pinnae 1–2 mm stalked; secondary veins on either side with 2 or 3 tertiary veins. Fertile fronds pinnate, 25–40 cm; stipe 20–25 cm; pinnae 10–30, 8–10 Χ 2–3 mm; two lowermost pinnae 1–9 mm stalked. Sporangia inserted throughout abaxial surface, arrangement fully acrostichoid.

Creeping on rocks of dense forest understories, on soil in evergreen forests; 100–1300 m. Guangdong, Guangxi, Hainan, Taiwan, Yunnan [Bangladesh, Bhutan, India, Indonesia, Japan, Laos, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, Vietnam].

3. Bolbitis medogensis (Ching & S. K. Wu) S. Y. Dong, Acta Phytotax. Sin. 43: 105. 2005.

墨脱刺蕨  mo tuo ci jue

Egenolfia medogensis Ching & S. K. Wu, Fl. Xizang. 1: 278. 1983.

Rhizome shortly creeping, densely scaly; scales brown, linear-lanceolate, entire. Fronds close together or tufted; sterile fronds pinnate, ca. 75 cm; stipe 20–35 cm, sparsely scaly; lamina green when dry, broadly lanceolate, ca. 48 Χ 20–25 cm, both surfaces glabrous, apex caudate-acuminate, bulbil subterminal; pinnae ca. 12 pairs, alternate, shortly stalked, 3–5 cm apart, 10–13 Χ ca. 3.5 cm, base nearly truncate, margin crenate-serrate, with a fleshy spine in sinus; secondary veins slightly obvious, dichotomous. Fertile fronds pinnate; stipe ca. 50 cm; lamina narrowly lanceolate, ca. 46 Χ 10–12 cm; pinnae 10–12 pairs, subsessile, linear, 4–6 Χ ca. 0.8 cm, entire. Sporangia inserted throughout abaxial surface.

* Evergreen forests on slopes; 700–900 m. Xizang (Mκdog).

4. Bolbitis hookeriana K. Iwatsuki, Acta Phytotax. Geobot. 18: 49. 1959.

虎克实蕨  hu ke shi jue

Bolbitis appendiculata (Willdenow) K. Iwatsuki subsp. vivipara (Buchanan-Hamilton ex Hooker) Hennipman; Polybotrya vivipara Buchanan-Hamilton ex Hooker.

Rhizome long creeping, 5–12 cm in diam., with usually 3 rows of fronds, without or with few scleraenchyma strands; sterile fronds pinnate, 20–100 cm; stipe and rachis usually throughout with spreading scales or sometimes glabrescent; stipe 5–25 cm; lamina 20–85 Χ 4–20 mm, terminal segment 3–10 cm; pinnae 33–60 pairs, 3.5–10 Χ 1.2–2.5 cm, base ± symmetrical, margin usually entire, sometimes lobed to 1/5 toward costa; [??pinnules] 3–5 pairs, alternate, linear-lanceolate, usually widest at (to below) middle, 15–25 Χ 2–3 cm, base ± symmetrical, broadly attenuate or truncate, margin slightly serrate-crenate, with a short spine in each sinus, apex acuminate; two lowermost pinnae with short stalk; secondary veins on either side with 2 or 3 tertiary veins. Fertile fronds 40–90 cm; stipe 16–45 cm; lamina 14–40 Χ 4–10 cm; pinnae 35–50 pairs; central pinnae 2–8.5 cm; lowermost two pinnae 0–4 mm stalked. Sporangia inserted on all sides of costa and costules, usually arranged moniliform.

On rocks in dense forests; ca. 500 m. Yunnan (Yingjiang) [Bangladesh, India, Indochina Peninsula[??countries], Myanmar].

5. Bolbitis sinensis (Baker) K. Iwatsuki, Acta Phytotax. Geobot. 18: 49. 1959.

中华刺蕨  zhong hua ci jue

Acrostichum sinense Baker, Bull. Misc. Inform. Kew 1906: 14. 1906; Egenolfia crassifolia Ching; E. sinensis (Baker) Maxon.

Rhizome shortly creeping, ca. 1 cm in diam., with 2 or 3 rows of fronds; scales brownish or blackish, ovate-lanceolate, subclathrate, ca. 3 mm. Fronds close together; sterile fronds 20–110 cm; stipe 10–35 cm, base ca. 3 mm in diam., grooved adaxially; lamina green to blackish, dark green when dry, ovate-lanceolate, 40–110 Χ 10–35 cm, herbaceous, both surfaces glabrous, pinnate; terminal segment 8–30 cm; rachis with a narrow wing except for lowest part, with sparse dark brown, ovate scales; pinnae 14–30 pairs, alternate to opposite, elliptic, sessile, 2–3 cm apart, 10–14 Χ 2–2.5 cm, base truncate, ± symmetrical, margin lobed to 1/3–2/3 toward costa, apex rounded or acuminate; lobes 16–20 pairs, ca. 1 mm spaced, 8–12 Χ 4–5 mm, margin usually entire or finely crenate-serrate, apex rounded, with a spine in sinus; veinlets 6 or 7 pairs, pinnate, simple or biforked. Fertile fronds 16–80 cm; stipe as long as sterile ones; lamina lanceolate, 22–30 Χ 4–5 cm, apex caudate; pinnae 10–14 pairs, 2–3 cm apart, narrowly lanceolate, 4–5 Χ ca. 0.5 cm, base truncate to slightly cordate, apex acuminate, basal pairs 3–4 mm stalked. Sporangia inserted throughout abaxial surface. 2n = 82.

In soil or on rocks in dense forests; 600–1900 m. 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: 434. 2006.

长耳刺蕨  chang er ci jue

Acrostichum appendiculatum Willdenow var. costulatum Hooker; Bolbitis bipinnatifida (J. Smith) K. Iwatsuki, Acta Phytotax. Geobot. 18: 49. 1959, nom. illegit., non (Kuhn[??Mett.]) Ching, 1934; B. sinensis (Baker) K. Iwatsuki var. costulata Tagawa & K. Iwatsuki; Egenolfia bipinnatifida J. Smith.

Rhizome shortly 　creeping, ca. 1 cm in diam., densely scaly; scales dark brownish, ovate-lanceolate, ca. 3 mm. Fronds close together; sterile fronds 40–70 cm; stipe 11–15 cm, base ca. 3 mm in diam., grooved adaxially, sparsely scaly; lamina dark green when dry, elliptic, 50–60 Χ 10–35 cm, widest at base, papery, both surfaces glabrous, bipinnate; terminal segment long attenuate to flagelliform; rachis shallowly grooved, glabrous; pinnae ca. 20 pairs, 14–16 Χ 2.5–3.5 cm, lanceolate, basal pairs nearly opposite, with short stalk, two lowermost pinnae 3–6 mm stalked, apex caudate-acuminate, base asymmetrical with basiscopic 2 or 3 lobes undeveloped, other lobes longer than acroscopic ones, other pairs of pinnae lobed to 1/2 to costa, base truncate and symmetrical; lobes 18–20, elliptic, 1–1.5 Χ ca. 0.6 cm, margin sparsely rounded dentate, with a spine in sinus; veinlets 6 or 7 pairs, free, biforked or simple. Fertile fronds: stipe 20–40 cm; lamina lanceolate, pinnate, 18–20 Χ 4–5 cm; pinnae 12–14 pairs, elliptic, 2–3 Χ ca. 0.5 cm, base rounded-truncate, apex obtuse, basal pairs with short stalk. Sporangia inserted throughout abaxial surface.

* On rocks in dense forests; ca. 1200 m. Yunnan.

This species is similar to Bolbitis sinensis, but differs in its rachis without wing, the base of the two lowermost pinnae asymmetrical with its basiscopic 2 or 3 lobes undeveloped, and the other lobes longer than the acroscopic ones.

7. Bolbitis tonkinensis (C. Christensen ex Ching) K. Iwatsuki, Acta Phytotax. Geobot. 18: 49. 1959.

镰裂刺蕨  lian lie ci jue

Egenolfia tonkinensis C. Christensen ex Ching, Bull. Fan Mem. Inst. Biol. 2: 306. 1931.

Rhizome shortly creeping, ca. 1 cm in diam., densely scaly; scales fuscous, ovate, ca. 3 mm, margin erose. Fronds close together; sterile fronds ca. 80 cm; stipe 20–30 cm, ca. 3 mm in diam. at base, grooved adaxially, covered with ovate-lanceolate fuscous scales; lamina dark green when dry, triangular-ovate, 50–60 Χ 20–30 cm, both surfaces glabrous, bipinnate, apex caudate-acuminate, bulbil subterminal; pinnae 12–16 pairs, alternate, with short stalk 2–4 mm, lanceolate, 13–15 Χ 3–3.5 cm, base circularly truncate and slightly asymmetrical, margin lobed to 2/3 to costa, apex caudate-acuminate, bulbil subterminal and small; terminal segment prolonged to flagelliform and pinnatifid, bulbil subterminal and large; lobes 16–18 pairs, 2–3 mm apart, elliptic-lanceolate, falcate, 10–15 Χ 5–6 mm, with a long spine in sinus; veinlets 6 or 7 pairs, pinnate, simple or biforked. Fertile fronds pinnate; stipe 40–45 cm; lamina narrowly lanceolate, ca. 25 Χ 6 cm; pinnae ca. 10 pairs, elliptic-lanceolate, 2.5–3 Χ 0.5 cm, entire. Sporangia inserted throughout abaxial surface.

On wet rocks in forests; 200–1200 m. S Yunnan [Thailand, Vietnam (Tonkin)].

8. Bolbitis fengiana (Ching) S. Y. Dong, Acta Phytotax. Sin. 43: 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 in diam., densely scaly; scales blackish brown, lanceolate, ca. 2 mm. Fronds close together; sterile fronds 40–60 cm; stipe 15–25 cm, base 2–3 mm in diam., grooved adaxially, sparsely scaly; lamina dark green when dry, broadly lanceolate, 22–30 Χ 10–15 cm, herbaceous, both surfaces glabrous, base rounded-truncate, bipinnate, apex long caudate, bulbil subterminal and small; rachis with a narrow wing; pinnae 10–15 pairs, alternate, basal pairs with short stalk 2–3 mm, linear-lanceolate, ca. 1 cm apart, 5–9 Χ 1–1.5 cm, base truncate, lobed to 1/3 toward costa; lobes 8–12 pairs, triangular, 5–6 Χ 5–6 mm, margin with sparse short teeth, with long acute spine in sinus; veinlets 3 pairs, simple pinnate. Fertile fronds pinnate; stipe 25–40 cm; lamina lanceolate, ca. 16 Χ 4–5 cm; pinnae 10–12 pairs, elliptic to elliptic-lanceolate, 1.5–2 cm apart, 2–2.5 Χ 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 throughout abaxial surface.

* Dense forests; 1300–1800 m. Yunnan.

9. Bolbitis angustipinna (Hayata) H. Ito, J. Jap. Bot. 14: 443. 1938.

多羽实蕨  duo yu shi jue

Leptochilus angustipinnus Hayata, Icon. Pl. Formosan. 5: 297. 1915; Acrostichum contaminans Wallich, nom. nud.; Bolbitis contaminans (Wallich[??]) Ching; L. cuspidatus (C. Presl) C. Christensen var. crenatus Ros.[??author name].

Rhizome long creeping, 10–16 cm, robust, 1–2 cm in diam., densely scaly, with 2–4 rows of fronds; scales dark brown or blackish, lanceolate, 4–6 mm, subclathrate. Fronds close together; sterile fronds pinnate, 55–145 cm; stipe 40–60 cm, near base ca. 8 mm in diam., sparsely scaly, with 7–16 vascular bundles; lamina gray-green when dry, elliptic, 60–85 Χ 30–45 cm, widest somewhat below middle, hard herbaceous; pinnae 12–20 pairs, alternate, linear-lanceolate, usually widest at (to below) middle, 15–25 Χ 2–3 cm, base ± symmetrical, broadly attenuate or truncate, margin slightly serrate-crenate, with a short spine in each sinus, apex acuminate, two lowermost pinnae with short stalk; terminal segment joined to rachis, usually narrowly triangular, ± conform[??] to central pinnae, 10–20 cm, bulbil subterminal; rachis not winged; secondary veins prominent; veins forming a costal areole and 3 rows distal areoles, without included free veinlets. Fertile fronds 50–150 cm; stipe 30–85 cm; lamina 70–90 Χ 15–25 cm; pinnae 80–120 Χ 5–8 mm. Sporangia inserted usually throughout abaxial surface, arrangement fully acrostichoid, sometimes either with a sterile strip along costa.

On rocks or near streams in dense forests; 200–1400 m. Hainan, Taiwan, Yunnan [Bhutan, India, Myanmar, Nepal, Sri Lanka, Thailand].

10. Bolbitis christensenii (Ching) Ching in C. Christensen, Index Filic., Suppl. 3: 47. 1934.

贵州实蕨  gui zhou shi jue

Campium chistensenii Ching, Bull. Fan Mem. Inst. Biol. 2: 214. 1931.

Rhizome shortly creeping, robust, ca. 1 cm in diam., densely scaly; scales blackish brown, lanceolate, ca. 9 mm, margin maple[??colored], entire. Fronds tufted; sterile fronds pinnate; stipe 30–40 cm, base ca. 4 mm in diam., base densely covered with lanceolate scales; lamina grayish brown when dry, elliptic, 40–60 Χ20–25 cm, thinly herbaceous, apical part pinnatipartite into lanceolate lobes, bulbil subterminal; rachis not winged, densely covered with lanceolate brown scales especially when young; pinnae 6–10 pairs, basal ones nearly opposite, with short stalk 2–3 mm, lanceolate, 6.5–12 Χ1.2–2 cm, base circular, margin crenate-serrate, sometimes with an obtuse tooth in sinus, apex long acuminate; secondary veins inconspicuous; veins forming 3 rows of long and narrow areoles, without included free veinlets. Fertile fronds as long as sterile fronds; lamina obviously narrower, linear-lanceolate; pinnae subsessile, 3–4 Χ ca. 5 mm, base circularly cuneate, entire, apex acuminate. Sporangia inserted throughout abaxial surface.

Near streams in forests; 400–1100 m. Guangxi, Guizhou [Vietnam (Tonkin)].

11. Bolbitis tibetica Ching & S. K. Wu, Fl. Xizang. 1: 276. 1983.

西藏实蕨  xi zang shi jue

Rhizome short and robust, ca. 1.5 cm in diam., densely scaly; scales nut brown, 4–5 mm, narrowly lanceolate, entire. Fronds tufted, 80–110 cm; sterile fronds pinnate; stipe stramineous, ca. 25 cm, base 4–5 mm in diam., grooved adaxially, sparsely scaly; lamina sap green when dry, broadly lanceolate, 50–65 Χ25–30 cm, herbaceous, both surfaces glabrous, apex pinnatilobate and elongate to flagelliform, bulbil terminal; rachis not winged, with sparse nut brown scales; pinnae ca. 15 pairs, alternate, 1–1.5 cm apart, lanceolate, 16–18 Χ 2.5–3 cm, lower pairs of pinnae shortly stalked, upper pairs basally connate with rachis, base truncate, margin with obtuse tooth, apex caudate-acuminate; veins prominent on both surfaces; veins forming a costal areole and 3 or 4 rows tetragonal to hexagonal areoles, without included free veinlets. Fertile fronds 95–115 cm; stipe 50–55 cm; lamina 50–60 Χ 10–12 cm, apex pinnatilobate; pinnae narrowly lanceolate, 6–8 Χ 1.5–1.8 cm, base circularly truncate, apex acuminate. Sporangia inserted throughout abaxial surface. [Chromosomes.]

* Broad-leaved forests; ca. 800 m. Xizang (Mκdog).

12. Bolbitis hekouensis Ching, Acta Phytotax. Sin. 21: 212. 1983.

河口实蕨  he kou shi jue

Rhizome robust and shortly creeping, 1–1.5 cm in diam., densely scaly; scales dark brown, lanceolate, ca. 4 mm, entire. Fronds close together, 28–51 cm; sterile fronds pinnate; stipe pale brown, 8–17 cm, base ca. 3 mm in diam., grooved adaxially, densely covered with lanceolate, brown scales; lamina grayish brown when dry, elliptic, 18–32 Χ9.5–15 cm, thickly papery, apical part pinnatipartite into lanceolate lobes, bulbil subterminal; rachis not winged, densely covered with lanceolate, brown scales especially when young; pinnae 6–10 pairs, basal ones nearly opposite, with short stalk 2–3 mm, lanceolate, 6.5–12 Χ 1.2–2 cm, base circular, margin crenate-serrate, sometimes with an obtuse tooth in each sinus, apex long acuminate; secondary veins inconspicuous; veins forming 3 rows of long and narrow areoles, without included free veinlets. Fertile fronds as long as sterile fronds; lamina obviously narrower, linear-lanceolate; pinnae 3–4 Χ ca. 5 mm, base circularly cuneate, subsessile, entire, apex acuminate. Sporangia inserted throughout abaxial surface.

* Rocky limestone crevices in rainforests; 400–1100 m. Hainan, Yunnan.

13. Bolbitis subcordata (Copeland) Ching in C. Christensen, Index Filic., Suppl. 3: 50. 1934.

华南实蕨  hua nan shi jue

Campium subcordatum Copeland, Philipp. J. Sci. 37: 369. 1928; Bolbitis media Ching & C. H. Wang; B. subcordata var. media (Ching & C. H. Wang) S. H. Wu.

Rhizome shortly creeping, 8–15 mm in diam., covered with lanceolate, clathrate, subentire, brown or blackish scales. Fronds ± close together, 60–120 cm; sterile fronds pinnate, 20–110 Χ 15–28 cm; stipe 8–45 cm, densely scaly near base, bearing small scales at apex, with 5–9 vascular bundles; lamina dark green or brownish, 12–65 cm, widest below middle, herbaceous, terminal segment narrowly triangular, longer to shorter than central pinnae, base trifid; rachis not winged; pinnae 6–14 pairs, opposite below middle, base broadly cuneate to subcordate, margin usually lobed to 1/3 toward costa, crenate-serrate, with a prominent tooth in each sinus between 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 fronds 30–125 cm; stipe 15–70 cm; pinnae 6–25, 2–6 cm apart, 4–8 Χ ca. 1 cm; two lowermost pinnae 1–7 mm stalked. Sporangia inserted throughout abaxial surface, arrangement acrostichoid. 2n = 82.

In forests, near streams; 100–600 m. Fujian, Guangdong, Guangxi, Hainan, Jiangxi, Taiwan, 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 shortly creeping, 5–9 cm, 1–1.5 cm in diam., densely scaly; scales dark brown, lanceolate, ca. 3 mm, entire. Fronds close together, 4–7 mm apart; sterile fronds pinnate; stipe 20–40 cm, near base ca. 3.5 mm in diam., densely scaly, glabrous upward; lamina grayish brown when dry, ovate, 25–35 Χ 20–25 cm, hard papery, both surfaces glabrous; pinnae 3–5 pairs, alternate, elliptic-lanceolate, 16–25 Χ 4.5–6 cm, base narrowly cuneate, margin wrinkled-undulate, apex caudate-acuminate, two lower pairs of pinnae with stalk ca. 1 cm; terminal segment ± conform[??] to central pinnae; rachis not winged; secondary veins tenuous[??], prominent abaxially; veins inconspicuous, forming a costal large areole and 3 rows of small areoles, sometimes with included free short veinlets. Fertile fronds 35–75 cm; stipe 30–55 cm; lamina narrowly ovate, 12–25 Χ 10–14 cm; pinnae 2–5 pairs, lanceolate, 6–10 Χ 1–1.5 cm, base narrowly cuneate, apex acuminate. Sporangia inserted usually throughout abaxial surface, arrangement acrostichoid.

* Dense forests; 700–1100 m. Hainan, Yunnan.

[New Name: Wang Faguo, K. Iwatsuki & Xing Fuwu. 2008. A new name of Bolbitis from China. Amer. Fern J. 98: 96-97.
Bolbitis x multipinna F. G. Wang, K. Iwatsuki & F. W. Xing, Amer. Fern J. 98: 97. 2008, nom. nov.
(Bolbitis yunnanensis Ching)]

15. Bolbitis scalpturata (Fιe) Ching in C. Christensen, Index Filic., Suppl. 3: 50. 1934.

红柄实蕨  hong bing shi jue

Heteroneuron scalpturatum Fιe, Mιm. Foug. 95. 1845.

Rhizome long creeping, up to 10 cm, up to 10 mm thick, with 2 rows of fronds, densely scaly; scales blackish, lanceolate, subclathrate. Fronds close together; sterile fronds pinnate; stipe 15–24 cm, sparsely scaly near base, with 5–9 vascular bundles; lamina purplish or purplish brown, 20–55 Χ 7–30 cm, widest below middle, herbaceous to subleathery, terminal segment usually subarticulate, 9–20 cm, bulbil subterminal; rachis not winged; pinnae 2–12 pairs, opposite or alternate, 1.5–4.5 cm apart, 10–12 Χ 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 areoles with few to several excurrent included free veins. Fertile fronds 30–60 cm; stipe 15–40 cm; pinnae 5–20, 2–6 cm apart, 4–8 Χ ca. 1 cm; two lowermost pinnae sessile or up to 6 mm stalked. Sporangia inserted throughout abaxial surface, or along margin only, arrangement pteridoid.

On rocks in forests; sea level up to 1200 m. Hainan, Taiwan [Indonesia, Malaysia, Myanmar, Philippines, Thailand, Vietnam].

16. Bolbitis deltigera (Beddome) C. Christensen, Index Filic., Suppl. 3: 48. 1934.

间断实蕨  jian duan shi jue

Poecilopteris costata (C. Presl) Moore var. deltigera Beddome, Ferns Brit. India 1: t. 114. 1865; Bolbitis virens (Wallich ex Hooker & Greville) Schott var. deltigera (Beddome) Hennipman.

Rhizome 10–15 cm, up to 14 mm in diam., with 2 or 3 rows of fronds, covered with lanceolate (5–7 Χ ca. 1.5 mm), subclathrate, blackish scales. Sterile fronds pinnate, 40–115 cm; stipe 20–60 cm, 2–7 mm in diam. near base, with 7–14 vascular bundles; lamina purplish or purplish brown, 20–75 Χ15–50 cm, widest at or below middle, herbaceous to leathery, terminal segment 14–40 cm, subarticulate or joined to rachis, bulbil subterminal; rachis not winged; pinnae 6–20 pairs, opposite or alternate, up to 6 cm apart, 10–30 Χ 2.5–6 cm, base ± symmetrical, usually narrowly, sometimes broadly attenuate or cuneate, margin usually ± entire or serrate-crenate, apex acuminate, two lowermost pinnae 1–10 mm stalked; veins forming a costal areole and several equally large distal areoles, 4 or 5 veins arising from each side of lateral veins anastomosing. Fertile fronds 40–120 cm; stipe 25–60 cm; pinnae 7–18, up to 6 cm apart, central pinnae 5–17 Χ 1.5–4.5 cm; two lowermost pinnae 1–9 mm stalked. Sporangia inserted on veins and near margin only, arrangement not acrostichoid.

Understories of rainforests in valleys or on slopes; 300–700 m. Hainan, Yunnan [Bangladesh, Bhutan, India, Myanmar, Nepal, Thailand].

17. Bolbitis costata (C. Presl) Ching in C. Christensen, Index Filic., Suppl. 3: 47. 1934.

紫轴实蕨  zi zhou shi jue

Campium costatum C. Presl, Tent. Pterid. 239. 1836.

Rhizome up to 10 cm, up to 15 mm in diam., with 2–4 rows of fronds, covered with lanceolate (ca. 5 Χ 1 mm), subclathrate, light brown scales. Sterile fronds pinnate, 55–120 cm; stipe 20–60 cm, 2–7 mm in diam. near base, with 5–20 vascular bundles; lamina purplish when dried, 35–75 Χ 20–40 cm, widest at or below middle, subleathery, terminal segment 20–40 cm, bulbil subterminal on terminal segment or pinnae; rachis not winged; pinnae 3–19 pairs, opposite or alternate, up to 8 cm apart, 16–45 Χ 4–9 cm, base symmetrical, acute to subcordate, margin entire or crenulate, sometimes crisped, apex rather abruptly narrowing into acute or acuminate tip, two lowermost pinnae 1–6 mm stalked; veins for greater part forming 4 or more regular rows of areoles, within part of areoles 1–3 excurrent included free vein ending in a thickening. Fertile fronds 40–170 cm; stipe 20–100 cm; lamina 15–85 Χ 7–30 cm, terminal segment 8–20 cm; pinnae 5–18, up to 11 cm apart, ca. 22 Χ 0.6–2 cm; two lowermost pinnae 1–6 mm stalked. Sporangia inserted mainly on veins, arrangement usually acrostichoid, rarely gymnogrammoid.

In forests near streams; sea level up to 1200 m. Yunnan [Bangladesh, India, Myanmar, Nepal, Thailand].

Bolbitis costata is similar to B. scalpturata in its fronds usually turning purplish when dry; it differs by having broader sterile pinnae (to 4–9 cm), 5–8 veinlets arising from each side of lateral veins, and fertile pinnae linear (14–16 Χ as long as wide).

18. Bolbitis virens (Wallich ex Hooker & Greville) Schott, Gen. Fil. ad t. 13. 1834.

宽羽实蕨  kuan yu shi jue

Acrosticum virens Wallich ex Hooker & Greville, Icon. Filic. t. 221. 1831; Bolbitis latipinna Ching.

Rhizome up to 10 cm, up to 13 mm in diam., with 2 or 3 rows of fronds, densely scaly; scales brownish, narrowly lanceolate, ca. 4 Χ 1.5 mm, subclathrate. Sterile fronds pinnate, 40–110 cm; stipe 20–60 cm, 2–7 mm in diam. near base, with 7–14 vascular bundles; lamina 25–75 Χ 20–50 cm, widest at or below middle, herbaceous to leathery, terminal segment 15–40 cm, subarticulate or joined to rachis, somewhat prolonged, bulbil subterminal on terminal segment or pinnae; rachis not winged; pinnae 6–22 pairs, opposite or alternate, up to 6.5 cm apart, 10–30 Χ 2.5–6 cm, base ± symmetrical, sometimes broadly attenuate or cuneate, margin entire or serrate or serrate-crenate, apex acuminate, two lowermost pinnae 1–10 mm stalked; veins forming a costal areole and several to many equally large distal ones. Fertile fronds 40–120 cm; stipe 25–70 cm; lamina 15–65 Χ 7–30 cm, terminal segment 4–19 cm; pinnae 7–18, up to 6 cm apart, 4–20 Χ 0.2–4.5 cm; two lowermost pinnae 1–6 mm stalked. Sporangia inserted throughout abaxial surface, arrangement various.

On rocks in forests; 800–900 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. 12 cm, ca. 1.5 cm in diam., densely scaly; scales medium brown, ovate-lanceolate, ca. 3 cm, margin brown, subentire. Fronds densely clustered on upper side of rhizome; sterile fronds elliptic, pinnate; stipe 20–25 cm, densely covered with scales as on rhizome, grooved adaxially; lamina dark green when dry, 35–45 Χ ca. 15 cm, herbaceous, terminal segment elongate to long flagelliform, terminal part with bulbil; pinnae 4 or 5 pairs, subsessile, lanceolate, 12–15 Χ 2.5–3 cm, base cuneate, margin repand, apex long acuminate; secondary veins prominent abaxially; veins forming nearly hexagonal areoles, without included free veinlets. Fertile fronds not seen.

* Rainforests. 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 more than 2–3 m in height, 1–1.5 cm in diam., densely scaly; scales nut brown, lanceolate, 3–4 mm, margin maple[??colored], entire. Fronds close together; sterile fronds pinnate; stipe 16–20 cm, near base ca. 4 mm in diam., grooved adaxially, sparsely scaly below middle; lamina gray-brown when dry, ovate, 35–40 Χ 20–25 cm, widest somewhat below middle, thickly papery; pinnae 2 or 3 pairs, subsessile, two lowermost pinnae nearly opposite, elliptic-lanceolate, 15–18 Χ 4–5 cm, base circularly cuneate, margin slightly undulate, apex caudate-acuminate; terminal segment larger than central pinnae; secondary veins prominent abaxially, parallel and 6–7 mm apart; veins inconspicuous on both surfaces, forming a narrow costal areole and 3 or 4 rows of distal areoles, sometimes with included free veinlets. Fertile fronds 40–60 cm; lamina narrowly ovate, 25–30 Χ 10–12 cm; pinnae 3 or 4 pairs, subsessile, broadly lanceolate, 10–12 Χ 2–2.5 cm, apex acuminate. Sporangia inserted throughout abaxial surface, arrangement fully acrostichoid.

* Climbing on tree trunks in forests; ca. 700 m. S Yunnan.

Bolbitis scandens is similar to B. heteroclita; it differs in having the rhizome climbing on tree trunks, lateral pinnae of sterile fronds usually 3-paired, terminal pinnae never obviously prolonged, and leaf texture thickly papery.

21. Bolbitis heteroclita (C. Presl) Ching in C. Christensen, Index Filic., Suppl. 3: 48. 1934.

长叶实蕨  chang ye shi jue

Acrostichum heteroclitum C. Presl, Rel. Haenk. 1(1): 15, t. 2. 1825[??1885]; Bolbitis annamensis Tardieu & C. Christensen.

Rhizome robust and long creeping, 3–6 mm in diam., densely scaly; scales brownish or blackish, ovate-lanceolate, 3–9 Χ 0.5–1.5 cm, usually subclathrate, subentire. Fronds ± spaced, 1–3 cm apart; sterile fronds simple or pinnate, 15–80 cm; stipe 10–30(–40) cm, sparsely scaly, grooved adaxially; lamina blackish when dry, 10–60 Χ 5–25 cm, thinly herbaceous, apex elongate to long flagelliform, terminal part with bulbil; rachis not winged, with sparse nut brown scales; pinnae 1–6 pairs, alternate to opposite, subsessile, broadly lanceolate, 10–19 Χ 3–6 cm, base circularly cuneate, margin subentire or crenate-serrate, apex caudate-acuminate; secondary veins prominent on both surfaces; veins forming 3 rows of regularly tetragonal to hexagonal areoles, without included free veinlets. Fertile fronds 15–70 cm; stipe 10–45 cm; lamina 10–26 Χ 5–20 cm. Sporangia inserted throughout abaxial surface. 2n = 82.

On rocks or at base of trees near streams in broad-leaved forests; sea level up to 1500 m. Chongqing, Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, Yunnan [Asian tropics: list countries].

Putative Hybrids

The following three forms are putative hybrids recorded from China.

H1. Bolbitis Χyunnanensis (Ching & C. H. Wang) F. G. Wang, K. Iwatsuki & F. W. Xing, comb. nov. [not Bolbitis yunnanensis Ching, Acta Phytotax. Sin. 21: 214. 1983]

云南刺蕨  yun nan ci jue

Egenolfia Χ yunnanensis Ching & P. S. Chiu ex Ching & C. H. Wang, Acta Phytotax. Sin. 1: 216. 1983; Bolbitis angustipinna (Hayata) H. Ito [Χ] sinensis Hennipman.

Rhizome creeping, 1–1.5 cm in diam., densely scaly; scales dark brownish, ovate-lanceolate, 2–3 mm. Fronds close together; sterile fronds 70–90 cm; stipe 10–35 cm, base covered with scales, ca. 3 mm in diam., grooved adaxially; lamina dark green when dry, ovate-lanceolate, 50–65 Χ 15–20 cm, papery, glabrous, bipinnate, terminal segment prolonged to flagelliform, bulbil subterminal and large; rachis with a narrow wing on upper part; pinnae ca. 20 pairs, alternate, lanceolate, 1–1.5 cm apart, 20–22 Χ ca. 3 cm, basal pairs with short stalk 3–4 mm, base broadly cuneate, margin lobed to 1/2 toward costa, apex long caudate; lobes 22–26 pairs, elliptic, ca. 1 mm spaced, 8–12 Χ ca. 5 mm, margin usually entire or slightly undulate, apex rounded, with a long spine in sinus; veinlets 5 or 6 pairs, simple pinnate, sometimes veins forming a costal areole. Fertile fronds: stipe ca. 35 cm; lamina ca. 20 cm; pinnae lanceolate, 4–6 Χ 0.6–0.9 cm, with short stalk, apex obtuse. Sporangia inserted throughout abaxial surface.

* In soil in dense rainforests; 800–900 m. Guangxi, Yunnan.

H2. Bolbitis Χlaxireticulata K. Iwatsuki, Acta Phytotax. Geobot. 18: 50. 1959.

网脉实蕨  wang mai shi jue

Bolbitis appendiculata (Willdenow) K. Iwatsuki [x] B. subcordata (Copeland) Ching; Egenolfia laxireticulata (K. Iwatsuki) C. M. Kuo.

Rhizome shortly creeping, ca. 4 mm in diam., with scleraenchyma strands, sparsely scaly; scales brown-black, ovate-lanceolate to lanceolate, subclathrate, 3–4 Χ 1.5–2 mm. Fronds dimorphic; sterile fronds pinnate, 30–75 cm; stipe 14–30 cm, densely scaly; lamina grayish green when dry, oblong-elliptic, widest below or about middle, 20–30 Χ 10–13 cm, papery, with a spherical subterminal bulbil; rachis sparsely scaly, with a narrow wing; pinnae alternate or opposite, 12–20 pairs, 4.5–7 Χ1.5–2 cm, both surfaces glabrous, base sometimes symmetrical, variously cuneate, usually asymmetrical, margin ± entire to lobed to 2/5 toward costa, lobes finely crenate-serrate, with a distinct spine in sinuses, apex acute to acuminate, two lowermost pinnae up to 1.5 mm stalked, terminal segment triangular; veins very irregular, forming a costal areole at least in part of pinnae of one plant[??]. Fertile fronds: stipe 25–40 cm; lamina 14–28 cm; pinnae 10–12 pairs, oblong-lanceolate, ca. 2 Χ 0.4 cm, entire or undulate. Sporangia inserted mainly on veins.

On rocks along streams in rainforests; 500–700 m. Guangdong, Hainan, Taiwan [Japan (Ryukyu Islands)].

H3. Bolbitis Χnanjenensis C. M. Kuo, Stud. Ferns Ken-Ting Nation. Park (Taiwan) 53. 1986; Bot. Bull. Acad. Sin. 31: 308. 1990.

南仁实蕨　  nan ren shi jue

Bolbitis appendiculata (Willdenow) K. Iwatsuki [Χ] B. heteroclita (C. Presl) Ching.

Rhizome creeping, ca. 4 mm in diam., densely scaly; scales dark brown, ovate-lanceolate, ca. 3 Χ 1 mm, margin subentire, apex long acuminate. Sterile fronds stramineous, 30–35 cm, sparsely scaly; lamina pinnate; rachis usually with a narrow wing; pinnae 11–13 pairs, sessile, oblong-lanceolate, ca. 5 Χ 1–1.2 cm, abaxially sparsely scaly, adaxially glabrous, base asymmetrical, usually oblique and broadly cuneate, margin ± entire, apex acute; superior pair of pinnae connate, terminal part with a bulbil; veins reticulate and forming a costal areole. Fertile fronds not seen.

* In ravines in semi-original [??old growth] dwarf forests, on outermost mountain slopes, on seaside exposed to wind. Taiwan (Pingtung).

Bolbitis Χnanjenensis is very close to B. Χlaxireticulata, both having on their rachis narrow wings, but the latter have larger pinnae 4.5–7 Χ1.5–2 cm, margin ± entire to lobed to 2/5 toward costa, with a distinct spine in each sinus.


1 South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China.

2 Museum of Nature and Human Activities, Hyogo, 669-1546 Japan.