全 文 ：植 物 分 类 学 报 43(2): 97–115(2005)
Acta Phytotaxonomica Sinica
———————————
Received: 8 December 2003 Accepted: 15 September 2004
Supported by the grant for surveying the biodiversity in South China from the Kadoorie Farm and Botanic Garden, Hong Kong
Special Administrative Region, China, and the Special Project for Taxonomic and Floristic Researches from the Chinese
Academy of Sciences.
* Author for correspondence. E-mail: .
A taxonomic revision of the fern genus Bolbitis
(Bolbitidaceae) from China
1, 2DONG Shi-Yong 2ZHANG Xian-Chun*
1 (South China Botanical Garden, the Chinese Academy of Sciences, Guangzhou 510650, China)
2 (Laboratory of Systematic and Evolutionary Botany, Institute of Botany, the Chinese Academy of Sciences, Beijing 100093, China)
Abstract A taxonomic revision of the Chinese species of Bolbitis based on studies of
herbarium specimens and SEM observation of spores is presented in this paper. Twenty
species and three hybrids are recognized in China. Among these, B. costata (C. Presl) Ching
and B. hookeriana K. Iwats. are new records to China, B. fengiana (Ching) S. Y. Dong and B.
medogensis (Ching & S. K. Wu) S. Y. Dong are new combinations. Eight new synonyms are
proposed: B. latipinna Ching, B. media Ching & Chu H. Wang, B. yunnanensis Ching,
Egenolfia crassifolia Ching, E. crenata Ching & P. S. Chiu, E. fengiana Ching, E. medogensis
Ching & S. K. Wu and E. ×yunnanensis Ching & P. S. Chiu. Based on the perispore features,
the spores of Chinese Bolbitis can be clearly divided into three types: type A with reticulate
perispore, type B with cristate-undulate perispore and type C with smooth, undulate perispore.
The perispore feature, as well as venation pattern and the pattern of apical part of a lamina, are
the most important and useful characters for the delimitation of species in Bolbitis.
Key words Bolbitis, Bolbitidaceae, taxonomic revision, spore morphology, China.
The genus Bolbitis and the genus Egenolfia were established by Schott in 1834 (cf.
Christensen, 1906). Iwatsuki (1959) merged Egenolfia into Bolbitis considering that the two
genera cannot be satisfactorily defined. Hennipman (1977) accepted Iwatsuki’s generic
delimitation. He recorded seven species, one hybrid and a dubious species with distribution in
China in his comprehensive worldwide monograph. Ching (1931) monographed the genus
Egenolfia and recognized nine species and one variety. Two of the nine species are present in
China. Ching and Wang (1983) described nine new species of Bolbitis and five new species of
Egenolfia from Hainan, Xizang and Yunnan; Kuo (1990) reported a new hybrid, B.
×nanjenensis from Taiwan, and a species, B. scalpturata (Fée) Ching, as a new record to
Taiwan; Chu and Zhou (1994) reported B. deltigera (Bedd.) C. Chr. as a new record to China.
In the treatment of the Flora Reipublicae Popularis Sinicae 6 (1), 13 species of Bolbitis and 10
species of Egenolfia were included (Wang, 1999), but B. ×nanjenensis, B. scalpturata and
B. deltigera were not mentioned.
Early in 1931, both Christensen and Ching recognized the close affinity of Egenolfia to
Bolbitis (previously as Campium C. Presl). Christensen (1931) suggested placing the species
of both in a single genus and dividing them into two sections: Egenolfia with free veins and
Bolbitis with anastomosing veins. Ching’s work in 1931 came to the same conclusion that the
two genera might well be united, but he still maintained Egenolfia and Bolbitis as two genera
for practical purposes, in view of that Egenolfia is exclusively a tropical Asiatic genus and it
distinguishes itself from Bolbitis by free venation, dentate or erose scales and caespitose
Acta Phytotaxonomica Sinica Vol. 43 98
leaves. When Iwatsuki (1959) observed a hybrid (Bolbitis×laxireticulata) with the inconstant
venation and elucidated that none of the characters used by Ching can discriminate Egenolfia
from Bolbitis, he merged Egenolfia into Bolbitis. In 1977, Hennipman’s elaborate work on the
genus Bolbitis shows that evidence from the habitat, morphology, karyology, gametophytes
and juvenile leaves does not support Egenolfia as a distinct genus separated from Bolbitis.
In the present paper, the generic delimitation of Bolbitis follows Iwatsuki (1959) and
Hennipman (1977). Our studies are based on both herbarium and field observations, as well as
SEM observation of the spores. Twenty-two species and three hybrids of Bolbitis are
recognized from China. Among these, B. costata (C. Presl) Ching and B. hookeriana K.
Iwats. are new records to China.
The spores of 28 samples belonging to 17 species were examined under SEM. Three
distinct types of spores in the Chinese Bolbitis can be distinguished, and the spore
morphology proved very useful in species delimitation. Hennipman (1997) recognized four
types of spores. Our observations agree with the description and division of spores given by
Hennipman, but type D spores were not found in Chinese Bolbitis.
1 Material and methods
Herbarium materials for the morphological study were from PE, PYU, IBSC, KUN and
SYS. Some taxa were studied in the field during two botanical expeditions to Hainan and one
to Yunnan in 2002 and 2003. For the SEM-photography spores were stuck on aluminum stubs
with double-sided tape and sputter-coated with gold. Spores were observed and photographed
under a Hitachi S-800 scanning electron microscope. All materials for spore SEM study were
from PE and the voucher specimens were listed in Table 1.
Table 1 Source of materials for SEM observation of spores
Species Locality and voucher Figure Spore type
Bolbitis appendiculata
(Willd.) K. Iwats.
Mt. Damaoshan (大帽山), Hong Kong (香港); K. Y. Chan 1279
(PE)
1: A A
B. appendiculata Mt. Limushan (黎母山), Hainan (海南); S. Y. Dong (董仕勇) 836
(PE)`
A
B. appendiculata Wanning (万宁), Hainan (海南); S. Y. Dong et al. (董仕勇等) 559
(PE)
Abortive

Egenolfia crenata Ching &
P. S. Chiu
(=B. appendiculata)
Jinping (金平), Yunnan (云南); Sino-USSR Yunnan Exped. (中苏
联合云南考察队) 892 (PE)
A
B. medogensis (Ching & S.
K. Wu) S. Y. Dong
Mêdog (墨脱), Xizang (西藏); Plateau Ecology Exped. (高原生态
组) 11252 (PE)
1: B, C A
B. angustipinna×sinensis
Hennipman
Jinghong (景洪), Yunnan (云南); Sino-USSR Yunnan Exped. (中
苏联合云南考察队) 5775 (PE)
Abortive
B. annamensis Tardieu & C.
Chr. (=B. heteroclita (C.
Presl) Ching)
Annam, Vietnam; Cadiere 149 (PE) B
E. bipinnatifida J. Sm.
(=B. sinensis (Baker) K.
Iwats.)
Jinghong (景洪), Yunnan (云南); C. W. Wang (王启无) 7902
(PE)
Abortive
B. fengiana (Ching) S. Y.
Dong
Malipo (麻栗坡), Yunnan (云南); K. M. Feng (冯国楣) 13971
(PE)
1: H B
B. hekouensis Ching Hekou (河口), Yunnan (云南); S. K. Wu (武素功) 4056 (PE) 1: L B
B. heteroclita (C. Presl)
Ching
Mt. Limushan (黎母山), Hainan (海南); S. Y. Dong (董仕勇) 867
(PE)
2: C B
B. heteroclita Without precise locality, Sichuan (四川); Z. Y. Zhu (祝正银) 768 (PE) B
B. rhizophylla (Kaulf.)
Hennipman
Luzon, Philippines; Anonymous s.n. (PE) 1: D, E B
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 99
Table 1 (continued)
Species Locality and Voucher Figure Spore type
B. scandens W. M. Chu Lüchun (绿春), Yunnan (云南); W. M. Chu et al. (朱维明等)
6733 (PE)
1: J, K B
B. sinensis Without precise locality, Yunnan (云南); Sino-USSR Yunnan
Exped. (中苏联合云南考察队) 7300 (PE)
1: F B
B. subcordata (Copel.)
Ching
Mt. Wuzhishan (五指山), Hainan (海南); Hainan Exped. (海南队)
1816 (PE)
2: A, B B
B. subcordata Wanning (万宁), Hainan (海南); S. Y. Dong et al. (董仕勇等) 558
(PE)
B
B. subcordata Wanning (万宁), Hainan (海南); S. Y. Dong et al. (董仕勇等) 557
(PE)
B
B. subcordata Jinxiu (金秀), Guangxi (广西); Y. J. Wang (王燕杰) 5190 (PE) Abortive
B. subcordata Rongshui (融水), Guangxi (广西); S. H. Chun (陈少卿) 15679
(PE)
B
B. tibetica Ching & S. K.
Wu
Mêdog (墨脱), Xizang (西藏); Qinghai-Xizang Exped. (青藏队)
74-4551 (PE)
1: I B
B. tonkinensis (C. Chr.) K.
Iwats.
Tonkin, Vietnam; Hanoi 3396 (PE) 1: G B
B. angustipinna (Hayata) H.
Ito
Jinghong (景洪), Yunnan (云南); Sino-USSR Yunnan Exped. (中
苏联合云南考察队) 5699 (PE)
2: I, J C
B. angustipinna Mt. Qixianling (七仙岭), Hainan (海南); S. Y. Dong (董仕勇)
920 (PE)
C
B. costata (C. Presl) Ching Yingjiang (盈江), Yunnan (云南); Yunnan Univ. West Yunnan Pl.
Exped. (云南大学滇西植物调查组) 10664 (PE)
2: K, L C
B. deltigera (Bedd.) C. Chr. Yingjiang (盈江), Yunnan (云南); Yunnan Univ. West Yunnan Pl.
Exped. (云南大学滇西植物调查组) 10901 (PE)
2: F C
B. hainanensis Ching & Chu
H. Wang
Simao (思茅), Yunnan (云南); R. C. Ching (秦仁昌) 595 (PE) 2: G, H C
B. scalpturata (Fée) Ching Mt. Qixianling (七仙岭), Hainan (海南); S. Y. Dong (董仕勇)
919 (PE)
2: D, E C
2 Spore morphology
2.1 Observation
Spores monolete, ellipsoid to spheroidal, 20-55 µm in diameter; perispores usually with
wide wings or gross ridges. According to the four spore types recognized by Hennipman
(1977), three types were observed in Chinese Bolbitis.
Type A—Perispore reticulate, with reticulate, thin and wide wings. This type was
observed in two species: B. appendiculata (Fig. 1: A) and B. medogensis (Fig. 1: B, C). In
addition, B. hookeriana K. Iwats. was reported having this type of spores (Hennipman, 1977).
Type B— Perispore cristate-undulate, with thin wings. This type is very common in
Bolbitis and was observed in the following eight species: B. rhizophylla (Fig. 1: D, E), B.
sinensis (Fig. 1: F), B. tonkinensis (Fig. 1: G), B. fengiana (Fig. 1: H), B. heteroclita (Fig. 2:
C), B. scandens (Fig. 1: J, K), B. hekouensis (Fig. 1: L), and B. tibetica (Fig. 1: I). Compared
with type A and type C, type B exhibits a wider range of variation.
Type C— Perispore undulate, without wings but with gross ridges. This type was
observed in five species: B. angustipinna (Fig 2: I, J), B. costata (Fig 2: K, L), B. deltigera
(Fig 2: F), B. hainanensis (Fig 2: G, H), and B. scalpturata (Fig 2: D, E). Among these
species, the spores of B. costata are distinct by having perispore without gross ridges.
2.2 Discussion
The spore morphology in Bolbitis is distinct and significant in the species delimitation.
There are no marked differences in spore size in Chinese Bolbitis, but the diverse and stable
perispore features provide useful evidence in the subdivision of this genus and the delimitation
Acta Phytotaxonomica Sinica Vol. 43 100

Fig. 1. SEM photographs of Bolbitis spores. A, B. appendiculata. B, C, B. medogensis. D, E, B. rhizophylla. F, B.
sinensis. G, B. tonkinensis. H, B. fengiana. I, B. tibetica. J, K, B. scandens. L, B. hekouensis.
Scale bar: 10 µm in A, B, D, F-J, L and 2 µm in C, E, K.
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 101

Fig. 2. SEM photographs of Bolbitis spores. A, B, B. subcordata. C, B. heteroclita. D, E, B. scalpturata. F, B.
deltigera. G, H, B. hainanensis. I, J, B. angustipinna. K, L, B. costata.
Scale bar: 10 µm in A, C, D, F, G, I, K and 2 µm in B, E, H, J, L.
Acta Phytotaxonomica Sinica Vol. 43 102
of species. Our preliminary spore examination shows that it is not reasonable to divide the
genus Bolbitis into Egenolfia and Bolbitis s.s. Because over half species of Egenolfia (Fig. 1,
D, E, G, H) and Bolbitis s.s. each (Fig. 1: F, I-L; Fig. 2: A-C) in China have spores with
perispore cristate-undulate (type B), although spores with reticulate perispore (type A) only
occur in Egenolfia (Fig. 1: A-C) and those with undulate perispore (type C) only occur in
Bolbitis s.s. (Fig. 2: D-L). Based on the differences of the three type spores, it appears
possible to divide Chinese Bolbitis into three sections. In the taxonomic treatment of some
dubious species, the perispore feature is very useful. Egenolfia crenata is morphologically
similar to B. appendiculata (Willd.) K. Iwats., but different by being larger in size. The spores
of these two species show no difference, all with reticulate perispore. Similarly, the perispore
feature supplies further evidence to treat B. annamensis Tardieu & C. Chr. as a synonym of B.
heteroclita (C. Presl) Ching.
It is difficult to draw a conclusion on the evolutionary trend of the three spore types
recognized herein. Though one of these spore types (type A) is easily distinguished from the
other two (type B and type C) by the perispore feature, the spore morphology has not been
found to be correlated with other gross-morphological characters in this genus. There are only
partial connections among spore types, venation pattern and the pattern of apical part of a
lamina, which are the most valuable diagnostic characters in Bolbitis. The type C spores only
occur in the species with anastomosing veins and with imparipinnate laminae; the type A
spores only occur in the species with free veins but with a pinnately divided apical part of
laminae; and the type B spores are not correlated with either the venation pattern or the
morphology of lamina apex. Therefore, the evolutionary trends of the spore types and thus the
evolutionary relationships of the species in Bolbitis cannot be revealed.
3 Systematic treatment
Bolbitis Schott, Gen. Fil. pl. 14. 1834; Ching in C. Chr., Ind. Fil. Suppl. 3: 47. 1934; Copel.,
Gen. Fil. 115. 1947; K. Iwats. in Acta Phytotax. Geobot. 18: 44. 1959; Hennipman, Monogr.
Gen. Bolbitis 123. 1977; et in Steenis & Holttum, Fl. Mal. Ser. II, 1 (4): 314. 1978; K. U.
Kramer in Kubitzki, Fam. Gen. Vas. Pl. 1: 167. 1990; C. M. Kuo in Bot. Bull. Acad. Sin. 31:
305. 1990; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 104. 1999. Type: Bolbitis serratifolia
(Kaulf.) Schott.
Egenolfia Schott, Gen. Fil. pl. 16. 1834; Ching in Bull. Fan Mem. Inst. Biol. Bot. Ser. 2:
297. 1931; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 115. 1999. Type: Egenolfia
hamiltoniana Schott (=Bolbitis appendiculata (Willd.) K. Iwats.).
A genus of about 80 species pantropical in distribution, 20 species and three hybrids
recognized from China.
Key to species of China
1. Veins all free.
2. perispore reticulate; lateral pinnae subentire or crenate, base asymmetrical (except B. hookeriana), apex
obtuse or acute.
3. Stipe and rachis subglabrous; base of pinnae asymmetrical; fertile pinnae ovate or oblong.
4. Rhizome creeping; sterile pinnae usually obtuse or rounded at apex; fertile pinnae 2-20 mm long
and 2-4 mm wide ……...……………………………………………….……… 1. B. appendiculata
4. Rhizome short-creeping or suberect; sterile pinnae acute at apex; fertile pinnae ca. 80 mm long and
8-10 mm wide. ………..….…………………………………………………….… 2. B. medogensis
3. Stipe and rachis densely scaly; base of pinnae symmetrical; fertile pinnae moniliform………………...
…………………..…………………………………………………………………… 3. B. hookeriana
2. perispore cristate-undulate; lateral pinnae pinnatifid, base symmetrical, apex acuminate (lateral pinnae of
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 103
B. rhizophylla usually margin serrate and apex rounded).
5. Bulbil terminal on the lamina; lateral pinnae subentire to serrate ……………………4. B. rhizophylla
5. Bulbil subterminal on the lamina; lateral pinnae pinnatifid.
6. Apex of lobes rounded, lobes 0-1 mm apart without spines on margin …………...…..5. B. sinensis
6. Apex of lobes obtusely acute, lobes 2-3 mm apart with fine spines on margin.
7. Stipe and rachis densely scaly, scales dark-brown, appressed; lobes of pinnae falcate, 6-15 mm
long …………………………………………………………………………….. 6. B. tonkinensis
7. Stipe and rachis subglabrous, scales brown, patent; lobes of pinnae triangular, 2-5 mm long…….
………………...…………………………………………………………………… 7. B. fengiana
1. Veins more or less anastomosing.
8. Venation pattern more or less irregular, veins along costae usually anastomosing and others free or rarely
uniting, included free veinlets never present; rachis narrowly winged (in B. angustipinna×sinensis,
rachis wingless).
9. Pinnae pinnatifid, base symmetrical; rachis wingless ……………….… 8. B. angustipinna×sinensis
9. Pinnae subentire or crenate, base more or less asymmetrical; rachis narrowly winged.
10. Pinnae subentire; more veins anastomosing ………………………………… 9. B. ×nanjenensis
10. Pinnae crenate; except those along costae, other veins all free ……………10. B. ×laxireticulata
8. Venation pattern regular, veins variously anastomosing, with or without included free veinlets; rachis
wingless.
11. 5-8 veinlets arising from either side of lateral veins uniting in sterile pinnae, veinlets near margin
more or less uniting; lamina imparipinnate; leaves often dark-brownish when dry.
12. Fronds in 2 series on rhizome; scales on base of stipe narrowly lanceolate, without obvious
brown margin.
13. Leaves herbaceous; rhizome 3-5 mm thick, terrestrial or climbing on bases of tree-trunks…..
…………………………………………………………………………...… 11. B. heteroclita
13. Leaves chartaceous; rhizome 8-10 mm thick, high-climbing ……………….12. B. scandens
12. Fronds in 4 series on dorsal surface of rhizome; scales on base of stipe ovate-lanceolate, with
obvious brown margin ………………………………………………………13. B. confertifolia
11. 1-4 veinlets arising from either side of lateral veins uniting in sterile pinnae, veinlets near margin all
free (excluding those in B. costata); lamina with a pinnate apical part or not; leaves greenish or
rarely purplish when dry.
14. Lamina with a pinnate apical part; lateral pinnae (4)7-15 pairs; base of sterile pinnae rounded,
truncate or subcordate; perispore cristate-undulate.
15. Stipe and rachis hardly scaly; venation pattern with included free veinlets or not.
16. Included free veinlets present; base of sterile pinnae rounded……...…14. B. subcordata
16. Included free veinlets absent; base of sterile pinnae subcordate……..15. B. christensenii
15. Stipe and rachis densely scaly throughout; venation pattern without included free veinlets.
17. Bulbil subterminal; sterile pinnae rounded at base ……………………16. B. hekouensis
17. Bulbil terminal; sterile pinnae truncate at base ……………………………17. B. tibetica
14. Lamina imparipinnate; lateral pinnae 2-7 pairs (except B. angustipinna with 8-24 pairs); base
of sterile pinnae cuneate to rounded; perispore smoothly undulate.
18. Fertile pinnae lanceolate, 3-8 times longer than wide.
19. Sterile pinnae 2-3 pairs, 4.5-6 cm wide; fertile pinnae acrostichoid.....18. B. hainanensis
19. Sterile pinnae 5-7 pairs, 2-4 (5) cm wide; either side of costae or lateral veins with a
narrow region not bearing sporangia in fertile pinnae.
20. 2-3 veinlets arising from lateral veins, most areoles without included free veinlets
and few areoles usually with one, free veinlets without thickened ends; costae
usually purplish or stramineous when dry …………………….….19. B. scalpturata
20. 4-5 veinlets arising from lateral veins, most areoles with 1-3 included free veinlets,
free veinlets with thickened ends; costae stramineous when dry…….20. B. deltigera
18. Fertile pinnae linear, 14-50 times longer than wide.
21. Venation pattern without included free veinlets; lateral pinnae (8-)15-24 pairs…………
…………………...………………………..………………………... 21. B. angustipinna
21. Venation pattern with included free veinlets; lateral pinnae 2-6 pairs.
Acta Phytotaxonomica Sinica Vol. 43 104
22. Leaves purplish when dry; 5-8 veinlets arising from either side of lateral veins
uniting in sterile pinnae; fertile pinnae 7-8 cm long ………...…….….22. B. costata
22. Leaves greenish when dry; 4 veinlets arising from either side of lateral veins uniting
in sterile pinnae; fertile pinnae ca. 20 cm long …………… ...…………23. B. virens

1. Bolbitis appendiculata (Willd.) K. Iwats. in Acta Phytotax. Geobot. 18: 48. 1959; Pic.
Serm., Ind. Fil. Suppl. 4: 140. 1965; Hennipman, Monogr. Gen. Bolbitis 185, figs. 49-51.
1977, p.p.; et in Steenis & Holttum, Fl. Mal. Ser. II, 1 (4): 322, fig. 29. 1978; Tagawa & K.
Iwats., Fl. Thail. 3 (3): 316. 1988.——Acrostichum appendiculatum Willd., Sp. Pl. 5: 114.
1810. ——Egenolfia appendiculata (Willd.) J. Sm., Ferns Br. For. 111. 1866; Ching in Bull.
Fan Mem. Inst. Biol. Bot. 2: 308. 1931; Tardieu & C. Chr. in Lecomte, Fl. Gén. Indo-Chiné 7:
426. 1941; J. L. Tsai & W. C. Shieh in T. C. Huang et al., Fl. Taiwan., ed. 2, 1: 356, pl. 142.
1994; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 117, pl. 22: 1-6. 1999. Type: India. Without
precise locality, Klein 912 (holotype, B).
Egenolfia crenata Ching & P. S. Chiu in Acta Phytotax. Sin. 21: 212. 1983; Chu H. Wang
in Fl. Reip. Pop. Sin. 6 (1): 118. 1999, syn. nov. Type: China. Yunnan (云南): Jinping (金平),
Sino-USSR Yunnan Exped. (中苏联合云南考察队) 892 (holotype, PE!; isotype, KUN!).
刺蕨 Fig. 1: A
Representative specimens examined:
China. Guangdong (广东): Huaiji (怀集), W. T. Tsang (曾怀德) 23204 (IBSC, SYS),
Y. G. Liu (刘英光) 02798 (PE, IBSC); Maoming (茂名), L. Deng (邓良) 1787 (PE, IBSC,
KUN), L. Deng (邓良) 2337 (IBSC, KUN); Shenzhen (深圳), Shenzhen Exped. (深圳队)
743, 783 (PE); Wengyuan (翁源), S. K. Lau (刘心祈) 2389 (PE, SYS); Xinyi (信宜), C.
Wang (黄志) 31155 (PE, IBSC, KUN), C. Wang (黄志) 31973 (PE, SYS), C. Wang (黄志)
37833 (IBSC); Yangchun (阳春), H. G. Ye et al. (叶华谷等) 121 (IBSC); Yu’nan (郁南), N.
Liu et al. (刘念等) 2728, 2735 (IBSC). Guangxi (广西): Fangcheng (防城), W. T. Tsang (曾
怀德) 26823 (IBSC, SYS); Luocheng (罗城), W. M. Chu et al. (朱维明等) 18378 (PYU);
Nanning (南宁), R. C. Ching (秦仁昌) 8238 (PE, SYS); Pingnan (平南), C. Wang (黄志)
4071 (PE); Rongshui (融水), S. H. Chun (陈少卿) 14129 (PE, IBSC, KUN); Rongxian (荣
县), S. H. Chun (陈少卿) 9819 (IBSC); Yaoshan (瑶山), K. K. Whong 140 (PE), S. S. Sin (辛
树帜) 3863 (PE, SYS). Hainan (海南): Baisha (白沙), W. M. Chu et al. (朱维明等) 18133
(PYU); Baoting (保亭), S. K. Lau (刘心祈) 28113 (PE, IBSC, KUN); Changjiang (昌江), S.
Y. Dong et al. (董仕勇等) 56, 110 (PE); Danzhou (儋州), W. T. Tsang (曾怀德) 756 (SYS);
Lingshui (陵水), W. M. Chu (朱维明) 5913 (PYU), Hainan Exped. (海南队) 1873 (PYU),
Diaoluoshan Exped. (吊罗山队) 2309, 3309 (PE, IBSC); Mt. Jianfengling (尖峰岭), S. Y.
Dong et al. (董仕勇等) 262 (PE), Z. X. Li et al. (李泽贤等) 1240 (IBSC), Hainan Exped. (海
南队) 1449 (PYU); Mt. Limushan (黎母山), S. Y. Dong (董仕勇) 836 (PE), S. H. Chun (陈
少卿) 10667 (PE, IBSC); Mt. Wuzhishan (五指山), E. Hainan Exped. (海南东队) 552 (PE,
IBSC), C. Wang (黄志) 35443 (PE), 35625 (IBSC), N. K. Chun (陈念劬) 44026 (PE, IBSC,
KUN), F. A. McClure 1978, 1984 (SYS), F. W. Xing et al. (邢福武等) 5406 (IBSC);
Qiongzhong (琼中), L. Deng (邓良) 3424 (IBSC); Sanya (三亚), C. Wang (黄志) 34603 (PE,
IBSC), 34299B (IBSC), H. Y. Liang (梁向日) 62714, 63186 (PE, IBSC); Wanning (万宁), L.
Deng (邓良) 2929 (IBSC), S. Y. Dong et al. (董仕勇等) 559 (PE); Without precise locality,
917 Group (917组) 042 (SYS), P. Zeng (曾沛) 12514 (SYS). Hong Kong (香港): Mt.
Damaoshan (大帽山), K. Y. Chan 1279 (PE). Taiwan (台湾): Xinzhu (新竹), D. E. Boufford
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 105
et al. 25213 (PE, KUN); Miaoli (苗栗), S. J. Mou (牟善杰) 17722 (PYU). Yunnan (云南):
Cangyuan (沧源), W. M. Chu et al. (朱维明等) 15417, 17417 (PYU); Hekou (河口), W. M.
Chu (朱维明) 5761 (PYU), Dept. Biol. Yunnan Univ. Exer. Exped. (云南大学生物系实习
队) 1699, 1700 (PYU), K. H. Cai (蔡克华) 215 (PE), Z. X. Zhou (周政贤) 42 (PE); Jinping
(金平), W. M. Chu (朱维明) 4051 (PYU), W. M. Chu et al. (朱维明等) 6525 (PYU), S. K.
Wu (武素功) 3997 (PE, PYU); Lüchun (绿春), W. M. Chu et al. (朱维明等) 6643 (PYU);
Mengla (勐腊), W. M. Chu et al. (朱维明等) 18664 (PYU); Pingbian (屏边), W. M. Chu (朱
维明) 51 (PYU); Yingjiang (盈江), Yunnan Univ. West Yunnan Pl. Exped. (云南大学滇西
植物调查组) 9918, 10767, 10871 (PYU).
Habitat: On moist rocks in rain forests, alt. 100-1200 m.
Distribution: A widespread species in tropical Asia, southward to Java and northward to
southern China, eastward to Philippines and westward to Sri Lanka.
Note: Egenolfia crenata was described as new for sterile leaves larger, fertile pinnae
more shortened and the apex of sterile pinnae sharply acute by comparison with B.
appendiculata (Ching & Wang, 1983). Our examination has shown that the type collection of
E. crenata is in the normal variation range of B. appendiculata in the size of sterile leaves,
and that there is no difference between these two species in the morphology of fertile pinnae
and in the spore morphology.
2. Bolbitis medogensis (Ching & S. K. Wu) S. Y. Dong, com. nov. ——Egenolfia
medogensis Ching & S. K. Wu in C. Y. Wu, Fl. Xizang. 1: 278. 1983; Chu H. Wang in Fl.
Reip. Pop. Sin. 6 (1): 116. 1999, syn. nov. Type: China. Xizang (西藏): Mêdog (墨脱),
Qinghai-Xizang Exped. (青藏队) 74-4335 (holotype, PE!; isotype, KUN!).
墨脱刺蕨 Fig. 1: B, C
Representative specimen examined:
China. Xizang (西藏): Mêdog (墨脱), Plateau Ecology Exped. (高原生态组) 11252 (PE).
Habitat: On slope in evergreen broad-leaved forests, alt. 900 m.
Distribution: Only known from the type locality.
Note: This species is close to B. appendiculata and may be an ecological form of the
latter. Considering the much larger size in leaves and the restricted distribution, it is
maintained as a separate species.
3. Bolbitis hookeriana K. Iwats. in Acta Phytotax. Geobot. 18: 49. 1959. —— Polybotrya
vivipara Buch.-Ham. ex Hook., Exot. Fl. 2: t. 107. 1825, non Bolbitis vivipara C. Chr., Ind.
Fil. Suppl. 3: 51. 1934. —— Egenolfia vivipara (Hook.) C. Chr., Ind. Fil. Suppl. 3: 102. 1934;
Tardieu & C. Chr. in Lecomte, Fl. Gén. Indo-Chiné 7: 423. 1941. —— Bolbitis appendiculata
ssp. vivipara var. vivipara (Hook.) Hennipman in Blumea 18: 147. 1970 et Monogr. Gen.
Bolbitis 196, fig. 50: a-h, fig. 51. 1977. Type: India. Assam: Goalpara, Wallich 29 p.p.
(holotype, K).
Representative specimen examined:
China. Yunnan (云南): Yingjiang (盈江), Q. Z. Huang (黄全忠) 783 (PE), new record.
Habitat: Creeping on rocks in forests, alt. 500 m.
Distribution: China (W Yunnan), Bangladesh, India, Myanmar, Indo-China Peninsula.
Note: This species is a member with type A spore but distinct in Chinese Bolbitis by the
moniliform fertile pinnae.
虎克实蕨
4. Bolbitis rhizophylla (Kaulf.) Hennipman in Blumea 18: 148. 1970 et Monogr. Gen.
Bolbitis 199, fig. 52: d-r, fig. 53. 1977; et in Steenis & Holttum, Fl. Mal. Ser. II, 1 (4): 32, fig.
Acta Phytotaxonomica Sinica Vol. 43 106
27g. 1978. ——Gymnogramma rhizophylla Kaulf., Enum. Fil. 78. 1824. ——Egenolfia
rhizophylla (Kaulf.) Fée, Gen. Fil. 48. 1852; J. L. Tsai & W. C. Shieh in T. C. Huang et al.,
Fl. Taiwan., ed. 2, 1: 356. 1994; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 118. 1999. Type:
Philippines. Manila: Chamisso s.n. (holotype, LE?; isotype, B).
根叶刺蕨 Fig. 1: D, E
Representative specimen examined:
China. Taiwan (台湾): Tainan (台南), T. Murakami et al. 236 (KUN).
Habitat: On rocks in forests, alt. 400-500 m.
Distribution: China (Taiwan), Philippines.
Note: This species is the only one with terminal bulbil on fronds among the species
which have a free venation pattern.
5. Bolbitis sinensis (Baker) K. Iwats. in Acta Phytotax. Geobot. 18: 49. 1959; Hennipman,
Monogr. Gen. Bolbitis 202, figs. 53, 54. 1977; et in Steenis & Holttum, Fl. Mal. Ser. II, 1 (4):
325, fig. 27h. 1978; Tagawa & K. Iwats., Fl. Thail. 3 (3): 318. 1988 ——Acrostichum sinense
Baker in Kew Bull. 1906: 14. 1906. ——Egenolfia sinensis (Baker) Maxon in Proc. Biol.
Soc. Wash. 36: 173. 1923; Tardieu & C. Chr. in Lecomte, Fl. Gén. Indo-Chiné 7: 424, fig. 48:
1, 2. 1941; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 120, pl. 22: 9-11. 1999. Type: China.
Yunnan (云南): Simao (思茅), Henry 12494 (holotype, K).
Egenolfia bipinnatifida J. Sm., Hist. Fil. 132. 1875; Chu H. Wang in Fl. Reip. Pop. Sin. 6
(1): 122. 1999. —— Bolbitis bipinnatifida (J. Sm.) K. Iwats. in Acta Phytotax. Geobot. 18:
49. 1959 (sphalm. bipinnata), non B. bipinnatifida (Mett.) Ching in C. Chr., Ind. Suppl. III.
1934. Type: Myanmar. Tenasserim: Dawna Range near Moulmein, parish 60 (holotype, K).
Egenolfia crassifolia Ching in Acta Phytotax. Sin. 21: 216. 1983; Chu H. Wang in Fl.
Reip. Pop. Sin. 6 (1): 120. 1999, syn. nov. Type: China. Yunnan (云南): Lushui (泸水),
Nanshuibeidiao Exped. (南水北调队) 8117 (holotype, PE!; isotypes, PYU!, KUN!).
中华刺蕨 Fig. 1: F
Representative specimens examined:
China. Guangxi (广西): Tianlin (田林), C. C. Chang (张肇骞) 10974 (IBSC). Yunnan
(云南): Cangyuan (沧源), W. M. Chu et al. (朱维明等) 15304, 15371 (PYU); Gengma (耿
马), W. M. Chu et al. (朱维明等) 15260 (PYU); Jingdong (景东), B. Y. Qiu (邱炳云) 52677
(KUN); Jinghong (景洪), W. M. Chu (朱维明) 3932 (PYU), W. M. Chu et al. (朱维明等)
479, 500 (PYU), Y. M. Feng (冯永明) 11 (PYU), J. M. Zeng & Q. Yang (曾觉明,杨泉) 140,
782 (PYU), Sino-USSR Yunnan Exped. (中苏联合云南考察队) 56097 (KUN), G. D. Tao et
al. (陶国达等) 43514, 43521 (PYU), Anonymous 56837 (KUN), C. W. Wang (王启无)
78219, 79027 (PE), 78750 (PE, KUN); Lüchun (绿春), S. K. Wu et al. (武素功) 821 (KUN);
Menghai (勐海), W. M. Chu et al. (朱维明等) 15764, 24592 (PYU), Sino-USSR Yunnan
Exped. (中苏联合云南考察队) 5605, 7121, 7300 (KUN); Mengla (勐腊), W. M. Chu (朱维
明) 2062 (PYU), X. W. Li (李锡文) 59-13479 (KUN), Sino-Japanese Exped. (中日队) 114
(KUN), H. T. Tsai (蔡希陶) 59-10985 (KUN); Simao (思茅), R. C. Ching (秦仁昌) 415, 524,
527 (PE), Sino-Japanese Exped. (中日队) 229 (KUN), Sino-USSR Yunnan Exped. (中苏联
合云南考察队) 56056 (KUN); Without precise locality, Sino-USSR Yunnan Exped. (中苏联
合云南考察队) 7120, 7300 (PE); Xinping (新平), W. M. Chu (朱维明) 357 (PE, PYU);
Yangbi (漾濞), W. M. Chu et al. (朱维明等) 9473 (PYU); Yingjiang (盈江), Q. Z. Huang (黄
全忠) s.n. (PE), Yunnan Univ. West Yunnan Pl. Exped. (云南大学滇西植物调查组) 10695,
10708, 10740, 10761, 10869, 10899 (PYU); Yongde (永德), W. M. Chu et al. (朱维明等)
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 107
15078 (PYU); Yun Xian (云县), W. M. Chu et al. (朱维明等) 14828 (PYU).
Habitat: In soil or on rocks in forests, alt. 650-1900 m.
Distribution: China, Bangladesh, Myanmar, Thailand, Cambodia, Vietnam, Java, Lesser
Sunda Islands.
Note: Egenolfia crassifolia was described based on a sheet of sterile specimen
(Nanshuibeidiao Exped. 8117, PE) and cannot be distinguished from B. sinensis. An isotype
(PYU) of E. crassifolia was identified as E. sinensis by Professor W. M. Chu of Yunnan
University in 1999.
6. Bolbitis tonkinensis (C. Chr.) K. Iwats. in Acta Phytotax. Geobot. 18: 49. 1959;
Hennipman, Monogr. Gen. Bolbitis 310. 1977; Tagawa & K. Iwats., Fl. Thail. 3 (3): 319.
1988. ——Egenolfia tonkinensis C. Chr. in Bull. Fan Mem. Inst. Biol. Bot. 2: 306. 1931;
Tardieu & C. Chr. in Lecomte, Fl. Gén. Indo-Chiné 7: 424. 1941; Chu H. Wang in Fl. Reip.
Pop. Sin. 6 (1): 119. 1999. Type: Vietnam. Tonkin: Lang-Son, Herb. Ecole Sup. Agric. &
Sylvic. Hanoi 3396 (holotype, BM; isotype, PE!).
镰裂刺蕨 Fig. 1: G
Representative specimens examined:
China. Yunnan (云南): Jinghong (景洪), W. M. Chu et al. (朱维明等) 24575, 24640
(PYU); Mengla (勐腊), W. M. Chu (朱维明) 5988 (PYU), B. G. Li (李保贵) 00690, 00756
(PYU); Simao (思茅), Z. H. Hu (胡志浩) s.n. (PYU).
Habitat: On moist rocks in forests, alt. 500-1260 m.
Distribution: China, Vietnam (Tonkin).
Note: This species resembles both B. fengiana and B. sinensis, and is distinguished from
the latter two by its densely scaly stipe and rachis.
7. Bolbitis fengiana (Ching) S. Y. Dong, com. nov. ——Egenolfia fengiana Ching in Acta
Phytotax. Sin. 21: 215. 1983; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 119, pl. 22: 7, 8.
1999, syn. nov. Type: China. Yunnan (云南): Malipo (麻栗坡), K. M. Feng (冯国楣) 13758
(holotype, PE!).
疏裂刺蕨 Fig. 1: H
Representative specimens examined:
China. Yunnan (云南): Jinping (金平), W. M. Chu et al. (朱维明等) 6513 (PYU);
Lüchun (绿春), W. M. Chu et al. (朱维明等) 6645 (PYU); Malipo (麻栗坡), K. M. Feng (冯
国楣) 13971 (PE, KUN), 22987 (KUN); Pingbian (屏边), Sino-USSR Yunnan Exped. (中苏
联合云南考察队) 3551 (PE), W. M. Chu et al. (朱维明等) 29221 (PYU), Y. Jiao (焦瑜)
96-49 (PYU).
Habitat: In valley under forests, alt. 550-1700 m.
Distribution: Endemic to China (Yunnan).
8. Bolbitis angustipinna×sinensis Hennipman, Monogr. Gen. Bolbitis 284, fig. 83: a-c.
1977. Type: China. Yunnan (云南): Xishuangbanna (西双版纳), between Jinghong (景洪)
and Mengxing (勐醒), Rock 2636 (holotype ?; isotypes, BM, C, US).
Egenolfia×yunnanensis Ching & P. S. Chiu in Acta Phytotax. Sin. 21: 216. 1983; Chu
H. Wang in Fl. Reip. Pop. Sin. 6 (1): 123. 1999, syn. nov. Type: China. Yunnan (云南):
Jinghong (景洪), Sino-USSR Yunnan Exped. (中苏联合云南考察队) 5775 (holotype, PE!;
isotype, KUN!).
Campium sinense auct. non (Baker) C. Chr.: C. Chr., Contr. U.S. Nat. Herb. 26: 292,
1931, p.p.
云南实蕨
Acta Phytotaxonomica Sinica Vol. 43 108
Representative specimens examined:
China. Yunnan (云南): Jinghong (景洪), W. M. Chu et al. (朱维明等) 480 (PYU);
Menghai (勐海), W. M. Chu et al. (朱维明等) 6856 (PYU).
Habitat: On ground or at base of tree-trunks in valleys under rain forests, alt. 850-1050 m.
Distribution: Endemic to China (southern Yunnan).
Note: The venation pattern in this species is irregular. One to three veinlets arising from
lateral veins can be observed anastomosing.
9. Bolbitis×nanjenensis C. M. Kuo in Bot. Bull. Acad. Sin. 31: 308, fig. 1. 1990; R. J.
Johns, Ind. Fil. Suppl. 6: 63. 1996. Type: China. Taiwan (台湾): Pingdong (屏东), Kuo & Yu
14856 (holotype, TAI).
南仁实蕨
Habitat: In a ravine in semi-original dwarf forests.
Distribution: Only known from the type locality.
Note: This hybrid is very close to B. ×laxireticulata, but their putative parents for the
two hybrids are different. The putative parents of B. ×nanjenensis are B. appendiculata and
B. heteroclita (Kuo, 1990), and those of B. ×laxireticulata are B. appendiculata and B.
subcordata (see the following note).
10. Bolbitis×laxireticulata K. Iwats. in Acta Phytotax. Geobot. 18: 50, figs. 7, 8. 1959;
Hennipman, Monogr. Gen. Bolbitis 307, fig. 86: i-q. 1977; C. M. Kuo in Bot. Bull. Acad. Sin.
31: 308. 1990. ——Egenolfia laxireticulata (K. Iwats.) C. M. Kuo in H. L. Li et al., Fl.
Taiwan. 1: 352. 1975; J. L. Tsai & W. C. Shieh in T. C. Huang et al., Fl. Taiwan., ed. 2, 1: 356.
1994. Type: Japan. Ryukyu: Isl. Amami-Ooshima, Tagawa & Iwatsuki 2918 (holotype, KYO).
网脉实蕨
Representative specimens examined:
China. Hainan (海南): Mt. Limushan (黎母山), S. Y. Dong (董仕勇) 858 (PE). Hong
Kong (香港): Without precise locality, Anonymous s.n. (IBSC, herb. no. 623194).
Habitat: On rocks along stream in secondary rain forests, alt. 650 m.
Distribution: China (Hainan, Hong Kong and Taiwan), Japan (Ryukyu Isl.).
Note: This hybrid was originally reported from Ryukyu and was suggested hybridity
between B. appendiculata and B. ×laxireticulata (Iwatsuki, 1959). The present fern was
recently collected from Mt. Limushan of the Hainan Island. It grows on rocks by a stream in
secondary rain forests. The outline of this fern is very similar to that of B. appendiculata.
which grows side by side with B. ×laxireticulata, but much larger than the latter. The plant
size, the number and shape of lateral pinnae, the venation pattern, and the number of scales on
stipe and rachis are intermediate between B. appendiculata and B. subcordata, two species
which are very common on Mt. Limushan and throughout the Hainan Island. In addition, B.
appendiculata and B. subcordata also grow in Hong Kong, Taiwan and Ryukyu where B.
×laxireticulata occurs. So the present fern may have originated from B. appendiculata and B.
subcordata by hybridization.
11. Bolbitis heteroclita (C. Presl) Ching in C. Chr., Ind. Fil. Suppl. 3: 48. 1934; Tardieu & C.
Chr. in Lecomte, Fl. Gén. Indo-Chiné 7: 434. 1941; K. Iwats. in Acta Phytotax. Geobot. 18:
57, fig. 12. 1959; Hennipman, Monogr. Gen. Bolbitis 221, fig. 60. 1977; et in Steenis &
Holttum, Fl. Mal. Ser. II, 1 (4): 325, figs. 25d, 31a-g. 1978; Tagawa & K. Iwats., Fl. Thail. 3
(3): 320. 1988; J. L. Tsai & W. C. Shieh in T. C. Huang et al., Fl. Taiwan., ed. 2, 1: 353, pl.
141. 1994; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 108, pl. 19: 1-4. 1999. ——Acrostichum
heteroclitum C. Presl, Rel. Haenk. 1: 15, pl. 2, fig. 2. 1825. Type: Philippines. Luzon:
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 109
Sorsogon, Haenke s.n. (holotype, PRC).
Bolbitis annamensis Tardieu & C. Chr. in Not. Syst. 7: 100. 1938; et in Lecomte, Fl.
Gén. Indo-Chiné 7: 436, fig. 50: 3, 4. 1941; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 106, pl.
19: 5. 1999. Type: Vietnam. Annam: Thanh Tan, Cadiere 149 (holotype, BM; isotype, PE!).
长叶实蕨 Fig. 2: C
Representative specimens examined:
China. Chongqing (重庆): Beibei (北碚), X. Zhou (周锌) 1202 (PE); Nanchuan (南川),
Z. Y. Liu et al. (刘正宇等) 4580 (PYU); Without precise locality, B. Y. Zhang et al. (张百誉
等) 0085 (PE), C. Z. Liu et al. (刘承泽等) 100121(PE). Guangdong (广东): Dongxing (东
兴), K. K. Tsoong (钟观光) s.n. (PE); Fengkai (封开), S. Wang (黄成) 164033 (PE).
Guangxi (广西): Fusui (扶绥), S. H. Chun (陈少卿) 12090 (PE, IBSC, KUN); Longsheng
(龙胜), P. S. Qiu (裘佩熹) 4580 (PE); Nanning (南宁), H. Li et al. (黎桦等) 504 (PYU);
Napo (那坡), S. P. Ko (高锡朋) 56007 (PE); Rongshui (融水), S. H. Chun (陈少卿) 15626
(PE, IBSC, KUN); Wuming (武鸣), Anonymous 138 (PE); Without precise locality, Guangxi
Exped. (广西队) 3477 (PE). Guizhou (贵州): Anlong (安龙), P. S. Wang (王培善) 76279
(PYU); Chishui (赤水), Z. Y. Cao et al. (曹子余等) 240, 285 (PE), P. S. Wang (王培善)
77787 (PYU); Dushan (独山), X. Y. Hou (侯学煜) 1904 (PE); Leishan (雷山), P. S. Wang
(王培善) 76970 (PE, PYU); Liping (黎平), F. Wang et al. (王峰等) 91609 (PYU). Hainan
(海南): Baisha (白沙), W. M. Chu et al. (朱维明等) 18107 (PYU); Danzhou (儋州), W. T.
Tsang (曾怀德) 382 (PE, SYS); Ledong (乐东), S. Y. Dong et al. (董仕勇等) 303 (PE); Mt.
Limushan (黎母山), S. Y. Dong (董仕勇) 837, 867 (PE). Sichuan (四川): Mt. Emeishan (峨
眉山), W. M. Chu (朱维明) 7205, 7843 (PYU), K. H. Shing et al. (邢公侠等) 1382, 1720,
1739 (PE); Qianwei (犍为), K. H. Shing et al. (邢公侠等) 0524-B (PE); Without precise
locality, Z. Y. Zhu (祝正银) 768 (PE). Yunnan (云南): Cangyuan (沧源), W. M. Chu et al.
(朱维明等) 15302, 15303, 15432 (PYU); Gengma (耿马), Y. H. Li (李延辉) 002276 (KUN);
Guangnan (广南), W. M. Chu et al. (朱维明等) 8347 (PYU); Hekou (河口), Dept. Biol.
Yunnan Univ. Exer. Exped. (云南大学生物系实习队) 704 (PE, PYU), 795, 1723, 1734
(PYU), K. H. Cai (蔡克华) 993 (PE); Jinghong (景洪), W. M. Chu (朱维明) 2086, 2102
(PYU), W. M. Chu et al. (朱维明等) 450, 24608 (PYU), H. Q. Guo (郭汉卿) 56 (PE),
Sino-USSR Yunnan Exped. (中苏联合云南考察队) 5701, 9612 (PE, KUN), 9759 (KUN), C.
W. Wang (王启无) 77910, 78094 (PE, IBSC, KUN), Yunnan First Group Exped. (云南一组)
92 (PE), G. F. Zhang (张光飞) 24669 (PYU), Anonymous 803 (PYU), 55859, 56779 (KUN);
Jinping (金平), Sino-USSR Yunnan Exped. (中苏联合云南考察队) 305, 886, 1714 (PE);
Lancang (澜沧), W. M. Chu et al. (朱维明等) 15547 (PYU); Luoping (罗平), W. M. Chu et
al. (朱维明等) 13254 (PYU); Lüchun (绿春), W. M. Chu et al. (朱维明等) 6732 (PYU);
Maguan (马关), W. M. Chu et al. (朱维明等) 23486, 29267 (PYU); Menghai (勐海), W. M.
Chu et al. (朱维明等) 6838 (PYU), C. W. Wang (王启无) 74857 (PE), 77117 (PE, KUN),
Sino-USSR Yunnan Exped. (中苏联合云南考察队) 5481, 5656 (PE, KUN); Mengla (勐腊),
W. M. Chu (朱维明) 2051, 2083 (PYU), J. W. Li et al. (李建伟等) 17854 (PYU),
Sino-Japanese Exped. (中日队) 248, 339 (KUN), C. W. Wang (王启无) 80051 (PE, KUN);
Mengzi (蒙自), T. N. Liou (刘慎锷) 018720 (PE); Pingbian (屏边), H. T. Tsai (蔡希陶)
60522 (PE, KUN); Shuangjiang (双江), W. M. Chu et al. (朱维明等) 15529 (PYU); Suijiang
(绥江), W. M. Chu (朱维明) 4855 (PYU); Xichou (西畴), W. M. Chu et al. (朱维明等)
21820 (PYU); Ximeng (西盟), W. M. Chu et al. (朱维明等) 15649 (PYU); Yingjiang (盈江),
Acta Phytotaxonomica Sinica Vol. 43 110
Q. Z. Huang (黄全忠) 780, 784, 789 (PE), Yunnan Univ. West Yunnan Pl. Exped. (云南大学
滇西植物调查组) 10665, 10750 (PYU).
Habitat: Usually on rocks or at base of trees near streams in forests, alt. 100-1250 m.
Distribution: Widespread in Asian tropics.
Note: This species has (0)1-3(5) pair(s) of lateral pinnae. The plant with simple leaf
(without lateral pinnae) was formerly referred to B. annamensis and reported only in central
Vietnam and Guangxi (Wang, 1999). Recently, a population of B. heteroclita without lateral
pinnae was also observed in Mt. Limushan of Hainan.
12. Bolbitis scandens W. M. Chu in Acta Phytotax. Sin. 21: 213. 1983; Chu H. Wang in Fl.
Reip. Pop. Sin. 6 (1): 113. 1999. Type: China. Yunnan (云南): Lüchun (绿春), W. M. Chu et
al. (朱维明等) 6733 (holotype, PYU!; isotypes, PE!, PYU!).
附着实蕨 Fig. 1: J, K
Representative specimen examined:
China. Yunnan (云南): Mengla (勐腊), W. M. Chu (朱维明) 2050 (PYU).
Habitat: Climbing on tree-trunks in valley under forests, alt. 700-800 m.
Distribution: Only known from southern Yunnan, China.
Note: This species is very similar to B. heteroclita. It is distinct by having rhizome
climbing on tree-trunks over two meters in height. In addition, it can be distinguished from
the latter by having lateral pinnae usually three-paired, terminal pinnae never obviously
prolonged and leaf texture thickly papyraceous. We suspect that it may be an ecological form
of B. heteroclita. More collections and further observation in the field are needed for a better
understanding of this species.
13. Bolbitis confertifolia Ching in Acta Phytotax. Sin. 21: 211. 1983; Chu H. Wang in Fl.
Reip. Pop. Sin. 6 (1): 108. 1999. Type: China. Yunnan (云南): Jinghong (景洪), Sino-USSR
Yunnan Exped. (中苏联合云南考察队) 1851 (holotype, PE!).
密叶实蕨
Habitat: In rain forests.
Distribution: Only known from the type locality.
Note: This species is poorly known to date because of only a type available for study. Its
rhizome which was originally described erect, is actually long-creeping. Many long roots are
present on the ventral surface of rhizome and four series of stipes on the dorsal surface of
rhizome. The stipes of fronds are jointed to the rhizome nearly at a 30º angle. This species is
close to B. heteroclita but differs by having ovate-lanceolate scales at the base of stipe. The
scales in B. heteroclita and other species of Bolbitis are usually narrowly lanceolate.
14. Bolbitis subcordata (Copel.) Ching in C. Chr., Ind. Fil. Suppl. 3: 50. 1934; Tardieu & C.
Chr. in Lecomte, Fl. Gén. Indo-Chiné 7: 433. 1941; K. Iwats. in Acta Phytotax. Geobot. 18:
54, fig. 11. 1959; Hennipman, Monogr. Gen. Bolbitis 280, figs. 81: a-c, 82. 1977; J. L. Tsai &
W. C. Shieh in T. C. Huang et al., Fl. Taiwan., ed. 2, 1: 355. 1994; Chu H. Wang in Fl. Reip.
Pop. Sin. 6 (1): 110, pl. 20: 1-12. 1999. ——Campium subcordatum Copel. in Philip. Journ.
Sci. 37: 369, fig. 23, pl. 16. 1928. Type: China. Hainan (海南): Mt. Wuzhishan (五指山), C.
C. C. McClure 9436 (holotype, P?; isotypes, BISH, BM, C, MO, P).
B. media Ching & Chu H. Wang in Acta Phytotax. Sin. 21: 212. 1983; Chu H. Wang in
Fl. Reip. Pop. Sin. 6 (1): 112. 1999, syn. nov. Type: China. Hainan (海南): Wanning (万宁),
Y. Chong (钟义) 3952 (holotype, PE!).
华南实蕨 Fig. 2: A, B

No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 111
Representative specimens examined:
China. Fujian (福建): Fuzhou (福州), W. M. Chu et al. (朱维明等) 17889 (PYU);
Nanjing (南靖), Fujian Exped. (福建队) 526, 545, 606 (PE), N. S. Zhou (周楠生) 441 (PE);
Yongchun (永春), Nandadui (南大队) 22485 (PE); Wuping (武平), Meihuashan Exped. (梅
花山队) 161 (IBSC); Without precise locality, S. G. Tang 13096, 16254, 16286, 16455,
16473 (SYS). Guangdong (广东): Dapu (大埔), L. Deng (邓良) 5435 (IBSC); Mt.
Dinghushan (鼎湖山), P. Zeng (曾沛) 10319 (SYS); Ruyuan (乳源), C. H. Tsoong (钟济新)
10768 (IBSC); Shenzhen (深圳), Shenzhen Exped. (深圳队) 189, 639 (PE); Shixing (始兴),
H. G. Ye et al. (叶华谷等) 1049 (IBSC); L. Deng (邓良) 6860 (PE, IBSC, KUN); Xinfeng
(新丰), H. G. Ye (叶华谷) 1041 (IBSC); Y. W. Taam 221 (SYS); Yangchun (阳春), H. G. Ye
et al. (叶华谷等) 400 (IBSC); Yu’nan (郁南), H. G. Ye et al. (叶华谷等) 2730 (IBSC);
Yingde (英德), C. Wang (黄志) 31559 (SYS), S. Wang (黄成) 163430 (PE, IBSC), 163811
(PE, IBSC, KUN); B. S. Wang (王伯荪) 12407 (SYS), H. T. Chang (张宏达) 7034 (SYS), P.
Zeng (曾沛) 12124 (SYS), Q. G. Wu (吴七根) 0029 (SYS), Y. K. Wang 31559 (PE).
Guangxi (广西): Cangwu (苍梧), S. H. Chun (陈少卿) 10173 (IBSC, KUN); Guiping (桂平),
H. Li et al. (黎桦等) 1019, 1021 (PYU); Jinxiu (金秀), Dayaoshan Exped. (大瑶山考察队)
13567 (IBSC), Y. J. Wang (王燕杰) 5190 (PE); Rongshui (融水), S. H. Chun (陈少卿) 15679
(PE, IBSC, KUN); Without precise locality, A. N. Steward et al. 1084 (PE). Hainan (海南):
Changjiang (昌江), S. Y. Dong et al. (董仕勇等) 168 (PE); Ledong (乐东), S. Y. Dong et al.
(董仕勇等) 218, 260 (PE); Lingshui (陵水), H. Feng (冯钦) 20146 (PE, SYS), S. Y. Dong
(董仕勇) 364, 378 (PE), Hainan Exped. (海南队) 1920 (PE), Z. X. Li et al. (李泽贤等) 1512
(IBSC); Mt. Wuzhishan (五指山), F. A. McClure 2789, 2880 (SYS), Hainan Exped. (海南队)
1816 (PE); Qionghai (琼海), E. Hainan Exped. (海南东队) 365 (PE, KUN), CAS Trop. For.
Exped. (中科院热带林调查队) 00365 (IBSC); Sanya (三亚), H. Y. Liang (梁向日) 62315
(PE); Wanning (万宁), S. Y. Dong et al. (董仕勇等) 557, 558 (PE), Z. X. Li et al. (李泽贤等)
4891 (IBSC); F. W. Xing et al. (邢福武等) 5404 (IBSC); P. Zeng (曾沛) 12864 (SYS); Y.
Chong (钟义) 3952 (IBSC); Without precise locality, P. Zeng (曾沛) 12522 (SYS), C. Wang
(黄志) 34232 (PE). Hong Kong (香港): Mt. Damaoshan (大帽山), S. Y. Hu (胡秀英) 9745
(PE), W. T. Tsang (曾怀德) 21228 (PE, SYS). Jiangxi (江西): Quannan (全南), J. F. Cheng
(程景福) 64430 (PYU), 64435 (PE, PYU). Taiwan (台湾): Without precise locality, Tanaka
s.n. (PE). Yunnan (云南): Menghai (勐海), W. M. Chu et al. (朱维明等) 6856 (PE);
Guangnan (广南), Z. R. Wang (王中仁) 403 (PE). Zhejiang (浙江): Yueqing (乐清), J. X.
Wang (王景祥) 1543 (PE), P. S. Qiu et al. (裘佩熹等) 6377 (PE).
Habitat: In rain forests, alt. 200-370 m.
Distribution: China (SE, S and SW China), Vietnam (Tonkin), Japan (southern Kyushu
to Ryukyu).
Note: The types of both B. media and B. subcordata were collected from the Hainan
Island. There are a lot of collections of B. subcordata but only a type specimen of B. media
has been seen in the major Chinese herbaria. Observation in the field has shown that B.
subcordata is common in Hainan yet B. media is a doubtful species. Ching and Wang (1983)
noted that B. media is similar to B. subcordata but differs by having plant smaller, stipe
slender, lateral pinnae smaller, subentire, coriaceous, and lateral veins beneath not distinct.
Examination of the type of B. media has shown that the leaves are not coriaceous but
herbaceous and the specimen is an immature plant of B. subcordata with slender and
Acta Phytotaxonomica Sinica Vol. 43 112
indistinct veins.
15. Bolbitis christensenii (Ching) Ching in C. Chr., Ind. Fil. Suppl. 3: 47. 1934; Tardieu &
C. Chr. in Lecomte, Fl. Gén. Indo-Chiné 7: 437. 1941; Hennipman, Monogr. Gen. Bolbitis
303, fig. 86: e, f. 1977; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 109. 1999. ——Campium
christensenii Ching in Bull. Fan Mem. Inst. Biol. Bot. 2: 214, pl. 31. 1931. Type: China.
Guizhou (贵州): Puding (普定), H. J. Esquirol 2672 (holotype, K?).
贵州实蕨
Habitat: In forests near streams.
Distribution: China (Guizhou), Vietnam (Tonkin).
Note: The present fern is poorly known because of lacking enough collections. It is close
to B. subcordata but differs in the venation pattern. The included free veinlets are present in
B. subcordata but not in B. christensenii.
16. Bolbitis hekouensis Ching in Acta Phytotax. Sin. 21: 212. 1983; Chu H. Wang in Fl.
Reip. Pop. Sin. 6 (1): 109. 1999. Type: China. Yunnan (云南): Hekou (河口), S. K. Wu (武素
功) 4056 (holotype, PE!; isotype, KUN!).
河口实蕨 Fig. 1: L
Representative specimens examined:
China. Yunnan (云南): Hekou (河口), K. H. Cai (蔡克华) 748 (PE), Dept. Biol.
Yunnan Univ. Exer. Exped. (云南大学生物系实习队) 2367 (PYU), W. M. Chu (朱维明)
5864 (PYU), W. M. Chu et al. (朱维明等) 19348, 21931 (PYU); Maguan (马关), W. M. Chu
(朱维明) 8413 (PYU).
Habitat: Usually in rocky crevices of limestone area in forests, alt. 400-1100 m.
Distribution: Only known from southern Yunnan, China.
Note: This species is slightly close to B. rivularis (Brackenr.) Ching from the Pacific
Islands, but differs in the venation pattern and geographical distribution. There are 1-2 pair(s)
of veinlets arising from each side of lateral veins in B. hekouensis but the similar veinlets are
3-4 pairs in B. vivularis. In distribution, B. hekouensis occurs only in southern Yunnan,
China, while B. vivularis has a wide distribution, occurring in New Guinea, Vanuatu, and Fiji.
17. Bolbitis tibetica Ching & S. K. Wu in C. Y. Wu, Fl. Xizang. 1: 276. 1983; Chu H. Wang
in Fl. Reip. Pop. Sin. 6 (1): 106. 1999. Type: China. Xizang (西藏): Mêdog (墨脱),
Qinghai-Xizang Exped. (青藏队) 74-4551 (holotype, PE!; isotype, KUN!).
西藏实蕨 Fig. 1: I
Habitat: In broad-leaved forests, alt. 800 m.
Distribution: Only known from the type locality.
Note: This species is distinct in the genus Bolbitis by having venation anastomosing,
bulbil terminal, stipe and rachis densely scaly.
18. Bolbitis hainanensis Ching & Chu H. Wang in Acta Phytotax. Sin. 21: 214. 1983; Chu H.
Wang in Fl. Reip. Pop. Sin. 6 (1): 115. 1999. Type: China. Hainan (海南): Without precise
locality, C. Wang (黄志) 35870 (holotype, PE!).
B. yunnanensis Ching in Acta Phytotax. Sin. 21: 214. 1983; Chu H. Wang in Fl. Reip.
Pop. Sin. 6 (1): 113. 1999, syn. nov. Type: China. Yunnan (云南): Simao (思茅), R. C. Ching
(秦仁昌) s.n. (holotype, PE!).
厚叶实蕨 Fig. 2: G, H
Representative specimens examined:
China. Yunnan (云南): Jinghong (景洪), Sino-USSR Yunnan Exped. (中苏联合云南
考察队) 8156 (PE); Simao (思茅), R. C. Ching (秦仁昌) 594, 595, 618 (PE).
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 113
Habitat: In dense forests.
Distribution: Endemic to China (Hainan and Yunnan).
Note: B. yunnanensis Ching is very similar to B. hainanensis in morphology. Both are
distinct among Chinese Bolbitis by having thick papyraceous laminae and wrinkled undulate
margin of leaves. In addition, the two species share many characters, such as scales on rachis
and costae narrowly lanceolate, terminal pinnae conforming to the lateral ones, a lamina
provided with 2-3 pairs of lateral pinnae, and venation pattern without included free veinlets.
As the two species cannot be clearly distinguished from each other, we reduce B. yunnanensis
herein as a synonym of B. hainanensis.
19. Bolbitis scalpturata (Fée) Ching in C. Chr., Ind. Fil. Suppl. 3: 50. 1934; K. Iwats. in Acta
Phytotax. Geobot. 18: 59. 1959; Hennipman, Monogr. Gen. Bolbitis 163, fig. 43: a-d. 1977;
et in Steenis & Holttum, Fl. Mal. Ser. II, 1 (4): 321. 1978; C. M. Kuo in Bot. Bull. Acad. Sin.
31: 308, fig. 1. 1990. —— Heteroneuron scalpturatum Fée, Hist. Acrost. 95, t. 56. 1845.
Type: Philippines. Manila: Without precise locality, Gaudichaud s.n. (holotype, P).
红柄实蕨 Fig. 2: D, E
Representative specimens examined:
China. Hainan (海南): Mt. Diaoluoshan (吊罗山), S. Y. Dong et al. (董仕勇等) 549
(PE); Mt. Qixianling (七仙岭), S. Y. Dong (董仕勇) 919 (PE).
Habitat: On rocks in forests, alt. 0-1200 m.
Distribution: China, Myanmar, Thailand, Vietnam, Malaysia, Indonesia, Philippines.
Note: The present species is close to B. deltigera. Both species are characterized by
having pinnae lanceolate and chartaceous, lateral pinnae 5-7-paired, terminal pinnae
conforming to the lateral ones and subarticulate to the rachis, fertile pinnae generally not
acrostichoid, and type C spore. But these two species is quite different in the venation pattern.
In B. deltigera, there are 4-5 veins arising from each side of lateral veins anastomosing,
areoles mostly with 1-3, usually 2 included free veinlets which thicken at the end. Whereas in
B. scalpturata, there are only 2-3 veins arising from each side of lateral veins anastomosing,
only a small part of areoles with 1, rarely 2 included free veinlets which do not thicken at the
end. In addition, the costae of pinnae in B. scalpturata, albeit not in every plant, turn purplish
when dry, yet the costae of pinnae in B. deltigera always remain stramineous.
20. Bolbitis deltigera (Bedd.) C. Chr., Ind. Fil. Suppl. 3: 48. 1934; Tagawa & K. Iwats., Fl.
Thail. 3 (3): 316. 1988. ——Poecilopteris costata var. deltigera Bedd., Ferns Br. Ind. 114, pl.
114. 1865. ——Bolbitis virens var. deltigera (Bedd.) Hennipman in Blumea 18: 149. 1970; et
Monogr. Gen. Bolbitis 184, figs. 47, 48: k-m. 1977. Type: Nepal. Between Katmandu and
Bhimpedy, Wallich 59 (holotype, K).
间断实蕨 Fig. 2: F
Representative specimens examined:
China. Hainan (海南): Mt. Qixianling (七仙岭), W. M. Chu et al. (朱维明等) 18174
(PYU). Yunnan (云南): Cangyuan (沧源), W. M. Chu et al. (朱维明等) 15438 (PYU);
Yingjiang (盈江), Yunnan Univ. West Yunnan Pl. Exped. (云南大学滇西植物调查组)
10901 (PE, PYU), 9921, 10606 (PYU).
Habitat: On slope or in valley under rain forests, alt. 340-700 m.
Distribution: China, Bangladesh, Nepal, Bhutan, Sikkim, India, Myanmar, Thailand.
21. Bolbitis angustipinna (Hayata) H. Ito in J. Jap. Bot. 14: 443. 1938; Pic. Serm., Ind. Fil.
Suppl. 4: 42. 1965; Hennipman, Monogr. Gen. Bolbitis 152, fig. 40: a-f. 1977; et in Steenis &
Holttum, Fl. Mal. Ser. II, 1 (4): 321, figs. 26c, 27a. 1978; Tagawa & K. Iwats., Fl. Thail. 3
Acta Phytotaxonomica Sinica Vol. 43 114
(3): 311. 1988; Chu H. Wang in Fl. Reip. Pop. Sin. 6 (1): 110, pl. 21: 4-6. 1999. ——
Leptochilus angustipinnus Hayata, Ic. Pl. Form. 5: 297, fig. 119. 1915. Type: China. Taiwan
(台湾): near Peikanghsi (北港溪), Owatari s.n. (holotype, TAI).
Acrostichum contaminans Wall., List no. 22. 1828, nom. nud.——Acrostichum
crispatulum var. contaminans Clarke, Trans. Linn. Soc. II, Bot. 1: 580, pl. 84, fig. 2A, C.
1880. ——Bolbitis contaminans Ching in C. Chr., Ind. Fil. Suppl. 3: 47. 1934; K. Iwats. in
Acta Phytotax. Geobot. 18: 53, fig. 9. 1959; J. L. Tsai & W. C. Shieh in T. C. Huang et al., Fl.
Taiwan., ed. 2, 1: 352. 1994. Type: Nepal. Between Helounda and Bhimpedy, Wallich 22
(holotype, K).
多羽实蕨 Fig. 2: I, J
Representative specimens examined:
China. Hainan (海南): Mt. Qixianling (七仙岭), S. Y. Dong (董仕勇) 920 (PE), new
record. Taiwan (台湾): Pingdong (屏东), Faurie 206, 281 (PE). Yunnan (云南): Jinghong
(景洪), W. M. Chu et al. (朱维明等) 525 (PYU), B. Y. Qiu (邱炳云) 57880 (KUN), J. F.
Rock 2427 (PE), Sino-USSR Yunnan Exped. (中苏联合云南考察队) 5699 (PE, KUN), 9541
(KUN), Yunnan First Group Exped. (云南一组) 95 (PE); Lüchun (绿春), S. K. Wu et al. (武
素功等) 888 (KUN).
Habitat: On rocks or at base of tree-trunks in dense forests, alt. 850 m.
Distribution: China, Nepal, Bhutan, Sikkim, India, Sri Lanka, Myanmar, Thailand.
Note: The terminal part of lamina is often a pinna which conforms to the lateral pinnae.
But sometimes a lamina with a narrowly triangular apical part instead of a pinna is also
present in some fronds.
22. Bolbitis costata (C. Presl) Ching in C. Chr., Ind. Fil. Suppl. 3: 47. 1934; Hennipman,
Monogr. Gen. Bolbitis 155, fig. 41. 1977; Tagawa & K. Iwats., Fl. Thail. 3 (3): 311. 1988.
——Campium costatum C. Presl, Tent. Pterid. 238. pl. X. 23. 1836. Type: Bangladesh.
Sylhet, Wallich 26 (holotype ?; isotypes, B, K, P, W).
紫轴实蕨 Fig. 2: K, L
Representative specimens examined:
China. Yunnan: Yingjiang (盈江), Yunnan Univ. W Yunnan Pl. Exped. (云南大学滇西
植物调查组) 10664 (PE, PYU), new record.
Habitat: In valley in forests, alt. 360 m.
Distribution: China, Bangladesh, Nepal, Sikkim, India, Myanmar, Thailand.
Note: The present fern was first found in China by Professor W. M. Chu of Yunnan
University from western Yunnan. A common character of this species and B. scalpturata is
that the fronds usually turn purplish when dry. However, the present fern is very different
from the latter by having sterile pinnae 4-9 cm wide, 5-8 veinlets arising from each side of
lateral veins, and fertile pinnae linear (14-16 times as long as wide). On the contrary, in B.
scalpturata, the sterile pinnae are 1.5- 4 cm wide, 2-3 veinlets arise from each side of lateral
veins, and the fertile pinnae are lanceolate (3-8 times longer than wide).
23. Bolbitis virens (Hook. & Grev.) Schott, Gen. Fil. pl. 13. 1834; Hennipman in Blumea 18:
149. 1970, p.p., et Monogr. Gen. Bolbitis 180, p.p., fig. 48: a-g. 1977. ——Acrostichum virens
Hook. & Grev., Ic. Fil., t. 221. 1831. Type: Myanmar. Tovag, Wallich 1033 (holotype, K).
Bolbitis latipinna Ching in Acta Phytotax. Sin. 21: 213. 1983; Chu H. Wang in Fl. Reip.
Pop. Sin. 6 (1): 112, pl. 21: 7-9. 1999, syn. nov. Type: China. Yunnan (云南): Jinghong (景
洪), C. W. Wang (王启无) 78807 (holotype, PE!; isotype, KUN!).
宽羽实蕨
No. 2 DONG Shi-Yong et al.: A taxonomic revision of the fern genus Bolbitis from China 115
Representative specimen examined:
China. Yunnan (云南): Jinghong (景洪), C. W. Wang (王启无) 78807A (IBSC).
Habitat: On rocks in forests, alt. 850 m.
Distribution: China (Yunnan), Bangladesh, Myanmar, Thailand.
Note: The type of B. latipinna Ching agrees well with the type of B. virens. Hennipman
(1977) had studied it and cited it under B. virens.
Acknowledgements We express our gratitude to the Wild Animals and Plants Conservation
Center of Hainan for the assistance in the field work. We also thank Mr. Y. H. XIAO
(Institute of Botany, the Chinese Academy of Sciences) for his help in the SEM observation
of spores. We are grateful to Prof. W. M. CHU and Dr. Z. R. HE (Yunnan University) for
assistance in checking some specimens in PYU. Finally, many thanks go to the curators and
staff of PE, PYU, IBSC, SYS, and KUN for the facilities, and to the curator and staff of K for
providing some type photographs.
References
Christensen C. 1906. Index Filicum. Hafniae: H. Hagerup. 161, 301.
Christensen C. 1931. Asiatic Pteridophyta collected by Joseph F. Rock 1920-1924. Contributions from the
United States National Herbarium 26: 265-337.
Ching R-C (秦仁昌). 1931. On the genus Egenolfia Schott. Bulletin of the Fan Memorial Institute of Biology
11: 297-317.
Ching R-C (秦仁昌), Wang C-H (王铸豪). 1983. Materiae ad floram filicum sinensium. Acta Phytotaxono-
mica Sinica (植物分类学报) 21: 211-218.
Chu W-M (朱维明), Zhou H-G (周厚高). 1994. Some pteridophytes new to Hainan and China. Acta Botanica
Yunnanica (云南植物研究) 16: 123-130.
Hennipman E. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden: Leiden University
Press.
Iwatsuki K. 1959. Taxonomic studies of Pteridophyta IV. Acta Phytotaxonomica et Geobotanica 18: 44-59.
Kuo C-M. 1990. Materials for the Lomariopsidaceae of Taiwan. Botanical Bulletin of Academia Sinica 31:
305-304.
Wang C-H (王铸豪). 1999. Bolbitidaceae. In: Flora Reipublicae Popularis Sinicae (中国植物志). Beijing:
Science Press. 6 (1): 104-124.
中国实蕨属的分类修订
1, 2董仕勇 2张宪春
1 (中国科学院华南植物园 广州 510650)
2 (中国科学院植物研究所系统与进化植物学重点实验室 北京 100093)
摘要 对实蕨属Bolbitis的17种植物的孢子进行了扫描电镜观察。根据孢子周壁特征,中国产实蕨属的孢
子明显可分为3种类型：A型孢子具网状周壁,B型孢子具鸡冠状-波状周壁,C型孢子具平滑的波状周壁。
孢子周壁特征、叶脉式样和叶片顶部的形态是实蕨属中最有价值的分类学性状。根据标本检查,结合野
外调查和孢子形态观察,对中国产实蕨属的分类进行了修订,确定中国有实蕨属植物20种和3杂交种,其
中包括2个新组合B. fengiana (Ching) S. Y. Dong和B. medogensis (Ching & S. K. Wu) S. Y. Dong,以及2个
中国新分布B. costata Ching ex C. Chr.和B. hookeriana K. Iwats.。将B. latipinna Ching、B. media Ching &
Chu H. Wang、B. yunnanensis Ching、Egenolfia crassifolia Ching、E. crenata Ching & P. S. Chiu、E. fengiana
Ching、E. medogensis Ching & S. K. Wu和E. ×yunnanensis Ching & P. S. Chiu等8个名称处理为新异名。
文中给出了分种检索表、每个种的生境和分布资料、大多数种的特征集要和孢子扫描电镜照片。
关键词 实蕨属; 实蕨科; 分类修订; 孢子形态; 中国