DAVALLIACEAE [Draft]
骨碎补科 gu shui bu ke
Xing Fuwu (Ch), Wang Faguo (Ch)*, Hans P. Nooteboom**
Small to moderate-sized ferns, epiphytes or rarely terrestrial. Rhizome long-creeping, dorsiventral, densely covered with peltate or sometimes basifixed scales, the fronds usually borne in two rows. Leaves remote, the stipes articulated at the base to short or long phyllopodia; lamina pinnatifid to 4-pinnate pinnatifid, firmly herbaceous to coriaceous, glabrous or casually covered with scales or hairs; veins free, usually forked. Sporangia borne in a small discrete sori terminal on the veins, submarginal or sometimes medial; indusium opening towards the margin, attached at base and sometimes at the sides, round, reniform or elongate towards the margin, pedicel long, 3-seriate, annulus longitudinal, consisted of 12–16 thickened cells; spores monolete, elliptic or long-elliptic, translucent, usually without perispore.
Three genera, ca. 45 species; mostly in tropical or subtropical Asia, 2 genera are represented by 14 species mostly distributed in SW and S China.
1. Terrestrial, sometimes epiphytic; lamina compound; scales basifixed along broad base, roots borne on all sides of rhizome; sori large, ca. 2.5 mm wide.............................................................. 2. Leucostegia
1. Epiphytic or epilitic; lamina compound or not; scales peltate, or basifixed with cordate base and much overlapping lobes, roots restricted to the ventral side of lateral buds; sori smalle, ca. 1 mm wide. 1. Davallia
1. LEUCOSTEGIA C. Presl, Tent. Pterid. 94, pl. 4, f. 11. 1836.
大膜盖蕨属 da mo gai jue shu
Rhizome bearing scales and hairs or only scales. Roots borne on all sides of rhizome. Scales glabrous or bearing multiseptate hairs, basifixed, base not cordate. Stipes articulated at the base to phyllopodia, grooved or not, glabrous. Lamina compound, tripinnate or quadripinnate (rarely in small plants bipinnate) towards base and in the middle part, deltoid and broadest towards base, glabrous (sometimes minute hairs present), not or slightly dimorphous (often fertile leaves more strongly dissected ). Pinnae deltoid or narrowly triangular. Pinnules of at least the larger pinnae anadromous. Pinnules or pinnalobes narrowly ovate. Leaf axes glabrous (sometimes a few minute hairs present). Veins in ultimate lobes simple, not reaching the margin. False veins not present. Sori indusiate, frequently single on a segment, terminal on the veins. Indusium scale-like, either attached at the narrow, cordate base only, or attached at the base and only part of the sides. pedicel very long, 3-seriate, annulus longitudinal, interrupted of ca. 16 thickened cells. Spores elliptic, monolete, closely tuberculate, translucent. n = 41.
Two species in India, Sikkim, Myanmar, Nepal, Polynesia, Thailand, Indochina Peninsula; one species in China.
Leucostegia immersa Wallich ex C. Presl, Tent. Pterid. 95, pl. 4, f. 11. 1836
大膜盖蕨 da mo gai jue
Acrophorus immersus (Wallich ex C. Presl) T. Moore; Davallia immersa [Wallich, nomen nudum] ex Hooker; Humata dryopteridifrons Hayata; H. immersa (Wallich ex C. Presl) Mett.
Rhizome robust, long-creeping, 3.5–5 mm thick, densely covered with scales and yellowish brown hairs; scales lanceolate or ovate-lanceolate, ca. 3 mm long, brown, membranous, apex long acuminate, margin nearly entire Leaves remote, 2–5 cm apart; stipe 25–35 cm long, stramineous; lamina deltoid, 25–35 cm long, nearly as wide, anadromous, papyraceous, slightly green when dry, apex acuminate, tripinnate, glabrous; pinnae 10–12 pairs, basal pair subopposite, 4–6 cm apart, long triangular and broadest towards the base, with stalk 5–20 mm long, base broadly cuneate, largest pinnae ovate-lanceolate, 12–22 × 6.5–12 cm, apex acuminate; pinnules 8–10 pairs, short stalked, alternate, ovate to elliptic, deflective, its basal pinnule 5–8 cm long, base asymmetrical; ultimate leaflets rhomboid, only shallowly lobed; ultimate segments (lobes) 3–4 mm wide; margin serrate; veins distinct, forking, in ultimate lobes simple, ending in submarginal round hydathodes. Sori separate, borne 1–2 on a segment, terminal on acroscopic veinlet; indusium attached by base only, oval or elliptic, 1–1.5 × 1–2 mm, thinly papyraceous.
Epiphytic and terrestrial on shaded hillsides; 1000–2800 m. Guangxi, Taiwan, Xizang, Yunnan [Bhutan, Cambodia, India, Indonesia, Malaysia, Myanmar, New Guinea, Philippines, Sikkim, Thailand; Pacific Islands (Polynesia)].
2. DAVALLIA Smith in Mém. Accad. Sci. Turin 5: 414. 1793.
骨碎补属 gu sui bu shu
Araiostegia Copeland; Davallia Sect. Cordisquama Kato; Davallia sect. Davallia Kato; Davallia sect. Wibelia Kato; Humata Cavanilles;Wibelia Bernhardi (non Fee 1852); Paradavallodes Ching; Trogostolon Copeland.
Roots restricted to the ventral side of lateral buds. Scales of rhizome peltate or basifixed with cordate base and greatly overlapping lobes, variously shaped: distinctly acicular, or flat and nearly acicular, narrowed evenly towards the apex, or narrowed abruptly from a broad base, or broad, ovate to oblong-subdeltoid with round to acute apex. Petioles usually well developed, in two ranks on the dorsal side of the rhizome; adaxial face sulcate, the groove usually raised in the middle; small species sometimes with subsessile leaves; petiole occasionaly persistently scaly. Lamina simple, imparipinnate, pinnate + pinnatifid, bipinnate + bipinnatifid, or tripinnate + tripinnatifid; if compound deltoid and broadest towards base or rarely elongate, glabrous or rarely bearing multicellular hairs, anadromous or rarely isodromous or catadromous and then the scales evenly narrowed towards the apex; lamina dimorphic or monomorphic, in dimorphic species with reduced leaf tissue and/or more dissected. Vein endings on sterile segments reaching the margin or not. False veins present in several species. Rhachis winged and therefore seemingly grooved adaxially but convex between the wings. In the dry state (re herbarium specimens) it is difficult to see whether the rhachis itself is grooved or flat. Sori typically separate but in D. undulata connate and elongate along leaf margins; sori near the margin, facing midveins at the forking point of veins or at the bending point of a vein.
About 35 species from India through continental SE Asia to China, Korea, and Japan; Malesia; the Pacific to Samoa and New Zealand; NE Australia; the islands in the Indian Ocean; Africa; one species in NW Africa, the Canary Islands, and SW Europe. 11 Species in China.
1a. Lamina elongate, often narrowing towards base.......................................................... 2
2a. Stipes bearing hairs and or scales when young.................. 6. Davallia membranulosa
2b. Stipes glabrous or with few scales............................................................................ 3
3a. Lamina simple, one pectinate or pinnatifid leaf, rhizome white waxy under the scales, scales with pale border from base to apex, bearing multiseptate hairs at least when young.... 8. Davallia pectinata
3b. Lamina compound, or pinnate towards base, rhizome not white waxy, scales without pale border, not bearing multiseptate hairs.................................................................................................... 4
4a. Scales broad, ovate to oblong-subdeltoid with round to acute apex, lacking marginal setae or teeth, or those rare, basifixed with cordate base and much overlapping lobes, 2 to 5 mm long, lamina tripinnate or quadripinnate, indusium scale-like, attached at the narrow, cordate base only.................. 9. Davallia pulchra
4b. Scales narrowed evenly towards the apex, toothed, peltate, 8 to 10 mm long, lamina pinnate with pinnatilobed to pinnatifid pinnae, indusium attached at the broad base and hardly or not at the sides 1. Davallia assamica
1b. Lamina deltoid and broadest towards base................................................................. 5
5a. Indusium also attached along the sides, pouch-shaped, oblong.................................. 6
6a. Sori frequently single on a segment............................. 12. Davallia trichomanoides
6b. Sori borne several on a segment............................................................................ 7
7a. Scales bearing multiseptate hairs at least when young............... 11. Davallia solida
7b. Scales not bearing multiseptate hairs.................................................................... 8
8. Scales basifixed with cordate base and much overlapping lobes, false veins not present. 4.Davallia divaricata
8. Scales peltate, false veins present...................................... 3. Davallia denticulate
5b. Indusium scale-like, attached at the narrow, cordate base only, at the base and only part of the sides, or attached at the broad base and hardly or not at the sides, reniform, semicircular, or more or less triangular to rhomboid 9
9a. Leaf axes and rhachises hairy........................................... 7. Davallia multidentata
9b. Leaf axes glabrous.............................................................................................. 10
10a. Scales broad, ovate to oblong-subdeltoid with round to acute apex 9. Davallia pulchra
10b. Scales narrowed evenly towards the apex........................................................ 11
11a. Scales basifixed with cordate base and much overlapping lobes, lamina compound, lowest pinnule inserted at very base of pinna rhachis.................................................................. 2. Davallia clarkei
11b. Scales peltate, if lamina compound lowest pinnule not inserted at very base of pinna rhachis 12
12. Rhizome 0.5--3 mm diam., white waxy under the scales, lamina from simple, trifoliolate to pectinate or compound................................................................................................ 10. Davallia repens
12. Rhizome 3--6 mm diam., not white waxy, lamina compound...............................
.................................................................................................... 5. Davallia griffithiana
1. Davallia assamica (Bedd.) Baker in Hook. & Baker, Syn. Fil. 452. 1868.
长叶阴石蕨 chang ye yin shi jue
Acrophorus assamicus Beddome; Davallia micans Mettenius ex Baker; Humata assamica (Beddome) C. Christensen; H. micans (Mettenius ex Baker) Diels; Leucostegia assamica (Beddome) J. Smith.
Rhizome without the scales 3--5.3 mm diam., not white waxy. Scales whitish or red brown, without pale border, narrowed evenly towards the apex, not or seldom curling backward, not bearing multiseptate hairs, toothed, peltate, 8--10 x 1.8--2.5 mm. Stipes adaxially grooved, 4--7 cm long, glabrous or with few scales. Lamina narrowly ovate, pinnate with pinnatilobed to pinnatifid pinnae towards base and in the middle part, or sometimes bipinnate, elongate, glabrous, 10--27 x 3.5--12 cm, not or slightly dimorphous. Longest petiolules 1 mm long. Pinnae narrowly ovate. Longest pinnae 3.5--6 x 1--2 cm. Pinnalobes of at least the larger pinnae anadromous, linear oblong, longest 7--15 x 3--4 mm. Leaf axes glabrous. Margins of the lamina of each leaflet thickened and decurrent on the edge of the grooved rhachis (in fertile leaves). Veins in sterile ultimate lobes pinnate, not reaching the margin. False veins not present. Sori separate, borne several or frequently single on a segment, at the forking point of veins. Indusium attached at the broad base and hardly or not at the sides, semicircular, wider than long or about as wide as long, 0.7--1 x 0.8--1.2 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin. Lamina generally extending into a tooth at both sides or only at the outside of a sorus.
Wet forests, climbing on rocks or tree trunks; 900–2300 m. Yunnan, Xizang [Bhutan, India, Myanmar].
2. Davallia clarkei Baker in Hooker & Baker, Syn. Fil. 91. 1874.
宿枝小膜盖蕨 su zhi xiao mo gai jue
Araiostegia faberiana Ching; A. hookeri Ching (non Davallia Hookeriana Wallich 1828; Hk. 1846); A. parvipinnula Ching; A. perdurans Copeland; Davallia clarkei var. faberiana C. Christensen; D. parvipinnula Hayata; D. perdurans Christ; D. subalpina Hayata; Humata hookeri Diels; Leucostegia clarkei C. Christensen; L. clarkei var. faberiana C. Christensen.; L. faberiana Ching; L. hookeri Beddome; L parva C. Christensen; L. parvipinnula Hayata; L. perdurans C. Christensen.
Rhizome without the scales 5 mm diam., not white waxy. Scales brown, without pale border, narrowed evenly towards the apex, not bearing multiseptate hairs, lacking marginal setae or teeth, or those rare, or toothed, basifixed with cordate base and much overlapping lobes, 7--10 mm long. Stipes pale, adaxially grooved, 7--35 cm long, glabrous or with few scales. Lamina compound, tripinnate, or quadripinnate, deltoid and broadest towards base, glabrous, 10--50 x 6--50 cm, not or slightly dimorphous. Longest petiolules 0 mm long (generally lowest pinnule at very base of pinna-rhachis). Pinnae deltoid or linear-triangular. Longest pinnae 2--30 x1.2--18 cm. Both lowest pinnules of at least basal pinnae inserted on pinna base, other pinnules anadromous (sometimes only one pinnule), deltoid or narrowly ovate. Longest pinnules 6--130 x 2--50 mm. Ultimate leaflets linear oblong, lobed almost to the midrib. Ultimate segments or lobes obtuse or acute without a tooth, 1--4 x 0.2--1 mm. Leaf axes glabrous. Veins in sterile ultimate lobes frequently simple, not reaching the margin. False veins not present. Sori separate, frequently single on a segment, at the forking point of veins. Indusium reniform or semicircular, attached at the narrow, cordate base only, or attached at the broad base and hardly or not at the sides, wider than long or about as wide as long, 0.3--1.2 x 0.4--1.5 mm.
Dense mixed forests, epiphytic on tree trunks; 1200–4000 m. Fujian, Guangxi, Guizhou, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Myanmar, Nepal, Sikkim, , Thailand].
3. Davallia denticulata (Burman) Mettenius ex Kuhn, Filic. Decken. 27. 1867.
假脉骨碎补 jia mai gu sui bu
Adiantum denticulatum Burman; Davallia bidentata Schkuhr; D. brevisora Ching; D. chaerophylloides Steudel; D. elegans Swartz; D. elegans var. bidentata Hooker; D. elegans var. coniifolia Hooker; D. elegans var. edentula Hooker; D. elegans var. polydactyla Moore; D. elegans var. pulchra Hooker; D. elegans var. subunidentata Hooker; D. impressa Copeland; D. patens Swartz; Humata chaerophylla Mettenius; H. chaerophylloides Desvaux; H. elegans Desvaux; H. patens Desvaux; Leucostegia chaerophylla J.Smith; L. pseudocystopteris Beddome; Parestia elegans C. Presl; Trichomanes denticulatum Houttuyn; T. chaerophylloides Poiret; T. elegans (Swartz) Poiret (1808), not Richard (1792); T. lucidum Roxburgh
In China only var. denticulata.
Rhizome without the scales 3--15 mm diam., not white waxy. Scales red brown or nearly black, with pale border from base to apex or without pale border, narrowed evenly towards the apex or flat and nearly acicular, narrowed abruptly from a broad base, often curling backward, not bearing multiseptate hairs, toothed, peltate, 4--8 x 0.5--1.5 mm. Stipes pale, adaxially grooved, 4--50 cm long, glabrous or with few scales. Lamina compound, bipinnate or quadripinnate towards base and in the middle part, deltoid and broadest towards base, glabrous, 16--90 x 13--50 cm, not or slightly dimorphous. Longest petiolules 4--35 mm long. Pinnae deltoid. Longest pinnae 8--45 x 5--30 cm. Pinnules of at least the larger pinnae anadromous, deltoid. Longest pinnules 70--200 x 40--110 mm. Ultimate leaflets linear oblong or narrowly ovate, lobed almost to the midrib or only shallowly lobed. Ultimate segments 5--27 x 2--6 mm. Leaf axes glabrous. Margins of the lamina of each leaflet not thickened. Veins in sterile ultimate lobes pinnate (or forked in very narrow lobes), reaching the margin. False veins present. Sori separate, borne several on a segment, at the forking point of veins. Indusium also attached along the sides, pouch-shaped, oblong, longer than wide or about as wide as long, 1--1.3 x 0.5--1 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin. Lamina generally extending into a tooth at both sides of a sorus.
Epiphyte on many different species of trees and in different types of forest incl. mangrove forest or on solitary trees, epilithic on granite, limestone, or sandstone, terrestrial on different kinds of soil; 0--2200 m. Hainan [Cambodia, India (Assam and Andaman and Nicobar is), Indonesia, Laos, Malaysia, Myanmar, Papua New Guinea, Philippines, Thailand, Vietnam; Australia (Queensland), Pacific (Samoa, Society Is. (Tahiti)),Africa and Indian Ocean (Madeira, Tropical and South Africa, Madagascar, Comores, Seychelles, Christmas I).
Note. Davallia brevisora Ching is a form with the false veins absent or
inconspicuous.
4. Davallia divaricata Blume, Enum. Pl. Javae 237. 1828
大叶骨碎补 da ye gu shui bu
Araiostegia divaricata Kato; Davallia amabilis Ching; D. austro-sinica Ching; D. divaricata var. orientalis Tardieu-Blot & C. Christensen; D. formosana Hayata; D. orientalis C. Christensen in Wu & al.
In China only var. divaricata
Rhizome without the scales 10--15 mm diam., not white waxy. Scales brown or red brown without pale border, narrowed evenly towards the apex, curling backward or not, not bearing multiseptate hairs, toothed, basifixed with cordate base and much overlapping lobes, 5--20 x 2--4 mm. Stipes pale, adaxially grooved, 15--60 cm long, glabrous or with few scales. Lamina compound, tripinnate towards base and in the middle part, deltoid and broadest towards base, glabrous, 60--100 x 40--70 cm, not or slightly dimorphous. Longest petiolules 4--35 mm long. Pinnae deltoid. Longest pinnae 8--45 x 5--30 cm. Pinnules of at least the larger pinnae anadromous, deltoid. Longest pinnules 70--200 x 40--110 mm. Ultimate leaflets linear oblong or narrowly ovate, lobed halfway towards midrib or only shallowly lobed. Ultimate segments 5--27 x 2--6 mm. Rhachis adaxially distinctly grooved (often with a groove at either side). Leaf axes glabrous. Margins of the lamina of each leaflet not thickened. Veins in sterile ultimate lobes pinnate (or forked in very narrow lobes), reaching the margin. False veins absent. Sori separate, borne several on a segment, at the forking point of veins. Indusium also attached along the sides, pouch-shaped, oblong, about as wide as long, 1 x 1 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin. Lamina generally extending into a tooth at both sides of a sorus.
Generally epiphytic, sometimes epilithic on limestone, or bedrock not specified; rarely terrestrial. Mostly in dense forest, sometimes on dry places; 0--1850 m. Fujian, Guangdong, Guangxi, Hainan, Hong Kong, Taiwan, Yunnan, [Cambodia, India, Indonesia, Laos, Myanmar, Malaysia, Thailand, Papua New Guinea, Philippines, Vietnam; Pacific (Solomon Is. (Bougainville, 1 coll.)).
Note. In TI I had the occasion to study the type of Davallia formosana Hayata. It fits perfectly well in the variability of D. divaricata.
5. Davallia griffithiana Hooker, Sp. Fil. 168, t. 49B. 1845.
杯盖阴石蕨 bei gai yin shi jue
Davallia henryana Baker; D. platylepis Baker; D. tyermannii T.Moore; Humata griffithiana C. Christensen; H. griffithiana var. tyermannii (T. Moore) Tagawa; H. henryana Ching; H. platylepis Ching; H. tyermanii T. Moore; Leucostegia griffithiana J. Smith.
Rhizome without the scales 3--6 mm diam., not white waxy. Scales whitish, brown, or red brown, without pale border, narrowed evenly towards the apex, curling backward or not, not bearing multiseptate hairs, with marginal setae at least in distal part, peltate, 6--9 x 1--1.5 mm. Stipes adaxially grooved, 6--24 cm long, glabrous or with few scales. Lamina compound, bipinnate or tripinnate towards base and in the middle part, deltoid and broadest towards base, glabrous, 10--32 x 8--27 cm, not or slightly dimorphous. Longest petiolules 2--10 mm long. Pinnae deltoid or linear. Longest pinnae 6--16 x 4--8 cm. Pinnules of at least the larger pinnae anadromous, linear oblong or narrowly ovate. Longest pinnules 25--100 x 7--60 mm. Ultimate leaflets narrowly ovate, lobed almost to the midrib or only halfway towards it. Ultimate segments or lobes obtuse or acute without a tooth, 2--5 x 2--3 mm. Leaf axes glabrous. Margins of the lamina of each leaflet not thickened. Veins in sterile ultimate lobes simple, forked or pinnate, not reaching the margin. False veins not present. Sori separate, borne several or single on a segment at the forking point of veins. Indusium attached at the base and only part of the sides, semicircular, about as wide as long, 1 x 1--2 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin. Lamina generally extending into a tooth at both sides or only at the outside of a sorus, or not extending into teeth beyond a sorus.
Wet forests; climbing on tree trunks or rocks; 400–2200 m. Zhejiang, Jiangxi, Hunan, Sichuan, Guizhou, Fujian, Taiwan, Guangxi, Guangdong, Yunnan, Xizang [Bhutan, India (Manipur, Assam), Japan (Okinawa) , Laos, Myanmar, (Kentung), Vietnam(Tonkin).
Note. Comparing the types of all the synonyms, which are all in K, convinced me that they all belong to the same characteristic species.
6. Davallia membranulosa [Wallich, Cat. (1828) 255, nomen], Hooker, Sp. Fil. 159. 1845.
膜叶假钻毛蕨 mo ye jia zuan mao jue
Acrophorus membranulosus (Wallich ex Hooker) T. Moore; Araiostegia membranulosa Holttum; Davallodes chingiae Ching; D. membranulosum Copeland; Leucostegia membranulosa (Wallich ex Hooker) J. Smith; Paradavallodes chingiae Ching; P. membranulosum Ching.
Rhizome without the scales 2--4 mm diam. Scales brown, red brown, or nearly black, with pale border from base to apex, narrowed evenly towards the apex, not bearing multiseptate hairs, lacking marginal setae or teeth or those rare, peltate, 5--6 mm long. Stipes pale, adaxially grooved, 3--15 cm long, bearing hairs and or scales when young. Lamina compound, bipinnate towards base and in the middle part, elongate, not or hardly narrowed towards base, bearing multicellular hairs, 12--27 x 5--14 cm, not or slightly dimorphous. Longest petiolules 0.5--1.5 mm long. Pinnae linear-triangular. Longest pinnae 2.6--7 x 1--3 cm. Pinnules of at least the larger pinnae catadromous or anadromous (but often opposite), linear oblong (pinnatipartite, the lobes entire or shallowly lobed). Longest pinnules 6--15 x 2--4 mm. Leaf axes, at least rhachises, hairy. Hairs 0.4--0.6 mm long. Veins in sterile ultimate lobes simple or forked, not reaching the margin. False veins not present. Sori separate, frequently single on a segment at the forking point of veins or (rarely) at the bending point of a vein. Indusium attached at the broad base and hardly or not at the sides, semicircular or oblong (to circular), longer than wide to wider than long, 0.5--0.8 x 0.5--0.8 mm.
Epilithic or epiphytic in forest; 600--2600 m. Yunnan, Sichuan [India (Darjeeling, Kumaon, Saharanpur), Myanmar (Kengtung state), Nepal, Sikkim, Thailand (north), Vietnam (Tonkin, Chapa)].
Note. Although Ching recognised two species, careful examination of the types could not but lead to the conclusion that there is only one, easily recognisable, species.
7. Davallia multidentata Hooker, Syn. Fil. 91. 1868
假钻毛蕨 jia zuan mao jue
[Aspidium multidentatum Wallich, nomen.]; Araiostegia multidentata Copeland; Leucostegia multidentata Beddome; Paradavallodes kansuense Ching; P. multidentatum Ching.
Rhizome without the scales 5 mm diam. (with scales c. 10), not white waxy. Scales brown without pale border, narrowed evenly towards the apex, often curling backward, not bearing multiseptate hairs, lacking marginal setae or teeth or toothed, basifixed with cordate base and much overlapping lobes, 6 mm long. Stipes pale or dark brown, adaxially grooved, 17--25 cm long, glabrous or with few scales. Lamina compound, tripinnate, deltoid and broadest towards base, glabrous, 30--45 x 17--34 cm, not or slightly dimorphous. Longest petiolules 8--10 mm long. Pinnae linear-triangular. Longest pinnae 10--19 x 6--9 cm. Pinnules of at least the larger pinnae anadromous, narrowly ovate. Longest pinnules 40--70 x 15--30 mm. Ultimate leaflets linear oblong, lobed halfway towards midrib. Ultimate segments or lobes obtuse or acute without a tooth, 2--4 x 1 mm (often shallowly lobed). Leaf axes, at least rhachises, hairy. Hairs 0.1--0.2 mm long. Veins in sterile ultimate lobes simple or forked, not reaching the margin. False veins not present. Sori separate, frequently single on a segment at the forking point of veins or at the bending point of a vein. Indusium reniform, attached at the narrow, cordate base only, wider than long, 0.5 x 0.6--0.8 mm.
Dense forests, epiphytic on rocks or tree trunks; 1200--2100 m. Gansu, Sichuan, Yunnan, [India (Darjeeling, Manipur, Assam), Nepal, Sikkim].
Note. In my paper Notes on Davalliaceae I (Blumea 37 (1992) 175) I reduced Paradavallodes kansuense Ching to Araiostegia pulchra on basis of the description. Later, however, I could study the type in PE and came to the conclusion that it belongs here.
8. Davallia pectinata J. Smith in Mém. Accad. Sci. Turin 5: 415. 1793.
马来阴石蕨 ma lai yin shi jue
Humata pectinata Desvaux; Davallia gaimardianum (Gaudichaud) C. Presl; D. multiflora Roxburgh; D. parallela Wallich ex Hooker; Humata gaimardiana John Smith; H. parallela (Wallich ex Hooker) Brackenridge; Nephrodium gaimardianum Gaudichaud; Pachypleuria pectinata C. Presl.
Rhizome without the scales 1.4--2.6 mm diam., white waxy under the scales. Scales red brown, with pale border from base to apex, narrowed evenly towards the apex, not or seldom curling backward, bearing multiseptate hairs at least when young, peltate, 5 x 1.1--1.5 mm. Stipes pale or dark brown, adaxially grooved, 5--18 cm long, glabrous or with few scales. Lamina simple, one pectinate or pinnatifid leaf, narrowly ovate, elongate, often narrowing towards base, bearing multicellular hairs or glabrous, 4--21 x 2.5--8 cm, not or slightly dimorphous. Longest pinnae 1.5--3.2 x 0.3--0.5 cm. False veins not present. Sori separate at the forking point of veins or at the bending point of a vein. Indusium attached at the broad base and hardly or not at the sides, semicircular, wider than long or about as wide as long, 0.6--0.8 x 0.6--1 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin.
Wet forests, epiphytic, epilithic, or sometimes terrestrial, on sand, lava, or limestone. Taiwan (Taitung) [Cambodia, India (S. Andaman, and Nicobar Is.), Indonesia, Malaysia, Papua New Guinea, Philippines, Thailand; Australia: (N. Queensland), Pacific Is (Truk Is., Solomons, New Hebrides, New Caledonia, Samoa, Carolines, Society Islands).
9. Davallia pulchra D. Don, Prodr. Fl. Nepal. 11. 1825.
美小膜盖蕨 mei xiao mo gai jue
Araiostegia athamantica Copeland; A. beddomei Ching; A. delavayi Ching; A. imbricata Ching; A. pseudocystopteris Copeland; Davallia athamantica Christ; D. beddomei Hope; D. pseudocystopteris Kunze; D. pulchra var. delavayi Beddome ex C.B. Clarke & Baker; D. pulchra var. pseudocystopteris Clarke; D. rigidula Baker (illegit. name); Humata yunnanensis Ching; Leucostegia delavayi Ching in C. Christensen; L. yunnanensis C. Christensen.
Rhizome without the scales 2--6 mm diam., not white waxy. Scales brown (often greyish), without pale border, broad, ovate to oblong-subdeltoid with round to acute apex, appressed to rhizome, usually crisped, margins recurved, not bearing multiseptate hairs, lacking marginal setae or teeth or those rare, basifixed with cordate base and much overlapping lobes, 2--5 mm long. Stipes pale, adaxially grooved, 10--20 cm long, glabrous or with few scales (sometimes with more scales). Lamina compound, tripinnate or quadripinnate, deltoid and broadest towards base, to elongate, often narrowing towards base, glabrous, 12--50 x 7--40 cm, not or slightly dimorphous. Longest petiolules 3--20 mm long. Pinnae deltoid or linear-triangular. Longest pinnae 5--21 x 3--12 cm. Pinnules of at least the larger pinnae anadromous, linear oblong or narrowly ovate. Longest pinnules 25--70 x 10--35 mm. Ultimate leaflets linear oblong, lobed almost to the midrib (each lobe bilobed again). Ultimate segments or lobes obtuse or acute without a tooth, 0.5--3 x 0.6--1 mm. Leaf axes glabrous. Veins in sterile ultimate lobes frequently simple, not reaching the margin. False veins not present. Sori separate, frequently single on a segment at the forking point of veins or at the bending point of a vein. Indusium reniform or semicircular, attached at the narrow, cordate base only, wider than long, 0.5--0.8 x 0.5--1 mm.
Wet forests, epiphytic and epilithic on granite and limestone; 450--3500 m. Guangxi, Sichuan, Guizhou, Yunnan, Xizang [Bhutan, India, Laos, Myanmar, Nepal, Sikkim, Sri Lanka, Thailand (north), Vietnam].
Note. Although the species is rather variable, especially because in different habitats the size may differ greatly, it is always recognisable by the broad, appressed, usually crisped rhizome scales.
10. Davallia repens Kuhn in Ann. Mus. Bot. Lugd.-Bat. 4: 286. 1869.
阴石蕨 yin shi jue
Adiantum repens Linnaeus fil., Suppl. Pl. 446. 1781; Davallia chrysanthemifolia Hayata; D. lepida C. [Presl, nomen] ex Goldmann; D. subalpina Hayata; D. vestita Blume; Humata chrysanthemifolia Hayata; H. lepida Moore; H. macrostegia Tagawa; H. pedata John Smith; H. repens Diels; H. trifoliata Cavanilles; H. vestita T. Moore; Pachypleuria lepida C. Presl; P. macrostegia (Tagawa) Kato; P. trifoliata C. Presl.
Rhizome without the scales 0.5--3 mm diam., white waxy under the scales. Scales brown or red brown, with pale border from base to apex or not, narrowed evenly towards the apex, not or seldom curling backward, bearing multiseptate hairs at least when young or, with marginal setae at least in distal part, peltate, 2.5--7 x 0.3--1.5 mm. Stipes adaxially grooved, 0.1--18 cm long, glabrous or with few scales. Lamina compound (pinnate with pinnatilobed to pinnatifid pinnae, or bipinnate to quadripinnate towards base and in the middle part), simple (one pectinate or pinnatifid leaf), 3-foliate (the leaflets more or less divided), or pinnate towards base, ovate, deltoid and broadest towards base, glabrous, 0.6--24 x 0.5--14 cm, strongly dimorphous or not or slightly dimorphous. Longest petiolules 0--4 mm long. Pinnae linear-triangular, narrowly ovate, linear, or ovate to deltoid. Longest pinnae 1--10 x 0.6--7 cm. Pinnules (if present) of at least the larger pinnae anadromous, linear oblong or narrowly ovate. Longest pinnules 5--55 x 5--20 mm. Ultimate leaflets (if present) lobed almost to the midrib or only shallowly lobed. Ultimate segments or lobes obtuse or acute without a tooth. In dimorphous plants lamina of fertile leaves pinnate with strongly dissected pinnae, bipinnate, or tripinnate towards base and in the middle part. Longest petiolules of fertile leaves 1--7 mm long. Pinnae deltoid, linear-triangular, or narrowly ovate, 1--8 x 0.3--2.5 cm. Pinnules or pinnalobes deltoid, or linear-oblong, 2--35 x 1.5--15 mm. Ultimate leaflets linear oblong. Ultimate segments of fertile leaves 1--15 x 0.5--2 mm. Leaf axes glabrous. Veins in sterile ultimate lobes simple, forked, or pinnate, reaching the margin. False veins not present. Sori separate, borne several on a segment, or in much divided leaves frequently single on a segment, at the forking point of veins. Indusium attached at the broad base and hardly or not at the sides, semicircular or more or less triangular to rhomboid, wider than long, about as wide as long, 0.3--1 x 0.3--1.3 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin. Lamina generally extending into a tooth at both sides or only at the outside of a sorus, or not extending into teeth beyond a sorus.
Low or high epiphytic, epilithic on various kinds of rocks, sometimes terrestrial, in very wet to dry sunny places; 0--3420 m. Fujian, Guangdong, Hongkong, Guangxi, Guizhou, Hainan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Cambodia, India, Indonesia, Japan, Malaysia, Myanmar (south), Philippines, Papua New Guinea, Sikkim, Sri Lanka, Thailand, Vietnam; Africa (Cameroun), Indian Ocean (Comores and Madagascar, Seychelles, Mascareignes, Reunion and Mauritius, Kerguelen, Bourbon Is.), Australia (Queensland) Pacific (Admiralty Is., Lashed is., Solomon Is., New Hebrides, New Caledonia, Fiji, and Samoa).
Note. This is a very variable species. All forms have in common the same spores which vary in size because of polyploidy
11. Davallia solida (G. Forst.) Swartz in J. Bot. (Schrader) 1800 87. 1801.
阔叶骨碎补 kuo ye gu sui bu
Trichomanes solidum G. Forster, Fl. Ins. Austr. 86. 1786; Davallia sinensis Ching; D. solida var. latifolia Hooker; D. solida var. ornata Mettenius ex Kuhn; D. solida var. sinensis Christ; D. subsolida Ching; Humata solida Desveaux; Stenolobus solidus C. Presl; S. ornatus C. Presl [Davallia ornata Wallich, nomen].
In China only var. solida.
Rhizome without the scales 4--14 mm diam., generally not white waxy. Scales red brown or nearly black (the peltate base black, persistent when the rest if the scales is shed) with pale border from base to apex, narrowed evenly towards the apex or above the much broader base evenly narrowed towards apex, not or seldom curling backward, bearing multiseptate hairs at least when young (hairs at least at apex of young scales, c. 1 mm long, woolly), peltate, 5--10 x 1--1.2 mm. Stipes pale, adaxially grooved, 9--35 cm long, glabrous or with few scales. Lamina compound, bipinnate or tripinnate towards base and in the middle part, deltoid and broadest towards base, glabrous (sometimes with hairs on junction of rhachis and petiolule), 15--90 x 21--40 cm, not or slightly dimorphous. Longest petiolules 5--25 mm long. Pinnae linear-triangular or narrowly ovate. Longest pinnae 11--28 x 6--15 cm. Pinnules of at least the larger pinnae anadromous, deltoid or rhomboid. Longest pinnules 40--100 x 15--80 mm. Ultimate leaflets linear oblong or rhomboid, lobed almost to the midrib, or only shallowly lobed (in bipinnate leaves the ultimate segments shallowly lobed). Ultimate segments 10--40 x 3--17 mm. Leaf axes glabrous (often hairs at junction of petiolules). Margins of the lamina of each leaflet not thickened. Veins in sterile ultimate lobes pinnate, reaching the margin or not. False veins not present. Sori separate, borne several on a segment at the forking point of veins. Indusium also attached along the sides, pouch-shaped, oblong, longer than wide, 1.2--2 x 0.5--1 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin. Lamina not extending into teeth beyond a sorus.
Epiphytic, epilithic on different kinds of rock, or terrestrial on different kinds of soil; as well in exposed places as in deep shadow, from open rocky places and savannahs to primary rainforest; 0--1500 m. Guangdong, Guangxi, Taiwan, Yunnan [Cambodia, India (Assam, Khasia, Andaman Is., Nicobar Is.), Indonesia, Malaysia, Myanmar, Philippines. Papua New Guinea, Sri Lanka, Thailand, Vietnam (Tonkin); Pacific (Bismarck Archipelago to the Santa Cruz Is., Samoa and the Society Is. and to the New Hebrides, New Caledonia, Fiji, and Tonga).
Note. Sometimes the leaf segments are very narrow and the plant resembles var. fejeensis. This is a widespread species which generally is easily recognisable from the black rhizome which bears hairy scales that soon fall off leaving only the bases.
12. Davallia trichomanoides Blume, Enum. Pl. Javae 238. 1828.
骨碎补 gu sui bu
Davallia bullata [Wallich] ex Hooker; Davallia mariesii T. Moore ex Baker; Davallia stenolepis Hayata; Davallia mariesii var. stenolepis Hoshizaki; Davallia petelottii Tardieu-Blot & C. Christensen; Trogostolon yunnanensis Ching; Davallia cylindrica Ching,
骨碎补 gu sui bu
In China only var. trichomanoides:
Rhizome without the scales 3--8 mm diam., not white waxy. Scales brown or red brown, with pale border from base to apex or not, flat and nearly acicular, narrowed abruptly from a broad base or above the much broader base evenly narrowed towards apex, often curling backward or appressed to rhizome, usually crisped, margins recurved, not bearing multiseptate hairs, with marginal setae at least in distal part or toothed, peltate, 4--8 x 1--1.5 mm. Stipes pale, adaxially grooved, 4.5--20 cm long, glabrous or with few scales. Lamina compound, tripinnate or quadripinnate towards base and in the middle part, deltoid and broadest towards base, glabrous, 10--35 x 9--25 cm, not or slightly dimorphous. Longest petiolules 1--6 mm long. Pinnae deltoid, longest 5--19 x 3--12 cm. Pinnules of at least the larger pinnae anadromous, narrowly ovate, longest 20--70 x 10--30 mm. Ultimate leaflets linear oblong or narrowly ovate, lobed almost to the midrib. Ultimate segments 5--27 x 2--6 mm. Leaf axes glabrous. Margins of the lamina of each leaflet not thickened. Veins in sterile ultimate lobes simple or forked, not reaching the margin. False veins present, rarely absent. Sori separate, frequently single on a segment at the forking point of veins. Indusium also attached along the sides, pouch-shaped, oblong, longer than wide, 1.2--2 x 0.5--1 mm, upper margin not elongated, truncate or slightly rounded, separated from or even with lamina margin. Lamina generally extending into a tooth at both sides of a sorus or only at the outside of a sorus.
Epiphytic and epilithic on different kinds of rock, mostly in wet places, sometimes on dry, exposed, places; 100--3500 m. Jiangsu, Liaoning, Fujian,Shandong, Taiwan, Yunnan [India (Kerala, Darjeeling, Assam and eastern Himalayas), Indonesia, Japan, Korea, Malaysia, Myanmar, Nepal, Papua New Guinea, Sikkim, Thailand, Vietnam.
Note. According to Hoshizaki (oral communication) several of the species treated here as synonyms behave as good species in culture. And, as I have seen in our botanical garden, they do indeed. However, after studying over 400 different collections of the entire area I could not but conclude that they all belong to one species. That does not exclude ,of course, that different forms from different localities intergrade in nature but behave different in culture. It would be best to give these forms cultivar names ('mariesii' and 'stenolepis'). As soon as a plant is cultivated and vegetatively propagated it forms a clone of similar plants that can be recognised from other clones of the same species. Formally naming the forms according to the rules of nomenclature means that quite a lot of collections can not be named. As the spores of all the forms are also extremely similar I have no doubt as to their conspecificity.
Acrophorus assamicus Beddome 7
Acrophorus immersus (Wallich ex C. Presl) T. Moore.......................................................................................................... 3
Acrophorus membranulosus (Wallich ex Hooker) T. Moore............................................................................................... 13
Adiantum denticulatum Burman 9
Adiantum repens Linnaeus fil 18
Araiostegia athamantica Copeland 17
Araiostegia beddomei Ching 17
Araiostegia Copeland 4
Araiostegia delavayi Ching 17
Araiostegia divaricata Kato 11
Araiostegia faberiana Ching 8
Araiostegia imbricata Ching 17
Araiostegia membranulosa Holttum 13
Araiostegia multidentata Copeland 15
Araiostegia parvipinnula Ching 8
Araiostegia perdurans Copeland 8
Araiostegia pseudocystopteris Copeland 17
Aspidium multidentatum Wallich, nomen 15
Davallia 1
Davallia amabilis Ching 11
Davallia assamica (Bedd.) Baker 7
Davallia athamantica Christ 17
Davallia austro-sinica Ching 11
Davallia beddomei Hope 17
Davallia bidentata Schkuhr 9
Davallia brevisora Ching 9
Davallia bullata [Wallich] ex Hooker 22
Davallia chaerophylloides Steudel 9
Davallia chrysanthemifolia Hayata 18
Davallia clarkei Baker 8
Davallia clarkei var. faberiana C. Christensen 8
Davallia cylindrica Ching 22
Davallia denticulata (Burman) Mettenius ex Kuhn 9
Davallia divaricata Blume 11
Davallia divaricata var. orientalis Tardieu-Blot & C. Christensen.................................................................................. 11
Davallia elegans Swartz 9
Davallia elegans var. bidentata Hooker 9
Davallia elegans var. coniifolia Hooker 9
Davallia elegans var. edentula Hooker 9
Davallia elegans var. polydactyla Moore 9
Davallia elegans var. pulchra Hooker 9
Davallia elegans var. subunidentata Hooker 9
Davallia formosana Hayata 11
Davallia gaimardianum (Gaudichaud) C. Presl 16
Davallia griffithiana Hooker 12
Davallia henryana Baker 12
Davallia immersa [Wallich, nomen nudum] ex Hooker........................................................................................................ 3
Davallia impressa Copeland 9
Davallia lepida C. [Presl, nomen] ex Goldmann 18
Davallia mariesii T. Moore ex Baker 22
Davallia mariesii var. stenolepis Hoshizaki 22
Davallia membranulosa [Wallich, Cat. (1828) 255, nomen], Hooker................................................................................... 13
Davallia micans Mettenius ex Baker 7
Davallia multidentata Hooker 15
Davallia multiflora Roxburgh 16
Davallia orientalis C. Christensen in Wu & al. 11
Davallia ornata Wallich, nomen 21
Davallia parallela Wallich ex Hooker 16
Davallia parvipinnula Hayata 8
Davallia patens Swartz 9
Davallia pectinata J. Smith 16
Davallia perdurans Christ 8
Davallia petelottii Tardieu-Blot & C. Christensen 22
Davallia platylepis Baker 12
Davallia pseudocystopteris Kunze 17
Davallia pulchra D. Don 17
Davallia pulchra var. delavayi Beddome ex C.B. Clarke & Baker..................................................................................... 17
Davallia pulchra var. pseudocystopteris Clarke 17
Davallia repens Kuhn 18
Davallia rigidula Baker (illegit. name) 17
Davallia Sect. Cordisquama Kato 4
Davallia sect. Davallia Kato 4
Davallia sect. Wibelia Kato 4
Davallia sinensis Ching 20
DAVALLIA Smith 3
Davallia solida (G. Forst.) Swartz 20
Davallia solida var. latifolia Hooker 20
Davallia solida var. ornata Mettenius ex Kuhn 20
Davallia solida var. sinensis Christ 21
Davallia stenolepis Hayata 22
Davallia subalpina Hayata 8, 18
Davallia subsolida Ching 21
Davallia trichomanoides Blume 22
Davallia tyermannii T.Moore 12
Davallia vestita Blume 18
Davallodes chingiae Ching 13
Davallodes membranulosum Copeland 13
Araiostegia hookeri Ching, (non Davallia Hookeriana Wallich 1828............................................................................. 8
hookeri parva C. Christensen 8
hookeri parvipinnula Hayata 8
hookeri perdurans C. Christensen 8
Humata assamica (Beddome) C. Christensen 7
Humata Cavanilles 4
Humata chaerophylla Mettenius 9
Humata chaerophylloides Desvaux 9
Humata chrysanthemifolia Hayata 18
Humata dryopteridifrons Hayata 3
Humata elegans Desvaux 9
Humata gaimardiana John Smith 16
Humata griffithiana C. Christensen 12
Humata griffithiana var. tyermannii (T. Moore) Tagawa................................................................................................... 12
Humata henryana Ching 12
Humata hookeri Diels 8
Humata immersa (Wallich ex C. Presl) Mett. 3
Humata lepida Moore 18
Humata macrostegia Tagawa 18
Humata micans (Mettenius ex Baker) Diels 7
Humata parallela (Wallich ex Hooker) Brackenridge 16
Humata patens Desvaux 9
Humata pectinata Desvaux 16
Humata pedata John Smith 18
Humata platylepis Ching 12
Humata repens Diels 19
Humata solida Desveaux 21
Humata trifoliata Cavanilles 19
Humata tyermanii T. Moore 12
Humata vestita T. Moore 19
Humata yunnanensis Ching 17
Leucostegia 1
LEUCOSTEGIA 2
Leucostegia assamica (Beddome) J. Smith 7
Leucostegia chaerophylla J.Smith 9
Leucostegia clarkei C. Christensen 8
Leucostegia clarkei var. faberiana C. Christensen 8
Leucostegia delavayi Ching in C. Christensen 17
Leucostegia faberiana Ching 8
Leucostegia griffithiana J. Smith 12
Leucostegia hookeri Beddome 8
Leucostegia immersa 2
Leucostegia immersa Wallich ex C. Presl 2
Leucostegia membranulosa (Wallich ex Hooker) J. Smith................................................................................................... 14
Leucostegia multidentata Beddome 15
Leucostegia pseudocystopteris Beddome 9
Leucostegia yunnanensis C. Christensen 17
Nephrodium gaimardianum Gaudichaud 16
Pachypleuria pectinata C. Presl 16
Pachypleuria trifoliata C. Presl 19
Pachypleuria. lepida C. Presl 19
Pachypleuria. macrostegia (Tagawa) Kato 19
Paradavallodes Ching 4
Paradavallodes chingiae Ching 14
Paradavallodes kansuense Ching 15
Paradavallodes membranulosum Ching 14
Paradavallodes multidentatum Ching 15
Parestia elegans C. Presl 9
Stenolobus ornatus C. Presl 21
Stenolobus solidus C. Presl 21
Trichomanes chaerophylloides Poiret 9
Trichomanes denticulatum Houttuyn 9
Trichomanes elegans (Swartz) Poiret (1808), not Richard (1792)....................................................................................... 9
Trichomanes lucidum Roxburgh 9
Trichomanes solidum G. Forster 20
Trogostolon Copeland 4
Trogostolon yunnanensis Ching 22
Wibelia Bernhardi (non Fee 1852); 4
*South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China; Correspondence to: XING Fu-Wu. E-mail:
**National Herabrium of the Netherlnads, Leiden branch. P.O. box 9514, 2300RA Leiden