Table Of ContentBLUMEA 37 (1992) 165-187
Notes on Davalliaceae I. The genera Araiostegia,
Davallodes, Leucostegia, and Gymnogrammitis
Hans+P. Nooteboom
Rijksherbarium/HortusBotanicus,Leiden,TheNetherlands
Summary
Arevision is givenofthe followinggenera: Araiostegiawith 4species, Davallodes with 7species,
Leucostegiawith 2species, andGymnogrammitiswith 1 species. The delimitation ofgeneraisac-
cordingtoKato(1985).
Araiostegia
AraiostegiaCopel., Philipp. J.Sci. 34 (1927)240, t. 1,2; Univ.Calif. Publ. Bot. 12 (1931)397,
t. 53a; Gen. Fil. (1947) 85; Holttum,Revis. Fl.Malaya2, sec. ed.(1966) 364; Copel.,Fem
Fl. Philipp. (1958) 167; Ching, Fl. Reipubl. Popul. Sin. 2 (1959) 285; Kato, J. Fac. Sci.
Univ.Tokyo, sect. 3 Bot., 13(1985)564. —Type:A. hymenophylloides(Blume)Copel.
Paradavallodes Ching,ActaPhytotax. Sin. 11 (1966)18.—Type:P.multidentatum Ching.
Rhizomebearing only scales. Rootsrestrictedtotheventralside oflateralbuds. Scales
withoutpale border, narrowedevenly towardstheapex, or broad, ovate tooblong-
subdeltoidwith roundto acuteapex, not bearing multiseptate hairs,with feworno
marginal setae,toothedorentire, smoothon theadaxial surface, basifixed with cor-
date,muchoverlapping base.Extra-axillary lateralbuds lateraltothephyllopodia, or
lowerandslightlyanterior. Stipe articulatedatthebaseonphyllopodia, pale ordark
brownwhendry,adaxially grooved, glabrous orwithfewscales. Laminacompound,
tripinnate orquadripinnate,deltoidandbroadesttowardsbaseor elongate, butlower
pinnae notvery small,aboutaslong as thelongestpinnae, bearing multicellularhairs
(A. multidentata)orglabrous. Laminanotorslightly dimorphous. Hairsbetweenveins
onbothsurfaces absentor nearly so. Pinnulesof atleastthe larger pinnae anadro-
mous. Leafaxes glabrous or atleastrachises hairy (only inA. multidentata). Veins
inultimatelobes simple orforked, notreaching themargin. False veinsnotpresent.
Soriindusiate, separate,frequently single onasegment,facing midveinsat thefork-
ing pointoftheveins, or facingmidveinsattheirbending point. Indusiumscale-like,
eitherattachedonly atthenarrow cordatebaseoralong abroadbase.
Distribution Ceylon, India, Nepal, Sikkim, Bhutan, Burma, Thailand, South
-
China; onespecies in Malesia.
166 BLUMEA Vol. 37, No. 1, 1992
Fig. 1. Indusia (SEM); scale bar = 1 mm. — a.Davallodes novoguineense(Rosenstock) Copel.
(Croxall & Parris 5768).—b.Araiostegiaclarkei Copel.(Ludlow elal. 16863);c.idem (Hennip-
man3406);d.idem (Y.F. Chen 2471).—e.Araiostegiapulchra(D.Don)Copel.(G. Mann,Aug.
1889); f.idem (MurataT15110).
H.P.Nooteboom: Notes on Davalliaceae I 167
Fig. 2. AraiostegiaclarkeiCopel. (M.T.Kao 4193).
168 BLUMEA Vol. 37, No. 1, 1992
Fig.3.AraiostegiaclarkeiCopel.(Polunin elat. 186).
H.P. Nooteboom: Notes on Davalliaceae I 169
Fig. 4.Araiostegiaclarkei Copel.(Hennipman3406).
170 BLUMEA Vol. 37, No. 1, 1992
KEY TO THE SPECIES
la. Scales broad, ovateto oblong-subdeltoid, withroundto acuteapex
4. A. pulchra
b. Scales evenly narrowedtowardstheacute apex 2
2a. Pinnae sessile 1. A. clarkei
b. Longest petiolules 2.5to 30mm 3
3a. Laminadeltoid,broadest atbase; leafaxes, atleastrachises, hairy
3. A. multidentata
b. Laminaelongate,not broadestatbase; leafaxes glabrous
2. A. hymenophylloides
ALTERNATIVE KEY TO THE SPECIES
la. Leafaxes, atleastrachises, hairy 3. A. multidentata
b. Leafaxes glabrous 2
2a. Pinnae sessile 1. A. clarkei
b. Longest petiolules 2.5to30mmlong 3
3a. Scales often curling backward, narrowed evenly towards apex, stipes dark
brown 2. A. hymenophylloides
b. Scales appressed, usually crisped, margins recurved, broad, ovate to oblong-
subdeltoidwithround to acuteapex, stipes pale 4. A. pulchra
1. Araiostegia clarkei (Baker) Copel. — Figs, lb-d, 2-4
AraiostegiaclarkeiCopel.,Philipp.J.Sci. 34 (1927)241. —[Acrophorushookeri Moore,Ind.Fil.
(1857)2, nom.nud.] —Davallia clarkeiBaker, Syn.Fil.(1874) 91.—Leucostegia hookeri
Bedd.,Handb. Ferns Brit.India (1883)32.—Humata hookeri Diels inE. &P.,Nat. Pflanzen-
fam. 1,4 (1899)209. —Leucostegia clarkeiC.Chr., Contr. U.S. Natl. Herb. 26 (1931) 294.
—Araiostegia hookeri Ching,Fl. Reipubl. Popul. Sin. 2 (1959)291; Panigrahi& Basu, J.
Econ. Tax.Bot. 5(1984) 845. —Type:Hookerf.& Thomson 315 (BM;iso K), India,Sirmur.
Davallia perduransChrist,Bull.Herb. Boissier6 (1898)970.—HumataperduransHieron.,Hedwigia
62 (1920) 12.—Leucostegia perduransC.Chr., Contr. U.S.Natl. Herb. 26 (1931)294. —
AraiostegiaperduransCopel.,Univ.Calif.Publ. Bot.12(1931)397.—Syntypes:Henry10086&
A +B(BM,Ain K), China,Yunnan.
Davallia parvipinnulaHayata,Mat.Fl.Formosa(1911)431.—LeucostegiaparvipinnulaHayata,
Icon.PI.Formos.4(1914)205,f. 139.—AraiostegiaparvipinnulaChing,Fl.Reipubl.Popul.
Sin. (1959) 292. —Type:Kawakami & Mori 1823(n.v.), in monteMorrison.
DavalliasubalpinaHayata,Mat.Fl.Formosa (1911)432. —Type:G.Nakahara,Nov. 1906 (n.v.),
Arizan.
DavalliaclarkeiBakervar.faberianaC.Chr.,ActaHorti Gothob. 1(1924)73.—Leucostegiaclarkei
var. faberianaC.Chr.,Contr. U.S.Natl. Herb.26 (1931)194.—LeucostegiafaberianaChing
in C.Chr.,Index Filic. Suppl. 3(1934) 120.—AraiostegiafaberianaChing,Fl.Reipubl.Popul.
Sin. 2(1959)293;Tagawa& Iwatsuki,ActaPhytotax. Geobot. 24 (1970) 180.—Type: Faber
1089 (W; isoK,P), Szechuan,Omei summit.
Araiostegiaparva Copel.,Univ.Calif.Publ. Bot. 12(1931) 399.—Leucostegiaparva C.Chr.,In-
dexFilic. Suppl.3 (1934) 121. —Type:Hooker(UC, n.v.), Sikkim.
Araiostegia hopeiPanigrahi&Basu,[J.Econ. Taxon.Bot. 5(1984)849,nom.nud.;]op. cit.6 (1985)
470. —Type:H.C.Levinges.n. (CAL,n.v.), Darjeeling,Sundukpho.
H.P.Nooteboom: Notes on Davalliaceae I 171
Rhizomewithoutthescales 5mmdiam.(with scales c. 10mm);scales lightbrown,
evenly narrowedtowardstheacuteapex,7-10mmlong. Stipespalewhen dry, 7-35
cm long. Laminatripinnate orquadripinnate,deltoidandbroadesttowardsbase, gla-
brous, 10-50cm long,6-50cm broad.Longest pinnae 2-30cmlong, 1.2-18 cm
broad. Bothlowestpinnules of atleastbasalpinnae insertedonpinna base(pinnae
sessile, sometimesonly one pinnule atbaseofpinna), otherpinnules anadromous.
Longest pinnules or pinnalobes 6-130mmlong,2-50mmbroad.Ultimateleaflets
linearoblong, lobedalmostto themidrib.Ultimatesegments 1-4mm long, 0.2-1
mm broad. Sori facing midveins atthe forking point ofthe veins. Indusium scale-
like, attachedatthenarrow,cordatebase only,orattachedatthebroadbaseandhard-
ly ornot along thesides, reniformor semicircular, widerthanlong orabout aswide
as long,0.3-1.2 mm long, 0.4-1.5 mmbroad.
Distribution NW India2coll., Bhutan4coll., Nepal6coll., Burma 1 coll., N
-
Thailand(Doi Inthanonand ChiengRai) 6 coll., China(Tibet 8 coll., Yunnan many
coll., Sichuan2coll., Gueizow1 coll.,Hongkong, Taiwan manycoll.).
Habitat&Ecology - Generally epiphytic,indenseevergreenormossy forest.Alti-
tude 1200-4250m.
Note- Araiostegia clarkeiis a veryvariablespecies,but mostofthevariability is
in thedimensions.
2. Araiostegia hymenophylloides (Blume) Copel. — Fig. 5
Araiostegia hymenophylloides (Blume)Copel.,Philipp. J. Sci. 34 (1927)241; Copel., Fem Fl.
Philipp. (1958)166. —Aspidium hymenophylloides Blume,Enum. PI.Javae (1828) 172.—
Leucostegia hymenophylloidesBedd.,Ferns S.India (1863)t.252; Hook.,Syn.Fil. (1868)92;
Bedd.,Suppl. Ferns S.India(1876)4;Handb. FernsBrit.India(1883) 54.—Davallia hymeno-
phylloidesKuhn,Ann. Mus.Bot.LugdunoBatavum 4(1869) 286;Christ,Bull. Herb. Boissier
6(1898)142.—Humata hymenophylloidesCopel.,Publ.Bur. Sci.Gov. Lab.Philipp. 28 (1905)
51.—Type:Blumes.n. (L,909.30-141),Java,MtBurangrang.
[Leucostegiaaffinis John Sm.,J.Bot. 3 (1841)416,nom.nud.]—Davallia affinis Hook.,Sp.Fil.
(1846) 158,t. 52B;Kunze,Bot.Zeitung(Berlin) 6 (1850) 236;Harrington,J.Linn.Soc. Bot.
16(1877)26.—Microlepiaaffinis Presl,Epim.Bot.(1851)97.—Acrophorusaffinis Moore,
Proc.Linn.Soc. London 2 (1854)286;IndexFil.(1857) 1.—Humataaffinis Mett.,Fil. Hort.
Bot.Lips. (1856) 102,t. 27 f. 5,6. —Type: Cuming 117(iso BM,K,L,P),Philippines.
Microlepia tenuifoliaPresl,Epim.Bot.(1851) 97.—Cuming215 (iso BM,K,L,PNH), Luzon.
Cystopterisdalhousiana Fde,M6m.Foug. 8(1857)108.—Type:LadyDalhousie (G-DEL;iso K),
Penang(acc. toFee Ceylan!).
Rhizome without the scales 3-20 mm diam.; scales lightbrown, membraneous,
evenly narrowedtowards the acute apex, often curledbackwards, 4-7 mm long.
Stipes darkbrown,9-45(-65) cm long. Laminatripinnate, elongate, oftennarrow-
ed towards base, glabrous (or nearly so), 20-80 cm long, 6-50(-90) cm broad.
Longest petiolules 2.5-30 mm long. Longest pinnae 4-30(-44) cm long, 1.5-15
(-18) cm broad. Longest pinnules or pinnalobes 10-80mm long, 5-20mm broad.
Ultimatesegmentslinear-oblong. Sorifacing midveinsatthebending pointofa vein.
Indusiumscale-like, attachedatthenarrow cordatebaseonly, reniform, widerthan
long, 0.1-0.4 mm long, 0.4-0.7 mm broad.
172 BLUMEA Vol. 37, No. 1, 1992
Fig. 5.Araiostegiahymenophylloides(Blume)Copel.(Ueda&Darnaedi B8740).
H.P.Nooteboom: Notes on Davalliaceae I 173
Distribution- Ceylon manycoll.,SIndia(Western Ghats7coll.);Malesia:Suma-
tra7 coll., MalayPeninsula, Java,andBorneo manycoll.,Philippines (Luzon many
coll., Mindanao7coll., Mindoro2coll., Biliran 1coll.).
Habitat&Ecology - Epiphytic or epilithic, rarely terrestrial, usually in shaded
places. Altitude500-2200m.
3. Araiostegia multidentata(Hook.) Copel.
Araiostegia multidentata (Hook.)Copel.,Philipp.J.Sci. 34 (1927)241;Ching,Fl.Reipubl.Popul.
Sin.2(1959)295. —[Aspidium multidentatumWall.,Cat. (1828)346, nom.nud.]—Daval-
lia multidentataHook, inHook. &Baker,Syn.Fil. (1868)91;C.B.Clarke,Rev.FemsN. India
(1880)442.—Leucostegia multidentata Bedd.,Ferns Brit.India (1869) t. 313 (n.v.); Handb.
FemsBrit. India(1888) 50.—Humata multidentataDielsin E.& P.,Nat. Pflanzenfam. 1, 4
(1899)209. —Paradavallodes multidentatum Ching, ActaPhytotax. Sin. 11 (1966) 20.—
Type;Wallich 346 (K; iso BM),Nepal, 1821.
MicrolepiapteropusBedd.,Fems Brit.India (1869)t.313,n.v., acc. toChing(1959).
[Acrophorusthomsonii Moore,Ind. Fil. (1857)4,nom.nud.: Hook.f. & Thomson 316].
Rhizomewithoutthescales 5 mmdiam.(with scalesc. 10mm); scaleslight brown,
evenly narrowedtowards theacute apex,6mmlong. Stipes whendry pale or dark
brown, 17-25 cm long. Laminatripinnate, deltoidand broadesttowardsbase, gla-
brous, 30-45 cm long, 17-34cm broad. Longest petiolules 8-10mmlong. Long-
estpinnae 10-19cm long, 6-9 cm broad.Longest pinnules or pinnalobes 40-70
mmlong, 15-30mmbroad.Ultimatepinnules linearoblong, lobedhalfway towards
midrib.Ultimatesegments2-4mmlong, 1 mmbroad, oftenshallowly lobed.Hairs
onleafaxes 0.1-0.2 mmlong. Sori facing midveins at the forking point or atthe
bending point. Indusiumscale-like, attachedat thenarrow, cordatebaseonly, reni-
form,widerthan long,0.5 mmlong,0.6-0.8 mm broad.
Distribution- N India(Manipur 1 coll., WBengal, Assam5 coll.), Nepal 1 coll.,
Thailand(Prachunburi 1 coll.),China(Yunnan, Shweli SalwinDivide 3coll.).
4. Araiostegia pulchra (D.Don) Copel. — Figs, 1e, f, 6
Araiostegia pulchra(D.Don)Copel.,Philipp. J. Sci. 34 (1927) 241;Ching, Fl.Reipubl. Popul.
Sin. 2 (1959) 288; Tagawa & Iwatsuki,ActaPhytotax. Geobot. 24(1970) 180.—Davallia
pulchraD.Don, Prodr. Fl.Nepal.(1825) 11; C.B. Clarke,Rev. Ferns N. India (1880)444;
Baker,Kew Bull. 1895 (1895)53.—LeucostegiapulchraJohn Sm., London J. Bot. 1 (1842)
426;Bedd.,Handb. Ferns Brit. India (1883)52.— Davallia chaerophyllaWall. [Cat. (1828)
259,nom.nud.] exPresl,Tent. Pterid. (1836) 129;Hook., Sp.Fil. (1846) 157, t. 51 A;Fee,
Mem.Foug.5,Gen. Filic. (1852)329. —LeucostegiachaerophyllaJohn Sm.,London J.Bot.
1 (1842)426.—Acrophoruschaerophyllus Moore,Proc.Linn. Soc. London 2(1854)286. —
Humata chaerophyllaMett.,Fil. Hort.Bot.Lips. (1856) 102,t. 27 f.9, 10.—Acrophoruspul-
cherMoore,IndexFil. (1857)3;Bedd.,FernsS.India(1863)t. 10.—Humata pulchraDielsin
E. & P.,Nat. Pflanzenfam. 1,4(1899) 209. —Type:Wallich 259(K; iso BM,L,P), Nepal,
1821.
Cystopterisdimidiata Decne. in Jacquem.,Voy.Inde 4 (1844)178,t. 178.—Type:Jacquemont(P).
Cystopterissquamata Decne. in Jacquem.,Voy. Inde 4 (1844) 178,t. 178.—Type:Jacquemont(P).
174 BLUMEA Vol. 37, No. 1, 1992
Fig.6.Araiostegiapulchra(D. Don)Copel.(Tagawaetal.T9478).