An account of the fern genus Belvisia Mirbel (Polypodiaceae)

17 Pages·1993·2.2 MB·English

by  Hovenkamp, P H

BLUMEA 37 (1993) 511-527
An account of the fern genus Belvisia Mirbel
(Polypodiaceae)
P.H. Hovenkamp & N.A.P. Franken
Rijksherbarium/HortusBotanicus,Leiden,TheNetherlands
Summary
Thegenus Belvisia isrevised.Twospecies aretransferredfromLemmaphyllumtoBelvisia,and two
species arereduced tovarieties.The genusnowincludeseightspecies, reachingfrom tropicalAfrica
toChina,Polynesia,and Australia.
Mirbel(1803a: 65; 1803b:473) establishedthegenusBelvisia forthatpartofthe
LinneangenusAcrostichumthatis characterizedby linearreceptacles onbothsides
ofthemidrib,reaching tothelaminamargin,andopening towardsthelowersurface.
Mirbelincludedfivespecies inBelvisia, butdidnot makeany ofthenecessary new
combinations(1803c: 114-116). Thefivespecies are "remarkably unrelated"(Cope-
land, 1947) anda"meaningless jumble" (Morton, 1946),but amongthemwas Acro-
stichumspicatum L. f. Thismeleeofunnamedspecies was conveniently forgotten
foralmost acentury.
Inthecourse ofthatcentury,Kaulfuss(1824) newlyestablishedthegenusHyme-
nolepis, basedonAcrostichumspicatum, forwhichhesubstitutedthenameHymeno-
lepis ophioglossoides. Kaulfuss' generic circumscription was based mainly on the
linearsoriandthepresenceofscalyparaphyses. Inthissense, thegenuswas accepted
as distinctfromAcrostichumL. by most 19thcenturyauthors(Blume, 1828; Kunze,
1842;Presl, 1851;Fee, 1852;Moore, 1857;Kuhn, 1868;J.Smith, 1875). Dissenting
views were expressed by Mettenius(1869), wholinked theferns underdiscussion
with Taenitis, and Hooker(1863), who adhered to a widerview ofAcrostichum.
However, Diels(1899) andChristensen (1906) succeededinfirmly establishing the
genus asdistinct.
By then, several species hadbeen addedto Hymenolepis and one ofthesewas
again removed by Presl (1851) and usedtoestablish Macroplethus. Kunze (1850)
substitutedHyalolepis forHymenolepis; however, Hyalolepis hadbeenusedearlier
for a genusofthe Compositae. That thesame was true forHymenolepis was later
pointed out byUnderwood(1899) andBecherer(1936). Bechererthereforerevived
Mirbel's nameBelvisia, andlectotypified it byAcrostichum spicatum, this being
,
simply thespecies thatcamefirst inMirbel'saccount. Tagawa (1942)didnot accept
this lectotypification, andsubstitutedPresl's nameMacroplethus, based onHymeno-
lepis platyrhynchos Kunze. Becherer's arbitrary lectotypification was also disputed
by Morton(1946), whochoseAcrostichum septentrionale L. as type species, based
512 BLUMEA Vol. 37, No. 2, 1993
Fig. 1. Scales. —Belvisia spicata (L.f.) Copel.,a. outline,b. detail ofmargin(vanSteenis 18445).
——B. mucronata(Fée)Copel., c.outline,d. detailofmargin(Philipson 10429). Dotted line in
a&b =marginofoverlappingbasal extensions; scalebars = 1mm.
on the factthat this species is theonly one forwhich Mirbelcited an illustration.
However, Copeland, inhisinfluentialGeneraFilicum (1947) accepted A.spicatum
as typeofthe genus,whichhe couldaccordingly nameBelvisia. He was followed
by Holttum(1949) and PichiSermolli(1954), who settledthe argument inan ex-
haustive study ofall theavailablenamesand typespecies forthis genus. Sincethen,
consensus exists thatBelvisia Mirbel,typified byAcrostichumspicatumL. f.,isthe
propername forthis groupoffems.
VARIABILITY
Fde (1852) accepted 5species, Moore(1857) 4,and J. Smith(1875) 3.Christensen
atfirst (1906) accepted 4 species, butlater, in a monograph ofthe genus (1929),
partlybased on unpublished work by Hieronymus, he recognized 11 species and a
considerablenumberofvarietiesandformae.Atthesametimeheadmittedthatonly
two orthreecharacterswere"ofvalueas specific ones."Thiswas perhaps abitpes-
simistic. In thepresent account 8 species arerecognized, and2 subspecies. This is
accomplished withtheaidofthefollowingcharacters.
Rhizome
Therhizomeis short-creeping in mostspecies, withfrondsmoreorless crowded,
but it is long-creeping with well-spaced fronds inBelvisia novoguineensis and
P.H.Hovenkamp& N.A.P.Franken: An account of Belvisia 513
B. abbreviata.Thefrondsare inserteddorsally in two(rarely more)rows. Thestele
isaperforated dictyostele, anda variablenumberofsclerenchyma strandsispresent.
Scales
Therhizomescales are basally attached(Fig. 1),pseudopeltate withoverlapping
basalextensions, clathrate, andhaveanentireto ciliatemargin. Thefollowing scale
charactershave diagnostic value:
Outline—Inmostspecies, the scales haveanarrow, sometimesalmostfiliform
acumen. However,inB.spicata andB. validinervis theacumen isrelatively short
andwide. Theapexis usually acute, butmay beobtusetorounded.
Margin — Themargin ofthescales mayberatherirregularlyentire, ormore or
less dentatetodistinctly ciliate, withteeththatare formed by protruding thick cell
walls (Fig. lb). Independently fromthis marginal configuration, a variablenumber
ofmulticellularhairs,usually tippedwith agland, mayalsobepresent, mainly inthe
basalandextremely apicalregion. Theselatterhairshavelittle,ifany, diagnostic value.
Colour— On macroscopic inspection, the colour may vary betweenreddish-
brownandblack. A distinctiridescenceis presentinB. validinervis.
Cellstructure — Inmostspecies,allcellsare clathrate.InB.spicata andB. vali-
dinervis, however, thereis usually amoreor less distinctmarginal zonewithnon-
clathratecells (Fig. la). Inaless distinctform,such a marginal zonemaybepresent
inB.abbreviata.Theluminaeofthecentralcellsvaryin widthfrom10-30pmwide
inB. henryi to30-110pmwidein B.platyrhynchos andB. dura.
Fronds(Fig. 2)
The frondsare uniformlyentireand usually monomorphic, with a distinctstipe
and usually afertilespike attheapex. A light degree ofsterile/fertile dimorphism
has developed only inB. abbreviata.Thesterilepart shows hardlyany diagnostic
characters. Itis very variableinsize, with alength-width indexranging from 4to
75.Theprimary veinsare immersedorprominent in driedmaterial,the secondary
andtertiary veins form irregular areoleswithmanyfree veinlets, which are mainly
directedtowards the primary vein. Mature fronds are usually glabrous, but a few
scalesmaybefoundonthelaminaandthemidrib.A majordiagnostic characteris the
shape ofthetransitionbetween thebasal sterilepartofthefrond('lamina') and the
apical fertilepart('spike'). Inmostspecies, thelaminais very gradually narrowed
towards thespike, whichitselfis sharply distinct, oftendistinctly separated from the
laminaby a constriction(Fig. 2c, e-g). InB. annamensis(Fig. 2a) andespecially
B. henryi (Fig. 2b)thelaminais moreabruptly truncated,butnot separated fromthe
spike by aconstriction.Especially in B.annamensis thespike is not sharply distinct
fromthelamina, withthesori oftenextending downwardsintothe upper,narrowed,
part ofthelamina. InB. novoguineensis (Fig. 2d), lamina and spike are not at all
distinct, withavery gradual transitionfromthewide, sterilepartofthelaminatothe
narrower fertilepart.
Sori
Thesori are locatedinthe apical spike, a strongly narrowedpartofthefrond. A
long narrow spike (length-width indexup to 50-120), with sori fully covering the
514 BLUMEA Vol. 37,No. 2, 1993
cm.
3376).
1
=
7920). —
(Hennipman—(Beaman
BrassScabarsle— 10647).
Copel.
Raymond
f.)
(L.
spicata
(C.Chr.) (
Franken
B.
henryi
&
B.PNH3044).—e.Hovenkamp
398b—7)..
(Celestinolongissima
(Hennipman
Copel. var.
Copel.
Tagawa(Kunze)
(Kunze)
(C.Chr.)
platyrhynchos
validinervis
annamensis
B.
B.
g.
—153d5)..
Belvisia
a. Schodde LAE 61604).—
(
(Croft
Copel.
L)sp.ecimens —
Copel.
(Rosenst.)
(all
(Kunze)
outlines
novoguineensis
validinervis
Frond
2.
Fig.B. B.
c. f.
P.H.Hovenkamp& N.A.P.Franken: An account of Belvisia 515
Fig. 3.Paraphyses (all x 125).— a.Belvisia abbreviataHovenkamp& Franken(Gaudichaud s.n.). —
b. B. annamensis (C.Chr.) Tagawa(Geesink& Santisuk 4916).—c.B. mucronata (Fée)Copel.
(vanBorssum Waalkes 1958).— d.B. novoguineensis (Rosenstock) Copel. (Hoogland& Pullen
5587).—e.B. platyrhynchos (Kunze)Copel.(Clemens 16300). —f.B. spicata(L. f.)Copel.(Wild
6571).
entirewidthwhenripe ispresentinmost species. Widerspikes witha distinct sterile
marginal zone occurinB. annamensisandB. platyrhynchos. InB.annamensis the
sori aresituatedcentrally betweenmidriband margin,witha distinctsterilezoneusu-
ally presentbetweenthemidribandthesorus; inthetransitionalzonethesorusinthis
species tends to follow the margin (Fig. 2a). The short, widespike (length-width
index2-12)ofB.platyrhynchos (Fig. 2d) hasadistinctsterilemargin 4-5mmwide.
Paraphyses (Fig. 3)
Theparaphyses are sterile, scale-like structures thatoccur interspersed between
thesporangia. They are best searchedforin youngsori.In oldersori they tendtofall
off,andparticularly inB.annamensisandB. henryi paraphyses areoftencompletely
absentwhenthesporangia areripe. However,inyoungsori on unfurling frondsthe
paraphyses areoftennotfully developed. Thereis muchvariationinthe shape ofthe
516 BLUMEA Vol. 37, No. 2, 1993
paraphyses, and differentparaphyses often occur in the same sorus. They consist
ofastalkwithalaterally orpeltately affixedblade, whichmaybeentireorvariously
incised.Like the scales, the bladesoftheparaphyses are clathrate.Smallbladesin
whichthewallsofthecentralcellsare so stronglythickenedthattheluminaarealmost
entirely filledare characteristic forB. annamensis(Fig. 3b) andB. henryi. Other
species usually havelargerblades,in whichthecentralcellshaveopenlumina.Para-
physes withpeltately andlaterallyaffixedbladesoftenoccurinthesamesorus inmost
species except B.annamensisandB. henryi, whichhave only peltate paraphyses.
Theregular occurrence ofirregularly branchedscale-likestructures (Fig. 3f)is char-
acteristic forB.spicataandB. validinervis.
Spores
ThesporesofBelvisiahavebeen treatedseparately by VanUffelen(1992).
SYSTEMATIC POSITION
Inthepresentcircumscription, Belvisiaincludestwospecies not includedby Chris-
tensen (1929), bothofwhichheincludedinLemmaphyllum (B.abbreviata,B.novo-
guineensis). For Christensen, theposition ofthesespecies was highly ambiguous,
and he expressed his doubts whetherany ofthe two involved generawas really
monophyletic. Whenwe nowplace thetwo species inBelvisiathis isbasedonthe
observationthatthe thenremaining species ofLemmaphyllum (L. accedensandL.
carnosum s.l.) bothhavestrictly peltate scales, whereasinBelvisiaall species now
includedhave basifixedscales withoverlapping auricles.Thischaracter, not seenby
Christensen, notonly separatesthe generareadily, butalsoarguesagainst a deriva-
tionofBelvisiafromLemmaphyllum. Following Hovenkamp (1986), we consider
pseudopeltate scales tobe intermediatein thetransformationofsimple hairstopeltate
scales. Underthatassumption, itis notlikely thata genuswithpseudopeltate scales
isderivedfromaspecies withpeltate scales. Thereare, inour opinion, noarguments
eitherto favourtheopposite, a derivationofLemmaphyllum fromBelvisia. Infact,
BelvisiaandLemmaphyllum sharefew, ifany, characterswitheachotherthatarenot
sharedalsowithLepisorus. Onlythecoenosorimightbeusedtounite thetwo genera,
butthe plethora offorms intermediatebetween separateand fusedsori inLemma-
phyllum carnosum in ouropinion clearly showsthatthecoenosorus inthat species is
not inheritedfromasingle coenosoroidancestor. Itis anindependent development,
possibly arrived atindependently severaltimeswithinLemmaphyllum. Thatcoeno-
sori arereadily formedindependently inunrelatedgroupshasbeenmadesufficiently
clear(Christensen, 1929;Ravensberg & Hennipman, 1986).
References cited above:
Becherer,A. 1936.Conservation d'homonymesg6n6riquesdans les fougferes.Candollea7: 137-139.
Blume,C.L. 1828.Enumeratio Plantarum Javae. II.Filices.
Christensen,C. 1906.IndexFilicum.
Christensen, C. 1929.Taxonomicfernstudies.I. Dansk Bot.Ark.6 (3): 54-73.
Copeland,E.B.1947.GeneraFilicum.
Diels, L. 1899.In:Engler&Prantl,Die natiirlichenPflanzenfamilien1,4.Polypodiaceae.
F6e, A.L.A. 1852.Genera Filicum. 5me Mem.
P.H.Hovenkamp& N.A.P.Franken: An account ofBelvisia 517
Holttum,R.E. 1949.The selection oftype-speciesofsome old genera offerns.Gard. Bull. Singa-
pore 12: 303-306.
Hooker,W.J. 1863. SpeciesFilicum V.
Hovenkamp,P. 1986.A monographofthe fern genusPyrrosia (Polypodiaceae).Leiden Bot.Series
9: 1-280.
Kaulfuss,G.F. 1824.EnumeratioFilicum.
Kuhn,M. 1868.Filices Africanae.
Kunze,G. 1842.Farmkrauter.
Kunze,G. 1850.Index Filicum.Linnaea 23.
Mettenius,G. 1869.In:Miquel,Ann. Mus. Bot.Lugd.-Bat.4.
Mirbel,C. 1803a. Histoirenaturelle g6neraleetparticulicre desplantes. 4.
Mirbel,C. 1803b. Histoirenaturelle des veg6taux3.
Mirbel,C. 1803c.Histoire naturelle desvegetaux 5.
Moore,T. 1857.Index Filicum.
Morton,C.V. 1946. The fern studies ofMirbel. J.Wash. Acad. Sc. 36: 166-169.
Pichi Sermolli,R.E.G. 1953.The nomenclature ofsomefem genera. Webbia 9:405-409.
Presl,C.B. 1851.EpimeliaeBotanicae.
Ravensberg,W.J. & E.Hennipman. 1986.ThePyrrosia species formerlyreferred toDrymoglos-
sum andSaxiglossum. Leiden Bot.Series 9: 281-310.
Smith,J. 1875. Historia Filicum.
Tagawa, M. 1942.Hymenolepis Klf.should be substituted by MacroplethusPr. Acta Phytotax.
Geobot. 11: 232-235.
Uffelen,G.A.van. 1992.Sporogenesis inPolypodiaceae(Filicales). II.The genera Microgramma
Presland Belvisia Mirbel. Blumea 36: 515-540.
Underwood,L.M. 1899.A review ofthe fernsproposedprior to 1832.Mem.TorreyBot.Club 6:
247-283.
BELVISIA
Belvisia Mirbel,Hist. Nat. Gen. 4 (1803)65;Hist. Nat. Veg.3 (1803)473;ibid. 5 (1803) 111;
L.M.Underw.,Mem.Torrey Bot.Club 6 (1899)276;Copel., Gen. Fil. (1947) 191;PichiSer-
molli,Webbia 9(1953) 387; Holttum,Fem Fl.Mai. (1954) 153,f.67 & 68; Copel., FemFl.
Philipp. 3 (1961)466;Pichi Sermolli,Webbia 31 (1977) 324. —Lectotypespecies: Belvisia
spicata (L.f.)Copel.
HymenolepisKaulf.,Enum. Fil. (1824) 146,pi. i: f.9, nom.illeg., non Cassini (1817); Blume,
Enum.PI.Jav.(1828)200;Endl.,Gen. PI.(1836)62; Presl,Epim. Bot.(1851)158;Fee,M6m.
Fougeres 5(1852) 81,pi. 6B; T. Moore, Ind.Fil. (1857)28,pi. 15A; Bedd.,Ferns S. India
(1863) 15,pi.26; J. Sm.,Ferns Br.& For. (1866) 91, pi. 19; HistFil. (1875) 118; Diels in
Engler& Prantl, Nat. Pflanzenfam. 1,4(1902) 305, f. 161F-H; Copel., Dept.Int. Bur.Gov.
28(1905) 110;Alderw.,Mai.Ferns (1908)727; Domin,Bibl. Bot. 85 (1915)168;Alderw.,Mai.
Ferns,Suppl.(1917)432;Goebel, Ann. Jard. BotBuitenzorg36 (1926) 108,pi.7, f. 34-43;
Bower,Ferns 3(1928)222,f.731;C.Chr.,DanskBot.Ark. 6(1929) 54,f. 1;Backer& Posth.,
Varenfl.Java (1939)228,f.53;C.Chr.&Tardieu,Notul.Syst 8(1939)185;Tardieu& C.Chr.,
Fl.Gen. Indochine 7(2) (1941)451,f. 53: 1-2; Tardieu,Mem.Inst. Fr.Afr.Noire 28 (1953)
207;Alston, FernsW.Trop.Afr.,Suppl.(1959)48.—Typespecies: Hymenolepisophioglos-
soides Kaulf. [=Belvisia spicata(L.f.) Copel.].
HyalolepisKunze,Linnaea 23 (1850)258,nom. illeg., non DC. (1837).—Typespecies: Hyalo-
lepisophioglossoides(Kaulf.)Kunze[=Belvisiaspicata(L.f.) Copel.].
MacroplethusPresl,Epim. Bot.(1851) 141;Tagawa, ActaPhytotax. Geobot. 11 (1942)232.—
Type species: Macroplethusplatyrhynchus (Kunze)Presl [=Belvisia platyrhynchos (Kunze)
Copel.].
Note - Species ofBelvisia havebeen namedinAcrostichum,Gymnopteris,Lomaria, Onoclea,
Schizaea,andTaenitis.
518 BLUMEA Vol. 37, No. 2, 1993
Epiphytic, rhizomeshort- tolong-creeping, approximately terete,coveredwithscales,
dorsally with2rowsoffronds, ventrally andlaterallydensely setwithroots, dictyo-
stelic, withscattered sclerenchyma strands.Rhizomescalesbasifixed, pseudopeltate,
ovateto linear-lanceolate,fully clathrateor withamembranaceousmargin,reddish
tobrown,entireto dentate, witha varying numberofglandularcellsonthemargin.
Fronds simple, entire.Stipes articulatedtotherhizome, basally terete,upwards ad-
axially grooved, sometimeswinged. Laminalinear-lanceolateto linear,base cuneate,
apex gradually narrowedorabruptly truncate tothenarrow fertilepart,olivaceousto
brownwhendry, dull, pergamentaceous,thinly covered withscattered, mostly ca-
ducousscales. Veins anastomosing, forming areoles withmany scatteredfreevein-
lets, freeveinletsin thecostaland marginal areoles all andin theotherareoles pre-
dominandy directedto thecosta; midribdistinctthroughout thesterileandthefertile
partofthelamina, secondary veins distinctorindistinct, immersed, tertiary veinsand
veinletsimmersed. Fertile partlinear-lanceolateto linear, withoneelongated, some-
timesinterrupted, sorus ateachsideofthemidribusually fully covering thelamina
between midriband margin, sometimesleaving anarrow zonefreealong themidrib.
Sporangia with 12-16induratedannuluscells, mixedwithpeltate or basally attached
paraphyses. Spores monolete,rugulate.
Distribution- TropicalAfricaeasttotropical AsiaandPolynesia, extending to South
China, Tahiti,Rapa andAustralia.
Note- Inthefollowing key, thetermlaminarefers tothesterilepartofthelamina
only, thetermspike referstothefertile apical part.
KEY TO THE SPECIES
la. Rhizomelong-creeping; frondsscatteredatintervalsofupto7 cm 2
b. Rhizomeshort-creeping; frondstuftedorspaced atintervalsofatmost2cm 3
2a. Sorishort, upto 1 cm long, onashortspike whichis nearly always sharply dis-
tinct from thelamina 1. B. abbreviata
b. Sorimuchlonger, onalonger spikewhich gradually mergesintotheupperpart
ofthe lamina 6. B. novoguineensis
3a. Paraphyses peltate only (often completely absentin oldsori), usually opaque
withluminaofcentralcellsalmostentirely filledwith wallmaterial;laminausu-
ally asymmetrically truncate belowthespike 4
b. Someparaphyses withlaterally affixedblades nearly always present, usually
translucent; laminausually gradually and symmetrically narrowedbelow the
spike 5
4a. Spike 4-7 mmwide; sorimedialbetween midriband margin
2. B. annamensis
b. Spike up to 4mmwide; sori close to themidrib 3. B. henryi
5a. Rhizomescales withdentatetociliatemargin, allcellsthick-walled 6
b. Rhizomescales entireoralmost so,centralcellsthick-walled, marginal cellsthin-
walled 7
6a. Sorileaving aclearmarginaland sometimesalso anarrow costalzonefree;spike
relatively wide, to 2 cm wide 7. B. platyrhynchos
P.H.Hovenkamp & N.A.P.Franken: An account of Belvisia 519
b. Soricompletely covering thespike; spike usually narrow, linear,0.1-0.2(rare-
ly up to 1) cm wide 4. B. mucronata
7a. Rhizomeshort; leavestufted; scalesdullwitha dark-browncentre andalighter
margin; lamina generally 1-2 cm wide 8. B. spicata
b. Rhizomemore widely creeping; fronds scattered at intervals ofup to2 cm;
scales uniformly lightbrown,glossy oriridescent; laminagenerally 3-4(some-
times upto 6) cm wide 9. B. validinervis
1. Belvisia abbreviata (Fée) Hovenkamp & Franken, comb. nov. - Fig. 3a
Drymoglossum abbreviatum F6e,Mem.Fougeres 5 (1852)26, t. 10:f. 2; C.Chr.,Ind.Fil. (1906)
246. — Hymenolepisabbreviatum F6e,Mem.Fougeres 10(1865) 12;C.Chr.,Ind.Fil. Suppl.
3 (1934) 113. —Polypodium abbreviatum C.Chr.,Ind. Fil. Suppl.2(1917)25,nom. illeg.,
n—onP.abbreviatum Mett.,Fil. Hon.Lips. (1856)33(=Pyrrosia). Lemmaphyllumabbrevia-
tumC.Chr.,Dansk Bot.Aik. 6(1929)50. —Macroplethusabbreviatus Tagawa,ActaPhytotax.
Geobot. 11 (1942)234;Pichi Sermolli,Ind.Fil.Suppl. 4(1965)185.—Type:Gaudichaud s.n.,
1839,VoyagedelaBonite,Cochinchina (P;iso B,G).
Polypodium hymenolepioidesChrist, J. de Bot. 19 (1905) 75; C.Chr.,Ind. Fil. (1906) 534. —
HymenolepishymenolepioidesChingex C.Chr.& Tardieu,Notul. Syst. 8 (1939) 185;Tardieu
& C.Chr.,Fl.G6n. Indochine 7 (2) (1941)452. —BelvisiahymenolepioidesChing,ActaPhy-
totax.Sin. 10(1965)301. —Type:Cadiere 97,Annam,Quang-Binh(P).
Rhizomevery slender,long-creeping, intemodes1-5cm long. Rhizomescales ovate
toovate-lanceolate, 1.5-2.7by0.7-1.3mm, index 1.2-4,marginentireor minute-
ly dentate,apex rounded; reddish-brown,wallsofcentralcells more strongly thick-
enedthanthoseofmarginal cells.Frondssomewhatdimorphic. Stipes to 1 cm long,
0.3-0.7mm thick, laminaofsterilefrondselliptic to linear-lanceolate,to 5 by 1cm,
offertilefronds mostly longer and sometimes narrower, to 12 by 0.5-1.5 cm to
linear, 4-10 by 0.3-0.8 cm, index 5-33, gradually narrowedto the base, often
slightly constrictedbelow thespike; spike ovate toovate-lanceolate, 0.2-1 by 0.2-
0.4 cm. Sori medialbetween midriband margin,often interrupted (best visible in
young sori), rarely a row ofseparateround soripresent, not covering marginand
costa whenripe. Paraphyses peltate, bladesflat, 0.4-0.45 mmwide, entire, with
thick-walled cells.Spores rugulate, 45-70by25-50pm, index 1.4-1.8.
Distribution- Vietnam.Annam: Cadiire97(P),Poilane 7939(P). Cochinchina:
Gaudichauds.n. (P).
Habitat—Epiphytic or epilithic;altitude425m(Poilane7939).
Note- Obviously arare species. Its systematic positionremains ambiguous. Itis
here includedinBelvisiaon accountofthecharacteristicsofthescales oftherhizome
andthe(rarely present)paraphyses.
2. Belvisia annamensis (C.Chr.) Tagawa - Figs. 2a, 3b
Belvisia annamensis Tagawa,ActaPhytotax. Geobot. 22 (1967) 107.—Hymenolepisannamensis
C.Chr., Dansk Bot. Ark. 6 (1929)68, f. le; C.Chr. & Tardieu,Notul. Syst. 8 (1939) 185;
Tardieu &C.Chr.,Fl.G6n. Indochine 7(2)(1941)452,f.35: 1,2.—Macroplethusannamen-
sis Tagawa,ActaPhytotax. Geobot. 11 (1942)234. —Type:Poilane 8010,Annam, Col des
nuages(P).
520 BLUMEA Vol. 37, No. 2, 1993
Gymnopterisspicata(L.f.) Bedd. var. BlatifronsBedd.,Handb.,Suppl.(1892)104.—Hymenolepis
spicata (L.f.) Presl var. latifrons Alderw.,Mai. Fems (1908) 729. —Syntypes: King 1100,
Perak(BO; isoM);Day s.n.,Perak(notseen).
Hymenolepiscallifolia Christ var.paltonioidesC.Chr.,DanskBot.Ark. 6 (1929)67. —Syntypes:
Haviland s.n.,Sarawak,Baram (K);Macleay s.n.,Borneo (K,not seen).
Rhizomeshort-creeping, intemodesnot elongated, 2-4mm thick.Rhizomescales
ovate-lanceolateto linear-lanceolate, 4.8-8.3by 0.9-2.1 mm,index 3.8-5.6, base
dentate,acumen narrowed,dentatenear the base, sometimes with a narrow entire
filiformapex,reddishbrownto black, allcellwalls strongly thickened. Stipes 1-4
cmlong, 1-2 mm thick, laminalinear-lanceolate, 15-30by 2-5 cm, index 6-9,
base gradually narrowed,apexmoreorless abruptly truncate; spike 5-25 by 0.4-
0.7 cm, index 8-60. Sorioften extending downwards to well below the narrowed
spike, medialbetweenmidriband margin, leaving amedialand marginal freezone,
paraphyses peltate, withcircularblade,0.1-0.4mmdiam.,brownish toblack, mar-
gin toothed, cell walls strongly thickenedinmature paraphyses. Spores rugulate,
40-60 by 25-40 pm.
Distribution- Thailand,Laos,Vietnam, Borneo.
Habitat- Epiphytic oronrocks inevergreenforest.Sealevelto 1900m.
Note- Theonly cleardifferencesbetweenthisspecies andB. henryi arethewidth
ofthespike andtherelativepositionofthesori.Thetwo species arereasonably well-
separated geographically. However,it may wellbe that whenmore materialfrom
theircontact zone becomesavailable, thetaxonomic statusofthesetwo species will
havetobereconsidered.
THAILAND. MtGoping: King 1100 (BO, M). MtKhao: Hennipman3745,3987 (L).Ranong:
Geesink & Santisuk4916 (L).
LAOS.Poilane 12210(P).
VIETNAM. Annam: Cadiere 11 (BM,P), 787(P); Poilane 8010, 11001 (P). —Tonkin: Petelot
4477(P).
BORNEO.Haviland & Hoses.n„Aug. 1894(K).
3. Belvisia henryi (C.Chr.) Raymond Fig. 2b
-
BelvisiahenryiRaymond, Mem. Jard. BoL Montreal 55 (1962)32;Panigrahi& Patnaik,Curr. Sci.
34 (1965) 127;Tagawa,Fl.E. Himalaya(1966)490. —Hymenolepis henryiC.Chr.,Dansk
Bot. Ark.6 (1929) 67,f. Id; C.Chr.& Tardieu,Notul. Syst. 8(1939) 185;Tardieu & C.Chr.,
Fl.Gen. Indochine 7 (2) (1941)451.—Macroplethus henryiTagawa,ActaPhytotax. Geobot.
11(1942)234. —Type:Henry 11461a,Yunnan (B,isoUS,W).
Rhizomeshort-creeping, intemodesnotelongated, 2-4 mmthick.Rhizome scales
ovate-oblong toovate-lanceolate, 2.7-5.2 by 0.8-1.4 mm, index 2.7-1.5, above
thebase constrictedtoalong acumen, basedentate, acumen narrowed,dentatenear
thebase, sometimeswith anentirefiliformapex, cellswithstrongly thickenedwalls.
Stipes to 4cm long, 1-2mm thick, laminalinear-lanceolate, 7-27 by 1.5-5cm,
index 4—9, base gradually narrowed, apex usually abruptly truncate (sometimes
asymmetrically) tothespike, sometimesnarrowedmore gradually, spike linear,5-35
by 0.1-0.4cm, index 15-115.Sori usually completely covering thelamina,para-
physes peltate, with circularblade,0.1-0.3 mm diam., margin toothed, cell walls
strongly thickenedinmature paraphyses. Spores rugulate, 40-75by 25-50pm.