582.547.

15

MEDEDELINGEN LANDBOUWHOGESCHOOL WAGENINGEN NEDERLAND 79-14(1979)

A REVISION OF ANUBIAS SCHOTT (ARACEAE)

( P R I M I T I A E A F R I C A N A E XII)

W. CRUSIO Laboratory ofPlant TaxonomyandPlant Geography, Agricultural University, Wageningen, The Netherlands
Received 28-VI-1979 Date ofpublication22-X-1979

H.VEENMAN &ZONEN B.V.- WAGENINGEN - 1979

CONTENTS

page INTRODUCTION HISTORY OF THE GENUS DISTRIBUTION AND ECOLOGY CULTIVATION GENUS DIAGNOSIS KEY TO THE SPECIES DESCRIPTIONS Anubias afzeliiSCHOTT
barteri SCHOTT

1 ' ' ,
2

^
..

var. barteri
var. angustifolia (ENGLER) CRUSIO

.,
., ,9
21 24 2g '
3)

var. caladiifolia ENGLER var.glabra N. E. BROWN

var. nana(ENGLER) CRUSIO gigantea CHEVALIERexHUTCHINSON gilletiiDE WILDEMANet DURAND gracilis CHEVALIERexHUTCHINSON

hastifolia ENGLER heterophylla ENGLER pynaertii DE WILDEMAN

35 43

. . . 46 ACKNOWLEDGEMENTS 47 LITERATURE 48 INDEX

INTRODUCTION The present revision of the genus AnubiasSCHOTT is based on a study of herbarium materials and literature, supplemented bytheinformation obtained from specimens cultivated at the Laboratory of Plant Taxonomy and Plant Geography of the Agricultural University at Wageningen. ThefollowingHerbariasuppliedspecimens;theDirectorsandKeeperskindly sent them on loan. B(Berlin),BM(London),BR(Brussels),COI(Coimbra),E (Edinburgh), G (Geneva), GRO (Groningen), HBG (Hamburg), K (Kew),L (Leyden), LE (Leningrad), M (Munich),P (Paris),S(Stockholm), UPS(Uppsala),WAG (Wageningen),Z(Zurich). HISTORY OF THE GENUS Anubiasisconfined towesternTropicalAfrica, SenegaltoAngolaandZaire. In 1857 SCHOTT established the genus Anubias,basing this on one species. AnubiasafzeliiSCHOTT.Inthecourseoftime, 18 speciesintotalweredescribed. In 1968 HEPPER removed Anubias hastifolia ENGLER and Anubias aunculata ENGLER from Anubiasto Amauriella RENDLE.Thislattergenushadbeenestablished in 1913,based on onenew species:Amauriellaobanensis RENDLE. Inthe present revision, Anubiasand Amauriella areunited. Only8spec.esin Anubias are recognised. D I S T R I B U T I O N AND ECOLOGY Anubias is most frequent in a comparatively "0\Z0^Xc^Z tropical African coast but can befound asfar asBamakoinM a i . ^ m Central Zaire.A. barteriisthemostwidelydistributedspecies, r o d ^ ^

Congo.A.afzelii, A.gigantea, andA.gracilisare ^ t ^ ^ ^ ^ h

ofWest Tropical Africa, whilethe other specieso c c u r J ^ J Anubias occurs inforests inwet,shadyplaces,mostlyonthebanks courses. Sometimes they growcompletely submerged. CULTIVATION u c Unwingsoecieswerecultivated: A. During the course of this revision the,fdow. g ^ . . ^ ^ . ( a va_ afzelii, A. gigantea, A. heterophylla, A. giUetu J> h can begr0wn rieties).Onlywhenkeptinaveryhumid^ a ^ b I ^ o r a b o v e 2TC successfully. The temperature of s 0 1 ' a m i x t u r e o fsand, clayandhumus. (-30C). Theplantswereplacedinpots ma m + ^ ^ ^ ^ Theplants Thepotswereplacedinbuckets,whichcoma _^ ^ ^ ^ ^ n Q t e v e n were kept inthe greenhouse, shaded,anu during winter. 1
Meded. Landbouwhogeschool Wageningen 79-14(1979)

A. barteriand A. heterophylla are sometimes kept by aquarists submerged in indoor tanks. Some of them report a satisfactory growth (ANSINK, 1977: 68, dealing with A. heterophylla; SADILEK, 1974: 129; 1978: 202; 1978: 238, all dealing with A. barteri; MOHLMANN, 1977:485,dealing with A. barteri).

GENUS DIAGNOSIS AnubiasSCHOTT

SCHOTT, 1857: 398; 1858: 42; 1860: 159; ENGLER, 1879: 433; BENTHAM & HOOKER, 1883:975; ENGLER, 1893:462; BAILLON, 1895: 50;BROWN, 1901: 182; ENGLER, 1915: 2; LEMEE, 1929: 188.

Type species: A. afzeliiSCHOTT(monotypic). Heterotypic synonym: Amauriella RENDLE, 1913:115;ENGLER, 1915:1; LEMEE, 1929: 188; HEPPER, 1968a: 454. Type species: Amauriella obanensis RENDLE(monotypic). Note: Amauriellaobanensis RENDLEisAnubiashastifolia ENGLER. Perennialherbs. Rhizomeprostrate,creeping and rooting. Top ofthe rhizome densely leaved, or leavesall overthe rhizome. Leavespetiolate. Leaf-blades very variable in form and size, from narrowly oblong-lanceolate with an obtuse or acute base,towidelyellipticwith a hastate base(sometimesnearlytripartiteinA.gigantea).Numerousparallelnerves,with many slender veinlets transversally between them. Above glabrous, below glabrous or more or less densely puberulous on the midrib and primary veins. Coriaceous or subchartaceous. Spathe oblong, oblong-elliptic or ovate, closed or open when flowering and then sometimes spreading backwards. After flowering closing again and persistent. Spadixcylindrical, thick or slender, more or lessdensely covered with naked unisexual flowers. <J flowers at the top, ? flowers at the base of the spadix. Between them sometimes sterile<Jflowersor rudimentary bisexual flowers. S flowers with 3-8(9) stamens. Stamens connate, united in an obpyramidal synandrium flowerssometimesdeformed, havingmorethantheusualnumber otstamens(2^flowersbecomingmoreorlessconnate).Thecaelateral,onedge, oronthetopofthesynandrium(inA.pynaertii coveringthesynandriumalmost entirely). Thecae oblong or ovate, opening by a longitudinal slit. $ flowers creamy white. Mowers withadiscoid stigma. Stigma on a short styleor (sub)sessile. Ovary U^-J-locular, depressed globose-ovoid. Numerous anatropous cylindrical ovules. Ovaries green. Stigma green, pink orwhite. 2
Meded. Landbouwhogeschool Wageningen 79-14(1979)

D. DEWIT, calidarioWAG,3.V.1978- {Crusio8, WAG). Meded. Landbouwhogeschool Wageningen 79-14(1979)

Berriesenclosedbythespathe,depressed-globose,2-3-locular, many-seeded. Seeds small,irregularly ovoid. Distribution: 8speciesinWest-Africa, mostly in the rainforest belt.

KEY TO T H E SPECIES la. Thecae only on thesideor on thetop ofthe synandrium 2 lb. Thecae covering the synandrium completely on the sides and (sometimes onlypartly)onthetop.Whenfloweringspatheopendowntothebaseand the spadix exserted 8. A. pynaertii 2a. Thecae on the side ofthe synandrium 3 2b. Thecae onthe top or at the edge ofthe synandrium 7 3a. Stamens (6)7-8(9); (thecae 14-16). Leaf-blades triangular-cordate in outline.Spathe up to 3cmlong 5. A. gracilis 3b. Stamens 36(8); (thecae6-12). Leaf-blades not asabove 4 4a. Spathe open down to thebasewhen flowering, but near the insertion with imbricate margins 5 4b. Spathe openingup tohalfway down during flowering 6 5a. Spathe oblong-ovate or oblong-elliptic, never reflexed, and more or less fleshy.Sometimesspadixalittleexserted.SpatheuptoA\cm long;almost twiceto4 | times aslong asbroad 7. A. heterophylla 5b. Spatheoblongtooblong-lanceolate, reflexed whenflowering, not fleshy, up to 6cmlong; more than 4x aslong asbroad 2. A. barteri 6a. Leaf-blade lanceolate, mostly longer than thepetiole . . . .1. A. afzelii 6b. Leaf-blade hastately lobed, mostly shorter than thepetiole. 3. A. gigantea 7a. Whenfloweringspatheopenattheupperpart only,upto4^cmlong.Thecae onthetop or at theedge ofthe synandrium, never on the sides of the synandrium 6. A. hastifolia 7b. When flowering spathe open down to thebase,at thevery base convolute, up to 3cm long. Thecae at the edge of the synandrium, ovate-elliptic. Spathe reflexed 4. A. gilletii (Notice: In A. heterophylla the thecae are sometimes only on the upper half of the side of the synandrium. Especially when dried, they may resemblethethecae ofA. gilletii strongly.) Notes: 1. For a correct determination adult flowering specimens are necessary. 2.Theposition ofthethecae on the synandrium isbest seenduring anthesis. 3.Sometimesdeformed$ flowersoccur,2neighbouringflowersbeingmoreor lessconnate, which leads to the presence of more stamens than usual. On the sameinflorescence however,normal flowers can alwaysbe found.

Meded. Landbouwhogeschool Wageningen 79-14(1979)

DESCRIPTIONS 1. Anubias afzelii SCHOTT Fig.1,Phot.2,Map1.

SCHOTT, 1857: 399; 1858: pi. 42; 1860: 159; ENGLER, 1879: 434; BROWN, 1901: 183; ENGLER, 1915: 3; HUTCHINSON & DALZIEL, 1936: 365; BERHAUT,

1967: 316;HEPPER, 1968b:120. Type: Afzelius s.n., Sierra Leone (UPS,holotype;isotypeBM). Rhizome prostrate, creeping and rooting, 1-4 cm thick. At the top densely leaved. Petiole up to 20cm long, sheathing till overthemiddle,at theend provided with an 1-1^ cm long geniculum, half to almost as long as the leaf-blade, glabrous. In cultivated specimens the petiole sometimes as long as or even slightly longer than the leaf-blade (upto29cmlong,sheath 8cmlong). Leaf-blade elongate-lanceolate to oblong-elliptic, 13-35 cm long, 3-13 cm broad, 2.5-8 x as long as broad, broadest about the middle, more or less narrowed at the base, both sides equal, coriaceous. Base subtruncate to cuneate. Top mucronulate orwith anacute oracuminate tip. Manyprominent lateralnerveswith5-8 lessprominentlateralnervesbetweenthem,nearthe edge anastomosing in 2-3 marginal nervesandwithverynumeroustransverse sender veinlets between them. Glabrous or sometimes below very sparsely peatncle(9-)13- 32cmlong,halftooneandahalftimesaslongas the petiole. Spathe3-7(-9)cmlong,oblongorlanceolate,apiculate, ^ o t a ^ n a n o v g tubular,green.Not spreadingatmaturity,butatthe topopened.Beforeanthe openinghalfway down,butnotreflexed(insectsmay haveaccessto the $ flowers) and closing tightly and entirely again after afewhours Spadix 5-8(-12) cmlong,veryshortlystipedorsessile,1.2-1.5 x z longa t h e U c up(per part exserted, S ^ ^ 2 ^ ^ ^ inflorescence almosttwiceaslongasthe?part.Sometimelowe 6 or less abortive (abortive part up to * cmlong -The o ^ J r f synandrium.Stamens 5-6. Ovariesdepressed-g ose gre.n, n o p lob very short or the discoid pink stigmasessile.S e e d ^ ^ ' J t ^ N o ivingseeds
VUh

broad(1.3-1.7 x

* * ^ > f f ^ ^ ^ L l
F

were observed.) Berries not observed. (Only oneirunnig completely eaten berries seen: Thomas2829.) D i s t r i b u t i o n : Senegal,Guinea, SierraLeone,Mali.

* Wvnlaces sometimescompletely submerged.

N,.:,_

nowenrfa,D-SL

" C * " * ^

Meded. Landbouwhogeschool Wageningen 79-14(1979)

FIG 1. Anubiasafzelii SCHOTT: 1. upper side leaf, jx ; 2. inflorescence, | x ; 3. synandrium, n '\ MUPP"Slf,e,s j; nand m> 20x ; 5.anther,front view,20x ; 6.part oftherhizome,f x . - (1-5. Morion&GledhillSL 1169, WAG; 6.J.V. 327,spirit coll.WAG). Meded. Landbouwhogeschool Wageningen 79-14(1979)

MJ**

~~"~ ' if. u/ith sliehtly opened spatne; ngnr. nii PHOT. 2. ,4m<ftto fl//ii SCHOTT - Inflorescence: left. witft " ' ^ j T j x I 9 7 g . . (Crusio 19. tightly closed spathe. Phot. J. W. MUGGE, calidar.oWAG,4.X.1978 and 5.X.1 WAG).

atKantora,iSster^^ Senegal, 2nd ed.) records A. afzelu for S e f g a ^ o ( M a l i ) d e n s e groupsofca stream,onthebank,inredlatent*m u d : ^ ^ . orest remnant. 1mtall specimens growing inthemargin of a gallery Specimens examined:
SENEGAL:Kantora, Mt.Casamance(st.)Elesse75(P). 12741 (P); Ma chexaUer Gu, NEA: Fouta Djaion (fl- Apr) Che.aUer ^ ^ ^ . ^ ^ D e k ^ m i i m Diaguissa and Ditinn(buds,fr.) Chevalier '&>"<?>'" Delesser, 17871(G); R>v. Katendeg. KindiaS.(ft. May)Z>efeer/ 77*(G); B e n * E o ^ 9 7 p a ) y V i m b o ( f r . , f l . A P r . ) M ^ / ^ i ( P ) . June) J D^ 0 w^ > (P);Rotouma(fr.,fl.July)^ W M W O y i n ' A/erferf. Landbouwhogeschool Wageningen 79-14(1979)

MAP 1. Anubiasafzelii.

MALI: Bamako(fl.) Chevalier44056(P);ibid,(fr.) Chevalier44086(P). SIERRALEONE:s.loc.(fr.)Afzeliuss.n.(UPS, holotype;BM,isotype);nearBatema(fr. May)Barter s.n. Nigerexp. (K);Baiima(Gbo),(fl. May)Deighton5767(K.);s.loc. (st.) Don s.n. (BM); s.loc. (fl. Apr.) Johnson732(K); No. 2River,Peninsula(fl. Apr.)Morton &GledhillSL1169 (WAG);Bafodeya (fr. Apr.) ScottElliot5560 (K);Jigaya (fr. Sept.) Thomas2829 (K). UNKNOWN LOCALITY:(fl.) Bogner 1298(WAG);(st.) collectorunknown (BM). CULTIVATED: Belgium,atGent(fl. Oct.)Boom21163 (L); Denmark,atCopenhagen(fl.)Jacobsen s.n. (WAG);Germany, at Munich (fl. Feb.) Bogner 1304 (WAG); Great Britain, at Kew(fr. May) Kew11-5-1912(K); ibid,(fl.) Kew entrynr. 64544(K); Netherlands,atWageningen(fl.Oct.)Crusio 19 (WAG);ibid.(fl.Oct.)J.V. 327 (WAG);ibid.(fl. May)/. V. 512(WAG);ibid.(fl.June) De Wit s.n. (WAG);ibid.(fl. Sept.)De Wit7788 (WAG).

2. Anubiasbarteri SCHOTT

Figs.2,3,Phot.1,3,4,Maps 2-6

SCHOTT, 1860: 159; ENGLER, 1879: 435; 1893: 463; BROWN, 1901: 185; ENGLER, 1915:5; HUTCHINSON& DALZIEL, 1936:365;HEPPER, 1968b:120.

Type: FernandoPo,Barter2045 (K,holotype). Rhizomecreeping,prostrate and rooting,0.2-1 cmthick. Petioleuptoover\ mlong,\-2\ x aslongastheleaf-blade. Sheath short or uptof ofthelengthofthepetiole.Geniculumupto 2cmlong,sometimesnearly visible. Leaf-bladeveryvariable,seebelow.More orlesscoriaceous. Peduncle\-A x aslongasthepetiole,upto over40cmlong.
Meded. Landbouwhogeschool Wageningen 79-14 (1979)


FIG. 2. Anubiasbarteri SCHOTTvar. bartervA- ^ "f inflorescence, f x ; 4.inflorescence, f x ; 5.upper s.desy view,20x; 7.part oftherhizome,f x. (1 m* ' spirit coll.WAG; 3-6. Bogner 1303, spintcoll.WAG,

A I M M X - 2.lower sideleaf, f x ; 3. ^ ^ . i , 20 x ; 6.synandrium side ^^ ^ R^ /i0i, Cr5w/0,spiritcoll.WAG), ?

Merferf. Landbouwhogeschool Wageningen 79-14(1979)

Spathe l | - 6 cm long, not fleshy, oblong-lanceolate, acute or acuminate, white,green,yellowish green orpurple, spreading and reflexed when flowering, laterclosingagain,4-16 x aslong as broad. Spadixcylindrical,slender,\-2 x aslongasthespathe, \\-l cmlong,$ part 1-6 x aslongasthe?part.Stamens4-6.Thecaeonthesideofthe synandrium, ovate or oblong-elliptic, $ and ? parts more or less densely flowered. Ovaries depressedglobose.Stigmadiscoid,(sub)sessileornot.Ovariesandstigmagreen. Berries depressed-globose. Seeds\ to up to 1mm long and 0.4-0.8 mm broad, 1.3-2.2 x aslongasbroad.Driedseedslightyellow;withoutanyothercoloured spots. Distribution: Guinea, Liberia, Ivory Coast, Nigeria, Fernando Po, Cameroon, Gabon, Congo. Eco1ogy:Inforests onshadyplaces,alongwater-coursesinsand,onrocksor onoldwood.Emersed ortemporarily submerged.Upto 1600m.Flowering and fruiting throughout the year. Note: Theleaf-blades varywidelyinsizeand shape.Although intermediates arenot rare,A. bartericanbemoreorlessarbitrarily divided into fivevarieties, which haveto someextent adifferent area ofdistribution and remain constant whencultivated.However,thedifferences betweenthetaxaareoflowtaxonomicalimportance, and donot merit the rank of subspecies.

KEY TOTHEVARIETIES

la. Leaf-basesagittate,sinusratherwide.Midribandmainveinspalmate(main veinsdirected downwards and passing intothe lobes) c.var. caladiifolia lb. Leaf-base otherwise 2 2a. Leaf-bladeupto2 x aslongasbroad.If2-2^ x aslongasbroad,thanleafbase obtuse-truncate or more or lesscordate 3 2b. Leaf-blade more than 2\ x aslong as broad. If 2-2^ x as long as broad, than leaf-base cuneate-obtuse .4 3a. Leaf-blade up to 6cmlong and 3cmbroad. Petiole up to 5cmlong. Leafblade longer than the petiole e.var.nana 3b. Leaf-blade longer than 7cm,4-11cmbroad. Petiole morethan 6cmlong. Mostly leaf-blade shorter than thepetiole a.var.barteri 4a. Leaf-blade morethan 5 x aslong asbroad, up to 3 | cmbroad . . . b.var.angustifolia 4b. Leaf-blade lessthan 5 x aslong asbroad, usually broader than 3^cm . . d.var.glabra Note: For acorrect determination adult leaves are necessary.
Meded. Landbouwhogeschool Wageningen 79-14(1979)

A 1af 2x 2 var. glabraN. E. BROWN, upper side leaf, f x ; 3 var. ( ^ ^ B ; angustifolia(ENGLER) CRUSIO,uppersidesteal,, Boger 1295, WAG; 4.5oger /J05,WAU). A/erferf. Landbouwhogeschool Wageningen 79-14(1979) 2.Bogner1302.WAG. 3. j,

a. var.barteri

Fig. 2,Map2.

Petiolealmostequallingtoupto 1^ x aslongastheleaf-blade, 6-23 cmlong. Leaf-bladeovate-lanceolate,acuminate-acute, sometimesapiculate,7-23 cm long,4-11cmbroad,uptoalmost2\ x as longas broad,usuallylessthan 2 x as long as broad, widest below the middle, glabrous or puberulous below, base truncate or more or lesscordate. Many prominent lateral nerves,with 3-6 less prominentlateralnervesbetweenthem,whichsometimesunitebeforetheyreach the margin. At the edge anastomosing in 2-3 marginal nerves and with many slender, transverse veinlets between them. Peduncle slightly longer orup to 4x aslong asthepetiole,up to 30cmlong. Spadixequallingorupto 1^x aslongasthespathe.S part 1.0-1.7 x aslong asthe? part. Distribution: S.E. Nigeria, Cameroon, Fernando Po. Specimens examined:
SENIGERIA: Prov. Ogoja,Distr.Ikom, Loc.BritishObokum (fr. May)FHI18880 (K); Calabar Province, Oban Forest Reserve,Orem(fl.Jan.) FHI 36037(K). CAMEROON:Env.ofLolodorf,Bikelegi(fl.)^ne?47P(P);Batanga(fl.,fr.)Bates289(BM); Efulen (fl. July)Bates289 (K);20kmSEof Mt. Elephant, 20kmSEofKribi(fr. Aug.)Bos 7251 (WAG); Moliki (fr. Nov.) Jacques-Felix 2510(P); km 30 Kumba-Victoria Road, S of Bombe (fr. Nov.) Leeuwenberg 10615 (WAG); between Fenda (60 km ESE of Kribi) and Kienke River (fl., fr.) Letouzey 4120 (P);Youngue,closetoNdom,40km SofNdikimeki(st.Dec.) Letouzey 10732(P); Kopongo-NgambeRd., 15 kmNNE ofEdea (fl., fr. Jan.) Letouzey 11034(BR,K, P);closeto the falls of Songlanon, 25 km SW of Ngambe (st. Jan.) Letouzey 11101(P); between Babong and Ogurong, 45 km SW of Mamfe (fr.) Letouzey 13581 (P); South Cameroon, 7.05 N, 3.46 E (st.) Mildbraed5725(HBG;partly:see alsovar. glabra);SouthCameroon,7.27N,3.47E(fl.)Mildbraed

10
12

MAP 2. Anubiasbarterivar.barteri

593Ibis (HBG; partly: see also var. glabra); Kumba river, NE of Barombi(fr.) Preuss 559 (K); SouthernCameroon,Barombi-Kang,Kumbas.l.(fr.June)SCA 40(E); Lolodorf(fr.)Staudt354(S; partly:seealsovar.caladiifolia);ca40 kmSofBadjob,ca50 km ofEseka(fl.Dec.) De Wildec.s. 1555 (WAG; partly: see also var.glabra); Bipinde, Mimfia (fr.) Zenker s.n. (S,P);Bipinde(fl.)Zenker 2853bis (M;partly: seealsovar. glabra). FERNANDO PO: S.loc.(fl.) Barter2045 (K,holotype). UNKNOWN LOCALITY:(fl.) Bogner 1301 (WAG);(fl.) Bogner 1303 (WAG). CULTIVATED: Netherlands, at Wageningen (fl. Apr.) Crush 1(WAG; cultivated plants from Bogner 1303);ibid. (fl. Sept.) Crusio 10 (WAG);ibid.(fl.June)J.V. 569 (WAG).

b. var.angustifolia (ENGLER) CRUSIO, stat.etcomb. nov. Fig.3,Phot.1,3,Map3 H o m o t y p i c synonym: A. lanceolata f. angustifolia ENGLER, 1915: 4. Type: Cameroon,Dusens.n.(S, neotype). Petiolei o r up to nearly aslongastheleaf-blade, 4-32cmlong. Leaf-bladenarrowlyoblong-lanceolateornarrowlyoblong-elliptic 5-y x as long as broad, up to 3J cm broad, 8-18 cm long, widest about the middle glabrous, obtuse or acute-apiculate, numerous more or lessprominentlateral nerves,withsometimes 1or 2lessprominentlateralnervesbetween hem which anastomosebefore themargin. Lateralnervesanastomosingin 2.<n-3^marginal nerves.Numerous slender,transverseveinletsbetweenthela eralnerves Peduncle equalling or upto overtwiceaslongasthepetiole,9-38cmlong. Spadix i or up to 1.3 x aslongasthespathe.S part 1-6 x aslongasthe? part.

MAP 3. AnubLarteri

var.angustifolia(alsofound inCameroon,, loc.).

13

Meded. Landbouwhogeschool Wageningen 79-14(1979)

PHOT. 3. Anubiasbarteri SCHOTT var.angustifolia (ENGLER) CRUSIO Inflorescence. Phot. H. C. D. DE WIT, calidario WAG, 3.V.1978. (Crush 8,WAG).

Distribution: Guinea, Liberia,Ivory Coast, Cameroon. Notes: 1.The two syntypes on which ENGLER based his description of A. lanceolata forma angustifolia {Dusen 87 and Schlechter12412,both from Cameroon) weremostprobably destroyed at Bduring the war. A neotype ishere designated. 2. ENGLER(1915: 4)described thisvariety as a forma of A. lanceolata N. E. BROWN. Thisspeciesisherereduced to avariety of A. barteri(seevar.glabra), whiletheforma angustifoliaisalsogivenvarietalstatusinA. barteri.Cultivated specimensproved to beconstant astothe shapeand sizeofleaves. Specimens examined:
Mfr^v^11^^^^111" S(P)
(fl (bU d> fl M a r ) Chemlier

'

12769

<P>;M t - Nzo > P ^ s desGuerzes (bud

Par0"Je69(P>'S-lTC-(flApr.)^e//7/i9(P);Mt.NimbUt.June)5c/,,//

L.BERIA:Nimbaexpedition(st., drawingofinfl.)Adam20296(K,UPS);ibid.(stbudMar.)Adam

14

Meded. Landbouwhogeschool Wageningen 79-14(1979)

21071(K,UPS); Jekepa, Nimba, Jiti River (fl.May) Adam27701 (BR, WAG);Jekepa, NimbaGangra (fr. May)Adam 27706(BR). IVORY COAST: Pays des Dyolas: Pied desMonts Nouba: Env.des sourcesduNuon(fr. Apr.) Chevalier21114(?). CAMEROON:s.loc.(fl.)Bogner 1309 (WAG);s. loc.(fl.Nov.)Dusen s.n.(S,neotype). CULTIVATED: Germany, at Munich (fl.Feb.) Bogner 1305 (WAG); ibid,(bud) Kress s.n. (M); Netherlands, atWageningen (fl. May) Crusio8(WAG).

c. var.caladiifolia ENGLER
ENGLER, 1915: 5.

Fig.3,Phot.4,Map 4

Type: Bipinde, Cameroon, Zenker 3343(B,lectotype; isotypes BM, BR, COI, E,G,K,L,M, P, S,Z). Petioleequalling oruptoovertwiceaslongastheleaf-blade, 10-54cm long. Leaf-blade ovate-elliptic, acute oracuminate,notapiculate, 10-23cmlong, 5-14 cm broad, l-2 x aslong aswide,puberulous orglabrous below,base sagittate,beloworaboutthemiddlewidest.Manyprominentlateralnerves,with 1-4 lessprominent lateral nervesbetweenthem,sometimesunitingbeforethey reachthemargin,attheedgeanastomosingin2-3marginalnerves.Oneofthese sometimes conspicuous atsome distance from themargin. Between thelateral nerves numerous slender, transverseveinlets. Peduncleslightlylongeroralmosttwiceaslongasthepetiole, 13-40cmlong.

inflorescence. Phot.J. W.
PHOT. 4. Anubias barteri SCHOTT var. caladiifolia ENGLER

MUGGE,

calidario WAG, 21.XI.1978.- (J.V. 333, WAG). Meded. Landbouwhogeschool Wageningen 79-14(1979)

15

Spadixequallingoruptoover \\ x aslongasthe spathe.<$ part l\-2\ x as longasthe? part. D i s t r i b u t i o n : SENigeria,Cameroon, Fernando Po. JO _0 JO _20_

10

MAP 4. Anubiasbarterivar. caladiifolia

Specimens examined:
CAMEROON:Env.ofLolodorf (fr. June)Annet241 (P);Bipindi(fl.June)Annet369(P); just Eof Kribi, tributary of Kienke River (fr. Dec.) Bos 5748(WAG; partly: see also var. glabra); Div. Victoria, Loc.Victoria (fr. Mar.) Brenan 9257 (B); Route des Mbos (fl. Mar.) CNAD 659(K, P); Prefecture Buea,Loc. Ekoma(st. Mar.)CNAD 1859 (K,WAG); s. loc. (fl.)Deistel129 (B, M); 10 km SSW of Zingui (fr. Mar.) Letouzey9093 (P);Nkol Tsia hill, 18km NW Bipindi, near Gonap (fl.) Letouzey 12741 (P);AmbasBay(fl.)MannXV (K,syntypeofvar.glabra);Rumba (fr.Jan.)Paysan s.n. (WAG);Lolodorf(fr.) Staudt354 (COI;partly:seealsovar. barteri);Victoria(fl.May)Winkler 19b (Z); SofN'kongsamba(fr.) De Wit 8039(WAG);s.loc.(st.) Zenker s.n. (BM);ibid,(fl.) Zenker s.n. (P);Mimfia (st.) Zenkers.n. (BR);Bipinde(fl.)Zenker90Ibis(G, P;partly:seealsovar.glabra); ibid,(fr.) Zenker2853(Z;partly:seealsovar.glabra); ibid,(fl.,fr. July)Zenker 3343(B,lectotype; isotypesBM,BR,COI,E,G,K, L,M,P,S,Z);ibid,(fr.)Zenker 4589(E,G,K, LE,iso-syntypes). NIGERIA: Ogoja, Ikom(st. May)FHI18880 (S;partly: seealsovar.barteri) FERNANDO PO: S.loc.(Minola?) (fr. Mar.) Guinea 1441 (K) m ^ f V u - r f D ; N e t h e r l a n d S ' a t W a S e n i n g e n ( Sept.) Crush 14(WAG);ibid. (fl. Sept.) J.V. 322 (WAG);ibid.(fl.Nov.)J.V. 333 (WAG).

d. var.glabraN. E.BROWN
BROWN, 1901:185; HEPPER, 1968b:120.

Fig. 3,Map 5

Type: Cameroon, Preuss422(K,lectotype). 16

Meded. Landbouwhogeschool Wageningen 79-14 (1979)

H e t e r o t y p i c synonyms: A. lanceolataN. E.BROWN, 1901: 183;ENGLER, 1915:4;HUTCHINSON&DALZIEL,1936:366;HEPPER, 1968b:120.Type:Nigeria, Holland167(K, lectotype). A. minima CHEVALIER, 1909: 134;HUTCHINSON& DALZIEL, 1936:366; HEPPER, 1968b: 120.Type: Ivory Coast, Chevalier 19649 (P, lectotype). Petiole\- \\ x aslong astheleaf-blade, 3-35 cmlong. Leaf-blade flat or undulate, lanceolate, ovate-elliptic or oblong-elliptic;obtuse,acuteoracuminate,sometimesapiculate,glabrousorpuberulousbelow,621 cm long, l-9 cm broad, 2 to over 4 x as long as broad, base cordate, truncate or narrowly cuneate. Widest about or below the middle. Manyprominent lateral nerves, with 2-6 less prominent lateral nerves between them, whichunitemostlybeforetheyreachthemargin.Lateralnervesanastomosingin 2-3 marginal nerves when they reach the margin. Many slender transverse veinlets between the lateralnerves. Peduncleca.\ to twiceaslong asthepetiole,6-43 cmlong. Spadix nearlyequallingtotwiceaslongasthespathe,$part2^-4 x aslongas the$ part. Distribution:Guinea,Liberia,IvoryCoast,Nigeria,Cameroon,Fernando Po, Gabon, Congo. N o t e s : 1.BROWNbasedhisdescriptiononthreespecimens: ^ ^ [ h e r e M designated asthelectotype),MannXFand Preuss 1223. FW* but doesnot matchtheoriginaldescription ofvan^ " " ^ J ^ J o w the leaf. It ishereassigned to var.caladiifolia. Preuss 1223wasn * s ^ H o w ever, the keeper of B informed me that the . f ^ . ^ ^ ^ L ^ d 'Schaupreparat' of A. afzelnSCHOTT. T h e s e ^ ^ ^ " ^ ^ f S mens occur frequently, as appeared from theherbarium ^ ^ ^ maybepresumedthat thethirdsyntype,Preuss1223,^f^Tallnceolata. It 2 Inthesamepublication BROWN(1901)also< f ^ ^ ^ being wasmerely distinguished from A. barteribythe^ ^ ^ ^ was glabrous,whileA. barteriwassaidtobepuberu lous.hradd-on,^ he g notedtobesessileinA. lanceolataandonashortstyle ^baen. presenceorquantityofpuberulousha^ vary widely even on one specimen. The sameapp f f A harmi, stigma on a short style.Onlyinleaf-shape A. f ^ ^ J ^ a t r a n A butinthat respectitfully agreeswith^ e s c n p t u , " ^ ^ ^ . ^ isclearlyconspecificwithPreuss422.\ h ^ e ^ t m lanceolata'. var.glabrahaving priority inthisrank tothes epun , l e c t i o n s from 3. in 1909 CHEVALIER described A ^^^^cZaUer 19746;Chevar Ivory Coast(Chevalier19649, herechosenasjectotyp , ^ t w o of the fer 79475; all in P). The <M^ OT ^ ^ . ^ L ^ t y p e mostprobably islost, syntypes (Chevalier19649 and 19746) heth dJ ^ . ^ ^ ^ s p a t h e s b e in g Unfortunately boththesyntypesareinaratne t i o n h o w e v e r i t became almostcompletelyeatenbyinsects.Alteracio ^
Meded. Landbouwhogeschool Wageningen 79-14(1979)

clearthat theybelonginA. barterivar.glabra.Thespecimensare smaller than usualbutsizealonehaslittletaxonomicalvalueinAnubias. 4. A. minima A. CHEV. was already reduced to A. lanceolata N . E . B R , by HUTCHINSON (1936: 366),but HEPPER (1968b: 120) segregated them again. A. barteri var.glabraN.E.BR, was united with A. lanceolata N . E . B R , by ENGLER (1915:4),whoalsoremarked,thatA. lanceolatawasdifficult todistinguish from A. nana ENGLERand A. barteri SCHOTT. ENGLERstated that A. nanamight bea variety of A. barteri.HUTCHINSON(I.e.) did not mention a var. glabra,but he placed one syntype{MannXV) inA. barteri,another syntype{Preuss422, here designatedasthelectotypeofvar.glabra)inA. lanceolata. HEPPER(I.e.)referred var.glabraagain to A. barteri while declaring that A. nanaappeared to be the sameasA. barterivar. glabra. Specimens examined:
GUINEA:Conakry (fl., fr.June)Alleizette 7810 (L). LIBERIA:E Province,Webodistr.,Mnanulu(st.June)Baldwin6026(K);E Province,Webodistr., Nyaake(Webo) (st.June) Baldwin 6118(K). IVORY COAST: Between Gabo and Fete (fr. May) Ake Assi 7861(K); Pays des Tepo, village de Graboetenv.(fl.)Chevalier19649(P,lectotypeofA. minima);ibid,(st.)Chevalier19746(P,syntype ofA.minima). NIGERIA: Oban district, cut lineW of mile 51(fr. Mar.) Coombe 168(B,BR, K); Prov. Ogoja, Distr.Ikorn,Loc.CrossRiver,Mfum, ferryonIkom-Mamferd.(fl.)FHI28315(K);Prov.Calabar, Orem, 66 ml. on Calabar-Mamfe Rd., S boundary of Orem enclave (fl. Jan.) FHI 36037(BR); Calabar, between Akor and Orem, Oban Forest Res. (fl.) FHI 36091 (K); Calabar, Distr. Orem, ObanForest Res.SNR4(fl.)FHI45823(K,WAG);NWofIkuru(fl.Jan.)Holland167(K,lectotype ofA. lanceolata);Prov. Ogoja, Loc.ObuduCattleRanch,south atcataract ravine(fl.Apr.)Medler 811(K); Oban distr. (fl.) Talbots.n.(K);Oban (fl.) Talbot768(K). CAMEROON: Reg.deDoume(fr. July) Alleizette 7810(L);region Lolodorf, Mt. Ngowayang (fr. June) Annet 203(P); Bipindi (fr. June) Annet 389(P); 14miles from Victoria in the direction of

10 18

10

20

30

MAP 5. Anubiasbarterivar.glabra. Meded. Landbouwhogeschool Wageningen 79-14 (1979)

Nigeria(fr.Jan.)P.Becks.n. (WAG);Dept.Odean,Nko'olong(= Nkoelon)(fr.)Bililong&Bullock 107a (K);5^kmSofKribi(fl.)Bos3121 (WAG);8kmfromKribi(fl.,fr.Nov.)Bos3397(WAG); 19 kmfromKribi, 5kmNofLolodorf(fl. Feb.)Bos3899(WAG);18km fromKribi(fl., fr.Mar.) Bos 4152 (WAG);ibid.(fr. Mar.) Bos 4153 (WAG);tributary ofKienkeR., justEofKribi(fl.,fr.Apr.) Bos4281 (WAG);just EofKribi,tributary ofKienkeR.(bud,fl.Sept.)Bos 5400(WAG);ibid.(fr. Dec.) Bos 5748bis(WAG;partly:seealsovar. caladiifolia);about 60 km N ofKribi(fl.,fr.Feb.) Bos 6349(WAG);s.loc.(st.)Brauns.n. (M);Lolodorf,Mt.Minn(fl.Nov.)Jacques-Felix 9170 (P); Rio delRey (fl.June)Johnson s.n. (K,syntypeofA. lanceolata); Bibundi(fr. Mar.)Jungner 48 (S); Mt. Kamerunensis, Isowi (fl., fr. Nov.) Jungner143 (S, UPS); between Bipundi and Isowi (fr. Jan.) Jungner 143 (UPS);closetoKienkeR.,NNWofNkolbawa (fl. Mar.)Letouzey8986(P); nearBodi, 20 kmSWEseka(bud,fr.Dec.)Letouzey12505(P); Victoria(fr.)Ludwigs36 (M); Ekuk,20 km E of Ebolowa (st.) Mildbraed5725 (HBG; partly: seealso var. barter!); Fenda, 58km Eof Kribi (fl.) Mildbraed5931 (HBG;partly:seealsovar.barter!); Barombi(fl.Aug.)Preuss422 (K, lectotypeof var.glabra);s.loc.(fl.)Preuss465 (M, withdesignationA.puberulaENGLER,nomennudum);43km 5ofbridge ontheroadconnectingKribiandEdea(fl.Jan.)Purzl 771 (B); Assok,65 kmESEKnbi (fl. Mar.)Raynal10379(P); betweenEdeaandKribi(fr.)Sanford5719(K);ca. 40 km SofBadjob ca 50km SWofEseka (fr. Dec.)De Wilde c.s. 1555bis(K,WAG,Z;partly:seealsovar.barteri);Ml facing thevillageofN'kolandom(fr. Oct.)De Wilde 7658(WAG);S. ofN'kongsamba(si:. Dec.)De Wit 8282(WAG);Bipinde(fl.,fr.)Zenker901 (E,K,L,LE,M, P); ibid,(fr.)Zenker2853(BR, E, G, K,L,S,Z;partly:seealsovar.caladiifolia;designatedasA.lanceolata.cordataENGLER,nomen nudum).
FERNANDO Po: s. loc. (fr. Dec.) Mann 104 (K).

GABON:Sibang(f,)i4-^(K);MWoum(^ 689(K, M,WAG);42km SEofLambarene(budSept.)BretekrSmfWAG), LanoR.,Mont,de CrisUl5kmSEofAsok(fr.Aug.)*retefc^ MbeRiver(fl.,fr,Aug.)^/er*ZW^^ Cristal,chutedeKinguele(fr.Jan.)Halle&Villiers4433(P); ^ ^ ^ ^ ^ i " f A 6 Villiers 4491 (P);ibid.,Nkam (fr. Feb.) Halle & Villiers 4841 (P);MontsdeCr.stal(fr.) Halle Villiers5218(P);env. of Libreville(fr. Nov.)Klaine2008(BR,P). CONGO:Fl.Ogooue (fr. July) Thollon 856 (P). . A > B ner 920 UNKNOWN LOCALITY:(fl., fr.) Anonymus s.n. (BM);y.acommerc.aldealer(fl. Aug) (M); (fl.) Bogner1302 (WAG) (fr.)^ J ' ^ % . Netherlands,atWageningen(fl.Aug.) CULTIVATED: Germany, at Munich (fl.) Bogners.n (K), r^ei mWAG); Denmark,at Crusio7(WAG);ibid. (fl. Aug.) Crusio 9(WAG) .tad. f ^ ^ ^ Copenhagen (fl.) N.Jacobsen s.n. (WAG);ibid,(fl.)N.Jacobsen3059(WAG).

. e. var.nana(ENGLER) CRUSIO,stat.etcomb. nov. H o m o t y p i c synonym: A. nana ENGLER 15'.

A

Fig. 3, Map 6 >

m

PMPIFR 1899- 423; BROWN, 1901: 186;

ENGLER, 1915: 4; HUTCHINSON& DALZIEL, 1936. 366, HEP

Type: Localityunknown,Bogner 1295 (WAG;neotype). Petiolesomewhat morethan half as long J ^ 1 ^ Zm^* Leaf-bladeovate-elliptic,acuminate-acute,api ^ ^ ^ ^ ^ wid^ m o s t l y cmbroad,glabrous,baseobtuse-truncate at ^ ^ ^ M a n y m o r e o r somewhatlessthan 2 x aslongaswide,widest ^ ^ ^ margin) at the lessprominent lateral nerves,which" ^ ^ w U h m a n y slender,transverse edge anastomosing in 2-3 marginalne veinletsbetweenthem. 19
Meded. Landbouwhogeschool Wageningen 79-14(1979)

Peduncle 2-4 x aslong asthepetiole,up to 16cm long. Spadix aslong asthe spathe.$part somewhat shorter than theS part. D i s t r i b u t i o n : Cameroon: Victoria. Notes: 1.ENGLERbasedhisdescription oncultivated specimens,whichwere collected by LEHMBACHatVictoria, Cameroon. HUTCHINSON& DALZIEL(1936) state that they have seen a conserved specimen, which should be the type. However,nosuchspecimencouldbetracedanditmaybeassumed,thatthetype specimen of A. nana(if any) was destroyed during the war. After the original collection, A. nana was only twiceagain collected in a natural growing station: byWINKLERin 1904at Muca (= ?Muyuka),Cameroon, and by PURZLin 1972 who secured one sterile specimen at the type locality. A color-slide of a spathe (not present on the herbarium sheet) was added. Furthermore J. BOGNER, Munich,provided mewithsomeexcellentlypreservedflowering specimens,from an unknown locality{Bogner 1295 and 1297),togetherwithlivingmaterial. Bogner 1295 ishere assigned asneotype. 2. Already in 1915, ENGLER suggested that A. nanamight be merely a small form ofA. barteri. The study ofthematerial at mydisposal (which fully agrees withthedescriptionofA. nana)provedtomethatthisopinioniscorrect.A.nana however can bemaintained asavariety inA. barteri. 3. Seealso note4. subvar.glabra. Specimens examined:
CAMEROON:N ofVictoria, inabrook flowing from Mt. Cameroon (st, fl.?Nov.) Purzl 741 (B); Muca (fl. July) Winkler 231 (Z). UNKNOWN LOCALITY:(fl.) Bogner 1295 (WAG, neotype); (fl.) Bogner1297 (WAG).

10
MAP 6. Anubiasbarterivar.nana.

10

20

30

20

Meded. Landbouwhogeschool Wageningen 79-14(1979)

CULTIVATED: Netherlands, at Wageningen (fl. Sept.) Crush16 (WAG, cultivated specimen of Bogner1295, neotype); ibid.(fl.Aug.) Crush17(WAG,cultivated specimenofBogner 7297);ibid, (fl. Oct.) Crush 18(WAG, plant received from MOHLMANN, GFR, as coming from Cameroon: Kumba); ibid. (fl. Sept.) J.V. 315 (WAG); ibid.(fl.June)J.V.568(WAG).

3. Anubiasgigantea CHEVALIERexHUTCHINSON

Fig.4,Phot.5,Map7

CHEVALIER, 1920: 683(nomentantum); HUTCHINSON&DALZIEL, 1936: 366

(nomennudum);HUTCHINSON,1939:246;HEPPER, 1968a:456; HEPPER,1968b: 120. Type: Guinea, Chevalier20857(P, holotype). H e t e r o t y p i c synonyms: A. giganteaCHEVALIERvar.tripartitaCHEVAUW, 1920:683(nomennudum).Type:Guinea,Chevalier20858;IvoryCoast, ChevaHer21606(both P; no lectotypechosen). A. hastifoliavzv. robustaEnaim, 1915:9; HEPPER,1968a:456.Type:Careysburg, Liberia, Dinklage2454 (B,holotype). Rhizome creeping, prostrate and rooting, 1-3 cmthicL ^ J ^ l T t o ^ P Petioleslightlyshorterthan orupto1\ x^ ^ ^ f ^ ' cmlong. Sheath short or upto themiddle.Geniculum -2jcrnlong. Leaf-bladehastately lobed sometimesnearly ^ ^ ^ ^ Z lanceolate, 13-30 cm long, 5-14 cm broad, 2-4 x a> long as , belowthemiddle,glabrousbe^ cmbroad.Manyprominentlateralnerves,with2-4lesipromi betweenthem,whichsometimesunitebefore ing in 2-3 marginal nerves and with numerous slender, transver between them. , ]ml(I Pedunclealittle shorter than thepetiole^14^0'ng. ^ Spathe 3 i - 8 cm long, oblong,^ ^ ^ ^ Z ^ ^ and the top open. Before anthesis opening halfway^downclosing tightly and entirely again after afewh o ^ bho 5_10 m m Spadix alittlelonger than the spathe u to9 cm to g , ^ ^ ^ flowered thick, <? part l i - 3 x aslong as the.? pari.-Van 6 V hitCj d i s c o i d > Stamens4-6(8).Lower^ flowers o m e t u n w ^ ^ 8 ^ ^ ^ ^ ^ ^ s subsessile or on a short contracted style.l h e c f . depressed-globose. Dried oblong, opening with a longitudinal fissure^erne^ j ^ Q ^ ^ m m b r o a d i seed yellow, with many brownish s P o t s ' ' f l y o n c e observed,Adams4551,B). 1.3-2 xaslongasbroad.(Seedsandberriesonly T ihpHa IvoryCoast,Togo. D i s t r i b u t i o n : Guinea, Sierra Leone,Libena, y t thebanksofriversorintheriverbed.Mostlyon Ecology: Semi-aquatic, at thebanksoi rocky places.Flowering from February-April.
Meded. Landbouwhogeschool Wageningen 79-14(1979)

t
: V d e ^ f ' T ^ T ' f ,CX H U T C H I ! ^wer side"leaf, *x ; 2. inflorescence, ~ d r i , l f ^ ' a C k S l d e l e a f ' e n l a r g e d ; 4.uppersidesynandrium,16x ; S.partofa synandnum,sideview, 16x . - (1-5. flos 79/4,WAG). 22 Meded. Landbouwhogeschool Wageningen 79-14(1979)

HOT. 5. Anubias gigantea C H E V I ^ T ^ N S O N _ Inflorescence (right: with spathe cut open).Phot. N. JACOBSEN,calidarioC, 1978. - (Jacobs* 3058,WAG).

frfcartifa' inA g/gotefl, which seemsthecorrectdecision. 2. In 1915, ENGLER described A. hastifohayu. ^ ^ ^ Z B ^ L B C B 2454from Liberia.Strangelyenough,thiss p e c i m e n ^ n o t _ ^ ^ description, being much smaller than ^d'catea ^ ^ ^ jn ^ ^ undoubtedly belongs in ^. gigantea,and X o/ s v n o n y m y ofA.gigantea HEPPER'S reduction of A. hastifoliavar.rotow tothesynony y (1968a: 456)iswarranted. , w h i c h j t differsmerely 3. ^ . giganteaisverycloselyrelatedtoA. fe,">* , e s s d i f T e r e n t from A. by the shape of the leaf-blade. Other charactersmo h o f t h e p e t j 0 ieand afielii are: petiole/leaf-blade ratiocolourofthestigm , ^ &^ . ^ QU the spadix/spathe ratio.It ispossible,that A .gig db o t h t a x a a r e ajfc/ff. However, asnointermediary specimenswereobserve ^
Meded. Landbouwhogeschool Wageningen 79^14(1979)

MAP 7. Anubiasgigantea.

very constant with regard to the shape of the leaf-blade, it seemspreferable to maintainthe twospecies. 4. Itwasoncereported{Adams 4551) that theinflorescence wassmellinglike fresh fish. Specimens examined:
GUINEA:Ziama(st. Febr.)Adam3580(P); Nionsomoridou,closetoBeyla(fl.) Chevalier20857(P, holotype);between Diendedou and Nionsomoridou (fr. Feb.) Chevalier20858(P,'syntype' ofvar. tripartita). SIERRA LEONE: Mano (fl.)Deighton3373(K,P);KondumbaiaN.P.(fl. Mar.) Morton & Gledhill SL 1024 (K,WAG). LIBERIA:GolaNat.Forest,NEofBomiHills(fl.Apr.)Bos1914 (K,WAG);halfway PeterstownRebbo(fl.Nov.)Bunting94 (BM); Monrovia,Careysburg(fl.Feb.)Dinklage2454(B, holotypeof A. hastifoliavar. robustaENGLER);BailabySt. JohnR.(fl.Feb.)Harley1478(K); Bensonville(fl. Mar.) Jansen1577 (WAG). IVORYCOAST:Troya(st. Feb.)AkeAssi6925(K); MontTonkoui,Gbepleu(fl.Feb.)Ake Assi7366 (BR, K); entre Dabou et Becedi (st. Mar.) Ake Assi8529(G); Ht. Sassandra, Pays Coura, entre Sanrou et Ouode(st.) Chevalier21606(P,'syntype' ofvar.tripartita). TOGO: Dutukfene (fl., fr. Dec.)Adams4551 (B,BR,P). CULTIVATED: Denmark, at Copenhagen (fl.) TV. Jacobsen 3058(WAG); Netherlands at Wageningen(fl.May)J.V. 520 (WAG).

4. AnubiasgilletiiD EWILDEMANet DURAND 1915:8. Type: Kimuenza,Zaire,Gillet1696 (BR,holotype). 24

Fig. 5,Phot.6,Map8

DE WILDEMAN & DURAND, 1901: 845; D E WILDEMAN, 1903: 13; ENGLER,

Meded. Landbouwhogeschool Wageningen 79-14 (1979)

4-

' i x - 2 upper sideleaf,

and C W , , p i r i l coll.WAO; 2.C M ' " Me<fe</.Ltmdbwwhogesckool mgeningen 79-1-1(1^1

'

25

 A. A. gilletiiis narrowly allied to A. hastifolia ENGLER. However, it can be easilydistinguished bytheposition ofthethecae,form and sizeofthespathe and by the spathe in anthesis being reflexed (not reflexed in A. hastifolia). Fruiting specimens are very hard to distinguish from A. hastifolia and A. pynaertii DE WILDEMAN,theycan onlybedistinguished when at least aremnant ofthe$ part of the spadix is left. Sterile specimens of these species cannot be segregated from each other. Specimens examined:
NIGERIA: Shasha Forest Reserve, Prov. Ijebu (fl.) Ross 121 (BM); Eket district (fl.) Talbot 3168 (BM). CAMEROON:s.loc.(fl.) Bogner 1320 (WAG); 8km SofKribi(fl.May) Bos 4588(WAG); SEslope of Mt. Elephant (fl. Nov.) Bos5667(WAG); 28km ENE Eta (bud, fl. Feb.) Letouzey 11972 (P). GABON: Booue (fl., fr. Nov.) Bogner720 (K, M, WAG); 30 km SW Makokou (fl. Oct.) Halle 2709 (P). CONGO: Route Mutamba-Kimpanzou (fr.) Descoings6210 (P); Env. of Brazzaville, between N'fonkama and Moutamba(fl. Nov.) Sita 1945 (P). ZAIRE: Befale, riv.Ifale(bud,fr. Dec.)Eward3139(BR,K);Kimuenza(fl. Oct.-Nov.) Gillet 1696 (BR, holotype); Binga cave(fl.June) Jans499(BR);Nsemakoo(fl. Mar.) Jans 1004 (BR). UNKNOWN LOCALITY:Africa (fl.June)Bogner54 (K);W.Africa (fl.) Bogner453 (M);(fl.)Bogner 1296 (WAG). CULTIVATED: Germany, at Munich (fl.) Bogner3a(K);Netherlands, at Wageningen (fl. May& Sept.) Crush 4(WAG);ibid.(fl.May)Crush6(WAG,cultivatedplantsfrom Bogner1296);ibid.(fl. Aug.)7.K.307(WAG).

5. AnubiasgracilisCHEVALIERex HUTCHINSON

Fig.6,Map9

CHEVALIER, 1920:683(nomen tantum); HUTCHINSON& DALZIEL, 1936:366 (nomen nudum); HUTCHINSON, 1939:246;HEPPER, 1968b:120.

Type: Guinea, Chevalier20800 (P,holotype). Petiole up to 33 cm long, \\-2\ x as long as the leaf-blade, very short sheathed. Geniculum 1-1^cmlong,glabrous. Leaf-blade7-12cmlong, 4-10cmbroad atthebase,glabrous,coriaceous, slightly hastate-trilobate, the outline ofthe leaf-blade triangular, lobes 7 cm long,2-3 cmwide,apexobtuse. Length of the blade equal toitswidth at the base.Numerousprominent lateralnerveswith2-3 lessprominent lateral nerves between them, at the margin anastomosing in 2 or 3 marginal nerves. Many slender transverse veinlets between them. Peduncle 8-15 cmlong, | a s long asthe petiole. Spathe1^-3cmlong,oblong or oblong-elliptic, apiculate.Colour unknown. Spadix a little shorter than or equalling the spathe, up to 3cm long, densely flowered. Stamens (6)7-8(9). Thecae on the side of the synandrium. Ovaries depressed-globose with many whitish spots. Stigma discoid, subsessile. $ part 4 x aslong asthe$ part.
^ Meded. Landbouwhogeschool Wageningen 79-14(1979)

*
i%; 2.upper sideleaf, ^ " H O N : 1..ower side>eaf i * ; * ^ FIG. 6. /4n6 bias gracilisCHEVALIER ex HUTCHINSON_ d r i u m , 1 4 x ; 6-f? p. 2<t x ; 3.bud, x x g ;4.inflorescence $A,x .. -rr-- * . _ (( , , 3 - . ^caues-^ 4.inflorescence j , * 2 * ' 1; ; view, 14x ; 7.detail of nervature,lowers.deleaf, g 29 Chevalier20578, P). Merferf. Landbouwhogeschool Wageningen 79-14(197

Distribution: Guinea, Sierra Leone. Ecology: Growing in wet places, sometimes in river beds. Flowering from February to May. Notes: 1. A. graciliswas named by CHEVALIER (1920: 683) without a description. This was provided by HUTCHINSON, 1939: 246, who described this speciesaccording to theCode. 2. A. gracilisisveryimperfectly known. Ihaveseenonly6collections(3 from Guinea, 3from Sierra Leone),and someoftheseweresterile.Thevariability of thisspeciesisinsufficiently known. Itisalsounknown whether thespathe opens when flowering, or remains closed (like in A. afzelii and A. gigantea). Some specimens suggest that the spathe remains closed, only opening at the top, but Jacques-Felix1561 shows an opened spathe with a somewhat exserted spadix. This may be an artefact, due to the drying-process. A. gracilis seems to be narrowly allied to A. giganteaand perhaps it ismerely a variety of thisspecies. The large number of stamens (Jacques-Felix1561)is, however, an important characteristic.Theremnants ofthespadixofthetypesuggest the same number. Nevertheless, this high number of stamens is occasionally also counted in A. gigantea.A. gracilis, then,isdistinguished from A. giganteaand A. afzeliibyits smallspathe,highnumber ofstamens andtheshapeoftheleaf.From A. barteri and A. heterophylla it is distinguished only by the number of stamens and the leaf-shape. Specimens examined:
GUINEA: Pays des Koniankes: Fassakoi'dou (bud) Chevalier 20800 (P, holotype); Zou-

30

10
MAP 9. Anubiasgracilis.

20

30

30

Meded. Landbouwhogeschool Wageningen 79-14(1979)

bouzoumaye(bud May)Jacques-Felix 906 (P); env.ofMacenta(fl. Mar.) Jacques-Felix 1561(P). SIERRALEONE:Regiondes sourcesduNiger:Souradou(st. Jan.) Chevalier20578(P);betweenJifin and Bandakarafaia (st.Apr.)Deighton 5066(K);TingiMountains, NKono(budApr.)Morton & GledhillSL 1929 (WAG).

6. Anubiashastifolia ENGLER
ENGLER,

Fi

8-7'MaP , 0

1889: 149 (nomen tantum, 'A. hastaefolid);ENGLER, 1893: 462;

BROWN, 1901:185; DEWILDEMAN, 1903: 13;ENGLER, 1915:9(excl.var.ra^

= A. gigantea CHEV.ex HUTCH.). H o m o t y p i c synonym: Amauriellahastifolia (ENGLER) HEPPER, 1968a: 454;HEPPER, 1968b: 120. Type: Gross-Batanga, Cameroon, Braun 6(B, holotype). H e t e r o t y p i c synonyms: Anubiashastifolia^. sub^ENM 1893: 463; BROWN, 1901: 186;ENGLER, 1915:9.Type:AttheJego,Togo, Klmg36(B, holotype; probably destroyed inthewar). m5:l\Amaum.FNGLER
A.auriculataENGLER, 1899:423;BROWN, 1901.184,ENGLER 1*13. ,

riella auriculata (ENGLER) HEPPER, 1968a: 454. Type: Batanga, Cameroon, Dinklage986(B,lectotype,isotype HBG) E n v of A. haullevilleana D E WILDEMAN, 1903: 13;ENGLER,m a . yv Kisantu, Zaire, Gillet1993 (BR, holotype). 7/ i i W ^ . laurentiiDEWILDEMAN,1910:169.Type:Env.ofKisantu,Za.r (BR, lectotype). HS-FNGLER 1915:1;LEMEE,1929:188; AmauriellaobanensisRENDLE, 1 9 1 3 : I I ^ N O U , , isotype; HEPPER, 1968a: 454. Type: Oban-district Nigeria Tdt^^1913:pl. holotype probably destroyed inthewar).Amaunella talbotu REND 16; in error. * i _ U cmthick,tipdenselyleaved. iM/zomecreeping,prostrate and rooting, 2 2 ^ leaf-blade. Sheathing Periofe 9-67 cmlong,equallingto 2 x asio g narrowed, for i - x of its length, sheath at the base w d e , u * ^ glabrous or puberulous. Geniculum0.8-2.Uc m i u * ^ broadly ovate. Leaf-blade varying from (sub)hastate (nearlyirp G , a b r o u s o r belowon oblong-lanceolatewithashortcordateorauncuiaw ^ ^ ,anceolate or the midrib and lateral nerves minutely P u b e r u ' o u , l 0 _ 3 3 c m long,3-14cm elliptic to obovate-elliptic; 2-4 x aslong asb r o 4 ^ Laterallobes wide,short acuteoracuminate,widestatorjus:o ^ % ^ , o n ga n dg smaller than themedian lobe,obtuse,sometimesacu v ^ prominent cm broad. Numerous prominent lateral nerves' ^ ^ y slen dertransverse lateralnervesbetween them.Betweenthelatere ^ nerves. 31 veinlets. Lateral nerves at theedge anastomosing
Meded. Landbouwhogeschool Wageningen 79-14(1979)


l l o w e r s i d e le ridev{J"iaS^,if0,ia^GLER:s d e af, f x ; 2.inflorescence, | x ; 3.synandrium, I ' , ? ; ! 4 R U P P : r ' syandrium,20x ; 5.detailofnervature,lowersideleaf,enlarged. U - A 5 . Braun6,B; 3-4. Bos4548,WAG).

32

Meded. Landbouwhogeschool Wageningen 79-14(1979)

Peduncle\ tooneandahalftimesaslongasthepetiole,mostlyshorter,8-24 cmlong, slender, sometimes minutely puberulous. Spatheoblong-lanceolate,convolute,notopeningwhen flowering, onlyat but theupper part open (oropeningasinA.afzeliiandA.giganteal), aslongas the spadix, 2-U cmlong,-1cmwide,brown, green-pinkish,orwhite Spadix H - 4 cmlong,2-5mmthick.< partslightlylongertoupto32times;as J long asthe ? part. Pistils notcrowded, depressed-globose,morethan 20. Style short, stigma discoid. Stamens4-6.Thecaeovate,placedentirelyoratleastthe upper half onthetopofthe synandrium. Seeds 1.7-2.5}& ^J* broad; 1.4-1.8 x aslong asbroad. Yellow,without anyothercoloured spots (dried, only observed once). D i s t r i b u t i o n : Ghana, Nigeria, Cameroon, Gabon,Zaire. Ecology: Onthebanks oflittle streams intheforest, on rocksorin mud. Flowering throughout theyear. Fruiting from SeptembertillJanuary. V e r n a c u l a r name: Batoiibangwa(Turumbu,Germain4854). N o t e s : 1.Together withthespecies,J * * ^ * *' ^ var. sublobata,based onKling36,asterile^ W ; afld c h a n g e s The shape of the leaf-blade in A. ^ ^ ^ ^ J ^ J ^ n leaf-shapeare moreover withtheageoftheplant.No * > ^ S v K e war.Itwasa recognized here.Thetypeofvar.sublobata P ^ ^antea CHEV. ex sterile specimen and soit ispossible, that it oeio g ^ m a y a,s0 rec rded lr HUTCH. Nevertheless, A. hastifoliawas is h e r e placedinto occur inTogo. Forthese reasons thevar.sublobata ENGLER the synonymy ofA. hastifolia. ninklase986andZenker 2. In 1899ENGLERdescribed^,auriculata basedon^^ ^ ^j m p o s s i b l e t 0 1174. Both these collections onlyhadsomebuds ss> t h e posit ionof ascertain thecharacters ofthe spathe inantnesis. i ^ unjted here, the thecae is identical with that inA. has%olia. " n o w taxonomicimportance becausethedifference inleaf-shape ,sconsidered ollow (see above). . D E WILDEMANandbasedonone 3. A.haullevilleanawasdescribed^ 1 9 0 3 ^ E ^ d h i s speci esfrom A. has* fructificating collection {Gillet 1993)He^ ^ J o * innote 1,th folia bytheleaf-shape andthesizeoftheP ^ * ^ s o m e difficulties asto insufficient evidence.However,A.^ f * ^ r large, which prevent. . its specific identity. The spathe on to type sra ^ n on reduction of A. haullevilleanatoA.pile*. Because c ^ . ^ o f ^ s p a t h n the type and no observations were ^ ^ ^ ^ ^ . n ^ anthesis,A. haullevilleanamaybelongto,A.ha ' A h tlfolmVo OnthetvPelocality(orvery^ thatreasonandalsotofurther stability nno placed into thesynonymy ofA. hastifolia. 33
Meded. Landbouwhogeschool Wageningen 79-14(19

10
MAP 10. Anubiashastifolia.

4. In 1910D EWILDEMANdescribedA. laurentii. It wasdistinguished from A. hastifoliaonly by the shape ofitsleaves.Both the syntypes wereexamined and proved to beconspecific withA. hastifolia. 5. Amauriella obanensiswas described in 1913 by RENDLE. The holotype (Talbot 1532) wasprobably destroyed at theBM,but at K anisotypeisextant. This collection, together with the drawing RENDLE made (there designating erroneously thespeciesasAmauriella talbotii RENDLE)prove,that thisspeciesis identical with A.hastifolia. HEPPER (1968a) placed A. hastifoliaand A. auriculatain Amauriella, on the strength ofthedifference intheposition ofthethecae on the synandrium. ENGLER(1893) haddividedAnubiasintotwosections:Synanubias,containing A. afzelii,A.heterophylla,andA.barteri;and Cylindranubias,containingonly A. hastifolia. This subdivision was based on the same character as adopted by HEPPER to separate Anubiasfrom Amauriella. Section Cylindr anubiasand the genus Amauriella are the same taxon. Later (1915: 2) ENGLER canceled his sections, because according to him both kinds of$ flowers might occur in A. hastifolia. Thespecimenswiththethecaeatthesideofthesynandrium however, are here assigned to A. gigantea. Unfortunately, both ENGLER and HEPPER overlookedA.gilletiiDEWILDEMAN& DURANDandA.pynaertii D EWILDEMAN. Theposition ofthethecaeinA.gilletiimaybeconsidered intermediary between A. hastifoliaandtheotherspeciesofthegenus(excl. A. pynaertii).Thepositionof thethecaeinA. pynaertiicouldbeconsidered acombination ofbothtypes.It has to beconcluded, that there exists no demarcation in the position of the thecae within thegenusAnubias SCHOTT,and soAmauriella RENDLEisnot maintained here. ENGLER'Ssections are in consequence also rejected. No valid other argument wasfound to subdivide the genusAnubias. 34
Meded. Landbouwhogeschool Wageningen 79-14(1979)

Specimens examined:
GHANA:VaneT.V.T.(fr.June)MortonA3439(K);Chilinga&Skiare,Nkwanta,Krachi(fl.May) MortonA 3977(K). NIGERIA:Div. Benin,Loc.OkumuForestReserve,Compt69(fl., fr.Jan.) Brenan8762(Q, BM, K, P); Prov.Benin,distr.Benin,Sapoba,JamiesonRiver(fl., fr.Sept.)FHI34268(K); SNigeria,Oban district(fr.) Talbot1297bis(K); SNigeria,Obandistrict(bud) Talbot 1532 (K, isotypeoUmauriella
obanensis RENDLE?).

CAMEROON: 12km from Kribi (fr. Jan.) Bos 3641 (WAG);8kmSof Kribi(fr. Feb.) Bos 3V72 (WAG); 13kmfromKribi(fl. May)Bos4548(WAG);17kmfromKribi(fr.May)Bos4600(VIAG); Gross Batanga(fl.Oct.)Braun 6(B, holotype);Batanga(fl.Sept.)Dinktage986 (B, IcctotypcofA. auriculata ENGLER,isotypeHBG); Bipinde(bud,fl.)Zenker 1174(BM, E,G,GRO, K,L, LE, M.!', S,Z;syntypeof^. auriculata BUGLER). ZAIRE: Yangambi (fr. Apr.) Germain 4854 (BR); LukenzuRiver,Ikela(fr.June)Germain 7449 (BR); env.ofKisantu (fr.) Gillet 1993 (BR,holotypeofA. haullevilleana DEWILDEMAN);ibid (fl.) Gillet3381(BR, lectotype of A. laurentii DE WILDEMAN); Kassai (fr. Nov.) Laurent s.n.(BR); Batempa falaises du Sarkuru (fl. Nov.) Laurents.n. (1903) (BR, syntype of A. laurentii Dfc
WILDEMAN).

CULTIVATED: Belgium,attheJardinColonialdeLaeken(fl.March)Seret 129 (BR).

7. Anubiasheterophylla ENGLER

Fig.8,Phot.7-11, Map11

ENGLER, 1879:435;1893:463;BROWN, 1889:67;1901:184; ENGLER,1915:5.

Type:GolungoAltodistriet,Angola, ^/w/^237(K,lectotype;isotypc S at B, BM, COI, LE,P). H e t e r o t y p i c synonyms: A. congensisK. E. BROWN, 1901:184; D^WILDEMAN, 1907-17; ENGLER, 1915: 7. Type: Boma, Za.re, Bull s.n. 1889 (K, lectotype). 1 9 1 5 . 7 T v p e : Cultivatedspecimenin A. congensisv&r. crassispadixENGLER,1VO. ' iyp<=. . _ trnVK i i n t h e the Berlin Botanic Garden Berlin-Dahlem (not conserved or destroyed in ^A.affinis DEWILDEMAN, 1907: 16; ENGLER,1915:3; DE W.LDEMAN, 1922: 176. Type: Anonymus s.n. (B^holotype?^ seenote -A) ^ ^ A. engleriDEWILDEMAN, 1907: 17; ENGLER, m a . *. m Laurent s.n. 1895 (BR, holotype, ^ atB); Waljkalc, Zairc, ^ . bequaertiDEWILDEMAN, 1922:176.Type, tnire Bequaert6450(BR, holotype). nlu inil 1 u n g a t e HORT. MOHLMANN, 1977:488(nomennudum). Rhizome creeping, prostrate and r o t i n ^ " 1 7 s X teaf-bladc, glabrous or Petiole 3-66 cm long, 0.7-1.7 x as long a . , Gcniculumi-2Jcin minutelypuberulous.Sheathsmalloruptoj ofthepet.o lon , i i ^ m hrnad 2to 6times as long as broad. Leaf-blade 10-38 cm long, 3 - U cm oro ' o r ( s o m e l i m e s verysparelliptic-ovate or elongate-lanceolate u s J ' b b e l o W , fiat or more or less t a sely) puberulous on the midrib ^ * b a s e acute, cuneatc-obtuse, undulate, coriaceous, apex acute oi ^
Meded.Landbouwhogeschool Wageningen 79-14(1979)

t
FIG. 8. Anubiasheterophylla ENGLER: 1.lowersideleaf,\ x ; 2.inflorescence, completely opened, f x ; 3.inflorescence, somewhat more closed, \ x ; 4. synandrium, side view, 24x ; 5. upper side synandrium, 24x ; 6. berry, 6x ; 7. seed, 14x.-(1-5.Arends71,spirit coll. WAG; 6-7. Bogner 1300, spiritcoll.WAG).

36

Meded. Landbouwhogesehool Wageningen 79-14(1979)

PHOT. 7. Anubias heterophylla ENGLER- Flowering plant. Phot.J. BOGNER,calidario M. .978. (Bogner 1300, WAG).

subtruncate, short sagittate or hastate. Basal lobes * < * J * ^ ^ Z * rounded, separated by a very wide sinus. Numerous prommert latend ncms with 2-4 less prominent lateral nervesbetween them, ^**Z*themoreprominent nervesbeforetheyreachthe ^ ' ^ J ^ L , . ingin2-3 marginal nerves.Betweenthelateralnervesnumerousslender, verseveinlets. . . . ntn?l cmlone,slender, Peduncleonethirdaslongastoequallingthepetiole,upto27 cmlong, glabrous or minutely puberulous. j o n g as 41 ; ^ l - ^ c m l o n g ^ . ^ l i c m b ^ wide,oblong-elliptic or ovate-oblong, g l a b r o u * ' j " l^JrinR b u t not renexed. green or dark purple, opening to & ^ ^ *%*** again tightly Sometimes spadix more or less exserted. After antne around the spadix. . t i c e a s i o n g asthespathe, Spadix c y c l i c a l , somewhat thick,equaUngto ^ ^ ^ , 0 . 6 - 2 1 U-4$ cmlong.$partsomewhatshorterorupio rf somet imesonlyin cmlo2ng.Anthers4-6. Thecaeonthesideo ^ ^ n to beplaced onthe the upper part (especially whendried thet h e c a e y ^ ^ ^ & ^ ? edge of the synandrium, as inA. gilletu), ODIO S r s o m e w h a t l e s s den sely part sometimes sterile flowers c? part denseJ| ^ ^ s t j g m a d i s c o i d , green, flowered. Ovaries depressed-globose, green,a y Q5 J m m b r ) a d ; Berries depressed-globose. Seeds small, 0.8-1.5 mm ^
M e . Landbouwhogeschool Wageningen 79-14(1979)

' " * * : . . . :

PHOT. 8. Anubiasheterophylla
ENGLER Inflorescence.

Phot. H. C. D. DE WIT, calidarioWAG, 6.V.1978.-(Crush 5, WAG).

about 1.3-1.7 x aslongasbroad, yellowish,without any other coloured spots when dried. D i s t r i b u t i o n : Cameroon, Equatorial Guinea, Gabon, Cabinda, Congo, Zaire, Angola. Ecology: Growing asarule onrocky grounds onthebanks oforinwatercourses,onshadyplacesintheforest, 300-1100m.Leonard2856bearsthe note 'plante epiphyte'. Flowering from July toJanuary, fructificating from July to March. Vernacular names: Biranansungu (Kitembo, Pier 1194);Bofokoko bo lot libande (Turumbu, Germain 250). Use: Theplant isused asastomachic forchildren (Pierlot1194). Notes:1. Intheoriginaldescription, ENGLERcitedastypematerialforhis A. heterophylla 'Africa tropica occidentalis, Angola distr. Golungo alto alt.30038
Meded. Landbouwhogeschool Wageningen 79-14(1979)

P5^B!BSBPiiPpP

?$*&:* J

PHOT. 9. Anubiasheterophylla
ENGLER Inflorescence.

Phot. H. C. D. DE WIT, calidario WAG, III. 1977. (Arends71,WAG).

800 m insilvis(We,,Usck .236-23W. Lateron m 1893(p.463) h,;ctal as wereplacedbyhimin^ . a/ze/.In 1915,however,up. J

236, 237and238 asbelongingtoA.*Wt<J%Tw

n o . ^ l . . l . t B M ^ ^ ofthe opinion,that ENGLERin 1879,whendescnwng

T)Am
^ / w / w A 2 i7.

to himby BULLfrom Boma,Zaire in 188/, law an puberulous, c W J from ^ . te^^/fa only because^ ^ / ^ J s h o u l d be especially onthelower surface ofthe leaf-blade whileA g glabrousThetypesofA. congensiswere^ ^ ^ T b a r t e r i ^ r . glabra, puberulousbeneath.FurthermoreIa l r e a d y . ^ f ^ ^ o f puberulous note 2),that Idonotassignany ^ ^ " ^ T c a n g e J ^ w f t o * * hairs.Nootherdifferential charactersbecomingevident,A inthesynonymy of A. heterophylla. ^
Meded. Landbouwhogeschool Wageningen 79-14(1979)

PHOT. 10. Anubias heterophylla ENGLER Infructescence. Phot. J. BOGNER, calidario M, 1978.-{Bogner 1300, WAG).

3. In 1915, ENGLERdescribed A. congensisvar. crassispadix. Thisvariety was described as having a much reduced 9 part. The original material (cultivated plants at Berlin-Dahlem)could not betraced. Theywerenotconserved or were possiblydestroyed inthewar.As1haveseennomaterialwithareduced9partof thespadix,described by ENGLER,aneotypefor thisvarietywasnot appointed. I consider var. crassispadix merely a monstruosity. However,thelengthofthe 9part isveryvariableinA. heterophylla,sometimes even in one individual. The specimens Bogner1299 and Bogner1300had the 9 part longer than the spathe.Inthesespecimens spadix/spathe ratio was 1.9 and 2.0. However,oneofthesespecimens(Bogner 1300) wascultivated intheWageningengreenhouseandflowered severaltimes(Crusio5). Thespadix/spatheratio proved tobevariable from 1.7 to 2.0,thedifference mainly duetothe length of the 9 part. In all other features, they were similar to other specimens of A. heterophylla. As thisfeatureprovedtobe variablealsowithinthesamespecimen, it was not consideredjustified to divide A. heterophyllain several infraspecific taxabasedonthelengthofthe9part.Therefore,A. congensisvar. crassispadixis hereplaced in the synonymy ofA.heterophylla.
40 Meded. Landbouwhogeschool Wageningen 79-14(1979)

i upwi^n,MMIInpiirJKI'HW **

"WW'

%*.4*ap^&

**

\ -r J
' * &

v. %.

"

PHOT. ILAnubias heterophylla ENCLER- Poller, REMmicrographW. BARTHLurr,uJJ-no M. 1979.- (Bogne/-7500,WAG).

4. A. affinis was described in 1907 by " ^ J * 7 ^ onlystated: 'Congo, s.loco'.However,from BR1recei ^ ^ affinis sp . without a collector's name or number. It was annou ^ ^ ^ M W]T nov.'. Furthermore, it was provided with a " a n n ^ w i i n D E WILDEMAN' stating that 'this specimen obviously s e ^ . ^ s p e c i m e n andthe drawing (1907).Iamnot surethat DEWITIScorrect,buta s t n e p ^ ^ ^ / t o j J ^ A in D E WILDEMAN both are, in my opinion, part o affinisin the synonymy of A. heterophylla. kH T h e typ e 5. Togetherwith,., affinis,DE WILDEMAN^ ^ ^ b u d f . Considering collection onlyconsisted ofthreeleavesandsomeim ^ . ^ t h a t DE the variability of the leaf-shape in Anubias, i d (the asym. WILDEMAN'S reason for describing &\^** areimmature,itisquite metricleaf)isinsufficient. Althoughthebudsonthe type he certain that A. engleribelongsinA. !ZZ/l^aScdoriBequaert6450,^ch 6. Anubiasbequaertimy/i^E^^l922)^^rt645obiscl^riy\x\ori&iri consists ofthree smallfloweringspecimens. J heterophylla. A. heterophylla.A. bequaertiisa small form o I A . , some t 7.M ^of this as( l ^ JACOB^ ^' C hlla_ ^ 1s e n1 m e0 ^ ^ O A N did Mr. N. ^ EN MN ^ specimens confirmed, that these plants belonged inA.
Meded. Landbouwhogeschool Wageningen 79-14(197

41

Specimens examined:
CAMEROON: S.loc. (fl.) Bogner1299(WAG); ibid, (fl.) Bogner 1300(WAG); Piste AkwayaMamfe,presNyang,25 kmNNEMamfe(fl.,fr.July)Letouzey14145 (BR,K,P, WAG); Kumba (fr. Jan.) Paysan s.n. (WAG). EQUATORIALGUINEA: Bebar, Camposgebiet (fl. Dec.) Tessmann 697(K). GABON: S.loc. (fl., fr.) Bogners.n. (M); Gamba (fl. Sept.) Bretekr 5605 (WAG); Chute de Tchimbele(fr. Aug.)Bretekr &De Wilde38 (WAG);Lonkandon (fr.Jan.)Le Testu 1278 (BM,P). CONGO: Fouet, surletracedela route delaMudongo,WSibiti(fr. Aug.) Farron 4357(P). CABINDA: Beligen Lualy-Maiombe(fr. Dec.) Gossweiler 7609(BM,COI,K: thissheet indicating 'Portugese Maiombe,Chiluango'). ZAIRE:Between MasisiandWalikale(fl.Jan.)Bequaert6450(BR,holotypeofA. bequaerti);ibid, (fl. Jan.) Bequaert 6450bis (BR);between Walikale and Lubutu (fl., fr. Jan.) Bequaert6590(BR); pont sur la Mpioka entre Zundu et Timasi, Terr. Ngombe-Matadi (fr. Dec.) Breyne 2236(BR); Kisantu (fr. Mar.) Callens 75 (BR);Irangi, 10(or 70?)kmW of L. Kivu (fl., fr. Aug.) Cambridge Congo Expedition 1959, 300(BM, BR); Mvuazi (fr. Jan.) Devred448 (BR); M'vuazi-Ngombe (fr. Oct.) Devred803(BR);Yanonghe(st. Mar.) Germain250(BR);betweenTumba and Kinpesri (fl.) Gillets.n. 1903 (BR);M.Pioka(budsOct.)Laurents.n., 1895 (BR,holotypeofA. engleri;isotypeat B); betweenWalikaleandKaleke(Kivu)(fr.Mar.)Lebrun5296(BR); Orega(Mamtma)(fl.,fr.July) Lebrun 5659 (BR); Kembe, Terr. Walikale (fl., fr. Nov.) Leonard 1581 (BR); Kampala, Terr. Walikale (buds Nov.) Leonard 1646 (BR);Bunyakiri Terr., Kalehe (fr. Feb.) Leonard2856 (BR); Dundusana (fl., fr. Sept.) Mortehan513 (BR); Mingazi, Terr. Kaleke, km 85 route KavumuWalikale(fr.Dec.)Pierlot 1194 (BR); Gimbi,Valleede laFuka(fr.Nov.)Toussaint644 (BR); Kivoe, Irangi,IRSAC-domein(fr. Mar.) Vanderveken 9718 (BR);Kizu(budsOct.) Vanderyst27072(BR). ANGOLA: Distr.Cazengo,inthemountainscalledMuxaulo(fr.Jan.) Welwitsch236 (BM,syntype of A. heterophylla); distr.Golungo Alto(fr. Mar.) Welwitsch237(K, lectotype of A. heterophylla; isotypes at B,BM,COI, LE,P);ibid.(st. Mar.) Welwitsch 238 (BM). UNKNOWN LOCALITY:Cultivated specimen?(fr.) Anonymus s.n. (BR,holotype of A. afflnis); (fl.) Bogner 1321 (WAG);(st.)deBrieys.n., 1913 (BR). CULTIVATED:Belgium,atBrussels(fl.,fr.June)Hort.Brux.juin 1902 (BR);Denmark, at Copenhagen(fl.) Jacobsens.n.(WAG);ibid, asA. 'undulatd (fl.) Jacobsen s.n.(WAG);Great Britain, at Kew(fl. May)Kew, entryno. 190/1904 (K);ibid. (fl.June)Kewjune 1914 (K);ibid.? from Boma,

10
MAP 11. Anubiasheterophylla.

10

20

30

42

Meded. Landbouwhogeschool Wageningen 79-14 (1979)

Zaireimported (st.Aug.)Bulls.n. 1887(K,syntypeofA. congensis);ibid.(fr.July)Bulls.n. 1889(K, lectotype of A. congensis);ibid. (fr. July) Bulls.n. 1894 (K, syntype of A. congensis); Germany, cultivated byA. BLASS,Munich (fl.,fr. May)Bogner1308(WAG);Netherlands,atWageningen(fl. Dec.)Arends71 (WAG); ibid. (fl. May) Crusio5(WAG,cultivated specimen ofBogner 1300);ibid, (fl. Aug. & Sept.) Crusio13(WAG); ibid. (fl. Sept.) Crusio 15 (WAG); ibid. (fl. May) J.V. 467 (WAG).

8. Anubiaspynaertii D E WILDEMAN
DEWILDEMAN, 1910: 168.

Fig.9,Phot. 12,Map12

Type: Paku, Zaire, Seret861(BR,lectotype). Rhizomecreeping,prostrateandrooting,sometimesbranched,\- \\ cmthick. Petiole 10-45cmlong,shortly sheathed,alittleshorterthanorupto1\ x as long as the leaf-blade, glabrous or minutely puberulous. Geniculum 1-2 cm long. . Leaf-bladehastate,longauriculate,cordatewithaverywidesinusorobtuseat thebase,glabrous above,midribandlateralnervesminutelypuberulousbelow, apexacuteoracuminate,middlelobeovatetooblong-ell.ptic,9-29cmlong,414cmbroad, almost twiceto morethan 3timesaslongasbroad,widestbelow the middle. Lateral lobes obtuse, ovate-elliptic, up to 18cm long, 7cmwide, sometimesalmostortotallyabsent.Numerousprominentlateralnerves,witnu5less prominent lateral nervesbetween them, withvery numerous,tran verse slenderveinletsbetweenthem.Lateralnervesneartheedgeanastomos.ngin 1-5 T Z l / T s l T n d e r , 0.4-1.1 x aslongasthepetiole,7-27cmlong,glabrousor
P

Xate lanceolate-elliptic,whenflowering spreadingbackward,after floweri i i i ^tr, inna acute-mucronu ate, white, green or onvc ing closing again, 2-3^ cm long, acute iu>-'u TadiX slender cylindrical, 2X-3Jcm long 1.2-L4 ^ ^ shortstipedorsubsessile.Spartmoreorless^nsely flowered, ?pa lessden* y flowered J part l i * as long asthe? part Pis.Is d ^ f ^ ^ some whitish spots, and avery short " ^ ^ X of the 4-6. Thecae oblong or elliptic. Sflowerswith theae cove^ng the * ^ synandrium entirely and alsothe largerpart ofthetop of hsyna thecae disappear afterfloweringand t e ^ ^ ^ T ^ U o . 9 - L S closetogether.Berriesdepressed-gbbose.Dneseeds whjtish spots. mm broad; 1.5-2.0 x aslongasbroad, light brownwn D i s t r i b u t i o n : Gabon, Congo,Zaire. * , . : On we,p b - i. or * ,he wa,e, a, ,he b - * - * - .
h

Jl/e(ferf. Landbouwhogeschool Wageningen 79-14(1979)

Fio. 9. AnubiaspynaertiiDEWILDEMAN: 1.lowersideleaf,j x ; 2.inflorescence, f x ; 3.upper side synandnum, 30 x ; 4. synandrium, side view, 30x ; 5. upper side synandrium, 30 x .- (1, 3. Seret861, BR;2.Seret861, BRand Crusio2, WAG;4-5. Van Tilborgs.n., 1914, BR).

*44

Meded. Landbouwhogeschool Wageningen 79-14(1979)

PHOT. 12. Anubiaspynaertii D E WILDEMAN

Inflorescence. Phot. J. W.
MUGGE, calidarioWAG,

25.IV.1978. - (Crush 2, WAG).

brooks, mostly onshadyplacesinforests; 20-1450m.Floweringand fruiting throughout theyear. Vernacular n a m e s : ^ / * ^ ^ 11934 12145 13474 15607, 15158); Batoie baNgwa(Turumbu, LouuUWO), 3793:nom generiquedesOrchideesetdes Fougeres). Note:Thisspeciesislongoverlooked. typical. Consequently,thespeciesstandsrelatrvelytsolatedwithinthegenus. Specimens examined:
WAG).IkoyBandja(fl.,fr.Oct.)arP

GABON:CapEsterias(fl.,fr.Ooi-^ g ner603(K MWAGM y ^ J ^ fr_^ (K,M,WAG);M'Voum(fl.Nov.)Bogneri 8 8 B ^ 7 ^ ' ^ J . mmgua(fl. Sept.)Breleler& De ! K ) ; 4kmWofLastoursville (fl., ^ ^ S K w i S S ; Be>"ga (buds, Nov.) fr. fl. Wilde574(WAG); Abauga, chantier CETA(buasJune, i?a//e3224(P);ibid,(budsJune) Halle3915 (P). p) CONGO: Mayombe,LesSaras,routedel ^ ^ 1 Feb)Zrd34,3 (BR,K); Befale (fl., fr ZAIRE: Bolumbuloko,EkekeliR. (Terr Be fate)budsFeb * > ^ ^ Y a n g a m b i and Apr.) W 4 f t f c (BR,K);KU(fl.Mar.)German 261 (BR), ^ Afeferf. Landbouwhogeschool Wageningen 79-14(1979)

10
MAP 12. Anubiaspynaertii.

Yakusu(budsNov.)Germain4660(BR);L.Yandja (Isangi)(fr. Aug.)Germain5162(BR); Wendji (env.ofCoquilhatville) (buds,fr. May) Lebrun308(BR);Yangambi,8kmN ofYaosuka (fl. Apr.) Louis3793(BR);Yalibwa,22kmNofYangambi(fl.June)Louis9925(BR);Yangole,20kmWof Yangambi (fr. Oct.) Louis11934 (BR, K, P); ibid, (buds, fl. Oct.) Louis 12145 (BM, BR, WAG); Lileko, between Yangambi and Basoko (fl., fr. Jan.) Louis 13474 (BR); Yangambi, valley of the Lusambila (buds Mar.) Louis14358 (BR); Yangambi, along the river Isalowe (fl., fr. Apr.) Louis 14635(BR, K); Yangambi, Island Booke wa Mbole (fl., fr. June) Louis 15158 (BR, WAG); Yangambi, rivegauche(fl., fr.July)Louis15607(BR);Eala(fr. June) Pynaert1437(BR,syntype); Paku (fl. June) Seret861(BR, lectotype); ibid, (fl., fr. June) Seret862(BR); Ingia (fl. July) Van Tilborgs.ni,1914(BR);Kisantu-Kwango(fl.) Vanderyst 16490(BR). CULTIVATED: Netherlands, at Wageningen (fl. Apr.) Crusio 2 (WAG, collected by BOGNER in Gabon).

ACKNOWLEDGEMENTS IwouldliketothankallmembersofthepersonnelintheLaboratory ofPlant Taxonomy and Plant Geography of Wageningen for their cooperation and hospitality.IacknowledgeespeciallyProf.Dr.H.C.D.DEWIT'Ssupervisionand manyvaluableadvices.Mr.G. BOELEMAassistedcorrectingthemanuscript and proofs Miss IKEZEVVALDskilfully prepared thedrawings.MissD.M.WASSINK typed the manuscript. Also Iamverygrateful to Mr.J. BOGNER,Botanischer Garten Munich,who providedmewithmanyfineherbariumandlivingmaterial,withmany beautiful photographs and color-slides and whose continuing interest in my work was highlystimulating.Thephotographs6and 11 werekindlyput atmydisposalby Mr.N.JACOBSEN(Copenhagen)and Dr.W. BARTHLOTT(Munich)respectively. Mr. F. MOHLMANN, Einbeck,G. F.R., Mr.N. JACOBSEN,University ofCopen46
Meded. Landbouwhogeschool Wageningen 79-14(1979)

hagen,and MissM. KNECHT,IvoryCoast,kindlyprovidedme withmanyliving specimens of Anubias. Their help facilitated andimproved therevision very considerably. ThisrevisionofAnubiasisthe 12thinstalmentofPrimitiaeAfricanae,ascries ofpapers, introducingthefirstresultsofresearch-workersinAfrican botanical taxonomy, prepared under supervision ofProf. Dr.H.C.D.on WIT. burlier instalments: 1. Acta Bot. Neerl. 5:171-178.1956. 2. Blumea 10:607-624.1960. 3. Acta Bot. Neerl. 11: 231-265.1962. 4. Blumea 12:209-239. 1964. 5. Acta Bot.Neerl. 13: 161-174.1964. 6. Bull.Jard. Bot.del'etat Bruxelles35:411-433.1965. 7. Journal S.Afr.Bot. 33:269-292.1967. 8. Acta Bot. Neerl. 19:227-248.1970. 9. Taxon 22:57-91. 1973. 10. Acta Bot. Neerl.24:69-82.1975. 11. Meded. Landb.hogesch.Wageningen 79(3):1-115. 19 n. LITERATURE
ANS.NK,J. 1977.EenAnubias geschiktvoor hetaquarium enhe,paludarium.He.Aquarium 48. BAILLON,H. 1895.Histoire desPlantes XIII,Pans. BENTHAM,G.&J.D.HOOKER F. 1883.Genera Plantarum 3,London. BERHAUT,J. 1967.FloreduSenegal2nded., Dakar. BROWN,N.E.1889.NeworNoteworthy Plants.Gard.Chror.be .s, . ^ BROWN,N. E. 1901. In:W.T.TH.SELTON-DYER Flora fTrop.calAfnca p ^ ^ CHEVALIER,A.1909.Plantes deI'Afrique trop.cale f r a n c a ^ [ ^ ' p J s CHEVALIER,A. 1920.Exploration botaniqueA f ^ ^ ^ , 2 , P a r i s .
ENOLER, A. 1879. in: A. DECANDOLLE, M o n o g r a p , a ^ K T S Gclehr.cn au* den

ENGLER, A. 1889. Araceae. Mitteilungen vonForschungsre.sen DeutschenSchutzgebieten(Danckelman)2. ENGLER,A. 1893.Beitragezur Floravon Afnka III. Bot.Jahrb.XV. ENGLER,A.1899.Araceae africanae II.Bot.Jhrb.XXVI. ENGLER,A.1915.Anubiadeae. DasPflanzenre.ch Hef:H U ' P | HEPPER, F.N. 1968a.Tropical African MonocotyWons 11.* c
HEPPER,F. N. 1968b.In: J. HUTCH.NSON&J. M. DALZ.EL,

i c a lAfri ca 2nded,.III.

part 2,London. YVII KewBull.1939. HUTCHINSON,J. 1939.Tropical Afncan PlantsX ^ ^ ^ HUTCHINSON,J.&J.M.DALZIEL. 1936.Florao ^ T ^ ^ p n a n e r o g a r o e s I.Brol. LEMEE,A. 1929.Dictionairedescriptifctsynonym.qu de G e n D a s Aquarium I . MOHLMANN, F.1977.Empfehlenswerte ^ f%H& F . WEXNHAM,Catalogueof.he
RENDLE,A. B. 1913. Araceae.In:A. B U N D L E
g

^ . ^

London.

plantscollectedby Mr.&Mrs"^ ^ J "ra Von Kamerun. DieAquancn undTcrrancn SADILEK, V. 1974.Kostbarke.ten der Wassemor Zeitschrift 27. v o n Kar nerun: I. Anubias nana Engler. D* SADILEK, V. 1978. Kostbarke.ten der Wassernor Aquarien undTerrarien Zeitschnft 31. 47 Meded. Landbouwhogeschool Wageningen 79-14(1979)

SADILEK,V. 1978. Kostbarkeiten derWasserflora vonKamerun: 2.Anubiaslanceolata. DieAquarien und Terrarien Zeitschrift 31. SCHOTT, H.W. 1857. Aroideen Skizzen.Oesterreichisches BotanischesWochenblatt 7. SCHOTT, H.W. 1858.Genera aroidearum, Wien. SCHOTT, H.W. 1860.Prodromus systematisAroidearum, Wien. WILDEMAN, E. DE. 1903. Etudes sur la Flore du Bas-et du Moyen-Congo. Annates du Musee du Congo Ser.V, I. WILDEMAN, E. DE. 1907. Etudes sur la Flore du Bas-et du Moyen-Congo. Annales du Musee du CongoSer. V,II. WILDEMAN, E. DE. 1910. Etudes sur la Flore du Bas- et du Moyen-Congo. Annales du Musee du Congo Ser. V, III. WILDEMAN, E. DE. 1922.Plantae Bequaertianae 1,fasc. II,Paris. WILDEMAN,E.DE&TH. DURAND. 1901. PlantaeGilletianaeCongolenses.Bull.Herb.Boiss.Ser.2,1.

INDEX The main entries of the correct taxon names are indicated by bold printed pagenumbers.Allfigures andphotographsareindicated byanasterisk attached totheir indexed pagenumbers.Synonymsareprinted initalics. Amauriella1,2, 34 - auriculata31 - hastifolia31 - obanensis 1, 2, 31, 34,35 -talbotii3l,34 Anubias 2, 34 - section Cylindranubias34 - section Synanubias34 -affinis 35* 41,42 - afzelii 1,2,4,5, 6*, 7*,8, 17,23, 30, 33,34,39 - auriculata1,27, 31,33,34,35 - barteri 1,2,4,8, 17, 18,20,30,34 - - var.angustifolia,stat.etcomb. nov. 3*, 10,11*,13, 14* - - v a r . barteri9*, 10,12, 16,19 - - var.caladiifolia 10,11*,13,15*,16, 17,19 - - v a r . glabra 10, 11*,12, 13,14,16, 18,20,39 - - var. nana, stat. et comb. nov. 10, 11*, 19,20 - bequaerti35,41,42 - congensis 35, 39 - var.crassispadix35,40
48

- engleri35, 41,42 - gigantea 1,2,4, 21, 22*,23*, 24,30,

31,33,34

- - var. tripartita21, 23, 24 - gilletii 1, 4,24,25*, 26*,27,33, 34, 37 -gracilis 1,4,28,29*,30 - hastaefolia31 - hastifolia 1,2,4,23,27,28,31, 32*, 34 - - var.robusta21,23, 24,31 - - var.sublobata31, 33 - haullevilleana 31, 33, 35 - heterophylla 1,2,4, 30, 34,35,36*, 37*,38*, 39*,40*,41*,42 - lanceolata14, 17, 18,19 - f. angustifolia 13,14 - var.cordata19 - Iaurentii3l,34, 35 - minima17,18 -nana 18, 19,20 - puberula 19 - pynaertii 1,2,4, 28, 33,34,43,44*, 45*, 46 - undulata 35,41,42
Meded. Landbouwhogeschool Wageningen 79-14(1979)