BOOK-ARACEAE

H EF D G B C J A Plate 27. Lasimorpha. A, base of plant showing stolons × 2/3; B, leaf × 2/3; C, inflorescence × 2/3; D, detail of spadix × 4; E, flower, tepals removed × 8; F, gynoecium, longitudinal section × 8; G, stamens, top view × 8; H, infructescence × 2/3; J, seed, side view × 5. Lasimorpha senegalensis: A, Bogner 691 (K & Kew spirit collection 29047.472); B, Leeuwenberg 1873 (K); C, Bogner 691 (Kew spirit collection 57526); D–J, Meikle 642 (Kew spirit collection 25024). 140 T H E G E N E R A O F A R A C E A E

niate; tepals 4(–5), free, fornicate. STAMENS: 4(–5), fila- veins mostly arising near petiole insertion, long-arcuate ments free or partially to completely connate, anthers towards division apex and running into margin, higher order partially exserted from tepals at anthesis, connective slen- venation reticulate. INFLORESCENCE: solitary. PEDUNCLE: der, thecae dehiscing by short apical slit. POLLEN: subequal to petiole and similar in appearance. SPATHE: ovate- monosulcate, ellipsoid, small (22 µm.), exine subreticulate, lanceolate, fully expanded, persistent, red brown. SPADIX: apertural exine psilate. GYNOECIUM: ovoid-ellipsoid to shorter or equalling spathe, cylindric, stipitate, stipe basally oblong, ovary 1-locular, ovules 4–6(–8), campylotropous, adnate to spathe, flowering sequence basipetal. FLOWERS: placenta single, ± prominent, parietal and basal, stylar bisexual, perigoniate; tepals 4–6, fornicate. STAMENS: 4–6, region short, somewhat narrowed, stigma discoid-hemi- free, filaments oblong, flattened, connective slender, thecae spheric. BERRY: irregularly globose, 1–4-seeded, red. SEED: ellipsoid, dehiscing by longitudinal slit. POLLEN: monosulcate, strongly curved, weakly strophiolate, testa hard, brown, ellipsoid-oblong, small (mean 18 µm.), exine scabrate or ver- warty to spiny, embryo rather large, curved, endosperm ruculate, minutely foveolate between verruculae. present but only as a few cell layers. See Plates 27, 112B. GYNOECIUM: cylindric to obovoid, ovary 1-locular, ovule 1, CHROMOSOMES: 2n = 26. anatropous, placenta parietal to subbasal, stigma discoid- DISTRIBUTION: 1 sp.; tropical west and central Africa:– hemispheric. BERRY: ovoid to ellipsoid, smooth, strongly Angola, Benin, Cameroon, Central African Republic, Chad, exserted at maturity, very prominent and thicker than spadix, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, red. SEED: curved, ± spherical, testa thin, hard, smooth, dark Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, brown, embryo curved, endosperm present but only as a few Nigeria, Senegal, Sierra Leone, Zaïre. cell layers. See Plates 28, 112C. ECOLOGY: tropical swamp forest, open wet areas; helo- CHROMOSOMES: 2n = 26. phyte along streams, in ditches and ponds, in forest gaps, DISTRIBUTION: 1 sp.; Malay Archipelago:– Indonesia often very abundant. (Sumatra), Malaysia (Sarawak, Peninsula). ETYMOLOGY: Lasia and Greek morphê (shape). ECOLOGY: tropical humid forest, usually peat swamp forest; TAXONOMIC ACCOUNTS: Engler (1911, as Cyrtosperma), helophyte, along streams. Knecht (1983), Hay (1988, 1992a). ETYMOLOGY: Greek pous, podos (foot) and Lasia; refers to the ± basal placentation of the ovule and the resemblance to Lasia. TAXONOMIC ACCOUNTS: Engler (1911), Hay (1988, 1992a). C 28. Podolasia Podolasia N.E. Brown Gard. Chron., ser.2, 18: 70 (1882). 29. Lasia C TYPE: P. stipitata N.E. Brown Lasia Loureiro, Fl. Cochinch. 64, 81 (1790). TYPE: L. aculeata HABIT: evergreen herbs, solitary or forming small clumps, rhi- Lour. (= L. spinosa (L.) Thwaites). zomatous, aerial stem erect to decumbent, internodes distinct, unarmed. LEAVES: several. PETIOLE: long, geniculate api- SYNONYM: [Lasius Hasskarl, Cat. Bogor. 59 (1844), cally, aculeate, spines either patent or pointing downwards, orth.var.] sheath short. BLADE: unarmed, sagittate to hastate or almost tripartite, coriaceous, posterior divisions ± equalling anterior, HABIT: clump- and colony-forming evergreen herbs, sometimes longer; basal ribs well-developed, primary lateral stoloniferous, stem thick, aculeate or unarmed (L. concinna), erect to decumbent, green, epigeal or submersed, internodes 28. Podolasia 29. Lasia L A S I O I D E A E : L A S I A 141

C DE A G FB Plate 28. Podolasia. A, habit × 2/3; B, detail of spadix × 5; C, flower × 10; D, flower, nearside tepals removed × 10; E, gynoecium, longi- tudinal section × 10; F, infructescence, × 2/3; G, seed, side view × 3. Podolasia stipitata: A, Corner 30898 (K); Kunstler (Dr King’s collector) 5499 (K); Parris 10979 (K); B, Corner 30898 (K); C–G, Parris 10979 (K). 142 T H E G E N E R A O F A R A C E A E

K J F B H AD G EC Plate 29. Lasia. A, leaf × 2/3; B, leaf × 2/3; C, leaf and base of plant with adventitious shoots, large portion of petiole removed × 2/3; D, habit × 1/15; E, inflorescence × 2/3; F, tranverse section of spiral portion of spathe × 2/3; G, detail of spadix × 5; H, flower, longitudinal sec- tion × 7; J, infructescence × 2/3; K, seed, side view × 5. Lasia spinosa: A, Cult. Kew (K); B, Hooker & Thomson s.n. (K); C, Clarke 7934 (K); Bogner 428 (Kew spirit collection 46984); D, Forman & Blewett 975 (Kew slide collection); E–H, Mayo 133 (Kew spirit collection 47735); J–K, Hooker & Thomson s.n. (K); Boyce 340 (Kew slide collection). L A S I O I D E A E : L A S I A 143

relatively long or short. LEAVES: several. PETIOLE: long, aculeate, weakly geniculate apically, sheath relatively short. BLADE: sagittate to hastate-sagittate when juvenile, adult blade deeply pinnatifid in anterior division, posterior divi- sions pedatifid, sometimes simple, adult blade rarely entire, or bipinnatifid (L. concinna), major veins aculeate on lower surface; primary lateral veins pinnate in anterior division, pedate in posterior divisions, primary lateral veins of each pinna forming submarginal collective vein in pinnatifid leaves, or running into marginal vein in simple leaves, higher order venation reticulate. INFLORESCENCE: solitary. PEDUN- CLE: subequal to petiole, aculeate as petiole. SPATHE: linear, very long and narrow (L. spinosa) or broader (L. concinna), very thick and spongy, spirally twisted, marcescent or decid- uous, basal part enclosing spadix, gaping at anthesis. SPADIX: shortly cylindric, obtuse, sessile. FLOWERS: bisex- ual, perigoniate; tepals usually 4, more rarely 6, fornicate. STAMENS: 4(–6), free, filaments broad, connective slender, thecae ellipsoid, dehiscing by longitudinal slit. POLLEN: monosulcate, ellipsoid, medium-sized (mean 27 µm.), exine reticulate, psilate along aperture margins. GYNOECIUM: ovoid to ellipsoid, ovary 1-locular, ovule 1, anatropous, funi- cle very short, placenta apical, stylar region well developed, shortly attenuate to cylindric, stigma discoid-hemispheric. BERRY: borne in cylindric infructescence, crowded, quad- rangular, apically densely muricate to spinose or smooth, 1-seeded, green. SEED: large, compressed-obovoid, testa 30. Urospatha thin, brown, hard, somewhat rugose, embryo large, some- what curved, endosperm present but only as a single cell layer. See Plates 29, 112D. apex and running into marginal vein, higher order venation CHROMOSOMES: 2n = 26. reticulate. INFLORESCENCE: solitary, rarely 2 in each floral DISTRIBUTION: 2 spp.; tropical southeast Asia, Malay sympodium. PEDUNCLE: equal to or longer than leaf, sim- Archipelago:- Bangladesh, Brunei, Burma, Cambodia, China ilar in appearance to petiole. SPATHE: erect, persistent, (Guandong, Guangxi, Hainan, Hong Kong, Taiwan, convolute below, gaping above, apically long-acuminate, Yunnan), India, Indonesia, Laos, Malaysia (Borneo, usually spirally twisted, sometimes flattened. SPADIX: Peninsula), Nepal, Papua New Guinea, Singapore, Sri Lanka, shortly stipitate or sessile, usually much shorter than spathe, Thailand, Vietnam. cylindric to subcylindric, obtuse. FLOWERS: bisexual, ECOLOGY: tropical humid forest; helophytes, wet places in perigoniate; tepals 4–6, fornicate. STAMENS: 4–6, free, fila- forest, open swamps, along streams, rice fields, tidal flats. ments broadish, a little compressed, connective slender, ETYMOLOGY: Greek lasios (shaggy), refers to the densely thecae ellipsoid, dehiscing by apical slit. POLLEN: mono- spiny stems. sulcate, ellipsoid, medium-sized (mean 26 µm., range 25–28 TAXONOMIC ACCOUNTS: Engler (1911), Sivadasan (1982), µm.), exine subreticulate, subrugulate to reticulate, apertural Hay (1988, 1990b, 1992a). exine psilate. GYNOECIUM: ovoid, ovary 1–2-locular, ovules (1–)2–4 or more per locule, anatropous, placenta C 30. Urospatha axile in 2-locular ovaries, basal in 1-locular ovaries, stylar region a little narrower than ovary, stigma broad, circular, Urospatha Schott, Aroideae 3 (1853). LECTOTYPE: U. discoid. BERRY: obovoid, 1–5(–8)-seeded, green to green- sagittifolia (Rudge) Schott (“sagittaefolia ”, Pothos sagittaefo- ish yellow. SEED: curved, testa hard, brown, thickish, lia Rudge; see Nicolson 1967). strongly warty, or spiny or crested, embryo curved, endosperm present but only as a very thin layer about 2 SYNONYMS: Urophyllum K. Koch in Berliner Allg. cells thick. See Plates 30, 113A. Gartenzeitung 25: 173 (1857, non Jack ex Wallich 1824); CHROMOSOMES: 2n = 52. Urospathella Bunting in Phytologia 65: 391 (1988). DISTRIBUTION: ca. 10 spp. (T. Croat pers. comm.); tropical HABIT: robust to slender evergreen herbs, solitary to clump- America:– Brazil, Colombia, Costa Rica, ?Ecuador, French forming, rhizome subterranean, horizontal or vertical, Guiana, Guatemala, Guyana, Nicaragua, Panama, Peru, spongy. LEAVES: few, basal, ± erect. PETIOLE: long, spongy, Surinam, Venezuela. smooth or scabrid-verrucose, sometimes angled, often varie- ECOLOGY: tropical humid forest and wetlands; helophytes, gated, apical geniculum absent or only very weakly open aquatic habitats, swamps, along streams, brackish water. developed, sheath long to short. BLADE: deeply sagittate or NOTE: Urospatha wurdackii (syn. Urospathella wurdackii) is subtripartite to hastate, posterior divisions usually longer notable for its small stature, linear-lanceolate leaves, and than anterior division, rarely blade lanceolate-linear and very slender spathe. lacking posterior divisions (U. wurdackii); basal ribs well- ETYMOLOGY: Greek “oura” (tail) and “spathê” (spathe). developed, primary lateral veins of both anterior and TAXONOMIC ACCOUNTS: Engler (1911), Bunting (1988, posterior divisions pinnate, long-arcuate towards division 1989a), Hay (1992a). 144 T H E G E N E R A O F A R A C E A E

N G M L K BA F D J HC E Plate 30. Urospatha. A, habit × 1/10; B, leaf × 2/3; C, inflorescence × 2/3; D, detail of spadix × 4; E, flower, longitudinal section × 10; F, infructescence showing persistent spathe × 2/3; G, seed, side view × 5; H, leaf × 2/3; J, leaf × 2/3; K, inflorescence × 2/3; L, diagrammatic longitudinal section of inflorescence to show partial spathe/spadix fusion × 2/3; M, detail of spadix × 4; N, flower, longitudinal section × 10. Urospatha sagittifolia : A, Mayo (Kew slide collection); U. sagittifolia: B, Jenman 5777 (K); C, Bogner s.n. (Kew spirit collection 29047.469); D–G, Bogner 580 (Kew spirit collection 29047.191); U. angustiloba : H, Spruce 3761 (K); U. wurdackii: J, Huber, Tillet & Davidse 3721 (K); K–N, Davidse, Huber & Tillet 17085 (K & Kew spirit collection 57275). L A S I O I D E A E : U R O S P A T H A 145

C VI. Subfamily Calloideae VII. Subfamily Aroideae C Subfamily Calloideae Endlicher, Gen. Pl. 239 (1837, Subfamily Aroideae “Callaceae”). Laticifers present (except Pistieae, Stylochaetoneae, Zamio- Laticifers present, simple, articulated; trichosclereids absent; culcadeae), usually simple and articulated, more rarely rooted aquatic, stem rhizomatous; leaves distichous; peti- anastomosing (in Caladieae, Colocasieae, Zomicarpeae); tri- ole not geniculate apically, sheath long; blade cordate, chosclereids absent; stem most frequently hypogeal, tuberous venation parallel-pinnate, uniformly fine; spathe fully or rhizomatous, less often aerial, rarely hemiepiphytic expanded, elliptic- to ovate-lanceolate, persistent; flowers climbers or epiphytes (Culcasieae, Philodendreae, bisexual, perigone absent, 3-merous; stamens 6 (or more), Syngonium), very rarely floating aquatics (Pistieae); petiole thecae dehiscing by longitudinal slit, pollen dicolpate; usually not geniculate apically (except Anubiadeae, Bognera, ovary 1-locular, ovules 6–9, anatropous, placenta basal; Culcasieae, Zamioculcadeae, rarely in Philodendreae and endosperm copious. Homalomeneae); higher order leaf venation more usually reticulate, less often parallel-pinnate (in Aglaonemateae, C 31. Calla Anubiadeae, Colocasieae, Dieffenbachia, Homalomeneae, Peltandreae, Philodendreae, Schismatoglottideae, Calla L., Sp. Pl. 968 (1753). TYPE: C. palustris L. Zantedeschieae); spathe usually differentiated into lower, convolute tube and upper, gaping blade; flowers unisexual, SYNONYMS: Provenzalia Adanson, Fam. 2: 469 (1763); perigone absent (except Stylochaetoneae, Zamioculcadeae); Aroides Heister ex Fabricius, Enum., ed. 2, p. 42 (1763); pollen inaperturate (except Zamioculcadeae and sometimes Callaria Rafinesque in Amer. Monthly Mag. & Crit. Rev. 2: 267 Stylochaetoneae). (1818). Laticifers present, simple, articulated. HABIT: seasonally Tribe Zamioculcadeae C dormant herb with repent or submersed, green, rhizoma- C tous stem, rooting at nodes. LEAVES: distichous. PETIOLE: Tribe Zamioculcadeae Engler in Nova Acta Acad. Leopold.- sheath long, with long, free, ligulate apex. BLADE: cordate Carol. 39: 141 (1876, “Zamioculcaseae ”). to broadly cordate, rounded, cuspidate-apiculate; primary lateral veins not differentiated, higher order venation par- Laticifers absent; seasonally dormant or evergreen allel-pinnate. INFLORESCENCE: solitary. PEDUNCLE: erect, (Zamioculcas), stem hypogeal; petiole geniculate; leaf blade as long or longer than petiole. SPATHE: fully expanded, compound; spathe convolute basally, blade reflexed at elliptic- or ovate-lanceolate, acuminate, shortly decurrent, anthesis; spadix subequalling spathe, female zone separated persistent, white within at anthesis, green without. SPADIX: from male by short, constricted zone of sterile flowers, male stipitate, cylindric, obtuse. FLOWERS: bisexual, often male zone cylindric to clavate, fertile to apex; flowers unisexual, at spadix apex, perigone absent. STAMENS: ca. 6, free, perigoniate; tepals 4, free, ± prismatic; stamens 4, sur- sometimes more, filaments somewhat flattened, anthers rounding ± clavate pistillode, filaments free, (connate in short, connective slender, short, thecae ellipsoid, oppo- Gonatopus), anther terminal, connective inconspicuous, site, dehiscing by longitudinal slit. POLLEN: diaperturate, pollen extended-monosulcate to fully zonate, extruded in globose, small (mean 23 µm.), exine foveolate, apertural strands; ovary 2-locular, ovules 1 per locule, hemiana- exine verrucate. GYNOECIUM: shortly ovoid, 1-locular, tropous, placenta axile to basal, style short, distinct, stigma ovules 6–9, anatropous, oblong, funicles short, placenta large, hemispheric-discoid; berries and seeds large, testa basal, stylar region attenuate, stigma small, subhemispheric. smooth, endosperm absent. BERRY: spheroid-conic, several-seeded, red. SEED: terete- oblong, testa thick, scrobiculate towards chalaza, 32. Zamioculcas sulcate-striate towards micropyle, raphe prominent, embryo axile, elongate, endosperm copious. See Plates 31, 113B. Zamioculcas Schott, Syn. Aroid. 71 (1856). TYPE: Z. loddi- CHROMOSOMES: 2n = 36, 54, 72. gesii Schott, nom. illeg. (Caladium zamiaefolium Loddiges, DISTRIBUTION: 1 sp.; circumboreal:– Austria, Belgium, Z. zamiifolia (Loddiges) Engler). Canada, China, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Germany, ?Hungary, ?Japan, Korea (N.), HABIT: seasonally dormant or evergreen herb with short, Netherlands, Norway, Poland, Romania, Russia, Slovak very thick rhizome. LEAVES: few to many, erect, leaflets Republic, Slovenia, Sweden, Switzerland, Turkey, USA. deciduous during dormancy leaving persistent petiole. PETI- ECOLOGY: in forest swamps, often between forest margins OLE: terete, base greatly thickened and succulent, geniculate and raised bogs, often with Sphagnum, up to 1270m alt.; at apex, sheath ligulate, free almost to the base, very short, creeping helophyte in streams and ponds. inconspicuous. BLADE: pinnatisect, leaflets oblong-elliptic, ETYMOLOGY: classical name, first used by Pliny for two dif- thickly coriaceous, capable of rooting at base once shed and ferent kinds of aroids; perhaps from the Greek kallos forming new plants; primary lateral veins of each leaflet pin- (beauty). nate, running into marginal vein, higher order venation TAXONOMIC ACCOUNTS: Krause (1908), Dudley (1937), reticulate. INFLORESCENCE: 1–2 in each floral sympodium, Huttleston (1953), Riedl (1977–1979), Topic & Ilijanic (1989). held at ground level. PEDUNCLE: very short. SPATHE: entirely persistent to fruiting stage, slightly constricted between tube and blade, green without, whitish within, tube convolute, blade longer than tube, expanded and horizontally reflexed 146 T H E G E N E R A O F A R A C E A E

B F CD AE Plate 31. Calla. A, habit × 2/3; B, spadix × 4; C, flower × 15; D, gynoecium, longitudinal section × 15; E, infructescence × 2/3; F, seed, side view × 10. Calla palustris: A, Bogner s.n. (Kew spirit collection 58031, 58904); B–F, Bogner 2117 (Kew spirit collection 57571). 31. Calla C A L L O I D E A E : C A L L A 147

C B D A E HJ KL G QP NF M Plate 32. Zamioculcas. A, habit × 1/3; B, leaflet × 2/3; C, leaflet with adventitious tuberlet × 2/3; D, first leafy shoot from adventitious tuber- let × 2/3; E, base of leaf showing swollen petiole × 2/3; F, inflorescence × 1; G, spadix × 1; H, male flower, nearside tepal removed × 8; J, detail of male flowers × 5; K, sterile flower, nearside tepal removed × 8; L, detail of sterile flowers × 5; M, gynoecium, longitudinal section × 8; N, detail of female flowers × 5; P, infructescence, longitudinal section × 1; Q, seed, side view × 6. Zamioculcas zamiifolia: A–M, Bogner s.n. Cult. Kew 1967–49401 (Kew spirit collection 49776, 29047.131 & 58712); P–Q, Faden & Faden 77/377 (K). 148 T H E G E N E R A O F A R A C E A E

32. Zamioculcas SYNONYMS: Heterolobium A. Peter, Nachr. Ges. Wiss. Göttingen, Math.-Phys. Kl. 1929 (3): 211, 221 (1930); at anthesis. SPADIX: sessile, female zone subcylindric, sepa- Microculcas A. Peter, Nachr. Ges. Wiss. Göttingen, Math.- rated from male zone by short constricted zone bearing sterile Phys. Kl. 1929 (3): 212, 222 (1930). flowers, male zone cylindric, ellipsoid to clavate, fertile to apex. FLOWERS: unisexual, perigoniate; tepals 4, in two HABIT: Seasonally dormant herbs, stem subterranean, a sub- whorls, decussate. MALE FLOWER: tepals subprismatic, apex globose tuber or a cylindric, horizontal rhizome. LEAF: thickened, stamens 4, free, shorter than tepals, filaments free, solitary, rarely pilose or scabrous, preceded by lanceolate oblong, thick, somewhat flattened, anthers introrse, connec- cataphylls. PETIOLE: geniculate basally or centrally. BLADE: tive slender, thecae ovate-ellipsoid, dehiscing by apical slit, usually trisect, rarely not (G. petiolulatus), primary divisions pistillode clavate, equalling tepals. POLLEN: extruded in trifid to trisect or pinnatifid, or pinnatisect to quadri-pinnat- strands, extended monosulcate to perhaps fully zonate, ellip- ifid, pinnae geniculate at junction with rachis, ultimate lobes soid, large (mean 60 µm.), exine thick, fossulate-foveolate, varying from linear to broad-elliptic, often decurrent; primary apertural exine verrucate. STERILE FLOWERS: each consisting lateral veins of each lobe pinnate, forming arching submar- of 4 tepals surrounding a clavate pistillode. FEMALE FLOWER: ginal collective vein, higher order venation reticulate. tepals strongly thickened apically, staminodes lacking, gynoe- INFLORESCENCE: 1–4 in each floral sympodium, appearing cium equalling tepals, ovary ovoid, 2-locular, ovules 1 per before or with leaves, subtended by several cataphylls. locule, hemianatropous, funicle very short, placenta axile PEDUNCLE: erect, very short to long. SPATHE: constricted near base of septum, stylar region attenuate, stigma large, between tube and blade, tube convolute, subglobose, cylin- discoid-capitate. BERRY: depressed-globose with furrow at dric or suburceolate, blade oblong to elliptic, reflexed at septum, 1–2-seeded, surrounded by persistent tepals, white, anthesis, marcescent. SPADIX: subequal to spathe, female infructescence ellipsoid. SEED: ellipsoid, testa smooth, brown, zone subcylindric, separated from male zone by very short, raphe conspicuous, embryo large, rich in starch, endosperm constricted zone of sterile flowers, male zone longer than nearly absent, present only as a few cell layers at chalazal end. female, cylindric to clavate, fertile to apex. FLOWERS: uni- See Plates 32, 113C. sexual, perigoniate; tepals 4(–6), in 2 decussate whorls, CHROMOSOMES: 2n = 34. fleshy, truncate to ± cucullate. MALE FLOWER: stamens with DISTRIBUTION: 1 sp.; tropical east and subtropical southeast connate filaments forming tube around central, cylindric to Africa:– Kenya, Malawi, Mozambique, South Africa (Natal), clavate pistillode, often exserted above tepals at anthesis, Tanzania (incl. Pemba, Zanzibar), Zimbabwe. connective slender, thecae dehiscing by apical pore. ECOLOGY: tropical moist forest, savannas; geophytes on POLLEN: extruded in strands, extended monosulcate or fully forest floor or in stony ground. zonate, hamburger-shaped, large (mean 76 µm., range 73–79 ETYMOLOGY: Zamia (genus of cycads) and Greek koloka- µm.), exine thick, foveolate, the foveolae scattered or sia (from Middle Eastern “qolqas”); perhaps in reference to the grouped in fossulae, apertural exine psilate to ± verrucate. pinnately compound leaves, in fanciful comparison to Zamia. FEMALE FLOWER: usually lacking staminodes, exceptionally TAXONOMIC ACCOUNTS: Engler (1905), Obermeyer & Strey 1 staminode present, ovary 2-locular, ovules 1 per locule, (1969), Mayo (1985a). anatropous, funicle very short to almost absent, placenta basal-axile, stylar region thick, somewhat attenuate, stigma large, discoid-hemispheric. BERRY: ovoid-ellipsoid, 1–2- seeded, red or orange to yellow, or whitish. SEED: ovoid-ellipsoid, testa thin, smooth, embryo large, plumule lateral, superficial, endosperm absent. See Plates 33, 113D. C 33. Gonatopus 33. Gonatopus Gonatopus J.D. Hooker ex Engler in A. & C. De Candolle, Monogr. Phan. 2: 208 (1879). TYPE: G. boivinii (Decaisne) Engler (Zamioculcas boivinii Decaisne). Z A M I O C U L C A D E A E : G O N A T O P U S 149

B EE DD D E M WY F Z X AA A V FF C K TU GG H S N P R Q CC J G L BB Plate 33. Gonatopus. A, habit × 1/2; B, stamens, perigone removed × 5; C, detail of male flower × 5; D, gynoecium, longitudinal section × 5; E, infructescence × 2/3; F, seed, front view × 5; G, habit × 1/2; H, stamens, perigone removed × 5; J, detail of female flower × 5; K, gynoecium, longitudinal section × 5; L, habit × 1/2; M, leaflet × 1; N, inflorescence × 1; P, detail of male flowers × 2; Q, detail of female flow- ers × 2; R, stamens, perigone removed × 5; S, detail of male flower × 5; T, gynoecium, longitudinal section × 5; U, single tepal, three quarter view × 5; V, habit × 1/2; W, stamens, perigone removed × 5; X, male flower × 5; Y, gynoecium, longitudinal section × 5; Z, infructescence × 2/3; AA, seed, side view × 2; BB, habit × 1/2; CC,, inflorescence, lower part of spathe removed to display spadix; DD, stamens, perigone removed × 5; EE, single tepal, three quarter view × 5; FF, male flower × 5; GG, gynoecium, longitudinal section × 5. Gonatopus angustus: A, Nuvunga & Conjo 385 (K); B–D, Cult. Kew 1971–02232 (Kew spirit collection 45233); E, Mogg 30003 (K); F, Bogner s.n. (Kew spirit col- lection 45229); G. petiolulatus: G, Vollesen 4788 (K); H–K, Bogner 241 (Kew spirit collection 37580); G. clavatus: L–M, Milne-Redhead & Taylor 7670 (K & Kew spirit collection 22034); N–U, Milne-Redhead & Taylor 7670 (Kew spirit collection 22034); G. marattioides: V–AA, Bogner 247 (Kew spirit collection 45222 & 52143); G. boivinii: BB, Bullock 1323 (K); CC–GG, Bogner 127 (Kew spirit collection 29047.481 ). 150 T H E G E N E R A O F A R A C E A E

CHROMOSOMES: 2n = 34, 68. bisexual flowers, male zone fertile to apex. FLOWERS: uni- DISTRIBUTION: 5 spp.; tropical east and subtropical south- sexual, perigoniate, borne in single basal whorl or in spirals; east Africa:– Kenya, Malawi, Mozambique, South Africa perigone a single cup-like (urceolate) structure. MALE (Natal, Transvaal), Tanzania (incl. Pemba, Zanzibar), Zaïre, FLOWER: perigone often with very thick, fleshy margins, Zambia, Zimbabwe. stamens 2–7, free, filaments filiform, long, rarely much thick- ECOLOGY: tropical evergreen forest; geophytes, on forest ened apically, connective slender, sometimes slightly floor, also in rocky crevices with humus deposits. thickened, thecae oblong, dehiscing by longitudinal slit, pis- NOTE: The leaf blade of G. petiolulatus, in which the lower tillode central, cylindric to conoid, sometimes absent. pinnae are reduced to irregular, linear rudimentary leaflets, is POLLEN: inaperturate, or sometimes vestigially monosulcate, intermediate in form between Gonatopus and Zamioculcas. In ellipsoid, large (mean 53 µm., range 43–58 µm.), exine fove- G. boivinii, the fallen leaflets may root and form new plants. olate-reticulate or subreticulate. FEMALE FLOWER: perigone ETYMOLOGY: Greek gony, gonatus (knee) and pous, podos usually greatly thickened and sticky-glandular or farinaceous (foot); in reference to the petiole geniculum in G. boivinii. on upper surface, ovary 1–4-locular, ovules 1–many per TAXONOMIC ACCOUNTS: Engler (1905), Obermeyer & locule, anatropous, placenta basal, parietal or axile, stylar Bogner (1979), Mayo (1985a). region thick, ± cylindric, exserted beyond perigone, stigma capitate to broadly discoid and massive. BERRY: borne at or below ground level in globose to cylindric infructescence, C Tribe Stylochaetoneae often rugose, fleshy, 1–few-seeded. SEED: ovoid to ellip- soid, slightly compressed, testa black, thin, costate, embryo Tribe Stylochaetoneae Schott, Syn. Aroid. 132 (1856, axile, elongate, endosperm copious. See Plates 34, 114A. “Stylochitoneae”). CHROMOSOMES: 2n = 28, 56. Laticifers absent; stem hypogeal, rhizomatous, roots often DISTRIBUTION: 17 spp.; tropical and southeast subtropical very thick, fleshy; higher order leaf venation reticulate; spathe Africa:– Angola, Benin, Burkina Faso, Cabinda, Cameroon, tube margins connate; spadix mostly hidden within spathe, Central African Republic, ?Chad, Equatorial Guinea (Bioko, fertile to apex; flowers unisexual, perigoniate; perigone a Rio Muni), Ethiopia, Gabon, Ghana, Guinea, Ivory Coast, single, cup-like (urceolate) structure; male flower usually Kenya, Liberia, Malawi, Mali, Mozambique, Nigeria, Senegal, with central peg-like pistillode, stamen filaments long, slen- Sierra Leone, Somalia, South Africa (Transvaal, Natal), Sudan, der, filiform, anthers terminal, dehiscing by longitudinal slits, Swaziland, Tanzania, Togo, Uganda, Zaïre, Zambia, Zim- connective slender, pollen sometimes monosulcoidate, oth- babwe. erwise inaperturate; ovary 1–4 locular, ovules 1–many per ECOLOGY: tropical humid forests (evergreen species, e.g. S. locule, anatropous, placenta basal, parietal or axile, style zenkeri), tropical savannas and deciduous forests (seasonally thick, exserted from perigone, stigma capitate to thickly dis- dormant species, e.g. S. natalensis); geophytes. coid; endosperm copious. NOTES: Engler (1920a) recognized 2 sections:– sect. Stylochaeton (syn. sect. Cyclogyne) and sect. Spirogyne. ETYMOLOGY: Greek stylos (style) and chitôn (tunic); refers C 34. Stylochaeton to connate perigone surrounding gynoecium. TAXONOMIC ACCOUNTS: Engler (1920a), Knecht (1983), Bogner (1984c), Mayo (1985a), Ntépé-Nyame (1988), Malaisse Stylochaeton Leprieur in Ann. Sci. Nat., ser. 2, Bot., 2: 184 & Bamps (1994). (1834). TYPE: S. hypogeum Leprieur SYNONYM: [Stylochiton Schott, Aroideae 10 (1855), orth.var.]. Laticifers absent. HABIT: seasonally dormant or evergreen 34. Stylochaeton herbs, rhizome subterranean, horizontal to erect, sometimes stoloniferous, roots thick, spindle-shaped, often very fleshy, almost tuberous. LEAVES: 1 to several, cataphylls often con- spicuously mottled and apically auriculate, sometimes persistent as fibrous mass. PETIOLE: sheath short to long, often purplish-spotted or banded. BLADE: lanceolate, ovate, ± rounded, cordate-sagittate, sagittate, hastate-sagittate or hastate; primary lateral veins pinnate or mostly arising basally, long-arcuate and running into marginal vein, lowermost pri- maries sometimes retrorse and then ascending, higher order venation reticulate. INFLORESCENCE: 1–4 in each floral sym- podium, appearing before or with leaves, borne at or partially below ground level. PEDUNCLE: short, much shorter than petiole. SPATHE: erect, marcescent; tube: margins connate, often ventricose at extreme base, sometimes constricted between a lower and an upper inflated zone, rarely entire spathe narrowly cylindric; blade: lanceolate-elliptic, ± gaping or opening only by narrow longitudinal slit, often much thickened. SPADIX: free, shorter than spathe, female zone densely flowered, sometimes contiguous with male zone, often separated by axis bearing a few (to many) sterile or S T Y L O C H A E T O N E A E : S T Y L O C H A E T O N 151

F D A J E K G B CH L Plate 34. Stylochaeton. A, habit × 1/2; B, flowering habit × 2/3; C, spadix × 2; D, habit × 1/2; E, habit × 1/2; F, infructescence × 1; G, habit × 1/2; H, spadix × 2; J, male flower × 8; K, female flower × 5; L, gynoecium, longitudinal section × 5. Stylochaeton lancifolius: A, Andrews 732 (K); B–C, Meikle 1294 (K & Kew spirit collection 27685); Schweinfurth 199 (K); Synge 220 (K & Kew spirit collection 58038); S. zenkeri: D, Letouzey 14413 (K); S. borumensis: E, Greenway & Kanuri 12751 (K); Verdcourt 3224A (K); S. bogneri: G, Bogner 145 (K); Lucas 253 (K); H–L, Bogner 148 (Kew spirit collection 7707). 152 T H E G E N E R A O F A R A C E A E

C Tribe Dieffenbachieae Tribe Dieffenbachieae Engler in Nova Acta Acad. Leopold.- Carol. 39: 148 (1876). SYNONYM: Tribe Bognereae Mayo & Nicolson in Taxon 33 (4): 689 (1984). Laticifers present, simple, articulated; terrestrial to helo- phytic, stem aerial, internodes distinct; spadix: female zone entirely adnate to spathe, laxly flowered, sterile male flow- ers (synandrodes) usually present between female and male zones; flowers unisexual, perigone absent; stamens connate into a truncate, prismatic synandrium, fused con- nectives strongly thickened, thecae lateral, pollen grains large; female flowers distant from one another, ovary 1–3 locular, ovules 1 per locule, anatropous, placenta basal or basal-axile. C 35. Dieffenbachia Dieffenbachia Schott in Wiener Z. Kunst 1829 (3): 803 (1829). TYPE: D. seguine (Jacquin) Schott (“seguinum”; Arum seguine Jacquin). SYNONYMS: Seguinum Rafinesque, Fl. Tell. 3: 66 (1837, “1836”); Maguirea A.D. Hawkes in Bull. Torrey Bot. Club 75: 635 (1948). HABIT: evergreen herbs, sometimes robust, stems erect to 35. Dieffenbachia decumbent, sometimes ± rhizomatous, rooting proximally, erect and unbranched distally, internodes distinct, green, FEMALE FLOWER: staminodes 4–5 in a whorl, white, clavate smooth, with conspicuous annular leaf scars. LEAVES: with rounded apices, spreading to erect, surrounding and numerous, forming an apical crown. PETIOLE: sheath more longer than gynoecium; ovary stout, subglobose to ovoid, than half as long as petiole or reaching blade. BLADE: thickwalled, 1–3-locular, locule walls bulging outwards giv- oblong-ovate, elliptic to oblanceolate, dark to light green or ing ovary distinctly lobed appearance when plurilocular, sometimes variegated with white, silver, yellow or various ovules 1 per locule, anatropous, placenta axile to basal, shades of green; midrib thick, sulcate or prominent on stylar region inconspicuous, stigma massive, almost as broad upper surface, primary lateral veins pinnate, sometimes to broader than ovary, 2–3-lobed or subhemispheric (when only weakly differentiated, running into margin, secondary unilocular), usually yellow, saturated with oily secretion at laterals parallel-pinnate, connected by transverse tertiary anthesis. BERRY: usually borne in arching infructescence, veins. INFLORESCENCE: (1–)2-several in each floral sym- berries globose to 2–3-furrowed, stigma remnants persistent, podium, cataphylls short and usually inconspicuous. 1–3-seeded, scarlet red to orange. SEED: globose to ovoid, PEDUNCLE: shorter than petiole. SPATHE: persistent, testa smooth, green to blackish green, embryo large, slightly or distinctly constricted between tube and blade, endosperm absent. See Plates 35, 114B. green, lower part convolute into a usually rather long, per- CHROMOSOMES: 2n = 34, 68. sistent tube which splits longitudinally in fruit, upper part DISTRIBUTION: 30 spp.; tropical and subtropical America, expanded into a short, erect or recurved blade. SPADIX: West Indies: Argentina (Corrientes, Misiones), ?Belize, slightly shorter than spathe, female zone entirely adnate to ?Bolivia, Brazil (Amazonia, Central West, South), Colombia, spathe, enclosed within tube, laxly flowered, separated Costa Rica, Dominican Republic, Ecuador, El Salvador, French from male zone by subnaked axis with a few, scattered Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Lesser sterile male flowers with reduced staminodes, rarely fertile Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto zones contiguous (D. humilis), male zone fertile to apex, Rico, Surinam, Trinidad, Venezuela. free, subcylindric, densely flowered, erect. FLOWERS: uni- ECOLOGY: tropical and subtropical humid forests, helophytes, sexual, perigone absent. MALE FLOWER: stamens 4–5, at edges of stream banks, or terrestrial in forest leaf litter. connate into a subsessile, rhomboid to hexagonal synan- ETYMOLOGY: named after J. Dieffenbach (1796–1863), head drium, truncate at apex, sulcate laterally, anthers lateral, gardener at the Imperial Palace of Schönbrunn, where Schott common connective thick, fleshy, thecae oblong-ellipsoid, was Director. dehiscing by short, apical, pore-like slit. POLLEN: extruded TAXONOMIC ACCOUNTS: Engler (1915), Young (1986). in strands, inaperturate, ellipsoid to oblong or nearly spher- ical, large (mean 79 µm., range 54–99 µm.), exine almost perfectly psilate to obscurely verruculate and/or sparingly punctate-foveolate to densely foveolate, rarely coarsely tuberculate (D. parlatorei). STERILE MALE FLOWERS: com- posed of a whorl of (3–)4–5(–6), ± flattened, irregularly globose-ellipsoid, sometimes ± connate staminodes. D I E F F E N B A C H I E A E : D I E F F E N B A C H I A 153

L B A H E K D J F GC Plate 35. Dieffenbachia. A, habit × 1/4; B, leaf × 1/3; C, inflorescence, nearside half of spathe removed × 2/3; D, detail of spadix showing sterile zone between male and female zones × 2; E, synandrium × 6; F, female flower × 6; G, gynoecium, transverse section × 6; H, infructes- cence × 2/3; J, habit, upper part of stem removed × 1/4; K, leaf × 1/2; L, detail of leaf venation × 5. Dieffenbachia seguine cv. Reginae: A, Cult. Kew 1950–48701; D. seguine: B, Bunting 2026A (K); D. seguine var. viridis: C–G, Cult. Kew 1973–13142 (Kew spirit collection 58927); D. seguine: H, Croat 65536 (Kew slide collection); D. elegans: J, Cremers 5730 (K); D. paludicola: K–L, Liesner 4006 (K). 154 T H E G E N E R A O F A R A C E A E

C 36. Bognera Bognera Mayo & Nicolson in Taxon 33: 690 (1984). TYPE: B. recondita (Madison) Mayo & Nicolson (Ulearum recon- ditum Madison). HABIT: evergreen herb with creeping rhizomatous stem, cataphylls marcescent, persistent and entire, drying red- dish-brown. LEAVES: erect, several. PETIOLE: shortly-sheathed, pilose, weakly geniculate at apex and base. BLADE: ovate, base rounded, apex acute; primary lat- eral veins pinnate, ca. 12 per side, glabrous adaxially, minutely pilose abaxially especially along veins, running into marginal vein, secondary laterals arcuate and ± paral- lel to primaries, finer veins reticulate. INFLORESCENCE: solitary, erect. PEDUNCLE: shorter than petiole, subtended by 3 cataphylls. SPATHE: erect, broadly ovoid, convolute in lower two-thirds, not constricted, apiculate, persistent. SPADIX: subequal to spathe, fertile to apex or most apical flowers sterile, female zone laxly flowered and entirely adnate to spathe, separated from male zone by very sparsely flowered sterile zone, male zone densely flowered. 36. Bognera FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4-androus, stamens connate into a synandrium, synandria truncate, irregularly rhombic, somewhat domed, usually with central slit, common connective very thick, thecae 37. Mangonia C ellipsoid-oblong, lateral, dehiscing apically by very broad slit. POLLEN: inaperturate, broadly ellipsoid, large (50–60 Mangonia Schott in Oesterr. bot. Wochenbl. 7: 77 (1857). µm.), exine scabrate to nearly smooth (psilate). STERILE TYPE: M. tweedieana Schott (“twedieana”). MALE FLOWERS: each usually consisting of 4 staminodes, those of lower flowers ± free and roundish, upper ones SYNONYMS: Felipponia Hicken in Anales Soc. Ci. Argent. nearly completely connate and irregularly formed, some- 84: 242 (1917), non Felipponea Brotherus (1912); times all 4 connate with a central slit. FEMALE FLOWER: Felipponiella Hicken in Darwiniana 2: 30 (1928). gynoecia arranged in somewhat distant spirals, ovary HABIT: seasonally dormant small herbs, tuber depressed-glo- depressed globose, 1-locular, ovule 1, anatropous, funicle bose, deeply embedded in soil. LEAVES: few to several. short, placenta basal, stigma sessile, discoid, broad. BERRY: PETIOLE: sheath long. BLADE: linear-elliptic or broadly ovate- unknown. SEED: unknown. See Plates 36, 114C. elliptic to oblong-sagittate; primary lateral veins pinnate, few, CHROMOSOMES: 2n = 34 (karyotype resembles that of running into margin, higher order venation reticulate. INFLO- Dieffenbachia). RESCENCE: solitary, appearing before the leaves. PEDUNCLE: DISTRIBUTION: 1 sp.; Brazil (Amazonas), ?Peru. fairly long, mostly subterranean. SPATHE: erect, not con- ECOLOGY: tropical humid forest (“terra firme”); terrestrial, stricted, lower part convolute into subcylindric tube, blade creeping in leaf litter on sandy soil. lanceolate to oblong, erect, gaping. SPADIX: subequal to NOTES: The mature shoot architecture seems to consist of spathe, sessile, female zone short, densely flowered, male repeating units of one small cataphyll, one larger cataphyll zone less densely to laxly flowered especially in basal portion, and one foliage leaf (T. Ray, pers. comm.). either contiguous with female or separated by short ± naked ETYMOLOGY: Named after Josef Bogner (born 1939). axis, apical third to half of spadix more densely flowered and TAXONOMIC ACCOUNTS: Madison (1980, as Ulearum covered with sterile male flowers. FLOWERS: unisexual, reconditum). perigone absent. MALE FLOWER: 2–5-androus, stamens with filaments connate, forming stipitate or sessile synandrium, C Tribe Spathicarpeae anthers free, connective inconspicuous, thecae ellipsoid to globose, dehiscing by a narrow slit or apical pore. STERILE Tribe Spathicarpeae Schott, Syn. Aroid. 214 (1856). MALE FLOWERS: synandrode stipitate, consisting of 3–5 sub- capitate, basally connate staminodes. POLLEN: inaperturate, Laticifers present, simple, articulated; seasonally dormant, ellipsoid, medium-sized (40 × 25 µm.), exine areolate to sub- stem tuberous, hypogeal; primary lateral veins of leaf or leaf rugulate. FEMALE FLOWER: gynoecium surrounded by whorl lobes forming single marginal vein (except Spathantheum of 3–4 clavate-spathulate staminodes, ovary globose to ovoid, and Spathicarpa), higher order venation reticulate; spathe 2–3-locular, ovules 2 per locule, anatropous, funicle short, unconstricted; spadix fertile to apex (except Mangonia); flow- placenta axile in upper part of septum, stylar region distinct, ers unisexual, perigone absent; male flower a synandrium of narrower than ovary, stigma 3-lobed or discoid, broader than partially or completely connate stamens (sometimes com- style, concave centrally. BERRY: depressed-globose, in dense, pletely free in Gorgonidium); in female flower the gynoecium subglobose to shortly cylindric infructescence. SEED: ellipsoid, surrounded by whorl of free staminodes or by cup-like synan- somewhat compressed, testa smooth, embryo and endosperm drode, ovules 1 per ovary locule (except Mangonia); embryo unknown. See Plates 37, 114D. relatively small, axile, endosperm copious. CHROMOSOMES: unknown. S P A T H I C A R P E A E : M A N G O N I A 155

B D E G F HJ CA Plate 36. Bognera. A, habit × 2/3; B, detail of leaf venation and trichomes on abaxial surface × 15; C, inflorescence, nearside lower half of spathe removed × 1; D, synandrium, top view × 8; E, synandrium, side view × 8; F, staminode, top view × 8; G, staminode, side view × 8; H, gynoecium × 8; J, gynoecium, longitudinal section × 8. Bognera recondita: A–B, Bogner 1995 (K); C–J, Bogner 1995 (Kew spirit collection 57525). 156 T H E G E N E R A O F A R A C E A E

F J Q D E G K H N L P AC B MS R Plate 37. Mangonia, A, habit showing inflorescence and petiole base only × 2/3; B, habit × 2/3; C, spadix × 1; D, synandrium, top view × 15; E, synandrium, side view × 15; F, synandrode × 15; G, gynoecium with associated staminodes × 15; H, gynoecium, longitudinal section × 15; J, infructescence × 1; K, leaf × 2/3; L, inflorescence × 1; M, spadix × 2; N, synandrium, top view × 15; P, synandrium, side view × 15; Q, synandrode × 15; R, gynoecium with associated staminodes × 15; S, gynoecium, longitudinal section × 15. Mangonia uruguaya: A, Darwiniana 18: 73, fig. 1,1; B, Darwiniana 18: 74, fig. 2,2; C–H, Felippone 5772 (SI); Felippone SI 297 (K, Kew spirit collection 58120 & SI); J, Darwiniana 18: 74, fig. 2,3 (1973); M. tweedieana: K, Waechter 2347 (K); L–S, Tweedie s.n. (K, LE); Waechter 2347 (K & Kew spirit collection 58090). S P A T H I C A R P E A E : M A N G O N I A 157

37. Mangonia 38. Taccarum DISTRIBUTION: 2 spp.; warm temperate South America:– and later deciduous (T. weddellianum). SPADIX: free or female Brazil (Rio Grande do Sul), Uruguay. zone adnate to spathe, sessile or stipitate, erect, much longer ECOLOGY: subtropical gallery forest; geophytes, stony, well than, subequal or shorter than spathe, male zone usually con- drained soils, flat areas, slopes near water courses. tiguous with female, rarely with a few bisexual flowers in NOTES: The first reliable record of M. tweedieana for Brazil between, fertile to apex. FLOWERS: unisexual, perigone absent. was recently made by Prof. Jorge Waechter (Waechter 2347 MALE FLOWER: 3–8-androus, stamens connate, synandrium [ICN, K!]), who also provided important new information very long-stipitate with apical whorl of anthers to short with contained in this generic treatment. anthers near base, stigmatoid apex inconspicuous or distinct ETYMOLOGY: Latin mango, mangonis (dealer, hence English and 4–6-lobed, or very large and dome-shaped, thecae oblong monger). or broadly ellipsoid, dehiscing by short apical slit or pore. TAXONOMIC ACCOUNTS: Engler (1920a), Herter (1943), POLLEN: extruded in strands, inaperturate, ellipsoid to oblong, Bogner (1973a). large (mean 63 µm., range 49–76 µm.), exine scabrate or ver- ruculate or spinulose-spinose, or smooth (psilate). FEMALE FLOWER: gynoecium surrounded by whorl of 4–6 free, erect, C 38. Taccarum filiform, clavate or oblong, often flattened staminodes, or by urceolate synandrode composed of connate staminodes (T. Taccarum Brongniart ex Schott in Oesterr. bot. Wochenbl. caudatum), ovary 3–6(–7)-locular, ovules 1 per locule, ana- 7: 221 (1857). TYPE: T. weddellianum Brongniart ex Schott tropous, funicle short, placenta axile, style very long and slender or very short to ± absent, always narrower than ovary, stigma SYNONYMS: Lysistigma Schott in Bonplandia 10: 222 thick, capitate or 5–7-lobed, lobes erect or stellate. BERRY: (1862); Endera Regel in Gartenflora 21: 226 (1872). borne in cylindric infructescence, depressed-globose, slightly HABIT: seasonally dormant herbs, often robust, tuber furrowed, stylar region persistent, 3–5-seeded. SEED: ellipsoid, depressed-globose. LEAF: solitary. PETIOLE: terete, mottled raphe conspicuous, testa granulate, light brown, embryo and variegated or not, sheath very short. BLADE: juvenile straight, elongate, endosperm copious. See Plates 38, 115A. leaves simply lobed, adult leaves subdracontioid, trifid to tri- CHROMOSOMES: 2n = 34. sect, anterior division usually deeply bipinnatifid, often DISTRIBUTION: 5 spp.; tropical and subtropical South tripinnatifid in lower pinnae, posterior divisions deeply pedat- America:– N. Argentina (Misiones), Bolivia, Brazil, Paraguay, ifid with segments themselves pinnatifid, ultimate lobes Peru. subtriangular to elliptic or lanceolate, acute to acuminate, ECOLOGY: open tropical woodland (cerrado), forests, humid broadly decurrent; basal ribs well-developed, primary lateral mountain valleys; geophytes, often in stony soils or between veins of ultimate lobes pinnate, running into inconspicuous rocks. marginal vein, higher order venation reticulate. INFLORES- ETYMOLOGY: Malay taka (name for Tacca in the Taccaceae) CENCE: 1, rarely 2 in each floral sympodium. PEDUNCLE: and Arum; refers to similarity of the leaf to that of Tacca leon- usually much shorter than petiole. SPATHE: not constricted, topetaloides. tube convolute, blade gaping to widely spreading, marcescent TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1989a). 158 T H E G E N E R A O F A R A C E A E

B J A L MN DC H EF GK Plate 38. Taccarum. A, leaf × 6; B, flowering habit, leaf removed × 1/8; C, spadix × 1/2; D, synandrium × 3; E, female flower × 3; F, gynoe- cium, longitudinal section × 3; G, gynoecium, tranverse section × 3; H, infructescence × 1/2; J, habit × 1/5; K, inflorescence, nearside half of spathe removed × 1/3; L, synandrium × 3; M, female flower × 3; N, gynoecium, longitudinal section × 3. Taccarum weddellianum: A, Pirani et al. 1946 (K); B, Cult. Hetterscheid (Kew slide collection); C–G, Bogner 458 (Kew spirit collection 29047.741); H, Harley 20436A (Kew spirit collection 29047.453); T. warmingii: J, Type drawing (Kew illustration collection); K–N, Cult. Kew 9 April 1890 (K & Kew spirit collection 58086). S P A T H I C A R P E A E : T A C C A R U M 159

C 39. Asterostigma short, placenta axile at base of septum, stylar region distinct, stigma usually deeply and stellately 3–5-lobed with each lobe Asterostigma F.E.L. Fischer & C.A. Meyer in Bull. Cl. Phys.- itself lobed, or 2–3-lobed with oblong, entire lobes. BERRY: Math. Acad. Imp. Sci. Saint Pétersbourg, ser. 2, 3: 148 (1845). subglobose to depressed-globose, deeply 4–5-sulcate, pale to TYPE: A. langsdorffianum F.E.L. Fischer & C.A. Meyer whitish green. SEED: oblong to ellipsoid, with swollen aril- like funicle, testa nearly smooth, embryo axile, elongate, SYNONYMS: Staurostigma Scheidweiler in Allg. endosperm copious. See Plates 39, 115B. Gartenzeitung 16: 129 (1848); Andromycia A. Richard in R. CHROMOSOMES: 2n = 34. de la Sagra, Hist. Fis. Cuba 11: 282 (1850); Rhopalostigmium DISTRIBUTION: ca. 7 spp.; tropical and subtropical South Schott in Oesterr. bot. Zeitschr. 9: 39 (1859); [Rhopalostigma America:– Bolivia, Brazil (Central, Northeast, Southeast, B.D. Jackson, Index Kew. 2: 713 (1895), orth. var., non R.A. South), Ecuador, Peru. Philippi (1860)]. ECOLOGY: tropical and subtropical humid forests; geo- HABIT: medium-sized, seasonally dormant herbs, tuber de- phytes, forest floor. pressed-globose. LEAVES: usually solitary. PETIOLE: smooth, NOTES: Engler (1920a) recognized 2 sections: sect. Astero- maculate, sheath very short. BLADE: usually pinnatisect, rarely stigma, sect. Rhopalostigma. entire, lobes oblong-lanceolate, acute to acuminate; primary ETYMOLOGY: Greek astêr (star) and stigma (a mark), refers lateral veins of ultimate lobes pinnate, running into marginal to the star-shaped stigmas. vein, higher order venation reticulate. INFLORESCENCE: 1–3 TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1969), in each floral sympodium, appearing with or before leaves, Bogner (in press). cataphylls often beautifully variegated. PEDUNCLE: rather long, similar to petiole. SPATHE: erect, hardly to not con- stricted, slender, persistent, lower part convolute into 40. Gorgonidium C narrowly cylindric tube, blade gaping to widely expanded at anthesis. SPADIX: female zone free or partly adnate to spathe, Gorgonidium Schott in Ann. Mus. Bot. Lugduno-Batavum 1: laxly flowered, contiguous with male zone, male zone longer, 282 (1864). TYPE: G. mirabile Schott fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium 3–4-androus, surrounding and con- HABIT: seasonally dormant herbs, tuber depressed globose. crescent with pistillode, subhexagonal to rounded, sometimes LEAF: solitary. PETIOLE: terete, sheath short. BLADE: pinnat- very shallow, umbonate at apex, shortly stipitate or sessile, ifid, pinnatisect or bipinnatifid, pinnae (9–)11–14, elliptic, entire thecae subglobose, distant, almost pendent from upper mar- or pinnatifid (with all intermediates), acute, upper lobes decur- gin of synandrium, dehiscing by transversely elongated pore. rent, lower ones sessile to shortly stalked; primary lateral veins POLLEN: inaperturate, ellipsoid to ellipsoid-oblong, medium- of ultimate lobes pinnate, running into margin, higher order sized (mean 36 µm., range 35–38 µm.), virtually psilate or venation reticulate. INFLORESCENCE: 1(–2) per floral sym- coarsely verrucate. FEMALE FLOWER: gynoecium surrounded podium, appearing before leaf. PEDUNCLE: much shorter than by cup-like synandrode, or by whorl of 3–5 short, some- petiole. SPATHE: erect or slightly fornicate, boat-shaped, very what thickened, subcuneate, truncate, often irregularly shortly convolute at base, not constricted, purple, persistent at connate staminodes, ovary pear-shaped to depressed-glo- least in lower part. SPADIX: sessile or stipitate (G. mirabile), bose, 3–5-locular, ovules 1 per locule, anatropous, funicle fertile to apex, female zone contiguous with and much shorter than male zone. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–7-androus, stamens free or connate to different degrees into a synandrium, connective slender, sometimes stipe-like between thecae (G. mirabile), otherwise inconspic- uous, thecae globular, sometimes remote from one another, dehiscing by apical pore or slit, pistillode present or absent, if present composed of 3–4 stylodia with slightly lobed or dis- coid-capitate apices (stigmatoids). POLLEN: inaperturate, ellipsoid, medium-sized (mean 34 µm, range 27–42 µm.), exine verrucose (G. mirabile) to retiverruculate, sometimes verrucae irregularly formed and flattened (G. vargasii). FEMALE FLOW- ERS: gynoecium surrounded by whorl of 6–8 filiform or subclavate staminodes, ovary subglobular to broadly ovoid, (2–)4–5(–7)-locular, ovules 1 per locule, orthotropous, ovoid, funicle half as long as ovule, placenta axile at base of septum, stylar region short to long, stigma 4–5-lobed or subcapitate. BERRY: globular to depressed-globular, blackish-purple to purple, sometimes upper surface somewhat warty (G. var- gasii), stigma and style remnants persistent. SEED: ovoid, globular to ellipsoid, testa brownish, smooth to rough, embryo elongate, endosperm copious. See Plates 40, 115C. CHROMOSOMES: 2n = 34. DISTRIBUTION: 3 spp.; Western and Andean South America:– N. Argentina, Bolivia, Peru. 39. Asterostigma ECOLOGY: tropical and subtropical forest or open places, between 900–3000m; geophytes, stony ground. 160 T H E G E N E R A O F A R A C E A E

B Q J A D G M F H R S L C KE PN Plate 39. Asterostigma. A, leaf × 1/2; B, detail of leaf venation × 5; C, base of plant × 1/2; D, juvenile infructescence enclosed by persis- tent spathe × 1/2; E, spadix × 2; F, female flower × 10; G, synandrium × 10; H, gynoecium, longitudinal section × 15; J, young plant × 2/3; K, spadix × 2; L, female flower × 10; M, synandrium × 10; N, female flower × 10; P, gynoecium, longitudinal section × 15; Q, infructescence × 1; R, leaf × 1/2; S, base of plant × 1/2. Asterostigma cryptostylum: A–H, Bogner 1237 (K & Kew spirit collection 48202); A. riedelianum: J–P, Harley et al. 18565 (K & Kew spirit collection 39166, 46592, 47693 & Kew slide collection); A. integrifolium: Q–S, Rawlins 204 (K). S P A T H I C A R P E A E : A S T E R O S T I G M A 161

K L A H D G J EB C F Plate 40. Gorgonidium. A, vegetative habit × 1/5; B, flowering habit, tuber longitudinally split; C, spadix × 2; D, synandrium × 8; E, female flower × 8; F, gynoecium, longitudinal section × 8; G, gynoecium, transverse section × 8; H, leaf × 2/3; J, infructescence × 2/3; K, synan- drium × 3; L, female flower × 3; Gorgonidium vermicidum: A, Bot. Jahrb. Syst. 109:550, fig. 23 (1988); B, Araque & Barkley s.n. (K); Aichinger 2 (Kew slide collection); C–G, Bogner 101(Kew spirit collection 45269); G. vargasii : H, Brunel 557 (K); J, Vargas s.n. (Kew spirit collection 49725); G. mirabile : K–L, Nicolson 3390 (K & Kew spirit collection 58918). 162 T H E G E N E R A O F A R A C E A E

40. Gorgonidium 41. Synandrospadix ETYMOLOGY: Greek Gorgo (the Grim One), -ides (descen- thecae dehiscing as above. FEMALE FLOWER: gynoecium dant) and -ion (diminutive); the name alludes to the filiform surrounded by 3–5 free, elongate-triangular, flattened, acute staminodia and stamens in G. mirabile, the appearance of staminodes, ovary ovoid-globose, 3–5-locular, ovules 1 per which recalls mythic Gorgo, whose hair was a mass of locule, orthotropous, funicle distinct, placenta axile-basal, writhing serpents. stylar region attenuate, stigma discoid-subcapitate, slightly TAXONOMIC ACCOUNTS: Bogner & Nicolson (1988). 3–5-lobed. BERRY: ± subglobose, 3–5-furrowed, 3–5-seeded, style persistent. SEED: rather large, ovoid to obnapiform, C 41. Synandrospadix testa ± rough, embryo elongate, straight, endosperm copious. See Plates 41, 115D. CHROMOSOMES: 2n = 34. Synandrospadix Engler in Bot. Jahrb. 4: 61 (1883). TYPE: DISTRIBUTION: 1 sp.; N. Argentina, Bolivia, Paraguay, Peru. S. vermitoxicus (Grisebach) Engler (Asterostigma vermitoxi- ECOLOGY: subtropical dry thorn forest; geophytes in clear- cum Grisebach). ings or on stony ground in shade. SYNONYMS: Lilloa Spegazzini, Pl. Nov. Crit. Argentina 3: ETYMOLOGY: Greek syn (together), anêr, andros (man) 10 (1897); [Synandriospadix Engler in Engler, Pflanzenreich and spadix (spadix). 73 (IV.23F): 49 (1920), orth. var.]. TAXONOMIC ACCOUNTS: Engler (1920a), Crisci (1971), HABIT: seasonally dormant herb, tuber subglobose. LEAVES: Croat & Mount (1988). several. PETIOLE: fairly long, with darker longitudinal stri- ations, sheath long. BLADE: ovate-cordate to -emarginate, 42. Gearum C somewhat glaucous; primary lateral veins pinnate, running into inconspicuous marginal vein, higher order venation Gearum N.E.Brown in J. Bot. 20: 196–197 (1882). TYPE: G. reticulate. INFLORESCENCE: 1, rarely 2 in each floral sym- brasiliense N.E.Brown podium, usually appearing with leaves. PEDUNCLE: short. SPATHE: ovate-lanceolate, boat-shaped, unconstricted, per- HABIT: seasonally dormant, tuberous herbs with aromatic sistent, convolute at base, gaping above, light green outside exudate when cut. LEAF: solitary. PETIOLE: sheath distinct, with dark longitudinal lines, inside purple and roughened. about half length of petiole. BLADE: subpalmatifid to peda- SPADIX: much shorter than spathe, subcylindric, obtuse, tisect, lobes or segments 5–9; primary lateral veins of each female zone partly or completely adnate to spathe, ± laxly lobe forming submarginal collective vein on each side, flowered, separated from male zone by a few bisexual or higher order venation reticulate. INFLORESCENCE: solitary. sterile flowers, male zone longer, fertile to apex. FLOWERS: PEDUNCLE: very short, much shorter than petiole. unisexual, perigone absent. MALE FLOWER: synandrium SPATHE: erect, slightly constricted, purplish, tube sub- 4–5-androus, long-stipitate, stipe elongate-conoid, whorl of cylindric, convolute, blade erect, gaping, oblong, cuspidate. anthers apical or shortly overtopped by central pistillode, SPADIX: sessile, shorter than spathe, fertile to apex, female thecae oblong-ellipsoid, dehiscing by longitudinal slit. zone shorter than male and separated from it by a sterile POLLEN: purple to pinkish, inaperturate, spherical to sub- zone of synandrodes. FLOWERS: unisexual, perigone spheroidal, medium-sized (mean 49 µm.), exine spinose. absent. MALE FLOWER: 4-androus, synandrium very shal- BISEXUAL FLOWERS: as for female but with whorl of 4–5 low, subrhombic, truncate, subpeltate or flat and not free stamens with flattened elongate-triangular filaments and subpeltate, thecae ± remote, situated near margin of synan- S P A T H I C A R P E A E : G E A R U M 163

A E D F GH J C K B Plate 41. Synandrospadix. A, habit × 1/4; B, juvenile plant × 1/2; C, leaf × 2/3; D, inflorescence × 2/3; E, spadix × 1; F, synandrium × 6; G, bisexual flower × 6; H, female flower × 6; J, gynoecium, longitudinal section × 6; K, infructescence × 1/2. Synandrospadix vermitoxicus: A, 184/10 (Kew slide collection); B, Cult. Kew 9 October 1891 (K); C, Araque & Barkley 440 (K); D–F, H–J, Bogner s.n. (Kew spirit collec- tion 56099); G, Burkart et al. 30412 (K); K, Badcock 815 (K). 164 T H E G E N E R A O F A R A C E A E

42. Gearum drium, globose to oblong-ellipsoid, dehiscing by apical pore or occasionally by an oblique slit. POLLEN: ellipsoid, inaperturate, large (mean 56µm, range 52–60 µm), exine smooth (psilate) to scabrous-granulate. STERILE MALE FLOWERS: composed of irregularly elongated, shallow, truncate synandrodes. FEMALE FLOWER: densely crowded, gynoecium surrounded by usually 4 staminodes, stamin- odes laterally compressed, thick, rounded to obovoid or subtrapezoid, ovary subglobose, 3–4-locular, ovules 1 per locule, orthotropous, rather elongate, placenta axile at base of septum, stigma subsessile, ± discoid, slightly (3–)4-lobed. BERRY: unknown. SEED: unknown. See Plate 42. CHROMOSOMES: unknown. DISTRIBUTION: ?2 spp.; Brazil (Central–West). ECOLOGY: tropical open herbaceous vegetation (cerrado , in campo limpo); geophytes in low-lying sites prone to periodic flooding. ETYMOLOGY: Greek gê (earth) and Arum. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner & Nicolson (1988), Mayo, Bogner & Boyce (1994). H D FG BC AE Plate 42. Gearum. A, leaf × 1/3; B, inflorescence × 1/3; C, spadix × 2/3; D, synandrium, top view × 8; E, synandrium, side view × 8; F, gynoecium with staminode, side view × 8; G, gynoecium, longitudinal section × 8. H, leaf × 1/3. Gearum sp .: A–G Araujo Dias 41 (HRB & Kew spirit collection 57663); Gearum brasiliense : H from Hatschbach 56028 (K). S P A T H I C A R P E A E : G E A R U M 165

C 43. Spathantheum suborthotropous, funicle distinct, placenta axile at base of septum, stylar region elongate-attenuate, stigma stellate, Spathantheum Schott in Bonplandia 7: 164 (1859). TYPE: 6–8-lobed, lobes subtriangular. BERRY: subglobose to S. orbignyanum Schott depressed-globose, 5–8-seeded. SEED: shortly ovoid to ± ellipsoid, testa ± smooth to slightly roughened, embryo SYNONYM: Gamochlamys J.G. Baker in Saunders Refug. elongate, endosperm copious. See Plates 43, 116A. Bot. 5: t. 346 (1873). CHROMOSOMES: 2n = 34. DISTRIBUTION: 2 spp.; Andean South America:– N. Argentina, HABIT: medium-sized, seasonally dormant herbs, tuber de- Bolivia, Peru. pressed-globose, sometimes large. LEAVES: solitary. ECOLOGY: tropical and subtropical uplands, mountain grass- PETIOLE: sheath very short. BLADE: outline ovate to ovate- lands, up to 2400m; geophytes on stony ground in shade, in cordate, entire or pinnatifid; primary lateral veins pinnate, deposits of humus on rocks. running into margin, higher order venation reticulate. ETYMOLOGY: Greek spathê, anthos (flower) and -eum (pos- INFLORESCENCE: usually 1, rarely 2 in each floral sym- sessing); refers to the fusion of the spadix to the spathe. podium, flowering before leaf. PEDUNCLE: short (S. TAXONOMIC ACCOUNTS: Engler (1920a), Crisci (1971), intermedium) to long (S. orbignyanum), slender. SPATHE: Bogner (in press). oblong-elliptic, boat-shaped, not constricted, cuspidate, at first convolute basally, then widely gaping at anthesis, green or purple (S. intermedium), entirely persistent. SPADIX: entirely adnate to spathe (S. orbignyanum) or male part free 44. Spathicarpa C (S. intermedium), slightly shorter than spathe, densely flow- ered, either female zone basal, contiguous with longer male Spathicarpa W.J. Hooker in Bot. Misc. 2: 146 (1831). TYPE: zone and spadix fertile to apex (S. intermedium, S. S. hastifolia W.J. Hooker orbignyanum), or female and male zones sometimes sep- arated by a zone of mixed male and female flowers (S. SYNONYMS: [Spaticarpa Schott in Oesterr. bot. Zeitschr. orbignyanum). FLOWERS: unisexual, perigone absent. 15: 34 (1865), orth. var.]; Aropsis Rojas Acosta in Bull. Acad. MALE FLOWER: 4–7-androus, stamens either connate into Int. Géogr. Bot. 28: 158 (1918). a long-stipitate, apically convex, peltate and shallowly 5–7- HABIT: small, seasonally dormant, sometimes evergreen lobed (stigmatoid) synandrium with anthers almost pendent herbs, tuber subterranean, rhizomatous, shortly oblong, hori- from thickened common connective, or filaments connate, zontal. LEAVES: several. PETIOLE: slender, sheath often rather anthers free and stigmatoids elongated, overtopping anthers long. BLADE: narrowly elliptic, oblong, ovate-emarginate, (S. intermedium), thecae linear-oblong and dehiscing by ovate-cordate, cordate, cordate-sagittate, auriculate-hastate longitudinal slit (S. orbignyanum) or subglobose and or strongly hastate to subtrisect; basal ribs short when pre- dehiscing by apical pore (S. intermedium). POLLEN: inaper- sent, primary lateral veins pinnate or mostly arising at petiole turate, ellipsoid-oblong, large (mean 54 µm.), exine nearly insertion, arcuate and running into margin or forming an psilate or shallowly uneven. FEMALE FLOWER: gynoecium irregular, submarginal collective vein, higher order venation surrounded by whorl of 5–8 terete-clavate, short stamin- reticulate. INFLORESCENCE: solitary, appearing with leaves. odes, ovary ovoid-ellipsoid, 5–8-locular, ovules 1 per locule, PEDUNCLE: relatively long, slender. SPATHE: not constricted, 43. Spathantheum 44. Spathicarpa 166 T H E G E N E R A O F A R A C E A E

J EF GH D K L M NP B C A Plate 43. Spathantheum. A, flowering habit × 2/3; B, juvenile leaf × 2/3; C, mature leaf × 2/3; D, inflorescence, nearside half of spathe removed × 2/3; E, synandrium × 4; F, female flower × 4; G, gynoecium, longitudinal section × 4; H, gynoecium, transverse section × 6; J, infructescence × 2/3; K, inflorescence, nearside half of spathe removed × 2/3; L, synandrium × 4; M, gynoecium with associated staminodes (= female flower) × 4; N, gynoecium, longitudinal section × 4; P, gynoecium, transverse section × 6. Spathantheum orbignyanum: A, Bang 1626 (K), Rauh 26024 (Kew spirit collection 37322); B, Bogner 900 (K); C, Cult. Kew 1883 (K); D–H, Rauh 26024 (Kew spirit collection 37322); Bogner 900 (K & Kew spirit collection 37369); S. intermedium: K–P, Munn 148 (Kew spirit collection 29047.761). S P A T H I C A R P E A E : S P A T H A N T H E U M 167

BC E K G D AF HJ Plate 44. Spathicarpa. A, habit × 2/3; B, synandrium and gynoecium with associated staminodes (= male and female flowers) × 10; C, gynoe- cium, longitudinal section × 16; D, leaf × 2/3; E, habit × 2/3; F, inflorescence × 2; G, detail of spadix × 6; H, synandrium and gynoecium with associated staminodes (= male and female flowers) × 10; J, gynoecium, longitudinal section × 16; K, infructescence, nearside half of spathe removed × 1. Spathicarpa burchelliana: A, Burchell 8335(K); Gardner 2447 (K); B–C, Cult. Kew 1981–388, Cristobal & Krapovickas s.n. (Kew spirit collection 45662); S. lanceolata: D, Balansa 579 (K); S. hastifolia (S. cf. gardneri ): E–K, Tressens et al. 1569 (K); Tressens et al. 3471 (K). 168 T H E G E N E R A O F A R A C E A E

oblong-lanceolate to narrowly elliptic or oblanceolate, acumi- HABIT: evergreen herbs, small to gigantic, stem repent to nate, fully expanded to subrevolute at anthesis, later closing, rhizomatous, climbing, arborescent or plant rosulate and persistent, green. SPADIX: entirely adnate to spathe, recurved acaulescent, internodes usually long, often short to very at anthesis, fertile to apex, ± laxly flowered, female flowers short, intravaginal squamules present, sometimes producing forming 2 outer longitudinal rows, enclosing 2 rows of male flagelliform shoots. LEAVES: numerous, small to gigantic, flowers, all in parallel. FLOWERS: unisexual, perigone absent. prophylls of mature stems caducous, marcescent and decid- MALE FLOWER: 3-4-androus, stamens connate, synandrium uous or persistent and membranaceous or decomposing to stipitate, apex formed by nectariferous, usually shallowly net-fibrous remains. PETIOLE: sometimes warty or covered lobed stigmatoid-connective, anthers short, partly covered with scale-like processes, sometimes swollen, rarely genicu- and overtopped by stigmatoid-connective, thecae 6-8, broadly late apically, sheath long and slilghtly ligulate in monopodial ellipsoid, lateral, dehiscing by subapical pore. POLLEN: leaves of all subgenera and in sympodial leaves of subgen. extruded in strands, inaperturate, ellipsoid-oblong to -elon- Pteromischum, otherwise very short and inconspicuous gate, medium-sized (mean 48 µm., range 45-51 µm.), exine except when subtending inflorescences. BLADE: very vari- almost perfectly psilate. FEMALE FLOWER: gynoecium par- ously shaped; simple and linear, cordate, sagittate or hastate, tially surrounded on one side by 3 small, very shortly stipitate, or trifid, trisect, pinnatifid, bipinnatifid, rarely pedatisect, discoid-umbonate, green staminodes, ovary oblong-ovoid, resin canals linear, short to long, obscured to very distinct on 1-locular, ovule 1, orthotropous, funicle very short to ± abaxial surface; basal ribs sometimes well-developed, pri- absent, placenta basal, stylar region attenuate, cylindric- mary lateral veins pinnate, rarely pedate, running into conoid, stigma discoid-subcapitate. BERRY: ovoid to ellipsoid, marginal vein, secondary lateral and higher order venation acute-attenuate, green. SEED: ovoid-obnapiform, strophiolate, parallel-pinnate, sometimes tertiaries and higher order veins testa smooth to ± rough, brownish to green, embryo axile, transversely reticulate between secondaries, sometimes all elongate, endosperm copious. See Plates 44, 116B. veins slender with no distinct primary laterals. FLOWERING CHROMOSOMES: 2n = 34. BRANCHES: sympodial articles of three main patterns:– sub- DISTRIBUTION: ca. 5 spp.; tropical and subtropical South gen. Pteromischum: prophyll, many foliage leaves, 1–2(–3) America:– N. Argentina, Bolivia, Brazil (Northeast, Central inflorescences; subgen. Philodendron: prophyll, following West, Southeast, South), Paraguay, Uruguay. internode suppressed, 1 foliage leaf, 1–11 inflorescences, ECOLOGY: tropical dry forest, humid and marshy forest, internode to prophyll of continuation shoot elongated; sub- seasonally wet places; geophytes; in dry forest areas the gen. Meconostigma: prophyll, following internode developed plant has a dormant period and starts growth at the begin- or very short, 1 foliage leaf, 1(–2) inflorescences, internode ning of the rainy season. to prophyll of continuation shoot suppressed. INFLORES- ETYMOLOGY: Greek spathê (spathe) and karpos (fruit); CENCE: 1–11 in each floral sympodium, secreting resin at refers to the fusion of spathe and spadix (in fruit). anthesis, either from spathe or from spadix, rarely from both. TAXONOMIC ACCOUNTS: Engler (1920a), Crisci (1971), PEDUNCLE: usually much shorter than petiole. SPATHE: Uhlarz (1983), Croat & Mount (1988). erect, entirely persistent, deciduous only at ripening of fruit (caducous after anthesis in P. surinamense), fairly thick, C Tribe Philodendreae Tribe Philodendreae Schott, Syn. Aroid. 71 (1856). Laticifers present, simple, articulated, resin canals present in roots, stems, leaves and inflorescences, sclerotic hypodermis present in roots; diminutive to gigantic, climbing hemiepi- phytes, epiphytes or terrestrial, stem usually epigeal, intravaginal squamules present; petiole only rarely geniculate apically; primary lateral veins pinnate (very rarely pedate) forming 1 marginal vein, higher order venation parallel-pin- nate; spathe persisting until fruit maturity, then deciduous at base; flowers unisexual, perigone absent; anther lacking cell wall thickenings in endothecium (except P. goeldii, P. leal- costae); endosperm copious. C 45. Philodendron 45. Philodendron Philodendron Schott in Wiener Z. Kunst 1829 (3): 780 (1829), nom. et orth. cons. (‘Philodendrum’). LECTOTYPE: P. grandifolium (Jacq.) Schott (see Britton & Wilson 1923). SYNONYMS: Philodendrum Schott in Wiener Z. Kunst 1929 (3): 780 (1829), orth. rej.; Arosma Rafinesque, Fl. Tell. 3: 66 (1837, “1836”); Telipodus Rafinesque, Fl. Tell. 3: 66 (1837, “1836”); Thaumatophyllum Schott in Bonplandia 7: 31 (1859); Elopium Schott in Oesterr. bot. Zeitschr. 15: 34 (1865); Baursea Post & O. Kuntze, Lexicon Gen. Phanerog. 62 (1903). P H I L O D E N D R E A E : P H I L O D E N D R O N 169

E D C A K B G J F H Plate 45 (i). Philodendron. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2; F, leaf × 1/2; G, leaf × 1/2; H, leaf × 1/2; J, leaf × 1/2; K, leaf × 1/2. Philodendron callosum: A, Granville et al. 10487 (K); P. crassinervium : B; Hatschbach 45979 (K); P. fibrillosum: C, Plowman et al. 11406 (K); P. calatheifolium: D, Bunting 11659 (K); P. heterophyllum: E, Whitmore 748 (K); P. blanchetianum: F, Storr 13 (K); P. grazielae: G, Dodson 2718 (K); P. frits-wentii: H, Madison 4175 (K); P. aromaticum: J, Croat 68423 (K); P. sp. cf. verrucosum: K, André 2629 (K). 170 T H E G E N E R A O F A R A C E A E

B C A DE Plate 45 (ii). Philodendron. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/3; D, leaf × 1/3; E, leaf × 1/6. Philodendron bipennifolium: A, Engler 237 (K); P. anisotomum: B, Heyde & Lux 4283 (K); P. goeldii: C, Stevenson 829 (K); P. angustisectum: D, Cult. Kew 1984–1076; E, P. bipinnati- fidum: Mayo et al. 574 (K). P H I L O D E N D R E A E : P H I L O D E N D R O N 171

A C B D EF Plate 45 (iii). Philodendron. A, habit × 1/6; B, leaf × 1/12; C, habit × 1/12; D, habit × 1/3; E, detail of lower stem showing intravaginal squamules × 1/2; F, habit × 1/12. Philodendron insigne: A, Cult. Kew. 1980–2229; P. verrucosum : B, Boyce s.n. (Kew slide collection); P. bip- innatifidum: C, Cult. Kew 1983–2024; P. scandens: D, Cult. Kew 1953–43201; P. uliginosum: E, Kew Bull. 46(4): 601–681, f.3, A (1991); P. melinonii: F, Cult. Kew 1970–84. 172 T H E G E N E R A O F A R A C E A E

B A D C Plate 45 (iv). Philodendron. A, inflorescence and associated leaf × 1/2; B, infructescences and associated stem and petiole bases × 1/2; C, floral sympodium and associated stem and petiole base × 1/2; D, inflorescence and associated petiole base × 1/2. Philodendron rigidifolium: A, Sugden 585 (K); P. leal-costae: B, Harley et al. 19428 (Kew slide collection); P. verrucosum: C, Knapp 4959 (K); McPherson 9033 (K); P. bipinnatifidum: D, Boyce s.n. (Kew slide collection). P H I L O D E N D R E A E : P H I L O D E N D R O N 173

B L CD G E MN H K J Q DD A F RS T EE AA V X W FF CC BB P Z YU Plate 45 (v). Philodendron. A, spadix × 1; B, stamen × 10; C, gynoecium × 10; D, gynoecium, longitudinal section × 10; E, gynoecium, transverse section × 10; F, spadix × 1; G, stamens × 10; H, gynoecium × 10; J, gynoecium, longitudinal section × 10; K, spadix × 1; L, stamen × 10; M, gynoecium × 10; N, gynoecium, longitudinal section × 10; P, spadix × 1; Q, stamen × 10; R, gynoecium × 10; S, gynoecium, longi- tudinal section × 10; T, gynoecium, transverse section × 10; U, spadix × 1; V, stamen × 10; W, gynoecium × 10; X, gynoecium, longitudinal section × 10; Y, gynoecium, transverse section × 10; Z, spadix × 2/3; AA, gynoecium × 5; BB, gynoecium, longitudinal section × 5; CC, spadix × 2/3; DD, stamen × 8; EE, gynoecium × 8; FF, gynoecium, longitudinal section × 8. Philodendron bipennifolium: A–E, Mayo 599 (Kew spirit col- lection 46286); P. melinonii: F–J, Evemy & Burgess 161 (Kew spirit collection 29047.428); P. blanchetianum: K–N, Storr 13 (Kew spirit collection 48471); P. eximium: P–T, Boyce s.n. (Kew spirit collection 51441); P. inaequilaterum: U–Y, Cult. Kew 1964–39505 (Kew spirit collection 49977); P. goeldii: Z–BB, Bogner 233 (Kew spirit collection 29047.740); P. bipinnatifidum: CC–FF, Boyce s.n. (Kew spirit collection 29047.785). 174 T H E G E N E R A O F A R A C E A E

sometimes (subgen. Meconostigma) extremely thick, usually Tribe Homalomeneae C constricted between tube and blade, tube convolute, cylin- C dric to ventricose, often coloured purple or red within, blade Tribe Homalomeneae (Schott) M. Hotta in Mem. Fac. Sci. usually boat-shaped, widely gaping at anthesis, later closing, Kyoto Univ., ser. Biol. 4: 89 (1970). usually white within, rarely red. SPADIX: sessile to stipitate, Laticifers present, simple, articulated, resin canals present in female zone free, rarely basally adnate to spathe, usually roots, stems and leaves, sclerotic hypodermis present in shorter than male zone and separated from it by intermedi- roots; terrestrial or rheophytic, diminutive to robust, stem ate sterile zone of staminodial flowers, intermediate sterile usually epigeal; petiole geniculum absent (very rarely pre- zone cylindric or constricted or ellipsoid and thicker than sent in Homalomena); primary lateral veins pinnate forming male zone, usually shorter than male zone, sometimes longer 1 marginal vein, higher order venation parallel-pinnate; (subgen. Meconostigma), a terminal staminodial appendix spathe often boat-shaped, constricted or unconstricted, per- sometimes also present. FLOWERS: unisexual, perigone sistent, closing after anthesis; flowers unisexual, perigone absent. MALE FLOWER: 2–6-androus, stamens free, prismatic absent; anther with cell wall thickenings in endothecium; to obpyramidal, sometimes very elongated and slender (sub- endosperm copious. gen. Meconostigma), anthers sessile to subsessile, connective thick, apically truncate, overtopping thecae, thecae ellipsoid 46. Furtadoa to oblong, dehiscing by short lateral slit or by subapical pore, endothecial thickenings lacking (except P. goeldii, P. leal- Furtadoa M. Hotta in Acta Phytotax. Geobot. 32: 142 (1981). costae). POLLEN: extruded in strands or mixed with resin TYPE: F. sumatrensis M. Hotta secretion or exuded in amorphous masses, inaperturate, ellip- HABIT: small evergreen herbs, stem repent. LEAVES: several to soid to oblong or occasionally elongate, medium-sized (mean many. PETIOLE: sheath to half as long as petiole or more. 40 µm., range 28–54 µm.), mostly perfectly psilate, sometimes BLADE: elliptic; primary lateral veins pinnate, running into from minutely verruculate, scabrate or fossulate to clearly margin, higher order venation parallel-pinnate. INFLORES- punctate, subfossulate, subfoveolate or subverrucate, rarely CENCE: 1–3 in each floral sympodium. PEDUNCLE: shorter densely and coarsely verrucate (P. leal-costae). STERILE MALE than or subequal to petiole. SPATHE: green, ellipsoid, not con- FLOWERS: staminodes usually prismatic, truncate, sometimes stricted, boat-shaped, persistent. SPADIX: subcylindric, tapering clavate, often somewhat similar to stamens. FEMALE apically, female zone a third to nearly half spadix length, male FLOWER: gynoecium ovoid, subcylindric, cylindric or zone contiguous with female. FLOWERS: unisexual, perigone obovoid, ovary (2–)4–8(–47)-locular, ovules 1–50 or more absent; each male and female flower with a stamen or sta- per locule, usually hemiorthotropous, rarely hemianatropous minode situated basally to pistillode or gynoecium respectively. to nearly anatropous, funicle long to very short, placenta MALE FLOWER: consisting of a single free stamen overtopped axile to basal, stylar region usually as broad as ovary, some- by single flask-shaped pistillode with subglobose stigmatoid times slightly broader, sometimes attenuate, rarely elongate, apex, pistillodes absent from apical flowers, stamen apex trun- lobed in subgen. Meconostigma, stigma sometimes also lobed cate, connective thick, thecae ovoid, dehiscing by short, or discoid-hemispheric, often as broad as style. BERRY: sub- longitudinal slit. POLLEN: inaperturate, ellipsoid-oblong, small cylindric to obovoid, 1–many-seeded, white, whitish- (mean 17 µm.), exine virtually psilate. FEMALE FLOWER: con- translucent, red or orange-red. SEED: tiny to fairly large, sisting of gynoecium with single, shorter, obovoid, apically ovoid-oblong to ellipsoid, rarely arillate (in P. goeldii funicle truncate staminode, lowermost flowers sometimes apparently thick, swollen, much larger than seed itself), testa thick, with more than one staminode, gynoecium ovoid, ovary 1- costate, rarely sarcotestate, embryo axile, straight, elongate, endosperm copious. See Plates 45i–v, 116C. 46. Furtadoa CHROMOSOMES: 2n = 28, 30, 32, 34, 36 (26, 48). DISTRIBUTION: over 500 spp.; tropical and southern sub- tropical America, West Indies:– Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Lesser Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Surinam, Trinidad & Tobago, Uruguay, Venezuela. ECOLOGY: usually tropical humid forest, more rarely in open woodland, swamps, streamsides; climbing hemiepiphytes, rosulate acaulescent epiphytes, rhizomatous terrestrials, litho- phytes (also on cliffs), helophytes, mostly shade-loving, sometimes arborescent. NOTES: Mayo (1986b, 1989b, 1990a, 1991) recognized 3 sub- genera: subgen. Philodendron, subgen. Pteromischum (partially revised by Grayum 1996), subgen. Meconostigma (revised by Mayo 1991). ETYMOLOGY: Greek philos (fond of) and dendron (tree); refers to the predominantly epiphytic or hemiepiphytic habit. TAXONOMIC ACCOUNTS: Krause (1913), Bunting (1968, 1977, 1980, 1984), Mayo (1986b, 1989a, b, 1990a, 1991), Grayum (1992b, 1996), Croat & Grayum (1994), Sakuragui (1994), Nadruz Coelho (1995). H O M A L O M E N E A E : F U R T A D O A 175

C D EF B LK N M J H AG Plate 46. Furtadoa. A, habit × 2/3; B, spadix × 3; C, stamens with associated pistillodes, × 15; D, stamen and associated pistillode, top view × 15; E, gynoecia with associated staminodes, × 15; F, gynoecium, longitudinal section × 15; G, habit × 2/3; H, detail of leaf tip tubule × 8; J, spadix × 3; K, stamen with associated pistillode, gynoecium in longitudinal section × 15; L, stamen and associated pistillode, top view × 15; M, gynoecia with associated staminodes, upper gynoecium in longitudinal section × 15; N, gynoecium with associated staminode, top view × 15. Furtadoa mixta: A, Nur 11091 (K); B–F, Hay 9131 (Kew spirit collection 58934); F. sumatrensis: G–H, Hotta s.n. (K); J, Bogner s.n. (Kew spirit collection 56675); K–N, Nerz s.n. (Kew spirit collection 56142). 176 T H E G E N E R A O F A R A C E A E

locular, ovules many, hemianatropous, placenta basal to intru- often becoming green, more rarely white or yellow-green or sive basal, stylar region short and attenuate to inconspicuous, red, persistent, usually not constricted, ellipsoid to boat- stigma discoid-subcapitate. BERRY: subcylindric, light green, shaped, more rarely constricted between tube and blade and bearing old stigma remains. SEED: ellipsoid to ovoid, testa then tube convolute, blade gaping at anthesis and afterwards smooth, thin, cream-coloured to very light green, embryo closing. SPADIX: shorter or subequal to spathe, stipitate or straight, endosperm copious. See Plates 46, 116D. sessile, female zone cylindric, male zone usually entirely fer- CHROMOSOMES: 2n = 40. tile, contiguous with and longer than female zone, rarely DISTRIBUTION: 2 spp.; Malay Archipelago:– Indonesia bearing staminodes basally, or very rarely separated by a ± (Sumatra), Malaysia (Peninsula). naked interstice. FLOWERS: unisexual, perigone absent. ECOLOGY: tropical humid forest; rheophytes on rocks in MALE FLOWER: 2–4-androus, rarely 5–6-androus, very rarely streams (F. sumatrensis), or forest floor terrestrials (F. mixta). 1-androus (H. monandra), stamens free, truncate apically, fil- ETYMOLOGY: named after C.X. Furtado (1897–1980). aments absent or distinct, connective thick, thecae ovoid, TAXONOMIC ACCOUNTS: Hotta (1981, 1985). ellipsoid or oblong, opening by longitudinal slit, rarely by transversal slit. POLLEN: extruded in strands, inaperturate, ellipsoid to oblong, small (mean 22 µm., range 12–31 µm.), C 47. Homalomena exine perfectly psilate in most species, rarely obscurely fos- sulate. STERILE MALE FLOWERS: 2–4-androus, sometimes Homalomena Schott in Schott & Endlicher, Melet. Bot. 20 present at base of fertile male zone, staminodia subprismatic, (1832). LECTOTYPE: H. cordata Schott (Dracontium corda- somewhat rounded apically. FEMALE FLOWER: gynoecium tum Houttuyn 1779, non Aublet 1775; see Nicolson in Taxon ovoid or oblong or subglobose, usually with single, anterior 16: 517. 1967). staminode (rarely 2, very rarely 3), equalling or half as long as ovary, sometimes absent (H. lindenii), ovary incompletely SYNONYMS: Homalonema Endlicher, Gen. Pl. 238 2–4–(–5)-locular, ovules many, hemianatropous, funicle long, (1837), orth. var.; Spirospatha Rafinesque, Fl. Tell. 4: 8 (1838, placenta parietal and axile, stylar region shortly narrowed or “1836”); Cyrtocladon Griffith, Notul. Pl. Asiat. (Posthum. Pap.) inconspicuous, stigma discoid, subhemispheric, subcapitate 3: 147 (1851); Chamaecladon Miq. in Bot. Zeitung (Berlin) or slightly 2–4-lobed. BERRY: obovoid or subglobose or 14: 564 (1856); Adelonema Schott, Prodr. syst. Aroid. 316 cylindric, locules many-seeded, rarely few-seeded. SEED: (1860); Curmeria E.F. André, Ill. Hortic. 20: 45 (1873); ellipsoid or elongate ellipsoid, testa thick, distinctly or only Diandriella Engler, Nova Guinea 8: 250 (1910). slightly costate, embryo axile, elongate, endosperm copious. HABIT: evergreen, usually aromatic (anise-scented) herbs, See Plates 47i–ii, 117A. stem shortly aerial, more rarely arborescent or hypogeal. CHROMOSOMES: 2n = 38, 40, 42, 80. LEAVES: several, rarely distichous (H. geniculata). PETIOLE: DISTRIBUTION: ca. 110 spp.; tropical southeast Asia, Malay rarely aculeate or pubescent, rarely geniculate apically (H. Archipelago, tropical America:– Bangladesh, Bolivia, Brazil geniculata), sheath usually less than half as long as petiole. (Amazonia, Central-West), Brunei, Burma, Cambodia, China BLADE: lanceolate, elliptic, oblong, subtriangular or cordate to (Guandong, Guangxi, Hainan, Taiwan, Yunnan), Colombia, sagittate, rarely peltate, usually glabrous, rarely pubescent on Costa Rica, Ecuador, French Guiana, Guyana, India (Assam), midrib and veins; primary lateral veins pinnate, running into Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, marginal vein, secondary and tertiary lateral veins parallel- Sumatra, Sunda Is.), Laos, Malaysia (Borneo, Peninsula), pinnate. INFLORESCENCE: 1–6 (or more) in each floral Panama, Papua New Guinea, Peru, Philippines, Singapore, sympodium. PEDUNCLE: shorter than petiole. SPATHE: erect, Solomon Is., Surinam, Thailand, ?Venezuela, Vietnam. 47. Homalomena H O M A L O M E N E A E : H O M A L O M E N A 177

G E A C B F D H Plate 47 (i). Homalomena. A, habit × 1/5; B, leaf × 1/2; C; leaf × 1/2; D, base of plant showing inflorescences × 1/2; E, leaf × 1/2; F, base of plant showing inflorescences × 1/2; G, leaf × 1/2; H, leaf × 1/2. Homalomena sagittifolia: A, Boyce 252 (Kew slide collecttion); H. pic- turata: B, de Granville 5467 (K); H. rubescens: C–D, Keenan s.n. (K); H. havilandii: E, Ilias & Azahari S 35677 (K); F, Hetterscheid s.n. (Kew slide collection); H. consobrina: G, de Wilde & de Wilde-Duyfjes 13495 (K); H. propinqua: H, Haviland 3134 (K). 178 T H E G E N E R A O F A R A C E A E

C H F L A ED KJ B N R T W M G P Q S UV Plate 47 (ii). Homalomena. A, infructescence, lower part of spathe removed to reveal berries × 1/1/2; B, inflorescence × 4; C, stamen × 15; D, gynoecium and associated staminode × 15; E, gynoecium, longitudinal section × 15; F, gynoecium, transverse section × 24; G, inflo- rescence, spathe partly removed × 2/3; H, stamen × 10; J, gynoecium × 10; K, gynoecium, longitudinal section × 10; L, gynoecium, transverse section × 16; M, inflorescence × 1; N, stamen × 10; P, gynoecium and associated staminode × 10; Q, gynoecium, longitudinal section × 10; R, gynoecium, transverse section × 16; S, inflorescence × 1; T, stamen × 10; U, gynoecium × 10; V, gynoecium, longitudinal section × 10; W, gynoecium, transverse section × 16. Homalomena vagans: A, Poulsen & de la Motte 273 (Kew spirit collection 58113); H. humilis: B–F, Sands 180 (Kew spirit collection 32886); H. speariae: G–L, Spear s.n. (Kew spirit collection 58911); H. rubra: M–R, Burkhill & Haniff 12792 (K & Kew spirit collection 59026); H. hostifolia: S–W, Poulsen 268 (Kew spirit collection 59027). H O M A L O M E N E A E : H O M A L O M E N A 179

ECOLOGY: tropical humid forest, swamp forest, rarely in open swamps; terrestrial in leaf litter on forest floor, along for- est streams, on road banks, on well-drained slopes in moist forest, rarely rheophytes. NOTES: Engler (1912) recognized 3 sections:– sect. Homalomena, sect. Chamaecladon, sect. Curmeria (tropical America); Furtado (1939) recognized a fourth section, sect. Cyrtocladon and Hotta (1967) a fifth, sect. Geniculatae. ETYMOLOGY: Greek homalos (flat) and mênê (moon), trans- lation of a vernacular name. TAXONOMIC ACCOUNTS: Engler (1912), Furtado (1939), Hotta (1967, 1982, 1984, 1985, 1986a, 1993), Bogner (1976b, 1986a), Bogner & Moffler (1984, 1985). C Tribe Anubiadeae Tribe Anubiadeae Engler in Nova Acta Acad. Leopold.- Carol. 39: 147 (1876). Laticifers present, simple, articulated, roots with sclerotic 48. Anubias hypodermis; helophytic or rheophytic, stem rhizomatous, creeping; petiole geniculate apically; leaf blade elliptic, lance- olate to hastate-tripartite, primary lateral veins pinnate, obpyramidal, filaments connate, sometimes fairly long, fused forming single marginal vein, higher order venation parallel- connectives thick, fleshy, sometimes covered by thecae and pinnate, finest transverse veins ± distinct; spathe boat-shaped, inconspicuous, often only incompletely connate at apex, unconstricted, persistent; flowers unisexual, perigone absent; with shallow fissures in between, thecae lateral or marginal stamens connate into ± prismatic synandria, fused connec- or covering nearly the whole synandrium from apex to base tives thickened, ± truncate, thecae lateral or marginal, rarely (A. pynaertii), dehiscing by longitudinal slit. POLLEN: inaper- covering whole synandrium, dehiscing by longitudinal slit; turate, subspheroidal to spherical, small (mean 24 µm., range ovary (1–)2–3-locular, ovules many per locule, anatropous, 20–31 µm.), exine perfectly psilate to obscurely verrucate placenta axile, style narrower than ovary, stigma broad; berry and/or dimpled. FEMALE FLOWER: gynoecium depressed- depressed-globose to obovoid; seeds small, ± ovoid to sub- globose to ovoid, ovary (1–)2–3-locular, ovules many per cylindric, endosperm copious. locule, anatropous, placenta axile, stylar region narrower than ovary, stigma broad, discoid, green, pink or white. C 48. Anubias BERRY: depressed-globular to obovoid, green to pale green, many-seeded. SEED: small, irregularly ovoid to subcylin- Anubias Schott in Oesterr. bot. Wochenbl. 7: 398 (1857). dric, testa rough, thickish, embryo axile, elongate, TYPE: A. afzelii Schott endosperm copious. See Plates 48, 117B. SYNONYM: Amauriella Rendle, Cat. Talbot’s Nigerian CHROMOSOMES: 2n = 48, 72. Pl. 115 (1913). DISTRIBUTION: 8 spp.; tropical west and central Africa:– Angola, ?Benin, Cabinda, Cameroon, ?Central African HABIT: evergreen herbs, rhizome thick, creeping, internodes Republic, Congo, Equatorial Guinea (Bioko, Rio Muni), short. LEAVES: several. PETIOLE: usually smooth, rarely Gabon, ?Gambia, Ghana, Guinea, ?Guinea-Bissau, Ivory Coast, shortly and sparsely spiny, geniculate apically, sheath rela- Liberia, Mali, Nigeria, Senegal, Sierra Leone, Togo, Zaïre. tively short. BLADE: lanceolate, ovate, elliptic, nearly ECOLOGY: tropical humid forest; helophytes or rheophytes, triangular to subsagittate, subcordate, auriculate, hastate to forest swamps, in rocky places along streams, sometimes trifid, mostly coriaceous, completely glabrous or midrib and completely submerged. primary lateral veins densely pilose abaxially; primary lateral ETYMOLOGY: Classical name of an unknown herb, veins pinnate, secondary lateral veins parallel-pinnate, ter- anoubias (according to Schott). tiaries transverse between them. INFLORESCENCE: 1–3 in TAXONOMIC ACCOUNTS: Engler (1915), Crusio (1979, each floral sympodium. PEDUNCLE: ± as long as petiole or 1987), Kasselmann (1995). shorter. SPATHE: elliptic-ovate or ovate, not constricted, only weakly differentiated into tube and blade, widely expanded Tribe Schismatoglottideae C at anthesis, but always slightly convolute at base, closing after anthesis and persistent to fruiting stage, uniformly Tribe Schismatoglottideae Nakai, Ord. Fam. Trib. Nov. 218 coloured, mostly green or cream to reddish tinged, paler (1943); Hotta in Acta Phytotax. Geobot. 33: 127–139 (1982). within. SPADIX: cylindric, shorter or sometimes much longer than spathe, sessile or stipitate, female zone free, usually Laticifers present, simple, articulated; terrestrial or rheophytic, densely flowered, rarely somewhat laxly so, shorter than stem usually epigeal; petiole sheath usually with long, male and contiguous with it or rarely with few sterile or marcescent, apical ligule (except most Schismatoglottis spp.); irregular, bisexual flowers in between, male zone fertile to blade apex with tubular mucro (except most Schismatoglottis apex. FLOWERS: unisexual, perigone absent. MALE spp.), primary lateral veins pinnate, forming single, usually FLOWER: 3–8-androus, stamens connate, synandrium ± prominent marginal vein, higher order venation parallel-pin- 180 T H E G E N E R A O F A R A C E A E

Q S PR V TU F E H A GJ CM D L B W KN Plate 48. Anubias. A, habit × 1/2; B, inflorescence × 1; C, detail of male zone of spadix × 5; D, detail of female zone of spadix × 5; E, synan- drium, side view × 10; F, synandrium, top view × 10; G, gynoecium, longitudinal section × 10; H, gynoecium, transverse section × 10; J, habit × 1/2; K, habit × 1/2; L, synandrium, side view × 10; M, synandrium, top view × 10; N, gynoecium, longitudinal section × 10; P, synandrium, side view × 10; Q, synandrium, top view × 10; R, gynoecium, longitudinal section × 10; S, inflorescence × 1; T, synandrium, side view × 10; U, synandrium, top view × 10; V, gynoecium, longitudinal section × 10; W, leaf × 1/2. Anubias afzelii: A, Morton & Gledhill SL 1169 (K); B, Hepper 2504 (K); C–H, Cult. Kew 1963–02202 (Kew spirit collection 50101); A. gracilis: J, Morton & Gledhill SL 1929 (K); A. barteri var. barteri: K, Onochie & Okafor IFH 36037 (K); L–N, Brenan 9257 (Kew spirit collection 25469); A. hastifolia: P–R, Bogner s.n. (Kew spirit collection 7100); A. pynaer- tii: S, W. Crusio, Die Gattung Anubias Schott, p. 38 (1987); T–V, Bogner 699 (K & Kew spirit collection 56680); A. gigantea: W, Bos 1914 (K). A N U B I A D E A E : A N U B I A S 181

nate; inflorescence usually 1 (except Schismatoglottis); and sterile terminally, or sterile below and fertile male ter- peduncle usually elongating in fruit; spathe tube persistent, minally, or sterile below, centrally fertile male and sterile blade usually white (except Piptospatha), deciduous at anthe- terminally, lowermost sterile zone sometimes very laxly flow- sis, rarely marcescent (in Schismatoglottis beccariana, ered, often constricted. FLOWERS: unisexual, perigone Hottarum lucens); flowers unisexual, perigone absent; sta- absent. MALE FLOWER: 2–3-androus, stamens very short to mens usually free, sometimes connate by fused filaments, long, mostly free, filaments usually well-developed, some- filaments often ± elongated, thecae truncate or horned, nearly times connate basally, always with distinctive tannin cells always dehiscing by apical pore; ovary 1-locular, ovules (dark in dried specimens), connective usually rather slender, orthotropous to hemiorthotropous (except Schismatoglottis), sometimes thicker apically, anthers truncate, often concave stigma usually sessile (except Schismatoglottis); seed testa apically, thecae opposite, cylindric or obconic to ovoid, usually costate, embryo axile, endosperm copious. dehiscing by apical, broadly elliptic, or bilobed pore. POLLEN: extruded in strands, inaperturate, ellipsoid to C 49. Schismatoglottis oblong, small (mean 20 µm., range 15–26 µm.), exine per- fectly psilate, rarely rugulate to verruculate (S. spruceana). Schismatoglottis Zollinger & Moritzi in Moritzi, Syst. STERILE MALE FLOWERS: staminodes less compressed than stamens, obpyramidal to clavate, usually truncate, short to Verzeichnis Zollinger, 83 (1846). TYPE: S. calyptrata (Roxburgh) Zollinger & Moritzi (Calla calyptrata Roxburgh). long. FEMALE FLOWER: gynoecium sometimes accompa- nied by 1–4 clavate, rarely peltate staminodes with generally SYNONYMS: Philonotion Schott in Oesterr. bot. distinctly swollen apices, or rarely with sterile flowers scat- Wochenbl. 7: 421 (1857); Apoballis Schott in Oesterr. bot. tered among gynoecia, ovary 1-locular, ovules (1–)few to Zeitschr. 8: 318 (1858); Apatemone Schott, Gen. Aroid. t. 57 many, anatropous to hemianatropous, funicle rather long, (1858); Colobogynium Schott in Oesterr. bot. Zeitschr. 15: 34 placentae 1–4, parietal, usually extending from base to apex (1865); Nebrownia O. Kuntze, Rev. Gen. 2: 742 (1891). of locule, stylar region inconspicuous or shortly conoid, HABIT: small to large, evergreen herbs, rarely shortly, densely stigma discoid to capitate, small to as wide as ovary. BERRY: or long-pubescent, stem rhizomatous or epigeal, shortly erect. oblong to globose, green or dull yellow or deep red, few- to LEAVES: numerous, rarely distichous. PETIOLE: sheath less many-seeded. SEED: ellipsoid, testa costate, embryo straight, than half petiole length, sometimes with long apical ligule. elongate, endosperm copious. See Plates 49i–iii, 117C. BLADE: narrow-elliptic, elliptic, lanceolate, oblanceolate, CHROMOSOMES: 2n = 26, 39, 52. ovate, obovate, cordate, cordate-sagittate, sometimes varie- DISTRIBUTION: ca. 120 spp.; tropical Asia, Malay Archipelago, gated with paler or silvery green, white or yellow; primary tropical South America:– Brazil (Amazonia), Brunei, Burma, lateral veins pinnate, running into distinct marginal vein, sec- Cambodia, China (Guandong, Guangxi, Hainan, Yunnan), ondary and tertiary laterals parallel-pinnate, higher order Colombia, French Guiana, ?Guyana, Indonesia (Borneo, Irian venation transverse-reticulate. INFLORESCENCE: 1–3 (or Jaya, Java, Moluccas, Sulawesi, Sumatra, Sunda Is.), Laos, more) in each floral sympodium. PEDUNCLE: shorter than Malaysia (Borneo, Peninsula), Papua New Guinea, Peru, petiole. SPATHE: constricted between tube and blade, some- Philippines, Solomon Is., Surinam, Thailand, Vanuatu, times only slightly so, rarely not at all, tube convolute, Venezuela, Vietnam. persistent, blade thinner, erect, broadly boat-shaped, gaping ECOLOGY: tropical humid forest; terrestrial, forest floor, and then caducous at anthesis, rarely marcescent, usually sometimes rheophytes. white to cream, sometimes greenish-yellow, very rarely pink, NOTES: Hotta (1966a) recognized 5 informal groups in the cuspidate to acuminate. SPADIX: shorter than or equalling Bornean species:– S. homalomenoidea group, S. monopla- spathe, lower part consisting of cylindric to conoid female centa group, S. barbata group, S. acutifolia group, S. zone, free or partially adnate to spathe, sometimes bearing calyptrata group; the neotropical species earlier recognized sterile organs at the very base, upper part of spadix usually as the genus Philonotion now form sect. Philonotion ± clavate, sometimes subcylindric, either fertile male below (Bunting 1960b). 49. Schismatoglottis 182 T H E G E N E R A O F A R A C E A E

B C D E A Plate 49 (i). Schismatoglottis. A, habit × 1/2; B, detail of leaf tip tubule × 8; C, habit × 1/2; D, habit × 1/2; E, habit × 1/2. Schismatoglottis gillianae: A–B, Coode 6313 (K); S. spruceana: C, Plowman 13527A (K); S. hottae: D, Johns 6872 (K); S. convolvula: E, Mamit S42102 (K). S C H I S M A T O G L O T T I D E A E : S C H I S M A T O G L O T T I S 183

D AB C E FG Plate 49 (ii). Schismatoglottis. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2; F, leaf × 1/2; G, leaf × 1/2. Schismatoglottis hastifolia: A, Clemens & Clemens 29493 (K); S. calyptrata: B, Beaman 10632 (K); S. ferruginea: C, Dransfield 6871 (K); D, S. hottae: Johns 6872 (K); S. crispata: E, Cult. Veitch, June & July 1881 (K); S. spruceana: F, Plowman 13527A (K); S. gillianae: G, Coode 6313 (K). ETYMOLOGY: Greek schisma, schismatos (separating) and zone of sterile male flowers, male zone cylindric, equal in glôtta (tongue); refers to the deciduous spathe blade. thickness to female, obtuse, fertile to apex or with a few TAXONOMIC ACCOUNTS: Engler (1912), Bunting (1960b), sterile terminal flowers. FLOWERS: unisexual, perigone Hotta (1965, 1966a), Bunting & Steyermark (1969), Bogner & absent. MALE FLOWER: 1–2-androus, stamens free, com- Hotta (1983b), Bogner (1988a). pressed, anthers truncate, connective ± flat or expanded apically or with conical beak (P. insignis), overtopping the- cae, thecae oblong-ellipsoid, dehiscing by apical pore. C 50. Piptospatha POLLEN: inaperturate, ellipsoid, small to medium-sized (mean 25 µm.), exine psilate. STERILE MALE FLOWERS: Piptospatha N.E. Brown in Gard. Chron., ser. 2, 11: 138 apparently composed of a single truncate, subclavate, pris- (1879). TYPE: P. insignis N.E. Brown matic staminode. FEMALE FLOWER: gynoecium free or cohering to neighbouring ones, ovary 1-locular, ovules SYNONYMS: Rhynchopyle Engler in Bot. Jahrb. 1: 183 many, hemiorthotropous to almost orthotropous and erect, (1880) (“1881”); Gamogyne N.E. Brown in J. Bot. 20: 195 funicle long, placentae 2–4, parietal or parietal and basal, (1882). stigma ± sessile or stylar region inconspicuous, as broad as HABIT: small to medium-sized evergreen herbs, stem erect ovary, contiguous with adjacent ones. BERRY: obovoid, or decumbent. LEAVES: several. PETIOLE: sheath short with green. SEED: elongate-ellipsoid to cylindric, with long, long, marcescent ligule. BLADE: elongate-lanceolate to ellip- curved micropylar appendage, testa slightly costate, embryo tic or oblanceolate, coriaceous, apex with tubular mucro; elongate, endosperm copious. See Plates 50, 117D. primary lateral veins pinnate, running into distinct marginal CHROMOSOMES: 2n = 26. vein, secondary laterals and higher order venation parallel- DISTRIBUTION: 10 spp.; Brunei, Indonesia (Borneo), pinnate. INFLORESCENCE: solitary, usually nodding. Malaysia (Borneo, Peninsula), Thailand. PEDUNCLE: subequal to or longer than petiole. SPATHE: ECOLOGY: tropical humid forest; rheophytes. stoutly ellipsoid, not constricted, often pink, lower part per- NOTES: sect. Piptospatha, sect. Gamogyne (N.E. Brown) M. sistent and cup-like, upper part slightly gaping at anthesis, Hotta. The latter section is only weakly defined, by the pres- caducous or deliquescent, cuspidate to acuminate. SPADIX: ence of superficially connate gynoecia. sessile with ± oblique insertion, sometimes with sterile ETYMOLOGY: Greek piptô (I fall) and spathê (spathe); refers female flowers at extreme base, female zone cylindric, to the deciduous spathe blade. shorter and contiguous with male, or separated by a short TAXONOMIC ACCOUNTS: Engler (1912), Hotta (1965). 184 T H E G E N E R A O F A R A C E A E

D K EH L BP A FG MN S C J T BB Y W R Q Z UV AA X Plate 49 (iii). Schismatoglottis. A, infructescences with associated petiole × 2/3; B, inflorescence × 1; C, spadix × 2; D, staminode × 20; E, stamens × 20; F, gynoecium × 20; G, gynoecium, longitudinal section × 20; H, gynoecium, transverse section × 30; J, spadix × 2; K, sta- minode × 20; L, stamen × 20; M, gynoecium × 20; N, gynoecium, longitudinal section × 20; P, gynoecium, transverse section × 30; Q, inflorescence × 1; R, spadix × 2; S, staminode × 20; T, stamen × 20; U, gynoecium × 20; V, gynoecium, longitudinal section × 20; W, gynoe- cium, transverse section × 30; X, spadix, spathe partly removed × 2; Y, stamen × 20; Z, gynoecium × 20; AA, gynoecium, longitudinal section × 20; BB, gynoecium, transverse section × 30. Schismatoglottis neoguinensis: A, Floyd 6445 (K); S. tecturata: B–H, Bogner 1554 (Kew spirit collection 45228); S. spruceana: J–P, Plowman 13527A (K & Kew spirit collection 58039); S. crispata: Q–W, Boyce 672 (Kew spirit collection 59035); S. calyptrata: X–BB, Hay 2007, Cult. Kew 1982–04973 (Kew spirit collection 46560). S C H I S M A T O G L O T T I D E A E : S C H I S M A T O G L O T T I S 185

E B FD C H K N A LM G J Plate 50. Piptospatha. A, habit × 1/2; B, seed × 15; C, habit × 1/2; D, spadix × 2; E, stamen × 15; F, gynoecia, right hand gynoecium longi- tudinally sectioned × 10; G, habit × 1/2; H, detail of leaf tip tubule × 6; J, spadix × 2; K, stamen × 15; L, gynoecium × 15; M, gynoecium, longitudinal section × 15; N, staminode × 15. Piptospatha elongata: A, Chew, Corner & Stainton 2501 (K); B, Bogner 2153 (Kew spirit collec- tion 59089); P. burbidgei: C–F, Richards 1091 (K & Kew spirit collection 58020); P. ridleyi: G, Burkill 2577 (K); H, Sinclair 10577 (K); J–M, Bogner 2120 (Kew spirit collection 57278). 186 T H E G E N E R A O F A R A C E A E

50. Piptospatha C 51. Hottarum posed entirely of sterile male flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: (1–)2–3(–4)-androus, sta- Hottarum Bogner & Nicolson in Aroideana 1: 72 (1979, mens truncate at apex, filaments distinct, sometimes connate “1978”). TYPE: H. truncatum (M. Hotta) Bogner & Nicolson basally, connective broad, thick, thecae lateral, opposite, not (Microcasia truncata M.Hotta). horned, dehiscing apically by single pore or two pores con- fluent into a short transverse slit. POLLEN: inaperturate, SYNONYM: Based on Microcasia sect. Truncatae M. Hotta ellipsoid or globose, small (mean 14 µm., range 10–17 µm.), in Mem. Coll. Sci. Univ. Kyoto, ser. B, Biol., 32: 21 (1965). exine perfectly psilate or irregularly dimpled. STERILE MALE HABIT: evergreen herbs, stem short, ± erect, epigeal, inter- FLOWERS: composed of truncate, prismatic staminodes. nodes short. LEAVES: numerous. PETIOLE: sheath with long, FEMALE FLOWER: usually consisting of gynoecium alone, free ligule. BLADE: elliptic, oblong-elliptic, or narrowly ellip- sometimes also with a clavate staminode in basal flowers tic, coriaceous, apex with tubular mucro; primary lateral (or lateral flowers when female zone adnate to spathe), ovary veins pinnate, running into distinct marginal vein, higher depressed-globose, 1-locular, ovules 10–15, orthotropous, order venation parallel-pinnate. INFLORESCENCE: 1, rarely 2 funicle long, placenta basal, stigma sessile, discoid or sub- in each floral sympodium. PEDUNCLE: subequal to petiole capitate. BERRY: ± globose, many-seeded, whitish green with in flower, elongating in fruit. SPATHE: constricted or not, brownish stigma remnants. SEED: ellipsoid to elongate, testa uniformly coloured, or green below and white above, ellip- whitish to brownish, costate, embryo straight, elongate, soid to obovoid at anthesis, lower part convolute, cup-shaped endosperm copious. See Plates 51, 118A. and persistent after anthesis, upper part only opening slight- CHROMOSOMES: 2n = 26. ly at anthesis, caducous, or marcescent and then evanescent DISTRIBUTION: 6 spp.; Brunei, Indonesia (Borneo), Malaysia (H. lucens), apex acuminate to cuspidate. SPADIX: subcy- (Borneo). lindric, usually free or sometimes adnate to spathe for ECOLOGY: tropical humid forest; rheophytes. two-thirds of length (H. lucens), female zone subcylindric, NOTES: Hottarum lucens is rather different from the other sometimes with sterile flowers at base, either contiguous species. with male, or with a few sterile male flowers in between, ETYMOLOGY: named after Mitsuru Hotta (born 1935) and male zone as thick or thicker than female, apical zone com- Arum. posed of sterile male flowers, in H. lucens fertile only where TAXONOMIC ACCOUNTS: Bogner (1983b,1984b), Bogner & exposed at anthesis by spathe opening and otherwise com- Hotta (1983a). 51. Hottarum S C H I S M A T O G L O T T I D E A E : H O T T A R U M 187

C E D A L B N F G H JM P K Q Plate 51. Hottarum. A, habit × 2/3; B, spadix × 2; C, stamen × 16; D, staminode × 16; E, gynoecium, longitudinal section × 16; F, spadix × 2; G, stamen × 16; H, staminode × 16; J, gynoecium with associated staminode; K, habit × 2/3; L, detail of leaf tip tubule × 6; M, spadix × 3; N, stamen × 16; P, gynoecium × 16; Q, gynoecium, longitudinal section × 16. Hottarum lucens: A, Elsener 184 (K); B–E, Bogner 1439 (Kew spirit collection 45223); H. sarikeense: F–J, Bogner 1553 (Kew spirit collection 45681); H. kinabaluense : K, M–Q, Edwards 2162 (Kew spirit collection 54425); L, Clemens & Clemens 29135 (K). 188 T H E G E N E R A O F A R A C E A E

C 52. Bucephalandra 53. Phymatarum C Bucephalandra Schott, Gen. Aroid. t. 56 (1858). TYPE: B. Phymatarum M. Hotta in Mem. Coll. Sci. Kyoto Imp. Univ., motleyana Schott ser. B, 32: 29 (1965). TYPE: P. borneense M. Hotta SYNONYM: Microcasia Beccari in Bull. Soc. Tosc. Ortic. HABIT: small evergreen herbs, stem creeping to decum- 4: 180 (1879). bent. LEAVES: several. PETIOLE: sheath fairly short with long marcescent ligule. BLADE: narrowly elliptic, some- HABIT: minute to medium-sized evergreen herbs, stem creep- what coriaceous, apex with tubular mucro; primary lateral ing, apex upright. LEAVES: numerous. PETIOLE: sheath with veins pinnate, running into conspicuous marginal vein, sec- long marcescent ligule. BLADE: elliptic, elliptic-oblong, linear- ondary and tertiary laterals parallel-pinnate, higher order oblanceolate to obovate, coriaceous, punctate below, apex venation inconspicuously transverse-reticulate. INFLORES- with tubular mucro; primary lateral veins pinnate, running into CENCE: solitary. PEDUNCLE: erect, shorter or subequal to distinct marginal vein, higher order venation parallel-pinnate. petiole. SPATHE: constricted between tube and blade, tube INFLORESCENCE: solitary. PEDUNCLE: subequal to petiole at convolute, persistent, green, blade longer, boat-shaped and anthesis, elongating later. SPATHE: ellipsoid, cuspidate, not gaping at anthesis, whitish, cuspidate, caducous after anthe- constricted, lower part light green, convolute, broadly funnel- sis. SPADIX: extreme base bearing a few pistillodes or not, shaped, persistent, enclosing developing fruits, upper part female zone conoid to subcylindric, basally adnate to white, gaping at anthesis, caducous immediately afterwards. spathe, separated from male zone by cylindric to ellipsoid SPADIX: sessile, shorter than spathe, with a few pistillodes at zone of sterile male flowers, male zone very short and extreme base, female zone cylindric, narrower than upper slightly narrower, terminal appendix much longer, elon- parts, with gynoecia in 2–6 spirals, separated from male zone gate-conoid, bearing sterile male flowers. FLOWERS: by a few rows (usually 2) of flattened, smooth, scale-like sta- unisexual, perigone absent. MALE FLOWER: apparently 1- minodes, male zone with 2–5 rows of flowers, terminal androus, free, filament short, connective inconspicuous, appendix globose or ellipsoid to subcylindric, composed of thecae tuberculate, ending in curved horn, dehiscing by truncate, obpyramidal to subcylindric, apically papillose sta- apical pore. STERILE MALE FLOWERS: staminodes sub- minodes, the uppermost ± connate. FLOWERS: unisexual, prismatic, tuberculate, flattened or excavated, lowermost perigone absent. MALE FLOWER: 1-androus, filament distinct either with or without central, short, subulate projection, but short, flattened, connective ± inconspicuous, thecae ellip- uppermost more slender, relatively longer, otherwise simi- soid, extrorse, dehiscing by pore at tip of conspicuous apical lar but never with projections. POLLEN: inaperturate, horn. POLLEN: extruded in a droplet, inaperturate, ellipsoid, ellipsoid, small (mean 19 µm., range 17–22 µm.) exine psi- medium-sized (mean 29 µm., range 28–30 × 20–24 µm.), exine late. FEMALE FLOWER: gynoecium depressed- globose, smooth (psilate). FEMALE FLOWER: gynoecium depressed- ovary 1-locular, ovules many, hemiorthotropous, funicle globose, 1-locular, ovules many, orthotropous, attenuate long, placenta basal, stigma sessile, slightly concave cen- towards micropyle, funicle distinct, placenta basal, stigma ses- trally, narrower than ovary, very thinly discoid. BERRY: sile, discoid, slightly concave in centre, narrower than ovary. many-seeded, depressed-obovoid, slightly furrowed, green- BERRY: globose to ellipsoid with numerous seeds. SEED: nar- ish-white. SEED: ellipsoid, with long micropylar appendage, row-ellipsoid, with long, curved micropylar appendage, testa testa costate, embryo elongate, straight, endosperm copious. very slightly longitudinally ribbed to scabrous, embryo straight, See Plates 53, 118C. elongate, endosperm copious. See Plates 52, 118B. CHROMOSOMES: 2n = 26. CHROMOSOMES: 2n = ca. 26. DISTRIBUTION: 3 spp.; Brunei, Indonesia (Borneo), Malaysia DISTRIBUTION: 3 spp.; Brunei, Indonesia (Borneo), Malaysia (Borneo). (Borneo). ECOLOGY: tropical humid forest; rheophytes. ECOLOGY: tropical humid forest; rheophytes. ETYMOLOGY: Greek phyma, phymatos (tumour, growth) ETYMOLOGY: Greek bous (bull or cow), cephalê (head) and and Arum. anêr, andros (man). TAXONOMIC ACCOUNTS: Bogner (1984a). TAXONOMIC ACCOUNTS: Engler (1912), Bogner (1980a, 1984b), Boyce (1995a), Boyce, Bogner & Mayo (1995). 52. Bucephalandra S C H I S M A T O G L O T T I D E A E : P H Y M A T A R U M 189