BOOK-ARACEAE

C P Q A D B MF G N L E KH J Plate 52. Bucephalandra. A, habit × 2/3; B, habit × 2/3; C, detail of leaf tip tubule × 6; D, inflorescence × 1; E, spadix × 5; F, upper spadix staminode × 15; G, stamen × 15; H, mid–spadix staminode × 15; J, gynoecium, longitudinal section × 15; K, lower spadix pistillode × 15; L, base of plant × 2/3; M, leaf × 2/3; N, spadix × 5; P, habit × 2/3; Q, spadix × 5. Bucephalandra motleyana: A, Mamit S 33470 (K); B, Chew 1111 (K); C, Marsh & Simpson 21 (Kew spirit collection 57279); D–K, Bogner 1447 (Kew spirit collection 45261); B. gigantea: L–M, Endert 4580 (K); N, Endert 4580 (Kew spirit collection 58050); B. catherineae: P–Q, de Vogel & Cribb 9210 (Kew spirit collection 57575). 190 T H E G E N E R A O F A R A C E A E

AD B E K F GH J C Plate 53. Phymatarum. A, habit × 2/3; B, detail of leaf tip tubule × 6; C, spadix × 2; D, staminode from appendix × 8; E, stamen × 8; F, staminode from lower zone of sterile male flowers × 8; G, gynoecium–associated staminode × 8; H, gynoecium × 8; J, gynoecium, longitu- dinal section × 8; K, seed × 4. Phymatarum borneense: A–B, Bogner 1506 (K); C–J, Boyce 341 (Kew spirit collection 56097); K, Bogner 2159 (Kew spirit collection 58007). 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 191

53. Phymatarum C 54. Aridarum on one side and situated inside or outside connective cavity, apically narrowed into long or short horn, dehiscing by api- Aridarum Ridley in J. Bot. 51: 201 (1913). TYPE: A. mon- cal pore. POLLEN: inaperturate, ellipsoid-oblong, small (mean tanum Ridley 23 µm., range 16–31 µm.), exine psilate. STERILE MALE FLOWERS: consisting of prismatic to obpyramidal, truncate HABIT: small to medium-sized, evergreen herbs, stem decum- staminodes with or without a small, central, apical cavity. bent, erect distal part sometimes rather long. LEAVES: several, FEMALE FLOWER: gynoecium shallow, laterally compressed, spiral or rarely distichous (A. annae). PETIOLE: sheath with subhexagonal to subglobose, ovary 1-locular, ovules many, long, marcescent ligule. BLADE: coriaceous, linear to elliptic, orthotropous to hemiorthotropous, funicle distinct, erect, pla- apex with tubular mucro; primary lateral veins pinnate, weakly centa basal, stigma sessile, slightly concave centrally, as broad or not differentiated, running into distinct marginal vein, higher as ovary, contiguous with neighbouring stigmas. BERRY: glo- order venation parallel-pinnate. INFLORESCENCE: solitary, bose or ellipsoid to cylindric, stigma remnant persistent, sometimes ± nodding. PEDUNCLE: subequal or longer than infructescence globular to slightly elongate. SEED: ellipsoid, petiole, sometimes ± deflexed at apex. SPATHE: stoutly ellip- elongate, testa longitudinally costate, embryo elongate, soid, not constricted, convolute and gaping at apex only or endosperm present. See Plates 54, 118D. broadly boat-shaped and widely gaping to base, lower part CHROMOSOMES: 2n = 24. persistent, green, upper part caducous, white, cuspidate to DISTRIBUTION: 7 spp.; Brunei, Malaysia (Borneo). acuminate at apex. SPADIX: sessile, cylindric, normally with a ECOLOGY: tropical humid forest; rheophytes. few sterile flowers at extreme base, female zone cylindric, NOTES: 2 sections recognized by Hotta (1965):– sect. shorter than male and separated from it by equally thick zone Aridarum, sect. Caulescentia. of sterile male flowers, male zone equally thick, with shorter ETYMOLOGY: Latin aridus (dry) and Arum (the genus); a apical zone of sterile flowers, apex obtuse. FLOWERS: uni- misnomer as Aridarum species are rheophytes and do not sexual, perigone absent. MALE FLOWER: 1–2-androus, occur in dry habitats. filaments distinct to very short, free to connate, connective TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), slightly to deeply excavated, thecae either opposite or paired Hotta (1965), Bogner (1979b, 1981a, 1983a). 54. Aridarum 192 T H E G E N E R A O F A R A C E A E

K N P L B M E J F U AC R GH D ST Q Plate 54. Aridarum. A, habit × 2/3; B, spadix × 6; C, detail of stamens × 6; D, detail of gynoecia × 6; E, stamen × 10; F, stamen, three quarter view × 10; G, gynoecium × 10; H, gynoecium, longitudinal section × 10; J, habit × 2/3; K, detail of leaf tip tubule × 3; L, detail of stamens × 6; M, detail of gynoecia × 6; N, stamen × 10; P, gynoecium × 10; Q, habit × 2/3; R, detail of stamens × 6; S, detail of gynoecia × 6; T, gynoe- cium × 10; U, stamen × 10. Aridarum nicolsonii: A–H, Bogner 1440 (K & Kew spirit collection 56427); A. burttii: J–K, Chai S34072 (K); L–P, Burtt 5116 (Kew spirit collection 34396); A. caulescens var. caulescens : Q–U, Boyce 250 (K & Kew spirit collection 56095). S C H I S M A T O G L O T T I D E A E : A R I D A R U M 193

C 55. Heteroaridarum of slender, basally connate, clavate, truncate staminodes. FEMALE FLOWER: gynoecium depressed-globose, ovary Heteroaridarum M. Hotta in Acta Phytotax. Geobot. 27 1-locular, ovules many, hemiorthotropous, placentae 2, (3–4): 63 (1976). TYPE: H. borneense M. Hotta basal and apical, apical one bearing smaller and apparently sterile ovules, stigma sessile, as broad as ovary, concave HABIT: evergreen herb, stem short, erect. LEAVES: sev- centrally. BERRY: unknown. SEED: unknown. See Plate 55. eral, distichous. PETIOLE: sheath with long, narrowly CHROMOSOMES: unknown. triangular ligule. BLADE: narrowly elliptic-oblanceolate, DISTRIBUTION: 1 sp.; Malaysia (Borneo). coriaceous; primary lateral veins pinnate, running into mar- ECOLOGY: tropical humid forest; rheophyte. ginal vein, higher order venation parallel-pinnate. ETYMOLOGY: Greek heteros (different) and Aridarum. INFLORESCENCE: solitary. PEDUNCLE: subequal to petiole. SPATHE: slightly constricted, tube convolute, green, per- sistent, blade white, acuminate, caducous. SPADIX: sessile, Tribe Cryptocoryneae C female zone short, separated from male by short zone of sterile flowers, male zone much longer, subcylindric-ellip- Tribe Cryptocoryneae Blume, Rumphia 1: 83 (1836). soid, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3-androus, filaments as long as anthers, Laticifers absent in foliage leaves but present in stem, roots connate, with conspicuous tannin cells, connective thick, and cataphylls (Sivadasan, pers. comm.); aquatic to amphibi- all three stamens parallel to one another, 2 outermost much ous, stem rhizomatous, intravaginal squamules present; larger with shortly horned thecae, central one much smaller primary lateral veins pinnate, forming single marginal vein, with hornless thecae, all thecae opening by apical pore. higher order venation reticulate; inflorescence 1; spathe tube POLLEN: unknown. STERILE MALE FLOWERS: composed with margins connate, tube usually ± swollen basally to C A BG EF H D JK Plate 55. Heteroaridarum. A, habit × 1/3; B, leaf × 1/3; C, detail of leaf tip tubule × 4; D, spadix × 2; E, stamens, side view × 10; F, stamens, front view × 10; G, stamens, longitudinal section × 10; H, staminodes × 10; J, gynoecium × 10; K, gynoecium, longitudinal section × 10. Heteroaridarum borneense: A–K, Abbe et al. 9845 (SAR). 194 T H E G E N E R A O F A R A C E A E

55. Heteroaridarum form a “kettle”, blade lanceolate to cordate, twisted or long- ered with shortly branched processes, distinct “collar” spiralled, often long-caudate; spadix entirely enclosed in around mouth of kettle present or absent. SPADIX: slender, spathe tube “kettle”, sterile terminal appendix apically adnate female zone usually with 3–5(–7) spirals of gynoecia, rarely to spathe at apex of kettle; flowers unisexual, perigone in pseudowhorl (L. nairii) or true whorl (L. gomezii), sepa- absent; male flower 1(–2)-androus, stamens free, anther ses- rated from male zone by slender naked axis, male zone sile or with short filament, excavated at apex with prominent cylindric to ellipsoid, terminal appendix small, conoid, api- margins, connective inconspicuous, thecae ending in promi- cally adnate to spathe behind spathe flap; olfactory bodies nent horn, dehiscing by apical pore, pollen exine smooth present or absent above female flowers. FLOWERS: unisex- (psilate); ovary 1-locular, ovules orthotropous; embryo elon- ual, perigone absent. MALE FLOWER: 1(–2)-androus, gate, endosperm copious. stamens free, anthers sessile or with short thick filaments, apex somewhat excavated with prominent thickened mar- gin, connective inconspicuous, thecae opposite, each C 56. Lagenandra narrowed into an erect horn usually exceeding, rarely equalling thickened margin and dehiscing by apical pore. Lagenandra Dalzell in Hooker’s J. Bot. Kew Gard. Misc. 4: POLLEN: inaperturate, ellipsoid to ellipsoid-oblong, medium- 289 (1852). TYPE: L. toxicaria Dalzell sized (mean 36 µm., range 35–38 µm.), exine perfectly Laticifers absent in foliage leaves but present in stem, roots 56. Lagenandra and cataphylls (Sivadasan, pers. comm.). HABIT: small to medium-sized evergreen herbs, rhizome creeping, thick, rarely stoloniferous (L. nairii). LEAVES: several, cataphylls conspicuous. PETIOLE: sheath fairly long. BLADE: ptyxis involute, ovate, lanceolate, elliptic to almost linear, usually glabrous (hairy in L. nairii), coriaceous; primary lateral veins pinnate, weakly differentiated, running into inconspicuous marginal vein, secondary laterals parallel to primaries, higher order venation transverse-reticulate. INFLORESCENCE: soli- tary. PEDUNCLE: short, rarely long (L. bogneri), erect at anthesis, deflexed in fruit. SPATHE: marcescent, very thick- walled, outer surface green to purple, completely smooth or very warty on blade, margins basally connate into cylindric or ellipsoid tube (“kettle”), tube sometimes narrowing api- cally within (by thickening of walls, thus not apparent from outside), inner surface of lower region usually longitudi- nally furrowed, mouth of tube occluded by transverse, centrally perforated septum with separate transverse flap situated immediately below septum and partly or completely covering male zone of spadix, blade straight or twisted, opening only slightly by narrow longitudinal or spiral slit, or widely gaping, apex acute to long-caudate, inside smooth, uniformly warty or with warts in transverse bands, or cov- C R Y P T O C O R Y N E A E : L A G E N A N D R A 195

JK H A B FG E D C Plate 56. Lagenandra. A, habit × 2/3; B, infructescence × 1; C, habit × 2/3; D, leaf × 2/3; E, inflorescence with spadix detail viewed through opened kettle × 2; F, inflorescence with spadix detail viewed through opened kettle × 1; G, spadix × 3; H, stamen × 20; J, gynoecium × 10; K, gynoecium, longitudinal section × 10. Lagenandra toxicaria: A, Barnes 641 (K); Meebold 8877 (K); B, Barnes 1003 (Kew spirit collection 22447); L. koenigii: C–D, Bogner 564 (K & Kew spirit collection 57394); L. nairii: E, Bogner 1847 (Kew spirit collection 53950); L. dewitii: F–K, Bogner 1749 (Kew spirit collection 49709). 196 T H E G E N E R A O F A R A C E A E

psilate. FEMALE FLOWER: gynoecium free, broadly ellip- almost contiguous, male zone ellipsoid to subcylindric, termi- soid to globose, ovary 1-locular, ovules 1–12(–15), nal appendix short, conical or irregularly oblong to subclavate, orthotropous, tapering towards micropyle, funicle short, apically adnate to spathe behind flap, often breaking free dur- bearing very long trichomes, placenta basal, stylar region ing anthesis. FLOWERS: unisexual, perigone absent. MALE thick, usually short, rarely long, stylar region and upper part FLOWER: apparently 1-androus, stamens free, anthers sessile of ovary often conspicuously warty, stigma discoid to hemi- or with short filaments, apex excavated with very prominent, spheric, sometimes oblique, relatively broad. BERRY: free, thickened margins, connective inconspicuous, thecae opposite, obovoid to prismatic-ellipsoid, often apically warty, rarely ending in short to long, attenuate horn usually projecting smooth, at maturity pericarp splitting and revolute basally to beyond thickened margins, dehiscing by apical pore. POLLEN: release seeds, infructescence usually globose, deflexed, pros- extruded in a droplet, inaperturate, ellipsoid to oblong, trate. SEED: ellipsoid to narrowly ellipsoid or subcylindric, medium-sized (mean 34 µm., range 27–42 µm.), exine psilate. testa longitudinally costate, dark brown, embryo elongate, FEMALE FLOWER: ovaries connate, 1-locular, ovules 5 to endosperm copious. See Plates 56, 119A. many, orthotropous, funicle short, usually bearing numerous CHROMOSOMES: 2n = 36, 72. long trichomes extending between ovules, placenta subbasal DISTRIBUTION: 14 spp.; tropical south Asia:– Bangladesh, to obliquely parietal (adaxial, morphologically basal), stylar India (Assam, south India), Sri Lanka. region free, usually curved away from spadix axis, stigma dis- ECOLOGY: tropical humid forest; usually helophytes, rarely coid, rounded or elongated, often emarginate, often centrally rheophytes, streams, marshes. concave. FRUIT: not a berry, all connate into an ovoid, smooth NOTES: L. nairii and L. gomezii are quite distinct from the to verrucose syncarp dehiscing apically at maturity and becom- other species. ing star-shaped. SEED: ellipsoid-oblong, straight or slightly ETYMOLOGY: Greek lagenos or lagynos (flask, bottle) and curved, testa brown, not very thick, rough to slightly costate, anêr, andros (man). or smooth, embryo elongate, straight, endosperm copious; TAXONOMIC ACCOUNTS: Engler (1920a), de Wit seed sometimes viviparous and then seedling with many fili- (1978,1990), Sivadasan (1982), Bogner & Jacobsen (1987), form cataphylls (C. ciliata). See Plates 57, 119B. Nicolson (1988a), Kasselmann (1995). CHROMOSOMES: 2n = 20, 22, 28, 30, 33, 34, 36, 42, 54, 66, 68, 72, 85, 88, 90, 102, 132. DISTRIBUTION: ca. 50 spp.; tropical Asia, Malay C 57. Cryptocoryne Archipelago:– Bangladesh, Brunei, Burma, Cambodia, China (Guandong, Guangxi, Guizhou, Yunnan), India, Indonesia Cryptocoryne Fischer ex Wydler in Linnaea 5: 428 (1830). (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Sumatra), Laos, TYPE: C. spiralis (Retzius) Fischer ex Wydler (Arum spirale Malaysia (Borneo, Peninsula), Papua New Guinea, Retzius). Philippines, Singapore, Sri Lanka, Thailand, Vietnam. ECOLOGY: tropical humid forest, freshwater tidal zone, rarely Laticifers absent in foliage leaves but present in stem, roots and in brackish water (C. ciliata), submerged or emergent aquat- cataphylls (Sivadasan, pers. comm.). HABIT: small or medium- ics, helophytes or facultative rheophytes. sized herbs, evergreen or rarely dormant in dry season (C. NOTES: Jacobsen (1982) recognized 26 informal subgroups. nevillii, C. consobrina), rhizome creeping, stoloniferous. C. spiralis is distinguished from other species of the genus in LEAVES: many, cataphylls conspicuous when present. PETI- its spathe, which lacks the tube between kettle and blade and OLE: sheath fairly long. BLADE: ptyxis convolute, cordate, ovate, lanceolate, elliptic, linear, rarely filiform (seasonally so in C. retrospiralis), mostly glabrous, rarely pubescent, some- times bullate; primary lateral veins pinnate, usually weakly differentiated, running into inconspicuous marginal vein, higher order venation transverse-reticulate. INFLORESCENCE: solitary, usually appearing with the leaves (except C. consob- rina). PEDUNCLE: short, elongating in fruit. SPATHE: tube with connate margins, at base swollen into ellipsoid to cylin- dric “kettle”, kettle rarely constricted, sometimes alveolar within, entrance with a lateral flap partially or completely cov- ering male zone of spadix, upper part of tube long or short (in some species length depending on water depth), rarely absent (C. spiralis), sometimes widening or narrowing at mouth, blade cordate to lanceolate, erect or reflexed at anthesis, spreading, slightly twisted or long-spiralled, apex cuspidate to long-cau- date, inner surface smooth, rough, uniformly warty or with warts in transverse bands, or covered with shortly branched protuberances, white, yellow, red, purple, brown, sometimes spotted, junction with tube usually marked by inconspicuous to very prominent “collar” which normally differs in colour and texture from blade. SPADIX: slender, female zone consisting of a single basal whorl of 4–8 upright gynoecia, with single whorl of rounded, subclavate or irregular olfactory bodies (?pistil- lodes) usually present immediately above, female and male 57. Cryptocoryne zones separated by a long naked axis , or rarely fertile zones C R Y P T O C O R Y N E A E : C R Y P T O C O R Y N E 197

D EF J BG H A K L CM Plate 57. Cryptocoryne. A, habit × 2/3; B, inflorescence × 1; C, inflorescence with spadix detail viewed through opened kettle × 2; D, sta- men × 30; E, female zone of spadix × 6; F, gynoecium, longitudinal section × 6; G, syncarp × 1; H, habit × 2/3; J, inflorescence × 1; K, inflorescence × 1; L, habit × 2/3; M, syncarp × 1. Cryptocoryne spiralis: A, Sivadasan CU 21406 (K); Sivadasan CU 21435 (K); B–G, Bogner 1829 (Kew spirit collection 53996); C. affinis: H, Bogner 1706 (K); J, Bogner 227 (Kew spirit collection 45235); C. ciliata: K, Cult. Kew 1953–203 (Kew spirit collection 16895); C. longicauda: L, Dransfield s.n. (Kew slide collection); M, Bogner 1735 (Kew spirit collection 50159). 198 T H E G E N E R A O F A R A C E A E

has a transverse perforated septum at the mouth of the ket- outer ones smaller; primary lateral veins of lobes pinnate, tle. It thus strongly resembles Lagenandra in this respect. joined into submarginal collective vein at least in upper However, C. spiralis shares with the other species of half of lobe, 1–2 marginal veins also present, higher order Cryptocoryne the connate gynoecia borne in a single whorl, venation reticulate. INFLORESCENCE: 3–5 in each floral the syncarpous infructescence and convolute ptyxis. sympodium, appearing before or with leaves. PEDUNCLE: ETYMOLOGY: Greek kryptos (hidden) and korynê (club); longer than petiole, slender. SPATHE: somewhat constricted, the spadix is entirely enclosed in the spathe tube and thus entirely persistent, tube only laxly convolute, slightly gap- hidden from view. ing at anthesis, obliquely held, blade longer than tube, TAXONOMIC ACCOUNTS: Engler (1920a), Rataj (1975), expanded at anthesis, distinctly or slightly bent forward at Jacobsen (1982, 1985, 1988, 1991), Sivadasan (1982), constriction, ± erect, sometimes fornicate, ovate-lanceolate Jacobsen & Bogner (1987), Jacobsen, Sivadasan & Bogner or lanceolate, acuminate. SPADIX: shorter than spathe, (1989), de Wit (1990), Kasselmann (1995). female zone few- and laxly flowered, adnate to spathe, contiguous with male zone, male zone cylindric, longer C Tribe Zomicarpeae than female, laxly to densely flowered, appendix erect or bent forward near base, clavate to subcylindric, smooth or ± covered with sterile flowers (Z. steigeriana). FLOWERS: Tribe Zomicarpeae Schott, Syn. Aroid. 33 (1856). unisexual, perigone absent. MALE FLOWER: 1–2-androus, Laticifers anastomosing; small, evergreen (Ulearum) or sea- stamens free, filaments distinct or very short, connective dis- sonally dormant (Zomicarpa, Zomicarpella); leaf blade tinctively coloured, thecae ellipsoid, opposite, dehiscing by cordate-sagittate (except Zomicarpa); primary lateral veins apical pore. POLLEN: inaperturate, spherical to subspher- often arising at petiole insertion, higher order venation retic- oidal, medium-sized (mean 32 µm., range 32–33 µm.), exine ulate; spadix longer than spathe (except Zomicarpa, spinose. STERILE FLOWERS: sparse when present, conic Zomicarpella amazonica), with sterile terminal appendix, and acuminate towards base of appendix, otherwise wart- female spadix zone adnate to spathe; flowers unisexual, like. FEMALE FLOWER: gynoecium cylindric or ovoid to perigone absent; stamens free; gynoecia few, ovary 1-locular, subglobose, ovary 1-locular, ovules 6–11, anatropous, funi- ovules anatropous, placenta basal. cle distinct, placenta basal, stylar region shortly attenuate, stigma broad, discoid-hemispheric. BERRY: globose to depressed-globose, whitish below, darker at apex, few–sev- C 58. Zomicarpa eral-seeded. SEED: ovoid to ellipsoid, testa smooth, thin, transparent, micropyle shortly rostrate, funicle strophiolate, swollen, white, embryo axile, elongate, endosperm copious. Zomicarpa Schott, Syn. Aroid. 33 (1856). TYPE: Z. pytho- See Plates 58, 119C. nium (Martius) Schott (Arum pythonium Martius). CHROMOSOMES: 2n = 20. HABIT: small, seasonally dormant herbs, tuber subglobose DISTRIBUTION: 3 spp.; Brazil (Northeast). to depressed-globose. LEAVES: few. PETIOLE: sheath short. ECOLOGY: tropical humid forest, upland gallery forest, BLADE: entire ovate-cordate when juvenile, adult blade tri- deciduous forest; geophytes on forest floor. sect to pedatisect, lobes oblong-elliptic or oblong-lanceolate, ETYMOLOGY: Greek zômê (woman’s girdle) and karpos (fruit); refers to spathe constriction. TAXONOMIC ACCOUNTS: Peyritsch (1879), Engler (1920a), 58. Zomicarpa Bogner (1980d). 59. Zomicarpella C Zomicarpella N.E. Brown in Gard. Chron., ser.2, 16: 266 (1881). TYPE: Z. maculata N.E. Brown HABIT: small herbs with creeping rhizome. LEAVES: 1–3. PETIOLE: sheath short. BLADE: cordate-sagittate to hastate- sagittate, often variegated with pale green blotches; primary lateral veins mostly arising at petiole insertion, running into margin or forming a submarginal collective vein, higher order venation reticulate. INFLORESCENCE: 1–2 in each floral sym- podium, appearing with leaves. PEDUNCLE: slender, longer or shorter than petiole. SPATHE: oblong-lanceolate, fully expanded and unconstricted (Z. maculata), or constricted between tube and blade, tube convolute, blade expanded, erect at first, then reflexed, persistent in fruit (Z. amazonica). SPADIX: longer or shorter than spathe, female zone adnate to spathe, contiguous or separated from male by short axis (Z. maculata), male zone subequal to female, cylindric, ter- minal appendix more slender, ± elongate. FLOWERS: unisexual, perigone absent. MALE FLOWER: 1-androus, sta- mens densely arranged, free, anthers subsessile, filaments short but distinct, connective inconspicuous, thecae opposite, Z O M I C A R P E A E : Z O M I C A R P E L L A 199

B C EF A GD Plate 58. Zomicarpa. A, habit with mature leaf × 1; B, juvenile leaf × 1; C, semi–mature leaf × 1; D, spadix × 5; E, stamen × 20; F, gynoe- cium, longitudinal section × 20; G, infructescence, nearside half of spathe removed × 2. Zomicarpa riedeliana: A, Noblick, Lemos & Valdomiro 3202 (K); B, Cult. Berkley (K); C, Mayo s.n. (K); D–F, Bogner 1213 (Kew spirit collection 42464); G, Schott, Icones Aroideae 3760 (K). 200 T H E G E N E R A O F A R A C E A E

A B D F GH M C L JK E Plate 59. Zomicarpella. A, leaf × 1; B, inflorescence, nearside half of spathe folded back × 3; C, habit × 1; D, leaf × 1; E, inflorescence, nearside half and upper part of spathe removed × 3; F, stamen, top view × 15; G, stamen, side view × 15; H, stamen, longitudinal section × 15; J, gynoecium × 15; K, gynoecium, longitudinal section × 15; L, fruit × 3; M, seed × 4. Zomicarpella maculata: A–B, Cult. Linden (K); Z. amazonica: C–D, Bogner 1985 (K & Kew spirit collection 57529 & 57400); E–M, Bogner 1985 (Kew spirit collection 57529 & 57400). Z O M I C A R P E A E : Z O M I C A R P E L L A 201

59. Zomicarpella insertion, forming a submarginal collective vein, 1–2 mar- ginal veins also present, higher order venation reticulate. subglobose, dehiscing by apical pore. POLLEN: extruded in INFLORESCENCE: solitary. PEDUNCLE: slender, subequal strands, inaperturate, spherical, small (18–20 µm. diam.), exine to or longer than petiole, base enclosed by a single rela- spinose. FEMALE FLOWER: gynoecium ellipsoid-oblong, or tively long cataphyll. SPATHE: oblong-lanceolate, greenish, depressed-bottle-shaped, somewhat laterally compressed, unconstricted, completely expanded, margins revolute, per- ovary 1-locular, ovules 1–6, anatropous, funicle short or very sistent and dark green in fruit. SPADIX: slender, subequal short, placenta basal, stylar region short, distinct, narrower to spathe, female zone adnate to spathe, separated from than ovary, stigma small, discoid-hemispheric. BERRY: male zone by a longer sterile zone, lower part of sterile depressed-globose, whitish, stigma remnant persistent, 1–3- zone naked or only sparsely covered with sterile flowers, seeded. SEED: irregularly ellipsoid, somewhat compressed upper part with short dense zone of staminodes, male zone laterally, testa thin, smooth, whitish, raphe conspicuous, hilum short, composed of 6–7 whorls of stamens, terminal appen- somewhat sunken, purple, embryo ellipsoid to pear-shaped, dix digitiform, base composed of a few whorls of apically small, endosperm copious. See Plates 59, 119D. rounded staminodes, becoming smooth above. FLOWERS: CHROMOSOMES: 2n = 26. unisexual, perigone absent. MALE FLOWER: apparently 1- DISTRIBUTION: 2 spp.; Brazil (western Amazonia), androus, stamens sessile, free, subtruncate, connective ± ?Colombia. broad, thecae ellipsoid to globose, dehiscing by apical ECOLOGY: tropical humid forest (“terra firme”); geophytes pore. POLLEN: inaperturate, spherical, medium-sized on forest floor. (diam. 26–32 µm.), exine spinose. STERILE FLOWERS: NOTES: Zomicarpella maculata N.E. Br. was described from depressed globose or cylindric; staminodes below male a cultivated plant known to have originated in tropical zone in 2–3 whorls, rounded apically. FEMALE FLOWER: America, but without a more exact provenance. No other col- gynoecium narrowly subcylindric, ovary 1-locular, ovule 1, lection of this species has been seen. anatropous, funicle short, placenta basal, stylar region ± as ETYMOLOGY: Zomicarpa and Latin -ella (diminutive); refers thick as ovary, stigma discoid, as broad as style. BERRY: to similarity to Zomicarpa. subcylindric to ellipsoid, stigmatic remnant persistent, 1- TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1979a), seeded, dark green and red-spotted. SEED: ellipsoid, testa Bogner (in press). thin, white, smooth, transparent, embryo ellipsoid, dark green with red spots externally, whitish internally, undif- ferentiated, endosperm absent. See Plates 60, 120A. CHROMOSOMES: 2n = 14. DISTRIBUTION: 1 sp.; Brazil (western Amazonia), Peru (Amazonia). ECOLOGY: tropical humid forest (“terra firme”); geophytes on forest floor. ETYMOLOGY: named after Ernst Ule (1854–1915) and Arum. TAXONOMIC ACCOUNTS: Engler (1920a), Boyce (1995e). C 60. Ulearum 60. Ulearum Ulearum Engler in Bot. Jahrb. 37: 95 (1905, “1906”). TYPE: U. sagittatum Engler HABIT: small evergreen herb, rhizome creeping horizon- tally just below the soil surface. LEAVES: 1–several, 1 marcescent cataphyll enclosing base of petiole. PETIOLE: sheath very short. BLADE: subreniform to broadly sagittate, often variegated; primary lateral veins arising from petiole 202 T H E G E N E R A O F A R A C E A E

F E CD B A Plate 60. Ulearum. A, habit × 2/3; B, inflorescence, nearside half of spathe removed × 3; C, gynoecium × 20; D, gynoecium, longitudinal section × 20; E, staminode × 20; F, stamen, side view × 10. Ulearum sagittatum: A, Ule 6323 (K); B, Jangoux et al. INPA 138864 (Kew spirit collection 56426); C–F, Bogner 1947 (K & Kew spirit collection 56424). Z O M I C A R P E A E : U L E A R U M 203

C 61. Filarum Tribe Caladieae C Filarum Nicolson in Brittonia 18: 348 (1967). TYPE: F. Tribe Caladieae Schott in Schott & Endlicher, Melet. Bot. 18 manserichense Nicolson (“manserichensis”). (1832). HABIT: small herb, tuber subglobose. LEAVES: few. PETI- Laticifers anastomosing; terrestrial, geophytic, rarely aquatic C OLE: sheath short. BLADE: entire, cordate; primary lateral (Jasarum) or climbing hemiepiphytes (Syngonium), stem veins 3 per side, arising from petiole insertion, lower 2 tuberous, rhizomatous or aerial; basal ribs well-developed, pairs retrorse, all arcuately curving upwards forming sub- primary lateral veins of leaf pinnate, forming submarginal marginal collective vein, marginal vein distinct, higher order collective vein, 1 or more marginal veins also usually pre- venation reticulate. INFLORESCENCE: 1–3 in each floral sent, higher order venation reticulate; spadix: female zone sympodium, appearing with leaves. PEDUNCLE: slender, often separated from male by subconical to attenuate zone longer or shorter than petiole. SPATHE: persistent, oblong- of synandrodes (sterile male flowers), male zone sub- lanceolate, pale green. SPADIX: very slender, much longer cylindric to subclavate, sterile appendix absent (except than spathe, female zone adnate to spathe, few-flowered, some Hapaline); flowers unisexual, perigone absent; sta- separated from male zone by longer sterile zone, male zone mens connate into synandria, synandria truncate, margins elongate, purple, sparsely flowered, with a very few sterile sinuous, thecae dehiscing by short apical slit, pollen rudimentary flowers occurring above and below male zone, extruded in strands (except Scaphispatha); ovules anat- appendix slender, naked, elongate. FLOWERS: unisexual, ropous to hemianatropous; endosperm copious (except perigone absent. MALE FLOWER: 1-androus, filaments Jasarum, Syngonium, Hapaline). short, connective extremely long and drawn out into a fil- iform flexuose thread, thecae globose, dehiscing by apical 62. Scaphispatha pore. POLLEN: inaperturate, spherical, small to medium- sized (mean 21 µm.), exine spinose. FEMALE FLOWER: Scaphispatha Brongniart ex Schott, Prodr. syst. Aroid. 214 ovary oblong, 1-locular, ovule 1, anatropous, funicle short, (1860). TYPE: S. gracilis Brongniart ex Schott placenta basal, stylar region attenuate, stigma small, dis- coid. BERRY: ellipsoid, smooth, bearing the style and stigma HABIT: seasonally dormant herb, tuber globose to de- remains. SEED: ellipsoid, testa thin, smooth, punctate, pressed-globose. LEAVES: usually solitary, sometimes 2. embryo ellipsoid, endosperm absent. See Plate 61. PETIOLE: slender, sheath very short. BLADE: ovate-sub- CHROMOSOMES: 2n = 28. cordate to ovate-sagittate, peltate, apex cuspidate- DISTRIBUTION: 1 sp.; Peru (Amazonia). acuminate; basal ribs well-developed, primary lateral veins ECOLOGY: tropical humid forest; geophytes on forest floor. pinnate, forming submarginal collective vein, 1–2 marginal ETYMOLOGY: Latin filum (thread) and Arum; named after veins also present, higher order venation reticulate. INFLO- the thread-like spadix and stamen connectives. 61. Filarum 62. Scaphispatha 204 T H E G E N E R A O F A R A C E A E

E B D F G A HC Plate 61. Filarum. A, habit × 1; B, leaf × 1; C, inflorescence, nearside spathe base folded back × 4; D, stamen, side view × 16; E, stamen, view from below × 16; F, gynoecium × 16; G, gynoecium, longitudinal section × 16; H, fruit × 10. Filarum manserichense A, D–G, Wurdack 2402 (K); B–C, H, Mexia 6353 (K). Z O M I C A R P E A E : F I L A R U M 205

H E FG DB C A Plate 62. Scaphispatha. A, habit × 1; B–C, inflorescences × 1; D, spadix × 5; E, synandrium, top view × 30; F, synandrium, side view × 30; G, gynoecium, longitudinal section × 30; H, infructescence × 1. Scaphispatha gracilis: A–C, Hatschbach & Koczicki 33081 (K); D–H, Bogner 1211 (Kew spirit collection 42451). 206 T H E G E N E R A O F A R A C E A E

RESCENCE: solitary, appearing well before leaf. PEDUNCLE: very long, slender. SPATHE: decurrent on peduncle, slightly constricted between tube and blade, light green without, white within, tube convolute before and after anthesis, gap- ing at anthesis, persistent to fruiting stage, blade fully expanded at anthesis, at first erect, later reflexed, marces- cent after anthesis. SPADIX: sessile, cylindric, hardly constricted, shorter than spathe, fertile to apex, densely flowered, male and female zones contiguous. FLOWERS: unisexual, perigone absent. MALE FLOWER: 4-androus, sta- mens connate into truncate, deeply lobed synandrium, connective not greatly thickened, thecae lateral, dehiscing apically by broad, pore-like slit. POLLEN: inaperturate, spherical to subspheroidal, medium-sized (mean 45 µm.), exine coarsely verrucate with large polygonal flat-topped verrucae. FEMALE FLOWER: ovary ovoid, 1-locular, ovules 3–5, anatropous, funicle short, placenta basal, stylar region distinct, attenuate, much narrower than ovary, stigma small, discoid-subcapitate, only slightly broader than style. BERRY: subglobose to obovoid, always 1-seeded, stigma remnants persistent, fruiting very rapidly (ca.10 days after anthesis), whitish-grey. SEED: subglobose, testa smooth and thin, greyish to brown, covered with minute brownish spots, raphe pronounced, swollen, embryo elongate, straight, endosperm copious. See Plates 62, 120B. CHROMOSOMES: 2n = 28. DISTRIBUTION: 1 sp.; Bolivia, Brazil (southern Amazonia, Central West, Northeast). ECOLOGY: tropical seasonal and dry forest, open wood- 63. Caladium land; geophytes, temporarily wet places, flowering at beginning of rainy season after first rains. ETYMOLOGY: Greek skaphê (boat, bowl) and -ion (diminu- more than twice as long as female zone. FLOWERS: unisex- tive) and “spathê” (spathe); refers to spathe shape. ual, perigone absent. MALE FLOWER: 3–5-androus, stamens TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1980d). connate into obpyramidal, truncate to ± convex, sinuously subhexagonal synandrium, common connective thick, thecae lateral, extending nearly to base of synandrium, oblong- C 63. Caladium lanceolate, dehiscing by short apical slit. POLLEN: extruded in strands, shed in monads, inaperturate, spherical to sub- Caladium Ventenat, Descript. Pl. Nouv. Jard. Cels, 30 (1801). spheroidal, medium-sized (mean 41 µm., range 38–43 µm.), LECTOTYPE: C. bicolor (Aiton) Ventenat (Arum bicolor Aiton, exine psilate or obscurely dimpled or verruculate. STERILE see Hubbard & Rehder, Bot. Mus. Leafl. 1: 3. 1932). MALE FLOWERS: synandrodes depressed-obpyramidal, com- pressed, truncate, lowermost often larger and prismatic, SYNONYMS: Phyllotaenium André in Ill. Hort. 19: 3 uppermost narrow and elongated. FEMALE FLOWER: gynoe- (1872); Aphyllarum S. Moore in Trans. Linn. Soc. London, cium cylindric to obconic, ovary 1–2(–3)-locular, ovules 1–20, Bot. ser. 2, 4: 501 (1895). anatropous, funicle short, placentae subbasal, pseudoaxile or HABIT: seasonally dormant or evergreen herbs, stem tuber- parietal, stylar region free or rarely coherent (C. paradoxum), ous, subglobose. LEAVES: several. PETIOLE: sheath distinct. as broad as ovary or broader in obconic gynoecia, stigma BLADE: usually peltate, sometimes not (C. lindenii), often nearly as wide as ovary. BERRY: 1–many-seeded (C. terna- variegated, cordate-sagittate or sagittate, rarely trisect (C. ter- tum usually 1-seeded), white. SEED: ovoid to ellipsoid, raphe natum); basal ribs well-developed, primary lateral veins somewhat prominent, integument succulent, testa thick, lon- pinnate, forming submarginal collective vein, 1–2 marginal gitudinally costate, costae angled, embryo axile, elongate, veins also present, secondary and tertiary laterals arising endosperm copious. See Plates 63, 120C. from the primaries at a wide angle, forming interprimary col- CHROMOSOMES: 2n = 22, 26, 28, 30, 32. lective vein, higher order venation reticulate. DISTRIBUTION: ca. 12 spp.; tropical America, West Indies:– INFLORESCENCE: 1–2 in each floral sympodium, appearing N. Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, with or before leaves. PEDUNCLE: rather long, often as long French Guiana, Guyana, Lesser Antilles, Panama, Peru, Puerto as petiole. SPATHE: constricted, tube with convolute margins, Rico, Surinam, Venezuela. usually ventricose-globose, persistent, green, eventually split- ECOLOGY: tropical humid forest, open woodlands; geo- ting in fruit, blade white, boat-shaped, gaping, marcescent phytes, forest floor, river margins and damp sites. after anthesis and deciduous. SPADIX: a little shorter than NOTES: Caladium bicolor and its many cultivated varieties spathe, stipitate or sessile, densely flowered, female zone are cultivated throughout the world as ornamental plants. cylindric-conoid or ellipsoid, separated from male by longer, This species occurs throughout tropical America. subconic to attenuate, basally thicker zone of sterile male ETYMOLOGY: Latinization of Malay word keladi, a vernacu- flowers, male zone fertile to apex, subcylindric to subclavate, lar name in Malaysia for several colocasioid genera. C A L A D I E A E : C A L A D I U M 207

B S J N A QR P M D G E LK H FC Plate 63. Caladium. A, leaf × 1/3; B, base of plant × 1/2; C, spadix × 1; D, synandrium × 10; E, upper synandrode × 5; F, lower synandrode × 5; G, gynoecium × 10; H, gynoecium, transverse section × 10; J, infructescence × 2/3; K, leaf × 1/3; L, spadix × 1; M, inflorescence × 2/3; N, synandrium × 10; P, synandrode × 10; Q, gynoecium × 10; R, gynoecium, longitudinal section × 10; S, habit × 1/5. Caladium aristeguietae: A–B, Bunting 2201 (K); C–H, Bunting 2201 (Kew spirit collection 58022); J, Bunting 4521 (K); C. coerulescens: K, Bunting 4980 (K); L, Bunting 4865 (K); C. lindenii: M–R, Cult. Kew 1982–02775 (Kew spirit collection 39521); C. bicolor: S, Lowe 3.1962, 7220 (Kew slide collection). 208 T H E G E N E R A O F A R A C E A E

TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), exine foveolate-reticulate. FEMALE FLOWER: ovary cylin- Madison (1976b, 1981), Bogner (1984d), Grayum (1986c), dric, 1-locular, ovules (1–)2–3(–4), anatropous, funicle Mayo & Bogner (1988). distinct, placenta basal, stylar region broader than ovary, stigma discoid-hemispheric. BERRY: subovoid, tightly packed, dull maroon- green apically, stigma remains per- C 64. Jasarum sistent, pericarp fleshy, usually 1-seeded, sometimes 2-seeded. SEED: large, obovoid, compressed when paired, Jasarum Bunting in Acta Bot. Venezuelica 10: 264 (1977, testa smooth, embryo globular, large, green, plumule well- “1975”). TYPE: J. steyermarkii Bunting developed, raphe prominent, endosperm nearly absent. See Plates 64, 120D. HABIT: evergreen aquatic herb, stem thick, hypogeal, CHROMOSOMES: 2n = 22. upright, annulate, terete, surface brown, white within, usu- DISTRIBUTION: 1 sp.; tropical South America: Venezuela, ally unbranched, roots thick, somewhat spongy. LEAVES: Guyana. 6–12, dark green in all parts except whitish hypogeal lower ECOLOGY: tropical uplands; submerged aquatic in oligo- part of petiole. PETIOLE: sheath long, fleshy. BLADE: lin- trophic (blackwater) streams. ear, somewhat bullate, acute to cuspidate, cuneate at base, ETYMOLOGY: named after Julian A. Steyermark (1909–1988). midrib pronounced; primary lateral veins pinnate, numer- TAXONOMIC ACCOUNTS: Bunting (1977), Bogner (1985b). ous, 35–60 per side, joined into submarginal collective vein, marginal vein also present, higher order venation reticulate. INFLORESCENCE: solitary, held above water 65. Xanthosoma C level. PEDUNCLE: much longer than petiole. SPATHE: erect, constricted between tube and blade, tube convolute, per- Xanthosoma Schott in Schott & Endlicher, Melet. Bot. 19 sistent, blade long-acuminate, never reflexed, marcescent (1832). LECTOTYPE: X. sagittifolium (L.) Schott (“sagittae- after anthesis and later deciduous, bronze-green outside, folium”; Arum sagittaefolium L.; see Nicolson 1975). cream-coloured within. SPADIX: shorter than spathe, sub- cylindric, basal half of female zone adnate to spathe, SYNONYMS: Acontias Schott in Schott & Endlicher, Melet. separated from male by zone of sterile flowers (synan- Bot. 19 (1832); Cyrtospadix K. Koch in Index Sem. Hort. drodes), male zone fertile to apex. FLOWERS: unisexual, Berol. 1853 App.: 13 (1853). perigone absent. MALE FLOWER: 3–4-androus, stamens Latex milky. HABIT: small to gigantic, sometimes arbores- connate into synandrium, synandrium truncate, irregularly cent, evergreen or seasonally dormant herbs, stem either a 4–6-angled, thecae 6–8, lateral, elongate, dehiscing by api- thick, subcylindric, hypogeal tuber often producing smaller cal pore. POLLEN: ellipsoid, medium-sized (mean 44 µm.), tubers on stoloniferous side branches or often with a dis- tal, epigeal, massive, arborescent upper part, or an entirely hypogeal subglobose tuber, bearing many very small tuber- cles in some spp. (X. pubescens, X. viviparum), tuber sometimes yellow within. LEAVES: several, rarely pubes- cent. PETIOLE: sheath usually rather long. BLADE: cordate, sagittate, hastate, trifid, trisect, pedatifid or pedatisect, rarely linear-lanceolate to ovate with emarginate base, rarely peltate; basal ribs well-developed, often denuded proxi- mally, primary lateral veins pinnate, forming submarginal collective vein, 1 or more distinct marginal veins also pre- sent, secondary and tertiary laterals arising from the primaries at a wide angle, forming interprimary collective vein, higher order venation reticulate. INFLORESCENCE: 1 to many in each floral sympodium, always appearing with leaves. PEDUNCLE: usually rather short, rarely long. SPATHE: strongly constricted, tube with convolute margins, ovoid to ellipsoid, usually ventricose, rather thick-walled in large species, persistent, blade boat-shaped-oblong to - oblong-lanceolate, gaping and erect or sometimes reflexed at anthesis, marcescent after anthesis and then deciduous. SPADIX: shorter than spathe, densely flowered, female zone cylindric-conoid, often obliquely inserted onto peduncle, separated from male zone by longer, conoid to attenuate, basally thicker zone of sterile male flowers, male zone cylin- dric-conoid, longer than female zone, usually fertile to apex, rarely with a few sterile flowers at extreme apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 4–6-androus, stamens connate into a truncate-obpyramidal, subpentago- nal or hexagonal synandrium, anthers lateral, nearly reaching base of synandrium, common connective thick, 64. Jasarum thecae oblong or tapering basally, dehiscing by subapical pore or short slit. POLLEN: extruded in strands, shed in C A L A D I E A E : X A N T H O S O M A 209

J B H E G F A DC Plate 64. Jasarum. A, habit, peduncle partly removed × 1/2; B, detail of leaf venation × 2/3; C, inflorescence × 2/3; D, spadix × 2; E, synan- drium, side view × 15; F, gynoecium × 15; G, gynoecium, longitudinal section × 15; H, infructescence × 2/3; J, berry, side view × 2. Jasarum steyermarkii: A, Bunting 4628 (K); Herkner & Jeschke s.n. (Kew spirit collection 29047.618); Aroideana 8(2): 55–63, f.4 (1985); B–J, Cult. Munich (Kew spirit collection 57524); Herkner & Jeschke s.n. (Kew spirit collection 29047.618). 210 T H E G E N E R A O F A R A C E A E

tetrads, inaperturate, ellipsoid to subspherical, medium- VERNACULAR NAMES AND USES: “cocoyam”, “tannia”; X. C sized (mean 42 µm., range 35–49 µm., tetrads:– mean 76 sagittifolium and its many varieties are important subsis- µm., range 62–97 µm.), exine minutely punctate- or fossu- tence food plants throughout the humid tropics of the world late-verruculate. STERILE MALE FLOWER: composed of due to their starch- and protein-rich tuberous stems (see obpyramidal truncate, laterally compressed synandrodes, chapter 16). lowermost larger, uppermost narrower and more elongated ETYMOLOGY: Greek “xanthos” (yellow) and “soma, (in sense of spadix axis). FEMALE FLOWER: ovary ovoid, somatos” (body); refers to the yellow colour of the stem tis- 2–4-locular, more rarely 1-locular, ovules (12–)20 to very sue present in several species. numerous, anatropous or hemianatropous, funicles rather TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), long, placentae usually pseudoaxile, or sometimes parietal Madison (1976b, 1981), Bogner (1986b), Grayum (1986c), or axile basally, stylar region broader than ovary, usually Mayo & Bogner (1988), Okeke (1992). discoid-thickened and coherent to weakly connate with those of neighbouring flowers, rarely free and swollen (X. 66. Chlorospatha plowmanii), stigma subhemispheric or 2–4-lobed, yellow, narrower than style. BERRY: cylindric, somewhat furrowed Chlorospatha Engler in Gartenflora 27: 97 (1878). TYPE: C. apically, whitish to orange, many-seeded. SEED: ovoid, kolbii Engler testa costate, embryo axile, subequal to endosperm, endosperm copious. See Plates 65, 121A. SYNONYM: Caladiopsis Engler in Bot. Jahrb. 37: 139 CHROMOSOMES: 2n = 22, 26, 39, 52. (1905). DISTRIBUTION: ca. 57 spp.; tropical and southern subtrop- ical America, West Indies:– N. Argentina, Bolivia, Brazil, Latex milky. HABIT: small to medium, usually evergreen Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, herbs, stem epigeal to ± subterranean, acaulescent to elon- El Salvador, French Guiana, Guatemala, Guyana, Haiti, gate, decumbent to erect. LEAVES: 1 to several. PETIOLE: Honduras, Jamaica, Lesser Antilles, Mexico, Nicaragua, sheath long. BLADE: cordate, sagittate, hastate, trifid, tri- Panama, Paraguay, Peru, Puerto Rico, Surinam, Trinidad, sect, pedatifid or pedatisect; basal ribs well-developed, Venezuela. primary lateral veins pinnate, forming submarginal collective ECOLOGY: tropical moist and humid forest, subtropical for- vein, 1 or more distinct marginal veins also present, sec- est; geophytes on forest floor, in wet places, swamps, river ondary and tertiary laterals arising from the primaries at a banks, seasonally flooded sites, grassy places, plantations, wide angle, forming interprimary collective vein, higher some species are weedy. order venation reticulate. INFLORESCENCE: 3–8 in each flo- NOTES: Engler in Engler & Krause (1920) recognized 2 sec- ral sympodium. PEDUNCLE: very slender, much shorter than tions:– section Xanthosoma, section Acontias. petiole and supported by sheath. SPATHE: constricted, tube 65. Xanthosoma 66. Chlorospatha C A L A D I E A E : C H L O R O S P A T H A 211

HC J A K P D F E QR S N B ML G Plate 65. Xanthosoma. A, leaf × 1/4; B, base of plant × 1/5; C, inflorescence × 1/5; D, detail of synandria, top view × 5; E, detail of gynoe- cia, top view × 5; F, synandrium, side view × 5; G, gynoecia, left hand gynoecium in longitudinal section × 5; H, infructescence × 1/4; J, leaf × 1/3; K, leaf × 1/2; L, inflorescence, nearside half of spathe removed × 2/3; M, detail of synandrodes, top view × 15; N, synandrode, side view × 15; P, synandrium, side view × 15; Q, gynoecium × 15; R, gynoecium, longitudinal section × 15; S, ovules × 25. Xanthosoma violaceum: A–B, Engler 245 (K); C, Vollesen (Kew slide collection); D–H, Skvortzov (Kew spirit collection 29047.189); X. striatipes: J, Krapovickas & Cristóbal 35351 (K); X. helleborifolium: K, Broadway 6340 (K); X. plowmanii: L–S, Plowman et al. 8460 (Kew spirit collection 49708). 212 T H E G E N E R A O F A R A C E A E

C K DE J F B M L A GH Plate 66. Chlorospatha. A, habit × 1/2; B, spadix × 2; C, synandrium, side view × 15; D, synandrode, side view × 15; E, gynoecium, side view × 15; F, gynoecium, longitudinal section × 15; G, leaf × 1/4; H, inflorescences with associated leaf × 1/2; J, inflorescence × 2; K, gynoe- cium, longitudinal section × 15. L, spadix × 2; M, gynoecium, side view × 15. Chlorospatha longipoda: A, Plowman 14063 (K); B–F, Plowman 3979 (Kew spirit collection 49717); C. croatiana subsp. enneaphylla: G, Lawrence 794 (K); C. atropurpurea: H, Dodson 5911 (K); C. corru- gata: J–K, Spear s.n. (Kew spirit collection 42452); C. kolbii : L–M, Franke s.n. (Kew spirit collection 62897). C A L A D I E A E : C H L O R O S P A T H A 213

with convolute margins, narrow, elongate, ± cylindric to narrowly ellipsoid, persistent, blade boat-shaped to more widely expanded, sometimes fornicate, rather narrowly ellip- tic to lanceolate, marcescent after anthesis and deciduous. SPADIX: female zone free or adnate to spathe, laxly or densely flowered, separated from male zone by longer or shorter laxly or densely flowered sterile zone, male zone densely flowered, fertile to apex. FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–5-androus, stamens con- nate into truncate synandrium, deeply or shallowly-lobed, fused connectives thickened, thecae extending almost to base of synandrium, oblong, dehiscing by short longitudinal slit. POLLEN: extruded in strands, shed in tetrads, inapertu- rate, spherical or subspheroidal, medium-sized (mean 26 µm., range 24–29 µm., tetrad mean 45 µm., range 41–48 µm.), exine psilate or very obscurely punctate to obscurely verruculate to foveolate-reticulate. STERILE FLOWERS: sta- minodes either free or partially or completely connate into irregular, fungiform or 3–4 lobed synandrodes, rarely pris- matic. FEMALE FLOWER: gynoecium ± ovoid to subhemispheric, ovary (1–)2–4(–5)- locular, ovules several per locule, anatropous to hemianatropous, funicle rather long, placentae pseudoaxile or axile, rarely basal (C. longipoda), stylar region short to relatively long, usually expanded into thin, spreading, evanescent mantle contigu- ous with neighbouring ones, usually containing numerous red chromoplasts, more rarely style with only a short, rim- 67. Syngonium like lateral outgrowth (C. longipoda, C. mirabilis) or with soft tuberculate outgrowth (C. kolbii), stigma subhemispheric or subcapitate or slightly lobed. BERRY: depressed-globose, arcuate and running into margin (e.g. S. schottianum), sec- somewhat 3–5-furrowed, stigma remnant persistent, many- ondary laterals ± parallel to primaries, higher order venation seeded, white to yellowish. SEED: minute, ovoid to ellipsoid, reticulate to transverse-reticulate. INFLORESCENCE: 1–8 in white, testa longitudinally costate, embryo elongate, axile, each floral sympodium. PEDUNCLE: much shorter than peti- endosperm copious. See Plates 66, 121B. ole, erect at anthesis, pendent in fruit. SPATHE: strongly CHROMOSOMES: 2n = 26. constricted between tube and blade, tube convolute, ovoid- DISTRIBUTION: ca. 16 spp.; tropical America:– Colombia, ellipsoid, or cylindric or globose, sometimes ventricose, Costa Rica, Ecuador, Panama, Peru. persistent, blade usually cream, boat-shaped to expanded ECOLOGY: tropical humid forest; terrestrial on forest floor, at anthesis, erect to spreading, usually marcescent after well shaded creek beds or in boggy areas. anthesis, later deciduous. SPADIX: sessile, much shorter NOTES: Phylogenetically, Chlorospatha may be a derived than spathe, female zone cylindric to conoid, separated offshoot of Xanthosoma, and C. longipoda is somewhat inter- from male zone by zone of sterile flowers, sterile zone con- mediate between the two genera. stricted, male zone clavate to cylindric or ellipsoid, longer ETYMOLOGY: Greek chlôros (green) and spathê (spathe). than female zone, fertile to apex. FLOWERS: unisexual, TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), perigone absent. MALE FLOWER: 3–4-androus, stamens Madison (1981), Bogner (1985a), Grayum (1986c, 1991a), connate into synandrium, synandrium obpyramidal, truncate Mayo & Bogner (1988). to rounded at apex, sometimes prominent above thecae, often deeply lobed, common connective thick, overtopping anthers, thecae oblong, dehiscing by apical pore or short C 67. Syngonium slit. POLLEN: extruded in strands, inaperturate, ellipsoid- oblong to spherical or subspheroidal, medium-sized (mean Syngonium Schott in Wiener Z. Kunst 1829 (3): 780 (1829). 48 µm., range 34–75 µm.), exine minutely fossulate or ver- TYPE: S. auritum (L.) Schott (Arum auritum L.). ruculate, narrowly rugulate and minutely scabrate, spinose or tuberculate. STERILE MALE FLOWERS: synandrodes ± SYNONYM: Porphyrospatha Engler in A. & C. de Candolle, prismatic. FEMALE FLOWER: gynoecia connate, rarely ± Monogr. Phan. 2: 289 (1879). free at anthesis and becoming connate later (S. steyermarkii), Latex milky. HABIT: root-climbing or shortly creeping, ever- ovary obovoid or oblong-obovoid, (1–)2(–3)-locular, ovules green herbs, internodes short to elongate, green. LEAVES: 1(–2) per locule, anatropous, funicle short, placenta axile numerous. PETIOLE: sheath long. BLADE: at maturity cor- near septum base to parietal-basal, stylar region as broad as date, oblong-cordate, lanceolate, sagittate, trifid to trisect, ovary and ± truncate or very slightly attenuate, stigma dis- pedatifid to pedatisect, rarely pinnatifid, when juvenile coid or 2-lobed, rarely globose, discoid-capitate or entire, ovate to sagittate or cordate-sagittate or hastate; basal cup-shaped, narrower than ovary. BERRIES: connate, form- ribs usually well-developed, primary lateral veins pinnate, ing ovoid to ellipsoid, usually brown, sometimes white, usually forming a submarginal collective vein, 1–2 distinct fleshy syncarp, usually exposed at maturity by opening of marginal veins also present, or sometimes primaries long- persistent orange, yellow, red or violet spathe tube. SEED: 214 T H E G E N E R A O F A R A C E A E

A D L MB J C H K FE G Plate 67. Syngonium. A, leaf × 1/3; B, inflorescence × 1/2; C, habit × 1/2; D, infructescences and associated stem × 1/2; E, leaf × 1/3; F, inflorescence × 1/2; G, leaf × 1/3; H, spadix × 1; J, synandrium, side view × 8; K, detail of gynoecia, top view × 4; L, gynoecium, longitudi- nal section × 6; M, gynoecium, transverse section × 6. Syngonium steyermarkii: A, 201/12 (Kew slide collection); B, Croat 47201 (Kew spirit collection 56140); S. armigerum: C, Grayum 6677 (K); S. podophyllum: D, Silverstone-Sopkin et al. 5468 (K); S. neglectum: E–F, Bunting 1658 (K); S. schottianum: G, Cult. Kew June 1878 (K); H–K, Sugden 425 (Kew spirit collection 29047.347); S. vellozianum: L–M, Harley et al. 17874 (Kew spirit collection 46608). C A L A D I E A E : S Y N G O N I U M 215

ovoid to ellipsoid, rather large, testa smooth, thin, black or dark brown, shiny, embryo large, ellipsoid to subglobose, endosperm absent. See Plates 67, 121C. CHROMOSOMES: 2n = 28 (24, 26). DISTRIBUTION: 35 spp.; tropical America, West Indies:– Belize, Bolivia, Brazil, Colombia, Costa Rica, ?Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, ?Lesser Antilles, Mexico, Nicaragua, Panama, Peru, Surinam, Trinidad, Venezuela. ECOLOGY: tropical humid forest and disturbed areas; climb- ing epiphytes and hemiepiphytes, creeping when juvenile. NOTES: Croat (1982) recognizes 4 sections:– sect. Oblongata, sect. Cordata, sect. Pinnatiloba, sect. Syngonium. ETYMOLOGY: Greek syn- (together), gonê (womb) and -ion (diminutive); refers to connate gynoecia. TAXONOMIC ACCOUNTS: Engler & Krause (1920), Croat (1982). C 68. Hapaline Hapaline Schott, Gen. Aroid. t. 44 (1858), nom. cons. TYPE: 68. Hapaline H. benthamiana Schott SYNONYM: Hapale Schott in Oesterr. bot. Wochenbl. 7: 85 (1857). HABIT: small to moderate, slender, seasonally dormant or ETYMOLOGY: modification of the feminine form of Greek C evergreen herbs, tubers small, depressed-globose. LEAVES: hapalos (soft), referring to the small tender nature of this C 1–few, usually solitary. PETIOLE: sheath short. BLADE: cor- genus. date-sagittate, sagittate or hastate, sometimes pale green or TAXONOMIC ACCOUNTS: Engler in Engler & Krause (1920), silvery variegated; primary lateral veins pinnate or mostly Li (1979), Bogner (1984a), Boyce (1996). arising at petiole insertion, forming arching submarginal col- lective vein, 1–2 marginal veins also present, higher order Tribe Nephthytideae venation reticulate. INFLORESCENCE: 1–2 in each floral sym- podium, appearing with leaf. PEDUNCLE: subequal or longer Tribe Nephthytideae Engler in Engler & Prantl, Nat. than petiole, slender. SPATHE: slender, not distinctly con- Pflanzenfam. II (3): 112, 128 (1887). stricted, tube very slender, persistent, tightly convolute around female flowers, blade longer, oblong-lanceolate, erect to Laticifers simple, articulated; geophytic, stem tuberous to reflexed and ± revolute at anthesis, marcescent. SPADIX: sube- rhizomatous; petiole geniculate apically; leaf blade sagittate qual or longer than spathe, slender, female zone adnate to or trifid to trisect and highly divided, primary lateral veins spathe, few-flowered (2–7), ± biseriate, separated from male of ultimate lobes or divisions pinnate, higher order vena- zone by short sterile zone, male zone subulate to cylindric, fer- tion reticulate; spathe boat-shaped or fully expanded, not tile to apex or with a few sterile flowers at apex or with a long, clearly differentiated into tube and blade; spadix cylindric, terminal appendix (H. appendiculata). FLOWERS: unisexual, female and male zones usually contiguous, sterile flowers perigone absent. MALE FLOWER: 3-androus, synandrium ± absent (except Pseudohydrosme); flowers unisexual, peltate, truncate, hexagonal, elongated in sense of spadix perigone absent; stamens free with filaments lacking or axis, shallow, stipitate, connective strongly dilated, thecae very short (except Nephthytis); ovules 1 per locule, anat- short, subglobose, remote, almost pendent from margin, ropous, placenta basal (except Pseudohydrosme); berries dehiscing by pore. POLLEN: inaperturate, spherical, medium- and seeds large, testa thin to absent, endosperm absent sized (mean 40 µm.), exine rather densely spinose. STERILE (fruit unknown in Pseudohydrosme). Chromosomes large to FLOWER: lower ones very few, ± remote, apiculiform, upper very large. ones very few, consisting of tiny, peltate synandrodes. FEMALE FLOWER: gynoecium oblong to lageniform, ovary 1- 69. Nephthytis locular, ovule 1, anatropous, funicle very short, placenta parietal to subbasal (morphologically basal), stylar region Nephthytis Schott in Oesterr. bot. Wochenbl. 7: 406 (1857). very short, stigma subcapitate. BERRY: ellipsoid to globose, TYPE: N. afzelii Schott style persistent, pericarp thin, 1-seeded, white. SEED: ellipsoid, testa smooth, very thin, embryo large, ellipsoid, light green, SYNONYM: Oligogynium Engler in Bot. Jahrb. 4: 64 endosperm absent. See Plates 68, 121D. (1883). CHROMOSOMES: 2n = 26, 28. DISTRIBUTION: 7 spp.; tropical southeast Asia, Malay HABIT: evergreen, rarely seasonally dormant, small to Archipelago:– Brunei, Burma, ?Cambodia, China (Yunnan), medium-sized herbs, rhizome creeping, subepigeal, rarely Laos, Malaysia (Borneo, Peninsula), Thailand, Vietnam. hypogeal, slender to thick, internodes very short, rarely long. ECOLOGY: tropical humid forest; geophytes on forest floor or humus deposits on limestone or basalt rocks. 216 T H E G E N E R A O F A R A C E A E

D A C GF E B K HL J Plate 68. Hapaline. A, habit × 2/3; B, leaf × 2/3; C, inflorescence × 2; D, synandrium, top view × 10; E, synandrium, side view × 10; F, gynoe- cium × 15; G, gynoecium, longitudinal section × 15; H, infructescence × 2; J, habit × 2/3; K, leaf × 2/3; L, leaf × 2/3. Hapaline celatrix: A–H, Boyce 417 (Kew spirit collection 57283 & Kew slide collection); H. benthamiana: J, Kerr 614 (K); H. ellipticifolia: K, Sun s.n. (YUKU); H. colaniae: L, Pételot 2919 (K). C A L A D I E A E : H A P A L I N E 217

LEAVES: several, rarely solitary. PETIOLE: geniculate apically TAXONOMIC ACCOUNTS: Bogner (1980c), Knecht (1983), C (geniculum very long in N. bintuluensis), usually smooth, Ntépé-Nyame (1988), Namur & Bogner (1994), Hay, Bogner rarely sparsely aculeate, uniformly green, sheath very short. & Boyce (1994). BLADE: cordate-sagittate to sagittate-trifid or subtriangular, posterior divisions usually longer than anterior, all three divi- 70. Anchomanes sions often long-acuminate; basal ribs usually well-developed, primary lateral veins of both anterior and Anchomanes Schott in Oesterr. bot. Wochenbl. 3: 314 posterior divisions pinnate forming submarginal collective (1853). TYPE: A. hookeri Schott, nom. illeg. (= Caladium vein, higher order venation reticulate. INFLORESCENCE: soli- petiolatum W.J. Hooker, A. petiolatus (W.J. Hooker) tary, usually flowering with leaves, rarely without (N. Hutchinson). bintuluensis). PEDUNCLE: usually relatively slender, shorter or equalling petiole. SPATHE: ovate- to oblong-elliptic, fully HABIT: herbs, often very robust, tuber small to gigantic, expanded, spreading to reflexed at anthesis, not constricted, erect or grossly rhizomatous, seasonally dormant. LEAF: soli- decurrent on peduncle, green, persistent or marcescent. tary, often gigantic. PETIOLE: very long, terete, aculeate, SPADIX: short- to long-stipitate, cylindric, female zone con- rarely smooth, sheath very short. BLADE: sagittate when tiguous with male, rarely with a few sterile flowers in juvenile, becoming dracontioid at maturity: i.e. trisect, pri- between, male zone longer than female, fertile to apex. mary divisions each further divided ± dichotomously or FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–4- pinnately, secondary divisions irregularly pinnatifid, ulti- androus, stamens free, obpyramidal, filaments thick, well mate lobes very variable in size and shape, distal ones larger, developed (except N. hallaei, N. mayombensis), connective trapezoid, apically broader, truncate or shallowly bifid, thick, thecae lateral, dehiscing by apical pore. POLLEN: pre- decurrent to sessile, proximal lobes ovate and acuminate; sented in amorphous mass, inaperturate, ellipsoid to primary lateral veins of ultimate lobes pinnate, long-arcuate, subsphaeroidal, medium-sized (mean 46 µm.), exine verru- mostly running into margin, sometimes forming irregular cate. FEMALE FLOWER: ovary ovoid, 1-locular, ovule 1, submarginal collective vein, higher order venation reticulate. anatropous, funicle very short, placenta basal, stylar region INFLORESCENCE: solitary, usually appearing before leaf. shortly attenuate or inconspicuous, stigma discoid-hemi- PEDUNCLE: aculeate, rarely smooth, shorter than petiole. spheric. BERRY: large, subglobose to ellipsoid, orange, SPATHE: erect, broadly ovate to narrowly oblong-lanceolate pericarp thick. SEED: obovoid or subglobose to ellipsoid, or oblong-elliptic, boat-shaped, not constricted, convolute smooth, testa absent at maturity, embryo large, outer cell lay- basally or not at all, apex sometimes fornicate, marcescent. ers with chlorophyll, endosperm absent. See Plates 69, 122A. SPADIX: much shorter or subequal to spathe, sessile to sub- CHROMOSOMES: 2n = 36, 40, 60, karyology very similar to sessile, cylindric, female zone subequal to male zone or that of Anchomanes. much shorter, male zone contiguous with female, fertile to DISTRIBUTION: 10 spp.; tropical west and central Africa:– apex. FLOWERS: unisexual, perigone absent. MALE ?Benin, Cameroon, Congo, Equatorial Guinea (Bioko, Rio FLOWER: stamens free, anthers sessile or filaments short, Muni), Gabon, Ghana, Ivory Coast, Liberia, Malaysia compressed, connective slender below, thickened and (Sarawak), Nigeria, Sierra Leone, ?Togo. dilated apically, thecae ovate-oblong, opposite, dehiscing by ECOLOGY: tropical humid forest; geophytes on forest floor. apical slit. POLLEN: extruded in strands, inaperturate, ellip- ETYMOLOGY: after mythic Nephthys, mother of Anubis and soid to ellipsoid-oblong, large (mean 64 µm., range 37–94 wife of Typhon. 69. Nephthytis 218 T H E G E N E R A O F A R A C E A E

A E LM F G B JK D C W Q V RS H NP TU Plate 69. Nephthytis. A, habit × 1/2; B, stamen × 10; C, stamen, longitudinal section × 10; D, gynoecium × 10; E, gynoecium, longitudinal section × 10; F, habit × 1/2; G, leaf × 1/2; H, spadix × 1; J, stamen × 10; K, stamen, longitudinal section × 10; L, gynoecium × 10; M, gynoe- cium, longitudinal section × 10; N, habit × 1/2; P, habit × 1/5; Q, inflorescence × 1; R, stamen × 10; S, stamen, longitudinal section × 10; T, gynoecium × 10; U, gynoecium, longitudinal section × 10; V, infructescence × 1; W, seed, hilum view × 1. Nephthytis hallaei: A, Bogner 750 (K & Kew spirit collection 46624); B–E, Bogner 750 (Kew spirit collection 54219); N. swainei: F, Swaine et al. GC 44621 (K); N. poissonii: G, Bogner 605 (K); H–M, Cult. Kew 1957–43801 (Kew spirit collection 51366); N. constricta: N, Brenan 8527 (K); N. afzelii: P, V–W, Knecht 6 (Cult. Kew 1982–4608); Q, Knecht 6 (Kew spirit collection 46572); R–U, Knecht 6 (Kew spirit collection 46103). N E P H T H Y T I D E A E : N E P H T H Y T I S 219

E F DD D EE R P FF K AA C H J Q CC BB M S T W X LG N AB ZY VU Plate 70. Anchomanes. A, habit in flower × 1/10; B, habit in fruit × 1/10; C, habit × 1/20; D, part of mature leaf × 2/3; E, seedling leaf × 1/3; F, juvenile leaf × 1/3; G, inflorescence × 2/3; H, stamen, top view × 6; J, stamen, longitudinal section × 6; K, gynoecium, longitudinal section × 6; L, infructescence × 1/2; M, seed × 2; N, inflorescence × 2/3; P, stamen, top view × 6; Q, stamen, longitudinal section × 6; R, gynoe- cium × 6; S, inflorescence × 2/3; T, stamen, top view × 6; U, stamen, longitudinal section × 6; V, gynoecium × 6; W, inflorescence × 2/3; X, stamen, top view × 6; Y, stamen, longitudinal section × 6; Z, gynoecium × 6; AA, stamen, top view × 6; BB, stamen, longitudinal section × 6; CC, gynoecium × 6; DD, stamen, top view × 6; EE, stamen, longitudinal section × 6; FF, gynoecium × 6. Anchomanes welwitschii: A, Bullock 1215 (Kew photograph collection); B, Cult. Kew 14/12/59 (Kew slide collection); A. difformis: C, Croat s.n. (Kew photograph collection); D, Lowe 2116 (K); A. welwitschii: E–F, Fanshawe 8549 (K); A. difformis: G–K, Cult. Kew 1961–38601 (Kew spirit collection 29047.636); L–M, Milne-Redhead 2676 (Kew spirit collection 18904); A. nigritianus: N–R, Cult. Kew 1976.00789 Bogner 640 (Kew slide collection & Kew spirit collection 29047.1); Bogner 662 (Kew spirit collection 37350); A. abbreviatus: S–V, Cult. Kew 1951–556 Greenway s.n. (Kew spirit collection 21975); Cult. Kew 1952.51801 Faulkner 878 (Kew slide collection); A. boehmii: W–Z, Bally 7558 (K & Kew spirit collection 56560); Shabani 11 (K); A. petiolatus: AA–CC, Boughey 163 (Kew spirit collection 16869); A. welwitschii: DD–FF, Cult. Kew (Kew spirit collection 22258). 220 T H E G E N E R A O F A R A C E A E

70. Anchomanes 71. Pseudohydrosme µm.), exine psilate or obscurely verruculate, very thin. innatifid to pinnatisect, ultimate (distal) lobes mostly truncate FEMALE FLOWER: ovary 1-locular, ovule 1, erect, anatropous, to shallowly bifid, sessile to decurrent, proximal lobes acute- funicle very short, placenta basal, stylar region shortly conic acuminate; primary lateral veins of ultimate lobes pinnate, or absent, sometimes strongly deflexed towards spadix base, forming irregular submarginal collective vein or running into stigma either 2-lobed and reniform to V-shaped or discoid margin, higher order venation reticulate. INFLORESCENCE: or depressed-globose. BERRY: large, oblong-ellipsoid, solitary, appearing before leaf. PEDUNCLE: aculeate, very fleshy, borne in cylindric spike, red, purplish or partly short, much shorter than petiole. SPATHE: large, somewhat white. SEED: obovoid to oblong-ovoid, testa very thin, resembling the horn of a euphonium, unconstricted, bright smooth, transparent, embryo large, green, endosperm yellow, dark purple within and at mouth of tube, tube con- absent. See Plates 70, 122B. volute, fleshy, obconic, blade very broad, thinner, with CHROMOSOMES: 2n = 40, very large. flaring, auriculate margins, fornicate. SPADIX: very short, DISTRIBUTION: 7–8 spp.; tropical Africa: Angola, Benin, sessile, female zone subcylindric, male zone cylindric, Burkina Faso, ?Cabinda, Cameroon, Central African Republic, obtuse, subequal to or longer than female, fertile to apex (P. Chad, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, gabunensis) or with appendix covered with sterile flowers Gambia, Ghana, ?Guinea, ?Guinea-Bissau, Ivory Coast, Kenya, (P. buettneri). FLOWERS: unisexual, perigone absent. MALE Liberia, ?Malawi, Mozambique, Nigeria, Senegal, Sierra Leone, FLOWER: 2–5-androus, stamens free, subprismatic, com- Sudan, Tanzania, Togo, Uganda, Zaïre, Zambia. pressed, anthers sessile, connective thick, broad, ECOLOGY: tropical humid forests, savannas (A. welwitschii), overtopping thecae, thecae oblong, long, lateral, dehiscing near swamp (A. boehmii); geophytes, in leaf litter between by apical pore. POLLEN: extruded in strands, inaperturate, rocks or on forest floor, seasonally dormant. ellipsoid-oblong, very large (mean 106 µm., range 93–114 NOTES. The only clear cut distinction between Anchomanes µm.), exine psilate to slightly scabrous. STERILE MALE and Pseudohydrosme is locule number. FLOWERS: composed of subprismatic, free staminodes. ETYMOLOGY: Greek agchein (to strangle) and mainesthai FEMALE FLOWER: ovary globose to broadly ellipsoid, some- (to rage); old plant name by Appuleius. times compressed, 2(–3)-locular, ovules 1 per locule, TAXONOMIC ACCOUNTS: Engler (1911), Knecht (1983), anatropous, funicle short, placenta axile at base of septum, Mayo & Bogner in Mayo (1985a), Ntépé-Nyame (1988). stylar region attenuate to cylindric, narrower than ovary, stigma thick, 2(–3)-lobed, concave centrally. BERRY: unknown. SEED: unknown. See Plates 71, 122C. C 71. Pseudohydrosme CHROMOSOMES: 2n = ca. 40. DISTRIBUTION: 2 spp.; tropical Africa:– Gabon. Pseudohydrosme Engler in Bot. Jahrb. 15: 455 (1892, ECOLOGY: tropical humid forest; geophytes on forest floor. “1893”). LECTOTYPE: P. gabunensis Engler (see N.E. Brown NOTES: Engler (1911) recognized 2 sections:– sect. in Thiselton-Dyer, Fl. Trop. Africa 8: 160. 1901, “1902”). Pseudohydrosme (“Chorianthera”), sect. Zyganthera. Pseudohydrosme buettneri (sect. Zyganthera) has never been SYNONYM: Zyganthera N.E. Brown in Thiselton-Dyer, Fl. recollected and the type no longer exists. N.E. Brown con- Trop. Africa 8: 160 (1901, “1902”). sidered this species to be sufficiently different to warrant HABIT: large, seasonally dormant herbs, tuber subglobose, generic recognition. subterranean, with annular leaf scars, growing continuously ETYMOLOGY: Greek pseudo (false) and Hydrosme, itself and not renewed with each growing period. LEAF: solitary, derived from hydra (water snake) and osmê (smell); in refer- large. PETIOLE: long, aculeate, sheath very short. BLADE: ence to its similarity to Hydrosme (now Amorphophophallus). dracontioid: i.e. trisect, primary divisions pinnatifid or bip- TAXONOMIC ACCOUNTS: Engler (1911), Bogner (1981b). N E P H T H Y T I D E A E : P S E U D O H Y D R O S M E 221

FG E H AJ C B D Plate 71. Pseudohydrosme. A, habit × 1/10; B, portion of leaf × 1/2; C, section of petiole × 2/3; D, habit in flower × 1/2; E, spadix × 2/3; F, stamen × 6; G, stamen, longitudinal section × 6; H, gynoecium × 6; J, gynoecium, longitudinal section × 6. Pseudohydrosme gabunensis: A, Aroideana 4(1): 32, fig.2 (1981); B–D, Bogner 664 (K); Aroideana 4(1): 33, fig.6, 34, fig.8 (1981); E–J, Bogner 664 (Kew spirit collection 45184). 222 T H E G E N E R A O F A R A C E A E

C Tribe Aglaonemateae ated into tube and blade, often apiculate, green to whitish, slightly to strongly decurrent, marcescent. SPADIX: cylindric Tribe Aglaonemateae Engler in Nova Acta Acad. Leopold.- to clavate, shorter or longer than spathe, stipe long to Carol. 39: 148 (1876, “Aglaonemeae”). almost absent, female zone rather few-flowered, contigu- Laticifers simple, articulated; leaf blade (linear-) ovate to ous with and much shorter than male zone, male zone elliptic-oblong, base sometimes subcordate, primary lat- fertile to apex; rarely with staminodes. FLOWERS: unisex- eral veins pinnate, forming single marginal vein, higher ual, perigone absent. MALE FLOWER: stamens free, not order venation parallel-pinnate; spathe boat-shaped, forming clear floral groups, filaments usually distinct, con- unconstricted; spadix with contiguous female and male nective thickened, thecae opposite, obovoid, short, zones; flowers unisexual, perigone absent; stamens free, dehiscing by apical pore or reniform transverse slit. thecae dehiscing by apical pore, pollen grains large, ovary POLLEN: inaperturate, ellipsoid, large (mean 52 µm.), range 1-locular, ovule 1, anatropous, style short, narrower than 37–67 µm.), exine essentially psilate. FEMALE FLOWER: ovary, stigma broad, discoid, concave; berry large; seed ovary subglobose, 1-locular, ovule 1, anatropous, shortly large, ellipsoid, testa smooth, thin, endosperm absent. ovoid, funicle very short, placenta basal, stylar region short, thick, stigma broad, discoid, concave centrally. BERRY: ellipsoid, outer layer fleshy green but turning yellow, rarely C 72. Aglaonema white and finally red. SEED: ellipsoid, almost as large as berry, testa thin, ± smooth, tegmen inconspicuous, embryo Aglaonema Schott in Wiener Z. Kunst 1829 (3): 892 (1829). large, endosperm absent. See Plates 72, 122D. TYPE: A. oblongifolium Schott, nom. illeg. (Arum integri- CHROMOSOMES: 2n = 40, 60, 80, 100, 120 (70, 110). folium Link, Aglaonema integrifolium (Link) Schott). DISTRIBUTION: 21 spp.; tropical Asia, Malay Archipelago, Papuasia:– Bangladesh, Brunei, Burma, Cambodia, China HABIT: evergreen herbs, sometimes robust, stem epigeal, (Guandong, Guangxi, Hainan, Yunnan), India (NE), erect and unbranched or creeping and often branched, Indonesia (Borneo, Java, Moluccas, Sulawesi, Sumatra), Laos, internodes green, smooth, often rooting at the nodes. Malaysia (Borneo, Peninsula), Papua New Guinea, LEAVES: several, forming an apical crown. PETIOLE: sheath Philippines, Thailand, Vietnam. usually long. BLADE: ovate-elliptic, narrowly elliptic, rarely ECOLOGY: tropical humid forest; terrestrial on forest floor, broadly ovate or sublinear, base often unequal, attenuate occasionally in deciduous forest or regrowth, also in humus to rounded, rarely cordate, often with striking, silvery and deposits on limestone and in peat deposits. pale green patterns of leaf variegation; primary lateral veins COMMON NAMES AND USES: chinese evergreen, very pinnate, often weakly differentiated, running into marginal widely cultivated as an ornamental plant. vein, higher order venation parallel-pinnate. INFLORES- ETYMOLOGY: Greek aglaos (splendid) and nema, nêmatos CENCE: 1–9 in each floral sympodium. PEDUNCLE: shorter (stamen). or longer than petiole, deflexing in fruit. SPATHE: ovate to TAXONOMIC ACCOUNTS: Engler (1915), Nicolson (1969), ± globose, erect, boat-shaped to convolute, not differenti- Jervis (1980). 72. Aglaonema A G L A O N E M A T E A E : A G L A O N E M A 223

E C D F B JH G A K L Plate 72. Aglaonema. A, habit × 1/4; B, leaf × 1/3; C, detail of leaf venation × 4; D, spadix × 1; E, stamen × 10; F, inflorescence × 1; G, sta- minode × 10; H, gynoecium × 10; J, gynoecium, longitudinal section × 10; K, habit × 1/4; L, inflorescence × 1. Aglaonema simplex : A–C, Cult. Kew 1963–43405; D–E, Giles & Woolliams PB109 (Kew spirit collection 37985); A. simplex : F–J, Hay 2001 (Kew spirit collection 59067); A. modestum: K, Cult. Kew 1968–38227; A. hookerianum: L, Cult. Kew (Kew spirit collection 22425). 224 T H E G E N E R A O F A R A C E A E

C 73. Aglaodorum Tribe Culcasieae C Aglaodorum Schott, Gen. Aroid. t. 58 (1858). TYPE: A. grif- Tribe Culcasieae Engler in Engler & Prantl, Nat. Pflan- C fithii (Schott) Schott (Aglaonema griffithii Schott). zenfam. II (3): 112, 116 (1887). HABIT: evergreen herb, stem rhizomatous, creeping, Laticifers present, simple, articulated (Cercestis) or absent aerenchymatous stem, much-branched, . LEAVES: several to (Culcasia), resin canals present in roots, stems and leaves, many. PETIOLE: terete, sheath short. BLADE: oblanceolate sclerotic hypodermis present in roots; usually climbing to oblong, thick; midrib very thick, primary lateral veins hemiepiphytes with slender stems (rarely robust), sometimes pinnate, weakly differentiated, higher order venation paral- erect or prostrate terrestrial plants; petiole geniculate api- lel-pinnate. INFLORESCENCE: solitary. PEDUNCLE: long, cally; primary lateral veins pinnate, higher order venation subequal to petiole, erect in fruit. SPATHE: oblong, cuspi- reticulate; female zone of spadix contiguous with male or date, convolute, erect. SPADIX: stipitate, a little shorter than separated by zone of sterile male flowers; flowers unisexual, spathe, female zone very short with a single whorl of flow- perigone absent; male flower 2–4 androus, stamens free, ers, male zone subcylindric, fertile to apex. FLOWERS: anthers subsessile, lacking endothecial thickenings, connec- unisexual, perigone absent. MALE FLOWER: stamens free, tive usually strongly thickened, broad, truncate, thecae short, prismatic, filaments distinct, connective thickened, dehiscing by short apical slit or pore; ovules 1 per locule, thecae oblong, adjacent or opposite, dehiscing by apical stigma sessile, large, discoid; berries ± globose, usually red; pore. POLLEN: inaperturate, ellipsoid, large (mean 59 µm.), seed testa thin, endosperm absent. exine apparently psilate or verruculate. FEMALE FLOWER: gynoecium surrounded by whorl of 1–3 shorter prismatic sta- 74. Culcasia minodes, ovary 1–2-locular, ovule 1 per locule, anatropous, funicle very short, placenta parietal in 1-locular ovaries, Culcasia Palisot de Beauvois, Fl. Oware 1: 3 (1805), nom. stigma broad, discoid, 4-lobed, concave at centre. BERRY: cons. TYPE: C. scandens Palisot de Beauvois (typ. cons.). obovoid to ellipsoid, large, pericarp very thick, green. SEED: large, ellipsoid, testa thin, smooth, embryo large, plumule SYNONYM: Denhamia Schott in Schott & Endlicher, well-developed, endosperm absent. See Plates 73, 123A. Melet. Bot. 19 (1832). CHROMOSOMES: 2n = 40. DISTRIBUTION: 1 sp.; Indonesia (Sumatra), Malaysia Laticifers absent. HABIT: erect, repent or climbing herbs, root- (Borneo, Peninsula), Vietnam. ing at least from lower nodes, branches slender. LEAVES: many, ECOLOGY: tropical open swamps, especially freshwater tidal often forming terminal crown in terrestrial species. PETIOLE: zones and brackish water, often growing with Nypa fruticans geniculate apically, geniculum often inconspicuous, sheath and Cryptocoryne ciliata; creeping rhizomatous helophyte. persistent, rather long. BLADE: lanceolate or oblanceolate to ETYMOLOGY: Greek aglaos (splendid) and dôron (gift), a ovate or ovate-oblong, rarely rounded, rarely pubescent below, poetic name. resin canals pellucid, linear or punctate; primary lateral veins TAXONOMIC ACCOUNTS: Engler (1915), Nicolson (1969). pinnate, often forming submarginal collective vein, otherwise running into marginal vein, higher order venation reticulate. 73. Aglaodorum C U L C A S I E A E : C U L C A S I A 225

EF B D A GC Plate 73. Aglaodorum. A, habit × 2/3; B, inflorescence × 1; C, spadix × 2; D, stamen × 8; E, gynoecium with associated staminode × 8; F, gynoecium, longitudianal section × 8; G, fruit × 1. Aglaodorum griffithii: A, C–G, Bogner 1767 (M); Bogner 1672 (Kew spirit collection 49830); B, Boyce (Kew slide collection). 226 T H E G E N E R A O F A R A C E A E

A B K T C FG LM HJ NP RS Q DE Plate 74. Culcasia. A, habit × 1/3; B, juvenile habit × 1/3; C, habit × 1/3; D, leaf × 1/3; E, habit × 1/3; F, inflorescence × 1; G, inflorescence × 1; H, stamens × 10; J, gynoecium, longitudinal section × 10; K, spadix × 5; L, detail of stamens, top view × 4; M, detail of gynoecia × 4; N, stamens × 10; P, gynoecium, longitudinal section × 10; Q, habit × 2/3; R, stamens × 10; S, gynoecium, longitudinal section × 10; T, seed × 1 1/2. Culcasia parviflora: A, Keay IFH 37529 (K); B, Bogner 697 (K); C. angolensis: C, Johnson s.n. (K); Robyns 1140 (K); C. panduriformis: D, Zenker 4572 (K); C. longevaginata: E, Baldwin 9226 (K); C. striolata: F, Milne-Redhead 5155 (Kew spirit collection 22442); C. orientalis: G–J, Bogner 140 (Kew spirit collection 22712); C. seretii: K–M, Cult. Kew 1982–04604 (Kew spirit collection 46485); C. saxatilis: N–P, Meikle 1275 (Kew spirit collection 25627); C. rotundifolia: Q–S, Bogner 749 (K & Kew spirit collection 56676); C. liberica: T, Cult. Kew 1982–04601 (Kew spirit collection 51491). C U L C A S I E A E : C U L C A S I A 227

INFLORESCENCE: 1–12(–20) in each floral sympodium, intern- 75. Cercestis C odes of floral sympodium sometimes relatively elongated. PEDUNCLE: short to relatively long. SPATHE: erect, green to Cercestis Schott in Oesterr. bot. Wochenbl. 7: 414 (1857). whitish, boat-shaped, not or hardly constricted, convolute TYPE: C. afzelii Schott basally, gaping apically at anthesis, deciduous to marcescent. SPADIX: subsessile to stipitate, cylindric-clavate, equal to or SYNONYMS: Alocasiophyllum Engler in Bot. Jahrb. 15: somewhat longer than spathe, female zone usually densely 449 (1892, “1893”); Rhektophyllum N.E. Brown in J. Bot. 20: flowered, rarely laxly, shorter than male zone and either con- 194 (1882). tiguous with it or separated by a laxly flowered zone of sterile male flowers, male zone fertile to apex, axis often persistent in Laticifers present, simple, articulated. HABIT: climbing herbs, fruit. FLOWERS: unisexual, perigone absent. MALE FLOWER: sometimes creeping, small to robust, stem long, producing fla- 2–4-androus, stamens free, short, obpyramidal, truncate api- gelliform shoots with long internodes and bearing only cally, anthers subsessile, connective thick, thecae oblong, cataphylls, followed by thicker, flowering stems with several dehiscing by short apical slit. POLLEN: inaperturate, sub- foliage leaves and short internodes. LEAVES: many. PETIOLE: spheroidal, medium-sized (mean 32 µm., range 27–40 µm.), geniculate apically, geniculum sometimes inconspicuous, exine verrucate to rugulate or subreticulate, usually with psilate sheath short to long. BLADE: oblong-lanceolate or oblong to patches of variable size, rarely spinose. STERILE MALE FLOW- cordate, sagittate, or hastate, or trifid with acuminate seg- ERS: with 3–4 obpyramidal, depressed staminodia. FEMALE ments, sometimes laciniate-pinnatifid with slit-like perforations FLOWER: ovary depressed, 1–3-locular, ovules 1 per locule, extending to margin forming large rhomboid to obtriangular anatropous, funicle short, placentae subbasal, stigma sessile, lobes (C. camerunensis, C. mirabilis), resin canals pellucid, hemispheric-discoid, relatively large, sometimes weakly lobed. linear or punctate; primary lateral veins pinnate, running into BERRY: globose to ellipsoid, 1–3-seeded, mostly red, sometimes marginal vein, higher order venation reticulate. INFLORES- orange to greenish-yellow, infructescence subglobose to cylin- CENCE: usually 1–4 in each floral sympodium (up to 16 in C. dric. SEED: ovoid to ellipsoid, testa thin, smooth, brown, camerunensis). PEDUNCLE: shorter than spathe. SPATHE: plumule lateral, superficial, with leaf primordia (C.liberica), erect, thick, boat-shaped to subcylindric, convolute basally endosperm absent. See Plates 74, 123B. into a tube, gaping apically at anthesis, not or hardly con- CHROMOSOMES: 2n = 42, 84. stricted, persistent to marcescent. SPADIX: sessile, shorter DISTRIBUTION: ca. 27 spp.; tropical Africa:– Angola, Benin, than spathe, female zone shorter and contiguous with male. Burkina Faso, ?Burundi, Cabinda, Cameroon, Central African FLOWERS: unisexual, perigone absent. MALE FLOWER: 2–4- Republic, Congo, Equatorial Guinea (Bioko, Rio Muni), androus, stamens free, prismatic, slightly narrowed basally, Gabon, ?Gambia, Ghana, Guinea, ?Guinea-Bissau, Ivory usually short and relatively broad, sometimes rather long and Coast, Kenya, Liberia, Mali, Niger, Nigeria, ?Rwanda, Senegal, narrow (C. camerunensis, C. mirabilis), filaments very short, Sierra Leone, Tanzania, Togo, Uganda, Zaïre. connective broad, thecae shortly oblong to almost linear, ECOLOGY: tropical moist and humid forest; usually climbing dehiscing by small apical slit. POLLEN: extruded in strands, hemiepiphytes, sometimes terrestrial, terrestrial species tend inaperturate, ellipsoid to oblong or spherical to subspher- to occur in damp forest sites. oidal, medium-sized (mean ca. 40 µm., range 36–55 µm.), ETYMOLOGY: variant of Middle Eastern name qolqas, (also exine obscurely fossulate-verrucate, or almost perfectly psilate the origin of the name Colocasia). (C. camerunensis, C. mirabilis). FEMALE FLOWER: ovary TAXONOMIC ACCOUNTS: Engler (1905), Bogner (1980b), obovoid, 1-locular, ovule 1, hemianatropous to anatropous, Knecht (1983), Mayo (1985a), Ntépé-Nyame (1988), Boyce rarely campylotropous (C. taiensis), funicle very short, pla- (1995b). centa parietal to subbasal, stigma sessile to subsessile, often rather broad, discoid, often somewhat concave centrally. 74. Culcasia 75. Cercestis 228 T H E G E N E R A O F A R A C E A E

JL KMN B QP A C H F GD E Plate 75. Cercestis. A, flowering shoot arising from leafless continuation shoot, attached to twig by aerial roots × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, inflorescences with associated stem and petiole base × 1/2; E, male flower × 6; F, gynoecium × 6; G, gynoecium, longitudinal sec- tion × 6; H, spadix, J, detail of male zone of spadix × 6; K, detail of female zone of spadix × 6; L, male flower × 9; M, gynoecium × 9; N, gynoecium, longitudinal section × 9; P, habit × 1/2; Q, fruiting habit × 1/2. Cercestis stigmaticus: A, Coombe 169 (K); C. kamerunianus: B, Jones 19526 (K); C. mirabilis: C, Bogner 601 (K); D, Bogner 705 (K); E–G, Bogner s.n. (K & Kew spirit collection 29047.471); C. dinklagei: H–N, Hepper 7508 (Kew spirit collection 49930); C. afzelii: P, Jones & Onochie FHI 18753 (K); Q, Adams 3729 (K). C U L C A S I E A E : C E R C E S T I S 229

BERRY: infructescence ellipsoid to oblong, berries obovoid to subglobose or ellipsoid, red, pericarp thick. SEED: obovoid to ellipsoid, testa smooth, thin, embryo large, outer cell layer green, endosperm absent. See Plates 75, 123C. CHROMOSOMES: 2n = 42, morphologically similar to those of Culcasia. DISTRIBUTION: 13 spp.; tropical Africa:– Angola, Benin, ?Burundi, ?Cabinda, Cameroon, Central African Republic, Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Nigeria, ?Rwanda, Senegal, Sierra Leone, ?Togo, Uganda, Zaïre. ECOLOGY: tropical humid forest; climbing hemiepiphytes and terrestrial plants. ETYMOLOGY: Greek Cercestes, one of the fifty sons of mythic Aegyptus. TAXONOMIC ACCOUNTS: Engler (1911), Knecht (1983), Mayo (1985a), Bogner (1986a), Ntépé-Nyame (1988), Bogner & Knecht (1995). C Tribe Montrichardieae Tribe Montrichardieae Engler in Nova Acta Acad. Leopold.- Carol. 39: 144 (1876). 76. Montrichardia Laticifers simple, articulated, sclerotic hypodermis present in roots; arborescent helophytes; leaf blade cordate-sagittate to tri- FLOWER: 3–6-androus, stamens free, obpyramidal-prismatic, fid, primary lateral veins pinnate, higher order venation ± truncate at apex, anthers sessile, connective thick, overtop- reticulate; spathe deciduous after anthesis, slightly constricted, ping thecae, thecae oblong-ellipsoid, dehiscing by longitudinal differentiated into lower tube and upper, ± boat-shaped blade; slit. POLLEN: inaperturate, spherical to subspheroidal, large spadix: female and male zone contiguous, sterile flowers (mean 94 µm., range 92–96 µm.), exine psilate. FEMALE absent; flowers unisexual, perigone absent; male flower 3–6- FLOWER: gynoecia prismatic-cylindric, ovary 1-locular, ovules androus, stamens free, ± sessile, connective overtopping thecae, 1–2, anatropous, funicle short, placenta subbasal to basal, thecae dehiscing by short, apical slit, pollen large; ovary 1-loc- stylar region prismatic, thick, excavated and rugulose at apex, ular, ovules 1–2, anatropous, placenta subbasal, style prismatic, stigma small, on low central boss, irregular to elliptic or truncate, longer than ovary, ± excavated apically; berry and roundish. BERRIES: free, large, subcylindric, somewhat com- seed large, plumule well-developed, endosperm absent. pressed, pericarp spongiose, at apex excavated and radiately furrowed, 1-seeded. SEED: large, obovoid to ellipsoid, testa C 76. Montrichardia smooth or rough, brown, embryo large, endosperm absent. See Plate 76, 123D. Montrichardia H. Crüger in Bot. Zeitung (Berlin) 12: 25 CHROMOSOMES: 2n = 48. (1854), nom. cons. TYPE: M. aculeata (G.F.W. Meyer) Schott DISTRIBUTION: 2 spp.; tropical America, West Indies:– (Caladium aculeatum G.F.W. Meyer). Belize, Brazil (Amazonia, Northeast, South East), Colombia, Costa Rica, French Guiana, Guatemala, Guyana, Honduras, SYNONYM: Pleurospa Rafinesque, Fl. Tell. 4: 8 (1838, Lesser Antilles, Nicaragua, Panama, Peru, Puerto Rico, “1836”). Surinam, Trinidad, Tobago, Venezuela. ECOLOGY: tropical humid forest; helophytes in tranquil HABIT: robust to sometimes gigantic arborescent evergreen freshwater habitats, forming dense, often extensive stands herbs, stem erect, multiplying at base from hypogeal rhi- along river margins. zomes, internodes well-developed, smooth or aculeate, ± ETYMOLOGY: named after G. de Montrichard. slender to massive. LEAVES: several, borne in terminal crown. TAXONOMIC ACCOUNTS: Engler (1911), Jonker-Verhoef & PETIOLE: sheath half as long as petiole or longer, with free Jonker (1953), Lins (1994). apical ligule. BLADE: cordate-sagittate to sagittate, hastate to trifid, rarely trisect, posterior divisions often longer than ante- rior division; basal ribs well-developed, primary lateral veins pinnate, running into marginal vein, secondary laterals ± par- Tribe Zantedeschieae C allel-pinnate, higher order venation reticulate. INFLORESCENCE: 1(–2) in each floral sympodium. PEDUN- Tribe Zantedeschieae Engler in Engler & Prantl, Nat. CLE: shorter than petiole. SPATHE: erect, thick, entirely Pflanzenfam. II (3): 113, 136 (1887). deciduous after anthesis, ± constricted between tube and blade, tube convolute, blade longer than tube, widely gap- Laticifers simple, articulated; stem a depressed-globose tuber ing at anthesis, ± boat-shaped. SPADIX: sessile, erect, or short rhizome (Z. aethiopica); leaf blade lanceolate to subequal to spathe, female zone cylindric, male zone sub- hastate-sagittate, primary lateral veins pinnate, forming single conoid, contiguous with and much longer than female zone, marginal vein, higher order venation parallel-pinnate; pedun- fertile to apex. FLOWERS: unisexual, perigone absent. MALE cle long, sometimes longer than leaves; spathe ± obconic, 230 T H E G E N E R A O F A R A C E A E

F Q R D P NE G A H J C B K ML Plate 76. Montrichardia. A, habit × 1/10; B, flowering shoot × 1/2; C, base of stem showing rhizome × 1/2; D, mid–portion of stem × 1/2; E, portion of stem showing prickles × 1/2; F, spadix × 1/2; G, stamens, top view × 3; H, stamen, side view × 5; J, gynoecium, top view show- ing irregular surface with stigma × 5; K, gynoecium, side view × 5; L, gynoecium, top view showing irregular surface with stigma × 3; M, gynoecium, longitudinal section × 5; N, infructescence × 1/2; P, seed × 2; Q, leaf × 1/2; R, leaf × 1/2. Montrichardia linifera: A, Mayo s.n. (Kew slide collection); M. arborescens: B, Philcox 8032 (K); Philcox et al. 8413 (Kew spirit collection 45821); C–E, Bogner 2188 (Kew spirit collection 29047.760 & Kew illustration collection); Nelson 1729 (K); F, Simmons s.n. (K & Kew spirit collection 58906); G-M, Bogner 2188 (Kew spirit collection 29047.760); M. linifera: N–P, Stannard & Arrais (Kew spirit collection 51643 & 51644); Q, Harley et al. 24747 (K); M. arborescens R, Nelson 1733 (K). M O N T R I C H A R D I E A E : M O N T R I C H A R D I A 231

unconstricted, tube subcylindric-obconic, blade gaping shorter than spathe, sessile or stipitate, male and female widely; spadix shorter than spathe, male and female zones zones contiguous, fertile to apex. FLOWERS: unisexual, contiguous; flowers unisexual, perigone absent; stamens free, perigone absent. MALE FLOWER: 2–3-androus, stamens connective thickened apically, thecae dehiscing by apical free, anthers oblong, somewhat compressed, subsessile, pore, pollen extruded in strands; ovary 1–5 locular, ovules connective truncate at apex, separating thecae, thecae (1–)4(–8) per locule, anatropous, placenta axile to subapical; opposite, oblong, dehiscing by apical pore. POLLEN: berry green, orange or yellow; seed testa striate, endosperm extruded in strands, inaperturate, ellipsoid to oblong, copious. medium-sized (mean 40 µm., range 31–49 µm.), exine per- fectly psilate to obscurely dimpled. FEMALE FLOWER: either C 77. Zantedeschia a naked gynoecium or rarely (Z. aethiopica, Z. odorata) sur- rounded by whorl of ca. 3 spathulate to clavate and apically Zantedeschia K. Sprengel, Syst. Veg. 3: 756, 765 (1826), truncate staminodes, ovary ovoid, 1–5-locular, ovules nom. cons. TYPE: Z. aethiopica (L.) K. Sprengel (Calla (1–)4(–8) per locule, anatropous, placenta axile to subapical, aethiopica L.), typ. cons. stylar region short- to long-attenuate, stigma rather small, discoid-subcapitate. BERRY: obovoid or depressed-globose, SYNONYMS: ? Aroides Heister ex Fabricius, Enum., ed. 2, 1–several-seeded, usually green, or orange, rarely yellow, 42 (1763); Houttinia Necker, Elementa Botanica 3: 291 (1790); and mucilaginous within (Z. aethiopica). SEED: ovoid to Colocasia Link, Diss. Bot. 77 (1795), nom. rej.; Richardia ellipsoid, strophiolate, testa costate, embryo axile, elon- Kunth in Mém. Mus. Hist. Nat. 4: 433, 437 (1818, non L. gate, endosperm copious. See Plates 77, 124A. 1753); Otosma Rafinesque, Fl. Tell. 4: 8 (1838, “1836”); CHROMOSOMES: 2n = 32. [Arodes O.Kuntze, Rev. Gen. 2: 739 (1891), orth. var.]; DISTRIBUTION: 8 spp.; southern Africa:– Angola, Botswana, Pseudohomalomena A.D. Hawkes in Madroño 11: 147 (1951). Lesotho, Malawi, ?Mozambique, Namibia, South Africa, HABIT: seasonally dormant, sometimes evergreen (Z. aethi- Swaziland, Tanzania, ?Zaire, Zambia, Zimbabwe. opica) herbs, tuber depressed-globose, or with thick ECOLOGY: tropical and subtropical damp open habitats; rhizome (Z. aethiopica). LEAVES: several, radical. PETIOLE: helophytes, seasonally wet places, along streams, swampy spongiose, sheath long. BLADE: lanceolate, narrowly ellip- ground along forest edges, dry rocky hills. tic, cordate, cordate-sagittate, sagittate, hastate, often NOTES: Zantedeschia aethiopica is morphologically distinct variegated by aggregations of pale diaphanous necrotic from the other species; it is naturalized in various parts of patches; primary lateral veins pinnate, running into distinct tropical America, in southern Europe, the Philippines, New marginal vein, secondary and tertiary laterals parallel-pin- Zealand and elsewhere. nate, higher order venation transverse-reticulate. ETYMOLOGY: named after G. Zantedeschi (1773–1846). INFLORESCENCE: solitary in each floral sympodium, TAXONOMIC ACCOUNTS: Engler (1915), Letty (1973), Perry appearing with leaves. PEDUNCLE: long, subequal or longer (1989), Singh, Wyk & Baijnath (1996). than petiole. SPATHE: unconstricted, persistent, pure white, cream, yellow, pink or rosy purple, lower part convolute C forming a short, stout, subcylindric to obconic tube, often Tribe Callopsideae dark purple within at base, upper part widely gaping, Tribe Callopsideae Engler in Engler & Prantl, Nat. shorter than or subequal to tube, suberect or recurved with Pflanzenfam., Nachtr. 3: 29, 34 (1906). somewhat revolute margins. SPADIX: erect, digitiform, Laticifers simple, articulated; small, evergreen herbs, rhizome slender, hypogeal; leaf blade cordate-ovate, primary lateral veins forming single marginal vein, higher order venation reticulate; spathe completely expanded at anthesis, persistent; spadix fertile to apex; female zone entirely adnate to spathe, contiguous with male zone or with short naked zone in between; flowers unisexual, perigone absent; stamens free, anthers sessile, thecae opposite, dehiscing by apical pore; gynoecia few, slender, ovary 1-locular, ovule 1, anatropous, placenta basal, style long-attenuate, stigma small; embryo elongate, endosperm copious. 78. Callopsis C 77. Zantedeschia Callopsis Engler in Notizbl. Königl. Bot. Gart. Berlin 1: 27 (1895). TYPE: C. volkensii Engler Laticifers simple, articulated. HABIT: small evergreen herb, stem a slender hypogeal rhizome, internodes very short. LEAVES: usually minutely hispid on petiole and abaxial sur- face of blade. PETIOLE: sheath very short. BLADE: cordate-ovate; primary lateral veins pinnate, running into marginal vein, higher order venation reticulate. INFLORES- CENCE: 2 in each floral sympodium, appearing with the 232 T H E G E N E R A O F A R A C E A E

J C B FE K L G HA D Plate 77. Zantedeschia. A, habit × 1/3; B, leaf × 2/3; C, inflorescence × 2/3; D, spadix × 2; E, stamens, top view × 9; F, stamen × 9; G, gynoe- cium × 9, H, gynoecium, longitudinal section × 9; J, infructescence × 2/3; K, leaf × 2/3; L, gynoecium with associated staminodes × 9. Zantedeschia albomaculata subsp. albomaculata: A, Scheepers 36 (K); B, Codd 10688 (K); C–H, Milne-Redhead 3812 (Kew spirit collection 199162); J, Prosser 1827 (K); Z. rehmannii: K, Cult. Kew 1958–11001 (K); Z. aethiopica: L, Cult. Kew (Kew spirit collection 7467). Z A N T E D E S C H I E A E : Z A N T E D E S C H I A 233

EF D A CB Plate 78. Callopsis. A, habit with spathe of infructescence flattened out by pressing × 1; B, inflorescence, nearside and upper part of spathe removed × 5; C, gynoecium, longitudinal section × 20; D, stamen × 20; E, berry, side view with hilum at top × 4; F, seed, side view × 4. Callopsis volkensii: A, Faulkner 1709 (K); Greenway 4631 (K); Robertson & Luke 6468 (K); B–D, Bogner 218 (Kew spirit collection 56422); E–F, Robertson & Luke 6468 (K). 234 T H E G E N E R A O F A R A C E A E

leaves. PEDUNCLE: slender, equalling or longer than peti- Tribe Thomsonieae C ole, sometimes exceeding entire leaf. SPATHE: broadly C ovate-elliptic to subcircular, acuminate, decurrent, white, ± Tribe Thomsonieae Blume, Rumphia 1: 138 (1837); Bogner fully expanded and sometimes arching backwards at anthe- et al. in Aroideana 8 (1): 14–25 (1985). sis, persistent, closing at fruiting stage. SPADIX: shorter than spathe, female zone entirely adnate to spathe, ± laxly flow- Laticifers simple, articulated; seasonally dormant, stem tuber- ered with gynoecia ± biseriately arranged, contiguous with ous, occasionally rhizomatous, hypogeal; leaf usually 1; male zone or with short, naked axis in between, male zone petiole long, terete, variously maculate; blade dracontioid, equalling female, cylindric, densely flowered, fertile to apex. first foliage leaf of seedling divided, never entire, the three FLOWERS: unisexual, perigone absent. MALE FLOWER: primary leaf divisions upright in bud, ultimate lobes usually probably 1-androus (2–3-androus according to Engler elliptic-lanceolate, acuminate, primary lateral veins of ulti- 1920a), stamens free, depressed subquadrate, anthers ses- mate lobes pinnate, forming submarginal collective vein, sile, thecae opposite with subovoid microsporangia higher order venation reticulate; inflorescence 1, subtended dehiscing by apical pore. POLLEN: inaperturate, spherical by cataphylls; spadix with sterile terminal appendix, female to subspheroidal, medium-sized (mean 37 µm.), exine papil- zone usually contiguous with male; flowers unisexual, late-spinose. FEMALE FLOWER: gynoecium slender, perigone absent; pollen extruded in strands, exine various; flask-shaped, falcate, yellowish, ovary 1-locular, ovule 1, ovules 1 per locule, anatropous; seed testa smooth, embryo anatropous, placenta basal, stylar region rather long-attenu- large, endosperm absent. ate, stigma small, discoid-subhemispheric, slightly wider than style. BERRY: ovoid-ellipsoid and slightly angled, style 79. Amorphophallus forming persistent mucro, green. SEED: ovoid-ellipsoid, testa ± smooth, thin, embryo elongate, straight to slightly Amorphophallus Blume ex Decaisne in Nouv. Ann. Mus. curved, endosperm copious. See Plates 78, 124B. Hist. Nat. 3: 366 (1834), nom. cons. TYPE: A. campanulatus CHROMOSOMES: 2n = 36. Decaisne (= A. paeoniifolius (Dennstedt) Nicolson). DISTRIBUTION: 1 sp.; Kenya, Tanzania. ECOLOGY: tropical humid forest; creeping, rhizomatous geo- SYNONYMS: Thomsonia Wallich, Pl. Asiat. Rar. 1: 83 phytes, in leaf litter. (1830), Bogner in Pl. Syst. Evol. 125 (1): 15–20 (1976), nom. NOTES: Callopsis resembles Ulearum in various respects but rej.; Pythion Martius in Flora 14: 458 (1831), nom. rej.; lacks anastomosing laticifers and has a narrower connective. Candarum Reichenb. ex Schott & Endlicher, Melet. Bot. 17 In Callopsis the thecae open by a pore which later becomes (1832), nom. illeg.; Pythonium Schott in Schott & Endlicher, a very broad slit. The two microsporangia of each theca thus Melet. Bot. 17 (1832), nom. illeg.; Kunda Rafinesque, Fl. Tell. have a single common opening. The stamens appear to be 2: 82 (1837, “1836”), nom. illeg.; Brachyspatha Schott, Syn. free and uniformly separated to a depth which reaches to the Aroid. 35 (1856); Conophallus Schott, Syn. Aroid. 34 (1856); base of the microsporangia. This makes it impossible to say Plesmonium Schott, Syn. Aroid. 34 (1856), Bogner in from mature material how many stamens form a male flower. Adansonia 20 (3): 305–308 (1980); Corynophallus Schott in Developmental studies will probably show that the male Oesterr. bot. Wochenbl. 7: 389 (1857); Allopythion Schott, flower is unistaminate. Gen. Aroid. t. 24 (1858); Hansalia Schott in Oesterr. bot. ETYMOLOGY: Calla and the Greek opsis (appearance). Zeitschr. 8: 82 (1858); Hydrosme Schott, Gen. Aroid. t. 33 TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1980c), (1858); Rhaphiophallus Schott, Gen. Aroid. t. 27 (1858); Mayo (1985a). Synantherias Schott, Gen. Aroid. t. 28 (1858); Proteinophallus J.D. Hooker in Bot. Mag. 101: t. 6195 (1875). 78. Callopsis HABIT: seasonally dormant (sometimes irregularly so) or rarely semi-evergreen herbs, often large, sometimes gigan- tic, tuber usually depressed-globose, sometimes irregularly ± elongate-cylindric, napiform or carrot-shaped, rarely rhi- zomatous or stoloniferous. LEAVES: usually solitary (rarely 2–3) in adult plants, sometimes 2–3 in seedlings. PETIOLE: long, usually smooth, rarely verrucose to asperate, some- times very thick, usually conspicuously spotted and marked in a variety of patterns, sheath very short. BLADE: dracon- tioide: i.e. trisect, primary divisions pinnatisect, bipinnatisect or dichotomously further divided, tubercles rarely present at junction of divisions, secondary and tertiary divisions ± regularly pinnatifid to pinnatisect, ultimate lobes oblong- elliptic to linear, acuminate, decurrent, rarely petiolulate; primary lateral veins of ultimate lobes pinnate, forming dis- tinct submarginal collective vein, higher order venation reticulate. INFLORESCENCE: always solitary, preceded by cataphylls, usually flowering without leaves, rarely with the leaves. PEDUNCLE: very short to long, similar to petiole. SPATHE: variously coloured, marcescent and finally decid- uous, boat-shaped and not or hardly convolute, or clearly differentiated into tube and blade, sometimes constricted between them; tube convolute, rarely connate (A. pusillus, T H O M S O N I E A E : A M O R P H O P H A L L U S 235

A DC G F HJ E B Plate 79 (i). Amorphophallus. A, habit × 1/20; B, habit × 1/18; C, habit × 1/0; D, part of leaf × 1/6; E, detail of leaf × 2/3; F, part of leaf × 1/6; G, part of leaf × 1/6; H, detail of petiole × 2/3; J, detail of petiole × 2/3. Amorphophallus decus-silvae: A, Java (Kew slide collection); A. maculatus: B, Bogner 600 (Kew illustration collection); A. sp.: C, Richards 741–64 (Kew slide collection); A. galbra: D–E, Brown 5801 (K); A. sp.: F, Cult. Kew. (K); A. dracontioides: G, Johnson 662 (K); A. paeoniifolius : H, Cult. Kew. (Kew slide collection); A. krausei : J, Cult. Kew (Kew slide collection). 236 T H E G E N E R A O F A R A C E A E

C B D A J N EF G H KL M Plate 79 (ii). Amorphophallus. A, inflorescence × 1/3; B, inflorescence × 1/3; C, spadix × 2/3; D, inflorescence × 1/3; E, inflorescence × 2/3; F, detail of fertile part of spadix × 3; G, inflorescence × 1/8; H, inflorescence × 1/6; J, detail of spadix appendix × 2/3; K, inflorescence × 2/3; L, inflorescence × 2/3; M, detail of fertile part of spadix and base of appendix × 3; N, inflorescence × 1/6; Amorphophallus pendulus: A, Cult. Kew 1968–200 (Kew spirit collection 7237); A. krausei: B, H.AM. 40 (Kew slide collection); A. dracontioides: C, I.F.H.19156 (Kew spirit collection 21971); A. lewallei: D, H.AM. 20B (Kew slide collection); A. sumawongii: E–F, Bogner 372 (Kew spirit collection 26332.44); A. paeoniifolius: G, Bogner s.n. (Kew spirit collection 34673); H.AM. 41A (Kew slide collection ); A. hirtus: H, H.AM. 132A (Kew slide col- lection); J, (Kew spirit collection 58005); A. pusillus: K, H.AM. 246 (Kew slide collection); A. pygmaeus: L–M, Bogner 1948 Cult. Hetterscheid (Kew spirit collection 59079); A. gomboczianus: N, H.AM. 3 (Kew slide collection). T H O M S O N I E A E : A M O R P H O P H A L L U S 237

AB C G DE F H J K S L MN P QR T UV W XY Z AA BB Plate 79 (iii). Amorphophallus. A, stamen × 10; B, gynoecium × 10; C, gynoecium, longitudinal section × 10; D, stamen × 10; E, gynoe- cium × 10; F, gynoecium, longitudinal section × 10; G, stamen × 10; H, gynoecium × 10; J, gynoecium, longitudinal section × 10; K, inflorescence, nearside half of spathe removed × 4; L, stamen × 10; M, gynoecium × 10; N, gynoecium, longitudinal section × 10; P, stamen × 10; Q, gynoecium × 10; R, gynoecium, longitudinal section × 10; S, gynoecium, transverse section × 10; T, stamen × 10; U, gynoecium × 10; V, gynoecium, longitudinal section × 10; W, stamen × 10; X, gynoecium × 10; Y, gynoecium, longitudinal section × 10; Z, stamen × 10; AA, gynoecium × 10; BB, gynoecium, longitudinal section × 10. Amorphophallus albispathus: A–C, H.AM 16 (Kew slide collection); A. pyg- maeus: D–F, Bogner 1948 Cult. Hetterscheid (Kew spirit collection 59079); A. pendulus: G–J, Cult. Kew 1968–200 (Kew spirit collection 7237); A. pusillus: K, H.AM. 246 (Kew slide collection); A. gomboczianus: L–N, Ash 1434 (Kew spirit collection 15077.14); A. hirtus: P–S, (Kew spirit collection 58005); A. sumawongii: T–V, Bogner 372 (Kew spirit collection 26332.44); A. paeoniifolius: W–Y, Bogner s.n. (Kew spirit collection 34673); A. dracontioides: Z–BB, I.F.H.19156 (Kew spirit collection 21971). 238 T H E G E N E R A O F A R A C E A E

C DE AB FG Plate 79 (iv). Amorphophallus. A, tuber × 1/6; B, tuber × 1/6; C, spathe base interior sculpturing × 4; D, spathe base interior sculpturing × 4; E, spathe base interior sculpturing × 4; F, spathe base interior sculpturing × 4; G, spathe base interior sculpturing × 4. Amorphophallus longituberosus: A, H.AM.120 (Kew illustration collection); A. corrugatus: B, H.AM. 229 (Kew illustration collection); A. johnsonii: C, H.AM. 41 (Kew slide collection); A. maculatus: D, Bogner 600 (Kew slide collection); A. parvulus: E, H.AM. 26A (Kew slide collection); A. henryi: F, H.AM. 97 (Kew slide collection); A. bulbifer: G, Bogner s.n., no other data (Kew slide collection). A. elliotii), campanulate to cylindric or ventricose, inner sur- smooth, thin, embryo large, somewhat green superficially, face smooth or longitudinally ribbed or near base endosperm absent. See Plates 79i–iv, 124C–D. verruculose, scabrate or densely covered with scale- or hair- CHROMOSOMES: 2n = 26, 28, 39. like processes or smooth; blade erect to spreading, smooth, DISTRIBUTION: ca. 170 spp. (W. Hetterscheid, pers. comm.); ribbed or variously undulate or frilled at margins. SPADIX: tropical Africa, Madagascar, tropical Asia, Malay Archipelago, sessile or stipitate, shorter or much longer than spathe; Melanesia, Australasia:– Angola, Australia, Bangladesh, Benin, female zone shorter, equalling or longer than male zone; Bhutan, ?Botswana, Brunei, Burkina Faso, Burma, ?Burundi, male zone cylindric, ellipsoid, conoid or obconoid, usually ?Cabinda, Cambodia, Cameroon, Central African Republic, contiguous with female, sometimes separated by a sterile Chad, S. China (incl. Taiwan), Congo, Equatorial Guinea zone which may be naked, or bear prismatic, subglobose or (Bioko, Rio Muni), Ethiopia, Gabon, Gambia, Ghana, Guinea, hair-like sterile flowers; terminal appendix usually present, Guinea-Bissau, India, Indonesia, Ivory Coast, Japan, Kenya, rarely absent or reduced to a stub, erect, sometimes hori- Laos, Liberia, Madagascar, Malawi, Malaysia, Mali, zontal, rarely pendent, very variable in shape, usually ± Mozambique, ?Namibia, Nepal, Niger, Nigeria, Papua New conoid or cylindric, rarely ± globose, sometimes ± stipitate Guinea, Philippines, ?Rwanda, Senegal, Sierra Leone, Somalia, or basally narrowed, usually smooth or bearing staminode- South Africa, Sri Lanka, Sudan, Tanzania, Thailand, Togo, like structures near base or entirely covered with staminodes, Uganda, Vietnam, Zaire, Zambia, Zimbabwe. sometimes corrugate or densely to sparsely hirsute, or ECOLOGY: tropical humid forest, seasonal forest, open grossly and irregularly crumpled. FLOWERS: unisexual, woodlands; geophytes, sometimes in humus deposits on perigone absent. MALE FLOWER: 1–6-androus, stamens free rocks (limestone), also in waste places or areas of human or sometimes connate in basal flowers or throughout male habitation (e.g. A. paeoniifolius). zone, short, filaments absent or distinct, connective fairly NOTES: About 14 sections or informal sectional groups are thick, sometimes projecting beyond thecae, thecae obovoid currently recognized, although many of these may be unnat- or oblong, opposite, dehiscing apically by an apical (rarely ural (W. Hetterscheid, pers. comm.). lateral) pore or transverse slit. POLLEN: extruded in strands, USES: The tubers of A. paeoniifolius and A. konjac are widely inaperturate, mostly ellipsoid to ellipsoid-oblong, occasion- used sources of carbohydrate foods in tropical Asia and Japan ally spherical or subsphaeroidal, medium-sized to large respectively. (mean 53 µm., range 34–82 µm.), exine striate, striate-retic- ETYMOLOGY: Greek amorphos (deformed) and phallus ulate, psilate, scabrate, areolate, fossulate, (penis), referring to the spadix appendix (especially of A. punctate-foveolate, verrucate, or spinose. FEMALE FLOWER: paeoniifolius). gynoecia usually crowded, sometimes ± distant, ovary sub- TAXONOMIC ACCOUNTS: Engler (1911), Gagnepain (1942), globose to ovoid or obovoid, 1–4-locular, ovules 1 per Hepper (1968), Hu (1968), Bogner (1976a), Li (1979), Sivadasan locule, anatropous, funicle very short to distinct, erect, pla- (1982, 1986, 1989), Knecht (1983), Bogner, Mayo & Sivadasan centa axile to basal, stylar region absent, short or very long, (1985), Mayo (1985a), Bogner (1986a), Nicolson (1988a), Ntépé- conoid to cylindric, stigma variably shaped, entire and sub- Nyame (1988), Hay (1990a, c), Bogner (1989b), Bogner & globose or 2–4-lobed or stellate or rarely punctiform, Hetterscheid (1992), Hetterscheid (1994), Hetterscheid & sometimes large and brightly coloured. BERRY: sometimes Serebryanyi (1994), Hetterscheid, Yadav & Patil (1994), very large, 1- to few-seeded, orange to red, rarely blue or Sivadasan, Mohanan & Rajkumar (1994), Bogner (1995), white, infructescence ± cylindric. SEED: ellipsoid, testa Hetterscheid & Peng (1995), Hetterscheid & Ittenbach (1996). T H O M S O N I E A E : A M O R P H O P H A L L U S 239