BOOK-ARACEAE

79. Amorphophallus C 80. Pseudodracontium boat-shaped, convolute basally, slightly fornicate, not con- stricted, green to yellow. SPADIX: sessile, shorter to subequal Pseudodracontium N.E. Brown in J. Bot. 20: 193 (1882). to spathe, female zone shorter than male, densely flowered, LECTOTYPE: P. anomalum N.E. Brown (= P. lacourii (Lind. male zone laxly flowered, contiguous with female zone, ter- & André) N.E. Brown)(see Nicolson in Taxon 16: 518. 1967). minal appendix usually shorter than male zone, with naked basal stipe, ± conic, covered with sterile male flowers. FLOW- HABIT: seasonally dormant herbs, tuber depressed-globose, ERS: unisexual, perigone absent. MALE FLOWER: 3–6-androus, napiform or irregularly elongate. LEAVES: often more than 1. flowers distinct, ± distant, filaments relatively long, connate to PETIOLE: long, usually mottled or otherwise variegated, sheath ± free, thecae subglobose, dehiscing by short slit or pore. very short and inconspicuous. BLADE: dracontioid: i.e. trisect POLLEN: inaperturate, ellipsoid to oblong, medium-sized with 3 ± equal primary divisions, anterior division sometimes (mean 48 µm.), exine narrowly striate. STERILE MALE FLOW- smaller than posterior divisions, primary divisions mostly pin- ERS: composed of clavate, irregularly ± connate staminodes, natisect or partly bipinnatisect, ultimate lobes decurrent, sessile not forming distinct floral groups. FEMALE FLOWER: ovary or shortly petiolulate, narrowly elliptic, acuminate; primary ovoid to subglobose, 1-locular, ovule 1, anatropous, funicle lateral veins of ultimate lobes pinnate, forming submarginal short, placenta basal, stylar region shortly attenuate, stigma dis- collective vein, 1 marginal vein also present, higher order coid-subcapitate. BERRY: ellipsoid, with stigma remnant venation reticulate. INFLORESCENCE: solitary, usually appear- persisting, 1-seeded. SEED: ellipsoid, testa smooth, brown, ing with leaves. PEDUNCLE: long, longer, shorter or subequal raphe conspicuous, embryo large, ellipsoid, somewhat truncate to petiole, similar in appearance to petiole. SPATHE: erect, at each end, endosperm absent. See Plates 80, 125A. 80. Pseudodracontium 240 T H E G E N E R A O F A R A C E A E

D EF C AB Plate 80. Pseudodracontium. A, habit × 1/4; B, leaflet × 1/2; C, spadix × 1; D, stamen × 8; E, gynoecium × 8; F, gynoecium, longitudinal section × 8. Pseudodracontium latifolium: A–B, Dransfield 6219 Cult. Kew. 1984–4105; C–F, Dransfield 6219 (Kew spirit collection 58899). T H O M S O N I E A E : P S E U D O D R A C O N T I U M 241

C F A D B KJ G EL H M Plate 81. Arophyton. A, habit × 1/6; B, leaf × 2/3; C, detail of leaf venation × 4; D, seedling × 2/3; E, inflorescence × 2/3; F, infructescence × 2/3; G, habit × 2/3; H, spadix × 3; J, synandrium, top view × 10; K, synandrode, side view × 10; L, gynoecium with associated synandrode × 10; M, gynoecium, longitudinal section × 10. Arophyton buchetii: A, 7542 (Kew slide collection); B–D, Bogner 207 (K); E–F, Bogner 207 (Kew spirit collection 32138 & 34394). A. tripartitum var. tripartitum: F, Bogner 205 (Kew spirit collection 34400); G, Bogner 205 (K!); H–M, Morat 3626 (K). 242 T H E G E N E R A O F A R A C E A E

CHROMOSOMES: 2n = 26. DISTRIBUTION: 7 spp.; tropical southeast Asia:– Cambodia, Laos, Thailand, Vietnam. ECOLOGY: tropical humid and deciduous forest, bamboo forest; geophytes, on forest floor. NOTE: Closely related to Amorphophallus, and character- ized by the staminodial, stipitate appendix, constantly 1-locular ovaries and male flowers with long, partially con- nate filaments. ETYMOLOGY: Greek pseudos (false) and Dracontium ; refers to the similarity to Dracontium. TAXONOMIC ACCOUNTS: Engler (1911), Gagnepain (1942), Serebryanyi (1995). C Tribe Arophyteae Tribe Arophyteae Bogner in Bot. Jahrb. 92: 9 (1972). SYNONYM: Tribe Synandrodieae Buchet in Bull. Soc. Bot. France 86: 279 (1939) Laticifers simple, articulated; seasonally dormant, stem tuber- 81. Arophyton ous (except most Arophyton spp.); primary lateral veins pinnate forming 1 submarginal collective vein and 1–2 mar- ginal veins, higher order venation parallel-pinnate to funicle usually short, placenta basal, stylar region short to rel- reticulate; spadix fertile to apex (except some Arophyton atively long and cylindric-conoid, or ± absent, stigma discoid. spp.); flowers unisexual, perigone absent; male flower a STERILE FLOWERS: consisting of a truncate synandrode with synandrium of partially or completely connate stamens central cavity. BISEXUAL FLOWERS: as for female flowers, (sometimes reduced to 1 stamen), pollen spinose; gynoe- but surrounded by synandrium bearing a few scattered the- cium surrounded by cup-like synandrodium, ovary 1-locular, cae. BERRY: ellipsoid to fusiform or clavate, red or green. ovule 1, orthotropous, placenta basal; testa thin, smooth, SEED: ellipsoid to globular, testa thin, smooth, embryo large, embryo large, endosperm absent. globular or ellipsoid, plumule subapical, endosperm absent. See Plates 81, 125B. C 81. Arophyton CHROMOSOMES: 2n = 38, 54, 76 (40). DISTRIBUTION: 7 spp.; Madagascar. ECOLOGY: tropical humid, seasonal forest or deciduous for- Arophyton Jumelle in Ann. Musée Colon. Marseille, 36 est on limestone; geophytes or epiphytes, litter-filled crevices année, sér. 4, 6 (2): 23 (1928). TYPE: Arophyton tripartitum and holes. Jumelle ETYMOLOGY: Greek aron (Arum) and phyton (plant). SYNONYMS: Synandrogyne S. Buchet in Bull. Soc. Bot. TAXONOMIC ACCOUNTS: Bogner (1972, 1975). France 86: 69 (1939); Humbertina S. Buchet in Bull. Soc. Bot. France 88: 848–849 (1941). 82. Carlephyton C HABIT: seasonally dormant herbs, rhizomatous or tuberous. LEAVES: 1–few. PETIOLE: sheath short. BLADE: cordate, has- Carlephyton Jumelle in Ann. Musée Colon. Marseille, 27 tate, trifid to trisect or pedatifid; primary lateral veins pinnate, année, ser.3, 7: 187 (1919). TYPE: C. madagascariense forming submarginal collective vein, secondary laterals some- Jumelle times parallel-pinnate, otherwise higher order venation reticulate. INFLORESCENCE: (1–)2–3 in each floral sym- HABIT: seasonally dormant herbs, tuber depressed-globose. podium. PEDUNCLE: subequal or shorter than petiole. LEAVES: 1–2(–3). PETIOLE: sheath short. BLADE: cordate; pri- SPATHE: not or slightly constricted between tube and blade, mary lateral veins pinnate, forming submarginal collective vein, tube persistent at maturity or entire spathe marcescent, blade 1–2 marginal veins also present, secondary and tertiary laterals widely spreading, white or cream to greenish within. SPADIX: mostly parallel-pinnate, higher order venation reticulate. INFLO- fertile to apex, or with short appendix, entirely free or female RESCENCE: 1–2(–3) in each floral sympodium, appearing zone partly adnate to spathe, female zone contiguous with before or with leaf. PEDUNCLE: shorter than petiole. SPATHE: male, or separated by synandrodes or a few bisexual flow- not constricted, upper half marcescent, lower part persistent, ± ers. FLOWERS: unisexual, perigone absent. MALE FLOWER: fully expanded at anthesis, later closing, cream to dull purple 2–7-androus, stamens connate, synandrium elongate-ellip- within. SPADIX: fertile to apex, lower part of female zone soid, often irregularly shaped, shallow, truncate, sometimes adnate to spathe, male and female zones contiguous or with a with a central slit, common connective broad, thecae mar- few bisexual flowers between them, sometimes (C. glauco- ginal, broad-ellipsoid to globular, dehiscing by slit or pore on phyllum) basal zone of spadix composed almost entirely of upper surface. POLLEN: inaperturate, spherical to subspher- bisexual flowers. FLOWERS: unisexual, perigone absent. MALE oidal, medium-sized (mean 32 µm., range 27–36 µm.), exine FLOWER: 1–6-androus, stamens connate, synandrium some- spinose. FEMALE FLOWER: gynoecium surrounded by cup- what angled, truncate apically, common connective broad, like synandrode, ovary 1-locular, ovule 1, orthotropous, thecae either marginal, projecting laterally and inverted, thus A R O P H Y T E A E : C A R L E P H Y T O N 243

E FH K D GJ C L B A N PM Plate 82. Carlephyton. A, habit in flower with leaf emerging × 2/3; B, leaf × 2/3; C, inflorescence × 2; D, synandrium, side view × 15; E, synandrium, top view × 15; F, gynoecium with associated synandrode × 15; G, gynoecium, longitudinal section × 15; H, bisexual flower × 15; J, bisexual flower, longitudinal section × 15; K, infructescence × 1; L, habit × 2/3; M, inflorescence × 2; N, synandrium, top view × 10; P, gynoecium with associated synandrode × 15. Carlephyton madagascariense: A, Bot. Jahrb. Syst. 92: 50, Fig. 4 (1972); B, Bogner 169 (K); C–J, Bogner 169 (Kew spirit collection 56421); K, Bogner 279 (Kew spirit collection 34032); C. diegoense: L, 6449 (Kew slide collection); M–P, Bogner 234 (Kew spirit collection 7700). 244 T H E G E N E R A O F A R A C E A E

82. Carlephyton 83. Colletogyne dehiscing by slit on lower surface, or stamens not completely gated, anther apical-oblique, thecae ellipsoid, dehiscing by connate, upper part of filaments ± free and thickened with slit. POLLEN: inaperturate, spherical, medium-sized (mean 39 thecae dehiscing by apical slit, sometimes remote. POLLEN: µm.), exine spinose, obscurely verruculate between spines, inaperturate, spherical or subspheroidal, medium-sized (mean spines long, stout. FEMALE FLOWER: gynoecium surrounded 34 µm., range 33–35 µm.), exine spinose. FEMALE FLOWER: by a cup-like, variegated synandrode, ovary ovoid-ellipsoid, gynoecium surrounded by cup-like synandrode, ovary 1-locu- 1-locular, ovule 1, orthotropous, funicle short, placenta basal, lar, ovule 1, orthotropous, funicle short, placenta basal, stylar stylar region shortly attenuate, stigma discoid-capitate. region, short, stigma discoid to subhemispheric. BISEXUAL BERRY: fusiform, stigma remnant persistent, reddish-spot- FLOWERS: gynoecium surrounded by synandrium bearing 1–4 ted. SEED: ellipsoid, testa thin, smooth, embryo with lateral thecae, ovary as above. BERRY: ellipsoid to fusiform, orange- plumule, endosperm absent. See Plates 83, 125D. red. SEED: ellipsoid, testa thin, smooth, embryo ellipsoid, CHROMOSOMES: 2n = 54. plumule lateral, endosperm absent. See Plates 82, 125C. DISTRIBUTION: 1 sp.; northern Madagascar. CHROMOSOMES: 54, 108. ECOLOGY: tropical deciduous forest, on limestone; geo- DISTRIBUTION: 3 spp.; northern Madagascar. phytes in holes or crevices with leaf litter. ECOLOGY: tropical deciduous forest on limestone or basalt; NOTES: The 1-androus male flowers are thought to have geophytes, in rock crevices and holes with leaf litter. been derived phylogenetically from multistaminate synan- NOTES: Bogner (1972) recognized 2 sections:– sect. dria (Bogner 1972). Carlephyton, sect. Pseudocolletogyne. ETYMOLOGY: Greek kollêtos (glued together) and gynê ETYMOLOGY: named for G. Carle with the Greek suffix (woman); the female part of the spadix is adnate to the spathe. -phyton (plant). TAXONOMIC ACCOUNTS: Bogner (1972, 1975). TAXONOMIC ACCOUNTS: Bogner (1972, 1975). C 83. Colletogyne Tribe Peltandreae C Tribe Peltandreae Engler in Nova Acta Acad. Leopold.- Colletogyne Buchet in Bull. Soc. Bot. France 86: 23 (1939). Carol. 39: 146 (1876). TYPE: C. perrieri Buchet SYNONYM: Tribe Typhonodoreae Engler in Nova Acta HABIT: seasonally dormant herb, tuberous. LEAVES: 1 (–2). Acad. Leopold.-Carol. 39: 146 (1876). PETIOLE: sheath short. BLADE: cordate; primary lateral veins Laticifers simple, articulated; rhizome hypogeal; primary lat- pinnate and also arising from petiole insertion, forming eral veins of leaf blade pinnate, forming submarginal submarginal collective vein, 1–2 marginal veins also present, collective vein, 1–2 marginal veins also present, higher order secondary and tertiary laterals mostly parallel-pinnate, higher venation ± parallel-pinnate; inflorescences usually 2, order venation reticulate. INFLORESCENCE: 1–3 in each flo- infructescence pendent; spathe ± constricted centrally, tube ral sympodium, appearing before or with leaves. PEDUNCLE: ellipsoid to oblong, persistent, blade gaping at anthesis, later shorter than petiole. SPATHE: obovate, not constricted, erect, marcescent; flowers unisexual, perigone absent; male flower fully expanded at anthesis, closing later and persisting to a truncate, prismatic synandrium of connate stamens, fused fruiting stage, red-spotted on greenish background. SPADIX: connectives very thick, thecae lateral, dehiscing by apical fertile to apex, female zone adnate to spathe, female and pore or short slit; ovary 1-locular, ovules orthotropous, style male zones contiguous. FLOWERS: unisexual, perigone narrower than ovary; berry and seeds large, embryo large, absent. MALE FLOWER: 1-androus, filament conoid, varie- plumule well-developed, endosperm vestigial to absent. P E L T A N D R E A E : P E L T A N D R A 245

B A D EF C Plate 83. Colletogyne. A, habit × 2/3; B, leaf × 2/3; C, inflorescence × 2; D, synandrium × 16; E, gynoecium with associated synandrode × 16; F, gynoecium, longitudinal section × 16. Colletogyne perrieri: A, 5489 (Kew slide collection); B–F, Bogner 165 (Kew spirit collection 42467 & photograph (M)). 246 T H E G E N E R A O F A R A C E A E

C 84. Peltandra ovary, stigma discoid-hemispheric. BERRY: obconical to irreg- ularly subglobose, green to blackish-purple (P. virginica) or Peltandra Rafinesque in J. Phys. Chim. Hist. Nat. Arts 89: 103 red (P. sagittifolia), 1–3-seeded, with mucilaginous contents. (1819), nom. cons. TYPE: P. undulata Rafinesque (= P. vir- SEED: large, ovoid to subglobose, laterally flattened, embryo ginica (L.) Rafinesque). large, plumule well-developed with 6–7 leaf primordia, endosperm vestigial or absent. See Plates 84, 126A. SYNONYMS: Lecontia W. Cooper ex Torrey, Compend. CHROMOSOMES: 2n = 56, 112. fl. n. middle stat. 358 bis (1826); Rensselaeria L.C. Beck, Bot. DISTRIBUTION: 2 spp.; eastern North America:– Canada N. Middle U.S. 382 (1833). (Ontario, Quebec), USA (Alabama, Arkansas, Carolinas, HABIT: seasonally dormant herbs, rhizome short, hypogeal, Connecticut, Delaware, Florida, Georgia, Illinois, Indiana, sometimes branching. LEAVES: several. PETIOLE: sheath Kansas, Kentucky, Louisiana, Maine, Maryland, rather long, up to at least half as long as petiole. BLADE: Massachusetts, Michigan, Mississippi, Missouri, New lanceolate to broadly ovate, sagittate, hastate or rarely cordate; Hampshire, New Jersey, New York, Ohio, Pennsylvania, basal ribs well-developed, primary lateral veins pinnate, some- Rhode Is., Tennessee, Texas, Vermont, Virginia, West Virginia, times only weakly differentiated, forming submarginal Wisconsin). collective vein, 2 or more distinct marginal veins also present, ECOLOGY: temperate wetland habitats; helophytes, marshes, secondary laterals parallel-pinnate, higher order venation par- along watercourses, brackish water. allel-pinnate near midrib, becoming reticulate towards margin. ETYMOLOGY: Greek peltê (small shield) and aner, andros INFLORESCENCE: 1–2 in each floral sympodium. PEDUN- (man). CLE: subequal to or shorter than petiole, erect at anthesis, bent TAXONOMIC ACCOUNTS: Engler (1915), Barkley (1944), downwards in fruit. SPATHE: constricted between tube and Huttleston (1953), Blackwell & Blackwell (1975). blade, tube convolute, externally green to yellow-green, ellip- soid, persistent, blade green to white, erect, widely expanded (P. sagittifolia) or only gaping (P. virginica), later deciduous. 85. Typhonodorum C SPADIX: sessile, cylindric, female zone shorter than male, partially adnate to spathe, either contiguous with male zone Typhonodorum Schott in Oesterr. bot. Wochenbl. 7: 70 or with a short zone of sterile flowers in between, male zone (1857). TYPE: T. lindleyanum Schott with short terminal appendix of sterile flowers. FLOWERS: unisexual, perigone absent. MALE FLOWER: synandrium 4–5- SYNONYM: Arodendron Werth in Mitt. Sem. Orient. androus, truncate, often slightly excavated centrally, anthers Sprachen 4 (Abt. 3): 112 (1901). lateral, thecae adjacent, oblong-ellipsoid, dehiscing by apical HABIT: evergreen, robust to gigantic herb (to 4m), rhizome pore. POLLEN: inaperturate, ellipsoid or spherical, medium- thick, subterranean, spreading, terminal shoot forming mas- sized (mean 33 µm., range 28–37 µm.), exine spinose (P. sive banana-like pseudostem formed by the lower part of the virginica) or almost psilate (P. sagittifolia). FEMALE FLOWER: petiole sheaths. LEAVES: several in terminal crown. PETIOLE: gynoecium closely surrounded by 3–5 free staminodes or very long, free apical part relatively short, sheath extending some staminodes free and others connate or all connate into for most of petiole length. BLADE: very large, subtriangular a truncate-urceolate synandrode, ovary ovoid, 1-locular, to sagittate, acute; basal ribs well-developed, primary lateral ovules 1–few, hemiorthotropous, funicle short, placenta pari- veins pinnate, forming submarginal collective vein very near etal to basal, stylar region shortly attenuate, narrower than to margin, 3–4 marginal veins also present, secondary lateral 84. Peltandra 85. Typhonodorum P E L T A N D R E A E : T Y P H O N O D O R U M 247

B D E A G F CH Plate 84. Peltandra. A, habit, rhizome cut through × 1/2; B, detail of leaf venation × 5; C, spadix × 1; D, synandrium, top view × 8; E, synan- drium, side view × 8; F, gynoecium with associated staminodes, three quarter view × 8; G, gynoecium, longitudinal section × 8; H, berry × 2. Peltandra virginica: A, Curtis 4572 (K); Cult. Kew 1969–19636 (Kew spirit collection 29047.739); B–G, Cult. Kew 1969–19636 (Kew spirit collection 29047.739); H, Herb. Careyanum s.n. (K). 248 T H E G E N E R A O F A R A C E A E

D B A F G H K A J E CL Plate 85. Typhonodorum. A, habit in × 1/30; B, habit in flower and fruit × 1/25; C, leaf × 1/5; D, detail of leaf venation × 4; E, spadix × 2/3; F, staminodes from upper sterile male zone × 5; G, synandrium × 5; H, synandrode from lower sterile male zone × 5; J, gynoecium with associated staminodes × 5; K, gynoecium, longitudinal section × 5; L, infructescence, spathe partly removed × 1/2. Typhonodorum lindleyanum: A, 5497 (Kew slide collection); B, 4M 4.91 (Kew slide collection); C, Cult. Kew 1979–4434; D, Vaughn 445 (K); E–L, Cult. Kew 1979–4434 (Kew spirit collection 29047.658); Lourance s.n. (Kew spirit collection 29047.356). P E L T A N D R E A E : T Y P H O N O D O R U M 249

and higher order venation densely parallel-pinnate. INFLO- ETYMOLOGY: Greek typhôn (personification of stormy C RESCENCE: 1–2 in each floral sympodium. PEDUNCLE: winds) and doron (gift); a poetic name. C erect, bent downwards in fruit. SPATHE: erect, constricted TAXONOMIC ACCOUNTS: Engler (1915), Bogner (1975), between tube and blade, tube convolute, oblong-ellipsoid, Mayo (1985a). green, persistent, blade longer than tube, narrowly boat- shaped, acuminate, gaping at anthesis, cream-coloured, Tribe Arisareae marcescent, later deciduous. SPADIX: shorter than spathe, sessile, free, differentiated into 4 zones, basal female zone Tribe Arisareae Dumortier, Fl. Belg. 162 (1827). with staminodes scattered between gynoecia, separated from male zone by narrower zone of sterile male flowers, male Laticifers simple, articulated; seasonally dormant, stem a tuber zone thicker, terminal zone composed of sterile male flow- or rhizome; leaves 1–3; petiole usually relatively long; pri- ers. FLOWERS: unisexual, perigone absent. MALE FLOWER: mary lateral veins forming submarginal collective vein, higher 4–6-androus, stamens connate into truncate synandrium, order venation reticulate; inflorescence 1, appearing with connective very broad, thecae lateral, sessile, dehiscing by leaves; spathe not constricted, tube margins connate, blade short longitudinal slit. POLLEN: inaperturate, ellipsoid, fornicate, gaping; spadix with usually smooth, sterile, termi- medium-sized (mean 45 µm.), exine subverrucate or nal appendix, female zone adnate to spathe, very short, obscurely foveolate or punctate. LOWER STERILE MALE contiguous with male, male zone laxly flowered; flowers FLOWERS: synandrode truncate, subhexagonal becoming unisexual, perigone absent; male flower 1-androus, filament narrowly rhomboid to trapezoid towards male zone. UPPER distinct, anther peltately attached, circular, thecae apically STERILE MALE FLOWERS: composed of 3–6 obpyramidal, confluent, dehiscing by 1 continuous slit, pollen ellipsoid, stri- truncate staminodes, irregularly connate or merely crowded ate-reticulate; gynoecia 2–5, ovary 1-locular, ovules several, together. FEMALE FLOWER: gynoecia surrounded by irreg- orthotropous, placenta basal; endosperm copious. ularly scattered, obpyramidal, truncate staminodes, ovary shortly ovoid to globose, 1-locular, ovule 1(–2), 86. Arisarum orthotropous, funicle thick, short, placenta basal, stylar region very shortly attenuate, stigma rather broad, shallowly Arisarum P. Miller, Gard. Dict. abr. ed., [121] (1754). LEC- 3–6-lobed. BERRY: borne in subcylindric, pendent infructes- TOTYPE: A. vulgare Targioni-Tozzetti (Arum arisarum L., cence enclosed in persistent spathe tube, berries very large, see Nicolson in Taxon, 24: 467. 1975). compressed-obovoid to -globose, bright yellow, pericarp fleshy. SEED: large, compressed-obovoid, testa thin, ± SYNONYMS: Arisaron Adanson, Fam. 2: 470 (1763); smooth, embryo very large, plumule exceptionally well- Balmisa M. Lagasca, Gen. Sp. Pl. Nov. 17 (1816). developed with several leaf primordia, endosperm absent. See Plates 85, 126B. HABIT: small, seasonally dormant herbs, stem an ovoid to CHROMOSOMES: 2n = 112. cylindric tuber or slender rhizome (A. proboscideum) with DISTRIBUTION: 1 sp.; Comores, Madagascar, Mauritius, stolons. LEAVES: 1–2(–3). PETIOLE: often sparsely spotted, Tanzania (Pemba, Zanzibar). sheath short. BLADE: cordate-sagittate to subhastate; pri- ECOLOGY: tropical wetland habitats; helophytes, freshwater mary lateral veins pinnate and also arising at petiole swamps, along rivers, coastal lagoons in brackish water, insertion, forming submarginal collective vein, 2 marginal forming huge populations. 86. Arisarum 250 T H E G E N E R A O F A R A C E A E

F BA GJ C DE HK Plate 86. Arisarum. A, habit × 2/3; B, spadix × 2; C, stamen × 15; D, gynoecium × 15; E, gynoecium, longitudinal section × 15; F, infructes- cence × 2; G, spadix × 2; H, habit × 2/3; J, spadix × 2; K, infructescence × 2. Arisarum proboscideum: A, Sievercori s.n. (K); B–E, Cult. Boyce (Kew spirit collection 53920); F, Cult. Meikle (Kew spirit collection 28255); A. simorrhinum: G, Cult. Salmon (Kew spirit collection 53914); A. vulgare : H, Mitchell, Cheese & Watson 3880 (K); J, Cook & Keesing (Kew spirit collection 51085); K, Cult. Kew Marr 1996 (Kew spirit collection 51128). A R I S A R E A E : A R I S A R U M 251

veins also present, higher order venation reticulate. INFLO- gynoecium 1, ovary 1-locular, ovules many, orthotropous, C RESCENCE: solitary, appearing with leaves. PEDUNCLE: placenta basal, style long, curved, stigma discoid; berry many- shorter or equalling leaf, often spotted. SPATHE: eventually seeded; seed subglobose, testa costate, strophiolate, evanescent, tube erect, margins connate, cylindric to endosperm copious. subventricose, slightly constricted at apex, white or with white to pale green longitudinal stripes, blade fornicate, 87. Ambrosina gaping, sometimes subtomentose, apex cuspidate or drawn out into a very long, erect to twisted thread, greenish or Ambrosina Bassi, Ambrosina Nov. Pl. Gen. 3 (1763); brownish or purple-brown. SPADIX: female zone adnate to Bononiensi Sci. Inst. Acad. Comment. 5 (2): 82–86 (1766, spathe, 2–5-flowered, contiguous with male zone, male zone “1767”). TYPE: A. bassii L. (“Ambrosinia ”). laxly flowered, extending for more than half spathe tube length, terminal appendix naked, either stipitate with mas- SYNONYM: [Ambrosinia L., Gen. pl., ed. 6, 579 (1764), sive apical knob, or stipitate with thick, clavate, basally orth. var.]. truncate, fungoid, apical region, or not stipitate and slenderly clavate. FLOWERS: unisexual, perigone absent. MALE HABIT: very small herb, seasonally dormant, stem a rhi- FLOWER: 1-androus, filament terete, as long or longer than zomatous tuber. LEAVES: 2–4. PETIOLE: sheath short. anther, anther peltately attached, circular, connective slender, BLADE: ovate or ovate-elliptic, obtuse, often spotted; pri- thecae apically confluent, dehiscing by single continuous slit. mary lateral veins 2–3 on each side, mostly arising at petiole POLLEN: pollen shed in amorphous mass, inaperturate, ellip- insertion, arcuate, running into apex, higher order venation soid-elongate, medium-sized (mean 45 µm., range 43–46 reticulate. INFLORESCENCE: solitary, lying horizontally on µm.), exine striate to foveo-reticulate. FEMALE FLOWER: ground. PEDUNCLE: short, hypogeal, elongating in fruit. ovary 1-locular, depressed-globose, ovules many, SPATHE: ellipsoid, boat-shaped, not constricted, basally con- orthotropous, funicle short, placenta basal, stylar region ± volute, gaping above, interior surfaces bearing hair-like abruptly narrowed, stigma small, subhemispheric. BERRY: processes, stellate hairs occurring on inner and outer sur- hemispheric, flattened at apex with elevated angled margins, faces of ventral (pistillate) chamber, apex forming curved few-seeded, pericarp carnose-leathery, style base persistent. beak. SPADIX: enclosed by the spathe, shortly appendicu- SEED: ovoid, with large, irregularly conoid strophiole (aril), late, adnate to internal wall of spathe by septum-like lateral testa longitudinally rugose, embryo terete, straight, axile, dilations thus forming two longitudinal chambers separating endosperm copious. See Plates 86, 126C. male flowers from female; ventral chamber (held upper- CHROMOSOMES: 2n = 28, 42, 56. most) containing single female flower, dorsal chamber (held DISTRIBUTION: 3 spp.; Mediterranean Europe, Macaronesia:– lowermost) containing usually 16 thecae arranged in two Albania, Algeria, Bosnia-Hercegovina, Croatia, Cyprus, Egypt, parallel rows of 8 each. FLOWERS: unisexual, perigone France (incl. Corsica), Greece (incl. Crete), Israel, Italy (incl. absent. MALE FLOWER: each row of 8 thecae represents 4 Sardinia, Sicily), Lebanon, Libya, Macedonia, Morocco, sessile stamens (see note), thecae oriented transversely, Portugal (incl. Azores), Serbia, Slovenia, Spain (incl. Balearics, opening by longitudinal slit. POLLEN: extruded in irregular Canary Is.), Syria, Tunisia, Turkey. masses, inaperturate, ellipsoid-oblong, medium-sized (44 ECOLOGY: warm temperate scrub and woodland; geophytes, µm.), exine striate-reticulate, with narrow undulate striae in on stony ground in macchie (“maquis”), between rocks or “football” pattern, breaking up into verrucae at ends. under trees and shrubs. FEMALE FLOWER: ovary 1-locular, ovules many, ETYMOLOGY: Greek word arisaron (as used by Dioscorides orthotropous, funicles rather long, placenta discoid, basal, in reference to aris, aridos, the name of a small herb men- stylar region long, attenuate, curved towards spadix axis, tioned by Pliny, possibly Arisarum itself and aron (Arum)). stigma discoid, flattened, held ± parallel to spadix axis. TAXONOMIC ACCOUNTS: Engler (1920a), Boyce (1990). BERRY: depressed-globose, many-seeded, style and old C Tribe Ambrosineae 87. Ambrosina Tribe Ambrosineae Schott in Schott & Endlicher, Melet. Bot. 16 (1832 “Ambrosinieae”). Laticifers simple, articulated; very small, seasonally dormant, stem a rhizomatous tuber; leaves several, blade ovate-ellip- tic, primary lateral veins mostly arising at petiole insertion, arcuate, running into apex, higher order venation reticulate; inflorescence 1, on recurved peduncle; spathe boat-shaped, unconstricted externally, margins free, basal part convolute, apical part narrowly gaping, rostrate, internally divided into 2 chambers, an upper, morphologically ventral one and a lower, morphologically dorsal one; spadix shortly appendic- ulate, adnate to spathe by longitudinal, septum-like lateral dilations, ventral chamber containing female zone, dorsal chamber containing 8–10 male flowers arranged in 2 longi- tudinal rows; flowers unisexual, perigone absent; male flower a 2-androus, sessile synandrium, thecae confluent, dehiscing by single continuous slit, pollen exine striate-reticulate; 252 T H E G E N E R A O F A R A C E A E

G B C A DEF Plate 87. Ambrosina. A, habit × 1; B, inflorescence, nearside half of spathe removed × 2; C, inflorescence, three quarter view, nearside half of spathe removed × 2; D, stamens × 8; E, gynoecium × 8; F, gynoecium, longitudinal section × 8; G, infructescence, nearside half of pericarp removed × 2. Ambrosina bassii: A, Besler s.n. (K); B–G, Bogner 2101 (Kew spirit collection K 57302). stigma persistent, whitish with reddish tinge. SEED: subglo- to reflexed; spadix with usually smooth, terminal, sterile bose to ellipsoid, testa costate with weak reticulation, brown, appendix; flowers unisexual, perigone absent; stamens usu- hard, with large, white, conical strophiole (aril), embryo ally free, connective slender, pollen exine spinose (except straight, elongate, endosperm copious. See Plates 87, 126D. Dracunculus, Biarum ditschianum; scabrous in Arum CHROMOSOMES: 2n = 22. korolkowii); ovary 1-locular, ovules orthotropous, style usu- DISTRIBUTION: 1 sp.; western Mediterranean Europe:– ally as broad as ovary (except some Biarum and Arisaema Algeria, Italy (incl. Lampedusa Is., Sardinia, Sicily). spp.), stigma hemispheric; berry subglobose; seed usually ECOLOGY: warm temperate scrub and woodland; geophytes, with rather large, fleshy strophiole (aril), testa usually rugose- usually in macchie (maquis) scrub, forest floor, humus reticulate (except Sauromatum and some Biarum spp.), deposits between rocks or in open stony ground. endosperm copious. NOTES: The morphological interpretation of the male flow- er remains uncertain. Benzing (1969: Tafel 27.I) presented an illustration which shows that in each longitudinal row, the 88. Arum C thecae are associated in pairs by their vascular supply. Both Engler (1920a) and Benzing state that the male flowers are Arum L., Sp. Pl. 964 (1753). LECTOTYPE: Arum maculatum 2-androus. However, in our opinion the evidence for this L. (see M.L. Green, Brit. Bot. 186. 1929). conclusion remains weak. ETYMOLOGY: named after B. Ambrosini (1588–1657), SYNONYMS: [Aron Adanson, Fam. 2: 470 (1763), orth. botanist at Bologna. var.]; Gymnomesium Schott in Oesterr. bot. Wochenbl. 5: 17 TAXONOMIC ACCOUNTS: Engler (1920a), Killian (1929, (1855). 1933), Benzing (1969). HABIT: seasonally dormant herbs, tuber hypogeal, subglo- bose or rhizomatous and horizontal. LEAVES: 3–4(–6) in each sympodial unit. PETIOLE: sheath long to short. BLADE: cor- C Tribe Areae date (A. pictum) or hastate-sagittate to sagittate; primary lateral veins pinnate, forming submarginal collective vein, at Tribe Areae least 1 marginal vein also present, higher order venation reticulate. INFLORESCENCE: 1, rarely 2 in each floral sympo- Laticifers simple, articulated; minute to robust, seasonally dium, usually appearing with leaves, rarely just before (A. dormant, stem a subglobose to rhizomatous tuber; leaves pictum), hidden within foliage or fully exposed above leaves. several (often 1 in Sauromatum), primary lateral veins of leaf PEDUNCLE: much shorter to much longer than petiole. blade or lobes or leaflets forming submarginal collective SPATHE: marcescent, usually strongly constricted between vein, higher order venation reticulate; inflorescence usually tube and blade, rarely not, tube convolute, cylindric to ellip- 1, appearing with or after leaves (except Sauromatum and soid, blade ovate- or oblong-lanceolate to lanceolate, ± some Biarum spp.); spathe usually strongly constricted, tube acuminate, at anthesis erect and boat-shaped or widely with convolute margins (partially to fully connate in expanded or patent and ± revolute. SPADIX: shorter to longer Sauromatum and in most Biarum spp.), blade gaping, erect than spathe, sessile, female zone cylindric, interstice sepa- A R E A E : A R U M 253

rating male and female zones usually short, rarely absent, ECOLOGY: temperate and warm temperate woodland, up C usually covered with sterile flowers (pistillodes), rarely naked, to 4400m alt.; geophytes, forest floor, hedges, orchards, male zone cylindric, conoid, ellipsoid or subglobose, inter- stony open ground, along rivers, open scrub, pastures, stice separating appendix and male zone usually covered abandoned areas. with sterile flowers (staminodes), rarely absent, terminal ster- NOTES: Boyce (1989, 1993) recognized 2 subgenera, subgen. ile appendix usually stipitate, then gradually or suddenly Arum (with 2 sections and 6 subsections) and subgen. dilated into a conoid or cylindric club, sometimes slender. Gymnomesium. Arum italicum is naturalized in Argentina FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4- and New Zealand. androus, stamens free, filaments very short but distinct, ETYMOLOGY: Latin form of the classical Greek name aron. connective slender, thecae shortly obovoid, opposite or sub- TAXONOMIC ACCOUNTS: Engler (1920a), Arrigoni et al. opposite, dehiscing by apical pore-like slit, rarely porose (A. (1982), Boyce (1989, 1993a,1995). pictum). POLLEN: usually presented in a loose, powdery mass, sometimes extruded in strands (A. pictum, A. nigrum), 89. Eminium grains inaperturate, spherical to subspheroidal, medium-sized (mean 32 µm., range 29–34 µm.), exine spinose, rarely Eminium (Blume) Schott, Aroideae 16 (1855, “1853”). LEC- scabrous (A. korolkowii). STERILE FLOWERS: (staminodes TOTYPE: E. spiculatum (Blume) Schott (Arum spiculatum and pistillodes) consisting of basal, ± hemispheric, swollen, Blume; see Nicolson 1967, p. 516). verrucose to smooth portion bearing usually one (sometimes more) erect, ± straight, subulate to filiform processes. SYNONYM: Helicophyllum Schott, Aroideae 20 (1855, FEMALE FLOWER: gynoecium cylindric, ovary 1-locular, “1853”), non Bridel (1827). ovules 6–more, orthotropous, lageniform, biseriate, funicle short, placenta parietal to subbasal, stylar region short, as HABIT: medium-sized, seasonally dormant herbs, tuber sub- broad as ovary or absent, stigma subhemispheric, exuding globose. LEAVES: 3–6 (–8). PETIOLE: sheath relatively long. nectar droplet at anthesis. BERRY: obovoid, pericarp juicy, BLADE: oblong-elliptic (E. lehmannii, E. regelii), linear- to many-seeded, bright glossy orange-red. SEED: ellipsoid to auriculate-hastate, or pedatifid-pedatisect with lobes of pos- ovoid, testa rugose, especially towards base, with large, suc- terior divisions held ± erect in a spiral on the twisted basal culent obconic strophiole, embryo axile, straight, endosperm ribs; primary lateral veins of lobes pinnate, forming submar- copious. See Plates 88, 127A. ginal collective vein, higher order venation reticulate. CHROMOSOMES: 2n = 28, 42, 56, 70, 84. INFLORESCENCE: solitary, appearing with leaves. PEDUN- DISTRIBUTION: 25 spp.; Central Asia, Europe, Macaronesia, CLE: much shorter than petiole of subtending leaf, apex often Mediterranean, Middle East:– Afghanistan, Albania, Algeria, greatly thickened. SPATHE: marcescent, tube with convolute Andorra, Armenia, Austria, Azerbaijan, Belgium, Belorussia margins, subventricose to oblong, blade oblong or ovate- (S), Bosnia–Hercegovina, Bulgaria, China (Tien Shan), oblong, erect, inner surface sometimes densely Croatia, Cyprus, Czech Republic, Denmark, France (incl. rugose-puckered. SPADIX: sessile, slender, shorter than Corsica), Georgia, Germany, Greece (incl. islands), Hungary, spathe, female zone short-cylindric, separated from male zone India (NW), Iran, Iraq, Ireland, Israel, Italy (incl. Sardinia, by longer zone bearing usually scattered sterile flowers, male Sicily), Jordan, Kazakhstan, Kirghizia, Lebanon, Libya, zone ellipsoid to cylindric, shorter, equalling or longer than Liechtenstein, Luxembourg, Macedonia, Moldavia, Morocco, female, appendix usually relatively short, elongate-clavate, Nepal (W), Netherlands, Pakistan, Poland, Portugal (incl. stoutly to narrowly cylindric, rugose or smooth. FLOWERS: Azores, Madeira), Romania, Russia (far S), San Morino, unisexual, perigone absent. MALE FLOWER: 2-androus, sta- Serbia, Slovak Republic, Slovenia, Spain (incl. Canary Is., mens free, anthers subsessile to sessile, connective slender, Balearics), Sweden (S), Switzerland, Syria, Tajikistan, Tunisia, thecae oblong-ellipsoid, dehiscing by apical slit. POLLEN: Turkey, Turkmenestan, Ukraine (S), United Kingdom (incl. presented in a loose, powdery mass, not extruded in strands, Channel Is.), Uzbekistan. grains inaperturate, spherical or subspheroidal, medium-sized 88. Arum 89. Eminium 254 T H E G E N E R A O F A R A C E A E

K C BF J E G D H AA XY A U Z V LM NP QR ST W Plate 88. Arum. A, habit × 1/5; B, tuber × 2/3; C, tuber × 2/3; D, leaf × 1; E, detail of leaf venation next to midrib × 6; F, detail of leaf vena- tion near margin × 6; G, leaf × 2/3; H, inflorescence, front of lower spathe removed × 2/3; J, spathe, front view × 2/3; K, spathe, side view × 2/3; L, spadix × 2/3; M, detail of fertile portion of spadix × 2; N, spadix × 2/3; P, detail of fertile portion of spadix × 2; Q, spadix × 2/3; R, detail of fertile portion of spadix × 2; S, spadix × 2/3; T, detail of fertile portion of spadix × 2; U, stamen × 10; V, stamens, lower one longi- tudinally sectioned × 10; W, gynoecia, upper one longitudinally sectioned × 10; X, pistillode × 15; Y, staminode × 15; Z, infructescence × 2/3; AA, seed × 2. Arum maculatum: A, Boyce s.n. (Kew slide collection); A. alpinum: B, Cook & Keesing 56 (Kew spirit collection 51396); A. concinnatum: C, L–M, U, X–Y, Boyce 10 (Kew spirit collection 53952); A. rupicola var. rupicola: D–F, Sintenis 5575 (K); A. italicum subsp. italicum: G, Herb. Churchill s.n. (K); A. orientale subsp. orientale: H, Cult. Tucker (Kew spirit collection 49773); A. rupicola subsp. rupi- cola: J, Boyce 87 (Kew spirit collection 51615); A. creticum: K, N–P, Cult. Kew (Kew spirit collection 51368); A. orientale subsp. orientale: Q–R, Cult. Tucker (Kew spirit collection 49773); A. pictum: S–T, V–W, Ferguson & Ferguson 4126, Cult. Kew 1985–01888 (Kew spirit collec- tion 51818); A. italicum subsp. italicum: Z, photo 1336 (Kew slide collection); A. alpinum: AA, Cook & Keesing 56 (Kew spirit collection 51898). A R E A E : A R U M 255

BC L D A F EG H M NK J Plate 89. Eminium. A, habit × 1/3; B, seedling showing contractile root × 2/3; C, detail of fertile portion of spadix × 1 1/2; D, habit × 2/3; E, spadix × 1 1/2; F, stamen × 15; G, gynoecium × 15; H, gynoecium, longitudinal section × 15; J, inflorescence × 2/3; K, spadix × 1 1/2; L, stamen × 15; M, gynoecium × 15; N, gynoecium, longitudinal section × 15. Eminium albertii: A, Furse 7390 (K); Gibbons & Gibbons 47 (K); B, Gibbons & Gibbons 47 (K); C, Grey-Wilson & Hewer 765 (Kew spirit collection 29047.120); E. koenenianum: D, Willdenowia 20: 47, f.4 (1991); E–H, Koenen 32*89 (Kew spirit collection 55074 & 59044); E. spiculatum subsp. spiculatum: J, Boyce 197 (Kew spirit collection 51645 & Kew slide collection); K–N, Boyce 197 (Kew spirit collection 51645). 256 T H E G E N E R A O F A R A C E A E

(mean 44 µm.; 15–20 µm in E. koenenianum), exine spinose. with male zone, male zone ellipsoid-cylindric, separated STERILE FLOWERS: composed of subulate, sometimes flat- from appendix by a short zone of subulate staminodes, tened, patent, straight to slightly curved processes. FEMALE appendix long, long-stipitate, then elongate-conoid. FLOWER: ovary ellipsoid-obovoid, 1-locular, ovules 2, FLOWERS: unisexual, perigone absent. MALE FLOWER: 3–4- orthotropous, funicle short, placenta basal to subbasal, stylar androus, stamens quadrate, filaments distinct, connate at region short to inconspicuous, stigma hemispheric. BERRY: base, connective slender and prominulent, thecae shortly subglobose, 1(–2) seeded. SEED: obnapiform to subglobose, ellipsoid, dehiscing by apical pore. POLLEN: extruded in testa leathery, rugose, with large strophiole, embryo small, strands, inaperturate, ellipsoid to spheroidal, medium-sized elongate, endosperm copious. See Plates 89, 127B. (mean 45 µm.), exine verrucate. FEMALE FLOWER: ovary CHROMOSOMES: 2n = 24, 28. ellipsoid, 1-locular, ovules few, orthotropous, placentae api- DISTRIBUTION: 8 spp.; Middle East, south central Asia: cal and basal, stylar region shortly conic, stigma hemispheric. Afghanistan, Egypt, Iran (N), Iraq (N), Israel, Jordan, BERRY: obovoid, few-seeded, red-orange. SEED: globose- Kazakhstan, Kirghizia, Lebanon, Syria, Tajikistan, Turkey, ovoid, testa rugose, embryo axile, short, endosperm copious. Turkmenestan, Uzbekistan. See Plates 90, 127C. ECOLOGY: warm temperate and subtropical habitats in fields, CHROMOSOMES: 2n = 28. savannas, semideserts, deserts; geophytes, stony or sandy DISTRIBUTION: 2 spp.; Mediterranean Europe, ground, consolidated sand. Macaronesia:– Albania, Bosnia–Hercegovina, Bulgaria, ETYMOLOGY: ancient name eminion, mentioned by Croatia, Greece (incl. Crete and islands), Italy (incl. Sardinia, Dodoens (1574). Sicily), Macedonia, Portugal (incl. Azores, Madeira), ? Serbia, TAXONOMIC ACCOUNTS: Engler (1920a), Riedl (1963, 1969, Slovenia, Spain (Canary Is.), Turkey. 1985), Lobin & Boyce (1991). ECOLOGY: temperate and warm temperate scrub, wood- land, macchie, olive groves, chestnut forest; geophytes, under trees and shrubs, also stony open ground. C 90. Dracunculus NOTES: The verrucate pollen exine of Dracunculus is unique in the tribe. The other genera have a spinose, spinulose or Dracunculus P. Miller, Gard. Dict., Abr. ed., [455] (1754). rarely smooth exine. The seed of Dracunculus lacks the LECTOTYPE: D. vulgaris Schott (see Nicolson 1975, p. 467). characteristic strophiole found in other genera of the tribe. SYNONYMS: Anarmodium Schott in Bonplandia 9: 368 ETYMOLOGY: ancient name, Latin draco (dragon, snake) (1861); Dracontium J. Hill, Brit. Herb. 336 (1756, non L. and suffix -unculus (diminutive). 1753). TAXONOMIC ACCOUNTS: Engler (1920a), Amaral Franco, Webb & Prime (1980), Boyce (1994). HABIT: large, seasonally dormant herbs, tuber subglobose, stoloniferous as well in D. canariensis. LEAVES: few. PETI- OLE: sheath very long, spotted or not, tightly convolute 91. Helicodiceros C around peduncle forming substantial erect pseudostem. BLADE: deeply pedatifid; primary lateral veins of each lobe Helicodiceros Schott ex K. Koch in Index Sem. Hort. Berol. pinnate, forming submarginal collective vein, higher order 1855, Appendix, p. 2 (1856), nom. cons. TYPE: H. crinitus K. venation reticulate. INFLORESCENCE: solitary, appearing Koch, nom. illeg. (Dracunculus crinitus Schott, nom. illeg., with leaves. PEDUNCLE: shorter than petiole. SPATHE: Arum muscivorum L. f., H. muscivorus (L.f.) Engler). marcescent, constricted, tube with convolute margins, erect, thick-walled, oblong-ellipsoid, shorter than blade, blade SYNONYM: Megotigea Rafinesque, Fl. Tell. 3: 64 (1837, ovate-lanceolate, acuminate, expanded, deep purple or “1836”), nom. rej. white, eventually bent backwards. SPADIX: subequal to HABIT: seasonally dormant herb, tuber depressed-globose. spathe, female zone ± sessile, cylindric to conic, contiguous LEAVES: several. PETIOLE: variously spotted, sheath long, 90. Dracunculus A R E A E : H E L I C O D I C E R O S 257

B G J KL H D A EF C Plate 90. Dracunculus. A, habit × 1/6; B, leaf × 1/3; C, detail of spadix fertile portion and appendix base × 1; D, stamen × 10; E, gynoe- cium × 10; F, gynoecium, longitudinal section × 10; G, infructescence × 2/3; H, spadix × 1; J, stamen × 10; K, gynoecium × 10; L, gynoecium, longitudinal section × 10. Dracunculus vulgaris: A, Fleming s.n. (Kew slide collection); B, Cult. Cheston (K); C–F, Boyce 19 (Kew spirit collection 51381); G, Cult. Kew (Kew spirit collection 25760); Zabeau 11/65 (Kew slide collection); D. canariensis: H–L, Cult. Boyce (Kew spirit collection 29047.679). 258 T H E G E N E R A O F A R A C E A E

B E A FG C D Plate 91. Helicodiceros. A, habit × 1/3; B, detail of hairs on spathe limb interior × 1; C, spadix × 2/3; D, detail of spadix fertile portion and appendix base × 1; E, stamen × 10; F, gynoecium × 10; G, gynoecium, longitudinal section × 10. Helicodiceros muscivorus: A, Boyce 36 (Kew slide collection); B–G, Boyce 36 (Kew spirit collection 29047.638). 91. Helicodiceros forming loose pseudostem. BLADE: deeply pedatifid, anteri- or division oblong, lobes of posterior divisions tightly spiralled on basal ribs and thus held ± erect; primary lateral veins of each lobe pinnate, weakly differentiated, forming submarginal collective vein, higher order venation reticulate. INFLORESCENCE: solitary, appearing with leaves. PEDUN- CLE: much shorter than petiole. SPATHE: spotted on outer surface, constricted between tube and blade, tube with con- volute margins, erect, green, thick, subcylindric, blade sharply bent backwards at constriction, large, oblong-ovate, widely spreading, pale purple, inner surface covered with purple hairs. SPADIX: shorter than spathe, bent backwards at spathe constriction, female zone sessile to shortly stipitate, sub- cylindric, sterile zone separating male and female zones short, covered with subulate pistillodes or naked, male zone subcylindric to ellipsoid, shorter than or subequal to female, appendix vermiform, much longer than rest of spadix, weakly stipitate, thicker at middle, attenuate towards apex and base, ± lying on spathe blade, entirely covered with upwardly curved, setiform staminodial processes becoming gradually larger and subulate towards appendix base. FLOWERS: uni- sexual, perigone absent. MALE FLOWER: 2–3-androus, A R E A E : H E L I C O D I C E R O S 259

stamens free, anthers sessile, connective slender, inconspic- uous, thecae ± ellipsoid, dehiscing by broad apical slit. POLLEN: extruded in strands, inaperturate, ± spherical, exine spinose. FEMALE FLOWER: ovary 1-locular, ovules up to 6, orthotropous, placentae 2, apical placenta with 3–4 ovules and basal placenta with 1–2 ovules, stylar region as broad or broader than ovary, stigma subhemispheric. BERRY: obovoid to ellipsoid, bearing small stigma remnant, (1–)2–3-seeded, orange. SEED: borne on either one or both placentae, broadly ovoid, with yellowish strophiole, testa leathery, strongly retic- ulate, pale brown when fresh, embryo axile, elongate, endosperm copious. See Plates 91, 127D. CHROMOSOMES: 2n = 56. DISTRIBUTION: 1 sp.; western Mediterranean Europe:– France (Corsica), Italy (Sardinia), Spain (Balearic Islands). ECOLOGY: warm temperate habitats; geophytes, in lime- stone and granite rock crevices beside the sea, rarely on stony ground. ETYMOLOGY: Greek “helix, helicos” (spiral), “dis” (twice) 92. Theriophonum and “keras” (horn), referring to the two basal lobes of the leaf that are twisted and erect like horns. TAXONOMIC ACCOUNTS: Arcangeli (1884), Engler (1920a); ovoid to ellipsoid, few-seeded. SEED: broadly ovoid to ellip- Arrigoni et al. (1982, as Dracunculus muscivorus), Boyce soid, apiculate, strophiolate, testa smooth to rugose, embryo (1994). axile, elongate, endosperm copious. See Plate 92. CHROMOSOMES: 2n = 16 (14, 18). C 92. Theriophonum DISTRIBUTION: 5 spp.; India, Sri Lanka. ECOLOGY: tropical forest; geophytes, forest floor, grassy Theriophonum Blume, Rumphia 1: 127 (1837, “1835”). places, ditches, damp sites on rocky, lateritic soil. TYPE: T. crenatum (R. Wight) Blume (Arum crenatum R. ETYMOLOGY: Greek name theriophonon from therion (wild Wight). beast) and phenô (I slay); means “beast-slayer”. TAXONOMIC ACCOUNTS: Engler (1920a), Sivadasan & SYNONYMS: Tapinocarpus Dalzell in Hooker’s J. Bot. Nicolson (1982). Kew Gard. Misc. 3: 345 (1851); Calyptrocoryne Schott in Oesterr. bot. Wochenbl. 7: 262 (1857); Pauella Ramamurthy & Sebastine in Bull. Bot. Surv. India 8: 348 (1967). 93. Typhonium C HABIT: small, seasonally dormant herbs, tuber small, Typhonium Schott in Wiener Z. Kunst 1829 (3): 732 (1829). depressed-subglobose. LEAVES: several. PETIOLE: sheath rel- LECTOTYPE: T. trilobatum (L.) Schott (Arum trilobatum L.; atively long. BLADE: sagittate-hastate or hastate, sometimes see Nicolson in Taxon 16: 519. 1967). linear-lanceolate or oblong-ovate; primary lateral veins pin- nate, forming submarginal collective vein, 1–2 distinct SYNONYMS: Desmesia Rafinesque, Fl. Tell. 3: 63 (1837, marginal veins also present, higher order venation reticu- “1836”); Heterostalis (Schott) Schott in Oesterr. bot. Wochenbl. late. INFLORESCENCE: solitary, appearing with the leaves. 7: 261 (1857). PEDUNCLE: usually short, erect in flower, deflexing in fruit. HABIT: very small to medium-sized, seasonally dormant or SPATHE: marcescent, constricted between tube and blade, evergreen, rarely pubescent (T. hirsutum) or glandular- tube convolute, blade much longer than tube, erect and gap- pubescent herbs, tuber globose, subglobose or irregular, ing to spreading, or sometimes ± reflexed. SPADIX: sessile, sometimes rhizomatous or stoloniferous, rarely an epigeal shorter or subequal to spathe, female zone few-flowered, stem (T. fultum). LEAVES: few to several. PETIOLE: apex and very short, separated from male by axis partly or completely middle portion rarely tuberculate, sheath rather short. covered with sterile flowers, sterile flowers either basal, cen- BLADE: usually cordate-sagittate, sagittate to hastate, trifid, tral or at apex of sterile zone, male zone cylindric, sometimes pedatifid or pedatisect, rarely linear, narrowly lanceolate, with zone of sterile flowers above, appendix stipitate or not, elliptic-oblong or cordate, apex rarely tuberculate; primary erect, shorter or much longer than rest of spadix, cylindric, lateral veins of blade or lobes pinnate, forming submarginal elongate-conoid to very long-subulate. FLOWERS: unisex- collective vein, 1–2 marginal veins also present, higher order ual, perigone absent. MALE FLOWER: 1–2-androus, stamens venation reticulate. INFLORESCENCE: solitary, appearing free, anthers subsessile, connective slender, sometimes elon- with or without (T. hayatae) or after the leaves. PEDUNCLE: gated and prominent, thecae ovoid, dehiscing by apical pore shorter than petiole. SPATHE: constricted between tube and or broad slit. POLLEN: inaperturate, spherical, medium-sized blade, tube with convolute or rarely basally connate (T. hir- (mean 25 µm.), exine spinose. STERILE FLOWERS: spread- sutum) margins, persistent or rarely evanescent (T. ing, somewhat decurved or suberect, long, slender and nudibaccatum), blade eventually bending backwards from filiform to shorter and subulate. FEMALE FLOWER: gynoecia constriction, broadly ovate to lanceolate, ± acuminate, usu- oriented vertically, ovary ellipsoid to obovoid, 1-locular, ally purple, rarely white within, tube persistent, blade ovules 3–9 (or more), orthotropous, funicle short, placentae marcescent. SPADIX: sessile, shorter, subequal or much 2, apical and basal, stigma ± sessile, discoid-hemispheric, longer than spathe, female zone cylindric, subconic to sub- sometimes rather massive with central depression. BERRY: globose, separated from male zone by rather long axis 260 T H E G E N E R A O F A R A C E A E

F H MC A J B K G L DE Plate 92. Theriophonum. A, habit × 2/3; B, detail of fertile portion of spadix and appendix base × 3; C, stamen with prominent connective × 15; D, gynoecium × 15; E, gynoecium, longitudinal section × 15; F, infructescence × 1; G, habit × 1; H, leaf × 1; J, leaf × 1; K, detail of fertile por- tion of spadix and appendix base × 4; L, detail of fertile portion of spadix and appendix base × 3; M, stamen × 15. Theriophonum minutum: A, Barnes 699 (K); Barnes 700 (K); Balasubramanian 2047 (K); Bogner s.n. (Kew slide collection); B–E, Barnes 698 (K & Kew spirit collection 58090); F, Nicolson 4266 (K); T. sivaganganum: G–K, Rajagopalan RHT 30294 (K & Kew spirit collection 58131); T. infaustum: L–M, Barnes 1333 (K & Kew spirit collection 58071). A R E A E : T H E R I O P H O N U M 261

HJ K E C A B DG F Plate 93. Typhonium. A, spadix × 1 1/2; B, habit × 1/2; C, leaf × 1/2; D, spadix × 1 1/2; E, infructescence × 2/3; F, habit, part of petiole removed × 1/2; G, detail of fertile portion of spadix × 1 1/2; H, stamen × 10; J, gynoecium, side view × 10; K, gynoecium, longitudinal sec- tion × 10. Typhonium brownii: A, Thomson 2360 (Kew spirit collection 57319); T. hirsutum: B, comm. Mayo ‘Arisaema 1’ (Kew spirit collection 29047.710); T. trilobatum: C, D, Hay 2045 (Kew spirit collection 55956); E, Walker s.n. (K); T. giganteum: F, Cult. Cambridge Bot. Gard. (K), Henry 6537 (K); Hetterscheid H.AR. 31 (Kew spirit collection 57592); G–K, Hetterscheid H.AR. 31 (Kew spirit collection 57592). 262 T H E G E N E R A O F A R A C E A E

NOTES: Some species are weedy and naturalized in various parts of the world outside Asia in tropical, subtropical and warm temperate regions. Sriboonma, Murata & Iwatsuki (1994) recognized 5 sections:– sect. Hirsuta, sect. Diversifolia, sect. Pedata, sect. Gigantea, and sect. Typhonium. ETYMOLOGY: cited by Dodoens (1574) as Greek Typhonion, an ancient plant name; Typhon, the youngest child of Gaia, the Greek Earth goddess, was a beast, half man, half reptile, later slain by Zeus (A. Hay pers. comm.) TAXONOMIC ACCOUNTS: Engler (1920a), Nicolson & Sivadasan (1981), Sivadasan (1982), Bogner (1988b), Murata (1990b), Murata & Mayo (1991), Hay (1993a), Sriboonma, Murata & Iwatsuki (1994). 94. Sauromatum C 93. Typhonium Sauromatum Schott in Schott & Endlicher, Melet. Bot. 17 (1832). LECTOTYPE: S. guttatum Schott (= S. venosum covered either entirely or only in basal part with sterile flow- (Aiton) Kunth, Arum guttatum Wallich 1831, non Salisbury ers of various shapes, rarely with sterile flowers above male 1796; see Nicolson 1967). flowers, male zone cylindric to ellipsoid, usually densely but rarely sparsely flowered (T. albispathum), appendix usually SYNONYMS: [Stauromatum Endlicher, Ench. 128 (1841), shortly stipitate, rarely with disc-like extension at the base or orth. var.]; Jaimenostia Guinea & Gómez-Moreno in Guinea, sessile (T. hirsutum), smooth, conoid to extremely slender Ensayo Geobot. 248 (1946) & in Anales Jard. Bot. Madrid 6 and filiform-subulate, usually long-exserted. FLOWERS: uni- (2): 465 (1946). sexual, perigone absent. MALE FLOWER: 1–3-androus but usually 1-androus, stamens free or sometimes ± connate, HABIT: small to medium-sized, seasonally dormant herbs, anthers subsessile, connective slender, sometimes prominu- tuber subglobose or depressed-globose, sometimes large. lent, thecae ovoid to ellipsoid, dehiscing by pore or lateral LEAVES: usually solitary, rarely up to 3. PETIOLE: often spot- slit extending to the middle or nearly to the base and con- ted, sheath very short. BLADE: deeply pedatifid to pedatisect; fluent apically. STERILE FLOWERS: either all similar or primary lateral veins of lobes pinnate, forming submarginal diverse in the same spadix, capitate, clavate to spathulate, collective vein, 1–2 marginal veins also present, higher order cylindric, filiform or subulate, rarely stout and flattened at the venation reticulate. INFLORESCENCE: solitary, appearing with tip, or reduced to verrucae, straight to flexuose, suberect, or without leaves, borne at ground level. PEDUNCLE: very spreading or decurved. POLLEN: extruded in amorphous short. SPATHE: marcescent, slightly constricted between tube mass, inaperturate, spherical to subspheroidal, medium-sized and blade, tube with connate margins, ± cylindric and usually (mean 32 µm., range 28–36 µm.), exine spinose (spines very somewhat ventricose basally, blade much longer, narrowly obtuse in e.g. T. trilobatum). FEMALE FLOWER: gynoecia ori- oblong-lanceolate, erect at first then reflexed and spiralled-rev- ented horizontally or vertically, ovary ovoid, ellipsoid or olute, margins undulate, conspicuously spotted on inner obovoid, 1-locular, ovules 1–3, orthotropous, funicle short, surface. SPADIX: subequal to spathe, free, sessile, female placenta basal, stigma sessile, discoid-hemispheric. BERRY: zone cylindric, separated from male zone by much longer axis ovoid, 1- or rarely 2-seeded, orange-red, green or white. bearing sterile flowers in lower part and naked above, male SEED: globose to obnapiform, testa thin, rugulose to smooth, zone short, cylindric, appendix elongated, very long-exserted, strophiolate, embryo axile, elongate, straight, endosperm relatively slender, cylindric, smooth, obtuse, suberect then copious. See Plates 93, 128A. somewhat forward-curving. FLOWERS: unisexual, perigone CHROMOSOMES: 2n = 16, 18, 20, 26, 36, 52, 54, 65, >100 (14). absent. MALE FLOWER: few-androus, stamens free, anthers ± DISTRIBUTION: 37 spp.; tropical south, southeast and east sessile, somewhat compressed, connective slender, thecae Asia, Malay Archipelago, Australasia:– Australia, Bangladesh, oblong-obovoid, dehiscing by broad, apical slit. POLLEN: Bhutan, Burma, Cambodia, Caroline Is., China (incl. Taiwan, extruded in amorphous mass, inaperturate, spherical to sub- Tibet), India, Indonesia (Borneo, Irian Jaya, Java, Moluccas, spheroidal, medium-sized (mean 33 µm.), exine spinose. Sumatra, Timor), Japan, Laos, Malaysia (Borneo, Peninsula), STERILE FLOWERS: consisting of distant, patent, terete, fili- Mariana Is., Mongolia, Nepal, Pakistan, Papua New Guinea, form, or clavate staminodes, apex rounded to obliquely Philippines, Singapore, Sri Lanka, Thailand, Vietnam. truncate-disciform. FEMALE FLOWER: gynoecium subcylin- Naturalized in Brazil (Northeast, Southeast), Comores, Cuba, dric, ovary 1-locular, ovules 1–4, orthotropous, funicle very Ghana, Ivory Coast, Madagascar, Martinique, Mauritius, South short, placenta basal, stylar region shortly attenuate or ± as Africa, Surinam, Tanzania (Zanzibar), Trinidad, Venezuela, broad as ovary, stigma discoid-hemispheric. BERRY: obpyra- ECOLOGY: warm temperate, subtropical and tropical humid midal, red-purple, darker and corrugated at apex, densely and seasonal forests, secondary forest, cultivated land; geo- congested in subglobose, sometimes partly hypogeal phytes, forest floor, among rocks, wet sites, streamsides, infructescence. SEED: obnapiform, testa thin, dark-spotted, grassy places. smooth to rough, strophiole present, embryo axile, elongate and curved, endosperm copious. See Plates 94, 128B. CHROMOSOMES: 2n = 26, 52, 104. DISTRIBUTION: 2 spp.; tropical Africa, Arabian peninsula, tropical and subtropical southern Asia:– Angola, Bangladesh, Bhutan, Burma, Cameroon, Central African Republic, China A R E A E : S A U R O M A T U M 263

B A L M D N PQ KC J EF G H Plate 94. Sauromatum. A, habit in flower × 1/5; B, habit × 1/5; C, leaf × 2/3; D, inflorescence × 2/3; E, spadix × 1; F, stamen × 20; G, gynoe- cium × 20; H, gynoecium, longitudinal section × 20; J, tuber in section, petiole base and mature infructescence × 1/2; K, juvenile plant × 1/2; L, inflorescence × 2/3; M, spadix × 1; N, stamen × 30; P, gynoecium × 30; Q, gynoecium, longitudinal section × 30. Sauromatum veno- sum: A, Barnes 1617; B, Andrews 6/80 (Kew slide collection); C, Andrews 27/6/82 (Kew slide collection); D, Robertson p/8/35/5 (Kew slide collection); E–H, Cult. Maw (Kew spirit collection 29047.122); J, Jarrett & Saldanha HFP 736 (K), Kultan s.n. (Kew spirit collection 6915); K, Aitchison 268 (K); S. brevipes: L–M, Templar s.n. (K); N–Q, Templar s.n. (Kew spirit collection 47636). 264 T H E G E N E R A O F A R A C E A E

94. Sauromatum (Tibet, Yunnan), Equatorial Guinea (Bioko) Ethiopia, India, appendix vermiform, slightly exceeding spathe, smooth, Kenya, Malawi, Nepal, Pakistan, Saudi Arabia, Tanzania, emerging from the mouth of the spathe tube and then Uganda, Yemen Republic, ?Zaire, Zambia. upturned. FLOWERS: unisexual, perigone absent. MALE ECOLOGY: tropical upland forest and grassland; geophytes FLOWER: 2-androus, stamens free, anthers sessile, thecae or occasionally epiphytes, shady, humid sites, also in humus opening by apical pore. POLLEN: inaperturate, ellipsoid, deposits on rocks. medium-sized (36–40µm. × 28–32µm.), exine spinose, spin- NOTES: Differs from Typhonium only in the consistently ulae 1.4– 1.6µm. long and apex somewhat rounded. STERILE fused margins of the spathe tube and the curved embryo. FLOWERS: slender, cylindric, obtuse, ascending at base, then ETYMOLOGY: Greek sauros (lizard) and shortened form of recurved and appressed against the wall of spathe tube. chrômaton (coloured); refers to lizard-like colour of the FEMALE FLOWER: lageniform, ovary ovoid, 1-locular, ovule petiole. 1, orthotropous, placenta basal, stylar region short, attenuate, TAXONOMIC ACCOUNTS: Engler (1920a), Sivadasan (1982), stigma subhemispheric. BERRY: ovoid, 1-seeded, minutely Mayo (1985a). beaked at apex, reddish, infructescence with long peduncle, with 5–7 berries in a whorl. SEED: ovoid to subglobose, micropyle prominent, strophiolate, testa minutely verrucu- C 95. Lazarum lose, embryo axile, endosperm copious. See Plate 94. CHROMOSOMES: 2n = c. 78. Lazarum A. Hay in Bot. J. Linn. Soc. 109 (3): 427–434 (1992). DISTRIBUTION: 1 sp.; Australia (Northern Territory, Melville TYPE: L. mirabile A. Hay Island). HABIT: very small, seasonally dormant herb; tuber subglo- ECOLOGY: tropical open forest; geophytes, forest floor, in bose, whitish. LEAVES: 1–3, appearing after flowering, dark laterite on ridges and in sandy soil. blueish green, subtended by deliquescent cataphylls. PETI- ETYMOLOGY: named after biblical Lazarus, in poetic refer- OLE: sheath relatively long. BLADE: elliptic, base obtuse, ence to the morbid colour of the partly buried inflorescence, apex apiculate; midrib distinct, primary lateral veins pinnate, which emerges from a leafless subterranean tuber. running ± arcuately into margin near apex, marginal vein pre- sent, higher order venation reticulate. INFLORESCENCE: 95. Lazarum solitary, subtended by long cataphylls, appearing before the leaves. PEDUNCLE: very short, elongating in fruit. SPATHE: marcescent, slightly constricted at position of an annular sep- tum through which the spadix passes and which separates the male zone from the female zone forming two chambers, swelling again for a short distance above the constriction; tube ± broadly ellipsoid, margins connate, not persistent in fruit; blade narrowly lanceolate, sharply bent up, erect, mar- gins inrolled. SPADIX: sessile, only slightly longer than spathe, female flowers free, 5–7 in a single whorl, zone of sterile flowers contiguous with female zone, male zone sep- arated from zone of sterile flowers by naked axis which passes through the central aperture of the spathe septum, male flowers thus situated in the upper spathe chamber; A R E A E : L A Z A R U M 265

A B CD Plate 95. Lazarum. A, habit × 2; B, inflorescence, nearside half and upper part of spathe removed × 6; C, habit, fruiting × 2; D, berry × 8. Lazarum mirabile: A–B, Jones 1695 (DNA spirit collection); C–D, Gallen 3 (DNA spirit collection). C 96. Biarum oblong-lanceolate, oblong-ovate or -obovate, or elliptic- oblong, margins smooth to strongly crispate; primary lateral Biarum Schott in Schott & Endlicher, Melet. Bot. 17 (1832), veins pinnate, ± forming submarginal collective vein, higher nom. cons. TYPE: B. tenuifolium (L.) Schott (Arum tenui- order venation reticulate. INFLORESCENCE: 1(–2) in each flo- folium L.), typ. cons. ral sympodium, usually appearing before leaves, rarely with or directly after the leaves at the end of the vegetative period. SYNONYMS: Homaïd Adanson, Fam. 2: 470 (1763), nom. PEDUNCLE: short, usually hypogeal. SPATHE: constricted rej.; [Homaida Rafinesque, Fl. Tell. 3: 63 (1837, “1836”), orth. between tube and blade, erect, eventually evanescent, tube var.]; Ischarum (Blume) Reichenbach, Deutsches Bot. cylindric to ventricose, hypogeal or held just above ground Herbarienbuch, Nom. 32 (1841); Leptopetion Schott, Gen. surface, margins connate near base or for most of length, Aroid. t. 8 (1858); Cyllenium Schott, Gen. Aroid. t. 9 (1858); rarely margins free (B. aleppicum), blade usually much longer, Stenurus Salisb., Gen. Pl. 5 (1866). rarely shorter and reduced, expanded to gaping, linear to HABIT: seasonally dormant herbs, tuber subglobose, smooth. oblong-lanceolate or oblong, rarely campanulate or fornicate LEAVES: few to many. PETIOLE: often only weakly differen- or subtriangular, usually dark brown-purple within, rarely tiated from blade, sheath long to short. BLADE: linear, white, pale yellowish or red-purple. SPADIX: sessile, shorter 266 T H E G E N E R A O F A R A C E A E

C A B DE P H Q F N RS KG J ML Plate 96. Biarum. A, leaf × 2/3; B, detail of fertile portion of spadix and appendix base × 2; C, stamen × 10; D, gynoecium × 10; E, gynoe- cium, longitudinal section × 10; F, berry, longitudinal section × 2; G, habit × 2/3; H, habit × 2/3; J, detail of fertile portion of spadix and appendix base × 2; K, habit × 2/3; L, inflorescence × 1; M, inflorescence × 1; N, detail of fertile portion of spadix and appendix base × 2; P, stamen, side view × 10; Q, stamen, view from below × 10; R, gynoecium × 10; S, gynoecium, longitudinal section × 10. Biarum pyrami var. pyrami: A, Davis 3844 (K); B–F, Mathew 11056 (Kew spirit collection 55694); B. tenuifolium subsp. tenuifolium: G, du Pavillon 189 (K); H, Davis 1138 (K); J, Archibald & Archibald 5904, Cult. Kew 1985–05490 (Kew spirit collection 49785); B. straussii: K, Hewer 1951 (K); B. davisii subsp. marmarisense: L, Baytop s.n. (Kew spirit collection 55936); B. ditschianum: M, Bonn accessions 22592 (Kew spirit collection 55925 & 59042); B. spruneri: N–S, Cult. Mathew (Kew spirit collection 51563). A R E A E : B I A R U M 267

96. Biarum to much longer than spathe, female zone free, short, hemi- Tribe Arisaemateae C spheric to subhemispheric, subglobose, separated from male zone by rather long axis bearing sterile flowers at base or Tribe Arisaemateae Nakai, Ord. Fam. Trib. Nov. 221 (1943, throughout or rarely naked, male zone 2–3 times longer than “Arisaematieae ”). female, rarely with a short zone of sterile flowers above, appen- dix smooth, either ± slender, elongate-vermiform and not or Laticifers simple, articulated; tuber depressed-globose, rarely hardly stipitate or thicker, cylindric to conoid and stipitate, rhizomatous; leaf blade usually compound, trisect, pedati- rarely procurved and filiform, rarely with a short, basal zone of sect or radiatisect, rarely cordate or trifid, primary lateral filiform processes. FLOWERS: unisexual, perigone absent. MALE veins of lobes or divisions pinnate, higher order venation FLOWER: 1–2-androus, stamens free, anthers sessile, connec- reticulate; spathe tube usually cylindric, blade usually for- tive slender, apiculate and slightly prominent or not, thecae nicate; spadix with terminal appendix, appendix often ellipsoid, with posterior microsporangia ± overtopping anterior cylindric, clavate or elongate-tapering; flowers unisexual, ones, dehiscing by pore or lateral slit extending to the middle perigone absent; ovary 1-locular, ovules orthotropous, pla- or nearly to the base and confluent apically. POLLEN: pre- centa basal, style ± attenuate, stigma small; seed testa rough, sented free or extruded in strands, grains inaperturate, spherical endosperm copious. to subspheroidal, medium-sized (mean 30 µm., range 26–56 µm.), exine spinose or smooth. STERILE FLOWERS: consisting 97. Pinellia of either a single filiform, acute or subulate-conic projection, or rarely composed of 2–3 uncinate, recurved projections from a Pinellia Tenore in Atti Accad. Sci. Napoli 4: 69 (1839), nom. single receptacle. FEMALE FLOWER: ovary ovoid or oblong- cons. TYPE: P. tuberifera Tenore, nom. illeg. (Arum subula- ovoid, 1-locular, ovule 1, orthotropous, funicle very short, tum Desfontaines, Cat. Pl. Horti Paris p. 385. 1829). placenta basal, stylar region very shortly conic to elongated, stigma subhemispheric. BERRY: obovoid, white to dull purple, SYNONYMS: Atherurus Blume, Rumphia 1: 135 (1837, pericarp carnose, not juicy, infructescence globose to hemi- “1835”); Hemicarpurus Nees, Delect. Sem. Horto Bot. Vratisl. spheric, borne at or slightly below soil level. SEED: obovoid 1839: [3] (1839) & in Linnaea 14 Litt.-Ber. 167 (1840). to subglobose, with obconic strophiole, testa smooth to irreg- ularly rugose, thickish, embryo axile, more than half as long HABIT: seasonally dormant herbs, tuber subglobose. LEAVES: as endosperm, endosperm copious. See Plates 96, 128C. few. PETIOLE: sometimes bearing tubercles near base and at CHROMOSOMES: 2n = 16, 20, 22, 24, 26, 32, 36, 74, 96, 98. apex, sheath fairly long. BLADE: usually deeply trifid or tri- DISTRIBUTION: 22 spp.; Mediterranean Europe, Middle East, sect or pedatisect, or simply cordate, ovate or oblong; primary North Africa:– Albania, Algeria, Croatia, Egypt, Greece (incl. lateral veins of each lobe or division pinnate, forming sub- Crete and islands), Iran, Iraq, Israel, Italy (incl. Sardinia, marginal collective vein, 1–2 distinct marginal veins also Sicily), Jordan, Lebanon, Libya, Macedonia, Morocco, present, higher order venation reticulate. INFLORESCENCE: Portugal, Spain, Syria, Tunisia, Turkey. solitary, appearing with leaves. PEDUNCLE: shorter or longer ECOLOGY: temperate and warm temperate scrub, wood- than petiole. SPATHE: slightly constricted between tube and land, open ground in fields, vineyards, pastures; geophytes, blade, tube convolute, narrowly ellipsoid to ovate, persistent, stony ground, under trees and shrubs, sandy areas, on lime- almost closed within by a transverse septum, gaping at base stone, rock crevices. (P. tripartita), blade oblong-elliptic, boat-shaped, gaping, for- NOTES: Boyce (in press 1) recognized 3 subgenera:– sub- nicate, green to purple. SPADIX: much longer than spathe, gen. Biarum; subgen. Cyllenium; subgen. Ischarum (with female zone adnate to spathe, separated from male zone by 3 sections). spathe septum and by short, free, naked part of spadix axis, ETYMOLOGY: Greek biaron, a name mentioned by Dodoens male zone cylindric, short, terminal sterile appendix smooth, (1574). elongate-subulate, often sigmoid, long-exserted from spathe. TAXONOMIC ACCOUNTS: Engler (1920a), Bogner & Boyce FLOWERS: unisexual, perigone absent. MALE FLOWER: 1- (1989), Boyce & Athanasiou (1991), Boyce (in prep.). androus, stamens sometimes united congenitally in pairs or groups of four, short, compressed laterally, anthers sessile, 268 T H E G E N E R A O F A R A C E A E

F H G B CD L E AK MN J Plate 97. Pinellia. A, habit × 2/3; B, stamen × 20; C, gynoecium × 20; D, gynoecium, longitudinal section × 20; E, petiole bulbil × 4; F, infructes- cence × 2/3; G, leaf × 2/3; H, infructescence × 2/3; J, habit × 2/3; K, inflorescence, nearside half of spathe removed × 3; L, stamen × 20; M, gynoecium × 20; N, gynoecium, longitudinal section × 20. Pinellia ternata: A–D, Kirkham, Flanagan & Boyce 146, (Kew spirit collection 57610); Wilson 4100 (K); E, Hickin s.n. (K); P. tripartita: F, Oldham 821 (K); P. pedatisecta: G, Wang x–333 (K); H, Cult. Kew 1962–24001 (Kew spirit collection 37614); P. cordata: J, Faber 82 (K); K–N, Cult. Boyce (Kew spirit collection 51852 & 58114). A R I S A E M A T E A E : P I N E L L I A 269

horizontal rhizome. LEAVES: 1–2, rarely 3, cataphylls often attractively mottled and spotted. PETIOLE: sheaths usually rather long and imbricate to form a long, usually mottled pseudostem, margins either free or fused from base to apex and then with a fringed, ligulate mouth. BLADE: usually compound, trisect, radiatisect, pedatisect, very rarely simple and ovate, lobes 3–19 or sometimes more, usually lanceo- late-elliptic, varying from linear to broadly ovate, elliptic or obovate, sometimes rhomboid, sessile or central leaflet often stalked, margin entire, serrate, erose or coarsely dentate; primary lateral veins of each lobe or division pinnate, form- ing submarginal collective vein, 1–2 conspicuous outer marginal veins also present, higher order venation reticulate. INFLORESCENCE: solitary, usually appearing with the leaves, sometimes before them, often subtended by conspicuous cataphylls, many species producing male and female inflo- rescences successively from the same plant at different seasons (paradioecy). PEDUNCLE: very short to longer than petiole, rarely tuberculate in upper portion (A. scortechinii), sometimes decurved in fruit. SPATHE: marcescent, usually unconstricted, rarely slightly constricted (e.g. A. flavum), lower part erect, convolute into cylindric, often longitudi- nally striped tube, tube mouth often with revolute to broadly 97. Pinellia or even grotesquely auriculate margins, blade usually strongly fornicate, sometimes erect, widely expanded to connective slender, thecae ellipsoid, dehiscing by apical slit. galeate, apex acute to long-acuminate, sometimes drawn POLLEN: extruded in amorphous mass, inaperturate, spheri- into very long filiform, erect or drooping thread. SPADIX: cal or subspheroidal, small to medium-sized (mean 25 µm., usually sessile, sometimes shortly stipitate, free, unisexual or range 21–29 µm.), exine spinulose. FEMALE FLOWER: ovary monoecious, female or monoecious spadices more robust ovoid to ovoid-oblong, 1-locular, ovule 1, orthotropous, funi- and differing from male in appendix shape and presence of cle very short, placenta basal, stylar region attenuate, stigma sterile organs, female zone densely flowered, usually conoid, small, hemispheric. BERRY: oblong-ovoid, green. SEED: male zone usually laxly flowered and contiguous with obnapiform to ellipsoid, testa irregularly verrucose-rugulose female in monoecious inflorescences, sterile terminal appen- or smooth, embryo axile, elongate or very small and subglo- dix erect, procurved or pendent, entirely hidden within bose, endosperm copious. See Plates 97, 128D. spathe to very long-exserted, usually somewhat longer than CHROMOSOMES: 2n = 26, 52. spathe tube, stipitate or not, cylindric, clavate, rounded, DISTRIBUTION: 6 spp.; temperate east Asia:– China (incl. rugose or apically echinate or drawn out into a sometimes Taiwan), Japan (incl. Ryukyu Is.), Korea N. and S.; P. ternata very long thread, rarely lacking altogether (A. exappendic- is occasionally naturalized in Europe and North America. ulatum), often with a few subulate to filiform projections in ECOLOGY: temperate woodland and forest; geophytes, on basal part, more rarely entirely composed of long filiform ground or on rocks, also on grassy banks and in fields as a projections. FLOWERS: unisexual, perigone absent. MALE weed (P. ternata). FLOWER: 2–5-androus, filaments connate forming synan- NOTES: Ulharz (1985) reports that P. tripartita is self-polli- dria, synandria ± distant from one another, sessile to nating, probably anemophilous. long-stipitate, connective slender, usually inconspicuous, ETYMOLOGY: named after Giovanni V. Pinelli (1535–1601). thecae shortly ovoid, dehiscing by short to long slit or pore, TAXONOMIC ACCOUNTS: Engler (1920a), Benzing (1969), Li sometimes confluent into ± lunate or even circular com- (1979), Ulharz (1985, 1986), Boyce (1988), Rugh (1990). pound thecae dehiscing by a single slit. POLLEN: very dry and powdery, inaperturate, spherical or subspheroidal, small (mean 22 µm., range 17–39 µm.), spinose. FEMALE FLOWER: C 98. Arisaema ovary 1-locular, ovoid or oblong-ovoid, ovules 3–10, orthotropous, erect, funicle short, placenta basal, stylar region Arisaema Martius in Flora 14: 459 (1831). TYPE: A. specio- short to attenuate, always narrower than ovary, stigma usu- sum (Wallich) Martius ex Schott (Arum speciosum Wallich), ally rather small, subhemispheric. BERRY: obovoid to lectotype selected by Pfeiffer, Nom. Bot. 1: 265. (1873). obconic, rounded apically, rarely conical, usually few-seeded, bright red, rarely yellow, glossy. SEED: ovoid to globose, SYNONYMS: Dochafa Schott, Syn. Aroid. 24 (1856); bearing a strophiole, testa hard, rough, light brown, embryo Muricauda J.K. Small, Fl. Southeast U.S. 227 (1903); axile, endosperm copious. See Plates 98i–iv, 129A. Flagellarisaema Nakai in J. Jap. Bot. 25: 5 (1950); Pleuriarum CHROMOSOMES: 2n = 20, 24, 26, 28, 39, 42, 48, 52, 56, 70, Nakai in J. Jap. Bot. 25: 5 (1950); Heteroarisaema Nakai in 72, 112, 140, 168 (64). J. Jap. Bot. 25: 6 (1950); Ringentiarum Nakai in J. Jap. Bot. DISTRIBUTION: ca. 170 spp.; temperate and warm temper- 25: 6 (1950). ate eastern and southeastern North America, northern HABIT: seasonally dormant or evergreen herbs, small to Mexico, tropical east and northeast Africa, Arabian Peninsula, fairly large, stem usually a depressed-globose tuber, pro- temperate east Asia, tropical south, southeast and east Asia, ducing tubercles or stolons, more rarely a branching, Malay Archipelago:– Afghanistan, Bhutan, Brunei, Burma, 270 T H E G E N E R A O F A R A C E A E

B A C ED Plate 98 (i). Arisaema. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2. Arisaema candidissimum: A, Cult. Kew 1986–3895 (Kew slide collection); A. grapsospadix: B, Cult. Kew 1987–2421 (Kew slide collection), Murata s.n. (Kew slide collection); A. consanguineum: C, Cult. Kew 1985–854 (Kew slide collection); A. heterophyllum: D, Cult. Kew 1987–2226 (Kew slide collection); A. abei: E, Cult. Kew 1987–3585 (Kew slide collection). A R I S A E M A T E A E : A R I S A E M A 271

C A B ED Plate 98 (ii). Arisaema. A, habit × 1/4; B, detail of inflorescence emergence from pseudostem ape× × 2/3; C, habit × 1/4; D, habit × 1/4; E, habit in flower × 1/4. Arisaema abei: A–B, Cult. Kew 1987–3585 (Kew slide collection); A. grapsospadix: C, Cult. Kew 1987–2421 (Kew slide collection); Murata s.n. (Kew slide collection); A. kiushianum: D, Murata s.n. (Kew slide collection); A. candidissimum: E, Cult. Kew 1986–3895 (Kew slide collection). 272 T H E G E N E R A O F A R A C E A E

C F E A H G B J D N QR LK M PS Plate 98 (iii). Arisaema. A, male inflorescence × 2/4; B, male spadix × 2; C, synandrium × 10; D, female spadix × 2; E, gynoecium × 10; F, gynoecium, longitudinal section × 10; G, male inflorescence × 2/3; H, male spadix, most of appendi× removed × 2; J, synandrium × 10; K, male inflorescence × 2/3; L, male spadix × 2; M, synandrium × 10; N, female inflorescence × 2/3; P, female spadix × 2; Q, gynoecium × 10; R, gynoecium, longitudinal section × 10; S, female inflorescence × 2/3. Arisaema nikoense var. nikoense: A–F, Cult. Kew 1987–2861 (Kew spirit collection 55951 & Kew slide collection); A. griffithii: G–J, Cult. Kew 1988–466 (Kew spirit collection 55695 & Kew slide collection); A. taiwanense: K–M, Cult. Kew 1987–3583 (Kew spirit collection 49912 & Kew slide collection); A. ringens: N–R, Cult. Kew 1987–3583 (Kew spirit collection 49912 & Kew slide collection); A. tortuosum: S, Cult. Kew 1984–4797 (Kew slide collection). A R I S A E M A T E A E : A R I S A E M A 273

C A D G M B EF H JK L Plate 98 (iv). Arisaema. A, male inflorescence × 2/3; B, male spadix × 1; C, stamen × 10; D, female spadix × 2; E, gynoecium × 10; F, gynoe- cium, longitudinal section × 10; G, monoecious inflorescence × 2/3; H, monoecious spadix × 2; J, stamen × 10; K, gynoecium × 10; L, gynoecium, longitudinal section × 10; M, female inflorescence × 2/3. Arisaema exappendiculatum: A–F, Bown 184/26 (Kew slide collection); B–F, Cult. Kew 1969–9202 (Kew spirit collection 49914); A. flavum subsp. flavum: G–L, Cult. Kew 1968–65 (Kew spirit collection 54924 & Kew slide collection); A. fimbriatum: M, Rubeli s.n. (Kew slide collection). 98. Arisaema 274 T H E G E N E R A O F A R A C E A E

Burundi, Cambodia, Canada, China (incl. Taiwan), Ethiopia, 99. Ariopsis India, Indonesia (Borneo, Flores, Java, Sulawesi, Sumatra, Timor), Japan, Kenya, N. and S. Korea, Laos, Malaysia (Borneo, Peninsula), Mexico, Nepal, Oman, Pakistan, Philippines, Russia (Far East), Rwanda, Saudi Arabia, Somalia, Sri Lanka, Sudan, Tanzania, Thailand, Uganda, USA, Vietnam, Yemen Republic, Zaïre. ECOLOGY: temperate, subtropical and upland tropical for- est, more rarely savanna, lowland tropical forest (Sarawak), subdesert or montane grassland (up to 4500m alt.); geo- phytes, forest floor, rocky slopes, rarely in wet places, very rarely epiphytic. NOTES: Murata (1990a) recognized 11 sections:– Fimbriata, Decipientia, Trisecta, Franchetiana, Pedatisecta, Clavata, Tortuosa, Dochafa, Tenuipistillata, Sinarisaema and Arisaema. ETYMOLOGY: Greek aris, aridos (name of a small herb mentioned by Pliny) and haima, haimatos (blood). TAXONOMIC ACCOUNTS: Engler (1920a), Hara (1971), Li (1979, 1980), Ohashi & Murata (1980), Sivadasan (1982), Murata (1984, 1987, 1990a), Mayo & Gilbert (1986), Pradhan (1990). dehiscing by oval pore, synandria all connate apically, form- C Tribe Colocasieae ing continuous surface punctured by cavities with somewhat prominent margins into which pollen is shed from the 6(–8) surrounding thecae (each pair of thecae belonging to a dif- Tribe Colocasieae Engler in Nova Acta Acad. Leopold.- ferent synandrium). POLLEN: inaperturate, spherical or Carol. 39: 149 (1876). subspheroidal, small (mean 20 µm.), exine spinose. FEMALE Laticifers anastomosing (except Ariopsis); primary lateral FLOWER: ovary ovoid to ovoid-oblong, 1-locular, ovules many, veins of leaf blade or leaflets (Protarum) pinnate, forming orthotropous, placentae 4–6, parietal, extending from base to submarginal collective vein, at least 1–2 other marginal veins apex of locule, stylar region absent, stigma stellate with 4–6- also present, higher order venation intermediate between laciniate lobes, lobes initially erect, later spreading and parallel-pinnate and reticulate; spathe usually constricted reflexed. BERRY: 4–6-angled, stigma persistent, many-seeded. centrally (except Ariopsis) with lower, convolute, persistent SEED: oblong, apically narrowed and obtuse, with indistinct tube and upper, gaping, ± boat-shaped, marcescent to strophiole, testa thickish, longitudinally costate, embryo axile, caducous blade; flowers unisexual, perigone absent; male small, endosperm copious. See Plates 99, 129B. flower a synandrium of several connate stamens, common CHROMOSOMES: 2n = 28, 84. connectives thick, fleshy, thecae lateral, pollen inaperturate; DISTRIBUTION: 2 spp.; tropical south Asia:– Bhutan, Burma, endosperm copious. India (Assam, Sikkim, Western Ghats), Nepal. ECOLOGY: tropical evergreen forest; geophytes on forest floor or in rock crevices. C 99. Ariopsis NOTES: The correct morphological interpretation of the male flowers remains unclear. Ariopsis Nimmo in J. Graham, Cat. Pl. Bombay 252 (1839). ETYMOLOGY: Greek aron (Arum) and -opsis (appearance). TYPE: A. peltata Nimmo TAXONOMIC ACCOUNTS: Engler (1920a), Nicolson (1976), Sivadasan (1982), Bogner & Boyce (in prep.). Laticifers simple, articulated. HABIT: small, seasonally dor- mant herbs, tuber ± subglobose, hypogeal. LEAVES: usually solitary, rarely few. PETIOLE: slender, sheath fairly short. 100. Protarum C BLADE: peltate, cordate-ovate or only emarginate basally, thin, glaucous below, posterior lobes very short; primary lat- Protarum Engler in Bot. Jahrb. 30, Beibl. 67: 42 (1901). eral veins pinnate and also radiating from petiole insertion, TYPE: P. sechellarum Engler forming submarginal collective vein, marginal vein also pre- sent, higher order venation reticulate. INFLORESCENCE: 1–3 HABIT: seasonally (?irregularly) dormant herb, tuber subglo- in each floral sympodium, appearing with or without leaves. bose, with numerous, very densely arranged, annular leaf PEDUNCLE: very slender, much longer than spathe, erect. scars. LEAF: solitary. PETIOLE: long, sheath short. BLADE: SPATHE: ovate, boat-shaped, fornicate, not constricted, gap- pedatisect to radiatisect, almost radiate, leaflets shortly stalked, ing widely, not convolute at base, marcescent. SPADIX: elliptic to narrowly elliptic, acuminate; primary lateral veins of shorter than spathe, female zone adnate to spathe, very short lobes pinnate, numerous, forming submarginal collective vein, and few-flowered, sometimes separated from male zone by 2 marginal veins also present, higher order venation reticulate. short, free, naked axis, male zone fertile to apex, relatively INFLORESCENCE: solitary, appearing with or without leaves. thick, cylindric-conoid, many-flowered. FLOWERS: unisexual, PEDUNCLE: short, much shorter than petiole. SPATHE: slightly perigone absent. MALE FLOWER: synandrium peltate, con- constricted between tube and blade, tube convolute, shorter nate filaments forming a stipe longer and narrower than than blade, blade erect, boat-shaped, marcescent. SPADIX: dilated common connective, thecae subglobose to ellipsoid, sessile, much shorter than spathe, female zone conoid, sep- C O L O C A S I E A E : P R O T A R U M 275

LK F D H C E AB GJ Plate 99. Ariopsis. A, habit × 1; B, spadix × 6; C, gynoecium × 15; D, gynoecium, longitudinal section × 15; E, section through male por- tion of spadix to show synandrium arrangement × 10; F, infructescence × 6; G, habit in flower, showing branching of tubers × 1; H, leaf × 1; J, spadix × 6; K, berry × 6; L, seed × 12. Ariopsis peltata: A, Talbot 496 (K); B–E, Bogner 1922 (Kew spirit collection 56425); F, Barnes 1087 (K); A. protanthera: G–H, Cult. Kew.1851 (K); J, Kurz s.n. (K); K–L, Cult. Kew. 1851 (Kew spirit collection 58040). 276 T H E G E N E R A O F A R A C E A E

K AJ B HG C E FD Plate 100. Protarum. A, leaf × 1/6; B, leaflet × 1/3; C, tuber × 1/2; D, inflorescence × 2/3; E, spadix × 2; F, detail of gynoecia, top view × 8; G, synandrium, top view × 15; H, gynoecium with associated staminode, longitudinal section × 15; J, infructescence, nearside half of spathe removed × 2/3; K, seed × 4. Protarum sechellarum: A, 5495 (Kew slide collection); B, Milne 14 (K); C, Jeffrey 485 (Kew spirit collection 25047); D, Thomasset s.n. (K); E–H, Whitehead 35 (Kew spirit collection 37323); J–K, Gardiner 110 (K & Kew spirit collection 58062). C O L O C A S I E A E : P R O T A R U M 277

100. Protarum 101. Steudnera arated from male zone by more slender sterile zone covered phylls. LEAVES: solitary or few to several. PETIOLE: sheath with narrowly elongated synandrodes or partly naked, male very short. BLADE: peltate, ovate or ovate-oblong, acumi- zone ± cylindric to obconic, terminal sterile appendix thicker, nate, often emarginate at base; basal ribs short, digitiform, obtuse, ± smooth. FLOWERS: unisexual, perigone well-developed, primary lateral veins pinnate, forming sub- absent. MALE FLOWER: synandrium 3–6-androus, sessile, marginal collective vein very near margin, marginal vein subprismatic, ± excavated centrally, fused connectives broad, also present, secondary and tertiary laterals arising from the margins lobed, thecae broadly ellipsoid to subglobose, primaries at a wide angle, then arching towards leaf margin dehiscing by broad apical slit. POLLEN: inaperturate, ellip- and forming ± conspicuous interprimary collecting vein, soid, small (21 µm., range 20–22 µm.), exine striate. FEMALE higher order venation reticulate. INFLORESCENCE: solitary. FLOWER: gynoecium surrounded by whorl of 4–6, free, ± PEDUNCLE: shorter than petiole. SPATHE: yellow or ± dark imbricate, thickish, oblong, subprismatic staminodes, ovary purple within, ovate or ovate-lanceolate, acuminate, not ovoid to ellipsoid, 1-locular, ovules 4, hemiorthotropous, constricted, barely convolute at base, ± fully expanded at micropyle elongate, funicle short, placenta basal, stylar region anthesis, lower part persistent to fruiting stage, upper part shortly attenuate or nearly absent, stigma 2–4-lobed, much becoming reflexed and revolute, marcescent. SPADIX: much broader than style. BERRY: ± ellipsoid to obovoid, with per- shorter than spathe, densely flowered, female zone cylindric, sistent stigma remnant, usually 1-seeded, orange, often longer than male, mostly adnate to spathe, male zone infructescence enclosed by persistent and much enlarged contiguous with female, cylindric, ellipsoid or subglobose, spathe tube. SEED: ovoid to ellipsoid-oblong, testa costate, fertile to apex, obtuse. FLOWERS: unisexual, perigone dark brown, embryo small, flattened and disc-like, absent. MALE FLOWER: 3–6-androus, stamens connate, endosperm copious. See Plate 100. synandrium strongly lobed, apically truncate, common con- CHROMOSOMES: 2n = 28. nective relatively small, impressed at apex, thecae oblong, DISTRIBUTION: 1 sp.; Seychelles Islands (Mahé, Praslin, contiguous, dehiscing by apical pore. POLLEN: inapertu- Silhouette). rate, ellipsoid-oblong, small (mean 22 µm.), exine striate. ECOLOGY: tropical (palm) forest, moss forest; geophytes, for- FEMALE FLOWER: gynoecium surrounded by whorl of 2–5, est floor, leaf litter in rock crevices, often on large granitic short, claviform staminodes, more rarely staminodes absent, rocks. ovary subglobose to ovoid, 1-locular, ovules numerous, ETYMOLOGY: Greek protos (first) and Arum. hemiorthotropous, funicle distinct, placentae 2–5, parietal, in TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1973b), basal part only or extending from base to apex, stylar region Robertson (1989). ± lacking, stigma strongly 2–5-lobed. BERRY: ovoid, many- seeded. SEED: ovoid to ellipsoid, testa costate, embryo axile, short, conoid, endosperm copious. See Plates 101, 129C. C 101. Steudnera CHROMOSOMES: 2n = 28, 56. DISTRIBUTION: 8 spp.; tropical south and southeast Asia:– Steudnera K. Koch in Wochenschr. Gärtnerei Pflanzenk. 5: Bangladesh, Burma, ?Cambodia, China (Guangxi, Yunnan), 114 (1862). TYPE: S. colocasiifolia K. Koch (“colocasiaefolia”). India (Assam), Laos, Thailand, Vietnam. ECOLOGY: tropical humid forest; geophytes on forest floor. HABIT: medium sized to robust herbs, usually evergreen, ETYMOLOGY: named after H. Steudner (1832–1863). sometimes seasonally dormant, stem epigeal, stout, erect or TAXONOMIC ACCOUNTS: Krause in Engler & Krause (1920), creeping, covered with fibrous remains of leaves and cata- Boyce (1995c). 278 T H E G E N E R A O F A R A C E A E

C D E H JG B FA Plate 101. Steudnera. A, habit × 2/3; B, spadix × 2; C, synandrium, top view × 15; D, synandrium, side view × 15; E, gynoecium, top view × 15; F, gynoecium, longitudinal section × 15; G, inflorescence, nearside half of spathe removed × 2/3; H, synandrium, side view × 15; J, gynoecium with associated staminode × 15. Steudnera henryana: A, Henry 11986 (K); B–F, S. discolor: Bogner 1814 (Kew spirit collection 57576); S. colocasioides: G–J, Cult. Kew (K & Kew spirit collection 58060). C O L O C A S I E A E : S T E U D N E R A 279

C 102. Remusatia cal pore-like slit. POLLEN: extruded in strands, inaperturate, spherical to subspheroidal, medium-sized (mean 32–33 µm.), Remusatia Schott in Schott & Endlicher, Melet. Bot. 18 spinose. STERILE MALE FLOWERS: each a ± elongated synan- (1832). TYPE: R. vivipara (Roxburgh) Schott (Arum vivipa- drode. FEMALE FLOWER: staminodes absent, ovary rum Roxburgh, “viviparium”). subcylindric to subglobose, 1-locular or partially 2–4-locular at apex, ovules many, hemiorthotropous, funicle short to SYNONYM: Gonatanthus Klotzsch in Link, Klotzsch & long, placentae 2–4 and parietal or placenta 1 and basal, sty- Otto, Icon. Pl. Rar. Hort. Berol. 1: 33 (1841). TYPE: G. sar- lar region very shortly attenuate or lacking, stigma mentosus Klotzsch discoid-subcapitate or slightly 3–4-lobed. BERRY: obovoid HABIT: small to medium-sized, seasonally dormant herbs, to globose, many-seeded, infructescence ellipsoid, borne tuber subglobose, producing erect to spreading, within persistent spathe tube. SEED: ellipsoid to subglobose, unbranched or branching stolons from axils of scarious, covered by thick, succulent sarcotesta or testa verruculose to deciduous cataphylls, stolons producing small, ovoid tuber- irregularly costate, embryo axile, short, ovoid to subglobose, cles at nodes, each invested by numerous, apically revolute endosperm copious. See Plates 102, 129D. or flexuose scales (minute cataphylls). LEAVES: 1–2. PETI- CHROMOSOMES: 2n = 28, 42, 56. OLE: sometimes slender, sheath relatively short. BLADE: DISTRIBUTION: 4 spp.; tropical Africa, tropical Asia, Malay peltate, cordate-lanceolate to cordate-ovate, acuminate; Archipelago, Australasia: Australia (N.), Bangladesh, Bhutan, basal ribs well-developed, primary lateral veins pinnate, Cameroon, China (Taiwan, Yunnan, Tibet), Christmas Is., forming submarginal collective vein very close to margin, Ethiopia, Guinea, India, Indonesia (Java), Ivory Coast, Liberia, marginal vein also present, secondary and tertiary laterals Madagascar, Nepal, Nigeria, Oman, ?Saudi Arabia, Sierra arising from the primaries at a wide angle, then arching Leone, Sri Lanka, Tanzania, Thailand, Vietnam, Yemen towards leaf margin and forming inconspicuous interpri- Republic (Socotra), Zaïre, Zambia. mary collective veins, higher order venation reticulate. ECOLOGY: tropical seasonal forest; epiphytes, lithophytes or INFLORESCENCE: solitary, appearing with or without leaf. geophytes, moss-laden boughs, rock crevices, damp banks, PEDUNCLE: shorter than petiole. SPATHE: strongly con- forest floor. stricted between tube and blade, sometimes with secondary NOTES: Li & Hay (1992a, b) recognize 2 sections, sect. constriction above the spadix, tube with convolute margins, Remusatia and sect. Gonatanthus. persistent, enclosing female zone and sterile zone of spadix, ETYMOLOGY: named after A. Rémusat (1788–1832). blade yellow or red, longer than tube, fully expanded or TAXONOMIC ACCOUNTS: Krause in Engler & Krause (1920), remaining convolute and opening only at base, sometimes Sivadasan (1982), Mayo (1985a), Li (1987a, b, 1991, 1992), Li becoming reflexed (R. vivipara, R. yunnanensis), later decid- & Hay (1992a, 1992b). uous. SPADIX: sessile or subsessile, much shorter than spathe, female zone subcylindric, about half as long as 103. Colocasia C spathe tube, separated from male zone by much narrower zone of sterile male flowers, male zone ellipsoid or sub- clavate, fertile to apex, obtuse. FLOWERS: unisexual, Colocasia Schott in Schott & Endlicher, Melet. Bot. 18 (1832), perigone absent. MALE FLOWER: 2–3-androus, stamens con- nom. cons. TYPE: C. antiquorum Schott (Arum colocasia nate into cuneate-clavate, 4–6-sulcate synandrium, fused L.), typ. cons. filaments distinct, common connectives somewhat excavated SYNONYM: Leucocasia Schott in Oesterr. bot. Wochenbl. at apex, thecae 4–6, oblong to ellipsoid, dehiscing by api- 7: 34 (1857). 102. Remusatia 280 T H E G E N E R A O F A R A C E A E

E D A B H J M C G K FP N L Q Plate 102. Remusatia. A, habit in flower with stolons × 1/4; B, habit with stolons × 1/4, C, leaf × 2/3; D, tubercle × 6; E, plantlet emerging from tubercle × 2/3; F, inflorescence × 2/3; G, spadix × 2; H, synandrium, top view × 12; J, synandrium, side view × 12; K, synandrode × 15; L, gynoecium, longitudinal section × 12; M, gynoecium, transverse section × 12; N, habit × 1/3; P, inflorescence × 2/3; Q, gynoecium, longitudinal section × 12. Remusatia vivipara: A, 299 (Kew slide collection); B, 300 (Kew slide collection); C–D, Kerr 1438 (K); E, Cult. Kew 1969–5188 (Kew spirit collection 59039); F–M, Bogner 2121 (Kew spirit collection 57570); Cult. Kew 1947–85 (Kew spirit collection 19155); R. pumila: N, Grey-Wilson & Phillips 128 (K); P, C.K. 227 (K); Q, Grierson & Long 2215 (K). C O L O C A S I E A E : R E M U S A T I A 281

C B D G K E J H F AL Plate 103. Colocasia. A, habit × 1/2; B, inflorescence × 2/3; C, synandria, top view × 5; D, gynoecium, longitudinal section × 11; E, floral sympodium with associated petiole sheath × 1/3; F, spadix × 1; G, synandria, top view × 5; H, synandrium, side view × 10; J, gynoecium, longitudinal section; K, gynoecium, transverse section; L, habit × 1/5. Colocasia fallax: A, Gamble 27041 (K); B–D Chantaranothai et al. 90/464 (Kew spirit collection 59076); C. gigantea: E, Bown 197/26 (Kew slide collection); F–K, Bogner 427 (Kew spirit collection 34491); C. escu- lenta: L, Townsend s.n. (Kew slide collection). 282 T H E G E N E R A O F A R A C E A E

HABIT: small, medium-sized or gigantic, seasonally dormant testa thickish, costate, embryo axile, cylindric, endosperm C or evergreen herbs, stem either a hypogeal, subglobose or copious. See Plates 103, 130A. subcylindric tuber or mostly epigeal, massive. LEAVES: sev- CHROMOSOMES: 2n = 28, 42, 56. eral, rosulate in acaulescent plants, forming terminal crown DISTRIBUTION: 8 spp.; tropical Asia, Malay Archipelago:– in arborescent species. PETIOLE: sheath rather long. BLADE: Bangladesh, Bhutan, Burma, Cambodia, China (Guandong, peltate, ovate-cordate to sagittate-cordate, posterior lobes Guangxi, Guizhou, Hainan, Hunan, Sichuan, Taiwan, rounded, shortly to almost entirely connate; basal ribs well- Yunnan), India, Indonesia (Java, Lesser Sunda Is.), Laos, developed, primary lateral veins pinnate, forming Malaysia (Peninsula), Nepal, Sri Lanka, Thailand, Vietnam. submarginal collective vein, 1–2 marginal veins also pre- The country list given here does not include C. esculenta. sent, secondary and tertiary laterals arising from the This species is cultivated and naturalized throughout the primaries at a wide angle, then arching strongly towards humid tropics and subtropics. Its natural area of distribution leaf margin and forming inconspicuous interprimary collec- remains somewhat obscure but is certainly tropical Asian, tive veins, higher order venation reticulate. extending into the Malay Archipelago and perhaps as far as INFLORESCENCE: 1–many in each floral sympodium, Papuasia and Australia. appearing with the leaves. PEDUNCLE: much shorter than ECOLOGY: tropical humid forest habitats; geophytes, ter- petiole. SPATHE: constricted between tube and blade, some- restrial or helophytes, wet places, along streams and ponds, times with a second constriction above male zone of spadix, forest floor in leaf litter, between rocks, sometimes on lime- tube with convolute margins, usually much shorter than stone (C. gigantea). blade, ovoid or oblong, persistent, enlarging in fruit and NOTES: Krause in Engler & Krause (1920) recognized 2 sec- then splitting open irregularly, blade white to yellow, oblong tions:– sect. Colocasia (“Tuberosae”), sect. Caulescentes. and boat-shaped to narrowly lanceolate, reflexing at anthe- VERNACULAR NAMES AND USES: taro, eddoe; C. esculenta sis, later deciduous. SPADIX: sessile, shorter than spathe, is an important food plant in tropical regions, especially the female zone short, separated from male by narrower zone Pacific. of sterile male flowers, male zone cylindric to fusiform, ter- ETYMOLOGY: classical name, Greek kolokasia, from an old minal appendix erect, ± smooth, elongate-conoid to fusiform Middle Eastern name qolqas. or subulate, sometimes reduced to small, mucronate stub, TAXONOMIC ACCOUNTS: Krause in Engler & Krause (1920); rarely absent. FLOWERS: unisexual, perigone absent. MALE Sivadasan (1982), Plucknett (1983), Shaw (1984), Sreekumari FLOWER: 3–6-androus, stamens connate into ± truncate & Mathew (1991a, b), Li & Wei (1993). synandrium, thecae lateral, oblong-linear, dehiscing by api- cal pore. POLLEN: extruded in strands, inaperturate, 104. Alocasia ellipsoid-oblong or spherical to subspheroidal (C. esculenta), small to medium-sized (mean 25 µm., range 25–26 µm), Alocasia (Schott) G. Don in Sweet, Hort. Brit., ed. 3: 631 exine finely striate, coarsely swirling-fossulate or fossulate- (1839), nom. cons. TYPE: A. cucullata (Loureiro) G. Don spinose. STERILE MALE FLOWERS: synandrodes (Arum cucullatum Loureiro), typ. cons. depressed-obpyramidal, truncate, laterally compressed. FEMALE FLOWER: ovary ovoid or oblong, 1-locular, ovules SYNONYMS: Colocasia sect. Alocasia Schott in Schott & many, hemiorthotropous, funicles relatively long, placentae Endlicher, Melet. Bot. 18 (1832); Ensolenanthe Schott in 2–5, parietal, stylar region short, narrowed or not, sometimes Bonplandia 9: 368 (1861); Xenophya Schott in Ann. Mus. ± absent, stigma discoid-capitate or weakly lobed. BERRY: Bot. Lugduno–Batavum 1: 124 (1863); Schizocasia Schott ex greenish to whitish or red(?), obconoid or oblong, stigma Engler in Bot. Jahrb. 1: 185 (1880, “1881”); Panzhuyuia Z.Y. remnants persistent, many-seeded. SEED: ovoid to ellipsoid, Zhu in J. Sichuan Chinese Medicinal School 4 (5): 49 (1985). 103. Colocasia 104. Alocasia C O L O C A S I E A E : A L O C A S I A 283

C A F D B E K J G M HL Plate 104 (i). Alocasia. A, leaf × 1/3; B, inflorescence × 1; C, synandrium, side view × 8; D, gynoecium, longitudinal section × 8; E, leaf × 1/3; F, habit × 1/6; G, infructescence with associated petiole and plant base × 1/8; H, spadix × 2; J, synandrium × 10; K, gynoecium × 10; L, gynoecium, longitudinal section × 10; M, habit × 2/3. Alocasia brancifolia: A, Cult. Veitch 10 June 1884 (K); B–D, Bogner 1736 (Kew spirit collection 58916); A. guttata: E, Forman 931 (K); A. lowii: F–G, Boyce 384 (Kew slide collection); H–L, Boyce 384 (Kew spirit collection 49923); A. beccarii : M, Simpson 2271 (K); Wood 798 (Kew spirit collection 52879). 284 T H E G E N E R A O F A R A C E A E

D EC F B A Plate 104 (ii). Alocasia. A, habit × 2/3; B, spadix × 2; C, synandrium, side view × 15; D, gynoecium × 15; E, gynoecium, longitudinal sec- tion × 15; F, infructescence. Alocasia beccarii: Alston 14176 (BM). HABIT: medium-sized to rarely arborescent and gigantic, towards leaf margin, sometimes forming interprimary veins, seasonally dormant to evergreen herbs, stem thick, often higher order venation reticulate. INFLORESCENCE: 2–many hypogeal, sometimes stoloniferous and bulbiferous, epigeal in each floral sympodium, appearing with the leaves. stem usually erect, rarely elongated and creeping, with PEDUNCLE: usually shorter than petiole. SPATHE: strongly clear to milky latex. LEAVES: few to several in terminal constricted between tube and blade, tube with convolute crown, sometimes each subtended by a cataphyll. PETI- margins, shorter than blade, ovoid or oblong, persistent OLE: long, sometimes asperate or glandular, sheath and then splitting irregularly in fruit, blade oblong, usually relatively long, sometimes deciduous. BLADE: sometimes boat-shaped, rarely fornicate, at anthesis at first erect, then pubescent, juvenile blade peltate, at maturity usually sagit- reflexing and later usually deciduous; in Xenophya group tate, less often ± hastate or cordate, peltate in some species, blade is persistent, erect, convolute, gaping only basally. margin entire, sinuate or slightly to deeply pinnatifid, pos- SPADIX: sessile, sometimes shortly stipitate, rarely obliquely terior divisions ovate or triangular; basal ribs adnate, shorter than spathe, female zone short, conoid- well-developed, glands present in axils of primary lateral cylindric, separated from male by a much narrower zone of veins and midrib, primary lateral veins pinnate, forming sterile flowers, male zone usually cylindric, appendix submarginal collective vein,1–2 closely adjacent marginal conoid to cylindric, surface with irregular, labyrinthine net- veins also present, secondary and tertiary lateral veins aris- work of fissures. FLOWERS: unisexual, perigone absent. ing from the primaries at a wide angle, then arching strongly MALE FLOWER: 3–12(–36)-androus, stamens connate into C O L O C A S I E A E : A L O C A S I A 285

an obpyramidal, subhexagonal, truncate, rarely linear (A. 105. Pistia C brisbanensis) synandrium, thecae oblong to linear-oblong, lateral, dehiscing by apical pore. POLLEN: extruded in Pistia L., Sp. Pl. 963 (1753). TYPE: P. stratiotes L. strands, inaperturate, spherical to subspheroidal, medium- SYNONYMS: Kodda-Pail Adanson, Fam. 2: 75, 541 sized (mean 35 µm., range 31–39 µm.), exine spinose. STERILE MALE FLOWERS: synandrodes shallow, obpyra- (1763); Zala Loureiro, Fl. Cochinch. 401, 405 (1790); midal, compressed, truncate. FEMALE FLOWER: ovary ovoid Apiospermum Klotzsch in Abh. Königl. Akad. Wiss. Berlin or oblong, 1-locular or partially 3–4-locular at apex, ovules 1852: 351 (1853); Limnonesis Klotzsch in Abh. Königl. Akad. 6–10, orthotropous, hemiorthotropous, hemianatropous or Wiss. Berlin 1852: 352 (1853). anatropous, funicle short, placenta basal, stylar region short, Laticifers absent. HABIT: small, free-floating evergreen herb stigma depressed-capitate, ± distinctly 3–4-lobed. BERRY: with pendent feathery roots, stem very short, acaulescent, generally reddish, ellipsoid or obconic-ellipsoid or subglo- stoloniferous. LEAVES: several in a rosette, densely pubes- bose, 1–5-seeded, stigma remnants persistent. SEED: cent. PETIOLE: very short, almost absent, sheath very short, subglobose to ellipsoid, testa thickish, smooth or scabrous, ligulate, very thin, scarious at base. BLADE: somewhat embryo broadly conoid, shortly cylindric or elongate, spongy, obovate-cuneate to obovate-oblong, apically endosperm copious. See Plates 104i–ii, 130B. rounded, truncate or retuse; midrib absent, primary veins CHROMOSOMES: 2n = 28, 42, 56, 70, 84. subparallel, all arising from base, diverging slightly and DISTRIBUTION: 60–70 spp.; tropical Asia, Australasia, Malay running into margin near apex, strongly prominent on Archipelago, Melanesia:– Australia (Queensland, New South lower surface, higher order venation reticulate. INFLO- Wales), Bangladesh, Brunei, Burma, Cambodia, China (incl. RESCENCE: solitary, very small, much shorter than leaves. Taiwan), Fiji, India, Indonesia (Borneo, Irian Jaya, Java, PEDUNCLE: very short, pubescent. SPATHE: pubescent Moluccas, Sulawesi, Sumatra), Japan (incl. Ryukyu Is.), Laos, without, glabrous within, somewhat constricted centrally, Malaysia (Borneo, Peninsula), Nepal, Papua New Guinea, lower margins connate with each other and with ovary Philippines, Solomon Is., Sri Lanka, Thailand, Vietnam. A. wall forming tube, free margins between tube and blade macrorrhizos, A. plumbea and A. cucullata are cultivated folded between stigma and male organs forming a partition and naturalized throughout the tropics. between an upper male partial chamber and a lower female ECOLOGY: tropical and subtropical humid forest; geophytes one, blade erect, ovate, expanded, acute-acuminate. or terrestrial, forest floor, in leaf litter, humus deposits on SPADIX: shorter than spathe, mostly adnate to spathe, only rocks, usually in deep shade, sometimes in exposed areas of the apical male zone free, female zone bearing single forest regrowth. gynoecium at base and a thin, green, pouch-shaped flap ETYMOLOGY: Greek a- (not) and Colocasia, the latter with- just below spathe partition, male zone subtended by a thin, out its first syllable, implying a genus close to but different marginally lobed, green, annular flap, composed of a from Colocasia. basally naked spadix axis supporting a single whorl of 2–8 TAXONOMIC ACCOUNTS: Krause & Engler in Engler & flowers, naked axis sometimes extending a little beyond. Krause (1920), Nicolson (1968a), Sivadasan (1982), Burnett FLOWERS: unisexual, perigone absent. MALE FLOWER: (1985), Hay (1990a), Hay & Wise (1991). synandrium composed of 2 connate stamens, thecae de- hiscing by single apical slit. POLLEN: inaperturate, ellipsoid-elongate to -oblong, medium-sized (mean 27 C Tribe Pistieae µm.), exine plicate with strongly undulate frills running the length of the grain. FEMALE FLOWER: gynoecium Tribe Pistieae Blume, Rumphia 1: 76 (1836, “Pistiaceae”). obliquely adnate to spadix axis, ovary ovoid, 1-locular, ovules numerous, orthotropous, placenta broad, appar- Laticifers absent; plant small, acaulescent; leaves several, ently parietal, probably morphologically basal, stylar region rosulate, densely pubescent; petiole very short; blade obo- attenuate, bending inwards towards male flowers, stigma vate-cuneate to oblong, apically rounded to retuse, midrib small, discoid-subcapitate. BERRY: thin-walled, utricular, absent, primary veins subparallel, higher order venation several-seeded, ellipsoid, irregularly breaking up and reticulate; inflorescence 1, very small; spathe ± constricted, decaying to release seeds. SEED: barrel-shaped, ± sub- lower margins connate forming tube, free margins between truncate and excavated at apex and base, testa thick, tube and blade folded to form a partition separating upper reticulate-alveolate, thicker and with operculum at micropy- male partial chamber from lower female one, blade gaping; lar end, embryo obovoid to conoid, endosperm copious. spadix:– female zone with single basal gynoecium and thin See Plates 105, 130C. pouch-shaped flap below spathe partition, male zone with CHROMOSOMES: 2n = 28. 2–8 flowers in a whorl, subtended by thin annular flap, DISTRIBUTION: 1 sp.; pantropical:– Afghanistan, Angola, spadix apex ± prominulent, naked, very short; flowers uni- Argentina, Australia, Bangladesh, Belize, ?Benin, Bolivia, sexual, perigone absent; male flower a sessile synandrium Botswana, Brazil, Brunei, Burkina Faso, Burundi, Cambodia, of 2 connate stamens, thecae dehiscing by single apical slit, Cameroon, Central African Republic, Chad, China (incl. pollen exine plicate with strongly undulate frills running Taiwan), Colombia, Comores Is., Congo, Costa Rica, Cuba, length of grain; ovary obliquely adnate to spadix axis, 1- Dominican Republic, Ecuador, Egypt, Equatorial Guinea locular, ovules numerous, orthotropous, placenta parietal (Rio Muni), Ethiopia, Gambia, Ghana, Guatemala, Guinea, (morphologically basal), style distinct, curved, stigma small; Guyana, Haiti, Honduras, India, Indonesia (Borneo, Irian berry thin-walled, several-seeded, pericarp eventually Jaya, Java, Moluccas), Ivory Coast, Jamaica, Kenya, Laos, decaying; seed subcylindric, testa thick, reticulate-alveolate, Lesotho, Lesser Antilles, Liberia, Madagascar, Malawi, with micropylar operculum formed by both integuments, Malaysia (Borneo, Peninsula), Mali, Mauritius, Mexico, embryo minute, endosperm copious. Mozambique, Nepal, Nicaragua, Niger, Nigeria, Pakistan, 286 T H E G E N E R A O F A R A C E A E

A ED CB Plate 105. Pistia. A, habit in flower with stolons × 2/3; B, inflorescence, front view × 6; C, inflorescence, side view, longitudinal section × 6; D, fruit × 4; E, seed × 8. Pistia stratiotes: A, FTEA Araceae 67, Fig. 17 (1985); B–E, Giles & Woolliams s.n., Cult. Kew 1963–41001 (Kew spirit collection 29047.737). P I S T I E A E : P I S T I A 287

105. Pistia Panama, Papua New Guinea, Paraguay, Peru, Philippines, tion of the structure of the male flowers, which is sup- Puerto Rico, Senegal, Sierra Leone, Somalia, South Africa, Sri ported by new developmental and anatomical studies Lanka, Sudan, Surinam, Swaziland, Tanzania, Thailand, (French 1986a, Buzgó 1994). Togo, Trinidad, Uganda, USA, Uruguay, Venezuela, Vietnam, ETYMOLOGY: Greek pister (hollow, trough, in the sense of Zaïre, Zambia, Zimbabwe. a drinking trough). ECOLOGY: tropical wetlands; floating aquatic in open, tran- TAXONOMIC ACCOUNTS: Engler (1920a), Bogner (1975), quil, freshwater habitats. Nicolson (1979), Sivadasan (1982), Mayo (1985a, 1986a), NOTE: We have followed Engler (1920a) in our interpreta- Hay (1990a). 288 T H E G E N E R A O F A R A C E A E

C 27 D E S C R I P T I O N O F A C O R A C E A E Family Acoraceae ing by longitudinal slit, connective inconspicuous. POLLEN: monosulcate to subulcerate, ellipsoid, small (mean 18 µm; Family Acoraceae C. Agardh, Aphor. Bot. 133 (1822, range 15–20 µm), exine shallowly and remotely or more “Acoroideae ”). densely foveolate, otherwise psilate, apertural exine subpsi- One genus, distributed throughout the northern hemisphere, late. GYNOECIUM: obconic-cylindric, slightly exceeding temperate to tropical latitudes. tepals, 2–3-locular, ovules several per locule, pendent, orthotropous, both integuments bearing trichomes, inner integument longer than outer and forming micropyle, pla- C Acorus centa apical, stigma minute, sessile. BERRY: oblong-obovoid with thin, leathery pericarp, enclosed by tepals, 1–5(–9)- Acorus L., Sp. Pl. 324 (1753). TYPE: A. calamus L. seeded, whitish with brownish stigma remnant when fresh, soon drying to straw-brown. SEED: oblong to ellipsoid, Laticifers absent, raphides absent. HABIT: herbs, stem a testa light brown, foveolate (A. calamus) or smooth (A. repent, much-branched, lacunose rhizome with aromatic oil gramineus), long integumentary trichomes present (A. cells. LEAVES: distichous, unifacial, ensiform, not dif- gramineus) or absent, embryo axile, cylindric or conoid ferentiated into petiole and blade, intravaginal squamules (A. gramineus), perisperm present, endosperm copious. present in leaf axils; primary veins parallel, higher order See Plates 106, 130D. venation parallel. INFLORESCENCE: solitary, terminal, borne CHROMOSOMES: 2n = 22, 24, 36, 44, 48. laterally on leaf-like scape, continuation shoot arising in axil DISTRIBUTION: 2(–4) spp.; temperate to subtropical Asia of leaf preceding spathe. SPATHE: much longer than spadix, and North America; introduced and naturalized in Europe. erect, persistent, appearing merely as a vertical extension of ECOLOGY: temperate to tropical wetlands, up to 1100m alt. the leaf-like peduncle. SPADIX: jutting out at an angle from in central Europe and up to 2600m alt. in China; helophyte, peduncle, conoid and finger-like or slender and tail-like. marshes, streams, ponds, swampy sites, pastures, meadows. FLOWERS: bisexual, perigoniate, densely arranged, bract- ETYMOLOGY: akoron, an ancient Greek plant name. less, 3-merous; tepals 6, in 2 whorls of 3, thin, fornicate. TAXONOMIC ACCOUNTS: Engler (1905), Huttleston (1953), STAMENS: 6, in 2 whorls of 3, free, filaments linear-oblong, Röst (1978, 1979a, b), Röst & Bos (1979), Riedl (1977–1979), anthers introrse, thecae rounded-elliptic, subopposite, dehisc- Sivadasan (1982), Grayum (1987). A C O R A C E A E : A C O R U S 289