BOOK-ARACEAE

100. Spathe tube narrow, elongate; female zone of spadix mostly adnate to C spathe; stylar region thin, spreading, diaphanous, mantle-like; synan- drodes (sterile flowers) betweeen male and female flowers usually irregular or fungiform, not prismatic · · · · · · · · · · 66. Chlorospatha 99. Pollen shed in monads; stylar region not laterally expanded; leaf blade usu- ally peltate, rarely trisect, never pedatifid or -sect 101. Spathe tube always convolute; stylar region as broad as ovary (Caladium paradoxum has discoid, coherent stylar regions); synandrodes (sterile flowers) between male and female flowers well-developed, prismatic; placentas 1–2(–3), parietal; seeds several (rarely 1–2) · · · 63. Caladium 101. Spathe tube gaping widely at anthesis; style much narrower than ovary; synandrodes (sterile flowers) lacking, male and female zones contigu- ous; placenta 1, basal; seed solitary · · · · · · · · · · · 62. Scaphispatha 98. Submerged aquatics; leaf blade linear · · · · · · · · · · · · · · · · · · · · · 64. Jasarum 97. Ovary clearly 1-locular, placentas not intrusive; palaeotropical plants 102. Ovules more than 1; leaf blade peltate 103. Female flowers with staminodes; spathe not constricted; stem trunk-like or creeping · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 101. Steudnera 103. Female flowers without staminodes; spathe with 1 or 2 constrictions; stem tuberous, producing erect or spreading stolons bearing small tubercles covered in hooked scales · · · · · · · · · · · · · · · · · · · · · · 102. Remusatia 102. Ovule solitary; leaf blade not peltate · · · · · · · · · · · · · · · · · · · · 68. Hapaline 96. Spadix with an appendix (occasionally absent in Colocasia esculenta); palaeotropical plants 104. Leaf blade pedatisect to radiatisect; female flowers each with several large stamin- odes · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · 100. Protarum 104. Leaf blade entire or pinnatifid; female flowers without staminodes (except single small ones in Colocasia esculenta) 105. Placentas parietal; ovules many; leaf blade always entire · · · 103. Colocasia 105. Placenta basal; ovules few; leaf blade entire or pinnatifid · · · 104. Alocasia 90 T H E G E N E R A O F A R A C E A E

26 D E S C R I P T I O N S O F T H E T R I B E S A N D G E N E R A O F A R A C E A E C I. Subfamily Gymnostachydoideae the floral sympodium. SPADIX: short- to long-stipitate, cylin- dric, many-flowered, erect in flower, pendent in fruit. Subfamily Gymnostachydoideae Bogner & Nicolson in FLOWERS: bisexual, perigoniate, often somewhat distant; Willdenowia 21: 37 (1991). tepals 4, ± as long as wide, fornicate, imbricate. STAMENS: 4, free, filaments somewhat flattened, anthers short, con- Laticifers absent; stem a short rhizome; leaves distichous, nective slender, thecae ellipsoid, dehiscing by a longitudinal bifacial (dorsiventrally flattened), linear, not differentiated slit, connective inconspicuous. POLLEN: monosulcate, ellip- into blade and petiole, primary veins parallel; flowering shoot soid, medium-sized (mean 31 µm., range 30–33 µm.), exine a complex synflorescence consisting of 3–6(7), short, peren- foveolate to slightly fossulate, apertural exine fossulate-ver- nating floral sympodia, borne on an erect peduncular axis, rucate. GYNOECIUM: ovary oblong, 1-locular, ovule 1, each sympodium axillary to a bract and composed of numer- orthotropous, funicle very short, placenta apical, stylar ous inflorescences; spathe inconspicuous; spadix region shortly attenuate, stigma small, subhemispheric. long-stipitate; flowers bisexual, perigoniate, 2-merous; tepals BERRY: ellipsoid, deep blue, apiculate, 1-seeded, strongly 4, fornicate, stamens 4, free, thecae dehiscing by longitudi- projecting beyond tepals. SEED: ellipsoid to obovoid, testa nal slit, pollen monosulcate; ovary 1-locular, ovule 1, absent at maturity, embryo axile, ± elongate, endosperm orthotropous, placenta apical, stigma small; berries deep copious. See Plates 1, 107A. blue, long-exserted beyond tepals; seed obovoid, testa absent CHROMOSOMES: 2n = 48. at maturity, embryo axile, endosperm copious. DISTRIBUTION: 1 sp.; Australia (Queensland, New South Wales). C 1. Gymnostachys ECOLOGY: southern temperate to subtropical or rarely tropical rainforest; understorey plant in moist hardwood forests, most common on cool southerly slopes and in cool Gymnostachys R. Brown, Prodr. 337 (1810). TYPE: G. moist gullies. anceps R. Brown ETYMOLOGY: Greek gymnos (naked) and stachys (spike). HABIT: acaulescent herbs, stem a short, thick subterranean TAXONOMIC ACCOUNTS: Engler (1905), Shelton (1980), rhizome. LEAVES: distichous, bifacial (dorsiventrally flat- Elliot & Jones (1990), Hnatiuk (1990), Hay (1993c). tened), somewhat plicate, linear, not differentiated into blade and petiole, margins erose-serrate; midrib not differentiated, primary lateral veins parallel, somewhat prominent, higher II. Subfamily Orontioideae C order venation parallel. INFLORESCENCE: borne on a long scape, terminating in 3–6(–7), short, perennating floral sym- Subfamily Orontioideae Mayo, Bogner & P.C. Boyce, podia separated from each other by a distinct peduncular Genera of Araceae p. 346 (1997). axis, each sympodium composed of several spadices and borne axillary to a leaf-like bract. SCAPE: subequal to leaves, Laticifers absent (except Orontium); helophytes, stem a stout, alate on each side, margins erose-serrate. SPATHE: a simple, erect rhizome, continuation shoot in axil of last leaf preced- short, keeled, inconspicuous bract, ± alternating with the ing spathe; petiole not geniculate apically, sheath long; leaf short, linear-lanceolate, 2-keeled prophylls of each article of blade entire, oblong-elliptic, higher order venation reticu- late; flowers bisexual, perigoniate; tepals free, fornicate, stamen with distinct filament, anther terminal, connective slender, thecae dehiscing by longitudinal slit, pollen mono- sulcate; ovary ± immersed in spadix axis; seed testa smooth, thin or absent at maturity, embryo large, endosperm very sparse to absent. 2. Orontium C 1. Gymnostachys Orontium L., Sp. Pl. 324 (1753). TYPE: O. aquaticum L. SYNONYMS: Amidena Adanson, Fam. 2: 470 (1763); Aronia J. Mitchell, Diss. Gen. Pl. 28 (1769), nom. rej. Laticifers present, simple, articulated. HABIT: seasonally dor- mant, aquatic herbs, continuation shoot of stem sympodium arising in axil of last foliage leaf of each article and begin- ning with a foliage leaf rather than a cataphyll, rhizome short, erect, buried deep in soil. LEAVES: several. PETIOLE: O R O N T I O I D E A E : O R O N T I U M 91

D A C H J K E L BF G M Plate 1. Gymnostachys. A, habit × 1/6; B, base of plant × 2/3; C, detail of leaf venation showing erose-serrate midrib and margin x 5; D, cross-section of leaf × 5; E upper part of inflorescence × 1/3; F, detail of peduncle showing erose margin × 5; G, spadix × 1; H, detail of spadix × 10; J, tepal, side view × 20; K, flower with perigone removed × 20; L, gynoecium, longitudinal section × 20; M, berry × 2. Gymnostachys anceps: A, Dransfield s.n. (Kew slide collection); B, Cunningham 337 (K); C–D, Forster 4851 (K); E, Brown s.n. (K); F–M, Covery 10432 (Kew spirit collection 29047.718). 92 T H E G E N E R A O F A R A C E A E

HJ C A F DB E G Plate 2. Orontium. A, habit × 1/5; B, habit × 2/3; C, detail of leaf venation × 5; D, detail of spadix × 5; E, flower × 10; F, stamen, adax- ial view × 10; G, gynoecium, longitudinal section × 10; H, fruit × 4; J, fruit, longitudinal section × 4. Orontium aquaticum: A, Boyce s.n. (Kew slide collection); B–C, Curtis s.n. (K); D–G, Mayo s.n. (Kew spirit collection 46576) (K); H, Hooker s.n. (K); J, Bogner 1896 (Kew spirit collection 56428). O R O N T I O I D E A E : O R O N T I U M 93

ETYMOLOGY: ancient Greek orontion referred to an unknown plant used for treating jaundice; according to some authors named after the River Orontes in Syria. TAXONOMIC ACCOUNTS: Huttleston (1953), Klotz (1991, 1993). 3. Lysichiton C 2. Orontium Lysichiton Schott in Oesterr. bot. Wochenbl. 7: 62 (1857). TYPE: L. camtschatcensis (L.) Schott (“camtschatcense”; terete, sheath long. BLADE: oblong-elliptic, held above water Dracontium camtschatcense L.) surface or floating on it; primary lateral veins arising at base of blade, arcuately ascending and running into apex, second- SYNONYMS: Arctiodracon A. Gray in Mem. Amer. Acad. ary laterals inconspicuous and parallel to primaries, higher Arts, ser. 2, 6: 408 (1859); [Lysichitum Schott in Oesterr. bot. order venation transverse-reticulate. INFLORESCENCE: 1–2 Wochenbl. 7: 62 (1857), orth. var.] in each floral sympodium. PEDUNCLE: absent, spadix sup- HABIT: large herbs with thick, hypogeal rhizome. LEAVES: ported on elongated stipe. SPATHE: a short, inconspicuous, numerous. PETIOLE: thick, flattened to deeply sulcate, sheath simple bract, inserted at point of attachment of inflores- long. BLADE: becoming large, ovate-elliptic to ovate-oblong; cence to rhizome, enclosing young spadix, later separated primary lateral veins pinnate, running into inconspicuous from it and disintegrating. SPADIX: held above water level, marginal vein, higher order venation ± regularly transverse- conical, slender, flowering acropetally, golden yellow, reticulate between primaries. INFLORESCENCE: solitary, peduncle-like stipe very long, often partly submerged, green- appearing just before or with the leaves in early spring. ish to red-brown with white, swollen apical part. FLOWERS: PEDUNCLE: absent, spadix supported on elongated stipe. bisexual except for some male flowers at spadix apex, SPATHE: marcescent, inserted at the point of attachment of perigoniate, tepals 6, sometimes 4, short, about as long as spadix stipe to rhizome, cucullate to boat-shaped, lower part wide, fornicate, subtruncate, irregularly imbricate. STAMENS: narrowly convolute and clasping spadix stipe, blade 6, sometimes 4, free, filaments flattened, connective slender, expanded, ovate-elliptic, white or yellow. SPADIX: subcylin- thecae ellipsoid, dehiscing by broad apical slit. POLLEN: dric, subacute at apex, stipe extremely elongated. FLOWERS: monosulcate, ellipsoid, large (mean 64 µm., range 55–73 bisexual, perigoniate; tepals 4, cucullate, imbricate. STAMENS: µm.), exine densely foveolate-fossulate. GYNOECIUM: ovary 4, free, filaments oblong, flattened, connective slender, thecae depressed-globose, broader than long, 1-locular, ovule 1, ellipsoid, dehiscing by longitudinal slit. POLLEN: monosulcate, hemianatropous, held horizontally, funicle short, placenta ellipsoid, medium-sized (mean 40 µm., range 38–43 µm.), basal, stylar region ± absent, stigma small, subsessile, dis- exine reticulate. GYNOECIUM: elongate-ovoid, attenuate api- coid-hemispheric. BERRY: depressed-globose, green, cally, ovary immersed in spadix axis, 2–locular or 1-seeded, partially immersed in spadix axis. SEED: ± glo- incompletely 2-locular, ovules 1–2 per locule, orthotropous, bose, testa thin, smooth, ± transparent, decaying at maturity, broad, conoid, funicle very short and thick, placenta axile, raphe conspicuous in furrow-like depression, hilum red- thick, stylar region attenuate, longer than tepals, stigma dis- dish, embryo large, almost globose to depressed-globose, coid-hemispheric. BERRY: ellipsoid, mostly 2-seeded, green, with a small internal cavity situated below the hilum, outer decaying or stylar portion breaking off with perigone at matu- cell layers green, plumule well-developed with 1(–2) filiform rity to expose seeds embedded in spadix axis. SEED: ± leaf primordia, endosperm absent. See Plates 2, 107B. ellipsoid, somewhat compressed, testa smooth, embryo large, CHROMOSOMES: 2n = 26. ellipsoid, endosperm nearly absent, present only as a single DISTRIBUTION: 1 sp.; temperate E. North America:– USA cell layer. See Plates 3, 107C. (Alabama, Carolinas, Connecticut, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Massachusetts, Mississippi, 3. Lysichiton New Jersey, New York, Pennsylvania, Rhode Is., Tennessee, Virginia, West Virginia). ECOLOGY: temperate wetlands, occurring from sea level to 900m alt.; partly submersed aquatic plant growing in open or partially shaded swamps, marshes, lakes and ponds, in dense stands or scattered colonies. 94 T H E G E N E R A O F A R A C E A E

HJ G KL M D B E F AC Plate 3. Lysichiton. A, flowering habit × 1/3; B, leaf × 1/3; C, detail of junction of spathe, stipe and rhizome, longitudinal section × 2; D, peduncle surrounded by spathe base, transverse section × 2; E, flowering habit × 1/10; F, vegetative and fruiting habit × 1/8; G, detail of spadix × 3; H, flower × 6; J, flower, longitudinal section × 6; K, seed, adaxial view × 3; L, seed, side view × 3; M, seed, abaxial view × 3. Lysichiton americanus: A-D, Cult. Kew 1969–18002; E, Ballard 364 (Kew slide collection); F, Townsend 87/142 (Kew slide collection); G–J, Cult. Jodrell (Kew spirit collection 29047.78 & 29047.764); K–M, Townsend 87/127 (Kew spirit collection 51789). O R O N T I O I D E A E : L Y S I C H I T O N 95

CHROMOSOMES: 2n = 28. coarsely verrucate. GYNOECIUM: ovary somewhat immersed DISTRIBUTION: 2 spp.; temperate E. Asia and W. North in spadix axis, 1-locular, ovule 1, orthotropous, funicle very America:– Canada (British Columbia), Japan, Russian Far East short, placenta apical-parietal, stylar region long-attenuate, (Kamchatka, Kuril Is., Okhotsk, Sakhalin, Udsk), USA (Alaska, stigma punctate-discoid. BERRY: tepals and style persistent to California, Idaho, Montana, Oregon, Washington). ripe fruiting stage, seed and base of berry immersed in spongy ECOLOGY: temperate wetlands; helophytes, in swamps and spadix axis, infructescence ± globose. SEED: globose, testa wet woodlands. thin, smooth, embryo globose, endosperm very sparse, only ETYMOLOGY: ancient Greek lysis (free) and chiton (tunic). a single cell layer thick. See Plates 4, 107D. TAXONOMIC ACCOUNTS: Krause (1908), Hultén & St. John CHROMOSOMES: 2n = 30, 60 (28). (1931,1956), Huttleston (1953, 1955), Nicolson (1981). DISTRIBUTION: 3 spp.; temperate E. Asia and E. North America:– Canada (New Brunswick, Nova Scotia, Quebec, C 4. Symplocarpus Ontario), China, Japan, Korea N. & S., Russian Far East, USA (Connecticut, Delaware, Indiana, Illinois, Iowa, Maine, Maryland, Massachusetts, Michigan, Minnesota, New Symplocarpus R.A. Salisbury ex Nuttall in W. Barton, Gen. Hampshire, New Jersey, New York, North Carolina, Ohio, 1: 105 (1817), (see Taxon 29: 601, Phytologia 72: 80–92) Pennsylvania, Rhode Is., Tennessee, Virginia, West Virginia, nom. cons. TYPE: S. foetidus (L.) Nuttall (Dracontium Wisconsin). foetidum L.) ECOLOGY: temperate damp woodlands, rarely open wet- SYNONYMS: Ictodes Bigelow in Amer. Med. Bot. 2: 41 lands, from near sea level to ca. 900m. alt.; usually in shaded (1818); Spathyema Rafinesque, Fl. Tell. 4: 13 (1838, “1836”); sites, frequent near water courses; S. foetidus and S. renifolius [Symplocarpos J.A. Schultes & J.H. Schultes, Mant. 3: 16 flower in early spring before the leaves appear and fruit in (1827), orth. var.] the summer of the same year; S. nipponicus flowers after the HABIT: fairly large, seasonally dormant herbs, rhizome thick, leaves appear and the fruits ripen in the following spring. erect with thick roots. LEAVES: few. PETIOLE: fairly broad, sul- ETYMOLOGY: Greek syn- (together), -plo (folded) and karpos cate, sheath short. BLADE: subcordate- to cordate-ovate; fruit); refers to immersion of gynoecia and fruits in spadix axis. primary lateral veins pinnate, arching towards apex, running TAXONOMIC ACCOUNTS: Huttleston (1953), Lee (1985). into inconspicuous marginal vein, secondary laterals and higher order venation reticulate to transverse-reticulate. INFLORESCENCE: 1–2 in each floral sympodium, appearing III. Subfamily Pothoideae C before or with leaves. PEDUNCLE: only shortly exserted above ground. SPATHE: thick, boat-shaped or conchiform, lower Subfamily Pothoideae Engler in Nova Acta Acad. Leopold.- part convolute, upper part somewhat to widely gaping, apex Carol. 39: 140 (1876). 2-keeled, rostrate, curving forwards. SPADIX: stipitate, sub- globose, hidden within spathe. FLOWERS: bisexual, Laticifers absent; stem usually aerial; petiole geniculate api- perigoniate; tepals 4, fornicate, imbricate. STAMENS: 4, free, cally; higher order leaf venation reticulate; spathe simple, filaments flattened, connective slender, thecae oblong, dehisc- spreading to reflexed, not enclosing spadix; flowers bisexual, ing by longitudinal slit. POLLEN: monosulcate, ellipsoid, perigoniate; stamen filament distinct, anther terminal, thecae medium-sized (mean 33 µm.), exine reticulate, apertural exine dehiscing by longitudinal slit, connective inconspicuous. 4. Symplocarpus 96 T H E G E N E R A O F A R A C E A E

D M F B L K J C E GA H Plate 4. Symplocarpus. A, habit showing vertical rhizome, longitudinally sectioned × 2/3; B, flowering habit × 1/3; C, leaf × 2/3; D, detail of leaf venation × 5; E, inflorescence, nearside half of spathe removed × 1; F, detail of spadix × 3; G, flower × 10; H, flower, nearside tepals removed × 10; J, gynoecium, longitudinal section × 10; K, infructescence × 1; L, seed, side view × 2; M, habit × 1/3. Symplocarpus foetidus: A, Herb Gray 7/82 (K) & Cult. Kew 1969–18003 (Kew spirit collection 51367); B, Cult. Kew 1969–18003 (Kew slide collection); C–D, W.D. s.n. (K); E–L, Cult. Kew 1969–18003 (Kew spirit collection 51367, 51616 & 55681); S. renifolius: M, Furuse 8737 (K). O R O N T I O I D E A E : S Y M P L O C A R P U S 97

C Tribe Potheae rescences, bearing 4–6 (sometimes more, e.g. P. insignis) rigid, coriaceous cataphylls at the base. PEDUNCLE: short to Tribe Potheae Engler in Nova Acta Acad. Leopold.-Carol. 39: long, sometimes reflexed. SPATHE: ovate to linear, rarely 140 (1876, Pothoeae). very long (P. mirabilis). SPADIX: globose, ovoid, cylindric, Laticifers absent; shrubby climbing herbs with tough woody ellipsoid or obovoid, sessile to long-stipitate, densely or laxly stems, main shoot monopodial, flowering shoots axillary or flowered. FLOWERS: bisexual, perigoniate; tepals 4–6, usu- infra-axillary; leaves distichous; petiole sheath long and often ally fornicate, free or partially to completely connate (e.g. P. broad, flattened and apically auriculate (reduced in rumphii). STAMENS: 4–6, free, filaments oblong, flattened, Pedicellarum and Pothos series Goniuri); flowers bisexual, connective slender, thecae ellipsoid, dehiscing by slit. perigoniate, (2–)3-merous; tepals (4–)6, stamens 6, free, pollen POLLEN: monosulcate, ellipsoid-oblong, small (mean 21 µm., monosulcate; ovules 1 per locule, anatropous, stigma sessile, range 16–25 µm.), exine foveolate to reticulate or subrugu- usually umbonate; embryo large, endosperm absent. late, muri psilate or minutely tuberculate. GYNOECIUM: ovary ovoid-oblong or depressed, (2?–)3-locular; ovules 1 per locule, anatropous, funicle short, placenta axile at base C 5. Pothos of septum, stylar region sometimes as broad as ovary, stigma discoid-hemispheric to umbonate. BERRY: ellipsoid to ovoid, Pothos L., Sp. Pl. 968 (1753). TYPE: P. scandens L. 1–3-seeded, red. SEED: ellipsoid, testa smooth, embryo large, SYNONYMS: Tapanava Adanson, Fam. 2: 470 (1763); endosperm absent. See Plates 5, 108A. CHROMOSOMES: 2n = 24, 36. Batis Blanco, Fl. Filip. 791 (1837); Goniurus Presl, Epim. DISTRIBUTION: ca. 70 spp.; south and southeast Asia, Bot. 244 (1851, “1849”); [Potha O. Kuntze, Rev. Gen. 2: 742 Australasia, Malagasy region, Malay Archipelago:– Australia (1891), orth.var.] (New South Wales, Queensland), Bangladesh, Brunei, Trichosclereids occasionally present. HABIT: climbing herbs, Burma, Cambodia, China (Guandong, Guangxi, Guizhou, stems rather woody, lower branches rooting, upper ones Hainan, Hunan, Sichuan, Taiwan, Yunnan), Comores Is., free and hanging, nodes rarely bearing short, clustered spines India, Indonesia (Borneo, Irian Jaya, Java, Moluccas, (P. armatus), buds of lateral shoots sometimes perforating the Sulawesi, Sumatra), Japan (Ryukyu Is.), Laos, Madagascar, leaf sheath or ± infra-axillary. LEAVES: distichous, juvenile Malaysia (Borneo, Peninsula), Nepal, Papua New Guinea, plants of some species with shingle form. PETIOLE: genicu- Philippines, Solomon Is., Sri Lanka, Thailand, Vanuatu, late (articulate) apically, either broad, completely flattened Vietnam. and usually auriculate apically, or morphology normal with ECOLOGY: tropical humid forest; usually climbing hemiepi- a long sheath, sometimes sheath reduced to a pair of hyaline phytes often in regrowth forest, rarely on rocks. ridges (series Goniuri). BLADE: linear-lanceolate to ovate or NOTES: Engler (1905) recognized 2 sections:– sect. Pothos elliptic, sometimes oblique; primary lateral veins either mostly (with 4 series), sect. Allopothos (with 3 series). arising near base of blade, long arcuate, and running into ETYMOLOGY: modified spelling of a Sinhalese vernacular marginal vein near apex, or primary lateral veins pinnate, name “potha”; P. scandens is still known as “pota-wel” in weakly differentiated, forming submarginal collective vein, Sri Lanka. 1–2 marginal veins also present, higher order venation retic- TAXONOMIC ACCOUNTS: Engler (1905), Sivadasan (1982), ulate in all types. INFLORESCENCE: axillary or infra-axillary, Nicolson (1988a), Sivadasan, Mohanan & Rajkumar (1989), solitary or forming short branching systems of several inflo- Boyce & Poulsen (1994), Boyce & Nguyen (1995). 5. Pothos 98 T H E G E N E R A O F A R A C E A E

C E B D J A M P F H S G R N KL Q Plate 5. Pothos. A, habit × 2/3; B, detail of leaf venation × 5; C, detail of spadix × 10; D, gynoecium, transverse section × 10; E, juvenile habit × 2/3; F, detail of spadix × 6; G, portion of flowering stem × 2/3; H, detail of petiole venation × 5; J, detail of leaf venation × 5; K, flow- ering shoot showing developing flagelliform shoot × 2/3; L, infructescence × 1; M, flower, top view × 10; N, flowering shoot × 2/3; P, detail of leaf venation × 5; Q, infructescence × 1; R, flower, top view × 10; S, gynoecium, longitudinal section × 10. Pothos beccarianus: A–D, Sumbing Jimpin SAN 110325 (K); P. motleyanus: E, Burbidge s.n. (K); P. barberianus: F, UNESCO (Kostermans) 192 (K & Kew spirit collection 58012); P. scandens: G–J, Cult. Kew (Kew spirit collection 19158); Soedarsono 280 (K); P. junghuhnii: K–M, Warnham s.n. Cult. Kew (K); P. rumphii: N–S, Forman 240 (K); Sands s.n., Cult Kew 1970–02267 (Kew spirit collection 29047.171). P O T H O I D E A E : P O T H O S 99

C 6. Pedicellarum Pedicellarum M. Hotta in Acta Phytotax. Geobot. 27 (34): 61 (1976). TYPE: P. paiei M. Hotta HABIT: climbing, shrubby herb, branches distichously leaved, 6. Pedicellarum C branching from below the nodes, juvenile shoots with shin- gle form, flowering branches producing flagelliform shoots. 7. Pothoidium LEAVES: several. PETIOLE: sometimes pubescent in young leaves, geniculate at apex, sheathed for the most part. Pothoidium Schott, Aroideae 6: 26, t. 57 (1857) & in Oesterr. BLADE: lanceolate to narrowly elliptic (nearly orbicular in bot. Wochenbl. 7: 70 (1857). TYPE: P. lobbianum Schott shingle form), acuminate; primary lateral veins pinnate, form- ing submarginal collective vein, 1–2 distinct marginal veins HABIT: climbing herb, stems somewhat woody, flowering also present, higher order venation reticulate. INFLORES- branches free and hanging. LEAVES: distichous, many. PETI- CENCE: arising below the nodes, bearing several short OLE: oblong, entirely flattened, resembling blade, venation cataphylls at base of peduncle. PEDUNCLE: shorter than parallel, joined to blade by constricted articulation. BLADE: spadix, subequal to or longer than petiole, very slender. much shorter than petiole, triangular-lanceolate; midrib SPATHE: small, fully expanded, ovate-cordate, membrana- absent, no primary veins differentiated, veins parallel, run- ceous. SPADIX: sparsely and very laxly flowered, ning into apex. INFLORESCENCE: several to many, borne in long-stipitate, axis flexuose, minutely hispid-papillose. FLOW- a terminal branching system, lower inflorescences axillary ERS: bisexual, perigoniate, on short pedicels, receptacle large to a foliage leaf, upper ones either subtended by cataphyll and conspicuous; tepals connate, forming cup-like structure. or without subtending leaf. PEDUNCLE: peduncular axis STAMENS: 6, free, filaments broad, flattened, connective slen- slender, composed of one to several internodes, sometimes der, thecae ellipsoid, latrorse, dehiscing by longitudinal slit. subtended by a prophyll, sometimes also bearing a cata- POLLEN: monosulcate, ellipsoid-oblong, small (mean 17 phyll ± halfway up. SPATHE: occurrence irregular, often µm.), exine reticulate, muri minutely tuberculate. GYNOE- absent, linear-lanceolate, widely spreading, margins usu- CIUM: obpyramidal, excavated at apex, ovary 3-locular, ally revolute. SPADIX: apparently often functionally ovules 1 per locule, anatropous, placenta axile at base of sep- unisexual, cylindric, sessile to long-stipitate when subtended tum, stigma sessile, umbonate. BERRY: obovoid, 1–3-seeded, red. SEED: compressed-ellipsoid, testa smooth, thin, embryo large, endosperm absent. See Plates 6, 108B. CHROMOSOMES: unknown. DISTRIBUTION: 1 sp.; Indonesia (Borneo), Malaysia (Borneo). ECOLOGY: tropical humid forest; climbing hemiepiphyte. NOTES: Appears closely related to Pothos series Goniuri, in which the spadix also has a slender axis and the flowers are distant. ETYMOLOGY: Latin pes, pedis (foot), -ella (diminutive) and Arum; refers to the unique character of pedicellate flowers. TAXONOMIC ACCOUNTS: Nicolson (1984b). 7. Pothoidium 100 T H E G E N E R A O F A R A C E A E

D E B A J HG C F Plate 6. Pedicellarum. A, habit × 2/3; B, petiole × 3; C, detail of petiole base × 5; D, detail of leaf venation × 5; E, juvenile shingle habit × 1; F, inflorescence × 2; G, flower × 10; H, gynoecium, transverse section × 10; J, infructescence × 2. Pedicellarum paiei: A–D, F, Paie SAN 16354 (K, L); E, Boyce 782 (K & Kew slide collection); G, Lassan SAN 107216 (K); H, Church et al. 303 (A, Kew spirit collection 58609); J, Lee S 54080 (L). P O T H O I D E A E : P E D I C E L L A R U M 101

C D B E G FA Plate 7. Pothoidium. A, habit × 2/4; B, detail of leaf blade and petiole venation × 2; C, infructescence × 1; D, detail of basal portion of spadix × 5; E, flower, top view × 10; F, flower, nearside tepal removed × 10; G, gynoecium, longitudinal section. Pothoidium lobbianum: A–B, de Vogel 3866 (K); C, Loher 7047 (K); D, Merrill 2293 (K & Kew spirit collection 58025); E–G, Herb. Lugd. Bat. (K). 102 T H E G E N E R A O F A R A C E A E

by spathe. FLOWERS: apparently usually unisexual, some- rotting to a fibrous mass (net-fibrous), sometimes com- times bisexual, perigoniate, male flowers with well pletely disappearing. PETIOLE: geniculate apically developed anthers and apparently sterile ovary, female (geniculum rarely well below blade, e.g. A. oerstedianum), flowers with large fertile ovary and lacking stamens; tepals variously shaped in cross-section, sheath long in juvenile 6, fornicate, membranaceous. STAMENS: 3–6, free, number (monopodial) leaves, very short in sympodial leaves. often varying on single spadix, filaments elongated and BLADE: small to very large (exceeding 2m), usually coria- overtopping perigone at anthesis, oblong-triangular, flat- ceous, more rarely membranaceous or stiff and brittle, tened, connective slender, thecae short, ellipsoid, dehiscing extraordinarily variable in shape, linear to orbicular in out- by broad slit. POLLEN: monosulcate, ellipsoid-oblong, line, rarely peltate, entire to trifid or trisect, or pedatifid or medium-sized (mean 26 µm.), exine reticulate with psilate subpalmatifid, or pedatisect to radiatisect, rarely the lobes muri, apertural exine shallowly fossulate or verrucate. or segments themselves pinnately lobed, blade base cuneate GYNOECIUM: broadly ovoid to subglobose or obovoid, to cordate, sagittate or hastate; primary lateral veins pinnate ovary 1-locular, ovule 1, anatropous, funicle short, placenta or more rarely all arising at the base, usually forming one subbasal, stylar region attenuate, stigma discoid-hemi- or more submarginal collective veins, basal ribs often pre- spheric. BERRY: ellipsoid to ovoid, apiculate (stigma sent in cordate leaves, higher order venation reticulate. remnant), prominently exserted when mature, red. SEED: FLOWERING BRANCHES: sympodial units usually compris- ellipsoid, testa smooth, embryo large, endosperm absent. ing one 2-keeled prophyll, one 1-keeled cataphyll, one See Plate 7. foliage leaf and terminal inflorescence. INFLORESCENCE: CHROMOSOMES: 2n = 24. always solitary. PEDUNCLE: usually rather elongated, rarely DISTRIBUTION: 1 sp.; southeast Asia, Malay Archipelago:– short. SPATHE: usually persistent, sometimes marcescent China (Taiwan), Indonesia (Moluccas, Sulawesi, Sumatra), or deciduous, usually linear to linear-lanceolate, more rarely Philippines. elliptic to ovate, broadly cordate to suborbicular, erect, ECOLOGY: tropical humid forest; climbing hemiepiphyte. spreading or reflexed. SPADIX: sessile to long-stipitate, usu- ETYMOLOGY: Pothos and Greek -idion (special); implies a ally cylindric to conic, more rarely clavate, rarely globose, plant distinct from Pothos. very short to very long (over 1m). FLOWERS: bisexual, TAXONOMIC ACCOUNTS: Engler (1905). perigoniate; tepals 4, fornicate, in 2 decussate whorls. STAMENS: 4, free, filaments somewhat flattened, usually equalling tepals at anthesis, sometimes exceeding them, C Tribe Anthurieae anthers short, connective slender, thecae ovate to oblong- ovate, dehiscing by longitudinal slit. POLLEN: forate (most Tribe Anthurieae Engler in Nova Acta Acad. Leopold.- often 3–4 pores), more rarely diporate, rarely inaperturate Carol. 39: 140 (1876). (sect. Polyphyllium), spherical to subspheroidal, small (mean 22 µm., range 14–29 µm.), exine foveolate to retic- Laticifers absent; climbers, hemiepiphytes, epiphytes, litho- phytes or terrestrial herbs, sympodial units of stem each normally composed of prophyll, cataphyll, foliage leaf, inflo- rescence (often aborted); petiole of sympodial leaves with very short sheath, geniculate apically; primary lateral veins pinnate (rarely all arising at petiole insertion) in blade of entire leaves and in lobes of compound leaves, usually form- ing one or more submarginal collective veins; spathe usually simple, spreading, reflexed or erect, linear to elliptic or ovate; flowers bisexual, perigoniate; tepals 4, fornicate, sta- mens 4, free, pollen usually forate (periporate) with 3–4 pores, more rarely diporate, rarely inaperturate (sect. Polyphyllium); ovary 2-locular, ovules 1–2 (rarely more) per locule, anatropous to campylotropous, placenta axile-sub- apical; mature berries exserted and usually dangling; seed often sticky, endosperm copious. C 8. Anthurium 8. Anthurium Anthurium Schott, in Wiener Z. Kunst 1829 (3): 828 (1829). LECTOTYPE: A. acaule (Jacquin) Schott (Pothos acaulis Jacquin, see Britton & Wilson, Sci. Surv. Porto Rico 5: 128. 1923). SYNONYMS: Podospadix Rafinesque, Fl. Tell. 4: 821 (1838, “1836”); Strepsanthera Rafinesque, Fl. Tell. 4: 13 (1838, “1836”). HABIT: evergreen herbs, stem erect, creeping, or short- to long-climbing, rarely rhizomatous, internodes very short (plant rosulate) to elongated. LEAVES: prophylls and cata- phylls usually ± persistent, entire (membranaceous) or P O T H O I D E A E : A N T H U R I U M 103

C F D E B A H J G Plate 8 (i). Anthurium. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2; F, leaf × 1/2; G, leaf × 1/2; H, leaf × 1/2; J, leaf × 1/2. Anthurium friedrichsthalii: A, Lehmann s.n. (K); A. vallense: B, Nee & Hale 9628 (K); A. carnosum: C, Valerio 181 (K); A. melastomatis: D, McPherson 7678 (K); A. smithii: E, Hahn & Grifo 3323 (K); A. antrophyoides: F, Lehmann 787 (K); A. peltigerum: G, Madison et al. 4599 (K); A. clidemioides: H, Stevens 24526 (K) & Bown 131/37 (Kew slide collection); A. puberulinervium: J, Croat 55033 (K). 104 T H E G E N E R A O F A R A C E A E

C B A D E Plate 8 (ii). Anthurium. A, leaf × 1/2; B, leaf × 1/2; C, leaf × 1/2; D, leaf × 1/2; E, leaf × 1/2. Anthurium watermaliense: A, Croat 36713 (K); A. rimbachii: B, Hepper 6445 (K); A. trisectum: C, Bogner 1088 (K); A. longissimum: D, Kalbreyer 853 (K); A. polyschistum: E, Paterson 121 (K). P O T H O I D E A E : A N T H U R I U M 105

A B CD Plate 8 (iii). Anthurium. A, habit × 1/16; B, habit × 1/8; C, habit × 1/2; D, habit × 1/2. Anthurium salviniae: A, Cult. Kew 1961–66404; A. oerstedianum: B, Cult. Kew 1981–3725; A. interruptum: C, Davidse & Herrera 31343 (K); A. melastomatis: D, McPherson 7678 (K). 106 T H E G E N E R A O F A R A C E A E

A B C D E Plate 8 (iv). Anthurium. A, habit × 1/3; B, habit × 1/8; C, habit × 1/16; D, habit × 1/3; E, habit × 1/6. Anthurium radicans: A, Cult. Kew. 1977–5366; A. andraeanum: B, Cult. Kew 1963–49801; A. warocqueanum: C, Cult. Kew. 1986–6028; A. affine: D, Harley et al. 19429 (Kew slide collection); A. wendlingeri: E, Cuadros et al. 3950 (K) & Nee et al. 8735 (K). P O T H O I D E A E : A N T H U R I U M 107

J K F A H G M P L B C DE N Plate 8 (v). Anthurium. A, inflorescence × 2; B, inflorescence, spathe mostly removed × 2; C, flower × 15; D, stamen, three quarter view × 15; E, gynoecium, longitudinal section × 15; F, infructescence × 1; G, inflorescence, spathe mostly removed × 2; H, flower × 15; J, stamen × 15; K, gynoecium, longitudinal section × 15; L, infructescence × 2; M, developing berry surrounded by persistent perigone × 8; N, inflores- cence × 1; P, flower × 10. Anthurium globosum: A, Croat 67966 (K & Kew spirit collection 58085); A. regale: B–E, Cult. Kew 1962–67111 (Kew spirit collection 51391); A. scandens subsp. scandens: F, Cult. Kew 1984–8010 (Kew spirit collection 52038); A. polyschistum: G–K, Cult. Kew 1976–1533 (Kew spirit collection 51386); A. sp.: L–M, (Kew spirit 59068); A. radicans: N–P, Cult. Kew. 1977–5366 & (Kew spirit collection 58028). 108 T H E G E N E R A O F A R A C E A E

ulate or subrugulate, rarely tuberculate, muri ± psilate or IV. Subfamily Monsteroideae C spinulose, apertural exine mostly psilate, rarely spinulose. C GYNOECIUM: ovary ovoid to oblong or obovoid, 2-locular, Subfamily Monsteroideae Engler in Nova Acta Acad. Leo- C ovules 1–2 per locule, rarely more, anatropous, hemiana- pold.-Carol. 39: 142 (1876). tropous or subcampylotropous, funicle short, placenta axile near apex of septum, stylar region inconspicuous to atten- Laticifers absent; trichosclereids (H- or T- shaped) abundant uated; stigma small, subcapitate, secreting conspicuous (except Anadendreae, Heteropsideae; in Amydrium present nectar droplet at anthesis. BERRY: variously shaped from only in certain parts); terrestrial, climbing hemiepiphytes or globose to elongate-fusiform, when mature exserted from more rarely epiphytes; petiole geniculate apically, sheath tepals and usually held dangling by tiny strips of inner tepal usually long; leaf blade never sagittate, base narrowed to epidermis, sometimes simply falling out of spadix, 2–4- subcordate, sometimes pinnatifid or perforated, outline seeded (more in sect. Tetraspermium), variously coloured, from always ± oblong-ovate to -elliptic or narrower, often oblique; con-spicuous reds and oranges to dull purplish green, white spathe expanded or boat-shaped, not constricted centrally; or blueish. SEED: ± oblong to ellipsoid or subglobose, spadix fertile to apex; flowers bisexual, perigone present or sometimes curved, testa usually smooth or somewhat ver- absent, 2-merous (2–3-merous in Spathiphylleae). rucose, thin, usually with sticky gelatinous mass adhering to raphe, small strophiole sometimes present, embryo axile, Tribe Spathiphylleae subcylindric to conoid, sometimes curved, endosperm copi- ous. See Plates 8i–v, 108C. Tribe Spathiphylleae Engler in Engler & Prantl, Nat. CHROMOSOMES: 2n = 30, 60, 90 (20, 24, 28, 40, 48, 56, 84) Pflanzenfam. II (3): 112, 121 (1887). DISTRIBUTION: over 800 spp.; tropical America, West Indies:– Argentina, Belize, Bolivia, Brazil, Colombia, Costa Laticifers absent; tissues with small trichosclereids occurring Rica, Cuba, Dominican Republic, Ecuador, El Salvador, French in bundles; leaves sometimes distichous, petiole sheath long; Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Lesser blade oblong, cuspidate-acuminate, higher order venation Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto parallel-pinnate; inflorescence solitary; spathe persistent or Rico, Surinam, Trinidad & Tobago, Uruguay, Venezuela. marcescent; flowers bisexual, perigoniate, 2–3-merous; tepals ECOLOGY: tropical humid forest, especially diverse in cloud free or connate, thecae dehiscing by longitudinal slit, pollen forests; climbing hemiepiphytes, terrestrial on forest floor, inaperturate, exine striate; ovules anatropous to hemianat- epiphytes, lithophytes, rarely helophytes or rheophytes. ropous; seed ± oblong, narrowed towards micropyle, NOTES: 19 sections are recognized by Croat & Sheffer endosperm copious. (1983):– Tetraspermium, Gymnopodium, Porphyrochit- onium, Pachyneurium, Polyphyllium, Leptanthurium, 9. Spathiphyllum Oxycarpium, Xialophyllium, Polyneurium, Anthurium (syn. Urospadix), Episeiostenium, Digitinervium, Cardiolonchium, Spathiphyllum Schott in Schott & Endlicher, Melet. Bot. 22 Chamaerepium, Calomystrium, Belolonchium, Semaeophyll- (1832). TYPE: S. lanceifolium (Jacquin) Schott (“lancae- ium, Schizoplacium, Dactylophyllium. folium”; Dracontium lanceaefolium Jacquin) ETYMOLOGY: Greek anthos (flower), oura (tail) and -ion (diminutive). SYNONYMS: Hydnostachyon Liebmann in Vidensk. TAXONOMIC ACCOUNTS: Engler (1905), Madison (1978c), Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1849: 23 (1849); Croat (1980), Mayo (1982), Croat & Sheffer (1983), Croat (1984, Massowia K. Koch in Bot. Zeitung (Berlin) 10: 277 (1852); 1986), Rodriguez (1987, 1989), Croat (1991), Sakuragui (1994). Spathiphyllopsis J.E. Teysmann & S. Binnendijk, Natuurk. 9. Spathiphyllum M O N S T E R O I D E A E : S P A T H I P H Y L L U M 109

Tijdschr. Ned.-Indië 25: 400 (1863); Amomophyllum Engler in (incl. New Britain, New Ireland), Peru, Philippines, Solomon C Gard. Chron., ser. 2, 7: 139 (1877, non Watelet 1866); Is., Surinam, Trinidad, Venezuela. [Massovia Bentham & J.D. Hooker, Gen. Pl. 3: 998 (1883), ECOLOGY: tropical humid forest, rarely cloud forest; forest orth. var.]. floor, in wet sites, sometimes on rocks in streams, rarely hemiepiphytic (S. solomonense). Trichosclereids present. HABIT: evergreen herbs usually NOTE: Bunting (1960a) recognized 4 sections:– sect. with short, erect to creeping stem, appearing acaulescent, Spathiphyllum, sect. Dysspathiphyllum, sect. Amomophyllum, sometimes stoloniferous, occasionally erect and climbing sect. Massowia; S. solomonense has recently been placed in (S. solomonense). LEAVES: several. PETIOLE: geniculate api- a separate section, sect. Chlaenophyllum (Nicolson 1994). cally, sheath long. BLADE: oblong to elliptic or narrowly ETYMOLOGY: Greek spathe (spathe) and phyllon (leaf). elliptic, cuspidate-acuminate; primary lateral veins pinnate, TAXONOMIC ACCOUNTS: Engler & Krause (1908), Bunting running into marginal vein, secondary and tertiary laterals (1960a), Nicolson (1968b, 1994), Williams & Dressler (1976). parallel-pinnate, higher order venation transverse-reticulate. INFLORESCENCE: solitary. PEDUNCLE: subequal to or 10. Holochlamys longer than petiole. SPATHE: oblong, elliptic, ovate or obo- vate, cuspidate-acuminate, ± decurrent at insertion, Holochlamys Engler in Beccari, Malesia 1: 265 (1883). TYPE: membranaceous to subcoriaceous, fully expanded, rarely H. beccarii (Engler) Engler (Spathiphyllum beccarii Engler). fornicate or clasping, persistent, with distinct midrib and pinnate primary lateral veins, usually white, rarely green, Trichosclereids present. HABIT: evergreen herbs, stem turning green in fruit. SPADIX: usually stipitate, rarely ses- short, upright. LEAVES: several. PETIOLE: geniculate api- sile, stipe often partially adnate to spathe, spadix cylindric, cally. BLADE: oblong-elliptic, or ovate to lanceolate, often erect, shorter than spathe. FLOWERS: bisexual, perigoniate; oblique, apex cuspidate to acuminate, base attenuate to tepals 4–6, free, fornicate and almost truncate at apex, or rounded; primary lateral veins pinnate, running into mar- partly or completely connate into a truncate cup. STAMENS: ginal vein, secondary and tertiary laterals parallel-pinnate, 4–6, free, filaments short, oblong, flattened, connective slen- higher order venation forming transverse cross connec- der, thecae oblong-ellipsoid to ovoid, dehiscing by tions, often obscured. INFLORESCENCE: solitary. longitudinal slit. POLLEN: inaperturate, ellipsoid to ellip- PEDUNCLE: shorter than petiole. SPATHE: white, with dis- soid-oblong, medium-sized (mean 32 µm., range 27–41 µm.), tinct midrib and pinnate primary lateral veins, tightly exine striate. GYNOECIUM: ovoid, subcylindric, obovoid or clasping the spadix, marcescent after anthesis, gradually flask-shaped, ovary 3-locular, more rarely 2- or 4-locular, decomposing. SPADIX: sessile to shortly stipitate, cylin- ovules 2, 4, 6 or 8 per locule, anatropous to hemianatropous, dric, fertile to apex. FLOWERS: bisexual, perigoniate; tepals placenta axile, stylar region usually long, conic and long- 4, fornicate apically and ± truncate, connate into a truncate exserted beyond perigone, sometimes shortly attenuate, cup. STAMENS: 4, free, filaments short, oblong, subequal sometimes ± truncate and not exserted and inconspicuous, to anthers, connective slender, thecae oblong, dehiscing by stigma 2–3-lobed or subcapitate to punctiform. BERRY: longitudinal slit. POLLEN: inaperturate, ellipsoid, medium- rounded, ovoid to obovoid, or conically attenuate apically, sized (mean 33 µm., range 32–34 µm.), exine striate. 1–8-seeded, greenish. SEED: oblong, ellipsoid to ovoid or GYNOECIUM: subcylindric to ovoid, ovary 1-locular, ovules slightly curved and ± reniform, pale yellow to brown, funi- many, anatropous, funicle long, placenta basal, stylar cle short, testa sparsely foveolate, otherwise smooth or region cylindric, ± as broad as ovary, stigma oblong or verrucose, embryo axile, elongate, slightly curved, 3–4-lobed. BERRY: 1–few-seeded. SEED: irregularly oblong- endosperm copious. See Plates 9, 108D. ellipsoid, narrowed towards micropyle, testa minutely CHROMOSOMES: 2n = 30, 60. verrucose or smooth, embryo elongate, endosperm copious. DISTRIBUTION: 41 spp.; tropical America, West Indies, east- See Plate 10. ern Malay Archipelago, Melanesia:– Belize, Brazil (Amazonia, CHROMOSOMES: 2n = 60. Central-West, Northeast), Colombia, Costa Rica (incl. Isla del DISTRIBUTION: 1 sp.; Indonesia (Irian Jaya), Papua New Coco), Ecuador, El Salvador, French Guiana, Guatemala, Guinea (incl. New Britain). Guyana, Honduras, Indonesia (Irian Jaya, Moluccas, Palau Is., ECOLOGY: tropical humid forest; on forest floor, particu- Sulawesi), Mexico, Nicaragua, Panama, Papua New Guinea larly along small streams, on river banks or on rocks. ETYMOLOGY: Greek holos (whole) and chlamys (short man- 10. Holochlamys tle, cloak); refers to the connate tepals of the flowers. TAXONOMIC ACCOUNTS: Engler & Krause (1908), Hay (1990a). Tribe Anadendreae C Tribe Anadendreae Bogner & French in Taxon 33(4): 689 (1984). Laticifers and trichosclereids absent; climbing hemiepiphytes; leaves distichous; petiole sheathed almost to apex; finer vena- tion reticulate; peduncle relatively long; spathe boat-shaped to reflexed, longer than spadix, marcescent or deciduous soon after anthesis; flowers bisexual, perigone membrana- 110 T H E G E N E R A O F A R A C E A E

A QR S C p J N D K E B F GH L M Plate 9. Spathiphyllum. A, habit × 1/5; B, leaf × 2/3; C, detail of leaf venation × 5; D, inflorescence × 2/3; E, detail of spadix × 3; F, flower × 6; G, flower, perigone removed × 6; H, gynoecium, longitudinal section × 6; J, inflorescence × 2/3; K, detail of spadix × 3; L, flower with developing fruit, longitudinal section × 6; M, fruit, transverse section × 6; N, inflorescence × 2/3; P, detail of spadix × 3; Q, flower × 6; R, flower, perigone removed × 6; S, gynoecium, longitudinal section × 6. Spathiphyllum ‘Clevelandii’ : A, Cult Kew 1962–43301 (Kew slide col- lection); S. laeve: B–D, Dressler 4465 (K); S. cannifolium: E–H, Jermy 2872, Cult. Kew 1976–56863 (Kew spirit collection 37366); S. humboldtii: J–K, Cremers 7813 (K); L–M, Jonker et al. 5845 (K); S. cochlearispathum: N–S, Cult. Kew 1972–68267 (Kew spirit collection 29047.83). M O N S T E R O I D E A E : S P A T H I P H Y L L U M 111

C A F GH D BE Plate 10. Holochlamys. A, habit × 1/6; B, leaf × 2/3; C, detail of leaf venation × 5; D, inflorescence, lower portion of peduncle removed × 1; E, detail of spadix × 5; F, flower × 10; G, flower, perigone removed × 10; H, gynoecium, longitudinal section × 15. Holochlamys beccarii: A–C, Cult. Kew 1970–1474; D, Sands 889 (Kew spirit collection 34304); E–H, Sands s.n. (Kew spirit collection 56122). 112 T H E G E N E R A O F A R A C E A E

ceous, urceolate, shorter than gynoecium; stamens 4, thecae DISTRIBUTION: ca. 7 spp.; southeast Asia, Malay Archipelago:- dehiscing by longitudinal slit, pollen inaperturate; gynoe- Brunei, Cambodia, China (Guandong, Hainan, Yunnan), cium truncate apically, ovary 1-locular, ovule 1, Indonesia (Borneo, Java, Sulawesi, Sumatra), Laos, Malaysia hemianatropous, placenta basal, stigma transversely oblong (Borneo, Peninsula), Philippines, Thailand, Vietnam. or subspheroid; embryo large, endosperm absent. ECOLOGY: tropical humid forest; climbing hemiepiphytes, sometimes on rocks. ETYMOLOGY: Greek ana (up) and dendron (tree), a tree C 11. Anadendrum climber. TAXONOMIC ACCOUNTS: Engler (1905), Bogner & French Anadendrum Schott in Bonplandia 5: 45 (1857). LECTOTYPE: (1984). A. montanum Schott (see Engler in De Candolle, Monogr. Phan. 2: 97, 250 (1879)). C C SYNONYMS: [Anadendron Schott in Oesterr. bot. Tribe Heteropsideae Wochenbl. 7: 118 (1857), orth. var.]. Tribe Heteropsideae Engler in Engler, Pflanzenreich 21 Trichosclereids absent. HABIT: climbing herbs. LEAVES: disti- (IV.23B): 20 (1905). chous. PETIOLE: geniculate apically, sheathed nearly to apex, Laticifers and trichosclereids absent; climbing hemiepiphytes, sheath persistent or marcescent. BLADE: obliquely ovate- main shoot monopodial, flowering articles usually short, axil- oblong, entire; primary lateral veins pinnate, running into lary; leaves distichous; petiole usually almost entirely adnate to marginal vein, higher order venation reticulate. INFLORES- succeeding internode, leaving only apical geniculum free; blade CENCE: 1–3 in each floral sympodium. PEDUNCLE: relatively oblong to lanceolate, primary and secondary lateral veins par- long. SPATHE: oblong-ovate, boat-shaped to reflexed, greenish allel-pinnate, forming submarginal collective vein, higher order white, rostrate apically and overtopping the spadix, deciduous venation reticulate; spathe boat-shaped, marcescent or decid- after anthesis. SPADIX: stipitate, cylindric. FLOWERS: bisexual, uous soon after anthesis; flowers bisexual, perigone absent; perigoniate; perigone membranaceous, a single cup-like struc- stamens 4, thecae dehiscing by apical slit, pollen zonate or ture, truncate, equalling or shorter than gynoecium. STAMENS: dicolpate; gynoecium truncate, ovary incompletely 2-locular, 4, free, filaments relatively short, broad, spathulate, connective ovules 2 per locule, anatropous, placenta axile at base of par- slender, thecae linear-elliptic, dehiscing by longitudinal slit. tial septum, stigma oblong or rounded; endosperm absent. POLLEN: inaperturate, subspheroidal, small (mean 22 µm.), exine psilate or subretipilate, pilae spinulose tipped and soli- tary, or united into groups of 2–4 or more. GYNOECIUM: ovary 12. Heteropsis obconic or obpyramidal, subquadrangular, 1-locular, ovule 1, ana-tropous, funicle short, placenta basal, stylar region as broad Heteropsis Kunth, Enum. Pl. 3: 59 (1841). LECTOTYPE: H. as ovary, stigma transversely oblong. BERRIES: distinctly trun- salicifolia Kunth (see Nicolson in Taxon 24: 468. 1975). cate apically, subglobose, orange red (A. microstachyum). SEED: rounded, subglobose, testa smooth, glossy, embryo Trichosclereids absent. HABIT: evergreen climbing herbs with large, endosperm absent. See Plate 11. woody-fibrous roots. LEAVES: numerous. PETIOLE: usually CHROMOSOMES: 2n = 60. very short, entirely geniculate, concave and somewhat flat- 11. Anadendrum M O N S T E R O I D E A E : H E T E R O P S I S 113

B C E A FG D Plate 11. Anadendrum. A, habit × 2/3, B, detail of leaf venation × 5; C, juvenile habit × 5; D, detail of spadix × 5; E, flower × 6; F, flower, perigone removed × 6; G, gynoecium, longitudinal section × 6. Anadendrum microstachyum: A, Cult. Kew 1982–4984, Scortechini 82 (K); B, de Wilde 14630 (K); C, ‘Native collector’ 2398 (K); D–F, Cult. Kew 1982–4984; G, Burkill 3231 (K). 114 T H E G E N E R A O F A R A C E A E

E B A F C GH J D L K Plate 12. Heteropsis. A, habit × 2/3; B, detail of leaf venation × 5; C, fruiting habit × 2/3; D, juvenile habit × 2/3; E, spadix × 2; F, detail of spadix × 5; G, flower × 10; H, gynoecium, longitudinal section × 10; J, berry × 2; K, berry, longitudinal section × 2; L, berry, transverse section × 2. Heteropsis spruceana: A–B, Traill 1135 (K); H. oblongifolia: C, E–L, Bell et al. 88–178 (K); Gentry & Young 31969 (K); Pinheiro & Santos 2266 (K) & Santos 1276 (K); H. cf. jenmanii: D, Harley et al. 17848 (K). M O N S T E R O I D E A E : H E T E R O P S I S 115

ETYMOLOGY: Greek heteros (different) and opsis (appear- ance); this aroid is different from all others in that the petiole sheaths are adnate to the internodes in most species, leaving only a very short free portion of the petiole. TAXONOMIC ACCOUNT: Engler (1905). 12. Heteropsis Tribe Monstereae C C tened, sheath adnate to subtended internode, rarely petiole Tribe Monstereae Engler in Nova Acta Acad. Leopold.- free with long sheath (e.g. H. melinonii). BLADE: oblong, ellip- Carol. 39: 143 (1876). tic or lanceolate, ± long-cuspidate, subcoriaceous; primary lateral veins pinnate, forming a submarginal collective vein Laticifers absent; trichosclereids abundantly present (sparse very close to margin, 1(-2) marginal veins also present, sec- in Amydrium), large, solitary and scattered in tissues; climb- ondary laterals ± parallel to primaries, higher order venation ing hemiepiphytes and epiphytes; leaves distichous; petiole reticulate. INFLORESCENCE: solitary, subtended by several sheath usually long, almost equalling petiole (except small cataphylls, borne terminally on free, axillary branches. Amydrium, Alloschemone); blade normally oblique; pedun- PEDUNCLE: very short. SPATHE: ovate-elliptic to ovate-oblong, cle relatively short (except Stenospermation), spathe usually cuspidate, convolute, opening at anthesis, then caducous. erect and boat-shaped (except Amydrium), marcescent or SPADIX: erect, free, stipitate, shorter than spathe, cylindric or deciduous soon after anthesis; spadix equalling or shorter ellipsoid. FLOWERS: bisexual, or lowermost flowers female by than spathe; flowers bisexual, perigone absent; stamens 4, abortion of stamens, perigone absent. STAMENS: 4 or fewer by thecae dehiscing by longitudinal slit not reaching base, pollen abortion, free, filaments short, flattened, rather broad, connec- usually fully zonate, dicolpate; style well developed, rather tive slender, thecae ovate-ellipsoid, overtopping connective a massive, usually prismatic with ± truncate apex, as wide or little, dehiscing by apical slit. POLLEN: zonate or dicolpate, wider than ovary, containing abundant trichosclereids, stigma ellipsoid-oblong or hamburger-shaped, medium-sized (mean 40 ± hemispherical to linear, appearing sessile; stylar region of µm., range 37–42 µm.), exine foveolate or foveolate-fossulate, berry deciduous at maturity. apertural exine psilate. GYNOECIUM: obpyramidal- prismatic, truncate, ovary incompletely 2-locular with conspicuous sep- 13. Amydrium tal aperture, ovules 2 per locule, anatropous, collateral, placenta axile at base of partial septum, stylar region dense and thick- Amydrium Schott in Ann. Mus. Bot. Lugduno-Batavum 1: ened, broader than ovary, stigma very small, ellipsoid, oblong 127 (1863). TYPE: A. humile Schott or subhemispheric. BERRY: shortly obovoid or obpyramidal, ± prismatic, stylar region forming a ± broad, flattened scar-like SYNONYMS: Epipremnopsis Engler, Pflanzenreich 37 structure, 1-4 seeded, orange or greenish-white with brown (IV.23B): 1–3 (1908). apex. SEED: obovoid to ellipsoid, testa thin, smooth, black, shiny, embryo large, endosperm absent. See Plate 12. Trichosclereids sparsely present in vegetative parts (petiole CHROMOSOMES: 2n = 28. and sheath only, Seubert 1996b), more abundant in style DISTRIBUTION: ca. 13 spp.; tropical America:- Bolivia, Brazil (Carvell 1989). HABIT: evergreen herbs, often very robust, (Amazonia, Atlantic region), Colombia, Costa Rica, Ecuador, stem climbing or prostrate, usually producing long flagelli- French Guiana, Guyana, Nicaragua, Panama, Peru, Surinam, form shoots with reduced cataphylls. LEAVES: many, often Venezuela. remote from one another. PETIOLE: geniculate apically, ECOLOGY: tropical moist and humid forest; climbing sheath usually less than half as long as petiole. BLADE: ovate- hemiepiphytes, sometimes on rocks. cordate or pandurate-trilobed or pinnatifid to pinnatisect, NOTES: Heteropsis melinonii differs from other species of sometimes with ± numerous round to oval perforations near the genus in having free petiole sheaths, but agrees in ovary midrib; primary lateral veins pinnate, running into marginal structure (incompletely 2-locular ovary with 2 ovules per vein, higher order venation reticulate. INFLORESCENCE: locule), leaf venation and lack of trichosclereids. 1–several in each floral sympodium. PEDUNCLE: erect, sube- qual or half as long as petiole. SPATHE: conchiform to ovate, apiculate, sometimes reflexed at anthesis and then deciduous. SPADIX: sessile to long-stipitate, sometimes very short. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, fil- aments short, broadly linear, anthers equalling or shorter than filaments, thecae ovoid, extrorse, dehiscing by longitu- dinal slit. POLLEN: fully zonate, hamburger-shaped, medium-sized (mean 39 µm., range 38–41 µm.), exine either densely and minutely punctate in one half and virtually psi- late in other, or uniformly foveolate-fossulate, apertural exine psilate or obscurely verrucate. GYNOECIUM: obpyramidal or obconoid, tetragonal, ovary 1-locular, ovules 2, anatropous, funicle short, placenta near base of deeply intrusive septum, stylar region broader than ovary, slightly prominent centrally below stigma, otherwise ± truncate, stigma small, hemi- spheric. BERRY: subglobose, truncate to domed at apex, white (A. medium, A. humile) or orange-red (A. zippelianum) 116 T H E G E N E R A O F A R A C E A E

L K H G B C J D A M EF Plate 13. Amydrium. A, partial habit × 2/3; B, leaf × 1/2; C, tip of flagelliform shoot × 2/3; D, detail of spadix × 3; E, flower × 6; F, gynoe- cium, longitudinal section × 6; G, infructescence × 1; H, fruit, transverse section × 4; J, leaf × 1/2; K, detail of leaf venation × 5; L, seed, side view × 2; M, leaf × 1/2. Amydrium medium: A, Synge 35 (K); B, Foxworthy 4605 (K); C, Boyce 463, Cult. Kew 1989–3217; D–F, Motley 778 (K); G, Boyce 763 (Kew spirit collection 59090); H, Sam & Dewol s.n. (K); A. zippelianum: J–K, Ridley 26 (K); A. humile: L–M, de Wilde 13078 (K). M O N S T E R O I D E A E : A M Y D R I U M 117

13. Amydrium when ripe. SEED: subglobose to heart-shaped, testa smooth, 14. Rhaphidophora C glossy, embryo curved and partly green, endosperm present (E. Seubert 1993). See Plates 13, 109A. Rhaphidophora Hasskarl in Flora 25 (2) Beibl. 1: 11 (1842). CHROMOSOMES: 2n = 60 TYPE: R. lacera Hasskarl, nom. illeg. (Pothos pertusa DISTRIBUTION: 4–6 spp.; tropical southeast Asia, Malay Roxburgh, R. pertusa (Roxburgh) Schott). Archipelago:– Brunei, Burma, China (Guandong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Sichuan, Yunnan), SYNONYMS: Afrorhaphidophora Engler in Engler & Indonesia (Borneo, Irian Jaya, Java, Moluccas, Sulawesi, Prantl, Nat. Pflanzenfam. Nachtr. 3: 31 (1906); [Raphidophora Sumatra), Malaysia (Borneo, Peninsula), Papua New Guinea, Hasskarl, Cat. Hort. Bogor. 58 (1844), orth.var.]. Philippines, Thailand, Vietnam. ECOLOGY: tropical humid forest; climbing hemiepiphytes on Trichosclereids abundant. HABIT: evergreen, usually climb- tree trunks or creeping on forest floor (A. humile). ing herbs, more rarely repent, often extremely robust, ETYMOLOGY: Greek amydron (obscure, faint) and ion climbing branches often thick, producing anchor and (diminutive). feeder roots, flagelliform shoots also produced, stem often TAXONOMIC ACCOUNTS: Nicolson (1968c), Carvell (1989a). square in cross section. LEAVES: many, distichous, juvenile shingle plants occur in some species. PETIOLE: geniculate 14. Rhaphidophora 118 T H E G E N E R A O F A R A C E A E

C D B F K A E M G HJ L Plate 14. Rhaphidophora. A, habit × 1/2; B, leaf × 1/2; C, detail of leaf venation × 5; D, leaf × 1/2; E, leaf × 1/2; F, detail of spadix × 5; G, flower × 8; H, gynoecium, longitudinal section × 8; J, gynoecium, transverse section × 8; K, flower × 8; L, gynoecium, longitudinal sec- tion × 8; M, seed, side view × 20. Rhaphidophora foraminifera: A, Boyce 235 (Kew slide collection); de Wilde 14592 (K); de Wilde & de Wilde-Duyfjes 18154 (K). R. glauca: B–C, Henry 12728 (K); R. tenuis: D, Haviland & Hose 3605 (K); R. sylvestris: E, Mamit S 33623 (K); F–J, Cult. Kew 1967–460 (Kew spirit collection 35951); R. africana: K–L, Bogner 100 (Kew spirit collection 32065); R. decursiva: M, J. Arn. Arb. 57(4): 185–201, pl. I, 12 (1976). M O N S T E R O I D E A E : R H A P H I D O P H O R A 119

F C G H J A E K B D Plate 15. Epipremnum. A, infructescences and associated stem × 1/2; B, leaf × 1/2; C, juvenile habit × 1/2; D, seed, side view × 5; E, inflo- rescence and subtending leaf × 1/2; F, detail of leaf venation × 5; G, detail of spadix × 3; H, flower × 5; J, gynoecium, longitudinal section × 5; K, gynoecium, transverse section × 5. Epipremnum pinnatum: A, Kostermans 18558 (K); B, Bloembergen 3827 (K); C, Powell & Chey 783 (K); D, J. Arn. Arb. 57(4): 185–201, pl. I, 3 (1976); E. nobile: E–K, Forman 289 (K & Kew spirit collection 6919). 120 T H E G E N E R A O F A R A C E A E

apically, sheath usually relatively long. BLADE: lanceolate or oblong, ± oblique, entire, perforate or pinnatifid to pin- natisect, often very large, lobes often subfalcately narrowed; primary lateral veins pinnate, running into mar- ginal vein, often not differentiated from secondaries, secondary laterals ± parallel-pinnate, higher order venation reticulate. INFLORESCENCE: usually solitary, rarely more. PEDUNCLE: relatively short. SPATHE: boat-shaped, decid- uous. SPADIX: subcylindric, conic, clavate, often extremely thick, sessile to stipitate, shorter than spathe. FLOWERS: bisexual, perigone absent. STAMENS: 4, free, filaments oblong-linear, anthers much shorter than filaments, con- nective slender, thecae ellipsoid, dehiscing by longitudinal slit. POLLEN: dicolpate, extended monosulcate to perhaps fully zonate, ellipsoid or hamburger-shaped, medium-sized (mean 33 µm., range 24–55 µm.), exine foveolate, sub- reticulate, rugulate, fossulate, scabrate, retiscabrate, verrucate, or psilate. GYNOECIUM: obconic-prismatic to oblong, truncate, ovary 1- to partially 2-locular, ovules few to many, anatropous, funicle long, placentae parietal to basal, sometimes ± subaxile, partial septa variably intrusive, stylar region well developed, usually broader than ovary, 15. Epipremnum usually truncate apically, rarely elongate-conic, stigma broadly elliptic or oblong and then transverse or longitu- dinal, or punctate-prominent. BERRY: usually many-seeded, minutely perforate (E. pinnatum); primary lateral veins pin- stylar region deciduous at maturity, red or yellow. SEED: nate, running into marginal vein, secondary and often oblong, testa thin, smooth, embryo axile, straight, tertiaries parallel-pinnate, tertiary and higher order venation endosperm copious. See Plates 14, 109B. often reticulate. INFLORESCENCE: 1(–2) in each floral sym- CHROMOSOMES: 2n = 60, 120 (42, 54, 56). podium. PEDUNCLE: relatively short. SPATHE: boat-shaped, DISTRIBUTION: ca. 120 spp.; tropical Africa, tropical south- withering after anthesis, usually deciduous. SPADIX: sub- east Asia, Malay Archipelago, Melanesia, Australasia, cylindric, conic, often quite thick, sessile or stipitate, shorter Pacific:– Australia (Queensland), Bangladesh, Bhutan, than spathe. FLOWERS: bisexual, or lowermost ones female, Brunei, Burma, Cambodia, Cameroon, Caroline Is., China perigone absent. STAMENS: 4, free, filaments linear, some- (Fujin, Guandong, Guangxi, Guizhou, Hainan, Sichuan, what broad, anthers much shorter than filaments, connective Taiwan, Xizang, Yunnan), Equatorial Guinea (Bioko, Rio slender, thecae oblong-ellipsoid, dehiscing by longitudinal Muni), Fiji, Gabon, Ghana, India, Indonesia (Borneo, Irian slit. POLLEN: fully zonate, hamburger-shaped, medium-sized Jaya, Java, Moluccas, Palau Is, Sulawesi, Sumatra), Ivory (mean 40 µm., range 36–44 µm.), exine foveolate-fossulate, Coast, Japan (Bonin Is., Ryukyu Is.), Laos, Liberia, Malaysia psilate at periphery, apertural exine coarsely verrucate. (Borneo, Peninsula), Nepal, New Caledonia, Nigeria, Papua GYNOECIUM: ovary subtetragonal-prismatic, truncate, 1-loc- New Guinea, Philippines, Samoa, Sierra Leone, Singapore, ular, ovules usually 2, more rarely 4 or 6–8 (E. amplissimum), Solomon Is., Sri Lanka, Thailand, Togo, Uganda, Vanuatu, anatropous, funicle short, placenta parietal or near base of Vietnam. parietal partial septa, stylar region prismatic, as broad or ECOLOGY: subtropical and tropical humid or rain forest or broader than ovary, stigma umbonate to oblong-linear and deciduous forest; climbing hemiepiphytes, rarely rheophytic longitudinal. BERRY: 1–8-seeded, stylar region deciduous at (R. beccarii). maturity. SEED: reniform, testa thickish, brittle, smooth, ETYMOLOGY: Greek rhaphis, rhaphidos (needle) and pherô embryo curved, endosperm copious. See Plates 15, 109C. (I bear); refers to the macroscopic (to 1cm long), needle-like CHROMOSOMES: 2n = 60 (56, 84). unicellular trichosclereids present in tissues. DISTRIBUTION: 20 spp.; tropical southeast Asia, Australasia, TAXONOMIC ACCOUNTS: Engler & Krause (1908), Sivadasan Pacific:– Andaman Is., Australia (Queensland, Northern (1982), Nicolson (1988a), Hay (1990a, 1993b). Territories), Brunei, Burma, Cambodia, Caroline Is., China (Guandong, Guangxi, Hainan, Taiwan,Yunnan), Fiji, Indonesia C 15. Epipremnum (Irian Jaya, Java, Moluccas, Sulawesi, Sumatra, Timor), Japan (Bonin Is., Ryukyu Is.), Malaysia (Borneo, Peninsula), Marshall Epipremnum Schott in Bonplandia 5: 45 (1857). TYPE: E. Is., Papua New Guinea, Philippines, Singapore, Solomon Is., mirabile Schott (1858). Sri Lanka, Thailand, Vanuatu, Vietnam. ECOLOGY: tropical humid forest; high-climbing hemiepi- SYNONYM: Anthelia Schott in Ann. Mus. Lugduno- phytes, on trees and rocks. Batavum 1: 127 (1863). NOTES: Engler & Krause (1908) kept Epipremnum separate Trichosclereids abundant. HABIT: evergreen climbing herbs, from Rhaphidophora because of the differences in seed producing flagelliform shoots. LEAVES: several to many, dis- structure. tichous. PETIOLE: geniculate apically, sheath long, ETYMOLOGY: Greek prefix epi- (on) and premnon (bottom marcescent to deciduous, often decomposing to conspicuous of tree trunk). net-fibrous mass. BLADE: entire, often oblique, lanceolate, TAXONOMIC ACCOUNTS: Engler & Krause (1908), Hay elliptic, elliptic- oblong, or pinnatipartite to pinnatisect, rarely (1990a). M O N S T E R O I D E A E : E P I P R E M N U M 121

E DF C JB M KL A HG N Plate 16. Scindapsus. A, habit × 1/3; B, stigma, top view × 6; C, juvenile habit × 1/3; D, habit × 1/3; E, detail of leaf venation × 5; F, seed, side view × 2; G, habit showing fertile shoot and flagelliform shoot × 1/3; H, detail of spadix × 4; J, stigma, top view × 9; K, flower × 6; L, gynoecium, longitudinal section × 6; M, stigma, top view × 6; N, flower with window cut in gynoecium to show ovule × 6. Scindapsus rupestris: A, Chew & Corner 4253 (K); B, Beaman 9984 (K); S. pictus: C, Boyce 225 (K & Kew slide collection); S. officinalis: D–E, Kerr 20536 (K); F, J. Arn. Arb. 57(4): 185–201, pl. II, 31 (1976); S. beccarii: G, Boyce 318 (Kew slide collection); Cult. Kew 1965–47801 (Kew spirit col- lection 40953); H–L, Cult. Kew 1965–47801 (Kew spirit collection 40953); S. perakensis: M–N, Fedilis & Sumbing 68774 (K). 122 T H E G E N E R A O F A R A C E A E

C 16. Scindapsus Bangladesh, Bhutan, Brunei, Burma, Cambodia, Caroline Is., China, Fiji, India (Assam, Bengal, Sikkim), Indonesia (Borneo, Scindapsus Schott in Schott & Endlicher, Melet. Bot. 21 Irian Jaya, Java, Moluccas, Sulawesi, Sumatra), Laos?, Malaysia (1832). LECTOTYPE: S. officinalis (Roxburgh) Schott (Pothos (Borneo, Peninsula), Nepal, Papua New Guinea, Philippines, officinalis Roxburgh; see Schott, Prodr. syst. Aroid. 395–397 Samoa, Solomon Is., Sri Lanka, Thailand, Vietnam. (1860)). ECOLOGY: tropical humid forest or dry, deciduous or ever- green forest; climbing hemiepiphytes, also creeping over rocks, Trichosclereids abundant. HABIT: evergreen climbing herbs, often terrestrial when juvenile, rarely rheophytic (S. rupestris). sometimes very robust, sometimes producing flagelliform ETYMOLOGY: Greek skindapsos, once used for an ivy-like shoots, shoots with leaves evenly spaced or forming rosulate plant. flowering zones separated by zones with elongated internodes TAXONOMIC ACCOUNTS: Engler & Krause (1908), Bogner and smaller leaves, or solitary leaf trapping bole epiphyte. & Boyce (1994). LEAVES: many, juvenile plants often of shingle form. PETIOLE: geniculate apically, sheath usually broad, rarely decomposing to form persistent net-fibrous mass with abundant, stinging 17. Monstera C sclereids. BLADE: always entire, lanceolate, elliptic or ovate to obovate, acuminate, rarely variegated; primary lateral veins Monstera Adanson, Fam. Pl. 2: 470 (1763), nom. cons. TYPE: hardly differentiated, pinnate, running into marginal vein, sec- M. adansonii Schott (Dracontium pertusum), typ. cons. ondaries and also sometimes tertiaries parallel-pinnate, higher order venation inconspicuous, transverse-reticulate. INFLO- SYNONYMS: Tornelia Gutierrez ex Schott, Gen. Aroid. t. RESCENCE: always solitary. PEDUNCLE: shorter than petiole. 74 (1858); Serangium W. Wood ex R.A. Salisbury, Gen. Pl. SPATHE: boat-shaped, gaping only slightly, caducous to decid- Fragm. 5 (1866). uous. SPADIX: sessile to shortly stipitate, cylindric, narrowly Trichosclereids abundant. HABIT: evergreen climbing herbs. ellipsoid or clavate, a little shorter than spathe. FLOWERS: LEAVES: distichous, juvenile leaves sometimes of shingle plant bisexual, perigone absent. STAMENS: 4, free, filaments oblong, form, rarely variegated. PETIOLE: geniculate apically, sheath flattened, broadish, connective slender, thecae oblong-ellip- usually long, persistent or decomposing to fibrous or mem- soid, dehiscing by apical slit. POLLEN: fully zonate, branous mass or entirely deciduous. BLADE: entire, oblique, hamburger-shaped, medium-sized (mean 38 µm., range 33–45 oblong to ovate-elliptic, often conspicuously and elaborately µm.), exine shallowly and sparsely punctate, scabrate or nearly perforated, more rarely deeply pinnatifid; primary lateral veins psilate. GYNOECIUM: ovary sometimes short, compressed ± pinnate, running into marginal vein, rarely forming an irregu- cylindric, 1-locular, ovules 1(–2), anatropous, funicle short, lar submarginal collective vein (M. obliqua), secondary laterals placenta basal, stylar region well-developed, prismatic, trun- often parallel-pinnate, sometimes reticulated (e.g. M. dubia), cate or with shortly conic central projection supporting stigma, higher order venation reticulate. INFLORESCENCE: 1–several stigma globose, elongate-globose, elliptic, linear, or puncti- in each floral sympodium. PEDUNCLE: shorter than petiole. form. BERRY: stylar region deciduous when mature, red. SEED: SPATHE: ovate or oblong-ovate, cuspidate, boat-shaped and rounded, subreniform, compressed, testa thickish, sparsely somewhat convolute basally, white to rose-coloured within, verruculose or smooth, embryo curved, endosperm present remaining open after anthesis, caducous. SPADIX: sessile, sub- (Seubert 1993). See Plates 16, 109D. cylindric, somewhat shorter than spathe. FLOWERS: bisexual, CHROMOSOMES: 2n = 60 (42, 56, 58, 64, 70, 112). perigone absent, lowermost flowers usually sterile. STAMENS: DISTRIBUTION: ca. 36 spp.; tropical Asia, Malay Archipelago, 4, free, filaments flattened, connective slender, thecae oblong- Melanesia, Pacific:– Andaman Is., Australia (Queensland), ellipsoid, dehiscing by longitudinal slit. POLLEN: fully zonate, 16. Scindapsus 17. Monstera M O N S T E R O I D E A E : M O N S T E R A 123

E D B F M G C N A P J K LH Plate 17. Monstera. A, leaf × 1/3; B, detail of leaf venation × 5; C, leaf × 1/3; D, gynoecium × 4; E, gynoecium, longitudinal section × 4; F, leaf × 1/3; G, infructescence × 1/3; H, inflorescence × 1/3; J, detail of spadix showing emerging stamens × 2; K, flower × 4; L, gynoecium, longitudinal section × 4; M, seed, side view × 3; N, flowering shoot × 1/3; P, juvenile shingle habit × 1/3. Monstera oreophila: A–B, Grayum et al. 6395 (K); M. subpinnata: C, Santos & Souza 1667 (K); D–E, Vasquez 1853 (K); M. adansonii var. laniata: F, Gomez 19565 (K); G, Whitmore 749 (Kew spirit collection 25790); M. lechleriana: H–L, Cult. Mason (Kew spirit collection 29047.113); M, J. Arn. Arb. 57(4): 185–201, pl. III, 40 (1976); M. tuberculata var. tuberculata: N, Mayo & Madison 341 (Kew spirit collection 29047.346) & Contr. Gray Herb. 207: 3–100, f. 63 (1977); P, Mayo & Madison 341 (K). 124 T H E G E N E R A O F A R A C E A E

hamburger-shaped, medium-sized (mean 48 µm, range 40–52 18. Alloschemone µm), exine densely foveolate to subreticulate, sparsely and shallowly foveolate or psilate, apertural exine verrucate or rugulate. STERILE FLOWERS: with 4 minute, conic stamin- odia, pistillode 2-locular, prismatic, lacking ovules. GYNOECIUM: obovoid to ellipsoid, prismatic, ovary 2-locular, ovules 2 per locule, anatropous, funicle short, placenta axile at base of septum; stylar region often massive, broader than ovary, apex truncate to shortly attenuate, stigma oblong-ellip- tic to linear and longitudinal or round. BERRY: 1–3-seeded, shedding prismatic stylar region at maturity, pulpy within. SEED: obovoid to ellipsoid, compressed, testa smooth, embryo large, endosperm absent. See Plates 17, 110A. CHROMOSOMES: 2n = 60 (24, 48, 56, 58, 70). DISTRIBUTION: ca. 40 spp. (T. Croat, pers. comm.); tropical America, West Indies:– Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Honduras, Lesser Antilles, Mexico, Nicaragua, Panama, Peru, Surinam, Trinidad & Tobago, Venezuela. ECOLOGY: tropical moist and humid forest, cloud forest; climbing hemiepiphytes, usually on tree trunks, also on rocks or ground (M. deliciosa). NOTES: Madison (1977a) recognized four sections:– sect. Monstera, sect. Marcgraviopsis, sect. Echinospadix and sect. Tornelia. ETYMOLOGY: Latin monstrum (monster), refers to the pecu- liar perforations of the leaves of many species. TAXONOMIC ACCOUNTS: Engler & Krause (1908), Madison (1977a). C 18. Alloschemone 19. Rhodospatha C Alloschemone Schott, Gen. Aroid. 99 (1858). TYPE: A. Rhodospatha Poeppig in Poeppig & Endlicher, Nov. Gen. poeppigiana Schott, nom. illeg. (Scindapsus occidentalis Sp. 3: 91 (1845). LECTOTYPE: R. latifolia Poeppig (see Poeppig, A.. occidentalis (Poeppig) Engler & Krause). Nicolson in Taxon 16: 518. 1967). Trichosclereids abundant. HABIT: evergreen climbing herbs, SYNONYMS: Anepsias Schott, Gen. Aroid. t. 73 (1858); stem epidermis becoming distinctly corky and longitudinally Atimeta Schott, Gen. Aroid. t. 71 (1858). furrowed with age. LEAVES: large, juvenile leaves entire, ovate to ovate-elliptic. PETIOLE: geniculate apically, thick- Trichosclereids abundant. HABIT: evergreen, usually climb- ened at base, sheath short. BLADE: pinnatifid, subcordate, ing herbs, producing flagelliform shoots. LEAVES: many, lobes acute, falcate, 4–6 per side, primary lateral veins 1 per distichously arranged. PETIOLE: geniculate apically, sheath lobe, secondary laterals ± parallel-pinnate, higher order vena- long, persistent to marcescent. BLADE: oblong-elliptic, ± tion reticulate. INFLORESCENCE: solitary. PEDUNCLE: shorter oblique, always entire; primary lateral veins pinnate, numer- than petiole. SPATHE: ovate-cymbiform, deciduous. SPADIX: ous, running into ± distinct marginal vein, secondary and stipitate, cylindric, apex obtuse. FLOWERS: bisexual, perigone tertiary laterals parallel-pinnate, higher order venation trans- absent. STAMENS: 4, free, filaments flattened, free or connate, verse-reticulate. INFLORESCENCE: usually solitary. shorter than ovary, thecae oblong, dehiscing laterally by PEDUNCLE: shorter to longer than petiole. SPATHE: broadly oblique, apical, pore-like slit. POLLEN: not fully zonate, ellip- ovate or oblong-ovate, abruptly cuspidate, yellowish white, soid, medium-sized (mean 46 µm), exine shallowly foveolate. cream, purplish or pink within, caducous after anthesis. GYNOECIUM: ovary prismatic, 1-locular, with abundant loc- SPADIX: long-stipitate to sessile, cylindric-conic, basal flow- ular mucilage, ovule 1, amphitropous, funicle with trichomes, ers sometimes sterile or female and scattered. FLOWERS: placenta basal, stylar region densely packed with trichoscle- bisexual, perigone absent. STAMENS: 4, free, filaments lin- reids, stigma sessile, elliptic. BERRY: unknown. SEED: ear-oblong, flattened, connective slender, thecae ovoid to unknown. See Plate 18. ellipsoid, dehiscing by longitudinal slit. POLLEN: extruded CHROMOSOMES: 2n = 84. in strands, fully zonate or inaperturate, hamburger-shaped or DISTRIBUTION: 2 spp.; Brazil (Amazonas, Pará, Rondonia). ellipsoid to oblong, medium-sized (mean 47 µm., range ECOLOGY: tropical humid forest; climbing hemiepiphytes. 34–57 µm.), exine densely to sparsely foveolate and nearly NOTES: An incompletely known genus, which has been psilate to obscurely fossulate or verrucate. GYNOECIUM: attributed sometimes to Asiatic Scindapsus. compressed obconic to cylindric, ovary 2-locular, ovules ETYMOLOGY: Greek allos (other), schema, schêmatos (form) usually numerous per locule, rarely few (R. venosa), anat- and ône (being). ropous to hemianatropous, funicle fairly long, placenta axile, TAXONOMIC ACCOUNTS: Engler & Krause (1908), Madison rarely subbasal, stylar region well-developed, broader than (1976a), Boyce & Bogner (in prep.). ovary, prismatic, truncate to convex apically, stigma elliptic to linear, usually longitudinal. BERRY: cylindric-prismatic, M O N S T E R O I D E A E : R H O D O S P A T H A 125

B A C E DF G HJ Plate 18. Alloschemone. A, leaf × 1/3; B, detail of leaf venation × 5; C, leaf × 1/3; D, detail of leaf venation × 5; E, mature main stem show- ing corky epidermis × 1; F, inflorescence, part of spathe removed, remainder flattened out × 1/2; G, detail of spadix × 1; H, flower × 5; J, gynoecium, longitudinal section × 5. Alloschemone sp.: A–B, E, Madison et al. 6310 (K); A. occidentalis: C–D, F, H–J, Plowman et al. 12207 (NY); G, Krukoff 7162 (NY). 126 T H E G E N E R A O F A R A C E A E

F K D HJ BC E A L G Plate 19. Rhodospatha. A, habit × 1/2; B, flower × 8; C, gynoecium, longitudinal section × 8; D, seed × 20; E, habit × 1/2; F, detail of leaf vena- tion × 5; G, detail of spadix × 5; H, flower × 8; J, gynoecium, longitudinal section × 8; K, gynoecium, transverse section × 16; L, flagelliform shoot × 1. Rhodospatha oblongata: A–C, Harley et al. 18234 (K); D, Harley et al. 18193 (K); R. rubropunctata: E, J. Arn. Arb. 57(4): 185–201, pl. I, 4 (1976); R. venosa: F–L, Montfort 155 (K). M O N S T E R O I D E A E : R H O D O S P A T H A 127

19. Rhodospatha 20. Stenospermation truncate, many- to few-seeded. SEEDS: rounded-reniform, running into margin, secondary laterals parallel-pinnate, flattened, testa brittle, very hard, smooth or with verrucose higher order venation inconspicuous. INFLORESCENCE: crest, embryo rather large, strongly curved, endosperm pre- solitary, often nodding. PEDUNCLE: relatively long. SPATHE: sent but sparse. See Plates 19, 110B. convolute, gaping at anthesis, boat-shaped or opening CHROMOSOMES: 2n = 28, 56. widely, white, caducous. SPADIX: usually stipitate, rarely DISTRIBUTION: ca. 75 spp. (T. Croat pers. comm.); tropical sessile, cylindric. FLOWERS: bisexual, perigone absent. STA- America:– Belize, Bolivia, Brazil (Amazonia, Atlantic MENS: 4, free, filaments oblong, flattened, connective region), Colombia, Costa Rica, Ecuador, French Guiana, slender, thecae ovoid-ellipsoid, dehiscing by longitudinal Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, slit. POLLEN: extruded in strands, fully zonate or inapertu- Peru, Surinam, Trinidad, Venezuela. rate, hamburger-shaped or subspheroidal, medium-sized ECOLOGY: tropical humid forest; climbing hemiepiphytes, (mean 42 µm., range 30–58 µm.), exine psilate to shallowly true epiphytes (?) or sometimes on rocks, juvenile plants and sparsely foveolate, fossulate-foveolate, verrucate or often on forest floor. baculate. GYNOECIUM: compressed obconic to cylindric, ETYMOLOGY: Greek rhodon (rose, roseate) and spathê ovary (1–)2-locular, ovules 4–many per locule, anatropous, (spathe); refers to the spathe colour of some species. arranged in 2 rows, funicles long, placenta basal, stylar TAXONOMIC ACCOUNTS: Engler & Krause (1908), Croat region well-developed, usually broader than ovary and trun- (1978). cate, stigma elliptic to punctiform. BERRY: obovoid, truncate apically, locules 3–many-seeded, white, orange (S. ulei) to reddish orange, or yellow. SEED: clavate to ellipsoid, raphe C 20. Stenospermation prominent, testa thickish, smooth, embryo axile, elongate, endosperm copious. See Plates 20, 110C. Stenospermation Schott, Gen. Aroid. t. 70 (1858). LECTO- CHROMOSOMES: 2n = 28. TYPE: S. mathewsii Schott (see Nicolson 1967). DISTRIBUTION: ca. 36 spp.; tropical America:– Bolivia, Brazil (Amazonia, Atlantic region), Colombia, Costa Rica, Ecuador, SYNONYM: [Stenospermatium Schott in Oesterr. bot. French Guiana, Guatemala, Guyana, Nicaragua, Panama, Zeitschr. 9: 39 (1859), orth. var.] Peru, Surinam, Venezuela. Trichosclereids abundant. HABIT: epiphytic, climbing ECOLOGY: tropical humid forest, especially cloud forests; hemiepiphytic or terrestrial evergreen herbs, stem rather epiphytes or terrestrial on forest floor. densely leaved, erect, often elongated. LEAVES: many. PETI- NOTES: Vestigial tepals occasionally present (S. ulei). OLE: geniculate apically, sheath long. BLADE: ETYMOLOGY: Greek stenos (narrow), sperma, spermatos oblong-elliptic or lanceolate, oblique, often rather thick, (seed) and -ion (diminutive); refers to the slender seeds. venation usually obscured; midrib narrowly sulcate above, TAXONOMIC ACCOUNT: Engler & Krause (1908), Pérez de primary lateral veins weakly or not differentiated, pinnate, Gómez (1983). 128 T H E G E N E R A O F A R A C E A E

E F G D H B CA KJ Plate 20. Stenospermation. A, habit, inflorescence removed × 1/2; B, inflorescence × 1/2; C, gynoecium, longitudinal section; D, habit × 1/2; E, detail of leaf venation × 5; F, detail of spadix × 5; G, flower × 10; H, gynoecium, longitudinal section × 10; J, infructescence × 1; K, habit × 1/2. Stenospermation rusbyi: A–B, Rusby 2609 (K); S. ulei: C, E–H, Cult. Kew 1990–2738; D, Ule 8490 (K); S. angustifolium: J–K, Gómez et al. 20533 (K). M O N S T E R O I D E A E : S T E N O S P E R M A T I O N 129

C V. Subfamily Lasioideae Subfamily Lasioideae Engler in Nova Acta Acad. Leopold.- Carol. 39: 144 (1876); Hay (1992). Laticifers and trichosclereids absent; terrestrial or rooted aquatics, stem tuberous or rhizomatous, usually geophytic (except Lasia, Podolasia); petiole usually aculeate or warty or with striking coloration, long, usually ± geniculate apically; primary lateral veins of leaf blade divisions pinnate to arcu- ate-parallel, higher order venation reticulate; spadix flowering and fruiting in basipetal sequence; flowers bisexual, perigo- niate (except Pycnospatha); tepals fornicate, ± truncate, free, in 2 or more whorls, stamens free (except Lasimorpha) with distinct filaments, anthers terminal, connective slender, pollen monosulcate; stigma capitate to subcapitate; embryo large, endosperm present (except Anaphyllum) and forming a thin but distinct layer. C 21. Dracontium Dracontium L., Sp. Pl. 967 (1753). LECTOTYPE: D. poly- 21. Dracontium C phyllum L. (Britton & Wilson 1923, p.130). subcampylotropous, funicle short, placenta axile to subbasal, SYNONYMS: Eutereia Rafinesque, Fl. Tell. 4: 12 (1838, stylar region as long or much longer than ovary, projecting “1836”); Echidnium Schott in Oesterr. bot. Wochenbl. 7: 62 beyond perigone, stigma small, subcapitate or slightly 3— (1857); Ophione Schott in Oesterr. bot. Wochenbl. 7: 101 lobed. BERRY: usually obpyramidal, bearing style rudiment, (1857); Chersydrium Schott in Oesterr. bot. Zeitschr. 15: 72 greenish to brown. SEED: rounded-reniform, a little com- (1865); Godwinia Seemann in J. Bot. 7: 314 (1869). pressed, testa verrucose to almost smooth, rather thick, embryo curved, endosperm present. See Plates 21, 110D. HABIT: usually seasonally dormant, often robust, sometimes CHROMOSOMES: 2n = 26 gigantic herbs, tuber hypogeal, depressed-subglobose, often DISTRIBUTION: 23 spp.: tropical America, West Indies:– bearing few to very numerous tubercles usually on upper Bolivia, Brazil (Amazonia, Central West), Colombia, Costa Rica, surface. LEAVES: usually solitary. PETIOLE: long, often ver- Ecuador, French Guiana, Guyana, Mexico, Nicaragua, Panama, rucose-asperate, sometimes bearing few to many longer, Paraguay, Peru, Puerto Rico, Surinam, Trinidad, Venezuela. prickle-like processes, covered with striking, transversely ECOLOGY: tropical moist and humid forest, rarely in savan- banded variegation, usually weakly geniculate at apex, nas (D. margaretae); geophytes on forest floor. sheathed only at very base. BLADE: dracontioid, i.e trisect, NOTES: Engler (1911) recognized 2 sections:– Sect. Dra- sometimes perforate, primary anterior division usually tri- contium, Sect. Godwinia; Sect. Echidnium is recognized by sect at middle and primary posterior divisions bisect below modern workers (G.H. Zhu pers. comm.). the middle, secondary and tertiary divisions further subdi- ETYMOLOGY: ancient name, Latin draco, dracontis (dragon, vided, ultimate lobes usually elliptic-acuminate, rarely linear snake) and suffix -ium (diminutive), referring to the similari- (D. margaretae); primary lateral veins of ultimate lobes pin- ties of the petiole markings to a snake; also perhaps from the nate, forming arching submarginal collective veins, higher Greek drakontion. order venation reticulate. INFLORESCENCE: 1(–2), preceded TAXONOMIC ACCOUNTS: Engler (1911), Bogner (1986a), by cataphylls, appearing before, with or after the leaf. Bunting (1986), Hay (1992a), Zhu (1996, 1997). PEDUNCLE: very short to long, epidermis similar to petiole. SPATHE: oblong, usually ± boat-shaped, erect, acute, cuspi- 22. Dracontioides date to acuminate, often fornicate, convolute basally, gaping above, sometimes pubescent within, marcescent to persistent, Dracontioides Engler in Pflanzenreich 48 (IV.23C): 36 usually dark brown-purple outside, more reddish-purple (1911). TYPE: D. desciscens (Schott) Engler (Urospatha within and often white at the base within. SPADIX: sessile to desciscens Schott). stipitate, shortly cylindric to ellipsoid, much shorter than spathe, apical flowers sometimes sterile or with enlarged, SYNONYM: Dracontium sect. Urospathopsis Engler in irregularly shaped, projecting structures. FLOWERS: bisexual, Bot. Jahrb. 5: 178 (1884). perigoniate; tepals 4–8, dilated apically, fornicate. STAMENS: usually 4–6, free, sometimes up to 19, usually longer than HABIT: unarmed herbs, sometimes robust (to 2m), rhizome tepals at anthesis, filaments somewhat dilated, ± compressed, subterranean, erect, sparsely branched, sometimes bearing a anthers longer than connective, connective slender, thecae few subterranean tubercles. LEAVES: several, ± erect. PETIOLE: oblong-ellipsoid, dehiscing by apical slit. POLLEN: extruded in a conglutinate mass, monosulcate, ellipsoid to oblong, medium-sized (mean 38 µm., range 29–48 µm.), exine fove- olate-fossulate to subreticulate, apertural exine rugulate or verruculate. GYNOECIUM: ovary ovoid, incompletely 1–6- locular, ovules 1 per locule, anatropous or 130 T H E G E N E R A O F A R A C E A E

H K RP J B T F C Q W D A G M L N SU EV Plate 21. Dracontium. A, habit × 1/10; B, detail of petiole × 2/4; C, rachis/petiole insertion × 2/3; D, tubercules at soil–level junction of petiole and tuber × 2/3; E, leaf segment × 1/2; F, leaf segment × 1/2; G, seedling × 1; H, partially buried inflorescence × 2/3; J, flower, lon- gitudinal section × 6; K, inflorescence × 1/5; L, inflorescence × 2/3; M, spadix × 1; N, detail of spadix × 3; P, flower, longitudinal section × 6; Q, spadix × 1; R, flower, longitudinal section × 6; S, leaf segment × 1/2; T, leaflet, transverse section × 1; U, infructescence × 1; V, berry, side view × 3; W, seed, side view × 4. Dracontium asperum: A–D, Cult. Kew 1977–5355; D. gigas: E, Cult. Bull 1878 (K) & Cult. Bull 1881 (K); D. spruceanum: F, Plowman 4490 (K); G, Plowman 4490 (Kew illustration collection); D. changuango: H, Boyce s.n.; J, Aristeguieta 12734 (Kew spirit collection 53986); D. prancei : K, Cult. Kew 1977–5372 (Kew slide collection); D. asperum: L, Cult. Kew 1979–3887 (Kew spirit collection 46575); M–P, Cult. Kew 1979–3887 (Kew spirit collection 29047.455); D. soconuscum: Q–R, Cult. Kew 1980–1628 (Kew spirit collection 51365); D. margaretae: S, Emmerich 4053 (K illustration collection); T–W, Emmerich 4053 (Kew spirit collection 29047.358 & 43972). L A S I O I D E A E : D R A C O N T I U M 131

F G K H L DJ C E AB Plate 22. Dracontioides. A, habit × 1/8; B, seedling × 1/3; C, perforate leaf × 2/3; D, eperforate leaf × 2/3; E, base of plant showing erect rhizome and leaf–axillary tubercles × 2/3; F, inflorescence × 2/3; G, inflorescence, nearside half of spathe removed × 1; H, detail of spadix × 5; J, flower, longitudinal section × 8; K, infructescence × 2/3; L, seed, side view × 3. Dracontioides desciscens: A, Harley et al. 18009 (Kew slide collection); B, Cult. Kew 1977–640 (Kew spirit collection 39121); C, Harley et al. 18009 (K); D, Lewis & de Carvalho 1060 (K); E, Harley et al. 18241A (Kew spirit collection 47695) & Lewis & de Carvalho 1060 (K); F–L, Harley et al. 18009 (Kew spirit collection 46561, 46564 & 47699). 132 T H E G E N E R A O F A R A C E A E

23. Anaphyllopsis C Anaphyllopsis A. Hay in Aroideana 11 (1): 25–31 (1989, 22. Dracontioides “1988”). TYPE: A. americana (Engler) A. Hay HABIT: seasonally dormant (A. americana), solitary herbs, long, smooth to roughened-verruculate, often transversely var- rhizome hypogeal, erect. LEAVES: solitary, rarely 2. PETI- iegated, geniculate at apex, sheath less than half petiole length. OLE: unarmed, smooth to tubercular, geniculate apically. BLADE: deeply sagittate to subtripartite, often with a few per- BLADE: pinnatifid and perforate or pinnatisect, juvenile forations of irregular size between primary lateral veins; basal leaves entire; basal ribs well-developed, primary lateral veins ribs very well-developed, primary lateral veins mostly arising of ultimate lobes pinnate, higher order venation reticulate. near petiole insertion, very long-arcuate towards apex of each INFLORESCENCE: solitary, rarely 2. PEDUNCLE: long, simi- division, running into margin, higher order venation reticulate. lar to petiole in colour and texture. SPATHE: membranaceous, INFLORESCENCE: solitary, appearing with leaves. PEDUN- convolute basally, spirally twisted apically, marcescent, CLE: shorter than petiole. SPATHE: marcescent, tube with papery when dry. SPADIX: stipitate, stipe mostly adnate to convolute margins, longitudinally white-striped, blade strongly spathe, flowering sequence basipetal. FLOWERS: bisexual, fornicate, usually obscuring mouth of tube, brown-purple. perigoniate; tepals 4, fornicate. STAMENS: 4, free, filaments SPADIX: sessile to shortly stipitate, cylindric, obtuse, shorter short, linear, connective slender, thecae dehiscing by short, than spathe tube, flowering sequence basipetal. FLOWERS: apical pore-like slit. POLLEN: monosulcate, ellipsoid, bisexual, perigoniate; tepals 4, fornicate, subtruncate. STA- medium-sized (31 µm.), exine foveolate, apertural exine psi- MENS: 4, free, strongly exserted from flower at anthesis, late. GYNOECIUM: ovary 1–locular, ovules 1–2, anatropous, anthers longer than connective, connective slender, thecae placenta basal, stylar region ± attenuate, stigma subcapitate. ovate-ellipsoid, dehiscing by pore-like apical slit. POLLEN: BERRY: ovoid to obpyramidal, reddish. SEED: campy- monosulcate, ellipsoid-oblong, small (mean 23 µm.), exine lotropous, testa thick, verruculose to channelled, embryo subreticulate. GYNOECIUM: ovate-conoid, ovary 2-locular, curved, endosperm present. See Plates 23, 111B. ovules 1 per locule, anatropous, placenta axile, stylar region CHROMOSOMES: 2n = 26. attenuate, longer than tepals, stigma small, button-like. BERRY: DISTRIBUTION: 3 spp., tropical South America:– Brazil obovoid, somewhat furrowed, 1–2-seeded, dark purplish-red. (Amazonia), French Guiana, Surinam, Venezuela. SEED: reniform, attenuate towards micropyle, testa strongly ECOLOGY: tropical humid swamp forest; helophytes in dentate-cristate, thick, hard, brown, embryo curved, sandy, partially flooded places along streams, swamp forest. endosperm present. See Plates 22, 111A. ETYMOLOGY: Anaphyllum and Greek opsis (appearance); CHROMOSOMES: 2n = 26. i.e. “like Anaphyllum”. DISTRIBUTION: 1 sp.; Brazil (Atlantic region); the record TAXONOMIC ACCOUNTS: Hay (1989, 1992a). from Tijuca, Rio de Janeiro (Peyritsch 1879) is almost certainly a mistake. 23. Anaphyllopsis ECOLOGY: tropical humid forest; helophytes in swamp for- est, marshes, stream margins, open or shaded sites, in peat or sand. ETYMOLOGY: Dracontium and Greek suffix -oides (resem- blance), i.e. similar to Dracontium. TAXONOMIC ACCOUNTS: Mayo (1978), Hay (1992a). L A S I O I D E A E : A N A P H Y L L O P S I S 133

B F E D A C K H L JG Plate 23. Anaphyllopsis. A, leaf × 1/4; B, gynoecium, longitudinal section × 5; C, leaf × 1/4; D, infructescence × 2/3; E, immature fruit, longitudinal section × 6; F, seed, side view × 4; G, leaf × 1/4; H, base of plant showing erect rhizome × 1/2; J, inflorescence, nearside half of spathe removed × 2/3; K, detail of spadix × 4; L, flower, nearside tepal removed × 6. Anaphyllopsis pinnata: A–B, Wessels Boer 2387 (U); A. cururuana: C–F, Anderson 10627 (NY); A. americana: G, J, Leprieur 152 (P); H, Aroideana 11(1) 30, fig. 3 (1988) (as A. cururuana); K–L, Mélinon 52 (P). 134 T H E G E N E R A O F A R A C E A E

C 24. Pycnospatha 25. Anaphyllum C Pycnospatha Thorel ex Gagnepain in Bull. Soc. Bot. France Anaphyllum Schott in Bonplandia 5: 126 (1857). TYPE: A. 88: 511 (1941). TYPE: P. palmata Thorel ex Gagnepain wightii Schott HABIT: seasonally dormant herbs with subglobose tuber. HABIT: evergreen herbs, clump- or colony-forming, rhi- LEAVES: 1–2. PETIOLE: smooth, rough or aculeate, mottled, zome creeping. LEAVES: solitary to few. PETIOLE: smooth sheath very short, inconspicuous. BLADE: dracontioid, i.e. to tuberculate, geniculate apically, sheath short. BLADE: trisect, anterior division trifid, segments simple to pinnatifid, sagittate-hastate to pedatifid when juvenile, trisect at matu- posterior divisions bifid to pedatifid, segments then simple to rity, anterior division remotely pinnatisect with ± pinnatifid, ultimate lobes decurrent, ovate-elliptic to triangu- oblong-lanceolate, acute lobes, the upper ones decurrent, lar, acute to acuminate; primary lateral veins pinnate, running posterior divisions either ± oblong-lanceolate or deeply into distinct marginal vein, higher order venation reticulate. divided into 3 coherent segments, rachis geniculate at inser- INFLORESCENCE: solitary, appearing before or with leaf. tion of anterior and posterior divisions; basal ribs PEDUNCLE: much shorter than petiole, similar in appearance well-developed, primary lateral veins of ultimate lobes pin- and texture to petiole. SPATHE: margins not overlapping, nate, running into marginal vein, higher order venation strongly fornicate, thick, ± purple, marcescent. SPADIX: much reticulate. INFLORESCENCE: solitary. PEDUNCLE: very long shorter than spathe, conic to ovoid-conoid, stipitate, fertile to and slender, similar in colour and texture to petiole. SPATHE: apex, flowering sequence basipetal. FLOWERS: bisexual, membranous to coriaceous, marcescent, either convolute perigone absent. STAMENS: ca. 6 or more per flower, free, basally and becoming spirally twisted and long-acuminate crowded densely together with those of neighbouring flow- apically, or oblong-ovate, ± flat and fully expanded. SPADIX: ers, filaments oblong, flattened, connective slender, thecae cylindric, much shorter than spathe, stipitate or sessile, flow- oblong-ellipsoid, dehiscing by apical, pore-like slit. POLLEN: ering sequence basipetal. FLOWERS: bisexual, perigoniate, monosulcate, ellipsoid to oblong, medium-sized (mean 34 tepals 3–4, fornicate. STAMENS: 3–5, free, filaments fairly µm.), exine subreticulate to rugulate. GYNOECIUM: elon- wide, connective slender, thecae ellipsoid, dehiscing by gated-flask-shaped, ovary 1-locular, ovule 1, anatropous to short, apical, pore-like slit. POLLEN: monosulcate, ellipsoid, hemianatropous, funicle short, placenta basal to subparietal, medium-sized (mean 29 µm., range 25–33 µm.), exine foveo- stylar region greatly elongated, straight or somewhat curved, late to reticulate, sometimes with elevated psilate regions, projecting well beyond stamens, stigma small, scarcely or no apertural exine psilate. GYNOECIUM: ovary ovoid, 1-locu- broader than style. BERRY: globose, pericarp densely covered lar, ovule 1, hemianatropous, funicle short, placenta parietal with conic prickles, with conspicuous, persistent style rem- on single intrusive septum, stylar region thick, attenuate, nant, dark green. SEED: reniform, black or very dark brown, stigma subcapitate, exuding droplet at anthesis. BERRY: testa hard, thick, verrucose, containing white druses, embryo ovoid, smooth, red. SEED: ovoid, funicle slender, testa mem- large, ellipsoid to slightly curved, endosperm very sparse, branaceous, smooth, embryo stout, straight, endosperm only a few cell layers thick. See Plates 24, 111C. absent. See Plates 25, 111D. CHROMOSOMES: 2n = 26. CHROMOSOMES: 2n = 26. DISTRIBUTION: 2 spp.; tropical southeast Asia:– Laos, DISTRIBUTION: 2 spp.; southern India (Kerala, Tamil Nadu). Thailand, Vietnam. ECOLOGY: tropical evergreen forest in leaf litter and in ECOLOGY: tropical humid forest; geophytes on forest floor swamp forest undergrowth; geophytes, rare. in sandy loam. ETYMOLOGY: Greek ana- (up, erect) and phyllon (leaf). ETYMOLOGY: Greek pyknos (compact, thick) and spathê TAXONOMIC ACCOUNTS: Engler (1911), Sivadasan (1982), (spathe). Hay (1992a). TAXONOMIC ACCOUNTS: Bogner (1973c), Hay (1992a), Boyce (1993b). 24. Pycnospatha 25. Anaphyllum L A S I O I D E A E : A N A P H Y L L U M 135

A C B Q P D F H G LM N E J K Plate 24. Pycnospatha. A–E, leaf developmental sequence from first leaf (A) to flowering size (E) × 1/3; F, detail of petiole × 1; G, habit × 1/8; H, inflorescence × 2/3; J, spathe sectioned to show spadix × 1; K, detail of spadix × 3; L, flower × 6; M, gynoecium, longitudinal sec- tion × 6; N, gynoecium, transverse section × 6; P, infructescence × 2/3; Q, seed, side view × 5. Pycnospatha arietina: A–F, Bogner 395 (K); G, Cult. Kew 1971–1039 (Kew slide collection 8021); H–J, Bogner 395 (Kew spirit collection 34429); K–Q, Bogner 395 (Kew spirit collection 29047.184). 136 T H E G E N E R A O F A R A C E A E

H A G J D B E CF Plate 25. Anaphyllum. A, habit × 1/8; B, leaf × 1/3; C, base of plant × 2/3; D, inflorescence × 2/3; E, detail of spadix × 5; F, flower, longi- tudinal section × 6; G, seed, hilum view × 3; H, leaf × 1/3; J, inflorescence × 2/3. Anaphyllum wightii: A–B, Cult. Kew 1984–04519; Fliegner 245 (Kew spirit collection 49762 & 51853); C, Sivadasan CU 9060 (K); D–F, Cult. Kew 1983–02560; Hay s.n. (Kew spirit collection 49796); G, Cult. Kew 1984–04519; Fliegner 245 (Kew spirit collection 51853); A. beddomei: H, Anon. s.n. (K); J, Blasco s.n. (K). L A S I O I D E A E : A N A P H Y L L U M 137

C 26. Cyrtosperma (Borneo, Irian Jaya, Java, Palau Is., Sumatra), Malaysia (Borneo, Peninsula), Mariana Is., Marquesas Is., Marshall Is., Papua New Cyrtosperma Griffith, Notul. Pl. Asiat. (Posthum. Pap.) 3: Guinea, Philippines, Samoa, Singapore, Society Is., Solomon 149 (1851); Icon. pl. Asiat. 3, t.169 (1851). TYPE: C. lasioides Is., Tahiti, Vanuatu, Vietnam (cultivated?). Griffith (“lacioides”), [= C. merkusii (Hassk.) Schott]. ECOLOGY: tropical humid forest, swamp forest, open swamps and cultivated areas; helophytes in streams, ponds SYNONYM: Arisacontis Schott in Bonplandia 5: 129 (1857). and other wet places. NOTES: Hay (1988) recognized 4 informal groups:– “Mer- HABIT: slender to gigantic evergreen herbs, usually solitary, kusii” group, “Cuspidispathum” group, “Carrii” group, sometimes clump-forming, rhizome thick, condensed, creep- “Macrotum” group. ing. LEAVES: several. PETIOLE: sometimes very long, aculeate, ETYMOLOGY: Greek kyrtos (curved) and sperma (seed); the geniculate apically, sheath short. BLADE: deeply sagittate, seeds are strongly curved. hastate-sagittate or ± tripartite (posterior divisions usually TAXONOMIC ACCOUNTS: Engler (1911), Thompson (1982), larger than anterior), veins sometimes aculeate on lower sur- Hay (1988, 1992a). face; basal ribs very well-developed, primary lateral veins pinnate, running into marginal vein, higher order venation reticulate. INFLORESCENCE: 1–2 in each floral sympodium, appearing with the leaves. PEDUNCLE: long, similar to peti- 27. Lasimorpha C oles. SPATHE: marcescent, erect, blackish purple to white, convolute or not in lower part, upper part rarely somewhat Lasimorpha Schott in Bonplandia 5: 127 (1857). TYPE: L. fornicate, long-acuminate and twisted in some species. senegalensis Schott (syn. Cyrtosperma senegalense (Schott) SPADIX: sessile or stipitate. FLOWERS: bisexual, perigoniate; Engler). tepals 4–6, somewhat thickened at apex, fornicate. STAMENS: 4–6, free, filaments free, flat and broad, connective slender, SYNONYM: [Lasiomorpha Engler in Engler, Pflanzen- thecae oblong-ovate, dehiscing by apical slit. POLLEN: mono- reich 48 (IV.23C): 14 (1911), orth. var.]. sulcate, ellipsoid, medium-sized (mean 29 µm., range 28–30 HABIT: robust to gigantic evergreen herbs forming large µm.), exine foveolate, apertural exine psilate. GYNOECIUM: colonies, rhizome short, thick, hypogeal, vigorously stoloni- 1-locular, ovules 1–many, campylotropous to subam- ferous. LEAVES: several. PETIOLE: very long, 4–6 angled, phitropous, placenta basal to parietal, stylar region short or aculeate along angled ridges, weakly geniculate at apex, inconspicuous, stigma subhemispheric, exuding droplet at sheath short. BLADE: deeply sagittate, sometimes weakly anthesis. BERRY: obovoid, 1–7-seeded, usually red when hastate, to over 1m long, coriaceous, ± erect; basal ribs mature, bearing remnant of stigma (in C. cuspidispathum ripe well-developed, primary lateral veins pinnate, long arcuate berries are extruded and dangle on strips of tepal epidermis; towards apex of each leaf division and running into mar- A. Hay pers. comm.). SEED: reniform to orbicular to helically ginal vein, higher order venation reticulate. twisted, cristate, warty or smooth, embryo curved, endosperm INFLORESCENCE: solitary, appearing with leaves. PEDUNCLE: present. See Plates 26, 112A. subequal to petiole and similar in appearance. SPATHE: CHROMOSOMES: 2n = 26. erect, ovate, ± convolute in lower third, gaping at anthesis, DISTRIBUTION: 11–12 spp.; tropical southeast Asia, Malay striped purple and yellow within, persistent. SPADIX: ses- Archipelago, Melanesia, Pacific:– Brunei, Caroline Is., China sile to stipitate, cylindric, obtuse, shorter than spathe, (cultivated?), Cook Is., Fiji, Gilbert & Ellice Is., Guam, Indonesia flowering sequence basipetal. FLOWERS: bisexual, perigo- 26. Cyrtosperma 27. Lasimorpha 138 T H E G E N E R A O F A R A C E A E

G B A NK H D M E J CF L Plate 26. Cyrtosperma. A, inflorescence × 2.3; B, flower, longitudinal section × 8; C, infructescence × 2/3; D, seed, side view × 6; E, leaf × 2/3; F, base of plant × 2/3; G, habit × 1/8; H, leaf × 2/3; J, base of petiole × 2/3; K, inflorescence × 2/3; L, detail of spadix × 4; M, flower, lon- gitudinal section × 8; N, infructescence × 2/3. Cyrtosperma cuspidispathum: A–B, NGF 10241 (K & Kew spirit collection 18946); C–D, Hay s.n. (Kew spirit collection 45943); C. beccarianum: E, Reksodihardjo 412 (K); F, van Royen 4792 (K); C. macrotum: G, Aroids of Papua New Guinea pl. x, 3 (1990); C. merkusii: H, Lobb s.n. (K); J, Nicolson 1217 (K); L–M, Bogner 1510 (Kew spirit collection 45187); N, Reksodihardjo 311 (K). L A S I O I D E A E : C Y R T O S P E R M A 139