4. ARRABIDAEA A. DC., Biblioth. Universelle Genéve ser 2, 17: 10. 1838.

Lianas or rarely shrubs or small trees, the branches terete, usually with interpetiolar glandular fields; pseudostipules small and inconspicuous. Leaves usually 3-foliolate, rarely ternate or bicompound; terminal leaflet often replaced by a simple tendril. Inflorescence usually a large panicle, terminal or axillary, sometimes reduced to few flowers. Calyx usually cupular, truncate or bilabiate, usually minutely 5-denticulate, pubescent; corolla magenta to red, purple, or white, tubular-campanulate to tubular-funnel-shaped or subsalverform, pubescent outside at least on the lobes. Anthers glabrous, the thecae straight or slightly curved, usually divaricate. Ovary narrowly cylindric to linear-tetragonal, usually minutely lepidote; ovules 2-seriate; disk cupular-pulviniform. Fruit a linear capsule, flattened, the valves parallel to the septum, usually smooth, rarely verrucose, usually with the midline slightly raised. Seeds 2-winged, the wings usually ± hyaline and sharply demarcated from seed body.

Neotropics from Mexico and the West Indies to Argentina; 70 species, 25 in Venezuela, 24 of these in the flora area.

Key to the Species of Arrabidaea

1. Leaves mostly bicompound; flowers magenta ..... A. inaequalis

1. Leaves simple or 2- or 3-foliolate (very rarely bicompound in part in A. bilabiata with white flowers); flowers of various colors ..... 2

2(1). Seeds with the wings reduced and woody or essentially absent; capsule slightly viscid and > 15 mm wide; inflorescence mostly axillary, usually with few flowers; calyx either > 15 mm long and strongly bilabiate or tiny (less than 2 mm long) ..... 3

2. Seed wings hyaline-membranous, at least the tips; capsule not viscid (except in A. lachnaea where less than 14 mm wide); inflorescence mostly terminal, usually multiflowered; calyx 3-11 mm long, very slightly or not at all bilabiate ..... 6

3(2). Leaves generally simple; calyx cupular, subtruncate, 2 mm long; fruit less than 9 × 1 cm ..... A. carichanensis

3. Leaves all 2- or 3-foliolate; calyx campanulate, bilabiate, > 15 mm long; fruit > 10 × 1.5 cm ..... 4

4(3). Capsule smooth, not viscid; seeds woody, essentially wingless; flowers white or cream, scarcely flattened dorsally, the mouth of tube open ..... A. bilabiata

4. Capsule viscid, the surface often slightly roughened; seeds with membranous wings, the bases brown, the apices hyaline; flowers magenta or wine-colored with white throat, dorsiventrally compressed, the mouth of tube almost closed ..... 5

5(4). Leaves and branchlets hispid ..... A. egensis

5. Leaves and branches puberulous to glabrate ..... A. japurensis

6(2). Leaves usually pubescent with dendroid trichomes [if ± glabrescent, the leaflets palmately 3(-5)-veined with the basal or subbasal lateral vein pair strongly ascending; capsule without prominent midline] ..... 7

6. Leaves with uniformly simple trichomes, at least in vein axils, the leaflets pinnately veined; capsule with or without raised midline ..... 11

7(6). Leaflets ± 3-veined from near base, the basal or subbasal vein pair strongly ascending; leaves simple or 2-foliolate, never 3-foliolate; corolla funnel-shaped-campanulate to almost salverform, either the corolla white or calyx and inflorescence grayish when dry ..... 8

7. Leaflets pinnately veined; leaves mostly 3-foliolate, never simple; corolla tubular-campanulate, magenta; calyx and inflorescence brownish to black when dry ..... 9

8(7). Corolla magenta, funnel-shaped-campanulate, less than 4 cm long; calyx less than 6 mm long; leaves usually with branched trichomes at least in vein axils below ..... A. brachypoda

8. Corolla white, nearly salverform, > 4.5 cm long; calyx > 7 mm long; leaflets glabrous even in vein axils ..... A. prancei

9(7). Capsule margins rounded; calyx and inflorescence black when dry; vegetative trichomes reddish ..... A. mollis

9. Capsule margins sharply angled, often subwinged; calyx and inflorescence brown to blackish when dry; vegetative trichomes usually short or tannish (or both) ..... 10

10(9). Calyx 8-15 mm long, drying brownish or tannish; leaves drying olive to tannish; capsule mostly > 2 cm wide ..... A. cinnamomea

10. Calyx 3-6 mm long, drying dark brown to blackish; leaves drying dark gray to blackish; capsule less than 2 cm wide ..... A. nigrescens

11(6). Calyx obtusely 5-lobed half way to base; corolla with conspicuous glandular pubescence outside; capsule glandular-pubescent ..... A. lachnaea

11. Calyx truncate to subbilabiate, usually minutely 5-denticulate; corolla puberulous with nonglandular trichomes outside; capsule without glandular pubescence ..... 12

12(11). Capsule valves usually without noticeably raised midline, softly puberulous or glandular-punctate with the margins acutely angled; corolla usually > 3.5 cm long, the tube pubescent outside; calyx usually dark with the margin lighter or thinner when dry ..... 13

12. Capsule valves with ± prominent midline, glabrous or lepidote, not glandular-punctate, the margins rounded; corolla usually less than 4 cm long (if longer, the tube essentially glabrous); calyx margin not noticeably different in color or texture from rest of calyx ..... 14

13(12). Capsule valves glabrous; calyx irregularly truncate to subbilabiate, irregularly denticulate, the margin thinner but not evenly white-fringed ..... A. corallina

13. Capsule valves pubescent; calyx uniformly 5-denticulate with a white marginal fringe ..... A. mollissima

14(12). Corolla actinomorphic, salverform-tubular, at least the lobes red, tube > 3 times as long as wide; leaves usually lead gray when dry, especially on the upper surface, the lower surface noticeably pubescent ..... 15

14. Corolla ± bilabiate, campanulate or funnel-shaped, magenta to white, tube less than 3 times as long as wide; leaves rarely grayish when dry, and only when the lower surface is essentially glabrous ..... 16

15(14). Corolla > 2.5 cm long, the tube > 2 cm long and almost glabrous; leaflets puberulous on lower surface only along veins with long, stiff, straight trichomes ..... A. grosourdyana

15. Corolla less than 2 cm long, the tube less than 1.5 cm long and densely puberulous; leaflets uniformly and densely puberulous on lower surface with short subappressed trichomes ..... A. traillii

16(14). Corolla tube glabrous outside; buds dark with a conspicuously white apex from the lobe pubescence; calyx reflexed from corolla base; leaves simple or 2-foliolate ..... A. patellifera

16. Corolla tube pubescent outside; buds not bicolored; calyx cupular, enclosing corolla base; leaves 3-foliolate in part, rarely simple ..... 17

17(16). Lower surface of leaflets conspicuously whitish from the dense short tomentum, the main veins (even in juvenile leaves) less puberulous and conspicuously darker than surface ..... 18

17. Leaflets variably pubescent or glabrate but the lower surface never conspicuously whitish with contrastingly darker main veins ..... 19

18(17). Corolla > 22 mm long, the tube > 8 mm wide ..... A. candicans

18. Corolla less than 11 mm long, the tube ca. 2 mm wide ..... A. fanshawei

19(17). Leaflets (including veins) uniformly and densely tannish puberulous below; corolla less than 18 mm long; capsule usually less than 9 cm long ..... A. pubescens

19. Leaflets glabrescent or pubescent mostly along the main veins below (if in part tannish puberulous, then the corolla > 22 mm long); mature capsule usually > 9 cm long ..... 20

20(19). Leaves drying brick-red, or at least with patches of this color, usually glabrous except in lateral vein axils below; interpetiolar glandular fields present ..... A. chica

20. Leaves not reddish when dry, mostly puberulous at least along the main veins; interpetiolar glandular fields present or absent ..... 21

21(20). Mature branchlets with loose, conspicuously wrinkled epidermis and inconspicuous lenticels; young branchlets drying black with contrasting pale lenticels; inflorescence contracted, 3-7(-25)-flowered, including buds) ..... A. oxycarpa

21. Mature branchlets with tight, finely and evenly striate epidermis with noticeable small lenticels; young branchlets drying grayish or brownish; inflorescence open, many-flowered ..... 22

22(21). Corolla less than 2 cm long; calyx grayish when dry, eglandular, 3-4 mm long; capsule when dry blackish, 0.8-1 cm wide; seed wings hyaline-membranous ..... 23

22. Corolla > 2 cm long; calyx whitish gray or brownish when dry, sometimes with submarginal glands, 4-9 mm long; capsule when dry brownish, > 1 cm wide at maturity; seed wings with the base brown and the apices hyaline ..... 24

23(22). Corolla tube ca. 2 mm wide ..... A. fanshawei

23. Corolla tube > 4 mm wide ..... A. florida

24(22). Calyx campanulate, 7-9 mm long, lepidote and sparsely puberulous, drying brownish, eglandular; capsule without raised midline ..... A. selloi

24. Calyx cupular, 4-7 mm long, densely puberulous, grayish white when dry, usually with conspicuously darker submarginal glands; capsule with prominent midline ..... A. tuberculata

Arrabidaea bilabiata (Sprague) Sandwith, Kew Bull. 22: 418. 1968. -Memora bilabiata Sprague, Bull. Herb. Boissier ser. 2, 6: 375. 1906.

Liana. Seasonally flooded or swamp forests, 50-300 m; Delta Amacuro (Santa Catalina), widespread in Bolívar and Amazonas. Apure, Guárico, Monagas; Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil, and Bolivia. Fig. 359.

Arrabidaea brachypoda (A. DC.) Bureau, Vidensk. Meddel. Naturhist. Foren. Kj¯benhavn 1893: 98. 1894. -Bignonia brachypoda A. DC., Prodr. 9: 145. 1845.

Liana. Seasonally dry forests, 50-400 m; Delta Amacuro (near Los Castillos), northern Bolívar. Carabobo, Cojedes, Guárico, Portuguesa; wide-ranging in dry areas of Peru, Brazil, Bolivia, and Paraguay.

Arrabidaea candicans (Rich.) A. DC., Prodr. 9: 185. 1845. -Bignonia candicans Rich., Actes Soc. Hist. Nat. Paris 1: 110. 1792.

Liana. Evergreen lowland to montane forests and open white-sand forests, 100-1200 m; Bolívar (eastern portions from the Serranía de Imataca to the Gran Sabana). Widespread elsewhere in Venezuela; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay.

Arrabidaea carichanensis (H.B.K.) Bureau & K. Schum. in Mart., Fl. Bras. 8(2): 62. 1896. -Bignonia carichanensis H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 137. 1818 [1819].

Bignonia verrucosa H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 137. 1818 [1819].

Shrub or vine. Seasonally flooded forests, 50-200 m; Bolívar (along Río Orinoco), Amazonas (Río Orinoco near Puerto Ayacucho). Anzoátegui, Apure, Guárico; adjacent Colombia.

Arrabidaea chica (Bonpl.) B. Verl., Rev. Hort. 40: 154. 1868. -Bignonia chica Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 107. 1808 [1807].

Liana. Nonflooded evergreen lowland to montane forests, 100-1000 m; Delta Amacuro (near Los Castillos), Bolívar (widespread), scattered in northern Amazonas. Widespread elsewhere in Venezuela; Mexico and the Antilles to northern Argentina. Fig. 361.

Arrabidaea cinnamomea (A. DC.) Sandwith, Candollea 7: 248. 1936. -Bignonia cinnamomea A. DC., Prodr. 9: 164. 1845.

Liana. Nonflooded evergreen lowland forests, 50-200 m; Bolívar (near Caicara), Amazonas (Río Mawarinuma). Amazonian Colombia, French Guiana, Amazonian Peru, Brazil, and Bolivia.

Arrabidaea corallina (Jacq.) Sandwith, Kew Bull. 1953: 460. 1954. -Bignonia corallina Jacq., Fragm. Bot. 37, t. 42. 1800-1809.

Liana. Seasonally dry forests, 100-300 m; northern Bolívar. Widespread elsewhere in Venezuela; Mexico to northern Argentina.

Arrabidaea egensis Bureau & K. Schum. in Mart., Fl. Bras. 8(2): 65. 1896.

Liana. Forest edges and slopes, 100-600 m; Amazonas (Cerro Aracamuni, Puerto Ayacucho, San Carlos de Río Negro). Amazonian Colombia, Guyana, Amazonian Colombia, Ecuador, Peru, northern Brazil.

It is possible that this is no more than a pubescent form of Arrabidaea japurensis.

Arrabidaea fanshawei Sandwith, Bull. Torrey Bot. Club 75: 662. 1948.

Liana. Nonflooded evergreen lowland to lower montane forests, 200-800 m; Delta Amacuro (Serranía de Imataca), Bolívar (near Kilómetro 88, Serranía de Imataca, Serranía Marutaní). Colombia, Guyana, Suriname, French Guiana, Peru.

Arrabidaea florida A. DC., Prodr. 9: 184. 1845.

Liana. Nonflooded evergreen lowland to lower montane forests, 100-800 m; Bolívar (Amaruay-tepui, El Mato on Río Caura), Amazonas (Culebra). Apure, Barinas, Cojedes, Distrito Federal, Mérida, Miranda, Portuguesa, Táchira, Yaracuy, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay.

Arrabidaea grosourdyana (Baill.) Sandwith, Kew Bull. 22: 418. 1968. -Paramansoa grosourdyana Baill., Hist. Pl. 10: 27. 1888.

Tecoma moritziana Kraenzl., Feddes Repert. Sp. Nov. Regni Veg. 17: 218. 1921. -Tabebuia moritziana (Kraenzl.) Schnee in Pittier et al., Cat. Fl. Ven. 2. 409. 1947.

Liana. Evergreen lowland to lower montane forests, 100-800 m; north-central Bolívar. Adjacent Colombia, Guyana, French Guiana.

This species is distinctive in its hummingbird-pollinated flowers.

Arrabidaea inaequalis (DC. ex Splitg.) K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(3b): 214. 1894. -Bignonia inaequalis DC. ex Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 1842.

Liana. Evergreen lowland to lower montane forests, 50-500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Gran Sabana, La Ceiba, Río Caroní, Río Cuyuní basin), Amazonas (widespread). Sucre; southern Mexico, Belize, Guatemala, northwestern Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia.

Arrabidaea japurensis (A. DC.) Bureau & K. Schum. in Mart., Fl. Bras. 8(2): 65. 1896. -Tabebuia japurensis A. DC., Prodr. 9: 214. 1845.

Liana. Evergreen lowland forests, 100-300 m; Delta Amacuro (Serranía de Imataca), northern Bolívar, Amazonas (widespread). Apure, Barinas, Táchira; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Fig. 358.

Arrabidaea lachnaea (Bureau ex Bureau & K. Schum.) Sandwith, Kew Bull. 22: 416. 1968. -Cremastus lachnaeus Bureau ex Bureau & K. Schum. in Mart., Fl. Bras. 8(2): 215. 1896.

Liana. Semideciduous forests, 50-100 m; northwestern Bolívar, northern Amazonas. Barinas; northeastern Brazil. Fig. 363.

Arrabidaea mollis (Vahl) Bureau ex K. Schum. in Engl. & Prantl, Pflanzenfam. 4(3b): 213. 1894. -Bignonia mollis Vahl, Eclog. Amer. 2: 46. 1798.

Liana. Evergreen lowland to lower montane forests, 100-500 m; Delta Amacuro (Serranía de Imataca), Bolívar (near Icabarú, Río Cuyuní basin, Serranía de Imataca, Tumeremo). Guyana, Suriname, French Guiana, northern Brazil.

Arrabidaea mollissima (H.B.K.) Bureau & K. Schum. in Mart., Fl. Bras. 8(2): 46. 1896. -Bignonia mollissima H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 133. 1818 [1819].

Liana. Semideciduous forests, 50-500 m; northern Bolívar. Widespread elsewhere in Venezuela; Mexico, Central America, Colombia. Fig. 357.

Arrabidaea nigrescens Sandwith, Lloydia 2: 209. 1939.

Liana. Nonflooded evergreen lowland to lower montane forests, 100-700 m; Bolívar (near El Dorado, Río Acanán basin, lower Río Caura), Amazonas (widespread). Colombia, Guyana, Suriname, French Guiana, northern Brazil. Fig. 362.

One collection (Berry 1657, MO, from San Juan de Manapiare) is intermediate between Arrabidaea nigrescens and A. cinnamomea. It differs from both in being indicated as a shrub.

Arrabidaea oxycarpa Urb., Repert. Spec. Nov. Regni Veg. 14: 301. 1916.

Liana. Semideciduous forests, 100-300 m; northern Bolívar, Amazonas (San Fernando de Atabapo, San Juan de Manapiare). Anzoátegui, Aragua, Cojedes, Guárico, Lara, Miranda, Nueva Esparta, Portuguesa, Sucre, Zulia; Panama, Colombia, Trinidad. Fig. 360.

Arrabidaea patellifera (Schltdl.) Sandwith, Kew Bull. 22: 413. 1968. -Bignonia patellifera Schltdl., Linnaea 8: 516. 1833.

Liana. Semideciduous and evergreen lowland to montane forests, 100-1000 m; Bolívar (scattered), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Yutajé, San Juan de Manapiare). Falcón, Mérida, Miranda, Táchira, Yaracuy, Zulia; Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

The 2-toned buds with white tips due to the corolla lobe pubescence are characteristic.

Arrabidaea prancei A.H. Gentry, Ann. Missouri Bot. Gard. 63: 50. 1976.

Liana. Evergreen lowland forests, 100-200 m; Amazonas (Sierra de la Neblina). Amazonian Peru and Brazil.

The long, narrow, almost salverform white flowers, presumably adapted to moth pollination, are unique in the genus.

Arrabidaea pubescens (L.) A.H. Gentry, Brittonia 25: 239. 1973. -Bignonia pubescens L., Sp. Pl. ed. 2, 2: 870. 1763.

Liana. Semideciduous forests, 100-300 m; northern Bolívar, Amazonas (Culebra, Puerto Ayacucho). Widespread in drier areas of northern Venezuela; Mexico, Central America, Colombia, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay.

Arrabidaea selloi (Spreng.) Sandwith, Kew Bull. 1953: 461. 1954. -Bignonia selloi Spreng., Syst. Veg. 2: 831. 1825.

Liana. Semideciduous forests, ca. 200 m; Bolívar (Serranía de Imataca). Falcón, Lara, Zulia; Colombia, southern Peru, Brazil, Bolivia, Paraguay, Argentina.

Arrabidaea traillii Sprague, Bull. Herb. Boissier ser. 2, 6: 372. 1906.

Liana. Somewhat seasonal, nonflooded evergreen lowland forests, 200-300 m; Bolívar (El Dorado to La Escalera). Guyana, Suriname, French Guiana, Peru, Amazonian Brazil, Bolivia.

The tiny, almost salverform, red flowers are unique in the genus.

Arrabidaea tuberculata A. DC., Prodr. 9: 184. 1845.

Liana. Semideciduous to evergreen lowland forests, 50-300 m; Bolívar (between Caicara and Ciudad Bolívar, Río Parguaza), Amazonas (Culebra, near Puerto Ayacucho). Apure; Colombia, Guyana, Suriname, French Guiana, Peru, Bolivia.

This species was included in Arrabidaea conjugata (Vell.) Mart. in Flora de Venezuela but differs from that species in the narrow elongate inflorescence, generally smaller calyx, and especially the thinner leaf texture with the lower surface of young leaflets usually conspicuously minutely puberulous.


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