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Flora of the Venezuelan Guayana

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by Alwyn H. Gentry

Trees, shrubs, or woody vines, rarely herbaceous; outer scales of axillary buds often pseudostipular and sometimes foliaceous; glandular fields often present between the petioles and rarely at the petiole apex. Leaves usually opposite, less often alternate or fasciculate, usually palmately or pinnately compound, sometimes simple; terminal leaflet often replaced by a tendril. Inflorescence terminal, axillary, or cauliflorous, paniculate, racemose, or reduced to a fascicle or a single flower. Calyx synsepalous, usually ± cupular, truncate to variously lobed or spathaceously split; corolla sympetalous with a conspicuous tube and 5 usually imbricate lobes. Stamens inserted on corolla tube, didynamous, (2)4(5), a posterior staminode usually present; anthers with (1)2 thecae, these usually divergent or divaricate and held against the adaxial side of corolla tube; filaments slender. Ovary superior, (1)2-locular; ovules numerous, attached to 4 vertical placental ridges, these axile in 2-locular ovaries and parietal in 1-locular ovaries (a single placental ridge in each locule in many species of Schlegelieae); style slender; stigma bilamellate; nectariferous disk usually conspicuous at base of ovary, sometimes lacking. Fruit a capsule with dehiscence perpendicular or parallel to the septum, or a berry or hard-shelled pepo (calabash). Seeds without endosperm, ± flattened, usually winged; cotyledons foliaceous.

Pantropics (concentrated in Neotropics), with a few temperate zone outliers; 113 genera and ca. 840 species, 38 genera and 131 species in the flora area.

The Bignoniaceae are predominantly woody and include several genera of large trees and many genera of lianas. Opposite, compound leaves are characteristic of many species in the family. The scaly buds ending in pseudostipules are frequently conspicuous and provide a useful taxonomic character. Presence or absence of zones of peltate glands at the nodes, or less frequently at the petiole apex is also useful; these glandular zones, however, disappear with age and do not occur in all species.

The major economic importance of the family is in horticulture, with many species cultivated for their showy flowers. Some species of Tabebuia and related genera are important lumber trees, and the hard shell of the fruit of Crescentia is widely used as a household utensil. Saritaea magnifica (Sprague ex Steenis) Dugand, which is native to Colombia and Ecuador, is cultivated in several states in Venezuela, including Bolívar.

For species descriptions, common names, and more extensive lists of synonyms for many of the species treated here, see Flora de Venezuela 8(4): 1-433. 1982.

Key to the Genera of Bignoniaceae
(see also: Key to the Genera of Bignoniaceae
based on fruiting material)

1. Hemiepiphytic lianas climbing by adventitious roots; leaves simple, without tendrils; corolla tubular (not funnel-shaped or campanulate) with reflexed lobes; fruit a globose berry ca. 3 cm diameter ..... 32. Schlegelia

1. Trees, shrubs, or lianas, not hemiepiphytic; leaves simple or compound (always compound and in part tendrillate when the plant scandent); corolla campanulate, tubular-funnel-shaped, or salverform; fruit indehiscent and > 5 cm long or dehiscent ..... 2

2(1). Calyx bilabiately split to base; corolla campanulate with a transverse fold in throat; flowers cauliflorous, borne singly or in a fascicle of 2 or 3 flowers; leaves in alternate fascicles; fruit indehiscent ..... 8. Crescentia

2. Calyx cupular or campanulate or spathaceously split; corolla without transverse fold in throat; inflorescence usually an axillary or terminal panicle or raceme, rarely cauliflorous; leaves opposite; fruit dehiscent ..... 3

3(2). Leaves palmately 5-7-foliolate, lacking tendrils; trees or shrubs ..... 4

3. Leaves simple, 2- or 3-foliolate, pinnately compound, or bicompound, frequently tendrillate; trees, shrubs, or lianas ..... 6

4(3). Flowers less than 1.8 cm long; corolla broadly campanulate from base; anthers pilose; fruit spirally twisted ..... 14. Godmania

4. Flowers > 2 cm long; corolla salverform to funnel-shaped or campanulate, broadly campanulate only when tube base narrowly cylindrical; anthers glabrous; fruit not spirally twisted ..... 5

5(4). Leaves (especially when young) and young shoots varnished; calyx membranous, 12-22 mm long, with thinner irregularly rupturing apex; corolla white or pale pink with purplish stripes in throat; seed wings replaced by terminal trichomes ..... 33. Sparattosperma

5. Leaves and young twigs not conspicuously varnished; calyx 4-40 mm long, the texture even throughout, usually coriaceous, apex often evenly denticulate; corolla color various, when white the throat uniformly white or yellow; seeds with membranous wings or wingless ..... 35. Tabebuia

6(3). Staminode longer than the stamens, subexserted, strongly glandular-villous; flowers blue or blue-purple; trees or shrubs ..... 7

6. Staminode minute, at base of corolla tube, not villous; flowers yellow, white, red, orange, magenta, purple, or lavender, never blue; trees, shrubs, or lianas ..... 8

7(6). Leaves simple or pinnate; calyx > 1 cm long ..... 11. Digomphia

7. Leaves bipinnate; calyx less than 0.7 cm long ..... 15. Jacaranda

8(6). Branchlets acutely tetragonal (sometimes subtetragonal in P. albiflora); tendrils trifid; leaves biternate to ternately bipinnate or tripinnate ..... 28. Pleonotoma

8. Branchlets terete to tetragonal, if tetragonal the tendrils simple; leaves mostly simple or 2-or 3-foliolate, biternate or bipinnate only in Arrabidaea and Memora, both with terete branches .....9

9(8). Corolla lobes narrow, not reflexed, appressed together at anthesis; calyx double with thick inner margin and wavy continuous outer margin ..... 2. Amphilophium

9. Corolla lobes reflexed at anthesis; calyx simple ..... 10

10(9). Corolla tube (and lobes except sometimes the margins) glabrous or lepidote outside ..... 11

10. Corolla puberulous outside (usually both tube and lobes, rarely only on lobes) ..... 26

11(10). Tendrils trifid-uncinate with claw-like apices; flowers yellow; stem cross section with many irregularly divided phloem arms ..... 12

11. Tendrils (when present) simple or 3- to multifid, the apices not uncinate or claw-like; flower color various; stem cross section with 4-16 radially symmetrical phloem arms or lacking anomalous structure ..... 13

12(11). Calyx apex obtuse or incurved; inflorescence bracts minute ..... 17. Macfadyena

12. Calyx with recurved apex; inflorescence bracts foliaceous ..... 21. Melloa

13(11). Calyx spathaceously split down one side to near the base, the tip somewhat recurved and forming a glandular hoodlike structure with sparse lepidote scales ..... 26. Phryganocydia

13. Calyx truncate or bilabiate, rarely spathaceous, but then not as above ..... 14

14(13). Leaves mostly pinnate or bipinnate ..... 22. Memora

14. Leaves simple or 2- or 3-foliolate ..... 15

15(14). Calyx large, > 2.2 × 1.2 cm, ± inflated ..... 16

15. Calyx small, less than 2.1 × 1.2 cm ..... 17

16(15). Tendril simple; calyx thick and spongy when fresh; branchlets not ribbed ..... 5. Callichlamys

16. Tendril multifid; calyx membranous; branchlets with thin detachable ribs .....18. Manaosella

17(15). Trees or shrubs; tendrils absent; leaves mostly simple; flowers white ..... 35. Tabebuia

17. Lianas; tendrils present; leaves mostly 2- or 3-foliolate; flowers yellow, magenta, cream, or white ..... 18

18(17). Anthers exserted; corolla orange, tubular, the lobes valvate ..... 30. Pyrostegia

18. Anthers included; corolla yellow, magenta, or white, tubular-campanulate to tubular-funnel-shaped ..... 19

19(18). Corolla yellow or cream ..... 20

19. Corolla magenta or white with purple markings in throat ..... 22

20(19). Branchlets acutely tetragonal ..... 23. Mussatia

20. Branchlets terete ..... 21

21(20). Calyx drying black, minutely 5-denticulate, 13-16 mm long; inflorescence an elongate raceme, drying black, with subfoliaceous bracts and bracteoles subtending the buds ..... 1. Adenocalymna (A. impressum)

21. Calyx usually drying olive, subtruncate to subulately 5-denticulate, less than 13(-15) mm long; inflorescence usually a contracted raceme or fascicle of few flowers, drying olive; bracts and bracteoles minute or absent ..... 3. Anemopaegma

22(19). Calyx > 11 mm long, irregularly bilabiate ..... 20. Martinella

22. Calyx less than 10 mm long, truncate or subbilabiate .....23

23(22). Leaves 3-foliolate in part; disk conspicuous; corolla tube lightly lepidote near apex ..... 29. Potamoganos

23. Leaves simple to 2-foliolate, never 3-foliolate; disk essentially absent; corolla tube densely lepidote outside ..... 24

24(23). Tendrils minutely trifid ..... 31. Roentgenia

24. Tendrils simple ..... 25

25(24). Ovary oblong, verrucose-muricate ..... 7. Clytostoma (C. sciuripabulum)

25. Ovary linear, lepidote ..... 10. Cydista

26(10). Corolla elongate, > 14 cm long, white ..... 36. Tanaecium

26. Corolla less than 8 cm long, variously colored ..... 27

27(26). Anther thecae villous ..... 28

27. Anther thecae glabrous or with a few scattered trichomes ..... 29

28(27). Thecae straight ..... 16. Lundia

28. Thecae reflexed forward near middle ..... 19. Mansoa (M. kerere)

29(27). Thecae reflexed forward 90°, the connective enlarged..... 30

29. Thecae straight or slightly curved, the connective not enlarged ..... 31

30(29). Corolla magenta, slightly bilabiate, the lobes almost equal; pollen in tetrads ..... 9. Cuspidaria

30. Corolla white, strongly bilabiate, the 2 upper lobes almost fused; pollen grains simple ..... 37. Tynanthus

31(29). Leaves pellucid-punctate; branchlets hollow ..... 34. Stizophyllum

31. Leaves not pellucid-punctate; branchlets not hollow ..... 32

32(31). Tendrils trifid-uncinate; calyx acutely 5-lobed, the lobes triangular, 4-5 mm long ..... 24. Parabignonia

32. Tendrils simple or trifid (but not uncinate); calyx subtruncate or bilabiate, often minutely 5-denticulate (the teeth subulate and to 5 mm long in Mansoa verrucifera)..... 33

33(32). Corolla yellow ..... 34

33. Corolla white to magenta ..... 36

34(33). Leaves 2- or 3-ternate ..... 22. Memora (M. tanaeciicarpa)

34. Leaves 2- or 3-foliolate ..... 35

35(34). Ovary glabrous to lepidote ..... 1. Adenocalymna

35. Ovary pubescent ..... 3. Anemopaegma (A. robustum)

36(33). Tendrils 3(-15)-fid; if an erect shrub without tendrils, the ovary always pubescent ..... 37

36. Tendrils simple or minutely bifid; ovary glabrous or lepidote, never puberulous ..... 41

37(36). Ovary glabrous or lepidote or ± subpuberulous; corolla often thin and sparsely puberulous; interpetiolar glandular fields often present ..... 38

37. Ovary villous; corolla thick, densely pubescent outside; interpetiolar glandular fields lacking ..... 39

38(37). Corolla white, rather thick; interpetiolar glandular fields present; petiolar glandular fields absent ..... 6. Ceratophytum

38. Corolla magenta, thin; both interpetiolar glandular fields and petiolar glands present or both absent ..... 19. Mansoa

39(37). Tendrils 3-15-fid; branchlets acutely hexagonal; vegetative trichomes uniformly simple; corolla white ..... 27. Pithecoctenium

39. Tendrils trifid; branchlets terete or hexagonal (if hexagonal the vegetative trichomes dendroid or the corolla magenta) ..... 40

40(39). Branchlets terete; corolla white, the tube bent forward near middle; leaves glabrous to puberulous with simple trichomes ..... 12. Distictella

40. Branchlets hexagonal; corolla white to magenta, the tube straight; leaves finely canescent below or with stellate trichomes ..... 13. Distictis

41(36). Corolla tube lepidote and sparsely puberulous; ovary oblong, muricate-verrucose; pseudostipules a group of subulate scales resembling a miniature bromeliad ..... 7. Clytostoma

41. Corolla tube not lepidote, glabrous (the lobes puberulous) or puberulous; ovary linear, not verrucose; pseudostipules not resembling a minute bromeliad, usually small and inconspicuous ..... 42

42(41). Tendrils minutely bifid; pseudostipules appressed-conical; interpetiolar glandular fields absent; petiolar glandular fields present ..... 25. Paragonia

42. Tendrils simple; pseudostipules inconspicuous; interpetiolar glandular fields often present; petiolar glandular fields absent ..... 43

43(42). Inflorescence with minute bracteoles or the bracteoles absent; vegetative trichomes usually simple, dendroid only when interpetiolar glandular fields absent ..... 4. Arrabidaea

43. Inflorescence with conspicuous linear bracteoles about as long as calyx; vegetative trichomes dendroid; interpetiolar glandular fields present ..... 38. Xylophragma

Key to the Genera of Bignoniaceae based on fruiting material
(The fruits of both Potamoganos and Parabignonia are unknown, so these two genera are not included in this key.)

1. Fruits indehiscent, subglobose; leaves simple; trees or hemiepiphytic lianas ..... 2

1. Fruits dehiscent; leaves simple or compound; trees, shrubs, or lianas ..... 3

2(1). Fruit > 5 cm long; leaves in alternate fascicles; small trees .....8. Crescentia

2. Fruit a globose berry less than 3 cm diameter; leaves opposite; hemiepiphytic lianas ..... 32. Schlegelia

3(1). Fruit dehiscent perpendicular to septum; trees or shrubs; leaves various but never 2-foliolate and always lacking tendrils (tribe Tecomeae) ..... 4

3. Fruit dehiscent parallel to septum (valves splitting in center and fruit thus apparently 4-valved in some species); lianas, shrubs, or treelets; leaves often 2-foliolate and usually tendrillate in part (tribe Bignonieae) ..... 8

4(3). Fruit rounded to elliptic-oblong, strongly flattened perpendicular to septum; leaves simple to pinnately or bipinnately compound ..... 5

4. Fruit linear or linear-oblong, terete or ± flattened parallel to septum; leaves simple to palmately compound ..... 6

5(4). Leaves simple or pinnate ..... 11. Digomphia

5. Leaves bipinnate ..... 15. Jacaranda

6(4). Fruit spirally twisted; leaflets 7-9, usually cuneate to base, the petiolules poorly differentiated ..... 14. Godmania

6. Fruit straight or slightly curved; leaflets 1-7, conspicuously petiolulate (at least the terminal ones) ..... 7

7(6). Seed wings fragmented into separate trichomes; leaves (especially when young) and young shoots varnished ..... 33. Sparattosperma

7. Seed wings membranous (or absent); leaves and young twigs not conspicuously varnished ..... 35. Tabebuia

8(3). Leaves mostly pinnate, bipinnate, or 2- or 3-ternate ..... 9

8. Leaves simple or 2- or 3-foliolate (occasionally 2-ternate toward base in Arrabidaea bilabiata) ..... 11

9(8). Branchlets acutely tetragonal; tendrils trifid ..... 28. Pleonotoma

9. Branchlets terete or subterete; tendrils simple (trifid in Memora flavida) ..... 10

10(9). Interpetiolar glandular fields present ..... 4. Arrabidaea (A. inaequalis)

10. Interpetiolar glandular fields absent ..... 22. Memora

11(8). Vegetative parts of plant with clove odor; fruit valves puberulous and winged or with acute squarish margins and less than 28 × 0.7 cm ..... 37. Tynanthus

11. Vegetative parts of plant without clove odor; fruit unwinged (except Adenocalymna purpurascens where glabrous), > 0.8 cm wide (except in Arrabidaea pubescens with glabrous fruit and Stizophyllum with terete fruit > 25 cm long) ..... 12

12(11). Fruit less than 7 mm wide, cylindric; leaflets pellucid-punctate ..... 34. Stizophyllum

12. Fruit > 6 mm wide, usually flattened; leaflets not pellucid-punctate ..... 13

13(12). Fruit linear, terete, each valve with a narrow, central, longitudinal furrow ..... 9. Cuspidaria

13. Fruit linear to oblong, lacking medial furrow, usually compressed or ± oblong ..... 14

14(13). Fruit oblong (linear-oblong and terete with smooth surface in Adenocalymna impressum), the valves ± woody (coriaceous in the species of Anemopaegma with wingless seeds), > 2.2 cm wide ..... 15

14. Fruit linear, distinctly flattened or the valves with sandpaper-like texture (Paragonia), less than 2.6 cm wide (except sometimes in Mansoa which also appears in preceeding key section); valves coriaceous to thinly subwoody; seeds thin, 2-winged ..... 32

15(14). Fruit conspicuously echinate or acutely tuberculate ..... 16

15. Fruit surface not echinate nor conspicuously tuberculate (surface rugose-verrucose in Mussatia) ..... 19

16(15). Branchlets acutely hexagonal with ribbed angles ..... 17

16. Branchlets terete or tetragonal ..... 18

17(16). Tendrils trifid; leaves usually with stellate or dendroid trichomes; fruit projections obtuse, often widely scattered ..... 2. Amphilophium

17. Tendrils 3-15-fid; leaves only with simple trichomes; fruit projections acute, close together ..... 27. Pithecoctenium

18(16). Tendrils simple; fruit projections thin and hook-tipped ..... 7. Clytostoma

18. Tendrils trifid; fruit projections thick, not curved ..... 19. Mansoa (M. verrucifera)

19(15). Branchlets acutely tetragonal; pseudostipules foliaceous ..... 23. Mussatia

19. Branchlets terete or hexagonal ..... 20

20(19). Capsule valves splitting in half at maturity, the fruit apparently 4-valved ..... 21. Melloa

20. Capsule valves not splitting, the fruit 2-valved ..... 21

21(20). Fruits conspicuously puberulous ..... 22

21. Fruits glabrous or lepidote ..... 24

22(21). Branchlets hexagonal ..... 13. Distictis

22. Branchlets terete ..... 23

23(22). Capsule stipitate ..... 3. Anemopaegma

23. Capsule not stipitate ..... 12. Distictella

24(21). Branchlets hexagonal; tendrils trifid ..... 2. Amphilophium

24. Branchlets terete or subterete; tendrils simple or trifid ..... 25

25(24). Fruits terete, > 6 cm diameter; seeds unwinged ..... 36. Tanaecium

25. Fruits usually ± flattened, if terete less than 4 cm diameter; seeds winged or unwinged ..... 26

26(25). Interpetiolar glandular fields present ..... 27

26. Interpetiolar glandular fields absent ..... 29

27(26). Tendrils simple; leaf trichomes mostly dendroid ..... 38. Xylophragma

27. Tendrils trifid; leaf trichomes uniformly simple ..... 28

28(27). Petiolar glandular fields absent; pseudostipules of 3 side-by-side subulate scales; fruit gibbous at base ..... 6. Ceratophytum

28. Petiolar glandular fields present; pseudostipules vertically 3-seriate; fruit acute at base ..... 19. Mansoa

29(26). Fruit stipitate; tendrils usually trifid ..... 3. Anemopaegma

29. Fruit inconspicuously stipitate; tendrils simple ..... 30

30(29). Fruit terete, subterete, or winged; seed body thicker than the wings or the seeds wingless ..... 1. Adenocalymna

30. Fruit flattened; seed body about as thick as the tannish or brownish wings ..... 31

31(30). Fruit > 5 cm wide, the surface smooth ..... 5. Callichlamys

31. Fruit less than 4 cm wide, the surface irregularly raised-striate ..... 10. Cydista (C. lilacina)

32(14). Fruit pubescent with nonglandular trichomes, the central vein and margins raised ..... 16. Lundia

32. Fruit glabrous or lepidote or pubescent (Manaosella, Arrabidaea mollissima, A. brachypoda, A. lachnaea) but central veins and margins uniformly flattened or with the trichomes in part gland-tipped ..... 33

33(32). Tendrils trifid or multifid ..... 34

33. Tendrils simple or minutely bifid (Paragonia) ..... 39

34(33). Tendrils much-divided, the apices in part enlarged and disk-like; fruit subtended by persistent calyx > 2 × 1.2 cm, trichomes in part dendroid and in part gland-tipped, the fruit pubescent ..... 18. Manaosella

34. Tendrils trifid, the apices not disk-like; calyx not persistent in fruit or less than 2 × 1.2 cm, the fruit glabrous or lepidote ..... 35

35(34). Tendrils trifid-uncinate; plants growing appressed to supporting tree ..... 17. Macfadyena

35. Tendrils not uncinate; stems free-climbing or hanging, not appressed to trunk of supporting tree ..... 36

36(35). Interpetiolar and petiolar glandular fields present ..... 19. Mansoa

36. Interpetiolar and petiolar glandular fields absent ..... 37

37(36). Capsule valves subwoody, > 1.6 cm wide ..... 31. Roentgenia

37. Capsule valves subcoriaceous, less than 1.8 cm wide ..... 38

38(37). Branchlets terete, smooth ..... 20. Martinella

38. Branchlets hexagonal, costate ..... 30. Pyrostegia

39(33). Stem cross section with 8-16 phloem arms; capsule with midline barely or not at all evident; seeds with brownish wings; interpetiolar glandular fields absent ..... 40

39. Stem cross section with 4 phloem arms; capsule with raised midline; seeds with the wings (except Paragonia) usually hyaline-membranous or absent; interpetiolar glandular fields present or absent ..... 41

40(39). Capsule valves scattered-lepidote, drying blackish; branchlets often tetragonal or subtetragonal ..... 10. Cydista

40. Capsule valves densely lepidote, drying grayish; branchlets terete ..... 26. Phryganocydia

41(39). Capsule valves flattened, the surface smooth to verrucose; tendrils simple; petiolar glandular fields absent; interpetiolar glandular fields often present ..... 4. Arrabidaea

41. Capsule valves convex, the surface sandpaper-like; tendrils minutely bifid; petiolar glandular fields present; interpetiolar glandular fields absent ..... 25. Paragonia


Scientific Comments:
Paul Berry (peberry@facstaff.wisc.edu) or Kay Yatskievych (kay.yatskievych@mobot.org).


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