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Flora of the Venezuelan Guayana

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CLUSIACEAE

4. CLUSIA L., Sp. Pl. 509. 1753.
Quapoya Aubl., Hist. Pl. Guiane 898. 1775.
Havetia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 203, t. 462. 1821 [1822].
Renggeria Meisn., Pl. Vasc. Gen. 1: 42. 1837.
Rengifa Poepp., Nov. Gen. Sp. Pl. 3: 12. 1845 [1840].
Havetiopsis Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 246. 1860.
Oedematopus Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 246. 1860.
Oxystemon Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 314. 1860.
Pilosperma Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 243. 1860.
Decaphalangium Melchior, Notizbl. Bot. Gard. Mus. Berlin-Dahlem 10: 946, f. 16. 1930.

by John J. Pipoly III, Denis M. Kearns, and Paul E. Berry

Free-standing, epiphytic, epipetric, or hemiepiphytic shrubs or trees, sometimes lianas. Branchlets terete or tetragonal, sometimes winged. Latex translucent, white, cream, yellow, or orange-yellow. Leaves decussate, with or without conspicuous latex canals; petioles with at most small basal adaxial margined pits. Inflorescence a terminal or lateral panicle of cymes, pyramidal or not, rarely flowers solitary. Flowers mostly petals 4-12, variously arranged, sometimes decussate or basally connate. Stamens 4-numerous, free or else connivent or connate basally by the filaments, or united into a frequently resiniferous, globular, umbrella-shaped or annular synandrium, when annular surrounding a pistillode or a staminodial mass; filaments of varying shape, sometimes lacking; anthers variable in shape, often short, muticous or not, dehiscent by a variety of slits or pores; staminodes mostly smaller than stamens, with or without anthers, sometimes early caducous. Pistil with 4-8(-21) carpels; styles short or rudimentary; stigmas free and radiate or connivent, large; pistillode somewhat similar to pistil or represented only by an undifferentiated central tissue mass producing waxy resin or absent; 1-many ovules per carpel. Fruit a septifragally dehiscent capsule, with persistent stigmas. Seeds with a reddish, nonvascularized aril; cotyledons minute.

New World tropics and subtropics; ca. 300 species, ca. 80 in Venezuela, 68 of these in the flora area.

Although the greatest number of taxa may be found in Colombia, a major radiation of endemic taxa in Clusia has occurred on the Guayana Shield, where many additional new species likely occur. This treatment takes an inclusive view of Clusia, treating a number of formerly segregate genera as sections. On the other hand, there is still considerable uncertainty over the circumscription of some of the sections in Clusia. Much more work is needed in this complex genus to better elucidate the basic taxonomy as well as the infrageneric classification.

Clusia troncosii Maguire was based on fruiting material only and cannot be assigned to a section; for this reason it is not included in the following keys. Clusia cuneata Benth. [Section Clusiastrum] is known from Guyana but has not yet been collected in Venezuela. It can be separated from C. pusilla by the long, ellipsoid fruit and the long, oblanceolate or oblong leaf blades.

Key to the Species and Sections of Clusia in the Venezuelan Guayana

by John J. Pipoly III and Paul E. Berry

1. Stigmas and carpels 4; stamens 4, 8, or 12 ..... 2

1. Stigmas and carpels (4)5-many; stamens > 12, or if 8 or 10, then with 5 or more carpels ..... 11

2(1). Stamens 4, anthers apically emarginate, staminodes basally united by a conspicuous tube; stigmas discoid (section Havetiopsis) ..... C. flavida

2. Stamens 8 or 12, anthers apically truncate, staminodes barely united basally by a very minute tube; stigmas triangular (section Oedematopus) ..... 3

3(2). Leaves sessile to subsessile, petioles, when present, less than 3 mm long ..... 4

3. Leaves petiolate, petioles 3-30 mm long ..... 6

4(3). Stems terete; leaf apex acute to subacuminate ..... C. sp. A

4. Stems tetragonal; leaf apex broadly rounded to obtuse ..... 5

5(4). Petioles absent; leaf blades widely oblong to obovate, upper surface dull, lower surface (when dry) without conspicuous latex canals, with round magenta glands ..... C. parvula

5. Petioles 1-3 mm long; leaf blades linear to very narrowly oblanceolate, lower surface with conspicuous latex canals, eglandular ..... C. sp. B

6(3). Lower surface of leaf blades lacking latex canals or glands ..... 7

6. Lower surface of leaf blades with numerous black latex canals or round, magenta glands ..... 9

7(6). Branchlets terete to angular, not tetragonal; mostly lowland habitats, along seasonally flooded river banks ..... C. octandra

7. Branchlets tetragonal; lowland to highland habitats, usually on rocks but not along flooded river banks ..... 8

8(7). Leaf blades with secondary veins prominently raised on both surfaces, shiny on upper surface when young, the margin flat toward base; staminodes 4 ..... C. obovata

8. Leaf blades with secondary veins inconspicuous on upper surface, prominulous on lower surface, the upper surface dull when young, the margin revolute toward base; staminodes 6 ..... C. ptaritepuiensis

9(6). Lower surface of leaf blades with numerous black linear canals, eglandular, the secondary veins impressed; petioles with adaxial margined pits equal to petiole diameter ..... C. huberi

9. Lower surface of leaf blades without latex canals, with round magenta glands, the secondary veins prominulous; petioles with adaxial margined pits wider than petiole diameter ..... 10

10(9). Branchlets 2.5-4 mm diameter, angular but not deeply sulcate; leaf blades rhombic to obovate, the apex subacuminate; petioles 10-15 mm long; fruit 6-8 mm diameter ..... C. sp. C

10. Branchlets 6-10 mm diameter, deeply sulcate; leaf blades widely obovate to suborbicular, the apex broadly rounded; petioles 20-30 mm long; fruit 15-25 mm diameter ..... C. sp. D

11(1). Stamens 10, united in a central umbrella-shaped or a lobed corona-like structure; stigmas 5 ..... 12

11. Stamens 10 to many, if 10, not as above; stigmas 4 or more ..... 14

12(11). Stamens in pairs on a 5-lobed corona-like structure; fruit conspicuously transversely ridged; leaf apex rounded (section Phloianthera, in part) ..... C. comans

12. Stamens in a central umbrella-like structure; fruits smooth or only slightly ridged; leaf apex acuminate or rounded (section Quapoya) ..... 13

13(12). Leaf apex acuminate; stems square; lianas ..... C. hammeliana

13. Leaf apex obtuse to rounded; stems terete; trees ..... C. sipapoana

14(11). Stamens and/or staminodes not forming a resiniferous structure, never immersed in resin; stigmas orbicular, elliptic, or, when triangular, with rounded angles ..... 15

14. Stamens and/or staminodes forming a resiniferous structure immersed in resin at anthesis; stigmas triangular with acute corners ..... 45

15(14). Filaments 2-5 times longer than the anthers; stigmas 8, concave, sessile (section Cochlanthera) ..... C. cochlitheca

15. Filaments at most slightly longer than the anthers; stigmas 4-many, convex, sessile or on short styles ..... 16

16(15). Stamens and staminodes with flat, nonfleshy filaments; stigmas on short (occasionally rudimentary), radiate styles ..... 17

16. Stamens and staminodes with terete, prismatic, or flat but fleshy filaments; stigmas sessile or on short, erect styles ..... 31

17(16). Stigmas wedge-shaped or with small, horn-like protuberances; anthers dehiscent by longitudinal slits (section Anandrogyne) ..... 18

17. Stigmas elliptic; anthers dehiscent by subterminal pores or to halfway down from the anther tip (section Clusiastrum) ..... 24

18(17). Branchlets 8-15 mm diameter; pedicels thick, 7-15 mm long; stigmas with small horn-like protuberances ..... 19

18. Branchlets 2-8 mm diameter; pedicels thin, 2.5-6.5 mm long; stigmas wedge-shaped ..... 21

19(18). Leaf blades chartaceous to coriaceous, obovate to oblanceolate, the bases acute, the margins plane; sepals stiffly coriaceous, the margins entire; styles thick, radiate, 3-5 mm long; ovary and fruit smooth, not sulcate ..... C. cerroana

19. Leaf blades cartilaginous, elliptic to suborbicular, the bases obtuse to truncate, the margins volute; sepals chartaceous, the margins variously incised; styles rudimentary; ovary and fruit deeply sulcate ..... 20

20(19). Leaf blades with secondary veins inconspicuous on both surfaces; floral bracts 2, 15-30 mm long ..... C. duartei

20. Leaf blades with secondary veins prominently raised on both surfaces; floral bracts 4, 8-12 mm long ..... C. pachyphylla

21(18). Branchlets verrucose, 2-3 mm diameter; leaf blades orbicular, 2-3.5 × 1.5-3.3 cm, the bases cordate; sepals 5; fruit globose ..... C. fabiolae

21. Branchlets smooth, 5-8 mm diameter; leaf blades 5-9 × 3.5-8 cm, the bases truncate or broadly acute; sepals 4; fruit urceolate or ovoid ..... 22

22(21). Petioles without conspicuous adaxial margined pits; leaf bases subacute; peduncle less than 1 cm long; petals 5; staminodes numerous, caducuous ..... C. phelpsiae

22. Petioles with large adaxial margined pits; leaf bases rounded to truncate; peduncle 1.2-5 cm long; petals 4; staminodes 4-10, persistent in fruit ..... 23

23(22). Inflorescence 3-4 times compound, 9-12-flowered; styles rudimentary; fruit ovoid; in Amazonas at 1200-2000 m ..... C. rotundifolia

23. Inflorescence simple, 1-3-flowered; styles thin, 1-1.5 mm long; fruit subglobose; in Gran Sabana, Bolívar, at ca. 1000 m ..... C. savannarum

24(17). Leaf blades obovate to suborbicular, apex rounded to truncate ..... 25

24. Leaf blades elliptic to narrowly oblanceolate, apex obtuse to acute ..... 28

25(24). Branchlets 3.5-5 mm diameter; leaf blades thinly coriaceous, not brittle, epunctate on lower surface, apex rounded to obtuse ..... 26

25. Branchlets 6-12 mm diameter; leaf blades thickly coriaceous to cartilaginous, brittle, bearing red glandular dots on lower surface, apex obtuse to truncate ..... 27

26(25). Peduncle 1.5-3.5 cm long; sepal margin narrow, opaque; leaf blades with secondary veins scarcely or not visible on lower surface, the margin revolute ..... C. schomburgkii

26. Peduncle 3-14 cm long; sepal margin broad, scarious; leaf blades with secondary veins visible on lower surface, the margin flat ..... C. pusilla

27(25). Leaf blades cartilaginous, 3.5-5(-6.8) cm wide, upper surface dull, costa prominent on lower surface for 4/5 of its length; petioles 3-10 mm long; styles less than 12 ..... C. amabilis

27. Leaf blades thickly coriaceous, (6-)7-15 cm wide, upper surface shiny, costa prominent on lower surface throughout its length; petioles 10-15(-30) mm long; styles 12-18 ..... C. crassifolia

28(24). Branchlets + 5 mm diameter; leaf blades to 4 cm wide, margin hyaline or scarious; peduncle 8-25 mm long ..... 29

28. Branchlets 5-12 mm diameter; leaf blades (4.2-)6-8 cm wide, margin opaque; peduncle 3-7 mm long ..... 30

29(28). Branchlets 1-3 mm diameter; leaf blades elliptic, 1.5-4.5 × 0.7-1.2 cm, apex and base acute; flower buds ca. 4 mm diameter; petals 1-1.2 cm long ..... C. parvifolia

29. Branchlets 4-5 mm diameter; leaf blades oblanceolate, 8-13 × 2.7-4 cm, apex obtuse, base rounded; flower buds ca. 20 mm diameter; petals 4-4.5 cm long ..... C. phelpsiana

30(28). Branchlets 5-7 mm diameter; leaf blades elliptic, 10-16 × (4.2-)5-7 cm; flowers solitary ..... C. tabulamontana

30. Branchlets 8-12 mm diameter; leaf blades oblanceolate, 25-30 × 7-8 cm; flowers 3 per inflorescence ..... C. steyermarkii

31(16). Stamens and staminodes on minute, terete or prismatic filaments; staminate flowers producing small quantities of resin (section Brachystemon) ..... 32

31. Stamens and staminodes on obvious flat and fleshy filaments; staminate flowers not producing resin ..... 33

32(31). Leaf blades oblong to elliptic, the base obtuse to broadly rounded, not decurrent on the petiole, the margins flat; petioles subterete, 3-5 cm long; branchlets 10-22 mm diameter ..... C. aymardii

32. Leaf blades oblanceolate to obovate, the base attenuate, decurrent on the petiole, the margins revolute; petioles broadly marginate, 1-2.2 cm long; branchlets 7-10 mm diameter ..... C. spathulaefolia

33(31). Lianas; petals fleshy, thick when dry; stigmas elliptic to oblong (section Criuvopsis) ..... 34

33. Hemiepiphytic or free-standing trees or shrubs; petals not fleshy, chartaceous when dry; stigmas orbicular, ovate, wedge-shaped, or pentagonal (section Criuva) ..... 35

34(33). Leaf base obtuse to broadly rounded, leaves coriaceous, adaxial latex canals inconspicuous, adaxial venation and submarginal collecting vein prominent ..... C. amazonica

34. Leaf base acute to cuneate, leaves subcoriaceous, adaxial latex canals conspicuous, adaxial venation and submarginal collecting vein inconspicuous ..... C. martiana

35(33). Leaf apex acute to acuminate; blades chartaceous to subcoriaceous, dull on upper surface, pallid on lower surface with round magenta glands ..... 36

35. Leaf apex obtuse, broadly rounded, or truncate; blades stiffly coriaceous to cartilaginous, shiny at least on upper surface, lower surface eglandular ..... 39

36(35). Apical portions of branchlets winged; leaves sessile or petioles 5-10 mm long, broadly marginate ..... 37

36. Apical portions of branchlets subterete to tetragonal, corners rounded; petioles 10-35 mm long, narrowly marginate or trigonal ..... 38

37(36). Leaves 1.5-4 cm wide, sessile; peduncle 20-35 mm long; sepal margins opaque, entire ..... C. duidae

37. Leaves 2-6 cm wide, petioles 5-10 mm long, broadly marginate; peduncle less than 3 mm long; sepal margins scarious, hyaline, incised-erose ..... C. grammadenioides

38(36). Branchlets subterete or weakly tetragonal; leaf blades symmetric, latex canals conspicuous; petioles marginate, thin, 1-1.5 cm long; sepals 7, without conspicuous latex canals, the outer sepals acute to acuminate; petals decussate, thickly fleshy, suborbicular; anthers obovoid ..... C. multilineata

38. Branchlets strongly tetragonal; leaf blades asymmetric, latex canals inconspicuous; petioles trigonal, thick, 2-3.5 cm long; sepals 4, with conspicuous latex canals, the outer sepals rounded; petals imbricate, coriaceous, obovate; anthers ovoid ..... C. asymmetrica

39(35). Leaf blades coriaceous, rarely thickly so, the secondary veins prominent on both surfaces; inflorescence a many-flowered, erect, compound cyme, the rachis flattened; bracteoles numerous, small, similar to the sepals ..... 40

39. Leaf blades thickly coriaceous to cartilaginous, the secondary veins not easily discernible without magnification; inflorescence pendent, the rachis ± terete, flowers solitary or in few-flowered compound cymes; bracteoles few, smaller than the sepals ..... 41

40(39). Petioles 3-10 mm long; leaf blades obovate to very widely obovate, the bases cuneate, fully decurrent on petiole to base; inflorescence rachis longitudinally ribbed, not winged; stigmas on persistent styles ca. 1 mm long; fruit ovoid at first, fusiform at maturity ..... C. melchiori

40. Petioles 15-25(-28) mm long; leaf blades oblong, elliptic or rhombic, the bases obtuse, not decurrent on petiole; inflorescence rachis narrowly winged; stigmas sessile; fruit subglobose at first, globose at maturity ..... C. cardonae

41(39). Flowers in (3-)6-18-flowered cymes; fruits globose, smooth, without ribs; petioles canaliculate ..... 42

41. Flowers solitary or in reduced, 3-6-flowered cymes; fruits ovoid to subglobose, strongly ribbed; petioles broadly marginate ..... 43

42(41). Leaf blade bases obtuse to broadly rounded, margins revolute; peduncle 3-5 mm long; sepals 4-6, suborbicular; petals coriaceous; staminodes numerous; stigmas orbicular, sessile ..... C. opaca

42. Leaf blade bases broadly acute to obtuse, margins plane; peduncle 5-12 mm long; sepals 2, oblate; petals cartilaginous; staminodes 4; stigmas pentagonal, on persistent styles ..... C. guayanae

43(41). Petioles less than 6 mm long; flowers in an axillary, branched, 3-6-flowered cyme; petals suborbicular; staminodes 4 ..... C. maguireana

43. Petioles thick, 6-15 mm long; flowers terminal, solitary; petals obovate to orbicular; staminodes 8 or 9 ..... 44

44(43). Leaf blades 2.5-4.5 cm wide; bracteoles 3.5-4 mm long; sepal margins not scarious; petals 6; carpels 6-8 ..... C. hexacarpa

44. Leaf blades 5-7 cm wide; bracteoles 12-14 mm long; sepal margins scarious; petals 8; carpels 7-10 ..... C. radiata

45(14). Filaments distinct in the apical 2/3, subulate or clavate; anthers dehiscent by a terminal pore or longitudinal slit; stigmas convex, apically connivent in fruit and forming a sharp point, crown-shaped, or strongly reduced ..... 46

45. Filaments fused and forming a globular mass, or else distinct and cylindrical; anthers dehiscent by a terminal annulus, radial slits, or subterminal pores; stigmas concave, free or laterally connivent in fruit, obtuse or rounded ..... 59

46(45). Androecium pentagonal; filaments clavate to rudimentary; connective rounded, anthers dehiscent by a terminal slit; stigma papillose; fruit smooth or usually transversely ridged or wrinkled (section Phloianthera, pro parte) ..... 47

46. Androecium circular, with central resiniferous staminodes; filaments subulate, basally connate in a ring; connective with an apical subulate extension, anthers dehiscent by a longitudinal slit; stigma not papillose; fruit smooth or longitudinally ridged, not wrinkled (section Clusia) ..... 51

47(46). Bracts, bracteoles, sepals, and staminodes persistent; fruit globose or rarely obovoid, not ridged or wrinkled ..... 48

47. Bracts, bracteoles, sepals, and staminodes early caducous, fruit ovoid or ellipsoid, markedly transversely ridged at maturity, or prominently wrinkled ..... 49

48(47). Leaf blades coriaceous, narrowly oblong to narrowly oblanceolate, without visible latex canals, base long-decurrent; petioles less than or equal to 9 mm long; fruit always globose ..... C. myriandra

48. Leaf blades, obovate to oblanceolate, with latex canals conspicuous on upper surface, base shortly decurrent; petioles 15-25 mm long; fruit globose to obovoid ..... C. microstemon

49(47). Leaf blades cartilaginous; fruit obovoid, 3.5-5 cm diameter at maturity; stigmas 8-10 mm long ..... C. leprantha

49. Leaf blades chartaceous to coriaceous; fruit ellipsoid, 1-2 cm diameter at maturity; stigmas 1.5-3 mm long ..... 50

50(49). Leaf blades with conspicuous secondary veins on lower surface, latex canals inconspicuous, bases acute; petioles 10-15(-35) mm long; fruit exocarp shiny, only slightly wrinkled transversely at maturity ..... C. renggerioides

50. Leaf blades without conspicuous secondary veins on lower surface, latex canals conspicuous, bases cuneate; petioles 2-6 mm long; fruit exocarp dull, with numerous transverse ridges at maturity ..... C. panapanari

51(46). Adaxial margined pit at base of petiole narrower than petiole width; petioles 0.8-3 cm long ..... 52

51. Adaxial margined pit at base of petiole as wide or wider than petiole or else petiole adaxially concave for whole length (C. imbricata); petioles 0.3-0.5 cm long or else mostly 3-5 cm long ..... 55

52(51). Leaf blades thinly coriaceous; latex canals and secondary veins numerous, highly conspicuous ..... 53

52. Leaf blades thickly coriaceous to cartilaginous; latex canals and secondary veins inconspicuous ..... 54

53(52). Petioles 2-2.5 cm long; leaves with revolute margins and attenuate base; petals obovate ..... C. macropoda

53. Petioles 1.5-2 cm long; leaves with flat margins and acute base; petals suborbicular ..... C. nemorosa

54(52). Leaf base cuneate ..... C. viscida

54. Leaf base acute to obtuse ..... C. insignis

55(51). Leaf blades thickly coriaceous, latex canals inconspicuous; branchlets angular ..... 56

55. Leaf blades thickly cartilaginous, latex canals prominent on upper surface; branchlets terete ..... 57

56(55). Fruit oblong, ca. 5 cm long; leaf blades with midrib flat on upper surface, prominently raised on lower surface, secondary veins inconspicuous, base broadly rounded, adaxial margined pit as wide as petiole, petiole 2-3 mm thick ..... C. chiribiquetensis

56. Fruit depressed-globose, ca. 12 cm diameter when mature; leaf blades with midrib raised on both surfaces, the secondary veins prominulous on both surfaces, base obtuse, adaxial margined pit wider than the petiole, petiole 4-5 mm thick ..... C. grandiflora

57(55). Leaves with petioles 30-50 mm long; epiphytic or terrestrial ..... C. palmicida

57. Leaves subsessile, petioles less than 15 mm long; terrestrial ..... 58

58(57). Leaf blades oblong, the margin revolute, apex broadly rounded, base cuneate, decurrent on petiole to base; fruit 2-3 cm diameter ..... C. imbricata

58. Leaf blades obovate, the margin flat, apex and base truncate, not decurrent on petiole; fruit 5-8 cm diameter ..... C. rosea

59(45). Staminate flowers with pistillode lacking or highly reduced; stamens forming a tall column; staminodes forming a ring; anthers 2-celled, obovoid, dehiscent by subterminal pores; fruit angled in cross section; stigmas laterally connivent (section Androstylium) ..... C. fockeana

59. Staminate flowers, when present, mostly with evident pistillode; stamens forming a low, compact mass, truncate cone, or annulus; staminodes similar to stamens or lacking; anthers not 2-celled, with varying dehiscence; fruit ovoid or oblongoid; stigmas free ..... 60

60(59). Pistillode present, resiniferous; anthers dehiscent by terminal radial slits; staminodes lacking or fewer than stamens; ovary 5-8-carpellate (section Retinostemon) ..... 61

60. Pistillode present or lacking; anthers variously dehiscent; staminodes or stamens 10-30, each with a terminal anther, some of these functional; ovary 5-carpellate (section Omphalanthera) ..... 63

61(60). Branchlets thick, (5-)7-15 mm diameter; leaf blades cartilaginous, 15-20 × 8-15 cm; inflorescence 10-20 cm long ..... C. candelabrum

61. Branchlets thin, 2.5-3.5 mm diameter; leaf blades thinly coriaceous, 6-12 × 3.5-5 cm; inflorescence 5-7 cm long ..... 62

62(61). Branchlets with exfoliating bark; leaf margins flat, the linear resin canals prominent ..... C. minor

62. Branchlets without exfoliating bark; leaf margins revolute to inrolled, the linear resin canals scarcely or not visible ..... C. lopezii

63(60). Fruit not beaked ..... 64

63. Fruit noticeably beaked ..... 65

64(63). Anthers annular; petals clawed at base, pinkish white with maroon claw ..... C. brachystyla

64. Anthers polytecate with an apical disc-shaped pit containing numerous elongate pollen sacs; petals not clawed at base, dark purple throughout on inner surface ..... C. schomburgkiana

65(63). Leaf blades very widely obovate to suborbicular; petioles less than or equal to 5 mm long; penduncle > 4 cm long ..... C. annularis

65. Leaf blades elliptic, oblanceolate or oblong; petioles (15-)20-40 mm long; peduncle 2-4 cm long ..... 66

66(65). Petals 20-30 mm long ..... C. columnaris

66. Petals 8-12 mm long ..... C. gratula

Synopsis of the sections of Clusia recognized in this treatment

Clusia sect. Anandrogyne Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 323. 1860. -Clusia sect. Criuva subsect. Anandrogyne (Planch. & Triana) Engl. in Mart., Fl. Bras. 12(1): 402. 1888.

-Clusia subg. Thysanoclusia sect. Anandrogyne (Planch. & Triana) Vesque in A. DC. & C. DC., Monogr. Phan. 8: 29. 1893.

A section of ca. 70 species, widespread in the Neotropics; 7 species in the flora area (C. cerroana, C. duartei, C. fabiolae, C. pachyphylla, C. phelpsiae, C. rotundifolia, C. savannarum).

Clusia sect. Androstylium (Miq.) Engl. in Mart., Fl. Bras. 12(1): 402. 1888. -Androstylium Miq., Stirp. Surinam. Select. 93. 1850 [1851].

A section with a single species (C. fockeana) occurring in Venezuela, Guyana, Suriname, French Guiana, and Brazil.

Clusia sect. Brachystemon (Engl.) Pipoly, Sida 16: 517. 1995. -Clusia sect. Criuva subsect. Brachystemon Engl. in Mart., Fl. Bras. 12(1): 412. 1888.

A section of 3 species from Colombia, Venezuela, Ecuador, Peru, Brazil, and Bolivia; 2 species in the flora area (C. aymardii, C. spathulaefolia).

Clusia sect. Clusia

A section of about 13 species, widespread in the Neotropics; 9 species in the flora area (C. chiribiquitensis, C. randiflora, C. imbricata, C. insignis, C. macropoda, C. nemorosa, C. palmicida, C. rosea, C. viscida).

The circumscription of this section follows R. Howard, J. Arnold Arbor. 43: 389-399. 1962. However, V. Bittrich and P. Stevens (personal communication) have a manuscript (Taxon, in press) that establishes one of the drawings of Clusia alba Jacq. as the lectotype for the generic type, Clusia major L. This will require the renaming of the section as circumscribed above because C. major will be excluded from it.

Clusia sect. Clusiastrum Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 322. 1860.

A section of some 17 species occurring mainly in Colombia, Venezuela, Guyana, Suriname, and Brazil; 8 species in the flora area (C. amabilis, C. crassifolia, C. parvifolia, C. phelpsiana, C. pusilla, C. schomburgkii, C. steyermarkii, C. tabulamontana).

Clusia sect. Cochlanthera (Choisy), Engl. in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 202. 1925. -Cochlanthera Choisy, Descr. Guttif. Inde. 16, t. 3. 1849. -Clusia sect. Euclusia subsect. Cochlanthera (Choisy) Engl. in Mart., Fl. Bras. 12(1): 403. 1888.

A section of 7 species in Colombia and Venezuela, with only C. cochlitheca in the flora area.

Clusia sect. Criuva Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 323. 1860. -Clusia sect. Criuva subsect. Eucriuva Engl. in Mart., Fl. Bras. 12(1): 406. 1888.

A section of about 25 species in tropical South America; 11 species in the flora area (C. asymmetrica, C. cardonae, C. duidae, C. grammadenioides, C. guayanae, C. hexacarpa, C. maguireana, C. melchiori, C. multilineata, C. opaca, C. radiata).

Clusia sect. Criuvopsis Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 322. 1860. -Clusia sect. Criuva subsect. Criuvopsis (Planch. & Triana) Engl. in Mart., Fl. Bras. 12(1): 401. 1888.

-Clusia subg. Criuva sect. Criuvopsis (Planch. & Triana) Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 225. 1895.

A section of 3 species from Central America and tropical South America; 2 species in the flora area (C. amazonica, C. martiana).

Clusia sect. Havetiopsis (Planch. & Triana) Pipoly, comb. & stat. nov. -Havetiopsis Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 246. 1860.

A section of 2 species from Central America and tropical South America; only Clusia flavida occurs in the flora area.

Clusia sect. Oedematopus (Planch. & Triana) Pipoly, comb. & stat. nov. -Oedematopus Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 249. 1860. Lectotype here designated: Havetia octandra Poepp & Endl., Nov. Gen, Sp. 3: 11, t. 209A. 1840. [Oedematopus octandrus (Poepp. & Endl.) Planch. & Triana, Ann. Sci. Nat. sér. 4, 14: 250. 1860].

A section of about 16 species from Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia; 9 species in the flora area (C. huberi, C. obovata, C. octandra, C. parvula, C. ptaritepuiensis, C. sp. A, C. sp. B, C. sp. C, C. sp. D).

Clusia sect. Omphalanthera Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 319. 1860.

Clusia sect. Gomphanthera Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 319. 1860.

Polythecandra Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 228. 1860. -Clusia sect. Polythecandra (Planch. & Triana) Vesque in A. DC. & C. DC., Monogr. Phan. 8: 134. 1893.

A section of 7 species in Central America and tropical South America; 5 species in the flora area (C. annularis, C. brachystyla, C. columnaris, C. gratula, C. schomburgkiana).

Section Polythecandra is often recognized as distinct from sect. Omphalanthera, based on the presence of a pistillode and its unusual anther morphology. Section Gomphanthera is also sometimes recognized as distinct based on the irregularly dehiscent anthers.

Clusia sect. Phloianthera Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13. 319. 1860.

Renggeria Meisn., Pl. Vasc. Gen., Commentarius 42. 1837.

Clusia sect. Cordylandra Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13. 321. 1860.

A section of about 20 species in tropical South America; 6 species in the flora area (C. comans, C. leprantha, C. microstemon, C. myriandra, C. panapanari, C. renggerioides).

Clusia sect. Quapoya (Aubl.) Pipoly, comb. & stat. nov. -Quapoya Aubl., Hist. Pl. Guiane 897. 1775. -Clusia sect. Pachystemon subsect. Quapoya (Aubl.) Engl. in Mart., Fl. Bras. 12(1): 425. 1888.

A section of 9 species known from Panama and tropical South America; 2 species in the flora area (C. hammeliana, C. sipapoana).

Clusia sect. Retinostemon Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 320. 1860.

A section of 5 species in the Neotropics; 3 species in the flora area (C. candelabrum, C. lopezii, C. minor).

Clusia Species Entries

Clusia amabilis Maguire, Mem. New York Bot. Gard. 10(1): 57. 1958. [Section Clusiastrum].

Shrub or small tree to 5 m tall; latex white; leaves subsessile; stamens numerous, male flowers in 1(3)-flowered cymes; petals 7. Upper montane shrublands, often on rock outcrops, 1200-2300 m; Bolívar (Acopán-tepui, Auyán-tepui, Cerro El Sol, Cerro Marutaní, Cerro Sarisariñama, between Luepa and Kavanayén, Sierra Pakaraima). Brazil (Roraima: Serra do Sol).

Clusia amazonica Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 358. 1860. [Section Criuvopsis].

Clusia oedematopoidea Maguire, Phytologia 39: 73. 1978.

Liana or scandent tree; latex white; peduncle characteristically wrinkled; leaves elliptic, apex acute to acuminate, base acute; petiole 1-2 cm long; petals 5, white, 3.5-4 × 2.5-3 mm; stamens numerous; fruit oblongoid, 2-3.5 cm long, 1.5-2.5 cm diameter. Riparian black-water habitats, 50-400 m; Amazonas (Capibara, Río Baría, Río Cuao, Río Cunucunuma near Culebra, Río Gavilán, Salto Yureba in lower Río Ventuari basin, 12 km southeast of San Fernando de Atabapo, Sierra de la Neblina base). Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. Fig. 184.

Clusia annularis Maguire, Mem. New York Bot. Gard. 10(4): 23. 1961. [Section Omphalanthera].

Small tree 2-10 m tall; latex white; leaves subsessile, widely obovate to suborbicular, 12-20 × 8-12 cm, apex rounded; cymes many-flowered; petals 5, white with red lacquer-like center. Meadows and rock outcrops, shrub islands, low montane forests, 1000-2100 m; Amazonas (Cerro Autana, Cerro Moriche, Cerro Sipapo, Cerro Yapacana, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina).

Clusia asymmetrica Pipoly, Sida 16: 657. 1995. [Section Criuva].

Small tree to 6 m tall; branches 4-angled; leaves oblanceolate, 12-22 × 5-9 cm, apex and base acute, the base decurrent on the petiole; staminate inflorescence pendent, pyramidally paniculate, 3-7 cm long, 7-10 cm wide; petals 4, white, coriaceous, obovate, 4.8-5 × 3-3.2; stamens 18. Granitic outcrops, montane forests, 200-1200 m; Amazonas (lower slopes of Cerro Aracamuni, slopes of Cerro Marahuaka). Endemic.

Clusia aymardii Pipoly, Sida 16: 524. 1995. [Section Brachystemon].

Free-standing tree to 3 m tall; latex white; leaves oblong-elliptic, 10-18 × 5-11 cm, widely rounded to truncate at the apex, base obtuse to widely rounded; petioles 3-5 cm long; inflorescence 4-12-flowered; petals 5, white, 13-17 mm long, + clawed; stamens numerous, free; fruit 7-carpellate, oblongoid, ca. 3 cm long, 1.45 cm diameter. Scrub forests on tepui summits and slopes, 700-1200 m; Bolívar (Cerro Guaiquinima). Endemic.

Clusia brachystyla Maguire, Mem. New York Bot. Gard. 10(4): 25. 1961. [Section Omphalanthera].

Clusia brachystyla var. guaiquinimensis Maguire, Mem. New York Bot. Gard. 10(4): 26. 1961.

Shrub or tree 1-15 m tall; latex whitish; stems 4-angled; leaves elliptic, apex acute, base attenuate; petioles 3-4 cm long; petals flushed pink and basally maroon; stigmas 6, stylar column 1-2 mm long; fruit oblong-ellipsoid. Exposed ridges on tepuis, meadows, along streams, low montane forests, 300-1700 m; Bolívar (Cerro Guaiquinima, near Kavanayén, Macizo del Chimantá, base of Ptari-tepui, Sororopán-tepui), Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yutajé). Guyana.

Clusia candelabrum Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 248. 1860. [Section Retinostemon].

Clusia grandifolia Engl. in Mart., Fl. Bras. 12(1): 429. 1888.

Tree to 20 m tall or epiphytic strangler; leaves 15-20 × 8-15 cm; inflorescence many-flowered, 10-20 cm long; petals 5, white; stigmas 5, large, concave; androecium pentagonal. Riparian forests, 50-1600 m; Delta Amacuro (Caño Araguabisi between Caño Araguao and Caño G¸iniquina, northeast of La Grulla), Bolívar (slopes of Amaruay-tepui, Quebrada Pacheco near Santa Elena de Uairén, Río Parg¸eni east of Gavilán), Amazonas (Caño Tamatama, Carmelitas, Maroa, Río Atabapo, Río Casiquiare, upper Río Cuao, Río Guainía, Río Pasimoni, Río Ventuari near Río Parú mouth). Colombia, Ecuador, Peru, Brazil (Rio Vaupés). Fig. 186.

Clusia cardonae Maguire, Mem. New York Bot. Gard. 9: 483, fig. 122. 1957. [Section Criuva].

Tree to 15 m tall; latex whitish or yellowish; branches 4-angled; leaves oblong or elliptic, 5.5-11 × 4-9 cm, apex rounded to broadly acute, base obtuse; petioles 15-35 mm long; staminate inflorescence erect, pyramidally paniculate, many-flowered, 5-20 cm long, 5-8 cm wide; petals 4 or 5, white, fleshy, obovate, 5-8 mm long; stamens numerous, free to base; fruits globose, 5-10 mm diameter. Tepui summits, montane forests, tepui scrub, dwarf forests, 1000-1900 m; Bolívar (Amaruay-tepuí, Ilú-tepui, Kamarkawarai-tepuí, Sierra de Lema, Uaipán-tepuí, Uei-tepui). Guyana. Fig. 187.

Clusia cerroana Steyerm., Fieldiana, Bot. 28: 386. 1952. [Section Anandrogyne].

Clusia sessilis Klotzsch ex Engl. in Mart., Fl. Bras. 12(1): 406. 1888, non G. Forst. 1786. -Clusia wurdackiana Pipoly, Sida 16: 747. 1995, nom. superfl.

Tree or epiphyte 4-15 m tall; leaves obovate to widely oblanceolate, apex broadly rounded to truncate, base acute, 9-24 cm long, 6-1 cm wide; staminate inflorescence a 6-9-flowered panicle; petals 6, 12-15 × 5-7 mm; stamens numerous; fruit subglobose, 15-20 mm long and thick, smooth, 5-carpellate. Rocky outcrops, river edges, gallery forests, montane dwarf forests, 700-2100 m; Bolívar (Auyán-tepui, Cerro Impacto, Cerro Guaiquinima, Ilú-tepui, Macizo del Chimantá [Chimantá-tepui], Ptari-tepui, Roraima-tepui, Sierra de Lema, Sierra de Maigualida), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yutajé, Serranía Tapirapecó, Sierra de la Neblina). Monagas; Guyana, Suriname, Brazil.

Clusia cerroana frequently forms large colonies and appears to be fire resistant.

Clusia chiribiquetensis Maguire, Bot. Mus. Leafl. 15: 56. 1951. [Section Clusia].

Tree 3-8 m tall; latex yellowish; leaves orbicular, petioles 1-2 cm long; fruit oblong-ellipsoid, ca. 5 cm long. Lowland riparian forests, dwarf montane forests, 0-2000 m; Delta Amacuro (Caño Guayo), Bolívar (near Roraima-tepui), Amazonas (Caño Caname, middle Caño Yagua, base of Cerro Aracamuni, Cerro Aratitiyope, Cerro Huachamacari, middle slopes of Cerro Marahuaka, Cerro Yutajé, Río Cuao, Río Yatúa, Salto Yureba in lower Río Ventuari basin, Serranía Uasadi). Colombia, Guyana. Fig. 188.

Clusia cochlitheca Maguire, Mem. New York Bot. Gard. 10(1): 61. 1958. [Section Cochlanthera].

Erect to scandent tree 6-30 m tall; stems sharply angled, latex cream-white; cymes 1-3-flowered; petals 8, white. Montane forests, scrub forests, 600-2000 m; Amazonas (Cerro Huachamacari, Cerro Marahuaka, Cerro Parú). Endemic. Fig. 191.

Clusia columnaris Engl. in Mart., Fl. Bras. 12(1): 432. 1888. [Section Omphalanthera]. -Paratayán.

Clusia columnaris var. magdalenensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 41. 1950.

Clusia columnaris var. vaupesana Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 41. 1950.

Clusia williamsii Steyerm., Fieldiana, Bot. 28: 392. 1952.

Shrub or small tree 2-12 m tall; latex clear-white to yellowish; leaves narrowly elliptic, base attenuate; petioles 2-5 cm long; flowers pendent in dichasia (9-21 flowers in staminate plants, 2-11 flowers in pistillate plants), petals rose-white with deep maroon stripes at the base; androecium and apices of staminodes covered by resin; petals persistent in fruit; stigmas on stylar column 2-3 mm high; fruits angled and + rugose. Semideciduous forest on granitic outcrops, shrub islands on white sands, forest edges, edges of evergreen lowland forests, Río Negro caatinga, 100-800 m; Bolívar (Amaruay-tepui, Corozal 6 km from Maniapure toward Caicara, Gran Sabana), Amazonas (widespread). Colombia (Meta, Vaupés), Brazil. Fig. 193.

Clusia comans (Meisn.) Pipoly, comb. nov. [Section Phloianthera]. -Renggeria comans Meisn., Pl. Vasc. Gen., Commentarius 42. 1837.

Small tree. Riparian forests, 0-100 m; the only known collection from Venezuela is from the Río Guanipa (in Monagas state), which is less than 20 km from Pedernales in Delta Amacuro. French Guiana, Brazil (Amapá, Pará). Fig. 192.

This species is similar to Clusia panapanari but has only 10 stamens.

Clusia crassifolia Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 351. 1860. [Section Clusiastrum].

Tree 2-18 m tall; latex white to yellowish; leaves obovate, sessile; flowers large, single or in 3-flowered cymes, petals 8, white, 3-5 cm long; stamens numerous; stigmas 9-18, umbrella-like, styles 3-4 mm long. Gallery forests amid upland and highland savannas or meadows, tepui scrub, 1300-2300 m; Bolívar (Auyán-tepui, Cerro Marutaní, Cerro Venamo, Ilú-tepui, Kamarkawarai-tepui, Macizo del Chimantá, Ptari-tepui, Roraima-tepui, Sororopán-tepui). Guyana (Ayanganna, Wokomung, Kaieteur Falls area).

Clusia duartei Maguire, Moscosoa 4: 215. 1986. [Section Anandrogyne].

Tree to 10 m tall; latex white; branches 4-angled; leaves suborbicular to widely elliptic, 20-27 × 19-22 cm, subsessile; staminate inflorescence 1-3-flowered, petals 8-10, coriaceous, 4.5-5 × 1.5-2 cm; stamens numerous, 1.5-2 cm long; female flowers with petals 6, suborbicular, 2.5-3 cm long and wide; carpels 10-12, deeply sulcate along the sutures; fruits subglobose, 5-6 cm long and wide. Montane scrub forests along rivers, 1500-1600 m; Amazonas (Cerro Cuao, Cerro Sipapo). Endemic.

Clusia duidae Gleason, Bull. Torrey Bot. Club 58: 402. 1931 [1932]. [Section Criuva].

Shrub or small tree to 12 m tall; leaves subsessile, obovate, nearly clasping the stem, 6-8 × 2-4 cm, apex acute; pistillate flowers yellow, petals 5, coriaceous, obovate, 4-4.5 mm long; staminodes 4; fruit oblong, 4-carpellate, 1.5-3 cm long, 0.6-0.9 cm thick, longitudinally ridged, often appearing axillary and solitary. Montane to upper montane mossy elfin forests, 1100-2200 m; Bolivar (Auyán-tepui, Cerro Venamo, 125 km south of El Dorado, Ilú-tepui, Macizo del Chimantá [Abacapá-tepui, Apacará-tepui, Churí-tepui, Sarvén-tepui]), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic.

Clusia fabiolae Pipoly, Sida 16: 740. 1995. [Section Anandrogyne].

Shrub or small tree to 4(-7) m tall; latex creamy yellow; leaves subsessile, orbicular, subcordate at base, 2.5-5 cm long, 1.5-3.5 cm wide; inflorescence terminal, 3-flowered; flowers pink, petals 4; stamens numerous; fruits 5-carpellate, globose, 10-12 mm diameter. Tepui scrub, rocky savannas below large rock walls, 1600-2500 m; Bolívar (Auyán-tepui, Macizo del Chimantá, Los Testigos). Endemic.

Some herbarium specimens of this species were labeled as "Clusia wurdackii Mag. ined.," a name that was never published.

Clusia flavida (Benth.) Pipoly, comb. nov. [Section Havetiopsis]. -Havetia flavida Benth., London J. Bot. 2: 369. 1843. -Havetiopsis flavida (Benth.) Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 247. 1860. -Tari-yek (Arekuna).

Havetiopsis flexilis Spruce ex Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 246. 1860.

Epiphytic shrub or tree, densely branched, sometimes scrambling, 2-15 m high; latex scanty, clear-whitish; leaves obovate; petals cream; fruits quadrangular. Lowland to montane evergreen forests, swamps, semideciduous forests, seasonally flooded forests, 50-1300 m; Delta Amacuro (Serranía de Imataca), Bolívar (widespread), Amazonas (widespread). Apure, Miranda, Táchira, Yaracuy, Zulia; Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. Fig. 194.

Clusia fockeana Miq. in Hoeven & de Vriese, Tijdschr. Nat. Gescheid. 10: 82. 1843. -Androstylium fockeanum (Miq.) Miq., Stirp. Surinam. Select. 93. 1850 [1851]. [Section Androstylium].

Shrub ca. 2 m tall. Upland shrublands, ca. 1300 m; Bolívar (Gran Sabana, Río Maremán south of Luepa). Guyana, Suriname.

Clusia grammadenioides Pipoly, Sida 16: 653. 1995. [Section Criuva].

Hemiepiphyte, attaching to tree trunks by adventitious roots; branches 4-angled, winged; leaves narrowly elliptic to obovate, 1--25 × 2-6 cm, apex acute, base long-attenuate and decurrent on the petiole; staminate inflorescence erect, pyramidal-paniculate, 8-9 × 11-17 cm; petals 5, coriaceous, 4-5 × 2.4-3.4 mm; stamens ca. 18. Along streambanks on terraces just above floodline at forest margins on deep white sands, 100-200 m; Amazonas (northeast of San Carlos de Río Negro at Huachica). Colombia (Caquetá).

Clusia grandiflora Splitg. in Hoeven & de Vriese, Tijdschr. Nat. Gescheid. 9: 101. 1842. [Section Clusia]. -Copei.

Shrub or tree 3-30 m tall; latex white; leaves orbicular to obovate, long-petiolate; flowers single or in 3-flowered cymes, 12-14 cm wide at anthesis; petals white throughout, persistent in fruit; stigmas numerous, + 15; fruits large, + 12 cm diameter. Gallery forests, montane forests, 0-1300 m; Delta Amacuro (Río Ibaruma near Río Cuyubini), Bolívar (widespread in the Gran Sabana, Río Caroní), Amazonas (Salto Yureba in lower Río Ventuari basin). Guyana, Suriname. Fig. 196.

Clusia gratula Maguire, Mem. New York Bot. Gard. 10(4): 23. 1961. [Section Omphalanthera].

Tree 5-25 m tall. Riparian forests, Río Negro caatinga, dense montane thickets, 100-1200 m; Amazonas (Cerro Marahuaka, Río Pasimoni, Río Yatúa, San Carlos de Río Negro, Sierra de la Neblina). Mérida, Táchira.

This might be better recognized as a subspecies of Clusia columnaris, but more study is needed.

Clusia guayanae Pipoly, Sida 16: 674. 1995. [Section Criuva].

Shrub to 2 m tall; branchlets 4-angled; leaves oblanceolate to obovate, 3-7 × 2-4 cm; apex obtuse to truncate, base obtuse to rounded; staminate inflorescence a pendent terminal panicle 2-3 × 2-3 cm; petals 4 or 5, oblong, 3.5-7 × 2-3 mm; stamens 8; fruit 5-carpellate, depressed-globose, 8-11 × 10-12 mm diameter. Montane scrub and cloud forests along sandstone bluffs, 1200-1900 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Kamarkawarai-tepui). Guyana.

Clusia hammeliana Pipoly, nom. nov. -Rengifa peruviana Poepp., Nov. Gen. Sp. 3: 12, pl. 210. 1845 [1840], non Clusia peruviana Szyszyl. 1894. -Quapoya peruviana (Poepp.) Kuntze, Revis. Gen. Pl. 1: 61. 1891. [section Quapoya].

Quapoya peruviana var. guayanensis Maguire, Mem. New York Bot. Gard. 10(4): 30. 1961.

Epiphytic liana to 25 m tall; leaves elliptic, apex acuminate. Evergreen lowland to montane forests, 100-1300 m; widespread in southwestern Amazonas. Colombia, Ecuador, Peru, Brazil. Fig. 198.

This species possibly should include Quapoya bracteolata Sandwith, from coastal Guyana.

Clusia hexacarpa Gleason, Bull. Torrey Bot. Club 58: 403. 1931 [1932]. [Section Criuva].

Clusia hexacarpa var. ptaritepuiana Steyerm., Fieldiana, Bot. 28: 387. 1952.

Shrub to small tree to 15 m tall; latex white; leaves suborbicular to obovate or elliptic, 4-7 × 2.5-5 cm; flowers solitary, pendent; petals 5-7, white, 6-9 × 3.5-5 mm; stamens numerous; stigmas 6-8, circular, depressed in center; styles 4-5 mm long; fruit ovoid, 2.5-3.5 cm long, 1.5-2.5 cm diameter. Tepui forests, rock outcrops and exposed slopes of tepuis, 1400-1800 m; Bolívar (Macizo del Chimantá [Chimantá-tepui, Churí-tepui, Toronó-tepui], Ptari-tepui), Amazonas (Cerro Aracamuni, Cerro Coro Coro, Cerro Duida, Cerro Huachamacari, Cerro Yutajé, Sierra de la Neblina). Endemic.

Clusia huberi Pipoly, nom. nov. [Section Oedematopus]. -Oedematopus duidae Gleason, Bull. Torrey Bot. Club 58: 406. 1931 [1932], non Clusia duidae Gleason. -Tari-yek (Arekuna).

Hemiepiphytic shrub or tree 1-15 m tall; petals cream; calyx magenta; buds greenish red. Upland forests, 1000-2200 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá [Tirepón-tepui], Ptari-tepui), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yutajé). Guyana. Fig. 200.

Clusia imbricata Steyerm., Fieldiana, Bot. 28: 387. 1952. [Section Clusia].

Tree 5-12 m tall; latex white; leaves elliptic-obovate, base attenuate, petioles ca. 1 cm long; petals white, ca. 12; stigmas and staminodes numerous. Dwarf montane forests, tepui summit forests, talus forests, 800-2100 m; Bolívar (La Escalera, Macizo del Chimantá [Churí-tepui], Ptari-tepui). Endemic. Fig. 204.

Clusia insignis Mart., Nov. Gen. Sp. Pl. 3: 64. 1829. [Section Clusia].

Clusia insignis var. hoffmannseggiana Engl. in Mart., Fl. Bras. 12(1): 424. 1888.

Liana or tree to 25 m tall; flowers showy, purple-red, ca. 12 cm across. Evergreen lowland forests, riparian forests, 100-200 m; Amazonas (upper Río Autana, Río Negro near Piedra Cocuy). Brazil (Amazonas, Rio Negro).

Clusia leprantha Mart., Nov. Gen. Sp. Pl. 3: 165. 1829 [1832]. [Section Phloianthera].

Clusia purpurea (Splitg.) Engl. in Mart., Fl. Bras. 12(1): 416. 1888. -Arrudea purpurea Splitg. in Hoeven & de Vriese, Tijdschr. Nat. Gescheid. 9: 101. 1842.

Strangler, epiphyte to 25 m tall; latex white; leaves spathulate, apex rounded to retuse; fruit pentagonal, transversely rugose. Riparian forests, ca. 100 m; Amazonas (Río Padamo). Brazil (Amazonas). Fig. 205.

Clusia lopezii Maguire, Bot. Mus. Leafl. 15: 61. 1951. [Section Retinostemon].

Terrestrial or epiphytic tree 5-20 m tall; latex cream; leaves oblanceolate-obovate; petals white, pink at base; stigmas 5, rounded-capitate, separate in fruit; fruits pear-shaped with beak + 5 mm long. Seasonally flooded (black-water) riparian forests, savanna edges, 50-200 m; Amazonas (Caño Cotúa to the base of Cerro Yapacana, near Piedra Cocuy, Río Atabapo basin, Río Baría, Río Negro, Río Pasimoni, Río Siapa, Río Yatúa). Colombia (Vaupés). Fig. 206.

Clusia macropoda Klotzsch ex Engl., Fl. Bras. 12(1): 406. 1888. [Section Clusia].

Tree 4-8 m tall; latex yellow; buds pink. Riparian forests, 0-50 m; Delta Amacuro (Caño Guayo near Misión San Francisco de Guayo). Guyana, Suriname, French Guiana.

The wood of this species is used locally for making furniture.

Clusia maguireana Pipoly, Sida 16: 689. 1995. [Section Criuva].

Shrub to small tree to 4(-10) m tall; branchlets 4-angled; leaves very widely obovate, 4-9 × 2.5-6 cm, apex rounded to truncate, base obtuse, subsessile; staminate inflorescence axillary, pendent, twice branched; petals 4, thick-fleshy, 5.2-6 × 2.1 mm; stamens ca. 26; fruit 10-carpellate, strongly ribbed, subglobose, 17-20 mm long, 14-16 mm diameter. Rocky tepui outcrops, usually on overhanging ledges, 1100-2300 m; Bolívar (Auyán-tepuí, Los Testigos, Macizo del Chimantá [Abacapá-tepuí, Agparamán-tepuí, Apacará-tepuí], Uei-tepui). Guyana.

Clusia martiana Engl. in Mart., Fl. Bras. 12(1): 411. 1888. [Section Criuvopsis].

Clusia uleana Engl., Bot. Jahrb. Syst. 58(Beibl. 4): 1. 1923.

Scandent shrub or liana to 5 m tall; latex clear; petals 5, fleshy, white; stamens + 25; stigmas 5; fruits subglobose to ovoid, 2-3.5 cm long, 1.5-2.5 cm diameter. Along riverbanks, 50-200 m; Amazonas (Caño Iguapo near La Esmeralda, Río Casiquiare, Río Jénita 7 km above mouth of Río Ocamo, Río Mavaca, upper Río Orinoco, Río Padamo). Amazon basin of Colombia, Peru, Brazil, and Bolivia.

Clusia martiana is one of the few weedy species of Clusia, forming mats over the canopies of low trees along rivers, eventually killing them. It can also withstand submersion for extended periods of time.

Clusia melchiori Gleason, Bull. Torrey Bot. Club 58: 403. 1931 [1932]. [Section Criuva].

Shrub or small tree 2-15 m tall; latex clear to yellowish; leaves thick-coriaceous, obovate to oblong, 7-17 × 3-11 cm, apex broadly rounded to truncate, base cuneate; inflorescence erect, pyramidally paniculate, many-flowered, 5-19 × 5-11 cm; flowers small, fragrant, yellowish white; petals 4-6; stamens numerous; stigmas and styles 4; fruits ovoid to fusiform, 10-13 mm long, 4-8 mm diameter. Montane to upper montane elfin forests often dominated by Bonnetia species, 600-2200 m; Bolívar (Auyán-tepuí, Cerro Guaiquinima, Cerro Jaua, Ilú-tepui, Macizo del Chimantá, Ptari-tepui, Uaipán-tepui, Uei-tepui), Amazonas (Cerro Duida, Cerro Parú, Cerro Sipapo, Cerro Yapacana, Cerro Yaví, Cerro Yutajé, Río Coro Coro basin west of Cerro Yutajé, Sierra de la Neblina, Sierra de Maigualida, Sierra Parima, Sierra Tapirapecó, Serranía Uasadi). Guayana Shield portions of Colombia, Guyana, Suriname, and Brazil. Fig. 190.

Clusia microstemon Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 331. 1860. [Section Phloianthera]. -Cupi.

Terrestrial or epiphytic tree 5-12 m tall; latex yellow; flowers in cymes of 3 or 5; petals 5 or 6, white with purple-maroon at base; fruit globose, + 1.5 cm diameter, 6-carpellate. Seasonally flooded riparian forests (usually black-water rivers), Río Negro caatinga, 50-200 m; Delta Amacuro (Río Acure), Amazonas (Caño Tamatama, Caño Yagua near Cerro Yapacana, 10 km upstream from San Fernando de Atabapo, Carmelitas, Río Atacavi, Río Baría, Río Casiquiare, Río Manapiare, Río Pasiba, Río Pasimoni, Río Ventuari, San Carlos de Río Negro to Solano, Santa Bárbara del Orinoco). Colombia (Amazonas), Brazil (Amazonas: Rio Uaupés). Fig. 189.

Clusia minor L., Sp. Pl. 510. 1753. [Section Retinostemon]. -Copei, Quiripití.

Clusia galactodendron Desv., Ann. Sci. Nat. Bot. sér. 2, 18: 315. 1842.

Tree 2-10 m tall, apomictic (flowers pistillate or apparently hermaphroditic, but staminate flowers lacking); latex white; stigmas 5, concave. Exposed igneous outcrops, semidry savannas and semideciduous scrub, 100-700 m; Delta Amacuro (Río Toro), northern Bolívar, Amazonas (Salto Yureba in lower Río Ventuari basin). Aragua, Anzoátegui, Barinas, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Portugesa, Sucre, Yaracuy; Panama, West Indies, Colombia, Guyana. Fig. 208 .

Clusia multilineata Pipoly, Sida 16: 655. 1995. [Section Criuva].

Tree to 3 m tall; leaves elliptic, 8-13 × 4-6 cm; apex and base acute, the base decurrent on the petiole, the margin revolute; staminate inflorescence erect, pyramidally paniculate, 2.5-5 × 3-7 cm; petals 4, thick-fleshy, ca. 4 × 3.2 mm; stamens 22-24. Montane river canyons, 700-800 m; Amazonas (Sierra de la Neblina). Endemic.

Clusia myriandra (Benth.) Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 332. 1860. -Tovomita myriandra Benth., London J. Bot. 2: 367. 1843. [Section Phloianthera].

Clusia jenmani Engl. in Mart., Fl. Bras. 12(1): 432. 1888.

Tree 5-10 m tall; latex whitish-clear; leaves oblanceolate to spathulate, sessile, apex rounded; petals 5, maroon inside; androecium pentagonal. Evergreen lowland to lower montane forests, granitic outcrops, 50-400 m; Bolívar (base of Amaruay-tepui, Cordillera Epicara, La Escalera, 16 km northwest of Kilómetro 88). Guyana.

Clusia nemorosa G. Mey., Prim. Fl. Esseq. 203. 1818. [Section Clusia].

Tree 5-15 m tall; leaves obovate with prominent and closely parallel secondary veins; petioles 2-4 cm long; petals 7 or 8, white with reddish center; stigmas 8; fruit elongate-ellipsoid. Rocky places, lowland forest edges, 50-1600 m; Delta Amacuro (Caño Jotajana, Curiapo, Río Toro), Bolívar (Cerro Guaiquinima). Táchira; Trinidad, Guyana, Suriname, French Guiana, Brazil (Bahia, Pará, Pernambuco). Fig. 207.

Clusia obovata (Spruce ex Planch. & Triana) Pipoly, comb. nov. [Section Oedematopus]. -Oedematopus obovatus Spruce ex Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 250. 1860.

Oedematopus quadratus Maguire, Bull. Torrey Bot. Club 75: 431. 1948.

Shrub or tree 1-8 m tall; latex white to yellowish. Dwarf tepui forests, granitic and sandstone outcrops, savannas, shrublands, evergreen lowland forests, 50-2000 m; Bolívar (Auyán-tepui, Cerro Bolívar and vicinity, Cerro Guaiquinima, Cerro Jaua, Cerro Sarisariñama, Cerro Toribio, Gran Sabana, Macizo del Chimantá, middle Río Orinoco, middle Río Paragua), Amazonas (Caño Fluta 35 km southeast of Puerto Ayacucho, Cerro Ualipano, Cerro Yapacana, Puerto Ayacucho, Río Autana, Río Puruname, Río Sipapo). Colombia, Guyana, Suriname, Brazil.

Clusia octandra (Poepp.) Pipoly, comb. nov. [Section Oedematopus]. -Havetia octandra Poepp., Nov. Gen. Sp. Pl. 3: 11. 1845 [1840]. -Oedematopus octandrus (Poepp.) Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 250. 1860.

Tree to 8 m tall, densely branched, often with aerial roots; latex yellow or white; leaves narrowly obovate; inflorescence lax, much branched; petals 4, white; fruit oblongoid, 1-2 cm long. Seasonally flooded riparian forests (especially along black-water rivers), river banks, montane forests, 100-200(-1400) m; Bolívar (km 128 south of El Dorado), Amazonas (Caño Hechimoni on Río Pasimoni, Caño Yapacana, base of Cerro Duida, Laja Catipán, Piedra Arauicaua, Río Atacavi, Río Pasimoni, Río Temi, Río Yatúa). Colombia, French Guiana, Ecuador, Peru, Brazil. Fig. 201.

Clusia opaca Maguire, Bot. Mus. Leafl. 15: 62. 1951. [Section Criuva]. -Baniha, Copei, Cupi, Upihi.

Clusia reducta Steyerm., Fieldiana, Bot. 28: 391. 1952.

Shrub or tree to 6 m tall; latex white; leaves + sclerophyllous, often revolute-margined, 7-9 × 2.5-3.5 cm, apex and base obtuse; petioles 5-12 mm long; flowers fragrant, in cymes of 3 or 5; petals white, 5, narrow, 1-1.3 × 0.4-0.6 cm; stamens yellow, numerous; stigmas small, 4-6; fruits subglobose, pendulous, ca. 2 cm long, 1.5 cm diameter. White-sand savannas, edge of meadows, Río Negro caatinga, white-sand scrub (bana), 50-200 m; Amazonas (Caño Pimichín, Caño San Miguel, Caño Yagua, Río Atabapo basin, Río Casiquiare, Río Guainía, upper Río Orinoco near San Antonio). Colombia (Caquetá, Guainía), Brazil (Amazonas). Fig. 203.

Clusia pachyphylla Gleason, Bull. Torrey Bot. Club 58: 405. 1931 [1932]. [Section Anandrogyne].

Clusia scariosa Lasser & Maguire, Brittonia 7: 81. 1950.

Shrub or tree 2-10 m tall; latex white; stems 4-angled; leaves thick, stiff, revolute, apex rounded, base acute; 5-18 cm long, 3-10 cm wide, petiole 10-18 mm long; staminate inflorescence 1-3-flowered; petals 6-8, white, 25-30 × 13-20 mm; stamens numerous; stigmas 8, depressed in center, styles ca. 1.5 mm high; fruits ca. 4 cm long, 2-3 cm diameter, longitudinally sulcate. Tepui meadows, on rocks, 1700-2200 m; Bolívar (Cerro Guaiquinima, Cerro Jaua, Sierra de Maigualida), Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Yaví, Cerro Yutajé, Serranía Uasadi, Sierra de Maigualida). Endemic.

Clusia palmicida Rich. ex Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 326. 1860. [Section Clusia]. -Copei.

Tree or strangler to 15 m tall; latex yellow; leaves elliptic, apex bluntly acute, base attenuate; stigmas ca. 12, sessile. Evergreen lowland forests, riparian forests, 50-1300 m; Delta Amacuro (Caño Atoiba north of Caño Araguao, Caño Joba-Suburu east of Caño Sacupana, Río Toro), Bolívar (vicinity of El Paují, La Escalera, Río Tonoro), Amazonas (Caño Caname, base of Cerro Marahuaka, San Antonio, San Carlos de Río Negro, base of Sierra de la Neblina, Sierra Parima, Yavita to Maroa). Mérida, Táchira; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Fig. 211.

Clusia panapanari Choisy in DC., Prodr. 1: 559. 1824. [Section Phloianthera]. -Copeicillo.

Clusia colorans Klotzsch ex Engl. in Mart., Fl. Bras. 12(1): 426. 1888.

Clusia microphylla Klotzsch ex Engl. in Mart., Fl. Bras. 12(1): 426. 1888.

Terrestrial to epiphytic tree 5-10 m tall or liana; latex white; fruits ovoid-oblong, ca. 1 cm long × 0.8 mm diameter, markedly transversely rugose. Evergreen lowland forests, riparian forests, 0-400 m; Delta Amacuro (Caño Joba-Suburu east of Caño Sacupana, Caño Jotajana, Curiapo, Río Amacuro, Río Ibaruma, Río Toro), Amazonas (base of Sierra de la Neblina). Guyana, Suriname. Fig. 210.

This species appears to be closely related to Clusia comans.

Clusia parvifolia Maguire, Mem. New York Bot. Gard. 10(1): 58. 1958. [Section Clusiastrum].

Flexuous tree to 3(-10) m tall; scanty clear latex; leaves narrowly elliptic, 3 × 1 cm; petals 8, white, narrowly obovate, ca. 15 mm long; styles ca. 10, elongate, stigmas cuneate. Clusia scrub forests, 1600-2000 m; Bolívar (Ilú-tepui, Roraima-tepui). Endemic.

Clusia parvula (Maguire) Pipoly, comb. nov. [Section Oedematopus]. -Oedematopus parvulus Maguire, Mem. New York Bot. Gard. 23: 869. 1972.

Shrub 2-3 m tall, with few erect branches; leaves sessile. Montane and high-tepui scrub, 1300-2100 m; Bolívar (Cerro Jaua, Cerro Sarisariñama). Endemic.

Clusia phelpsiae Lasser & Maguire, Brittonia 7: 81. 1950. [Section Anandrogyne].

Small tree 2-3 m tall; leaves subsessile, coriaceous, obovate, 5-9 × 3-6 cm; inflorescence terminal, 3-flowered; flowers yellow, petals 5, fleshy, 12-15 × 6-8 mm; carpels 5, fruit urceolate, ca. 25 mm long, 15 mm wide. Open rocky areas and tepui summit thickets, 1800-2200 m; Amazonas (Cerro Yaví). Endemic.

Clusia phelpsiana Maguire, Mem. New York Bot. Gard. 10(1): 51. 1958. [Section Clusiastrum].

Tree 10-15 m; scanty cream-colored latex; sepals 12-15, imbricate; petals 8, white, 3.5 × 2 cm; styles 12-14, elongate. Along streams in montane to upper montane forests, 1500-2300 m; Bolívar (Cerro Uroi, Ilú-tepui). Endemic.

Clusia ptaritepuiensis (Steyerm.) Pipoly, comb. nov. [Section Oedematopus]. -Oedematopus ptaritepuiensis Steyerm., Fieldiana, Bot. 28: 398. 1952.

Climbing shrub or tree to 15 m tall. South-facing slopes, 1700-1800 m; Bolívar (Ptari-tepui). Endemic.

Clusia pusilla Steyerm., Fieldiana, Bot. 28: 390. 1952. [Section Clusiastrum].

Spreading, densely branched low shrub to tree; latex white; leaves sessile to subsessile. Venezuela, Guyana, Suriname, Brazil; 2 subspecies, both in the flora area.

Key to the Subspecies of C. pusilla

1. Leaf blades narrowly obovate to obovate, 5-10 cm wide, the lower surface without visible latex canals; gallery forests in upland savannas ..... subsp. orinocensis

1. Leaf blades widely obovate to suborbicular, 2-5 cm wide, the lower surface with conspicuous latex canals; margins of shrub islands on rocks, savannas below tepuis ..... subsp. pusilla

C. pusilla subsp. orinocensis Maguire, Mem. New York Bot. Gard. 10(1): 56. 1958.

Tree to 6(-10) m tall; stigmas 14-16, separate; fruits pendent, oblong. Edges of montane forests, streamsides, rocky meadows, 600-2000 m; Bolívar (Cerro Guaiquinima) Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yutajé). Brazil (Amazonas: Serra Aracá).

C. pusilla subsp. pusilla

Clusia pusilla subsp. pakaraimae Maguire, Mem. New York Bot. Gard. 10(1): 56. 1958.

Clusia pusilla subsp. jauaensis Maguire, Mem. New York Bot. Gard. 23: 870. 1972.

Shrub 1-4 m tall; flowers in cymes of 3 or 5; petals pure white, staminate flowers with yellow stamens, pistillate flowers with 11 or 12 papillose stigmas. Tepui forests, rocky outcrops, 700-2000 m; Bolívar (Amaruay-tepui, Auyán-tepui, along base of Cerro Guaiquinima, Cerro Jaua, Cerro Sarisariñama, Kavanayén to Parupa, La Escalera, Macizo del Chimantá [Apacará-tepui, Chimantá-tepui], Ptari-tepui, Sierra de Maigualida, Sororopán-tepui). Guyana (Pakaraima Mountains, Kaieteur Plateau), Suriname. Fig. 202.

Clusia radiata Maguire & K.D. Phelps, Bol. Soc. Venez. Ci. Nat. 14: 14. 1952. [Section Criuva].

Shrub to small tree to 5 m; branchlets 4-angled; leaves widely elliptic to suborbicular, 6-10 × 5-7 cm, apex and base widely rounded; pistillate inflorescence of 1 or 3 pendent, white flowers; petals 8, 2-2.5 × 1.5-2.0 cm; carpels 7-10; fruit ovoid, 3.5-4 cm long, 3-4 cm diameter. Rocky outcrops along steep tepui slopes, 1800-2000 m; Amazonas (Cerro Guanay). Endemic.

Clusia renggerioides Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 350. 1860. [Section Phloianthera].

Tree 5-9 m tall; latex white; inflorescence 3-5-flowered, 2-4 cm long; petals 5, crassate, obovate; stigmas triangular, concave; fruits pear-shaped, 5-carpellate. White-sand savannas, riparian forests, 100-200 m; Amazonas (near Cerro Yapacana, Río Pacimoni). Colombia (Amazonas, Vaupés), Brazil (Amazonas).

Clusia rosea Jacq., Enum. Syst. Pl. 34. 1760. [Section Clusia].

Clusia rosea var. colombiana Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 53. 1950.

Large tree to 30 m tall, at least some populations apparently apomictic (only pistillate flowers known; stigmas 6 or 7). Evergreen lowland forests, semideciduous forests, 0-500 m; Bolívar (Río Cuyuni, lower Río Paragua, San Mateo). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Monagas, Nueva Esparta, Sucre, Zulia; Panama, Antilles, Colombia. Fig. 209.

Clusia rotundifolia Gleason, Bull. Torrey Bot. Club 58: 406. 1931 [1932]. [Section Anandrogyne].

Shrub or small tree 1-4 m tall; branchlets 4-angled; leaves orbicular, sessile, stiffly brittle, lustrous on upper surface, 6-9 × 4-8 cm; inflorescences of 9-12 flowers; petals 4, flesh-colored; stamens numerous; stigmas 4-7. Margins of scrub savannas, open rocky areas, 1000-2000 m; Amazonas (Cerro Duida, Cerro Parú, Sierra Parima). Endemic.

Clusia savannarum Maguire, Bull. Torrey Bot. Club 75: 422. 1948. [Section Anandrogyne].

Shrub or small tree to 3 m tall; leaves widely elliptic to suborbicular, 3-9 × 3-7 cm, margins revolute; petioles 0-5 mm long; pistillate inflorescence 1-3(-7)-flowered; petals 4(5), fleshy, 7-10 × 2.5-3.5 mm; fruit subglobose, 10-12 mm long, 13-15 mm diameter. Shrub islands among white-sand savannas, ca. 1000 m; Bolívar (north of Kamá-meru in the central Gran Sabana). Guyana (Kaieteur Plateau).

Clusia schomburgkiana (Planch. & Triana) Benth. ex Engl. in Mart., Fl. Bras. 12(1): 431. 1888. [Section Omphalanthera]. -Polythecandra schomburgkiana Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 229. 1860.

Clusia spruceana Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 13: 346. 1860.

Clusia planchoniana Engl. in Mart., Fl. Bras. 12(1): 432. 1888.

Clusia schomburgkiana var. lakei Maguire, Bol. Soc. Venez. Ci. Nat. 20: 366. 1959.

Shrub or tree 3-15 m tall; latex white; flowers hanging, petals purplish red inside, white outside. Savanna-forest ecotones, granitic outcrops, rocky forested slopes, along streams in rocky montane forests, near waterfalls, iron-rich outcrops; 100-2000 m; Bolívar (Canaima, Cerro Altamira, Cerro Bolívar, Cerro Guaiquinima, Ciudad Piar, El Pao, Gran Sabana, Guri, Ptari-tepui, Río Aro, Río Caroní, upper Río Paragua, Roraima-tepui), Amazonas (Cerro Coro Coro, near Cerro Yapacana, Cerro Yutajé, Culebra, La Esmeralda, Maroa, Piedra Cocuy, Piedra Nunca north of Piedra Cocuy, Río Casiquiare, Río Temi, Río Yatúa). Colombia (Vaupés), Guyana, Brazil (northern Amazonas). Fig. 195.

Clusia schomburgkii Vesque, Epharmosis 3: 7, pl. 14, 15. 1892. [Section Clusiastrum]. -Žru-yek (Arekuna).

Clusia roraimana Steyerm., Fieldiana, Bot. 28: 391. 1952.

Tree to 5 m tall; petals white. Along streams in forests on steep slopes, 1000-1300 m; Bolívar (road between Luepa and Santa Elena, Macizo del Chimantá, lower slopes of Roraima-tepui). Guyana (Mount Roraima, Kaieteur Plateau).

Clusia sipapoana (Maguire) Pipoly, comb. nov. [section Quapoya]. -Quapoya sipapoana Maguire, Mem. New York Bot. Gard. 10(4): 27, fig. 30 A-H. 1961.

Epiphytic scandent shrub or tree to 15 m high; leaves obovate-elliptic, rounded to slightly retuse at the apex. Evergreen lowland to montane forests, 100-800 m; Amazonas (Cerro Sipapo, Río Yatúa, Sierra de la Neblina). Guyana, Peru. Fig. 197.

Clusia spathulaefolia Engl. in Mart., Fl. Bras. 12(1): 412. 1888. [Section Brachystemon]. -Cupí, Cupí banero hoja gruesa.

Free-standing tree to 20 m tall; branches 4-angled; latex white; leaves stiffly coriaceous, oblanceolate to obovate, 12-28 × (3.5-)10-17 cm, apex obtuse to broadly rounded, base attenuate and decurrent on the petiole; petals 5, thick-fleshy, 5-7 mm long, 2-2.5 mm wide; stamens numerous; fruit 5- or 6-carpellate, depressed-globose, strongly ribbed, 1.5-2 cm long, 1.1-2.8 cm diameter. Gallery forests along black-water rivers, Río Negro caatinga, montane forests on tepui slopes, 100-1700 m; Amazonas (Caño San Miguel, Caño Yagua, near Cerro Yapacana, Guarinuma, Maroa, Pimichín, Río Atacavi, Río Mawarinuma, Cerro Marahuaka slopes, near San Carlos de Río Negro, Sierra Unturán). Colombia (Amazonas, Caquetá, Vaupés), Brazil. Fig. 185.

Clusia steyermarkii Maguire, Mem. New York Bot. Gard. 10(1): 52. 1958. [Section Clusiastrum].

Tree to 8 m tall; leaves oblanceolate, 25-30 × 7-8 cm; flowers 3 per inflorescence. Montane gallery forests, 1200-1400 m; Bolívar (Macizo del Chimantá [Río Tírica between Apacará-tepui and Toronó-tepui], Sierra de Maigualida). Endemic.

Clusia tabulamontana Maguire, Bull. Torrey Bot. Club 75: 420. 1948. [Section Clusiastrum].

Tree to 5 m tall. Scrub forests, 500-1100 m; Bolívar (Amaruay-tepui, La Escalera). Suriname (Tafelberg). Specimens from Amaruay-tepui [Liesner & Holst 20746 (MO); 20860 (MO)] are provisionally referred to this species because of their elliptic, prominently brochidodromous leaves, solitary flowers, terete branchlets, and sub-basally attached articulated stigmas. The Amaruay-tepui population, however, has fruits ca. 3 (not 1.5) cm diameter and 4-5 (not 3) cm long, peduncles to 7 (not 6) cm long, and styles 1 mm longer than the type collection from Suriname.

Clusia troncosii Maguire, Moscosoa 4: 215. 1986. [Section uncertain].

Tree 3-10 m tall; latex yellow; sepals in 3 pairs; stigmas flat and sessile, ovary with (8-)14-18(-20) carpels. Lowland savannas and forest edges, 100-200 m; Amazonas (base of Cerro Sipapo, La Esmeralda and base of Cerro Duida, Río Coro west to Cerro Yutajé Coro, base of cerro Yapacana). Endemic.

Clusia viscida Engl. in Mart., Fl. Bras. 12(1): 422. 1888. [Section Clusia].

Small tree 4-8 m tall; latex white to yellow; fruit globose with red longitudinal lines. Río Negro caatinga, seasonally flooded forests, 100-200 m; Amazonas (Río Manapiare, Río Mawarinuma, San Carlos de Río Negro). Colombia, Ecuador, Peru, Brazil.

Clusia sp. A [Section Oedematopus].

Tree ca. 8 m tall. Montane forests, ca. 1500 m; Amazonas (Caño Piedra on Cerro Sipapo massif). Endemic.

Clusia sp. B [Section Oedematopus].

Shrub to 2.5 m tall. Scrub forest, ca. 1600 m; Bolívar (Cerro Guaiquinima). Endemic.

Clusia sp. C [Section Oedematopus].

Shrub or tree to 10 m tall; latex white. Lower montane to montane forests, along streamsides, 500-1500 m; Bolívar (northeast of Cerro Marutaní, Icabarú, Raudales de Maihia on Río Paragua), Amazonas (Cerro Yutajé, Río Coro Coro west to cerro Yutajé). Endemic.

Clusia sp. D [Section Oedematopus].

Shrub or small tree 2-10 m tall; latex white. Gallery forests, scrub forests, talus slopes, savanna edges, 400-1100 m; widespread in Bolívar and Amazonas. Guyana. Fig. 199.

TOP

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

 
 
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