15. TOVOMITA Aubl., Hist. Pl. Guiane 956, t. 364. 1775.

Marialvaea Mart., Nov. Gen. Sp. Pl. 2: 82. 1826 [1827].
Tovomitidium Ducke, Arq. Inst. Biol. Veg. 2: 61. 1935.

by Nidia L. Cuello

Trees or shrubs, glabrous, the latex often white or yellowish. Stems sometimes buttressed or stilt-rooted. First internode of axillary shoots characteristically long, ± curved. Leaves loosely clustered, decussate, the secondary veins sometimes forming a submarginal vein; petioles usually with an adaxial margined pit. Inflorescences mostly terminal, often pyramidal, cymose panicles; staminate inflorescences often more floriferous than pistillate ones; bracteoles usually present; pedicels often thickening in fruit. Flowers usually unisexual (plants monoecious, dioecious, or rarely polygamo-monoecious or -dioecious), the buds often minutely apiculate, completely enclosed by the outer sepals which are usually completely fused, splitting lengthwise into 2-4 sepals, an inner whorl of sepals sometimes also present; petals sometimes fused in bud and splitting lengthwise into 2-8 segments, or the petals free and distinct in bud, decussate. Staminate flowers with numerous stamens; filaments free, ± linear; anthers small, the 2 lobes separated by a connective (produced into a minute apiculum); pistillate flowers with staminodes similar to the stamens of the staminate flowers. Ovary 4-6-carpellate, ovules 1 per carpel; styles usually short, or produced into prongs in fruit; stigmas ovoid to discoid. Fruit a fleshy, septicidal capsule, often brightly colored within, some locules frequently not developing. Seeds large, 1 per locule, enveloped by a fleshy venose aril; cotyledons minute.

Tropical America; ca. 45 species, ca. 30 species in Venezuela, 24 of these in the flora area.

Key to the Species of Tovomita

1. Secondary leaf veins well differentiated and usually > 10 mm apart, curving toward the margin with distinct but fainter intersecondary veins ..... 2

1. Secondary and intersecondary leaf veins similar, less than 3 mm apart, or if with stronger secondary veins, these less than 10 mm apart and either ascending and curving upwards or nearly straight and perpendicular to the midvein ..... 14

2(1). Petioles less than 5 mm long, blades mostly obovate, base attenuate and auriculate ..... T. sp. A

2. Petioles > 5 mm long; blades other than obovate, if attenuate at the base, then not auriculate ..... 3

3(2). Leaf blades broadly oblong-elliptic, 30-45 × 20-25 cm; margins of axillary pits conspicuously raised, 6-9 mm diameter ..... T. sp. B

3. Leaf blades various in shape and size, less than 26 cm long; margins of axillary pits less than 5 mm diameter ..... 4

4(3). Leaf blades normally less than 12 × 2-5 cm, mostly narrowly elliptic to oblanceolate ..... 5

4. Leaf blades > 12 × 5 cm, shape various ..... 8

5(4). Secondary leaf veins mostly straight, usually more than 7 per side ..... 6

5. Secondary leaf veins curving upwards, usually 4-6 per side ..... 7

6(5). Leaf blades with black-dotted latex canals on lower surface; lateral veins mostly straight, ascending at angles of 40-50°; flower buds oblong in pistillate specimens, staminodes filiform ..... T. aff. glazioviana

6. Leaf blades without black-dotted latex canals on lower surface; lateral veins barely ascending, at angles of 60-80° from the midvein; flower buds subglobose-apiculate in pistillate specimens, staminodes fusiform ..... T. brevistaminea

7(5). Leaf blades membranaceous, secondary veins forming a marginal vein 2-6 mm from the margin ..... T. gracilipes

7. Leaf blades coriaceous to subcoriaceous; secondary veins extending independently to the margin ..... T. elliptica

8(4). Margins of axillary pits with a raised border, the pits 4-5 mm diameter ..... 9

8. Margins of axillary pits not conspicuously raised, the pits usually less than 3 mm diameter ..... 10

9(8). Leaf blades broadly obovate-elliptic, 15-26 × 9-14 cm, densely covered with black-dotted latex canals on lower surface; flower buds oblong-apiculate, 8-10 mm long; stamens 6-8 mm long ..... T. macrophylla

9. Leaf blades narrowly elliptic, 12-18 × 5-8 cm, without black-dotted latex canals on lower surface; flower buds globose-apiculate, 4-5 mm long; stamens less than 1.5 mm long ..... T. fanshawei

10(8). Flower buds globose, 6-9 mm wide, pedicels stout ..... 11

10. Flower buds usually ovoid or oblong, if globose, then less than 6 mm wide and pedicels slender ..... 12

11(10). Leaf blades rounded at base, secondary veins strongly raised on lower surface, abruptly curving upwards from the midvein along their length and joining 2-3 mm from margin; pedicels and inflorescence rachis rugose but not lenticellate ..... T. atropurpurea

11. Leaf blades obtuse to cuneate at base; secondary veins obvious but not strongly raised on lower surface, ascending at angles of 55-70°, mostly straight, curving upwards close to the margin, but not joining; pedicels and inflorescence rachis conspicuously rugose-lenticellate ..... T. albiflora

12(10). Leaf blades mostly oblong to obovate-oblong; flower buds ovoid or oblong, sometimes subglobose-apiculate, mostly drying brown, 6-10 × 4-6 mm; stamens 5-7 mm long ..... T. longifolia

12. Leaf blades mostly oblong-elliptic to lanceolate; flower buds globose, or if subglobose, then not apiculate, mostly drying black, 3-6 mm long; stamens 3-4 mm long ..... 13

13(12). Leaf blades membranaceous, usually drying green, mostly oblong-elliptic to slightly obovate, obtuse to mucronate at apex ..... T. schomburgkii

13. Leaf blades subcoriaceous to coriaceous, drying reddish or blackish, mostly lanceolate, acuminate at apex ..... T. rubella

14(1). Secondary and intersecondary veins distinct, nearly straight, almost perpendicular to the midvein, or the secondary veins usually ascending and 4-10 mm apart ..... 15

14. Secondary and intersecondary veins nearly indistinguishable, closely parallel, less than 3 mm apart, sometimes inconspicuous on lower surface ..... 20

15(14). Secondary veins varying in both angle and direction, usually ascending and slightly curving upwards, sometimes not conspicuous on lower surface ..... 16

15. Secondary veins all nearly straight, almost perpendicular to the midvein, prominent on lower surface ..... 18

16(15). Leaf blades with conspicuous black-dotted latex canals on lower surface; secondary veins ascending at angles of 40-50° ..... T. aff. glazioviana

16. Leaf blades without black-dotted latex canals, or if present, then very few and distant; secondary veins nearly straight, ascending at angles of 60-80° ..... 17

17(16). Leaf blades mostly narrowly elliptic-lanceolate, 3.2-7.2 × 1.5-3 cm, apex acuminate to cuspidate; staminate flower buds oblong-ovoid, drying brown, 6-7 × 3.5-4 mm; pedicels 9-11 mm long ..... T. sp. C

17. Leaf blades mostly obovate to oblong-elliptic, 6-15 × 3-7 cm, apex obtuse to mucronate; staminate flower buds globose, drying black, pistillate buds sometimes drying brown, 2.5-3 mm diameter; pedicels 3-8 mm long ..... T. brevistaminea

18(15). Leaf blades mostly oblong; fruits pyriform, with capitate stigmas and 4-5 mm long styles ..... T. spruceana

18. Leaf blades mostly elliptic or narrowly elliptic to lanceolate; fruits oblong, with triangular-ovate, sessile stigmas ..... 19

19(18). Leaf blades mostly elliptic, base and apex obtuse; fruits mostly oblongoid, 12-15 × 7-9 mm, stigmas 2.5 × 3 mm ..... T. umbellata

19. Leaf blades mostly narrowly elliptic to lanceolate, apex acuminate, base attenuate; fruits ellipsoid, 25-33 × 12-15 mm, stigmas 1.5 × 2 mm ..... T. eggersii

20(14). Leaf blade truncate or slightly auriculate at the base; petiole short, appearing winged ..... T. weddelliana

20. Leaf blade acute to attenuate at the base, not auriculate; petiole obvious, not winged ..... 21

21(20). Leaf blades mostly obovate or oblanceolate ..... 22

21. Leaf blades mostly lanceolate or oblong-elliptic to lanceolate ..... 23

22(21). Leaf blades 7.5-15 cm long, midvein on lower surface drying concolorous; petioles stout, 2-3 mm wide, 18-20 mm long; flower buds globose, 5-7 mm diameter ..... T. sp. F

22. Leaf blades 4-9 cm long, midvein on lower surface discolorous; petioles slender, ca. 1 mm wide, 10-13 mm long; flower buds cylindric-apiculate, 5-6 × 2 mm ..... T. sp. D

23(21). Leaf blades thinly membranaceous or papyraceous ..... 24

23. Leaf blades subcoriaceous to coriaceus ..... 25

24(23). Trees; leaves 9-20 × 3-5.5 cm; fruits 2-2.2 × 1.5-2 cm; pedicels on fruits short and stout, 5-6 mm long, 2-2.5 mm wide ..... T. sp. E

24. Shrubs; leaves 6-7 × 1.8-2 cm; fruits 1-2 × 0.5-0.8 cm; pedicels on fruits long and slender, 19-20 mm long, ca. 1 mm wide ..... T. aff. brasiliensis

25(23). Leaf blades thick and stiffly coriaceous, broadly obovate-oblong, 15-25 × 6-10 cm, apex broadly obtuse, abruptly mucronate; petioles 3-3.5 mm wide; flower buds cylindric-cuspidate, 1.5-2 mm wide ..... T. tenuiflora

25. Leaf blades subcoriaceous, oblong, oblong-elliptic to oblong-lanceolate, 12-18 × 1.7-7 cm, apex subobtuse to acuminate; petioles 1.5-2 mm wide; flower buds oblong-ovoid, subglobose or conical, 3-4 mm wide ..... 26

26(25). Leaf blades 1.7-4 cm wide; flower buds narrowly conical, 10-13 mm long, 3-4 mm wide at the base, less than 2 mm wide at the middle ..... T. acutiflora

26. Leaf blades 4-7 cm wide; flower buds subglobose, oblong-ovoid to shortly apiculate, 6-8 mm long, 3-4 mm wide at the middle ..... T. carinata

Tovomita acutiflora M.S. Barros & G. Mariz, Acta Amazon. 12: 291. 1982.

Tree to 10 m tall. Montane forests, 400-1100 m; Bolívar (Gran Sabana, Río Pacairao near Kavanayén, Río Uaiparú near its mouth in Río Icabarú). Guyana, Brazil (Amazonas).

Tovomita albiflora A.C. Sm., Bull. Torrey Bot. Club 67: 293. 1940. Wakome (Arekuna).

Monoecious tree 7-12 m tall; flowers fragant; petals and stamens white; inflorescence branches stout, rugose, lenticellate; fruit rusty brown-rugose without, splitting open with recurved valves exposing dark red within; aril dark orange. Lower montane to montane forests, 500 1400 m; Bolívar (near Arabopó, Cerro Venamo, km 108 124 on El Dorado to Santa Elena road, Roraima-tepui). Guyana.

Tovomita atropurpurea Steyerm., Fieldiana, Bot. 28: 399. 1952. -Wakami-yek.

Tree to 8 m tall; sepals pale green with dull rose margins; petals, filaments, and anthers dull purple-red. Forested lower tepui slopes, 1500-1600 m; Bolívar (Ptari-tepui). Endemic. Fig. 225.

Tovomita aff. brasiliensis (Mart.) Walp., Repert. Bot. Syst. 1: 392. 1842. -Marialvaea brasiliensis Mart., Nov. Gen. Sp. Pl. 2: 83, t. 167. 1826 [1827].

Shrub to small tree 4-5 m tall; petals white, fruits green; aril orange. Shrubby savannas on white sand, edges of forests along black-water rivers, 100-200 m; Amazonas (Laguna Pasiba, Río Casiquiare and tributaries below Río Pasimoni and Río Siapa, Tamatama). Peru.

The material from the flora area closely matches the description of Tovomita brasiliensis (Mart.) Walp. However, it does not match the type material of that name very well and perhaps belongs to another species or is an undescribed species.

Tovomita brevistaminea Engl. in Mart., Fl. Bras. 12(1): 446, t. 96, fig. 3. 1888. -Bojoro, Coloradito, Coloradito negro, Mangle montañero.

Tovomita guianensis Planch. & Triana, Ann. Sci. Nat. Bot. sér 4, 14: 268. 1860, non Aubl. 1775.

Tovomita jenmanii Engl. in Mart., Fl. Bras. 12(1): 447, t. 96, fig. 3. 1888.

Tree 4-15 m tall with stilt roots; petals creamy white; fruits green. Riparian and evergreen lowland to lower montane forests, 0-600 m; Delta Amacuro (Caño Joba-Suburu east of Caño Sacupana, Curiapo, Río Amacuro, Río Cuyubini, Río Grande), Bolívar (Altiplanicie de Nuria, Río Nichare, Río Toro, road between Upata and San Félix). Guyana, Suriname, Brazil. Fig. 226.

Tovomita carinata Eyma, Polygon. Guttif. Surinam 42. 1932. -Naranjillo.

Tree 3-18 m tall; flowers fragrant; stamens yellow; latex yellow. Lowland and lower montane forests over lateritic crusts, 100-600 m; Amazonas (Caño Galipero, Cerro Vinilla, Serranía Batata in Cuao-Sipapo massif). Suriname, Brazil (Amazonas), Bolivia.

Tovomita eggersii Vesque, Epharmosis 3: 19, pl. 94. 1892. -Coloradito, Cupi, Naranjillo negro, Uakamiek (Arekuna).

Small tree 3-10(-15) m tall; latex yellow; flowers white; fruits green, aril red. Evergreen lowland, lower montane, and riparian forests, 100-900 m; Bolívar (Altiplanicie de Nuria, Amaruay-tepui, Quebrada Los Brasileros near Icabarú, upper Río Caroní, Río Kukenán basin, Río Paragua, Río Parguaza), Amazonas (Cerro Yapacana, Río Atabapo, Río Atacavi). Miranda, Sucre; Trinidad. Fig. 227.

Tovomita elliptica Engl. in Mart., Fl. Bras. 12(1): 453. 1888.

Tree 5 m tall. Evergreen lowland forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Endemic.

Tovomita fanshawei Maguire, Bull. Torrey Bot. Club 75: 436. 1948.

Tree 8-22 m tall; latex yellow; leaves coriaceous; flowers fragrant, light green; stamens white; fruits green. Seasonally flooded evergreen lowland to montane slope forests, 100-1100 m; Bolívar (east of Los Pijiguaos), Amazonas (Cuao-Sipapo massif, Río Asisa, Río Autana, Río Coro Coro west of Cerro Yutajé). Colombia, Guyana, Peru, Brazil (Mato Grosso), Bolivia.

Tovomita aff. glazioviana Engl. in Mart., Fl. Bras. 12(1): 445. 1888.

Small tree to 15 m tall; latex yellow; flowers white. Nonflooded evergreen lowland forests, 100-200 m; Amazonas (near La Esmeralda, upper Río Cunucunuma, Río Marueta, Río Ventuari).

Collections from Venezuelan Amazonas closely match Tovomita glazioviana, but until now the species has been reported only from Rio de Janeiro, Brazil. Further study is required to determine if this is a true disjunction or not.

Tovomita gracilipes Planch. & Triana, Ann. Sci. Nat. Bot. sér 4, 14: 280. 1860. -Tariji (Yekwana), Ua-kami-yek (Arekuna).

Tovomita duidae Steyerm., Fieldiana, Bot. 28: 400. 1952.

Shrub to small tree 2-10 m tall; latex yellow; sepals pale green; petals greenish white; fruits green. Riparian forests, nonflooded evergreen lowland to upland forests, secondary forests, open areas, 100-1100 m; Bolívar (Cerro Camarón near Cerro Guiaquinima, El Cácaro between Río Caura and Río Paragua, Entrerios, northern Gran Sabana, Río Anacapra in Río Paragua basin, Río Tabaro near Río Nichare, near Santa María de Erebato, Urimán), Amazonas (Cerro Duida, Río Asisa, Río Cunucunuma near Culebra, Río Yureba, San Carlos de Río Negro, Sierra de la Neblina along Río Mawarinuma). Brazil.

Tovomita longifolia (Rich.) Hochr., Annuaire Conserv. Jard. Bot. GenĖve 21: 66. 1919. -Clusia longifolia Rich., Actes Soc. Hist. Nat. Paris 1: 113. 1792. -Tovomita richardiana Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 273. 1860, nom. nov. illeg. for Clusia longifolia Rich. -Cupi, Hiel de agua dulce, Mangle negro, Wakamiyek (Arekuna).

Tree 5-20 m tall; latex cream, white, or yellow; flowers white, fragrant; fruits green to purple when ripe; aril orange. Riparian to montane forests, 50-1300 m; Delta Amacuro (Río Toro), Bolívar (Gran Sabana, Río Canaracuni, Río Caruay, Santa María de Erebato), Amazonas (widespread). Apure; Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amazonas). Fig. 228.

Tovomita macrophylla (Poepp.) Walp., Repert. Bot. Syst. 1: 392. 1842 -Marialvea macrophylla Poepp., Nov. Gen. Sp. Pl. 3: 14. 1845 [1840].

Tree ca. 16 m tall; latex white; flowers white, fragrant. Amazonas (Río Casiquiare, Río Pasiba, Río Pasimoni). Colombia, Guyana, Brazil.

Tovomita rubella Spruce ex Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 275. 1860.

Tree to 5 m tall; fruit green. Riparian forests, lowland to montane forests, 50-1200 m; Amazonas (Caño San Miguel, Cerro Marahuaka, San Carlos de Río Negro). Endemic.

Tovomita schomburgkii Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 274. 1860.

Tree 5-20 m tall; latex white, turning yellow; fruits green, red inside. Riparian forests, lower to upper montane and tepui slope forests, 400-1300 m; Bolívar (Amaruay-tepui, Aparurén, near El Paují, Quebrada Los Brasileros near Icabarú, Río Paragua, near Santa María de Erebato), Amazonas (Sierra Parima). Colombia, Guyana, Suriname, French Guiana, Brazil.

Tovomita spruceana Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 277. 1860. -Conoví (Baniva), Cupi, Cupi rebalsero, Palo perro de agua, Peramán.

Small tree 5-10 m tall; latex yellow; petals white, green, or yellow; stamens white. Rocky hills in open growth, riparian forests, seasonally flooded forests, borders of Mauritia palm swamps, 50-1200 m; Bolívar (between Luepa and Cerro Venamo, northeast of Maripa, Río Parguaza), Amazonas (widespread). Apure; Colombia, Brazil (Amazonas).

This species is used to treat diarrhea.

Tovomita tenuiflora Benth. ex Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 272. 1860.

Tree 10-15 m tall. Riparian to lower montane forests, 100-1100 m; Bolívar (Icabarú, Kavanayén, Río Uaiparú), Amazonas (Caño Colorado on Cuao-Sipapo massif, Río Casiquiare, Río Pasiba, Río Pasimoni, near Solano). Brazil.

Tovomita umbellata Benth., London J. Bot. 2: 367. 1843. -Guacamía, Palo culebra.

Tovomita nigrescens Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 276. 1860.

Tovomita stigmatosa Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 275. 1860.

Tovomita cephalostigma Vesque, Epharmosis 3: 207, pl. 96. 1892.

Tree 2-15 m tall; latex yellow; flowers pale green to white; fruits green. Riparian forests, nonflooded evergreen lowland to lower montane forests, often on white sand, 100-1100 m; Bolívar (Kavanayén, Río Uairén and Quebrada O-paru-má in Río Kukenán basin), Amazonas (Caño San Miguel, El Pozo near San Fernando de Atabapo, Isla Ratón, Río Atacavi, Río Casiquiare, Río Coro Coro west of Cerro Yutajé, Río Orinoco south of Isla Cárida). Miranda; Guyana, Peru, Brazil (Acre).

Tovomita weddelliana Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 14: 277. 1860. -Acami-yek (Arekuna).

Tovomita angustata Steyerm., Fieldiana, Bot. 28: 399. 1951.

Tree 4-20 m tall; fruiting racemes pendent. Montane forests, 700-1200 m; Bolívar (Kavanayén, Sierra de Lema). Aragua, Distrito Federal, Miranda, Yaracuy; Costa Rica, Panama, Colombia, Guyana, Ecuador, Peru, Bolivia. Fig. 229.

Tovomita sp. A

Tree 3-10 m tall; latex yellow; flowers green; stamens white; fruits brown. Evergreen lowland to lower montane forests over granite, 100-600 m; Amazonas (Culebra, Cuao-Sipapo massif, 78 km northeast of Puerto Ayacucho). Peru.

Tovomita sp. B

Tree 4-5 m tall; latex yellow. Riparian forests, 100-200 m; Amazonas (Río Sipapo). Endemic.

Tovomita sp. C. -Cafecillo.

Tree to 8 m tall. Evergreen lowland forests, 100-200 m; Amazonas (Santa Bárbara del Orinoco). Endemic.

Tovomita sp. D

Small tree 3 m tall; flower buds whitish. Lower montane forests, 700-800 m; Amazonas (Sierra de la Neblina). Endemic.

Tovomita sp. E

Small tree 4-6 m tall. Semidry montane forests, ca. 1200 m; Bolívar (Uaipán-tepui). Endemic.

Tovomita sp. F

Tree 4-6 m tall; petals white; anthers yellow; latex yellow. Seasonally flooded forests, 100-200 m; Amazonas (lower Río Casiquiare and tributaries, Río Negro). Colombia, Brazil (Minas Gerais).

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