Tovomita fructipendula

Description — Trees up to 12 m tall, prop roots inconspicuous; exudate yellow, abundant. Petioles 0.6–2.2 cm long, green or reddish, smooth, lenticels absent. Leaf blades 3–8(–13) × 1.3–4.5 cm, greenish in sicco, black dots absent, subcoriaceous, elliptic to ovate, base convex to decurrent, apex acuminate; papillae, lenticels and fungal spots absent; exudate canals inconspicuous. Venation: secondary veins 10–17 pairs, 1–3 mm apart from each other, forming angle 60°–70° to midvein, immersed in both surfaces, straight near margin; intersecondary veins present, one per intercostal area, similar to secondary veins, parallel to major secondary,> 50 % of subjacent secondary length; tertiary veins inconspicuous; intramarginal vein present. Inflorescences: ♂ lax cyme with 3 basal branches up to 9 flowers, lacking terminal flower, ♀ dichasium, lenticels absent. Pedicels 6–20 mm long, green, distal and proximal portion with same gauge, proximally articulated on lateral flowers of dichasia; ca- Fig. 38. Distribution map of Tovomita fructipendula View in CoL – Amazon Basin (light grey), Atlantic forest (dark grey), Guiana Shield (dark grey striped). lyptrae and lenticels absent. Floral buds 4–6 mm long, spheroid, apex rounded, len- 2006, fr., P. O. Garcia s.n. (CESJ no. 49168!). RIO DE ticels absent. Sepals 2–4, 4.5–8 × 3.5–7 mm, circular, JANEIRO: Teresópolis, Serra dos Órgãos, 20 Nov 2006, apex rounded, light green to light yellow; petals 4, 8–9 bud, E. J. Lucas & al. 596 (RB!). × 3–7 mm, oblong, deflexed, apex rounded, pale yellow. Staminate flowers: stamens 50–60, 3–4.5 mm long, 24. Tovomita gracilipes Planch. & Triana in Ann. Sci. heterodynamous; filaments terete, white; anthers 0.5– Nat., Bot., ser. 4, 14: 280. 1860. – Lectotype (des- 0.8 mm long, connective not exceeding thecae; pistillode ignated here): Brazil, Brésil septentrional, San Carc. 0.5 mm long, conic, greenish-white. Pistillate flowers: los sur le Rio Negro, 1853–1854, R. Spruce 3079 (P staminodes 50–60, 3–3.5 mm long, white; ovary 4–5 mm [P00089914]!; isolectotypes: BR [BR0000005915187] long, not costate or lobed, 4-locular, pale yellow, stig- photo!, G [G00355923] photo!, GH [GH00067910]!, K mas 4, sessile, c. 1 mm in diam. Capsules fleshy, 3.9–3.5 [K000488520]!, K [K000488521]!, NY [NY00579021]!, × 1.9–2.7 cm, 4-septate, spheroid when closed, slightly P [P00089915]!, W [no. W-1889-0309881]!). 4-costate when immature, not lobed, rostrum present, = Tovomita rubella Spruce ex. Planch. & Triana in c. 5 mm long, epicarp smooth, green when immature and Ann. Sci. Nat., Bot., ser. 4, 14: 275. 1860, syn. mature, mesocarp red; sepals and petals caducous, stami- nov. – Lectotype (designated here): Brazil, Brésil nodes and stigmas persistent. Aril orange. Fig. 41. boréal, San Carlos sur del Rio Negro, 1853–1854,

R. Spruce 3143 ( P [ P00093871 ]!; isolectotypes: BM Iconography — Illustration available in Marinho & al. [ BM000047392 ] photo!, BR [BR 0000005852802] (2016b: 761). photo!, G [ G00355926 ] photo!, G [ G00355927 ]

photo!, GH [GH00067915]!, GOET [GOET003980] Distribution — Brazil (Bahia, Minas Gerais, Rio de Ja- photo!, K [K000488534]!, K [K000488535]!, neiro). Fig. 42. MPU [MPU014283]!, NY [NY00578995]!,

NY [NY00578996]!, P [P00093870]!, RB Conservation status — Least Concern (LC, Marinho & [RB00539007]!, TCD [TCD0000898] photo!, W [no. Beech 2019). W-1889-0123962]!).

= Tovomita acuminata Engl. in Martius & al., Fl. Bras. Recognition and discussion — Tovomita glazioviana 12(1): 449. 1888. – Lectotype (designated here): shares with the Atlantic forest related species T. fructi- Brazil, Amazonas, Manaus, province de Alto Amapendula and T. iaspidis the small leaves, white stamens, zonas, Jul 1851, R. Spruce 1716 (M [M0086205]!; and 4-carpellate ovary (see Table 1). This species can be isolectotypes: K [K000488536]!, K [K000820676]!, distinguished from T. fructipendula by the spheroid floral W [no. W-1889-0309908]!). buds and smooth epicarp (vs oblong floral buds and as- = Tovomita duidae Steyerm. in Fieldiana, Bot. 28: 400. perous epicarp in T. fructipendula ), and from T. iaspidis 1952. – Holotype: Venezuela, Amazonas, southeastby the small and greenish-white pistillode (vs large red- ern-facing slopes along Caño Negro (tributary of dish pistillode in T. iaspidis ). Caño Iguapo), 305–1095 m, 25–26 Aug 1944, J. A.

Steryermark 57975 (F [F0054535F] photo!; isotype: Selected specimens examined — BRAZIL: BAHIA: Al- NY [NY00077363]!). madina, Serra do Corcovado / São Domingos , 25 Jan 2014, fl. ♂, L. C. Marinho & al. 602 ( CEPEC!, HUEFS!). Description — Shrubs or trees up to 6 m tall, prop MINAS GERAIS: Juiz de Fora , REBIO Santa Cândida, Nov roots inconspicuous; exudate yellow, scarce. Petioles 2.1–2.8 cm long, green, smooth, lenticels absent. Leaf blades (5.5–)7.8–21.5 × (0.8–) 3.4–8.5 cm, brown-orangish abaxially and dark brown to purplish-red adaxially in sicco, black dots absent, subcoriaceous to coriaceous, oblong to ovate, base convex to decurrent, apex acuminate, usually with drip tip; papillae, lenticels and fungal spots absent; exudate canals inconspicuous. Venation: secondary veins 5–10 pairs, 7–24 mm apart from each other, forming angle 65°–70° to midvein, prominent in both surfaces, arcuate and connecting near margin; intersecondary veins present, one to three per intercostal area, thinner than secondary veins, parallel to secondary or basiflexed, c. 50 % of subjacent secondary length; tertiary veins percurrent sinuous; intramarginal vein absent. Inflorescences: ♂ lax cyme with 5 basal branches and up to 125 flowers, lacking terminal flower, ♀ cyme with 5 basal branches and up to 60 flowers, dichasia sometimes reduced lacking central flower, lenticels absent. Pedicels 6–25 mm long, green, distal and proximal portion with same gauge, proximally articulated on lateral flowers of dichasia, sometimes in central flower; calyptrae and lenticels absent. Floral buds 4–5 mm long, spheroid, apex rounded. Sepals 4, 4.5–6 × 2.5–4 mm, circular to oblong, apex rounded, greenish; petals 4, 3–6 × 1.5–5 mm, oblong, deflexed, apex rounded, white. Staminate flowers: stamens 40–45, 2–3 mm long, heterodynamous; terete, white; anthers c. 0.5 mm long, connective not exceeding thecae; pistillode conic, white. Pistillate flowers: staminodes 40–45, 2–2.5 mm long, white; ovary c. 3 mm long, 4-locular, white, styles c. 1 mm long, stigmas 4, c. 0.5 mm in diam. Capsules fleshy, 1.3–2.4 × 1–2.1 cm, 4-septate, napiform when closed, not costate, slightly lobed, rostrum present, c. 1 mm long, epicarp smooth, green when immature and mature, mesocarp red; sepals, petals, staminodes and stigmas persistent. Aril orange. Fig. 43 .

Distribution — Brazil (Amazonas, Pará), Colombia (Amazonas, Antioquia, Caquetá), Peru (Loreto), Venezuela (Amazonas, Bolivar). Fig. 36.

Conservation status — Least Concern (LC, Marinho & Beech 2019).

Nomenclatural notes — For the lectotypes of Tovomita gracilipes and T. rubella , we chose the P specimens since this is the herbarium where Planchon and Triana handled the specimens. As for the lectotype of T. acuminata , the specimen deposited at B was destroyed during World War II (see photograph in GH), so we chose the M specimen, which is the best preserved among known duplicates and the only duplicate that has a “det. Engler”.

Recognition and discussion — Specimens of Tovomita gracilipes from the Dpto. Amazonas, in Venezuela, have smaller and narrower leaves (5.5–9.5 × 0.8–2.4 cm) than the specimens from other regions (7.8–21.5 × 3.4– 8.5 cm). Also, the fruits are smaller and slightly constricted at the base. Specimens with this characteristic resemble T. duidae , a species here treated as a synonym of T. gracilipes .

Tovomita gracilipes is part of a species group with small floral buds and flowers with reduced subclavate stamens. Its leaves and flowers are similar to those of T. guianensis , from which it can be distinguished by the 5–10 (vs 11–15) pairs of secondary veins, and the white (vs yellow, Fig. 45C) petals and stamens ( Fig. 43C). Leaf venation and floral bud size of this species are similar to those of T. schomburgkii (see discussion under the latter).

Selected specimens examined — BRAZIL: AMAZONAS: Manaus, Reserva Florestal Ducke, 02°53'S, 59°58'W, 17 Oct 1996, fr., M. A GoogleMaps . D. Souza & al. 265 ( GH!, INPA!) . — COLOMBIA: AMAZONAS: Araracuara, Río Caquetá, 00°39'S, 72°08'W, 9 Oct 1992, fr., A GoogleMaps . Duque & A . Posada 4121 ( COAH!). ANTIOQUIA: Anori, Corregimiento de Liberia , sector Usurá, 1 Nov 2010, fr., A . Taborda & L . Londoño 409 ( HUA!). CAQUETÁ: Araracuara , 00°37'S, 72°23'W, 8 Mar 1994, sterile, H GoogleMaps . Vester & O . Roman 880 ( COAH!) . — PERU: LORETO: Requena, Dpto. Jenaro Herrera, 1 km NE of Arboretum 04°55'S, 73°45'W, 20 Nov 1988, fr., D. C GoogleMaps . Daly & al. 5725 ( NY!) . — VENEZUELA: AMAZONAS: Vicinity of Cerro Neblina Project base camp, 00°55'N, 66°10'W, 140 m, 27 No 1984, fr., T. B GoogleMaps . Croat 59352 ( MO photo!, NY!). BOLÍVAR: Raul Leoni, 13 km de La base El Cerro Camarón, 05°43'N, 64°07'W, 250 m, 30 Oct – 2 Nov 1988, fr., G GoogleMaps . Aymard & A . Fernandez 7245 ( NY!) .