Tovomita fructipendula (Ruiz & Pav.) Cambess.

epicarp smooth, green when immature and mature, meso- 21. Tovomita fructipendula (Ruiz & Pav.) Cambess. View in CoL in carp purplish-red; sepals, petals, staminodes and stigmas Mém. Mus. Paris. 16: 419. 1828 ≡ Beauharnoisia fructipersistent . Aril orange. Fig. 35. pendula Ruiz & Pav. in Ann. Mus. Natl. Hist. Nat. 11: 72, pl. 9. 1808. – Lectotype (designated by Marinho & Distribution — Brazil (Amazonas, Rondônia), Colom- al. 2016b: 769) or perhaps holotype: Peru, “Chicoplaya, bia (Cauca, Caquetá, Vaupés), Venezuela (Amazonas). habitat in Peruviae Andium”, s.dat., F. P. c. l. 405 (K Fig. 36. [K000488587]!).

=

= Tovomita brasiliensis ( Mart.) Walp. View in CoL in Repert. Bot. Syst. 1: 392. 1842 ≡ Marialva brasiliensis Mart. View in CoL in Nov. Gen. Sp. Pl. 2: 83, t. 167. 1826. – Lectotype (designated by Marinho & al. 2016b: 769): Brazil, “ Bahia, habitat in sylvis ad Itahype fluv.”, s.dat., C. F. P. Martius s.n. (M [ M0086209 ]!; isolectotypes: M [ M0086206 ]!, M [ M0086207 ]!, M [ M0086208 ]!, M [ M0086210 ]!).

Tovomita pauciflora Cuatrec. View in CoL in Anales Inst. Biol. Univ. Nac. México 20: 104. 1949, syn. nov. – Holotype: Colombia, Norte de Santander, region del Sarare, hoya del Río Cubugón, vertientes Del Caraño , 500–700 m, 18 Nov 1941, J. Cuatrecasas 13302 (F [ F0054549 F] photo!; isotypes: COL [ COL000002850 About COL ] photo!, US [ US 00114299] photo!).

Description — Shrubs or trees up to 10 m tall, prop roots inconspicuous; exudate yellow, abundant. Petioles 0.6– 0.8 cm long, green or reddish, smooth, lenticels absent. Leaf blades 3.2–7(–12) × 0.9–3.5 cm, greenish in sicco, young leaves sometimes purplish-red adaxially in vivo and in sicco, black dots absent, chartaceous, narrowly elliptic, base convex to decurrent, apex acuminate; papillae, lenticels and fungal spots absent; exudate canals inconspicuous. Venation: secondary veins 13–17 pairs, 2–4 mm apart from each other, forming angle 60°–70° to midvein, immersed adaxially, slightly prominent abaxially, 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 percurrent sinuous; intramarginal vein absent. Inflorescences: ♂ a lax 3-basal branched cyme, up to 9 flowers, lacking terminal flower, or ♀ a dichasium, lenticels absent. Pedicels 10–18 mm long, green or reddish, distal and proximal portion with same gauge, proximally articulated on lateral flowers of dichasia, lenticels absent. Floral buds 5–8 mm long, oblong, apex rounded, lenticels absent. Sepals 2, 6–8 × 2–4 mm, oblong, apex rounded, green; petals 4, 6–11 × 2.5–5 mm, oblong to oblanceolate, reflexed, apex rounded, white to light yellow. Staminate flowers with 45–50 stamens, 4–7 mm long, isodynamous; filaments terete, white; anthers 0.5–0.7 mm long, connective not exceeding thecae; pistillode inconspicuous, white. Pistillate flowers with 45–50 staminodes, 3.8–4 mm long, white; ovary 6–7 mm long, not costate or lobed, 4-locular, white, styles free 3–3.2 mm long, stigmas 4, 1–1.3 mm in diam. Capsules fleshy, 2–2.4 × 1.2–2 cm, 4-septate, spheroid when closed, not costate or lobed, rostrum absent, free styles, 3.5–4.5 mm long, epicarp asperous, lenticels present, green when immature, brown when mature, mesocarp orangish to red; sepals, petals, staminodes and stigmas persistent;. Seeds aril orange. Fig. 37.

Distribution — Bolivia (Beni, La Paz, Santa Cruz), Brazil (Acre, Amapá, Amazonas, Bahia, Espírito Santo, Maranhão, Minas Gerais, Pará, Paraíba, Pernambuco, Rio de Janeiro, Rio Grande do Norte, Roraima, Tocantins), Colombia (Amazonas, Meta, Norte de Santander, Vaupés), French Guiana (Coralie, Kourou), Guyana (Barbice, Potaro-Siparuni), Peru (Cuzco, Huanuco, Loreto, Madre de Díos, San Martín), Venezuela (Carabobo, Táchira). Fig. 38.

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

Recognition and discussion — In the Atlantic forest, Tovomita fructipendula is commonly a shrub, whereas in the Amazon forest it is a tree up to 10 m tall. In herbaria, it is commonly confused with T. longirostrata and T. stergiosii , due to the usually small, narrowly elliptic leaves, but it can be easily differentiated from these two species by the floral buds with a rounded apex ( Fig. 37C, vs rostrate in T. longirostrata and mucronulate in T. stergiosii ) and mainly by the rugose epicarp ( Fig. 19C, vs asperous in T. longirostrata [ Fig. 19E] and smooth in T. stergiosii ).

Alencar & al. (2023) showed that some specimens of Tovomita fructipendula from Espírito Santo state, Brazil, had fruits with a smooth epicarp, contrasting with the typical asperous fruits of T. fructipendula . To confirm theses specimens as T. fructipendula , it is necessary to collect materials with floral buds or flowers. The specimens from Bolivia and the extreme northeast of Brazil have unusually large leaves up to 12 cm long. Table 1 compares T. fructipendula with T. longirostrata , T. stergiosii , and other species with small, narrowly elliptic leaves.

Selected specimens examined — BOLIVIA: BENI: Prov. Gral, Ballivián , 15°18'S, 67°05'W, 1040 m, 1 Oct 2000, fr., Beck St G GoogleMaps . 27631 (NY!); LA PAZ: Province of Larecaja , 6 km N Consata, 15°20'S, 68°31'W, 1300 m, 15 Dec 1981, bud, J. C GoogleMaps . Solomon & al. 6604 ( NY!) ; SANTA CRUZ: Prov. Ichilo, Parque Nacional Amboró , 17°39'S, 63°43'W, 400 m, 18 Jan 1988, fr., M GoogleMaps . Nee & M . Saldias P . 35934 (NY!). — BRAZIL: ACRE: Mâncio Lima , estrada para o lugar Barão, 24 Oct 1984, bud, C . A. Cid Ferreira & al. 5244 (INPA!); AMAPÁ: Rio Falsino, approx. 10 km upstream of confluence with Rio Araguari , 00°50'S, 51°45'W, 1 Oct 1983, fl. ♀, B. V GoogleMaps . Rabelo & al. 2395 ( MG!); AMAZONAS: Humaitá, near Tres Casas, 14 Sep–11 Oct 1934, fl. ♀, B. A . Krukoff 6316 ( B!, NY!) ; BAHIA: Uruçuca , 14°31'03"S, 39°02'37"W, 24 Feb 2019, fl. ♂, L. C GoogleMaps . Marinho & K. M . Pimenta 1529 ( HUEFS!) ; ESPÍRITO SANTO: Vitória , Jardim Caburi, 17 Oct 2000, fl. ♀, A. M . Assis 857 ( VIES!) ; MARANHÃO: Morros , ramal à margem da MA-110, 16 Jul 2022, fl. ♀, L. C . Marinho & al. 1805 ( MAR!) ; MINAS GERAIS: Caratinga, Estação Biológica de Caratinga, Mata do Jaó , 13 Nov 1985, bud, M. A . Lopes & P. M . Andrade 842 ( BHCB!); PARÁ : Belém, Mata da Reserva Mocambo , 31 Jan 2017, sterile, L. C . Marinho & al. 1241 ( CEPEC!) ; PARAÍBA: 28 Jan 1971, fr., O . Lima 2402 ( UFP!) ; PERNAMBUCO: Igarassú, Usina São José , 07°41'50"S, 34°57'37"W, 84 m, 12 Mar 2009, fl. ♀, J. A. N GoogleMaps . Souza & al. 391 ( UFP!) ; RIO DE JANEIRO: Angra dos Reis, Praia do Manduri , 23°02'S, 44°38'W, 17 Nov 1993, bud, M. T. Z GoogleMaps . Toniato & al. 30162 ( UEC!) ; RIO GRANDE DO NORTE: Baía Formosa, RPPN Mata de Bela , 06°25'28"S, 35°06'46"W, 34 m, 4 May 2012, fr., A. A GoogleMaps . Roque & al. 1367 ( UFRN!) ; RORAIMA: Serra do Surucucu , 28 Oct 1991, fl., S . Almeida & M . Cordeiro 750 ( MG!, NY!) . TOCANTINS: Wanderlândia , 06°51'12"S, 48°01'10"W, 207 m, 6 Aug 2010, fr., G. A GoogleMaps . Thomé & al. 21 ( HUTO) . — COLOMBIA: AMAZONAS: Río Caquetá, margen derecho, frente a Villa Azul, 25 Nov 1989, sterile, T. van Andel & al. 523 ( COAH!) . META: Sierra de La Macarena , central mountains north ridge, 1500 m, 21 Dec 1949, bud, W. R . Philipson & J. M . Idrobo 1836 ( NY!) ; VAUPÉS: Río Vaupés, between Mitú and Javareté , 14–24 May 1953, fl. ♂, R. E . Schultes & I . Cabrera 19273 ( GH!) . — FRENCH GUIANA: KOUROU: Fluve Kourou , à environ 7.7 km em amont de la crique Couy, 21 Sep 1967, bud, anonymous B-1356 ( K!, L photo!, NY!, P!) . — GUYANA: BARBICE: Brandtwegt, R. B . Berbice river , 31 Jan 1951, fr., F3022 ( NY!) . POTARO-SIPARUNI: Amatuk Portage , 19 May 1944, bud, B . Maguire & D. B . Fanshawe 23550 ( NY!) . — PERU: CUZCO: Dist. Camisea, Campamento Armihuari , 11°51'48"S, 72°46'44"W, 469 m, 27 Jan 1997, fr., P GoogleMaps . Acevedo-Rodriguez & al. 9201 ( K!, NY!) ; HUANUCO: southwestern slope of Río Llullapichis watershed , 09°25'S, 74°44'W, c. 1535 m, 28 Jul 1969, fr., F GoogleMaps . Wolfe 12405 ( W!) ; LORETO: Maynas, Iquitos, cerca a La boca del Río Nanay , 1 Nov 1976, bud, J . Revilla 1718 ( NY!) ; MADRE DE DÍOS: Parque Nacional del Manu, Río Manu, Cocha Cashu station , 28 Oct 1980, bud, R. B . Foster 5656 ( NY!) ; SAN MARTÍN: Prov. Lamas, Alonso de Alvarado , Cerro Blanco, 1000 m, 8 May 1973, fr., J . Schunke V . 6213 (NY!). — VENEZUELA: CARABOBO: Límite Distrito Bejuma / Distrito Montalbán, Cordillera de La Costa , 10°15'N, 68°17'W, 1000–1200 m, 1 Jan 2007, fr., W GoogleMaps . Meier & N . Flauger 14198 ( M!) ; TÁCHIRA: 10 km E of La Fundacíon around Represa Dorada, 600– 1000 m, 10–13 Mar 1981, bud, R . Liesner & A . González 10171 ( NY!) .