Coccoloba mollis Casar.

Ancona, Juan José, Ortiz-Díaz, Juan Javier, Gutiérrez-Alonso, Eduardo & Ledesma, Patricia Hernández, 2025, A taxonomic revision of Coccoloba sect. Paniculatae (Polygonaceae, Eriogonoideae), Phytotaxa 704 (2), pp. 106-152 : 134-136

11. Coccoloba mollis Casar. , Nov. Stirp. Bras. 72. 1844. Coccoloba polystachya var. mollis Meisner, DC. Prodr. 14: 151. 1856 ≡ Coccoloba polystachya var. pubescens Lindau, Bot. Jahrb. 13: 133. 1891. ( Figs. 14A–D ).

Lectotype (designated by Howard 1960c: 253 as “ holotype ”, here corrected according to the Art. 9.10 if ICN):— BRAZIL. Reperi in insula Itaparica ad ostium sinus Bahiensis, in maritimis arenosis, February 1844, Casaretto 2218 (TO! [photo]).

= Coccoloba polystachya Wedd., Ann. Sci. Nat., Bot. , sér. 3,13: 261. 1849.

Lectotype (here designated):— BRAZIL, Matto Grasso , July–August 1845, Weddell 1848 (P!, barcode P00734657 [photo], isolectotypes P00734658, P00734659 [Photos]) https://mediaphoto.mnhn.fr/media/1443547639018WT2Wufr05OJE78nK).

= Coccoloba paniculata Meisn., Fl. Bras. 5(1): 43. pls. 20, 21. 1855. Coccoloba polystachya var. glabra Lindau, Bot. Jahrb. Syst. 13: 133. 1891.

Lectotype (here designated):— BRAZIL, Goyazana , ad fluv. Richáo, Pohl s.n. (BR!, barcode: BR0000005782260 [photo], https://plants. jstor.org/stable/viewer/10.5555/al.ap.specimen.br0000005782260).

= Coccoloba dugandiana Á. Fernández. Mutisia 5: 1. 1952.

Type:— COLOMBIA. Boyacá: Caño Guira near mouth of Rio Meta, Haught 2624 (holotype COL! [photo], isotypes S!, US! [photos]).

Description:—Trees hermaphrodites, functionally unisexual, 5‒20 m tall; branches striate, sparsely to abundantly pubescent, brown to greyish, lenticelated; ochrea 1.5–4 (–5.5) cm long, densely, pubescent, puberulent, strigose to woolly, with yellow-brown trichomes, breaking irregularly, glandular-punctate. Leaves simple, alternate; petiole 1.3–2 cm long, 3–7 mm diameter, striated, strigose or tomentose, with yellow-brown trichomes, inserted at base of the ochrea. Leaf blade obovate to obovate-elliptical 13–17 (–19) cm long, 7–12(–14) cm wide, chartaceous, adaxially glabrous or sparsely puberulent, mainly on nerves, abaxially pubescent to densely or sparsely hirsute, pilose, puberulous, strigose or tomentose, with yellow‒brown trichomes, margin entire, apex acuminate, rarely obtuse or rounded, base slightly cordate, venation brochidodromous, reticulate, primary veins (9–)10–14 pair, prominent, glabrous. Inflorescence spiciform, arranged in a monopodial panicle, main axis elongated, 8–13 cm long, 2.5 mm diameter, ribbed, densely hirtellous, puberulent; spiciform portion 4–15 (–19) cm long, rachis ribbed, striate, 1.3 mm diameter, densely hirtellous, puberulent; bracteole 0.5 mm long, cymbiform, densely hirsutulous, pulverulente; ochreolae 0.8–1.2 mm long, campanulate, bilobulate, chartaceous, hirtellous-puberulent; pedicels in flower 0.5–0.8 mm long, pedicels in fruit 0.8 mm long, thicker, hirsutulous to strigose, pulverulent. Flowers in two whorls, outer tepals 3, inner tepals 2, alternate; flowers functionally male 2–3 × fascicle; hypanthium 0.7–1 mm long, hirtellous, campanulate, abruptly narrowed at the articulate with the pedicel; outer tepals strigose, coriaceous, inner tepals glabrous, papyraceous; tepals 1.2–1.5 mm long; stamens 7, 1.5–2 mm long, filaments filiform 1 mm long, anthers 0.5 mm long; ovary 0.8 mm long, trigonous, glabrous, style 3, 0.5 mm long; flowers functionally female not seen. Fruit acrosarcum, 11–13 (–15) × 9–12 mm, globose, subglobose, ovoid, sparsely pilose, apex acute to obtuse, base truncate or flat, abruptly contracted at junction with pedicel; hypanthium accrescent, succulent, almost completely enclosing achene, tepals accrescent, sparsely or densely pilose, succulent, appressed at the apex tuberculate of the achene.Achene 9–10 × 8–9 mm, globose-subglobose, base truncate or flat, apex tuberculate, conical, prominent, brown, smooth, glossy.

Etymology:—The specific epithet “mollis ” refers to plants with softly-hairy, or velvety hairs.

Phenology:—Flowers from May to December; and fruits from January to May and from August to November.

Distribution and ecology:— Coccoloba mollis has a wide geographic distribution in Colombia, Venezuela, Suriname, French Guiana, Ecuador, Bolivia and Brazil ( Fig. 14E ). In the Amazon rainforest, very humid and rainy forests, it is found on clayey, sandy or rocky soils, at elevations ranging from 0– 800 m.

Provisional conservation status:—The EOO is 109,477,357 km 2, within the threshold for Least Concern (LC) status under subcriterion B1, and the AOO is estimated to be 2442 km 2, which is below the threshold for Near Threatened (NT) status under subcriterion B2. Coccoloba mollis es la especie con mayor rango de distribución en Sudamérica, aun cuando la gran mayoría de los registros no están ubicado dentro de áreas naturales protegidas, muchas de ellas se encuentran dentro de la espesa selva amazónica. According to the IUCN methodology ( IUCN Standards and Petitions Committee 2024), if the EOO threshold is lower than the AOO threshold, the EOO should be set equal to the AOO to ensure consistency with the definition of AOO as an area within the EOO. Therefore, C. mollis is assigned a preliminary status of Endangered [NT B2ab(iii)].

Typification of Coccoloba mollis :— Coccoloba mollis is a species with a wide distribution in South America. It is also one of the species in the genus with a great morphological variability in the shape and size of the fruits, that are globose, subglobose or ovoid, 8–13 mm long, as well as in the indumentum of the leaves, ochrea, inflorescences and fruits, that can be densely to sparsely pubescent, pilose, strigose, or puberulous. Howard (1960c) discussed all of this variation and recognized several synonyms for C. mollis , with which we also agree.

In the original publication, Casaretto (1844: 72) did not cite the holotype. Casaretto’s collection, mainly preserved at the herbarium TO (HUH-Index of Botanists 2025), we traced a specimen collected on the island of Itaparica, near Bahia, Brazil, matching Casaretto’s protologue ; the label also includes the collection number (2218) and the catalogue number (80). This specimen was selected by Howard (1960c) as holotype; however, according to the Art. 9.10 of ICN, this indication should be corrected to lectotype.

Coccoloba dugandiana A.Fernández was described from a type specimen which is sterile. Howard (1960a) and Aymard & Howard (2004) acknowledged the existence of C. dugandiana , but disagreed with Fernández’s description, because this was based on what must be considered anomalous material, by have abnormally emarginated leaves of the type being uncharacteristic of the species. However, in none of Howard’s works there is an adequate description and diagnostic characters presented to delimit both species. Melo (2004) also recognized all the morphological variation in C. mollis and considered C. dugandiana as a new synonym of C. mollis . She said that the diagnostic characters presented by Fernández-Pérez for C. dugandiana presented discrepancies with the characters observed in C. mollis . We do not agree with her conclusion.

A Another species morphologically very similar to Coccoloba mollis is C. efigeniana . The characteristics of the fruit in both species are important for their delimitation, due to vegetatively they are very similar. The presence of triangular or obpyramidal fruits distinguishes C. efigeniana from C. mollis , which has globose, subglobose or ovoid fruits. In addition, C. efigeniana is distributed from Costa Rica to French Guyana in areas closer to the coasts of the Atlantic Ocean, while C. mollis is a species with greater affinity to the Amazon rainforest.

Typification of Coccoloba polystachya :—In the original description of Coccoloba polystachya, Weddell (1849: 261) observed a living plant that was assigned a catalogue number 3381 and also cited the collection Martius 1242. In Herbarium P, we have located three sheets of the living plant observed by Weddell (cat. No. 3381) with the herbarium collection number Weddell 1848, and we have selected the specimen with barcode P00734657 as the lectotype for the name C. polystachya .

Typification of Coccoloba paniculata :— Coccoloba paniculata was described by Meisner citing the specimens Pohl s/n, Poeppig 2649 and Martius 1242. Here we selected the specimen Pohl s/n from the BR herbarium as the lectotype of the name C. paniculata .

Specimens examined:— BOLIVIA. Beni: Gral. Ballivian, Rurrenabaque, cerro del Calvario, 14°26’52” S, 67°31’36” W, 411 m, 21 October 2003, Fuentes 5537 (MO); Gral. Ballivian, carretera Carinavi-San Borja, lado oeste de la Serranía del Pilón Lajas, 5 km de la cumbre, 15°19’ S, 67°03’ W, 765 m, 2 November 1989, Smith & García 13854 (MO); Cumbre de la Serranía del Pilón Lajas, carretera Carinavi-San Borja, 25 km de Yucumo, 15°17’ S, 67°04’ W, 850–900 m, February 1990, Smith et al. 14418 (MO). Gral. Ballivian, 14°26’S, 67°31’W, 411 m, 21 October 2003, Fuentes 5537 (LPB); Gral. Ballivian, 15°19’S, 67°03’W, 765 m, 2 November 1989, Smith & García 13854 (MO). Gral. Ballivian, 15°17’S, 67°04’W, 850 to 900 m, February–July 1990, D.N. Smith et al. 14418 (MO); Gral. Ballivian, 14°26’S, 67°31’W, 411 m, 21 October 2003, A.F. Fuentes 5537 (MO, NY); Itenez, 14°25’S, 62°03’W, 200 m, 19 July 1993, Quevedo et al. 961 (MO). La Paz: Nor Yungas, 4.5 km below Yolosa, then 10 km W, on road up the Río Huarinilla, 16°12’ S, 067°50’ W, 1450 m, 19–20 October 1982, Solomon 8525 (MO); Abel Iturralde, Alto Madidi, ridge top, 7 km NE of camp. Transect 1, 13°35’ S, 68°46’ W, 380 m, 25 May 1990, Gentry & Estenssoro 70552 (MO); Franz Tamayo, Región Madidi, Santo Domingo, arroyo Tintaya, 14°47’09” S, 068°35’00” W, 1437 m, 12–19 September 2012, Cayola et al. 4153B (MO), 4324B (MO), 4403B (MO), 4404B (MO), 4505 (MO); Franz Tamayo, Parque Nacional Madidi, Sumpulo. Parcela Permanente 34, 14°34’19” S, 68°46’46” W, 1223 m, 13 July 2008, Moya et al. 255 (LPB, MO); Franz Tamayo, Región Madidi, Santo Domingo, sector arroyo Tintaya, 14°47’23” S, 68°35’01” W, 1468 m, 25 October 2006, Chapi et al. 188 (LPB, MO); Franz Tamayo, Culi, en Parque Nacional Madidi, cercanías del río Pelechuco, bajando desde Pelechuco por el camino antiguo Pelechuco-Apolo, 14°43’44” S, 68°50’41” W, 1496 m, 30 September–03 Octtober 2009, Arellano et al. 845 (LPB, MA, MO); Franz Tamayo, Parque Nacional Madidi, Mojos, campamento Fuertecillo, sobre la senda a Queara, 14°35’31” S, 68°55’14” W, 1750 m, 30 April 2007, Cayola et al. 2742 (MO); Franz Tamayo, Región Madidi, Santo Domingo, Lechemayu, a 984 m N campamento Lechemayu, primera palca, 14°46’09” S, 68°37’11” W, 1522 m, 16 August 2010, Cornejo et al. 1350 (MO); Franz Tamayo, Santo Domingo, sector Tintaya, parcela temporal 2, 14°47’10” S, 68°34’58” W, 1400 m, 12 October 2006, Cornejo et al. 200 (MO); Larecaja, desde la comunidad de Quilapituni, subiendo por el camino antiguo Ingenio-Mapiri, hasta un lugar llamado Victopampa. Campamento a unas 3 horas a pie bajando hacia el río Chiñijo, 15°28’03” S, 68°21’17” W, 1489 m, 17–19 April 2010, Arellano et al. 1614 (LPB, MA, MO); Bautista Saavedra, Área Natural de Manejo Integrado Apolobamba, Wayrapata, 0,7 Km NE del caserío, 15°05’11” S, 68°29’19” W, 1334 m, 18 May 2004, Antezana et al. 359 (MO). Santa Cruz: Velazco, San José de Campamento, 20 km al oeste de la hacienda San Roque, 15°09’20” S, 60°59’29” W, 230 m, 29 August 1996, Guillén & Marmañas 4609 (MO); Provincia de Ichilo. Río Ibabo, Reserva Forestal Choré; Bosque Experimental “Elias Meneces”, 16°35’ S, 64°31’ W, 180, 5–10 August 1990, Neill et al. 9216 (MO); Velazco, alrededores de la Hacienda San Roque, 15°00’37” S, 61°07’42” W, 280 m. 06 August 1996, Guillén et al. 180 (MO). BRAZIL. Acre: Mun. de Rio Branco, Mata de terra firme, solo argiloso, 19 October 1980, Cid Ferreira & Nelson 2971 (MO); Mun. Porto Acre. Flat terrain not flooded or only shallowly and briefly, 9°45’S, 67°38’W, 3 November 1993, Daly et al. 8027 (MO); Tarauacá: Bacia do Rio Juruá, Rio Muru, Seringal Vitória Velha, Colocacão Vista Alegre, margem direita, área de transicão entre “Restinga” (floresta de terra firme em terraco alto) e “Tabocal”, 8°25’S, 70°48’W, 28 September 1994, Silveira et al. 946 (MO); Basin of Rio Purus, near mouth of Rio Macauhan (tributary of Rio Yaco), 9°20’S, 69°00’W, 17 August 1933, Krukoff 5550 (MO); Mun. Sena Madureira. Bacia do Rio Purus, Fazenda Nova Olinda, Carreador do Poti, ca. 8 km da sede, 10°07’S, 69°13’W, 21 October 1993, Silveira et al. 560 (MO). Amazonas: Basin of Rio Madeira, municipality Humayta, near Tres Casas, 14 September–11 October 1934, Krukoff 6114 (MO); Basin of Rio Solimoes, municipality of São Paulo de Olivenca, near Palmares, 11 September–11 October 1936, Krukoff 8314 (MO); Basin of Rio Solimoes, Municipality São Paulo de Olivenca, near Palmares, Terra firme; high land, 11 September.– 26 October 1936, Krukoff 8337 (MO); Basin of Rio Solimoes, Municipality São Paulo de Olivenca; basin of creek Belem, 26 October–11 December 1936, Krukoff 8841 (MO). Goiás: Mun. Luziania. Lagoa de Prata , 15 August 1980, Hatschbach 43146 (IBE, MO); States of Goiás and Mato Grosso. Goiaz: Jataí-Caiaponia road, 40 km. from Caiaponia, 600–1000 m, 27 June 1966, Hunt 6244 (MO). Maranhão: Campo on Fazenda do Senhor Miquel Pacheco. Km 176 of Br 135. 2 km N of São Mateus, 4°00’S, 44°30’W, 27 September 1980, Daly et al. D285 (MO); Maranhão, Maracassumé River Region, Campo de Boa Esperanca, 30 August 1932, de Lemos & Krukoff 1811 (MO). Mato Grosso: Vicinity of Xavantina. Creek margin, gallery forest, ca. 70 km N of Xavantina, 14°40’S, 52°20’W, 300–400 m, 10 October 1964, Irwin & Soderstrom 6719 (MO); Mato Grosso, Rib. Claro (Mun. Alto Araguaia), 22 September 1974, Hatschbach & Kummrow 35071 (MO). Pará: Serra dos Carajás. Rio Parauapebas; control point at entrance to Serra Norte, ca. 39 km east of AMZA camp N-5. Forest on low ground along river, high presence of babassu palm, 6°04’S, 49°55’W, 150 m, 23 June 1982, Sperling et al. 6324 (MO); Mun. Óbidos. 91 km de Oriximiná nos Campos de Ariramba , entre rio Jaramacaru e Igarapé Mutum, 1°10’S, 55°35’W, 4 December 1987, Cid Ferreira 9774 (MO); On varzia land near Cassipa, Tapajos River region, September 1931, Krukoff 1246 (MO). Sergipe: Santa Luzia do Itanhy, RPPN Mata do Crasto, Borda de mata, solo arenoso, 14 August 2012, Gomes et al. 616 (MO); Santa Luzia do Itanhy, RPPN Mata do Crasto, Trilha no interior da mata, solo arenoso. Mata Atlântica, 11°23’S, 37°25’W, 09 August 2011, Prata et al. 2613 (MO). Minas Gerais: Triángulo región, Cerrado, McGill University Expedition, 1967–1968, Goodland 822 (MO); Ituiutaba, 28 August 1948, Macedo 1186 (MO). São Paulo: Munic. José Bonifácio, estrada de terra Municipal José Bonifácio-Nova Alianca, Fazenda Jacaré (entre Córregos Jacaré-Fartura), 21°03’S, 49°41’W, 5 September 1992, Silva 294 (MO); Mpio. Penapolis, 18 September 1974, Hatschbach & Kummrow 34820 (MO). ECUADOR. Orellana: Estación Científica Yasuni, Río Tiputini, al noroeste de la confluencia con el R. Tivacuno, este de la carretera Repsol-YPF, km 7 desvío hacia el pozo Tivacuno, 30 November 2002, Villa & Alvia 1621 ( US); Manabí: Pedernales Cantón, Reserva Ecológica Mache-Chindul, comunidad Ambacha (vía marginal de la costa-Chindul), 00°15’00” N, 79°48’00” W, 270 m, 25 March 1997, Clark et al. 4206 (MO, US). VENEZUELA. Amazonas: Rio Orinoco between San Fernando de Atabapo and San Antonio, Silverio 41 (A); Sanariapo, Williams 15961 (A, F), 16042 (F). Bolívar: Alto Rio Paragua, Cardona 1183 (NY, US).

Figures

FIGURE 14. A. Specimen of Coccoloba mollis (W. Palacios et al. 501, MEXU); B. Portion of inflorescence and pedicels; C. Acrosarcum lateral view; D. Achene lateral view; E. Distribution model.