Authority:

Michelangeli, Fabián A. 2005. (Melastomataceae). Fl. Neotrop. Monogr. 98: 1-114. (Published by NYBG Press)

Family:

Melastomataceae

Synonyms:

Melastoma physophora Vahl, Melastoma tococo Desr., Tococa aubletii D.Don, Tococa formicaria Mart. ex DC., Tococa vesiculosa DC., Tococa barbata Benth., Tococa roreimi Hook., Tococa acuminata Benth., Tococa guianensis var. macrophysca Naudin, Tococa didymophysca Naudin, Tococa formicaria var. didymophysca (Naudin) Cogn., Tococa latifolia Naudin, Tococa occidentalis Naudin, Tococa cardiophylla Naudin, Tococa orcheophora Naudin, Tococa coriacea S.Moore, Tococa platyphysca Cogn., Tococa subglabrata Cogn., Tococa formicaria var. gardneri Cogn., Tococa discolor Pilg., Tococa juruensis Pilg., Tococa peckiana B.L.Rob. ex O.E.Schulz, Tococa loretensis Ule, Maieta glandulifera Standl., Tococa cachimbensis Brade, Tococa guianensis var. orinocensis Wurdack, Tococa mexiae Wurdack, Tococa chuivensis Wurdack, Tococa chuivensis var. ecuadorensis Wurdack

Description:

Species Description - Shrub or small tree, to 1-5 (-7) m tall; stems glabrous, sparsely glandular-pubescent or with a mix of glandular and nonglandular setae, often with a reddish or fuchsia tinge on the younger branches, the setae 2.5 mm long, occasionally up to 12 mm long, persistent, the nodes glabrous. Leaves anisophyllous, seldom isophyllous, ovate to oblong elliptic, the larger leaves (5.5-)8-30(-34) X (4.0-)5-15(-18) cm, the smaller leaves 3.5-10 X 2.5-7 cm, apex acute to acuminate, caudate or aristate, base rounded to acute, occasionally attenuate, adaxial surface adpressed nonglandular-setose, occasionally glandular-setose, rarely glabrous, abaxial surface glandular-setose, occasionally nonglandular-setose, rarely glabrous, the setae denser on the primary and secondary veins, usually light green, purple or dark red in some young individuals, 5(-7)-nervate, the veins reticulate underneath, membranaceous to chartaceous (slightly coriaceous in individuals growing in the sun), entire to ciliate-serrulate, occasionally ciliate-crenulate; petioles 0.5-2.0(-4.5) cm long, sparsely or densely glandular-setose; domatia free from the leaf blade, globose, ovoid or narrowly ellipsoid, (1.5-)2.0-3.0 (-4.5) X (0.7-)1.0-2.0(-3.0) cm, sometimes present only in the larger leaf, occasionally with a purple or light red coloration. Inflorescence a terminal panicle of 8-30 (-50) flowers, (3.5-)5-15(-21) cm long, the axis compressed, dark red to magenta, occasionally green or turning green after it matures. Flowers on pedicels 3-9 mm long, sometimes articulate 1-2 mm below the hypanthium, flared at the apex, subtended by caducous or persistent bracts, subulate to lanceolate, 0.5-1.5(-3.0) X 0.2-0.7 mm; hypanthium broadly conical to globose, (3.5-)5.0-7.5(-9.5) mm long, glabrous to densely glandular-setose, or with a mix of lepidote hairs and long glandular setae up to 3.5 mm long; outer calyx teeth reduced to well developed and deltoid, 0.3-2.0 mm long, projecting 0-1.0 mm above the inner calyx teeth, terminated in 1 or more glandular setae up to 3.5 mm long, occasionally glabrous, inner calyx teeth broadly deltoid to ovate, fused halfway to the apex (0.5-1.2 mm), 1.0-3.0 mm long; the ring inside the torus glabrous; petals 5, broadly oblong to ovate, (5.5-)6.5-15 X (3.0-)4.5-7.5(-9.0) mm, base attenuate, apex emarginate to obtuse, border entire, often undulate, pruinose to smooth, sometimes finely granulóse, glabrous, occasionally with a caducous, terminal, glandular seta, pink; stamens all the same size; filaments (5.0-)5.5-7.5(-8.5) mm long, glabrous; anthers yellow; the connective with a dorsal-basal blunt tooth; thecae (4.5-)5.0-7.5(-8.5) mm long, opening by a ventrally inclined pore; ovary 3-locular, 1/2 to totally inferior, the superior portion globose to broadly conical, terminated in a ring elevated 0.1-0.5 mm over the apex, the ring glabrous to densely pilose or with a corona of long glandular setae or small sessile glands; style glabrous, very rarely minutely stellate-pubescent at the base, 13-20(-23) mm long; stigma funnelform, l.5 mm wide, stigmatic surface finely papillate. Fruits globose, 10-19(-22) mm long, black, glandular-setulose, rarely glabrous; seeds not evident through the pericarp, triangular, 0.6-1.2 mm long, without capitate trichomes in the raphal area, without sculpturing, anticlinal walls straight, periclinal walls convex, boundaries between periclinal walls flat. Chromosome number n = 17.

Discussion:

Tococa guianensis is a highly variable species that is widely distributed throughout the entire geographical range of the genus. It is very common in both primary and disturbed habitats in the Amazon and Orinoco basins and less frequent in northern South America and Central America.

Due to the variation exhibited across the geographical range, several species have been described, most of them based on stem or ovary pubescence or on the morphology of the calyx. In many cases, the type specimens are clearly distinct from each other, but analysis of all the populations across the entire geographical range shows that most of the variation falls within a continuum, which makes it impossible to recognize all of these forms as separate species. Several authors (Cogniaux, 1888; Gleason, 1931b; Macbride, 1941; Wurdack, 1973b, 1980; Wurdack & Renner, 1993; Vasquez, 1997; Michelangeli, 2001b) have addressed this problem but seldom arrived at similar conclusions as to which species should be recognized. The most detailed of these analyses was performed by Gleason (1931b), who took into account the shape and pubescence of the apex of the ovary and of the outer teeth calyx. He outlines 12 possible combinations of calyx and ovary characters, although he did not observe all of them. However, in reviewing live populations and herbarium collections for this monograph, I found all of them to occur in natural populations. Many of these forms correspond to described names.

In general, individuals growing in the sun tend to be less pubescent than individuals growing in the shade. Likewise, individuals in wetter areas or soils tend to be more pubescent than those in drier areas. This variation can occur even within individuals. During the course of this study, two branches of the same individual, one growing in the shade and one in the sun, were keyed out to different “species” (T. guianensis and T. occidentalis) due to the variation in pubescence of the hypanthium and ovary.

The type of T. acuminata is labeled as coming from Ecuador, but Bentham’s description cites it as coming from “Colombia and Isle of Gorgana” (sic). There is a Barclay specimen at K (Hooker Herbarium) annotated as from “Island of Gorgana,” presumably by Bentham.

Of all the forms here considered synonyms of T. guianensis, T. formicaria seems to be the most distinct, due to its dense and long pubescence on vegetative parts as well as the hypanthium, and perhaps could be considered a separate taxon. However, the distribution of specimens with the typical pubescence of T. fornicaria falls within the range of distribution of specimens with characters that match the type of T. guianensis. Moreover, in this area (central Brazil), numerous specimens examined exhibited a continuum of density and size of pubescence between those typical of T. guianensis and T. formicaria.

The type of T. orcheophora has all the vegetative and floral characteristics of T. guianensis, with the exception that the apex of the ovary lacks the typical corona of setae present in typical populations of T. guianensis. The lack of the ovary corona is extremely rare, occurring in less than ten out of nearly 1000 specimens of examined during the course of this monograph. Toe oca orcheophora is here is treated as a synonym of T. guianensis, until more specimens from the type locality can be examined.

Tococa guianensis is closely related to T. bullifera, which differs in the single-branched paniculate inflorescence, and to T. aristata, which differs in the calyx morphology and color of the petals.

Tococa guianensis often seems to be associated with Azteca ants and more rarely with Crematogaster. Even though T. guianensis is found at elevations over 1600 m, specimens collected over 1000 m often lack any inhabiting ants. A notable exception seems to be open habitats of the Gran Sabana in SE Bolívar State in Venezuela, in which ants can be found in plants growing above 1400 m. In southern Ecuador and northern Peru the domatia can be inhabited by ants of the genus Myrmelachista. The workers of this genus create large clearings in the forest understory, which become monospecific stands of T. guianensis (Morawetz et al., 1992; Renner & Ricklefs, 1998).

Pemón, Piaroa, and Bare Amerindians in southern Venezuela occasionally consume the fruits of T. guianensis. A tea made of the leaves, or the domada alone, is given to pregnant women by Yekuana or Bare Amerindians to induce the birth of a boy. Piaroa Amerindians in northern Amazonas State use a decoction of the leaves as an unspecified remedy.

Distribution:

Amazonas Brazil South America| Goiás Brazil South America| Acre Brazil South America| Valle del Cauca Colombia South America| Distrito Capital Colombia South America| Cauca Colombia South America| Chiapas Mexico North America| Tabasco Mexico North America| Veracruz Mexico North America| Belize Central America| Honduras Central America| Zelaya Nicaragua Central America| Puntarenas Costa Rica Central America| San José Costa Rica Central America| Darién Panamá Central America| Panamá Panama Central America| Amazonas Colombia South America| Antioquia Colombia South America| Chocó Colombia South America| Cundinamarca Colombia South America| Meta Colombia South America| Nariño Colombia South America| Putumayo Colombia South America| Valle Colombia South America| Vaupés Colombia South America| Vichada Colombia South America| Colombia South America| Amazonas Venezuela South America| Anzoátegui Venezuela South America| Apure Venezuela South America| Bolívar Venezuela South America| Mérida Venezuela South America| Táchira Venezuela South America| Zulia Venezuela South America| Essequibo Guyana South America| Cuyuni-Mazaruni Guyana South America| Potaro-Siparuni Guyana South America| Suriname South America| French Guiana South America| Morona-Santiago Ecuador South America| Napo Ecuador South America| Pastaza Ecuador South America| Sucumbíos Ecuador South America| Amazonas Peru South America| Loreto Peru South America| Requena Peru South America| Madre de Dios Peru South America| San Martín Peru South America| Amapá Brazil South America| Distrito Federal Brazil South America| Maranhão Brazil South America| Mato Grosso Brazil South America| Minas Gerais Brazil South America| Pará Brazil South America| Pernambuco Brazil South America| Rondônia Brazil South America| Roraima Brazil South America| Beni Bolivia South America| La Paz Bolivia South America| Santa Cruz Bolivia South America| Kuna Yala Panama Central America|

Common Names:

pabojoni, e-mo-naweE, ma-eE-pa-he, Pucacuru-caspi, anayo-caspi, boyuyo, Boyuyo homiga, Buyo-Buyo