Taxon Details: Tococa guianensis Aubl.
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Family:

Melastomataceae (Magnoliophyta)
Scientific Name:

Tococa guianensis Aubl.
Primary Citation:

Hist. Pl. Guiane 1: 438, t. 174. 1775
Accepted Name:

This name is currently accepted.
Type Specimens:

Specimen 1: Paratype -- Y. E. J. Mexia
Common Names:

pabojoni, e-mo-naweE, ma-eE-pa-he, pucacuru-caspi, anayo-caspi, Boyuyo, Boyuyo homiga, Buyo-Buyo
Description:

Description Author and Date: Fabian A. Michelangeli, January 2010, based on Michelangeli, F. A. (2005). Tococa (Melastomataceae). Flora Neotropica Monographs 98: 1-114.

Type: French Guiana: Ad margines viarum Caiennae & Guianae, Aublet s. n. (holotype BM; isotypes P[2]).

Description: Shrub or small tree, to 1-5 (-7) m tall. Stem pubescence highly variable, glabrous, sparsely glandular pubescent, or with a mix of glandular and non-glandular setae, often with a reddish or fuchsia tinge in the younger branches, the setae 2.5 mm long, occasionally up to 12 mm long, persistent, the nodes glabrous. Petioles 0.5-2.0 (-4.5) cm long, sparsely or densely glandular setose. Leaves anisophyllous, seldom isophyllous, ovate to oblong elliptic, the large 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 attenuated, adaxial surface adpressed non-glandular setose, occasionally glandular setose, rarely glabrous, abaxial surface glandular setose, occasionally non-glandular setose, rarely glabrous, the setae denser in the primary and secondary veins, usually light green, purple or dark red in some young individuals, with 2(-3) pairs of secondary veins, basally nervate, the veins reticulate underneath, membranaceous to chartaceous, (slightly coriaceous in individuals growing in the sun), entire to ciliate-serrulate, occasionally ciliate-crenulate; 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; sepals broadly deltoid to ovate, fused halfway to the apex (0.5-1.2 mm), 1.0-3.0 mm long, calyx teeth reduced to well developed into a deltoid teeth 0.3-2.0 mm long, projecting 0 to 1.0 mm above the sepals, terminated in one or more glandular setae up to 3.5 mm long, occasionally glabrous; 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, pruinosus to smooth, sometimes finely granulose, glabrous, occasionally with a caducous terminal glandular seta, pink. Stamens all of the same size, the 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 on a ring elevated 0.1-0.5 mm over the apex of the ovary, the ring glabrous to densely pilose or with a corona of long glandular setae; style glabrous, very rarely minutely stellate pubescent at the base, 13-20 (23) mm long; stigma capitate, 1.5 mm wide (appearing funneliform when dry), the stigmatic surface finely papillate. Fruits globose, 10-19 (-22) mm long, black, glandular setulose, rarely glabrous. The 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.

Habitat and Distribution: Widely distributed from S Mexico to N Bolivia, to the E into E Brazil and French Guiana. Growing in forested and disturbed areas, or in forested savannas from 0 to 1600 m.

Phenology: Flowering year round, but peaking during the rainy season.

Ecology: Tococa guianensis seems to be often associated with Azteca ants, and more rarely with Crematogaster. Even though T. guianensis is found at elevations over 1600 m, often the specimens collected over 1000 m 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).

Taxonomy and Systematics: Tococa guianensis is a highly variable species widely distributed throughout the entire geographical range of the genus. It is very common in both primary and disturbed environments 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 while analyzing all the populations across the entire geographical range, it becomes evident that most of the variation falls within a continuum, which makes it impossible to recognize all of these forms as separate species. Different authors (Cogniaux, 1888; Gleason, 1931b; Macbride, 1941; Wurdack, 1973b, 1980; Wurdack & Renner, 1993; Vasquez, 1997; Michelangeli, 2001b) have addressed this problem, seldom arriving 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 the shape and pubescence of the outer teeth calyx. Twelve possible combinations of calyx and ovary characters were outlined by Gleason (1931b), 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 that in dryer 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” due to the variation in pubescence of the hypanthium and ovary (i.e. T. guianensis and T. occidentalis). 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 specimen at K (Hooker Herbarium) of Barclay annotated as from “Island of Gorgana”, which seems to be annotated by Bentham. Of all the forms here considered synonyms of T. guianensis, T. formicaria seems to be the most distinct one, 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. formicaria falls within the range of distribution of specimens with characters that match the type of T. guianensis. Moreover, in this area (central Brazil) a considerable number of the specimens examined exhibited a continuum of density and size of pubescence between T. guianensis and T. formicaria.

Flora and Monograph Treatment(s):

Tococa guianensis Aubl.: [Article] Michelangeli, Fabián A. 2005. (Melastomataceae). Fl. Neotrop. Monogr. 98: 1-114.
Tococa guianensis Aubl.: [Article] Maguire, Bassett, et al. 1953. The Botany of the Guayana Highland. Mem. New York Bot. Gard. 8 (2): 87-160.
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R. A. A. Schnell 9109, Brazil
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Tococa guianensis Aubl.
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Tococa guianensis Aubl.
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G. T. Prance 8372, Brazil
G. T. Prance 20536, Brazil
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G. T. Prance 15154, Brazil
G. T. Prance 8321, Brazil
G. T. Prance 8211, Brazil
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G. T. Prance 2872, Brazil
G. T. Prance 7320, Brazil
G. T. Prance 3208, Brazil
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P. J. M. Maas 4350, Guyana
P. J. M. Maas 8966, Brazil
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J. L. Luteyn 1143, Panama
J. J. Pipoly 7930, Guyana
J. J. Pipoly 6373, Ecuador
J. C. Solomon 3449, Peru
H. H. Rusby 2242, Bolivia
M. Bang 2668, Bolivia
M. Bang 2668, Bolivia
O. Buchtien 978, Bolivia
O. Buchtien 978, Bolivia
D. E. Atha 1199, Belize
M. H. Nee 30966, Venezuela
M. H. Nee 39658, Bolivia
B. A. Krukoff 10990, Bolivia
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B. A. Krukoff 10917, Bolivia
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B. A. Krukoff 8703, Brazil
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W. W. Thomas 5727, Bolivia
Tococa guianensis Aubl.
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J. A. Steyermark 103021, Venezuela
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G. Klug 1747, Colombia
E. P. Killip 37459, Venezuela
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J. L. Zarucchi 3352, Colombia
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L. Uribe-Uribe 6578, Colombia
R. E. Schultes 18076, Colombia
R. E. Schultes 12700, Colombia
R. E. Schultes 17939, Colombia
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T. B. Croat 19841, Peru
T. B. Croat 56836, Colombia
T. B. Croat 17360, Panama
T. B. Croat 59267, Venezuela
T. B. Croat 98314A, Ecuador
T. B. Croat 62427, Brazil
T. B. Croat 62413, Brazil
T. B. Croat 85669, Brazil
J. M. Idrobo 690, Colombia
H. García-Barriga 14269, Colombia
H. García-Barriga 14201, Colombia
H. García-Barriga 18257, Colombia
H. García-Barriga 18264, Colombia
W. A. Archer 1837, Colombia
W. A. Archer 7990, Brazil
W. A. Archer 8236, Brazil
W. A. Archer 8038, Brazil
J. Lanjouw 593, Suriname
T. W. Henkel 867, Guyana
T. W. Henkel 726, Guyana
W. J. Hahn 5628, Guyana
F. A. González 3391, Colombia
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
T. B. Croat 24218, Belize
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A. Fernandez 2697, Venezuela
R. L. Liesner 21349, Venezuela
A. J. Henderson 946, Venezuela
R. L. Liesner 16446, Venezuela
Tococa guianensis Aubl.
C. Cowan 3263, Mexico
W. R. Philipson 1605, Colombia
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N. A. Rosa 2726, Brazil
D. H. Linder 104, Suriname
Tococa guianensis Aubl.
E. Lleras P16954, Brazil
S. R. Hill 12888, Brazil
C. L. Lundell 3849, Belize
C. L. Lundell 6880, Belize
C. L. Lundell 6880, Belize
W. G. D'Arcy 6225, Panama
A. H. Gentry 5072, Panama
G. Davidse 18739, Venezuela
G. Davidse 2310, Nicaragua
G. Davidse 18323, Venezuela
G. Davidse 18574, Venezuela
G. Davidse 17555, Brazil
G. Davidse 15402, Venezuela
G. Davidse 17994, Brazil
J. Steinbach 2880, Bolivia
J. Steinbach 1450, Bolivia
S. Altamirano 3627, Bolivia
G. Torrico 1373, Bolivia
S. G. Beck 27645, Bolivia
M. Rimachi Y. 2175, Peru
S. T. McDaniel 16050, Peru
F. Ayala 2871, Peru
A. L. Monteagudo Mendoza 4260, Peru
A. H. Gentry 15618, Peru
J. Revilla 83, Peru
R. Scolnik 1169, Peru
G. Klug 415, Peru
S. Vilca C. 347, Peru
C. Díaz Santibañez 10569, Peru
Y. E. J. Mexia 6363, paratype; South America
F. R. Fosberg 29136, Peru
E. P. Killip 29996, Peru
E. P. Killip 29883, Peru
R. Vásquez Martínez 36889, Peru
J. Revilla 943, Peru
R. Vásquez Martínez 26006, Peru
R. Rojas 0959, Peru
Á. J. Pérez 2842, Peru
R. Rojas 439, Peru
H. H. van der Werff 16275, Peru
C. Díaz Santibañez 4122, Peru
J. G. Graham 4806, Peru
E. F. Rodríguez Rodríguez 1157, Peru
J. G. Graham 2702, Peru
J. M. Schunke-Vigo 10534, Peru
J. Revilla 1402, Peru
M. Aguilar 507, Peru
M. Aguilar 1031, Peru
R. Vásquez Martínez 23786, Peru
J. Revilla 3199, Peru
J. Revilla 2751, Peru
F. Ayala 2591, Peru
H. Murphy 321, Peru
I. Huamantupa Chuquimaco 10140, Peru
L. Valenzuela Gamarra 2542, Peru
L. Valenzuela Gamarra 7892, Peru
L. Valenzuela Gamarra 7916, Peru
M. Rimachi Y. 3727, Peru
M. Rimachi Y. 2620, Peru
F. A. Michelangeli 1921, Peru
J. Ruíz 242, Peru
R. Vásquez Martínez 6720, Peru
F. A. Michelangeli 496, Peru
F. A. Michelangeli 498, Peru
F. A. Michelangeli 500, Peru
F. A. Michelangeli 501, Peru
E. P. Killip 27292, Peru
E. P. Killip 27348, Peru
L. W. Williams 3657, Peru
G. Tessmann 3594, Peru
G. Tessmann 5042, Peru
E. P. Killip 26974, Peru
D. R. Simpson 726, Peru
J. D. Boeke 3283, Peru
J. D. Boeke 3291, Peru
R. Vásquez Martínez 2046, Peru
R. Vásquez Martínez 245, Peru
J. M. Schunke-Vigo 5451, Peru
S. D. Knapp 7924, Peru
J. M. Schunke-Vigo 8635, Peru
D. C. Daly 7002, Brazil
D. C. Daly 7105, Brazil
D. A. Neill 14981, Ecuador
C. Morales 1990, Ecuador
D. A. Neill 14659, Ecuador
T. Katan 267, Ecuador
J. P. Janovec 1385, Ecuador
D. E. Atha 1288, Belize
H. T. Beck 172, Brazil
S. A. Mori 9077, Brazil
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
L. O. Williams 28444, Costa Rica
A. Molina Rositto 14971, Nicaragua
A. Molina Rositto 15126, Nicaragua
A. Molina Rositto 14717, Nicaragua
S. A. Mori 8839, French Guiana
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
M. J. Balick 3554, Belize
L. Valenzuela Gamarra 14450, Peru
R. Vásquez Martínez 44492, Peru
Tococa guianensis Aubl.
R. G. Barbosa-Silva 260, Brazil
L. Buscalioni 4162, Brazil
S. Elcoro 694, Venezuela
L. A. Delgado 233, Venezuela
D. Cárdenas-López 24054, Colombia
D. Cárdenas-López 46246, Colombia
A. L. Weitzman 224, French Guiana
M. G. Bovini 3376, Brazil
J. Meirelles 729, Brazil
K. Ruokolainen 10785, Peru
K. Ruokolainen 12444, Colombia
K. Ruokolainen 5407, Peru
K. Ruokolainen 12425, Colombia
K. Ruokolainen 11599, Ecuador
K. Ruokolainen 12969, Peru
K. Ruokolainen 10220, Peru
K. Ruokolainen 9994, Peru
C. V. Martin 408, French Guiana
C. V. Martin 563, French Guiana
C. V. Martin 431, French Guiana
C. V. Martin 453, French Guiana
C. V. Martin 402, French Guiana
S. A. Antoniazzi s.n.
L. F. Sardelli s.n.
L. F. Sardelli s.n., Brazil
N. Castaño Arboleda 3795, Colombia
H. Medeiros 164, Brazil
I. Brasil 496, Brazil
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
P. H. Gentle 3614, Belize
I. G. Vargas Caballero 3456, Bolivia
I. G. Vargas Caballero 3491, Bolivia
I. G. Vargas Caballero 3902, Bolivia
D. C. Daly 11631, Brazil
B. M. Boom 6499, Venezuela
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
W. E. Broadway 710, French Guiana
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.
Tococa guianensis Aubl.