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Family:

Lecythidaceae (Magnoliophyta)

Primary Citation:

Linnaea 27: 444. 1854

Accepted Name:

This name is currently accepted.

Description:

Author: Scott A. Mori

Type: Colombia. Antioquia: near Turbaco, without date (fl), Humboldt & Bonpland 1411 (holotype, P; isotypes, B-Willdenow (2 sheets), F, GH photo, P)

Description: Trees, to 20 m x 25 cm, sparsely branched, the leaf-bearing branches 5-20 mm diam., often arching upwards, the leaves tightly compacted at ends. Bark brown, slightly fissured. Leaves: petioles 5-150 x 1-9 mm, semi-circular in cross section; blades oblanceolate, 25-128 x 6-25 cm, glabrous, chartaceous the base long tapered to acute or cuneate, the margins serrate, the apex acuminate; secondary veins in 16-36 pairs. Inflorescences cauline, racemose, puberulous, with 3-12 flowers, the rachis 10-64 mm long; pedicels 29-85 mm long, subtended by a single ovate to oblong basal bract 1.8-4 x 1.5-5 mm and bearing at or below middle 2 ovate, cucullate bracteoles 1-6 x 1.5-5.5 mm, the latter often fused at bases to form a cup-like structure which encircles pedicel, the cup often with inconspicuous extrafloral nectaries in axils (only easily seen in picked young flowers). Flowers 10-15 cm diam.; calyx entire or very slightly 4-lobed; petals (7-)8(-9), obovate, oblanceolate, or oblong, 33-67 x 12-37 mm, glabrous or puberulous, white with pink at apex and with pink speckles throughout; androecial base 8-15 mm high, the outermost filaments 10-23 mm long, yellow at base pink at apex, the anthers 2.5-4 mm long, yellow; ovary smooth, puberulous (5-) 6-locular, with 18-30 ovules per locule, puberulous at summit, the style 1-2 mm long. Fruits globose or depressed globose, 3-9 x 4-100 cm, the opercular region sunken below fruit surface and < 1/2 diam. of fruit diam., the calyx forming a raised, entire, circular rim, the exocarp yellow at maturity, the mesocarp orange. Seeds usually angled in cross section, 12-35 x 11-22 mm, the seed coat turns brown at maturity; aril poorly developed, yellow, surrounding funicle. x = 34. Also see Carrasquilla (2005) for description and color images.

Common names: Panama: membrillo, membrillo hembra, wild mango. Colombia: membrillo, paco.

Distribution: From central Panama into northwestern Colombia. It is very common and conspicuous in the Canal Zone.

Ecology: This species is found in lowland moist forest, semi-deciduous forest, and disturbed habitats. This and several other species of Gustavia do well in disturbed habitats, perhaps because they have the ability to sprout from cut stems. This species is a common and conspicuous (because of the large leaves clustered at the ends of the branches) tree along disturbed roadsides in the Canal Zone, Panama. In a study of Gustavia superba on Barro Colorado Island and the adjacent mainland (Sork 1985), it was concluded that this species germinates well in gaps, forest edges, and forest understory. The mean germination percentage was caluculated at 85.4% and the seeds took 40-70 days to germinate. Both buried and unburied seeds germinated well under the shade of the forest canopy while unburied seeds germinated poorly in a large light gap. The most advanced forest type studied (300 years old) had the lowest adult and seedling densities. It appears from this study that Gustavia superba does not colonize gaps but is most common in early successional forests. Oppenheimer and Lang (1969) have observed white-faced monkeys eating the terminal buds of this species. They claimed that individuals on Barro Colorado Island were more branched than those found on the mainland because monkeys are common on the island and rare on the mainland because of hunting. This may be true, but at the time they wrote their paper the monopodial or few-branched G. grandibracteata, which occurs on the mainland but not on the island, had not been described.

Phenology: A study of vegetative, flowering, and fruiting phenology has been published by Mori and Kallunki (1976) for a population of G. superba found in the Canal Zone, Panama. In this area, leaves fall all year long but there is a peak of leaf fall in the first month of the wet season (March). Leaf flush peaks at two times of the year, once at the beginning of the dry season (December) and again just after the onset of the rainy season (May and June). Flowering occurs during the latter three quarters of the dry season but may be prolonged into the wet season if the dry season is severe. Fruiting takes place from the end of the dry season into the first part of the wet season (April through August). Sork (1995) concludes that G. superba is a species that fruits and germinates seed during the rainy season. Johnston (1949) has studied the phenology of this species on San José Island, Panama where he noted that new shoots elongate mostly in January but some are flushing throughout the year, flowers are most abundant in May and June, and that July is probably the month in which most fruit is produced.

Pollination: The flowers are visited by bees belonging to at least the genera Melipona, Trigona, and Xylocopa. The species of the first two genera are probably too small to be efficient pollinators whereas species of Xylocopa and other large bees are more likely to be much more efficient pollinators of G. superba. Because the anthers open by apical pores, the species is probably most efficiently pollinated by buzz-pollinating bees.

Dispersal: The seeds of this species are dispersed by many mammals, including white-faced monkeys, red-tailed squirrels, agoutis, and probably pacas, collared peccaries, and spiny rats. These animals carry away the fruits to eat the mesocarp (Sork, 1987).

Predation: Agoutis and red-tailed squirrels eat the seeds. However, only agoutis are known to recover buried seeds and consume them (Sork, 1987). The leaves of this species are preyed upon by the larvae of a skipper buttefly (Entheus priassus) which is most successful eating tender, newly flushed leaves in comparison with older, lignified ones (Aide & Londoño, 1989).

Field characters: Gustavia superba is characterized by its pachycaul, sparsely-branched growth form; thick, leaf-bearing stems; large, membranous leaves that dry a light green; cauline inflorescences; small bracts and bracteoles subtending the flowers; yellow fruits with an orange mesocarp and an opercular area notably smaller in diameter than the overall fruit diameter; and large seeds without a distinctive funicle or aril.

Taxonomic notes: This and other species of Gustavia often emit a very unpleasant odor when the foliage and bark are crushed or when branches and other litter rot for an extended period of time (Johnston, 1949).

Conservation: This species is not listed as endangered by the IUCN Red List of Threatened Species verson 2011.2. It thrives in disturbed habits throughout its range.

Uses: Carrasquilla (2005) reports that this species is often used as an ornamental because of its beautiful flowers and that the fruit pulp is eaten after being cooked. He noted that the pulp is rich in carotenoids, fatty acids, and contains phosphorous. Johnston (1949) says that the pulp is edible but he did not find it very palatable and added that he tasted the seeds and found them to have "...a most unpleasant and very persistent bitter taste." According to Carrasquilla (2005), the Kuna Indians use a concoction of the cooked bark as a medicine to relieve body pains.

Etymology: The specific epithet refers to the beauty of this magnificient plant.

Source: Based on Mori in Prance & Mori, 1979.

Acknowledgements: The author is grateful to A. Berkovk, C. Galdames, and C. Gracie for allowing the use of their images.

Flora and Monograph Treatment(s):

Gustavia superba (Kunth) O.Berg: [Article] Prance, Ghillean T. & Mori, S. A. 1979. Lecythidaceae - Part I. The actinomorphic-flowered New World Lecythidaceae (Asteranthos, Gustavia, Grias, Allantoma & Cariniana). Fl. Neotrop. Monogr. 21: 1-270.