Habitat differences in cocoa tree flowering, fruit-set, and pollinator availability in Costa Rica

ABSTRACT Phenological patterns of flowering and fruit-set were studied in cocoa trees (Theobroma cacao) (Sterculiaceae) at monthly intervals in two contrasting habitats in Costa Rica for a one-year period. One of these habitats, a well-maintained plantation, had irregular and broken shade cover {Erythrinatrees in particular) while in the other habitat, a ‘cocoa forest’, cocoa trees were heavily shaded byHuara crepitans(Euphorbiaceae). ‘Matina’ variety cocoa trees of about the same age (50–60 years) were censused in both habitats. Cocoa-pollinating midge (Diptera: Ceratopogonidae and Cecidomyiidae) availability was estimated by sampling immature stages in experimentally-distributed ground-cover breeding substrates, comparing overall abundances and species numbers between the two habitats over three census dates (dry, mid-rainy, and late-rainy seasons), along with examination of fungal-rotted (diseased) cocoa pods both on trees and the ground. Although total flower production was much greater in the plantation habitat, total production of new pods was similar between habitats. Flowering followed a cyclic temporal pattern in the forest but not in the plantation. Sudden leaf drop of forest shade trees in the dry season probably triggered a cyclic response in which flowering peaked in the first half of the rainy season. There was an inverse relation for frequencies of mature cocoa pods killed by squirrels and pathogenic fungi (Monilia roreriandPhytophthora) between the two habitats: squirrel-killed pods were far more abundant in the plantation than in the forest, and the opposite for fungus-killed pods. Fungus-killed but otherwise intact pods, and not squirrel-killed pods rotting on trees, were a major breeding site for midges, particularly during the late rainy season. Ceratopogonidae were most abundant in the dry season and frequently encountered in cocoa pod husks and banana tree trunk sections in both habitats, and much more so in the forest habitat. The abandoned cocoa plantation (cocoa forest) supported a more diverse assemblage of pollinating midges than the plantation. In the plantation but not in the forest, a negative correlation was discovered between distance from shade trees and the numbers of pods on trees, suggesting greater pollinating activity by midges in cocoa trees beneath shade trees than away from them. The uniform dense shade cover in the adjacent forest probably obliterated such a pattern.