Hostname: page-component-cd9895bd7-q99xh Total loading time: 0 Render date: 2024-12-24T00:20:43.508Z Has data issue: false hasContentIssue false

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

Published online by Cambridge University Press:  10 July 2009

Allen M. Young
Affiliation:
Invertebrate Zoology Section, Milwaukee Public Museum, Milwaukee, Wisconsin 53233, USA

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 {Erythrina trees in particular) while in the other habitat, a ‘cocoa forest’, cocoa trees were heavily shaded by Huara 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 roreri and Phytophthora) 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.

Type
Research Article
Copyright
Copyright © Cambridge University Press 1986

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

LITERATURE CITED

Alvim, P. de T. 1977. Cacao. Pp. 279313 in Alvim, P. de T. & Kozlowski, T. T. (eds). Ecophysiology of tropical crops. Academic Press, New York.Google Scholar
Alvim, P. de T. 1984. Flowering of cocoa. Cocoa Growers' Bulletin 35:2331.Google Scholar
Azhar, I. & Wahi, S. M. 1984. Pollination ecology of cocoa in Malaysia: identification of taxonomic composition and breeding sites, ecology and pollinating activities, and season abundance. Proceedings of the 1984 International Conference on Cocoa and Coconuts: 115.Google Scholar
Borchert, R. 1980. Phenology and ecophysiology of tropical trees: Erythrina poeppigiana O.F. Ecology 61:10651074.Google Scholar
Bowden, J. 1976. Weather and the phenology of some African Tabanidae. Journal of the Entomological Society of South Africa 39:207245.Google Scholar
Bystrak, P. G. & Wirth, W. W. 1978. The North American species of Forcipomyia, subgenus Euprojoannisia (Diptera: Ceratopogonidae). US Department of Agriculture Technical Bulletin No. 1591.Google Scholar
De la Cruz, J. & Soria, S. de J. 1973. Estudio de fluctuaciones de polinizacion del cacao por las mosquitas Forcipomyia spp. (Diptera, Ceratopogonidae), en Palmira, Valle, Colombia. Acta agronomia 23:117.Google Scholar
Dessart, P. 1961. Contribution a l'étude des Ceratopogonidae (Diptera). Bulletin agricole de Congo Belgique 52:525540.Google Scholar
Downes, J. A. 1955. The food habits and description of Atrichopogon pollinivorus sp. n. (Diptera: Ceratopogonidae). Transactions of the Royal Entomological Society of London 106:439453.CrossRefGoogle Scholar
Downes, , 1958. The feeding habits of biting flies and their significance in classification. Annual Review of Entomology 3:249266.CrossRefGoogle Scholar
Entwistle, P. F. 1972. Pests of cocoa. Longmans, London.Google Scholar
Gibbs, D. G. & Leston, D. 1970. Insect phenology in a forest cocoa-farm locality in West Africa. Journal of Applied Ecology 7:519548.CrossRefGoogle Scholar
Hernandez, J. 1965. Insect pollination of cacao (Theobroma cacao L) in Costa Rica. Doctoral dissertation, University of Wisconsin, Madison.Google Scholar
Hutcheon, W. F. 1981. Physiological studies on cocoa (Theobroma cacao L.) in Ghana. Doctoral dissertation, University of Aberdeen, Scotland.Google Scholar
Janzen, D. H. 1967. Synchronization of sexual reproduction of trees within the dry season in Central America. Evolution 21:620637.CrossRefGoogle ScholarPubMed
Kaufmann, T. 1973a. Behavioural ecology of a cocoa pollinator Forcipomyia inomatipennis (Diptera: Ceratopogonidae) in Ghana. Journal of the Kansas Entomological Society 47:541548.Google Scholar
Kaufmann, T. 1973b. Preliminary observations on a cecidomyiid midge and its role as a cocoa pollinator in Ghana. Ghana Journal of Agricultural Science 6:10931098.Google Scholar
Kaufmann, T. 1974. Pollinators of Theobroma cacao in Ghana. Proceedings of the 4th International Cocoa Research Conference (West Africa).Google Scholar
Kaufmann, T. 1975a. Cocoa pollination by males of Forcipomyia squamipennis (Diptera: Ceratopogonidae) in Ghana. Tropical Agriculture 52:7174.Google Scholar
Kaufman, T. 1975b. Biology and behaviour of cocoa pollinating Ceratopogonidae in Ghana. Environmental Entomology 4:347351.CrossRefGoogle Scholar
Kaufman, T. 1975c. An efficient, new cocoa pollinator, Lasioglossum sp. (Hymenoptera: Halictidae) in Ghana, West Africa. Turrialba 25:9091.Google Scholar
Lounibos, L. P. 1981. Habitat segregation among African treehole mosquitoes. Ecological Entomology 6:125134.CrossRefGoogle Scholar
McGregor, S. E. 1976. Insect pollination of cultivated crops. US Department of Agriculture Handbook No. 496.Google Scholar
McLachlan, A. J. & Cantrell, M. A. 1980. Survival strategies in tropical rain pools. Oecologia 47:344351.Google Scholar
Opler, P. A., Frankie, G. W. & Baker, H. G. 1976. Rainfall as a factor in the release, timing, and synchronization of anthesis by tropical trees and shrubs. Journal of Biogeography 3:231236.Google Scholar
Porter, C. H. & Defoliart, G. R. 1981. The man-biting activity of phlebotomine sand flies (Diptera: Psychodidae) in a tropical wet forest environment in Colombia. Archivos de zoologia, Museo de zoologia, Universidad de Sao Paulo 30:81158.Google Scholar
Privat, F. 1979. Les Bromeliacèes, lieu de développement de quelques insectes pollinisateurs des fleurs de cacao. Brenesia 16:197212.Google Scholar
Saunders, L. G. 1959. Methods for studying Forcipomyia midges, with special reference to cacao-pollinating species (Diptera, Ceratopogonidae). Canadian Journal of Zoology 37:3351.CrossRefGoogle Scholar
Soetardi, R. G. 1959. De Betekenis van Insecten Bij Bestuiving van Theobroma cacao L. Archiv für koffiekultur, Indonesie 17:13.Google Scholar
Soria, J. V. 1963. An experimental attempt to repeat some of the spontaneous cacao hybrid populations of Mexico. Cacao (IIAS, Turrialba) 8:1921.Google Scholar
Soria, S. de J. 1970. Studies on Forcipomyia spp. midges (Diptera, Ceratopogonidae) related to the pollination of Theobroma cacao L.. Doctoral dissertation, University of Wisconsin, Madison.Google Scholar
Soria, S. de J. 1973. Locais de coleta e distribuicao de Forcipomyia (Diptera, Ceratopogonidae) relacionadas com a floracao e fruitificacao do cacauerio na Bahia, Brasil. Revista Theobroma 3:4149.Google Scholar
Soria, S. de J. 1975. O papel das abelhas sem ferrao (Meliponinae) na polinizacao do cacaueiro na America tropical. Revista Theobroma 5:1520.Google Scholar
Soria, S. de J. 1976. Tabelas etarias dos polinizadores do cacaueiro Forcipomyia spp. (Diptera, Cera topogonidae) em condicoes de laboratorio. Revista Theobroma 6:513.Google Scholar
Soria, S. de J. 1977a. Dinamica populacional de Forcipomyia spp. (Diptera, Ceratopogonidae) na Bahia, Brasil. 2 – variaveis bioticas relacionadas com a polinizacao do cacueiro. Revista Theobroma 7:1933.Google Scholar
Soria, S. de J. 1977b. Dinamica populacional de Forcipomyia spp. (Diptera, Ceratopogonidae) na Bahia, Brasil. 3 – variaveis climaticas relacionadas com a polinazacao do cacaueiro. Revista Theobroma 7:6984.Google Scholar
Soria, S. de J. 1977c. Las relaciones de variables climaticas y bioticas con la dinamica de poblaciones de Forcipomyia spp. (Diptera, Ceratopogonidae) y la polinizacion de cacaotero en Bahia, Brasil. Turrialba 27:424426.Google Scholar
Soria, S. de J. 1978. Laboratory rearing of Forcipomyia spp. midges (Diptera, Ceratopogonidae). 2. Determination of the reproductive and biotic potentials, preliminary tests. Revista Theobroma 8:6171.Google Scholar
Soria, S. de J. & Abreu, J. M. 1974. Mortalidade dos polinizadores Forcipomyia spp. (Diptera, Ceratopogonidae) causada pela applicacao de inseticidas nos cacauais Baianos. Revista Theobroma 4:1325.Google Scholar
Soria, S. de J. & Abreu, J. M. 1976. Dinamica populacional de Forcipomyia spp. (Diptera, Ceratopo gonidae) na Bahia, Brasil. 1. Fluctuacao estacional dos polinizadores do cacaueiro relacionada com chuva e bianco de agua (Thornthwaite). Revista Theobroma 6:4754.Google Scholar
Soria, S. de J. & Chapman, R. K. 1983. The floral biology of cacao. 2. Observations on foraging activity of Forcipomyia midges. Revista Theobroma 14:5359.Google Scholar
Soria, S. de J. & Wirth, W. W. 1974. Identidade e caracterizacao taxonomica preliminar das mosquinhas Forcipomyia (Diptera, Ceratopogonidae) associadas com a polinizacao do cacaueiro na Bahia. Revista Theobroma 4:312.Google Scholar
Soria, S. de J. & Wirth, W. W. 1975. Ciclos de vida dos polinizadores do cacaueiro Forcipomyia spp. (Diptera, Ceratopogonidae) e algumas anotacoes sobre o comportamento das larvas no laboratorio. Revista Theobroma 5:3–22.Google Scholar
Soria, S. de J. & Wirth, W. W. 1979. Ceratopogonid midges (Diptera: Nematocera) collected from cacao flowers in Palmira, Colombia: an account of their pollinating abilities. Revista Theobroma 9:7784.Google Scholar
Soria, S. de J., Wirth, W. W. &Flores, J. D. 1976. Identidad de las mosquitas Forcipomyia spp. (Diptera, Ceratopogonidae) relacionadas con la polinizacion del cacaotero en Ecuador. Revista Theobroma 6:101108.Google Scholar
Soria, S. de J., Wirth, W. W. & Besemer, H. A. 1978. Breeding places and sites of collection of adults of Forcipomyia spp. midges (Diptera, Ceratopogonidae) in cacao plantations in Bahia, Brazil: a progress report. Revista Theobroma 8:2129.Google Scholar
Soria, S. de J., Chapman, R. K. & Knoke, J. 1980. The floral biology of cacao. 1. Attractants and food substances for adult midges. Revista Theobroma 11:4752.Google Scholar
Strand, M. A. 1984. Comparative floral morphology of four Theobroma (Sterculiaceae) species in relation to pollination biology. Master's dissertation, University of Wisconsin, Milwaukee.Google Scholar
Vello, F. & Magalhaes, W. S. 1971. Studies on the role of the cacarema ant (Azteca chartifex spiriti Forel) on the pollination of cacao in Bahia. Revista Theobroma 1:367398.Google Scholar
Wellensiek, S. J. 1932. Flower-biological observations with cocoa. Archiv für koffiekultur, Indonesie 6:87101.Google Scholar
Winder, J. A. 1972. Cacao pollination: microdiptera of cacao plantations and some of their breeding places. Bulletin of Entomological Research 61:651655.Google Scholar
Winder, J. A. 1977a. Some organic substrates which serve as insect breeding sites in Banian cocoa plantations. Revista brasileño de biología 37:351356.Google Scholar
Winder, J. A. 1977b. Field observations on Ceratopogonidae and other Diptera: Nematocera associated with cocoa flowers in Brazil. Bulletin of Entomological Research 67:5763.CrossRefGoogle Scholar
Winder, J. A. 1977c. Recent research on insect pollination of cocoa. Cocoa Growers Bulletin 26:1119Google Scholar
Winder, J. A. 1978a. Cocoa flower Diptera: their identity, pollinating activity and breedings sites. PANS 24:518.Google Scholar
Winder, J. A. 1978b. The role of non-dipteran insects in the pollination of cacao in Bahia. Proceedings of the 4th International Cocoa Research Conference, Trinidad and Tobago.Google Scholar
Wirth, W. W. 1970. The Neotropical Forcipomyia midges of the subgenus Thyridomya Saunders (Dip tera, Ceratopogonidae). Studies in Entomology 13:429440.Google Scholar
Wirth, W. W. 1982. The cacao-pollinating midges of the Forcipomyia argenteola group (Diptera, Cera topogonidae). Proceedings of the Entomological Society of Washington 84:568585.Google Scholar
Wirth, W. W. & Soria, S. de J. 1975. A new Neotropical Forcipomyia midge closely related to F. (F.) genualis (Loew) (Diptera, Ceratopogonidae). Revista Theobroma 5:1927.Google Scholar
Wirth, W. W. & Soria, S. de J. 1979. Studies on the genus Forcipomyia. VI. The Neotropical species of the subgenus Warmkea (Diptera, Ceratopogonidae). Revista Theobroma 9:137161.Google Scholar
Wirth, W. W. & Waugh, W. T. 1976. Five new Neotropical Dasyhelea midges (Diptera: Ceratopo gonidae) associated with culture of cocoa. Studies in Entomology 19:223236.Google Scholar
Young, A. M. 1981. The ineffectiveness of the stingless bee, Trigonajaty (Hymenoptera: Apidae: Meli-poninae) as a pollinator of cocoa (Theobroma cacao L.). Journal of Applied Ecology 18:149155.Google Scholar
Young, A. M. 1982. Effects of shade cover and availability of midge breeding sites on pollinating midge populations and fruit set in two cocoa farms. Journal of Applied Ecology 19:4763.Google Scholar
Young, A. M. 1983. Seasonal differences in abundance and distribution of cocoa-pollinating midges in relation to flowering and fruit-set between sunny and shaded habitats of the La Lola Cocoa Farm in Costa Rica. Journal of Applied Ecology 20:801831.Google Scholar
Young, A. M. 1984a. Flowering and fruit-setting patterns of cocoa trees (Theobroma cacao L.) (Ster-culiaceae) at three localities in Costa Rica. Turrialba 34:129142.Google Scholar
Young, A. M. 1984b. Ecological notes on cacao-associated midges (Diptera: Ceratopogonidae) in the “atongo” cacao plantation at Turrialba, Costa Rica. Proceedings of the Entomological Society of Washington 86:185194.Google Scholar
Young, A. M. 1984c. Research on the natural pollination of cocoa in Central America: overview of current directions. Proceedings of the 9th International Cocoa Research Conference, Lome, Togo: 557565.Google Scholar
Young, A. M. 1985a. Studies of cecidomyiid midges (Diptera: Cecidomyidae) as cocoa pollinators in Central America. Proceedings of the Entomological Society of Washington 87:4979.Google Scholar
Young, A. M. 1985b. Pollen collecting by stingless bees on cacao flowers. Experientia 41:760762CrossRefGoogle Scholar
Young, A. M. 1985c. Cocoa pollination. Cocoa Grower's Bulletin 36: in press.Google Scholar
Young, A. M., Schaller, M. & Strand, M. 1984. Floral nectaries and trichomes in relation to pollination in some species of Theobroma and Herrania (Sterculiaceae). American Journal of Botany 71:466480.Google Scholar