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(1) During 1978–1981, marked individuals of 107 species of treelets and shrubs in three forest types between 1300–1650 m elevation at Monteverde, Costa Rica, were monitored at monthly intervals for behaviour of leafing, flowering, and fruiting.
(2) Although there was not a pronounced seasonal pattern of leafing activity, more species produced new leaves in the dry season. Species that flush large quantities of new leaves do so more commonly in the drier months. Leaf loss was gradual and unobtrusive in species observed.
(3) Flowering activity was greatest in the late dry season and early wet season. Most species exhibited extended flowering; only 15% of the species were massively flowering. Massive flowerers showed less seasonality than extended flowerers.
(4) Of the species studied, the majority had relatively unspecialized flowers which were visited by a variety of insects; small bee-pollination was the next most common, followed by hummingbird, beetle, settling moth, sphingid, butterfly, large bee and fly pollination (the pollination system of 18 species was unknown). Hummingbird pollinated species showed little seasonality of flowering when compared with species exhibiting small moth, and beetle pollination syndromes, as well as those with unspecialized flowers.
(5) The vast majority of species studied have fleshy fruits (sarcochores). Fruiting activity was less markedly seasonal than flowering. Species with fruit are more numerous in the second half of the year (the wet season and early dry season). The second year of the study saw substantially fewer species in fruit than the first year; this is attributed to the greater than usual rainfall and inclement weather during the peak flowering season.
(6) Cloud forest shrub and treelet phenology is compared with patterns of other forests that have been studied. In general, the greater the rainfall, the less seasonality of flowering and fruiting is seen. Although Monteverde is very wet, rainfall is intermediate between that of lowland dry and lowland wet forest in Costa Rica. Seasonality of flowering and fruiting at Monteverde is more pronounced than at La Selva (wet) and less obvious than in Guanacaste (dry).
Introduction
Conducting a study of the seasonality of flowering, fruiting, and foliation, often termed a phenological study, of species within a plant community is often not an end in itself. It may serve as a way to discover the component species of the community and when each species may be expected to be in a reproductive or vegetative stage suitable for study.
Knowledge of the seasonality of reproductive and vegetative activities by the species in a community is fundamental to other questions. How do the plants respond to environmental variables, notably the climate? Are the phenological patterns of some species affected by competition between plant species for pollinating or dispersal agents? Does foliation occur in such a way as to minimize herbivory? Are the flowering and fruiting of plants with particular pollinatory or dispersal strategies limited to particular seasons? Alternatively, the zoologist may wish to know the seasonal availability of nectar, fruit, or leaf resources for particular animals.
Our approach in this chapter is to emphasize the phenological patterns of reproduction and foliation by many locally sympatric climbers, with a review of the literature and from our own field work in northwestern Costa Rica. We also consider phenological studies of plants (climbers and others), that are adapted for certain groups of pollinators or frugivores.
Community studies
In addition to our work there have been only three other studies on the phenology of climber communities: those by Croat (240 species: 1975) and Putz & Windsor (43 species: 1987), both on Barro Colorado Island, Panama; and that of Hegarty (1988) on the climbers of a Queensland rainforest.
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