• Volume 18, Issue 4

      December 1993,   pages  425-564

    • Population dynamics of animals in unpredictably-changing tropical environments

      Tamiji Inoue Koji Nakamura Siti Salmah Idrus Abbas

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      We studied population dynamics of a solitary phytophagous beetle,Epilachna viqintioctopunctata and a social stingless bee,Trigona minangkabau, in Sumatra, Indonesia for 5 years from 1981.

      Population increase ofEpilachna vigintioctopunctata was suppressed in months of normal rainfall (≥300mm) but was released in the 1982–1983 El Nino-Southern. Oscillation when rainfall dropped to 50% of the long-term average. Mechanisms might be direct; rainfall lowered egg hatchability and the time of adult’s residence on host plants. When dry weather continued for more than three generations, theEpilachna vigintioctopunctata population reached a density at which food shortage due to defoliation occurred. Although parasitism of immature stages was high, it was not a population-regulating factor. Thus, there were two types of ecological crunch: competition for food resources at the end of favourable dry periods and high mortality during heavy rainfall periods that usually followed El Nino-Southern Oscillation dry conditions.

      By an experimental addition of artificial nest sites, colony density ofTrigona minangkabau increased 2.5 times the original density of natural colonies. One-half of artificial nest sites were occupied by arboreal ants and thus competition for nest sites with ants suppressed further increase ofTrigona minangkabau. Intermediate rainfall was favourable forTrigona minangkabau because the rate of colony foundation decreased both during dry El Niño-Southern Oscillation months and months with heavy rain. Colony death was independent from rainfall. Many colonies that survived for 6 months persisted for >2 years and colony density was quite stable.Trigona minangkabau colonies could survive even under unfavourable periods, by hoarding resources in the nest. There was no significant ecological crunch during the study period and colony density almost always tracked the carrying capacity of the habitat, which was basically determined by nest-site abundance.

      Climatic conditions, especially rainfall, changed with various periodicities, 4–5 years for El Nino-Southern Oscillation, and 2 years for the monsoon and other shorter periods. The contribution of periodicities of 1 and 0.5 years, that were linked to movement of the sun, were weak, indicating that animals could not use seasonal changes of environments,e.g. daylength, to predict environmental changes. We discuss traits adaptive to such unpredictably-changing tropical environments. Separation of predictability of temporal environmental change and synchronous changes among patches improves our understanding. Low oviposition rate and resulting prolonged life-span ofEpilachna vigintioctopunctata, usually associated withK-selected traits of life history, seem to be adaptations for unpredictable environmental changes.

    • Mass flowering of dipterocarp forests in the aseasonal tropics

      S Appanah

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      At irregular intervals of 2 to 10 years the aseasonal tropical rain forests in west Malesia come into heavy mass flowering, followed by mast fruiting. During a heavy flowering almost half the mature individuals and over 80% of the canopy and emergent tree Species in a forest may flower. This involves over 200 tree species in a forest flowering over a short period of 3–4 months. The pollination needs during a mass flowering appears to be overcome in several ways. A rapid increase in the number of pollinators seems to occur in the forest. This is partly caused by the migration of pollinators from the fringes of the forest to forage on the superabundance of flowers. At the same time, some groups of plants which share common pollinators appear to reduce pollinator competition by flowering in interspecific sequence. Many members of the family Dipterocarpaceae have evolved sequential flowering too. They also share unique pollinators, common flower thrips which appear to build up rapidly in numbers by feeding and breeding on the millions of dipterocarp flower buds which are present several weeks before the flowering. The environmental cue for this irregular, but widespread mass flowering can be traced to a small dip of about 2° C below mean night-time temperature for 4 or 5 nights. The conditions for such temperature drops occur during El Nino events.

    • Catastrophic and background disturbance of tropical ecosystems at the Luquillo Experimental Forest

      Ariel E Lugo Robert B Waide

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      The forests of the Luquillo Experimental Forest Long-Term Ecological Research site are subject to low-intensity, widespread disturbance that establishes levels of background mortality that contrast with periodic catastrophic mortality resulting from hurricanes and landslides. Although catastrophic mortality is more dramatic, background mortality is still more important in determining population turnover. However, catastrophic mortality may still be an important agent in determining ecosystem structure. Catastrophic disturbances affect forest function in many ways besides mortality, some of which are only apparent in the context of long-term studies. Since most ecosystems are subject to some form of catastrophic disturbance, general principles can be derived from comparative studies of disturbance in different systems.

    • Stable isotopic structure of aquatic ecosystems

      Eitaro Wada Yuko Kabaya Yasushi Kurihara

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      Isotopic, biogeochemical and ecological structure can provide a new dimension for understanding material flows, and the simultaneous function and structure of an ecosystem. Distributions ofδ13C andδ15N for biogenic substances in the Nanakita river estuary involving Gamo lagoon in Japan were investigated to construct isotope biogeochemical and ecological structure for assessing fate and transfer of organic matter, and food web structure. The isotopic framework of the ecosystem was successfully described in aδ15N–δ13C map. In this estuary the variations of isotope ratios of biogenic substances were clearly explained by the mixing of land-derived organic matter, and marine-derived organic matter.

      A trophic-level effect of15N enrichment was clearly observed. Organisms were classified into three groups depending upon the contribution of land-derived organic matter in a food chain. Almost all biota except mollusca in the lagoon depend on organic matter of marine origin. The contributions of both land and marine organic matter were comparable for mollusca in the lagoon.

    • The community ecology of Asian rain forests, in relation to catastrophic events

      P S Ashton

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      Although natural catastrophic disturbance of tropical forests in Asia can be caused by volcanism and earthquakes climate-induced catastrophes are most widespread. These are prevalent and most diverse at high tropical latitudes because of the single annual dry and wet monsoon. Comparative studies indicate that periodicity of catastrophes may influence forest physiognomy, structure and species richness but long-term research in sample plots suggests that a variety of other factors are locally influential. The importance of accounting for natural catastrophes in silvicultural protocols is stressed, and research priorities identified.

    • Environmental responses of plants and ecosystems as predictors of the impact of global change

      F Stuart Chapin Emmanuel Rincon Pilar Huante

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      An understanding of plant responses to fluctuations in environment is critical to predictions of plant and ecosystem responses to climate change. In the northern hemisphere, the northern limits of distribution of major biomes are probably determined by the tolerance of their dominant physiognomic types (e.g., deciduous hardwood trees) to minimum winter temperatures and can thus be predicted from long-term patterns of temperature fluctuations. At a more detailed level, the responses of functional groups of plants to altered climate can be predicted from their known responses to fluctuations in soil resources (nutrients and water) and the expected effect of climatic change on these soil resources. Laboratory and field experiments demonstrate the feasibility of this approach.

    • Evolution of mutualism in plant-pollinator interactions on islands

      Ken Inoue

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      The evolution and ecology of interactions between plants and pollinators are discussed based on the studies on the Izu Islands and mainland Honshu, Japan. The species assemblage is depauperate, and long-tongued pollinators are absent or rare on the islands. Bumblebees, one of the most important pollinators in Japan, are generally absent. Plants depending strongly on bumblebee pollination are absent on Izu Islands, but those depending on varied pollinators including bumblebees display smaller flower sizes and accommodate smaller pollinators than their mainland counterparts. Breeding systems of these species also shift to partial inbreeding, possibly an evolutionary result of the decrease in pollinator availability. Changes in flowering phenology between mainland and island populations also occur. Plants in the islands tend to reproduce vegetatively less frequently and produce greater numbers of smaller seeds than those in the mainland. The possibility of evolution on the side of island pollinator species is also discussed, although there are few data on this topic.

    • Direct costs of forest reproduction, bee-cycling and the efficiency of pollination modes

      David W Roubik

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      The bee guild represents direct primary costs of angiosperm reproduction. Tropical flower visitors take an amount comparable to herbivores, exceeding 3% of net primary production energy. Therefore herbivory and aboveground net primary production have been underestimated. Comparing pollinators to other herbivores, harvest in mature forest by tropical bees is greater than leafcutter ants, game animals, frugivores, vertebrate folivores, insect defoliators excluding ants, flower-feeding birds and bats, but not soil organisms. The ratio of total aboveground net primary production to investment in pollen, nectar and resin used by pollinators suggests wind pollination is several times more efficient in temperate forests than is animal pollination in neotropical moist forest. Animal pollination may be favoured by habitat mosaics and an unpredictable or sparse dispersion of conspecifics — consequences of fluctuating abiotic and biotic environments. Natural selection evidently favours diminished direct reproductive costs in forests, for example by wind pollination, regardless of latitude and disturbance regime. An example is “wind pollination by proxy” of dominant trees in seasonal southeast Asian forests. They flower only occasionally and their pollen is dispersed by tiny winged insects that are primarily carried by the wind — rather than the nectar-hungry bees, bats, birds and moths used by most tropical flora. Increasing evapotranspiration is associated with greater net primary production; I show its correlation with species richness of social tropical bees across the isthmus of Panama, which may indicate increasing forest reproductive effort devoted to flowering, and its monopolization by unspecialized flower visitors in wetter and less seasonal lowland forests.

    • Acknowledgements

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