widespread, usually transitional between semidesert or savanna and moist forest. About 40% of the earth's tropical and subtropical landmass is dominated by open or closed forest. Of this, 42% is dry forest, 33% is moist forest, and only 25% is wet and rain forest (sensu Holdridge, 42; 15). We will never know the true original or potential extent of dry forest because many savannas and scrub or thorn woodlands are thought to be derived from disturbed dry forest. Walter (110), for example, considers most or all of the grassland in India to have been derived from seasonal or dry forest. Some of the processes that cause this conversion are addressed later in this review. The largest proportion of dry forest ecosystems is in Africa and the world's tropical islands, where they account for 70-80% of the forested area. In South America they

Ecology of Tropical Dry Forest

Early studies of nutrient cycling in moist tropical forests described productive forests rich in nutrients (98, 114, 176) in which rates of primary production and the amounts of nutrients cycled clearly exceeded those in temperate zone forests. Reviews of global-scale patterns in biomass, production, and nutrient cycling reported these results as representative of tropical forests (130, 175). At the same time, tropical forest soils were described as acid, infertile clays that harden irreversibly to "laterite" when cleared (106), or as bleached quartz sands low in mineral nutrients (88). This apparent paradox was crystallized by Whittaker (174) in the statement "The tropical rain forest thus has a relatively rich nutrient economy perched on a nutrient-poor substrate" (p. 271). Reviews of more recent research on overall patterns of mineral cycling in the tropics (78, 124), and of important components such as biomass (20, 21), litterfall nutrients (123, 164), and decomposition (5), clearly show that patterns of nutrient cycling in tropical forests are diverse. It makes no more sense to describe a 'typical' tropical forest than a 'typical' temperate forest (33, 151). Variations in mineral cycling nonetheless follow coherent, explicable patterns in tropical forests. Our purposes in undertaking this review are: (a) to illustrate the patterns of nutrient cycling in moist tropical forests; (b) to identify the mechanisms which regulate those patterns; and (c) to show how those patterns affect the productivity, physiology, and population biology of tropical forests and their large-scale linkages with aquatic ecosystems and the atmosphere. We emphasize the cycling of nitrogen, phosphorus, potassium, calcium, and magnesium; these are the best-studied soil-derived nutrients, and they are the nutrients most likely to limit primary production and other ecosystem functions.

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Nutrient cycling in moist tropical forest

Despite uncertainty in all levels of analysis, recent and long-term changes in our climate point to the distinct possibility that greenhouse gas emissions have altered mean annual temperatures, precipitation and weather patterns. Modeling efforts that use doubled atmospheric CO2 scenarios predict a 1–7°C mean global temperature increase, regional changes in precipitation patterns and storm tracks, and the possibility of “surprises” or sudden irreversible regime shifts. The general effects of climate change on freshwater systems will likely be increased water temperatures, decreased dissolved oxygen levels, and the increased toxicity of pollutants. In lotic systems, altered hydrologic regimes and increased groundwater temperatures could affect the quality of fish habitat. In lentic systems, eutrophication may be exacerbated or offset, and stratification will likely become more pronounced and stronger. This could alter food webs and change habitat availability and quality. Fish physiology is inextricably linked to temperature, and fish have evolved to cope with specific hydrologic regimes and habitat niches. Therefore, their physiology and life histories will be affected by alterations induced by climate change. Fish communities may change as range shifts will likely occur on a species level, not a community level; this will add novel biotic pressures to aquatic communities. Genetic change is also possible and is the only biological option for fish that are unable to migrate or acclimate. Endemic species, species in fragmented habitats, or those in east–west oriented systems will be less able to follow changing thermal isolines over time. Artisanal, commercial, and recreational fisheries worldwide depend upon freshwater fishes. Impacted fisheries may make it difficult for developing countries to meet their food demand, and developed countries may experience economic losses. As it strengthens over time, global climate change will become a more powerful stressor for fish living in natural or artificial systems. Furthermore, human response to climate change (e.g., increased water diversion) will exacerbate its already-detrimental effects. Model predictions indicate that global climate change will continue even if greenhouse gas emissions decrease or cease. Therefore, proactive management strategies such as removing other stressors from natural systems will be necessary to sustain our freshwater fisheries.

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Potential impacts of global climate change on freshwater fisheries

We suggest here that large-scale natural forest disturbance and primary succession in the lowland rainforests of the Peruvian Amazon is caused by lateral erosion and channel changes of meandering rivers. Our results indicate that in the upper Amazon region, primary succession on newly deposited riverine soils is a major mode of forest regeneration. Landsat imagery analyses show that 26.6% of the modern lowland forest has characteristics of recent erosional and depositional activity; 12.0% of the Peruvian lowland forest is in successional stages along rivers. This successional development is used to classify the western Amazon rainforests according to their geomorphological erosion–deposition pattern. We also propose that by causing high site turnover, disturbance and variation in forest structure, the river dynamics may be a major factor creating and maintaining the high between-habitat (β-type) species diversity characterizing the upper Amazon1.

River dynamics and the diversity of Amazon lowland forest

The amount and spatial distribution of forest biomass in the Amazon basin is a major source of uncertainty in estimating the flux of carbon released from land-cover and land-use change. Direct measurements of aboveground live biomass (AGLB) are limited to small areas of forest inventory plots and site-specific allometric equations that cannot be readily generalized for the entire basin. Furthermore, there is no spaceborne remote sensing instrument that can measure tropical forest biomass directly. To determine the spatial distribution of forest biomass of the Amazon basin, we report a method based on remote sensing metrics representing various forest structural parameters and environmental variables, and more than 500 plot measurements of forest biomass distributed over the basin. A decision tree approach was used to develop the spatial distribution of AGLB for seven distinct biomass classes of lowland old-growth forests with more than 80% accuracy. AGLB for other vegetation types, such as the woody and herbaceous savanna and secondary forests, was directly estimated with a regression based on satellite data. Results show that AGLB is highest in Central Amazonia and in regions to the east and north, including the Guyanas. Biomass is generally above 300Mgha(sup 1) here except in areas of intense logging or open floodplains. In Western Amazonia, from the lowlands of Peru, Ecuador, and Colombia to the Andean mountains, biomass ranges from 150 to 300Mgha(sup 1). Most transitional and seasonal forests at the southern and northwestern edges of the basin have biomass ranging from 100 to 200Mgha(sup 1). The AGLB distribution has a significant correlation with the length of the dry season. We estimate that the total carbon in forest biomass of the Amazon basin, including the dead and below ground biomass, is 86 PgC with +/- 20% uncertainty.

Distribution of Aboveground Live Biomass in the Amazon Basin

The Amazon -that name was given to the biggest river on earth and is often used for the whole area of its basin too. This geographical region is currently referred to as Amazonia, thus emphasizing the peculiar character of its aquatic and terrestrial reaches. The Amazon embodied the dream of many a naturalist to explore what for a long time was a terra incognita. In recent years, however, Amazonia has emerged as a main centre for 'development' by some of the countries in which it lies and by foreign industrialized nations. The development projects and enterprises have aroused woridwide interest and have given rise to discussions on their aims and their consequences to the Amazonian nature. Limnological and ecological investigations in Amazonia started only about 40 years ago. The editor had the good fortune to partake in them from the very beginning. He spent his decisive years in Amazonia, and dedicated his life's work to that research and to that country and the Amazonian people. Nearing the end of his scicntific activities, hc is gratcful to bc ablc to summarizc in this book most of the knowledge we possess at present of Amazonian limnology and landscape ecology.

Harald Sioli

Papers

The Amazon : limnology and landscape ecology of a mighty tropical river and its basin