sustained by 100 centimeters or more of annual rainfall. Typically two or more other layers of trees and shrubs occur beneath the upper canopy. Because relatively little sunlight reaches the forest floor, the undergrowth is sparse and human beings can walk through it with relative ease.

The species diversity of rain forests borders on the legendary. Every tropical biologist has a favorite example to offer. From a single leguminous tree in the Tambopata Reserve of Peru, I recently recovered 43 species of ants belonging to 26 genera, about equal to the entire ant fauna of the British Isles (Wilson, 1987). Peter Ashton found 700 species of trees in 10 selected 1-hectare plots in Borneo, the same as in all of North America (Ashton, Arnold Arboretum, personal communication, 1987). It is not unusual for a square kilometer of forest in Central or South America to contain several hundred species of birds and many thousands of species of butterflies, beetles, and other insects.

Despite their extraordinary richness, tropical rain forests are among the most fragile of all habitats. They grow on so-called wet deserts—an unpromising soil base washed by heavy rains. Two-thirds of the area of the forest surface consists of tropical red and yellow earths, which are typically acidic and poor in nutrients. High concentrations of iron and aluminum form insoluble compounds with phosphorus, thereby decreasing the availability of phosphorus to plants. Calcium and potassium are leached from the soil soon after their compounds are dissolved from the rain. As little as 0.1% of the nutrients filter deeper than 5 centimeters beneath the soil surface (NRC, 1982). An excellent popular account of rain forest ecology is given by Forsyth and Miyata (1984).

During the 150 million years since its origin, the principally dicotyledonous flora has nevertheless evolved to grow thick and tall. At any given time, most of the nonatmospheric carbon and vital nutrients are locked up in the tissue of the vegetation. As a consequence, the litter and humus on the ground are thin compared to the thick mats of northern temperate forests. Here and there, patches of bare earth show through. At every turn one can see evidence of rapid decomposition by dense populations of termites and fungi. When the forest is cut and burned, the ash and decomposing vegetation release a flush of nutrients adequate to support new herbaceous and shrubby growth for 2 or 3 years. Then these materials decline to levels lower than those needed to support a healthy growth of agricultural crops without artificial supplements.

The regeneration of rain forests is also limited by the fragility of the seeds of the constituent woody species. The seeds of most species begin to germinate within a few days or weeks, severely limiting their ability to disperse across the stripped land into sites favorable for growth. As a result, most sprout and die in the hot, sterile soil of the clearings (Gomez-Pompa et al., 1972). The monitoring of logged sites indicates that regeneration of a mature forest might take centuries. The forest at Angkor (to cite an anecdotal example) dates back to the abandonment of the Khmer capital in 1431, yet is still structurally different from a climax forest today, 556 years later. The process of rain forest regeneration is in fact so generally slow that few extrapolations have been possible; in some zones of greatest combined damage and sterility, restoration might never occur naturally (Caufield, 1985; Gomez-Pompa et al., 1972).



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