harsh that many species of trees, vines, and herbs are deciduous for 2 to 6 months. Its rainy season, during which 1 to 3 meters of rain can fall, is as wet, if not wetter, than that of a rain forest. In the dry season, the sun penetrates to the forest floor, the leaf litter becomes very dry (and virtually ceases to decompose), watercourses dry up or greatly diminish in flow, and daytime relative humidity ranges from 20 to 60%. The dry forest may appear uniformly green during the rainy season, but during the dry season this homogeneity changes into a complex mosaic of tens of habitat types distinguished by the differential drying rates of different soils and exposures, different ages of succession, and different vegetation types. Many animals migrate to moist refugia (hollow logs and caves, moist riparian sites, north-facing slopes protected from the wind, and sites close to rain forests). During the dry season, most plants cease their vegetative activities, but many species of woody plants flower, mature their fruits, and disperse their seeds. Some species of animals feed on dry season fruits, seeds, and flowers; for them, the dry season is the bountiful time of year and the rainy season, inimical.

DIVERSITY IN THE DRY FOREST

What is the level of diversity in tropical dry forests? A lowland dry forest adjacent to a lowland rain forest (such as a portion of the Pacific dry and Atlantic wet sides of northern Costa Rica) sustains a fauna and flora about 50 to 100% as species-rich as does the neighboring rain forest (Janzen, 1986a). Floras are the least similar in richness of species, largely because the dry forest epiphytes and trees are substantially less rich in species. The greatest similarity in species richness is represented by mammals and major insect groups such as butterflies and moths (Lepidoptera) and Hymenoptera such as bees, wasps, and ants. Species overlap between the two areas, ranging from less than 5% (e.g., epiphytes, amphibians) to as high as 80% (e.g., sphingid moths, mammals). In the 11,000-hectare dry forest of Santa Rosa National Park in Costa Rica, I estimate that there are 13,000 species of insects, and fairly accurate counts indicate that there are 175 breeding species of birds (Stiles, 1983). There are also 115 species of nonmarine mammals (Wilson, 1983) and about 75 species of reptiles and amphibians (Savage and Villa, 1986). All the species of small herbs and grasses have not yet been collected, but the final list of angiosperms (which include vascular plants such as orchids and trees) will probably not exceed 700 species (Janzen and Liesner, 1980). When such a dry forest habitat is replaced by fencerows, ditchsides, unkempt pastures, and woodlots, the species richness of the breeding fauna and flora is reduced by 90 to 95%.

It is true that long lists of species have been used as criteria for identifying tropical habitats worthy of conservation. However, an approach that merely considers the number of species present is incomplete. This can be seen by examining the tropical dry forest, which is less rich than the rain forest in total species but is much richer in its variety of species’ activities. It contains many species that remain dormant in inclement (wet or dry) weather, and species that magically find enough water to develop flowers, fruits, and leaves at the height of the dry season (e.g., Janzen, 1967, 1982a,b). Its parasitoids and grazers range from absolutely monophagous to extremely polyphagous; that is, some subsist only on one type of



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