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56.14 Small, Distant Islands Have Fewer Bird Species The dots show the numbers of land and freshwater bird species on islands of different sizes in the Moluccas, Melanesia, Micronesia, and Polynesia. These islands have been divided into three groups according to their distance from the "mainland,"which in this case is New Guinea.

Species richness on islands and peninsulas is always less than that in an equivalent area on the nearest mainland. On islands, species richness is positively correlated with island size, but inversely correlated with distance from the mainland (Figure 56.14). An influential model relates this pattern to the island's history of immigrations and extinctions.

Species richness is related to rates of immigration and extinction

Over periods of a few hundred years (during which specia-tion is unlikely), the species richness of an area is influenced by the immigration of new species and the extinction of species already present. It is easiest to visualize the effects of these two processes if we consider, as did Robert MacArthur and Edward O. Wilson, an oceanic island that initially has no species.

Imagine a newly formed oceanic island that receives colonists from a mainland area. The list of species on the mainland that might possibly colonize the island is called the species pool. The first colonists to arrive on the island are all "new" species because no species live there initially. As the number of species on the island increases, a larger fraction of colonizing individuals will be members of species already present. Therefore, even if the same number of species arrive as before, the rate of arrival of new species should decrease, until it reaches zero when the island has all the species in the species pool. As we will see, however, the process is unlikely to proceed that far.

Now consider extinction rates. At first there will be only a few species on the island, and their populations may grow large. As more species arrive and their populations increase, the resources of the island will be divided among more species. Therefore, the average population size of each species will become smaller as the number of species increases. The smaller a population, the more likely it is to become extinct. In addition, the number of species that can possibly become extinct increases as species accumulate on the island. Furthermore, new arrivals on the island may include pathogens and predators that increase the probability of extinction for other species. For all these reasons, the rate of extinction increases as the number of species on the island increases.

Because the rate of arrival of new species decreases and the extinction rate increases as the number of species increases, eventually the number of species on the island should reach an equilibrium at which the rates of arrival and extinction are equal (Figure 56.15a). If there are more species than the equilibrium number, extinctions should exceed arrivals, and species richness should decline. If there are fewer species than the equilibrium number, arrivals should exceed extinctions, and species richness should increase. The equilibrium is dynamic because if either rate fluctuates, as they generally do, the equilibrium number of species shifts up and down.

MacArthur and Wilson's model can also be used to predict how species richness should differ among islands of different sizes and different distances from the mainland. We expect extinction rates to be higher on small islands than on large islands because species' populations are, on average, smaller there. Similarly, we expect fewer immigrants to reach islands that are more distant from the mainland. Figure 56.15b gives hypothetical relative species richnesses for islands of different sizes and distances from the mainland. As you can see, the number of species should be highest for islands that are relatively large and relatively close to the mainland.

The MacArthur-Wilson model has been tested

Major disturbances, which serve as "natural experiments," sometimes permit colonization and extinction rates to be estimated directly. In August 1883, Krakatau, an island in the Sunda Strait between Sumatra and Java, was devastated by a series of volcanic eruptions that destroyed all life on the island's surface. After the lava cooled, Krakatau was colonized rapidly by plants and animals from Sumatra to the east and Java to the west. By 1933, the island was again covered with a tropical evergreen forest, and 271 species of plants and 27 species of resident land birds were found there.

During the 1920s, when a forest canopy was developing, there were high rates of colonization by both birds and plants (Table 56.1). Birds probably brought the seeds of many plants because, between 1908 and 1934, both the percentage (from 20% to 25%) and the absolute number (from 21 to 54) of plant species with bird-dispersed seeds increased. Today the numbers of species of plants and birds are not increasing as fast

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