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Conservation biologists have long been concerned that species extinction could have significant consequences for the stability of entire ecosystems—groups of interacting organisms and the physical environment that they inhabit. An ecosystem could survive the loss of some species, but if enough species were lost, the ecosystem would be severely degraded. In fact, it is possible that the loss of a single important species could start a cascade of extinctions that might dramatically change an entire ecosystem. A good illustration of this occurred after sea otters were eliminated from some Pacific kelp (seaweed) bed ecosystems: the kelp beds were practically obliterated too because in the absence of sea otter predation, sea urchin populations exploded and consumed most of the kelp and other macroalgae.

It is usually claimed that species-rich ecosystems tend to be more stable than species-poor ecosystems. Three mechanisms by which higher diversity increases ecosystem stability have been proposed. First, if there are more species in an ecosystem, then its food web will be more complex, with greater redundancy among species in terms of their nutritional roles. In other words, in a rich system if a species is lost, there is a good chance that other species will take over its function as prey, predator, producer, decomposer, or whatever role it played. Second, diverse ecosystems may be less likely to be invaded by new species, notably exotics (foreign species living outside their native range), that would disrupt the ecosystem’s structure and function. Third, in a species-rich ecosystem, diseases may spread more slowly because most species will be relatively less abundant, thus increasing the average distance between individuals of the same species and hampering disease transmission among individuals.

Scientific evidence to illuminate these ideas has been slow in coming, and many shadows remain. [■] One of the first studies to provide data supporting a relationship between diversity and stability examined how grassland plants responded to a drought. [■] Researchers D. Tilman and J A. Downing used the ratio of above-ground biomass in 1988 (after two years of drought) to that in 1986 (predrought) in 207 plots in a grassland field in the Cedar Creek Natural History Area in Minnesota as an index of ecosystem response to disruption by drought. [■]In an experiment that began in 1982, they compared these values with the number of plant species in each plot and discovered that the plots with a greater number of plant species experienced a less dramatic reduction in biomass. [■]Plots with more than ten species had about half as much biomass in 1988 as in 1986, whereas those with fewer than five species only produced roughly one-eighth as much biomass after the two-year drought. Apparently, species-rich plots were likely to contain some drought-resistant plant species that grew better in drought years, compensating for the poor growth of less-tolerant species.

To put this result in more general terms, a species-rich ecosystem may be more stable because it is more likely to have species with a wide array of responses to variable conditions such as droughts. Furthermore, a species-rich ecosystem is more likely to have species with similar ecological functions, so that if a species is lost from an ecosystem, another species, probably a competitor, is likely to flourish and occupy its functional role. Both of these, variability in responses and functional redundancy, could be thought of as insurance against disturbances.

The Minnesota grassland research has been widely accepted as strong evidence for the diversity- stability theory; however, its findings have been questioned, and similar studies on other ecosystems have not always found a positive relationship between diversity and stability. Clearly, this is a complex issue that requires further field research with a broad spectrum of ecosystems and species: grassland plants and computer models will only take us so far. In the end, despite insightful attempts to detect some general patterns, we may find it very difficult to reduce this topic to a simple, universal truth.

OFFICIAL46 Look at the four squares [■] that indicate where the following sentence could be added to the passage. Where would the sentence best fit? Click on a square [■] to add the sentence to the passage.

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