Does diversity equal stability? Are diverse ecosystems the most stable ecosystems?

Animal and plant species are disappearing from the Earths diverse ecosystems at an increasingly alarming rate. However, apart from obvious ethical and aesthetic motivations can we justify concern, in fact obsession, with the loss of individual species? What is the ecological value of biodiversity? It was this question which first sparked the stability-diversity hypothesis, or rather debate, between scientists. Intuition tells us that biological diversity in an ecosystem would confer stability, and this was the consensus among early ecologists (Elton, 1958; MacArthur, 1955). However, theoretical studies in the 1970s would contradict these inferences (May, 1973). In this paper I review past and recent studies into this debate. Since the concept of an ecosystem was first coined scientists hypothesised that a positive linear correlation would exist between species diversity and ecosystem stability. This general belief was based on a variety of observations but not experimentally tested. Charles Eaton was a major advocate of this view, arguing that ecosystems with an abundance of species, representing a complex food web, demonstrated better resilience to invasions and flocculation in populations (Elton, 1958). He observed that cultivated land which had simplified ecological communities was more vulnerable to species invasions than similar areas that hadnt had human intervention. He also observed populations on small islands and found that they were more susceptible to invasion than are those on continents. Theoretical studies in the early 1970s challenged this motion, concluding that more complex ecosystems are less likely to be stable than simpler ones. Notably, in 1973 Robert May published a theoretical study in which randomly constructed communities were analysed (May, 1973). To allow the mathematical processes to be precise May assumed that population numbers of each species were at equilibrium. In complex ecosystems, however, populations all experience at least some degree of flux and may therefore depend on this for functionality (McCann, 2000). It has been found in subsequent studies that high biodiversity decreases the stability of each species population, thus supporting theoretical results. However, the positive relationship between biodiversity and the stability of the entire ecosystem productivity supports the earlier hypotheses (Tillman, 1996a). Recent investigations have sought to resolve this debate. Some of the strongest evidence for the hypothesis is provided by David Tilman and his colleagues (Tillman, 1996b). In 1982 Tilman divided grasslands into more than 200 plots and monitored the species richness and ecosystem productivity in each field over two decades. He was able to delineate experimentally that plant cover and thus stability is an increasing function of species richness and lower concentration of inorganic soil nitrogen due presumably to greater nitrogen uptake in more diverse communities. Another study on coral reef colonies across the Philippines produced a similar response. Laurie Raymondo and his team investigated the effects of overfishing (diversity reduction) on coral health, studying the disease status of thousands of coral colonies. Their results demonstrated a clear link between functionally diverse, species rich communities and low disease prevalence (Raymondo, 2009). A meta-analysis conducted by Bradley Cardinale of 111 experiments over more than a decade manages to add yet another dimension to the debate, a question of number of species vs. functionality of species (Cardinale, 2006). The results of his research suggest that whilst species loss does indeed affect the stability of numerous ecosystems, it is the identity of the species going extinct that affects the magnitude. Logic would tell you that certain species are more important to ecosystem function than others. As a consequence, the addition or deletion of a single species could, depending on their functionality, result in system collapse or have no overall impact. Species redundancy, several species performing similar

functions, is one possible explanation. If this were true of an ecosystem, when one species is lost from an ecosystem, another of similar function may increase in abundance and thus compensate for the lost species. Therefore simple biological systems could retain stability despite loss of biodiversity. One of the most recent theoretical propositions argues that the stability-diversity relationship is statistically inevitable (Doak, 1998). On the basis of simple probability theory, stability will always increase with species diversity because of statistical averaging of flocculation. In other words, the more diverse the ecosystem, the fewer things (biotic and abiotic interactions) are left out of it, thus the fewer the possible outside influences. This results in greater dynamic stability. Thus, stability is (almost) a necessary consequence of diversity (Tillman, 1998). Although the reasoning detailed above may seem sound, it is flawed in that it only considers diversity (number of species) when in fact numerous features of ecosystems affect stability. For instance: the strength of interactions among species (affecting food web relations), the resilience of functional species to environmental stresses and even abiotic factors (i.e. soil chemistry) (Ives, 2007). Diversity is easier to measure and manipulate, but investigating this factor alone may lead to misleading conclusions about the value of biodiversity.

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