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Human-dominated marine ecosystems are experiencing accelerating loss of populations and species, with largely unknown consequences. We analyzed local experiments, long-term regional time series, and global fisheries data to test how biodiversity loss affects marine ecosystem services across temporal and spatial scales. Overall, rates of resource collapse increased and recovery potential, stability, and water quality decreased exponentially with declining diversity. Restoration of biodiversity, in contrast, increased productivity fourfold and decreased variability by 21%, on average. We conclude that marine biodiversity loss is increasingly impairing the ocean's capacity to provide food, maintain water quality, and recover from perturbations. Yet available data suggest that at this point, these trends are still reversible.

The invasion paradox describes the co-occurrence of independent lines of support for both a negative and a positive relationship between native biodiversity and the invasions of exotic species. The paradox leaves the implications of native-exotic species richness relationships open to debate: Are rich native communities more or less susceptible to invasion by exotic species? We reviewed the considerable observational, experimental, and theoretical evidence describing the paradox and sought generalizations concerning where and why the paradox occurs, its implications for community ecology and assembly processes, and its relevance for restoration, management, and policy associated with species invasions. The crux of the paradox concerns positive associations between native and exotic species richness at broad spatial scales, and negative associations at fine scales, especially in experiments in which diversity was directly manipulated. We identified eight processes that can generate either negative or positive native-exotic richness relationships, but none can generate both. As all eight processes have been shown to be important in some systems, a simple general theory of the paradox, and thus of the relationship between diversity and invasibility, is probably unrealistic. Nonetheless, we outline several key issues that help resolve the paradox, discuss the difficult juxtaposition of experimental and observational data (which often ask subtly different questions), and identify important themes for additional study. We conclude that natively rich ecosystems are likely to be hotspots for exotic species, but that reduction of local species richness can further accelerate the invasion of these and other vulnerable habitats.

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Motivated by recent global reductions in biodiversity, empirical and theoretical research suggests that more species-rich systems exhibit enhanced productivity, nutrient cycling, or resistance to disturbance or invasion relative to systems with fewer species. In contrast, few data are available to assess the potential ecosystemlevel importance of genetic diversity within species known to play a major functional role. Using a manipulative field experiment, we show that increasing genotypic diversity in a habitat-forming species (the seagrass Zostera marina) enhances community resistance to disturbance by grazing geese. The time required for recovery to near predisturbance densities also decreases with increasing eelgrass genotypic diversity. However, there is no effect of diversity on resilience, measured as the rate of shoot recovery after the disturbance, suggesting that more rapid recovery in diverse plots is due solely to differences in disturbance resistance. Genotypic diversity did not affect ecosystem processes in the absence of disturbance. Thus, our results suggest that genetic diversity, like species diversity, may be most important for enhancing the consistency and reliability of ecosystems by providing biological insurance against environmental change.

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The spread of exotic species and climate change are among the most serious global environmental threats. Each independently causes considerable ecological damage, yet few data are available to assess whether changing climate might facilitate invasions by favoring introduced over native species. Here, we compare our long-term record of weekly sessile marine invertebrate recruitment with interannual variation in water temperature to assess the likely effect of climate change on the success and spread of introduced species. For the three most abundant introduced species of ascidian (sea squirt), the timing of the initiation of recruitment was strongly negatively correlated with winter water temperature, indicating that invaders arrived earlier in the season in years with warmer winters. Total recruitment of introduced species during the following summer also was positively correlated with winter water temperature. In contrast, the magnitude of native ascidian recruitment was negatively correlated with winter temperature (more recruitment in colder years) and the timing of native recruitment was unaffected. In manipulative laboratory experiments, two introduced compound ascidians grew faster than a native species, but only at temperatures near the maximum observed in summer. These data suggest that the greatest effects of climate change on biotic communities may be due to changing maximum and minimum temperatures rather than annual means. By giving introduced species an earlier start, and increasing the magnitude of their growth and recruitment relative to natives, global warming may facilitate a shift to dominance by nonnative species, accelerating the homogenization of the global biota. P redicted increases in global temperature (1) likely will have dramatic effects on the structure and function of ecosystems worldwide (2-4). One approach to predicting the biological effects of climate change has been to examine the direct physiological consequences to resident species of changes in temperature or CO 2 concentration (5-8). Other studies have focused on the indirect consequences of altered temperature on interactions among resident species and the cascading effects on local community composition (9-12). Climatic warming on the time scale of decades also may alter the composition of the resident biota by facilitating the poleward spread of species characteristic of warmer temperature regimes (9, 13-15). However, it is also possible that climate change could facilitate quantum leaps in the range of species across ocean basins or continents. Humans already inadvertently transport countless species around the globe (16), and, although many of these inoculations presumably fail because of inhospitable climate in the recipient region (17, 18), global warming may relax this constraint. Despite considerable interest in predicting the spread and success of ''invasive'' species because of their significant effect on native communities (19), there are few data to evaluate the effects of climate change on this type of invasion...

Theory predicts that systems that are more diverse should be more resistant to exotic species, but experimental tests are needed to verify this. In experimental communities of sessile marine invertebrates, increased species richness significantly decreased invasion success, apparently because species-rich communities more completely and efficiently used available space, the limiting resource in this system. Declining biodiversity thus facilitates invasion in this system, potentially accelerating the loss of biodiversity and the homogenization of the world's biota.