On Islands

Predation by exotic species has caused the extinction of many native animal species on islands, whereas competition from exotic plants has caused few native plant extinctions. Exotic plant addition to islands is highly nonrandom, with an almost perfect 1 to 1 match between the number of naturalized and native plant species on oceanic islands. Here, we evaluate several alternative implications of these findings. Does the consistency of increase in plant richness across islands imply that a saturation point in species richness has been reached? If not, should we expect total plant richness to continue to increase as new species are added? Finally, is the rarity of native plant extinctions to date a misleading measure of the impact of past invasions, one that hides an extinction debt that will be paid in the future? By analyzing historical records, we show that the number of naturalized plant species has increased linearly over time on many individual islands. Further, the mean ratio of naturalized to native plant species across islands has changed steadily for nearly two centuries. These patterns suggest that many more species will become naturalized on islands in the future. We also discuss how dynamics of invasion bear upon alternative saturation scenarios and the implications these scenarios have for the future

*Brown University, Department of Ecology and Evolutionary Biology, Box G-W, Providence, RI 02912; ^Department of Ecology, Evolution and Marine Biology and ^Marine Science Institute, University of California, Santa Barbara, CA 93106.

86 / Dov F. Sax and Steven D. Gaines retention or extinction of native plant species. Finally, we identify invasion-motivated research gaps (propagule pressure, time-lags to extinction, abundance shifts, and loss of area) that can aid in forecasting extinction and in developing a more comprehensive theory of species extinctions.

Species invasions have contributed to the extinction of many species worldwide (Clavero and Garcia-Berthou, 2005), particularly on islands (Sax et al., 2002; Steadman, 2006). These extinctions have occurred disproportionately among taxonomic groups. For example, birds have lost many species, both in absolute terms and relative to their total number of species, whereas plants have lost few species (James, 1995; Sax et al., 2002; Blackburn et al., 2004). The lack of recorded extinctions in plants does not appear to be due to a lack of knowledge, particularly on well-studied islands like New Zealand, where both historic and fossil records suggest that few native plant species have been lost (Sax et al., 2002). Why so few plant species have been lost is somewhat of a mystery, particularly considering the thousands of exotic plant species that have been introduced to islands (Sax et al., 2002). This near-lack of plant extinctions to date raises the obvious question of whether this trend will continue. In particular, should we expect few additional plant species to go extinct—even on individual islands where hundreds or thousands of exotic plant species have invaded? Should ongoing and future invasions eventually lead to mass extinction events among island plant taxa?

Our ability to accurately forecast future extinction events is limited by shortcomings in current ecological and evolutionary theory. In particular, we have a relatively poor understanding of the processes that ultimately limit how many species can inhabit any given place or area. One important concept, however, that arises from several prominent theories of species diversity is that for any given set of environmental conditions there is a ''saturation point'' that bounds the number of species a place or region can support (MacArthur and Wilson, 1963, 1967; Hubbell, 2001; Tilman, 2004). There are two basic ways that a saturation point for species richness could operate. First, the total number of species present in an area could be maintained as a balance between extinction and colonization. At the saturation point, the addition of new colonizing species results in the local extinction of a like number of previously established species. We refer to this as ''extinction-based saturation,'' which is consistent with Island Bio-geography Theory (IBT) (MacArthur and Wilson, 1963, 1967). Second, the total number of species in a place could be maintained by suppression of colonization. A saturation point is reached if the invasion of new species is inhibited by species already present. We refer to this as ''colonization-based saturation,'' which is consistent with the resistance to invasion in niche-based models of species diversity, such as Stochastic Niche Theory (SNT), where the total number of species in a place is determined by how finely resources can be partitioned among species (Tilman, 2004).

These alternative pathways for species saturation have very different implications for the impact of invading species on native plant extinctions. Extinction-based saturation (like IBT) explicitly predicts species turnover, i.e., change in species composition, with the addition of species to islands (MacArthur and Wilson, 1963, 1967). Therefore, once a saturation point has been reached, the continued addition of exotic species should result in the extinction of native species, and the rate of extinction increases as the rate of colonization increases. If this holds true for islands, then we may eventually have dramatic increases in the number of native plant extinctions—particularly if exotics continue to be added in large numbers. In contrast, colonization-based saturation (like SNT) explicitly predicts that the addition of exotic species will become progressively less likely to occur as species richness increases (Stachowicz and Tilman, 2005). Colonization-based saturation (like SNT) also predicts that existing species should benefit from a priority effect that makes them unlikely to be displaced by incoming species—as long as invading species are more or less equivalent to native species in their efficiency in using resources (Stachowicz and Tilman, 2005). If this holds true on islands, then we should expect the rate of naturalization of exotic species to diminish dramatically and few currently established species (both native and exotic) to be at risk of extinction in the future. These are two very distinct views of the future, and distinguishing among these and other alternatives is important. Doing so will help us to advance our basic understanding of ecological and evolutionary theory while simultaneously advancing our understanding of a pressing applied issue, namely the future of plant diversity on islands worldwide.

Here, we evaluate the role of species invasions on the extinction of native species. We begin by examining the International Union for Conservation of Nature database for patterns of extinction in plants and terrestrial vertebrates that have occurred worldwide over the past 500 years. We then focus on patterns of invasion and extinction on islands over the past few hundred years. Next, we consider, with a focus on plants, whether islands are saturated with respect to the total number of species they can support. We also consider what the implications of species saturation are for future plant extinctions. Finally, we explore several research gaps that currently limit our ability to forecast species extinctions.

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