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of large fauna on most continents of the world (Barnosky, Chapter 12, this volume). Over the past few thousand years, human colonists and their commensals (such as the Polynesian rat) have contributed to the extinction of thousands of bird species across oceanic islands of the world (Steadman, 2006). Over the past 500 years, humans have reduced the amount of natural habitat worldwide, directly exploited species, introduced exotic species and exotic pathogens, and created many other conditions conducive to species extinction. The total number of recent extinctions is unknown, because many species have likely gone extinct before ever being recorded by science (Wilson, 1992). Estimates of global species loss vary, but based on rates of tropical deforestation and the species–area relationship a fairly typical estimate is 27,000 species lost per year; this is based on a species–area relationship with a z value of 0.15 and an estimate of 10 million species globally (Wilson, 1992). Even with a more conservative estimate of 5 million species on the planet (Primack, 2006), this would still equate to ≈13,500 species lost, or at least committed to extinction, per year. In sharp contrast to such estimates, the number of documented extinctions during the past 500 years is much lower; the International Union for Conservation of Nature, as of November 2007, reports 785 extinctions worldwide. Many other extinctions, not included in this number, have likely occurred, but they have not yet been documented adequately enough to be listed as extinct by the International Union for Conservation of Nature. Consequently, although species listed as extinct by the International Union for Conservation of Nature are likely an underestimate of the total number of extinctions over the past 500 years, those listed provide the most detailed evidence on extinction available.

There has been recent disagreement in the literature about how best to interpret extinction data from the International Union for Conservation of Nature, particularly with regards to the role species invasions play in causing extinctions (Gurevitch and Padilla, 2004; Ricciardi, 2004; Clavero and Garcia-Berthou, 2005). This disagreement has largely been due to the difficulty in ascribing precise causes to species extinctions. The precise mechanisms for any individual extinction are difficult to confidently determine for two reasons. First, extinctions are often caused by multiple factors, such as species invasions, habitat destruction, human exploitation, pollution, and infectious disease (KF Smith et al., 2006). Second, most “documented” extinctions actually involve some speculation about the factors responsible (because few species have been carefully monitored from the point of initial population decline to the point of final extinction). Additionally, it is worth noting that disagreement over species concepts, and disagreement over phylogenetic classifications of individual species, although not an issue for most extinct species, is an important point of debate in some cases. Given these limitations, it seems most appropriate



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