Ecology Letters, (2006) 9: 51–60 doi: 10.1111/j.1461-0248.2005.00845.x LETTER Quantifying the extinction vortex
Abstract William F. Fagan1* and E. E. We developed a database of 10 wild vertebrate populations whose declines to extinction Holmes2 were monitored over at least 12 years. We quantitatively characterized the final declines 1Department of Biology, of these well-monitored populations and tested key theoretical predictions about the University of Maryland, College process of extinction, obtaining two primary results. First, we found evidence of Park, MD 20906, USA logarithmic scaling of time-to-extinction as a function of population size for each of the 2 Cumulative Risk Initiative, 10 populations. Second, two lines of evidence suggested that these extinction-bound Northwest Fisheries Science populations collectively exhibited dynamics akin to those theoretically proposed to occur Center, National Marine in extinction vortices. Specifically, retrospective analyses suggested that a population size Fisheries Service, 2725 Montlake Blvd E, Seattle, WA 98112, USA of n individuals within a decade of extinction was somehow less valuable to persistence *Correspondence: E-mail: than the same population size was earlier. Likewise, both year-to-year rates of decline [email protected] and year-to-year variability increased as the time-to-extinction decreased. Together, these results provide key empirical insights into extinction dynamics, an important topic that has received extensive theoretical attention.
Keywords Decline to extinction, extinction risk, extinction vortex, population extinction, time-to- extinction, vertebrate extinctions.
Ecology Letters (2006) 9: 51–60
tions have declined in numbers or spatial distribution (e.g. INTRODUCTION Kunin & Gaston 1997, Akc¸akaya 2004), quantitative data on In coining the term Ôextinction vortexÕ, Gilpin & Soule´ the dynamics leading up to actual extinction events in (1986) argued that as populations decline, an insidious vertebrate populations remain rare. Instead, almost all mutual reinforcement can occur among biotic and abiotic quantitative analyses of population-level extinction processes such as environmental stochasticity, demographic dynamics for vertebrates are either wholly theoretical or stochasticity, inbreeding, and behavioural failures, driving tied to populations that may have declined but have not population size downward to extinction. In the two decades gone extinct (e.g. Tuljapurkar 1990; Dennis et al. 1991; since that work, extinction-related issues have continued to Ludwig 1999; Brook et al. 2000; Ripa & Lundberg 2000; motivate research by ecologists and conservation biologists, Holmes & Fagan 2002; Lande et al. 2003). In those few leading to the development of population viability analyses cases where ecologists have actually witnessed extinction and related approaches where the aim is to characterize the events in vertebrate populations, available data tend to risk of extinction that particular populations face (Morris & afford only short or highly patchy time series, neither of Doak 2003). In contrast to the large body of research, which permits detailed quantitative investigation of a especially by theoreticians, that has focused on risks of population’s dynamics as it declines to extinction. extinction, far fewer studies have investigated the dynamics To address the critical need for a multispecies summary of actual extinction events. Studies by Belovsky et al. (1999) of the dynamics by which well-monitored vertebrate on laboratory populations and by Schoener et al. (2003) on populations decline to extinction, we combed published island invertebrates are among the rare exceptions that have and unpublished sources to develop a small database with provided much needed empirical perspectives on the which we could quantitatively characterize the final declines dynamics of extinction. of wild vertebrate populations. We were also interested in For vertebrate populations, ecologistsÕ understanding of using the collected time series to test two primary extinction dynamics is far less clear. Indeed, even though theoretical predictions about the process of extinction. ecologists can cite a plethora of empirical examples of The first prediction was that, for populations that go extinct, situations in which species with previously robust popula- population lifetimes should scale logarithmically with