Extinction Rates in North American Freshwater Fishes, 1900–2010 Author(S): Noel M

Extinction Rates in North American Freshwater Fishes, 1900–2010 Author(s): Noel M. Burkhead Source: BioScience, 62(9):798-808. 2012. Published By: American Institute of Biological Sciences URL: http://www.bioone.org/doi/full/10.1525/bio.2012.62.9.5 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Articles Extinction Rates in North American Freshwater Fishes, 1900–2010 NOEL M. BURKHEAD Widespread evidence shows that the modern rates of extinction in many plants and animals exceed background rates in the fossil record. In the present article, I investigate this issue with regard to North American freshwater fishes. From 1898 to 2006, 57 taxa became extinct, and three distinct populations were extirpated from the continent. Since 1989, the numbers of extinct North American fishes have increased by 25%. From the end of the nineteenth century to the present, modern extinctions varied by decade but significantly increased after 1950 (post-1950s mean = 7.5 extinct taxa per decade). In the twentieth century, freshwater fishes had the highest extinction rate worldwide among vertebrates. The modern extinction rate for North American freshwater fishes is conservatively estimated to be 877 times greater than the background extinction rate for freshwater fishes (one extinction every 3 million years). Reasonable estimates project that future increases in extinctions will range from 53 to 86 species by 2050. Keywords: North America, freshwater fishes, extinction rates, E/MSY, aquatic biodiversity nderstanding the underlying causes of the extinction of become extinct, and three unique populations have been Umodern organisms is, in most cases, relatively simple. extirpated (Miller et al. 1989, Jelks et al. 2008). Extinction is the antithesis of speciation and a natural outcome of evolutionary change (Darwin 1859). Under natural conditions, including catastrophic events, extinction “Do not tell fish stories where the people know you; is one of nature’s best ideas: It enables life to adapt to ever- but particularly, don’t tell them where they know the fish.” changing environments. A fundamental concern among —Mark Twain. biologists is that contemporary rates of extinction due to human activities are orders of magnitude greater than background rates evidenced in the fossil record, and these rates Declining biodiversity and increasing rates of extinction appear to be increasing (May et al. 1995, Pimm et al. 1995, are fundamentally important metrics of natural resource 2006). Catastrophic events that caused the five well-known status and are the subjects of intensive investigation. In mass extinctions, including the cataclysmic extermination of particular, the mitigation and prevention of biodiversity loss the dinosaurs, are not responsible for the majority of extinc- are among the overarching goals of conservation biology tions in geological history. Rather, 90%–96% of all extinct (Helfman 2007, Pereira et al. 2010). The threat of extinc- species variably disappeared during the normal give and take tion may be greatest in freshwater ecosystems, where the of speciation and extinction over geological time scales (May proportion of threatened and endangered species is gener- et al. 1995). In phylogenetic theory, ancestral species may ally greater than that in terrestrial ecosystems (Pimm et al. become extinct in the evolutionary transition to new species, 1995, Strayer and Dudgeon 2010) and where imperilment or conversely, derived taxa may split from the ancestral taxa levels are similar to those of tropical rain forests (Ricciardi and remain extant. Both modes are theoretically plausible and Rasmussen 1999). Inferences that the rates of contem- under allopatric speciation scenarios (Wiley and Mayden porary extinction will increase are based on large numbers 1985). In the present study, I am not concerned with meth- of organisms considered to be critically endangered world- odological intricacies; rather, I focus on the human-caused wide (IUCN 2011), the increasing negative effects of human extinction of North American freshwater fishes from the activities on the Earth’s biosphere (Vitousek et al. 1997, close of the nineteenth century to the present. In the context Naiman and Turner 2000, McKee et al. 2004, Davies et al. of human life spans, observations of extinction should be 2006), and the conclusion that such activities will result extraordinarily rare. Nonetheless, since 1900, at least 57 spe- in higher near-future extinction rates (Pereira et al. 2010, cies and subspecies of North American freshwater fishes have Barnosky et al. 2011). BioScience 62: 798–808. ISSN 0006-3568, electronic ISSN 1525-3244. © 2012 by American Institute of Biological Sciences. All rights reserved. Request permission to photocopy or reproduce article content at the University of California Press’s Rights and Permissions Web site at www.ucpressjournals.com/ reprintinfo.asp. doi:10.1525/bio.2012.62.9.5 "IO3CIENCEsSeptember 2012 / Vol. 62 No. 9 www.biosciencemag.org Articles North American freshwater fishes: A brief synopsis and the filling of numerous basins gouged by the giant North America is geographically defined as Canada, the ice sheets (e.g., the Great Lakes; Hocutt and Wiley 1986). United States, and Mexico, including the coastal islands off The evolution and biogeography of the North American Canada and Alaska but excluding Hawaii and the southern fish fauna is intimately linked to transitional changes to Mexican states of Quintana Roo and Campeche (Jelks et al. landforms and their influences on continental watercourses 2008). The diversity of fishes is equivalent to that of all other over time (Mayden 1988, Smith GR et al. 2002). Generally, vertebrate groups combined (Nelson 2006); therefore, the species richness per drainage (or ecoregion) increases from number of descriptions of fishes new to science exceeds that Canada southward, but over two-thirds of species richness of descriptions of new tetrapods each year. At the close of occurs east of the Great Continental Divide, especially in 2010, the cutoff point for this study, 31,769 valid fish species the Interior and Appalachian Highlands (Hocutt and Wiley had been described worldwide (William N. Eschmeyer, 1986, Mayden 1992). Fishes in the western United States and California Academy of Sciences [retired], personal com- Mexico exhibit relatively lower species richness per drainage munication, 28 August 2011). About 43% of the world’s unit but generally have higher endemism and dispropor- fishes are freshwater or diadromous species (Nelson 2006). tionate numbers of extinct species (Minckley and Douglas Comparatively, the freshwater fish diversity of North America 1991, Burr and Mayden 1992). is less than those of Africa (about 2945 species), Asia (3533), and South America (4035), but higher than those of Europe Rates of faunal documentation and evidence of (330), Russia (206), and Oceania (260) (Lévêque et al. 2008). extinction North America has the most diverse nontropical freshwater Our knowledge of North American fishes is facilitated by fish fauna in the world (Lundberg et al. 2000). At the end of a long tradition of faunal study and public access to most 2010, the freshwater fish fauna of North America consisted river and lake systems. From 1758 to 2010, the average of 53 families, 214 genera, and 1213 species, or about 8.9% rate of fish descriptions for North America was 4.8 species of the Earth’s freshwater fish diversity (Nelson et al. 2004). described per year, but as is evident in figure 1a, the rates Continental fishes primarily consist of obligate freshwater varied over time and logically increased as more biologists species, diadromous fishes (species in which part of their life studied the fauna. From 1970 through 2010, species docu- cycle occurs in the marine realm), and marine fishes that mentation averaged 6.7 species described per year, which is occasionally or regularly enter freshwater during their lives. considered the modern rate of faunal discovery and docu- The tally through 2010 is based on independent counts of mentation (figure 1a). The discovery and description of species described after 2004, aided by comments from Larry North American freshwater fishes continues unabated, and Page (University of Florida Museum of Natural History, unlike birds (see Pimm and colleagues’ [2006] figure 1), personal communication, 14 October 2010). there is no evidence of an asymptote in the rates of species descriptions. Patterns of species richness Modern extinctions of North American fishes were first Until recently, most species of the modern North American observed at the dawn of the twentieth century, and the trend fish fauna were considered

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