Bullfrogs: Introduced predators in southwestern wetlands

Abstract

In the American Southwest, much of the native fish fauna is facing extinction (Minckley and Deacon 1991); frogs in California (Fellers and Drost 1993) and frogs and garter snakes in Arizona (Schwalbe and Rosen 1988) are also in critical decline. Habitat destruction and introduced predators appear to be primary causes of native frog declines (Jennings and Hayes 1994), and habitat modification often yields ponds and lakes especially suitable for introduced species. Introduced bullfrogs ( Rana catesbeiana ) have been blamed for amphibian declines in much of western North America (e.g., Hayes and Jennings 1986; Leonard et al. 1993; Vial and Saylor 1993). Extensive cannibalism by bullfrogs renders them especially potent predators at the population level. The tadpoles require only perennial water and grazeable plant material; hence, transforming young can sustain a dense adult bullfrog population even if alternate prey are depleted. This may increase the probability that native species may be extirpated by bullfrog predation. Introduced predatory fishes are apparently an important cause of frog declines (Hayes and Jennings 1986). They have been strongly implicated in one important case of decline of native ranid frog (family Ranidae, the "true" frogs; Bradford 1989). Some introduced crayfish may also be devastating in some areas (Jennings and Hayes 1994). In our study region, however, neither introduced fishes nor crayfish are dominant. We present results that sustain a "bullfrog hypothesis" for some native ranid declines, and we present our study as an example of how evidence accumulates to support such a hypothesis. In 1985 we began documenting historical localities for wetland herpetofaunas (reptiles and amphibians), based on museum records and personal interviews, then revisited these and additional areas to determine current species' status. Results of this process, plus circumstantial evidence, suggested that the bullfrog was a primary cause for declines of leopard frogs and garter snakes in southern Arizona (Schwalbe and Rosen 1988). In 1986-89 and 1992-93 we conducted removal censuses of bullfrogs at San Bernardino National Wildlife Refuge (SBNWR), Cochise County, Arizona. We simultaneously monitored native Chiricahua leopard frogs ( R. chiricahuensis ) and Mexican garter snakes ( Thamnophis eques ) at the sites of bullfrog removal. A control site, with no bullfrog removal, was established in comparable habitat at Buenos Aires National Wildlife Refuge (BANWR), Pima County, Arizona.