River–watershed Exchange: Effects of Riverine Subsidies on Riparian Lizards and Their Terrestrial Prey

Resource subsidies from external habitats can enhance the performance or population density of local consumers, altering their effects on in situ prey. Indirect effects of subsidies may be either positive or negative depending on the behavior of the shared consumer. Here we document strong links between riverine insects, riparian lizards (Sceloporus occidentalis), and terrestrial invertebrates. We hypothesized that aquatic insects subsidize riparian lizard populations leading to higher growth rates of these lizards in nearriver habitats, and that subsidies exert short-term positive effects on terrestrial resources as a result of diet shifts by lizards to aquatic insects. To test these hypotheses, we used 2 m high fences, or ‘‘subsidy shields,’’ to experimentally reduce aquatic insect flux to large (91 m2) enclosures of lizards. Subsidy shields reduced aquatic insect flux by 55–65%. Growth rates of lizards were 73 higher in subsidized (no-shield) enclosures during the early summer but were not significantly different later in the summer, when ambient fluxes of aquatic insects dropped to 20% of their early season levels. Within the watershed, lizard growth rates (in mass) were positively correlated with the numerical abundance of aquatic insects. Thus, lizard growth rates tracked both seasonal and spatial availability of riverine insect subsidies during our experiment. Subsidies also had indirect effects on the grounddwelling, terrestrial prey of lizards. Declines of diurnal terrestrial invertebrates were significantly higher in shield than no-shield enclosures, and the most common ground spider (Arctosa sp. [Lycosidae]) disappeared completely from shield enclosures by the end of the experiment. Declines in terrestrial invertebrate abundance did not differ between no-shield enclosures and lizard exclosures. These data suggest that riverine insects subsidize riparian Sceloporus and, in the short term, reduce their predation on terrestrial arthropods.