A basal aquatic-terrestrial trophic link in rivers: Algal subsidies via shore-dwelling grasshoppers

Rivers provide important resources for riparian consumers, especially in arid or seasonally arid biomes. Pygmy grasshoppers (Paratettix aztecus and P. mexicanus; Tetrigidae) graze river algae stranded along shorelines of the South Fork Eel River in northern California (39°44′N, 123°39′W) as the river recedes during the summer drought. Densities of tetrigids during the mid to late summer were highest (1 individual/m² in July) within 1 m of the river margin, and declined to near zero at 4 m from the margin, especially during peak temperatures in the afternoon. These observations suggested that the distribution of tetrigids was determined by the availability of algae, water, or both. We manipulated the presence/absence of water and beached algae (Cladophora glomerata) in a 2x2 factorial design. All treatments were positioned 2 m upslope from the river's edge (about 30 cm above the water table), where the cobble bar was naturally dry and devoid of algae and densities of tetrigids were lower than at the river margin (0.4 individuals/m² in July). Tetrigids responded only to the wet Cladophora treatment, which had 30x higher densities than other treatments. Stable isotopic signatures (δ¹³C) of tetrigids (-19.7‰) collected from the same cobble bars were more similar to those of epilithic algae (-20.4‰) than terrestrial plants (-28.2‰), and higher than those of acridid grasshoppers (-27.9‰) from the same habitats. Mixing models suggest that 88-100% of the C in tetrigid grasshoppers at our study site is derived from riverine algae. A preliminary analysis suggests that tetrigids ingested sufficient quantities of algae to easily meet their energetic demands during the summer. This study supports the idea that algae, produced in stream systems, can determine the distribution and relative abundance of a common terrestrial scavenger and provide an additional pathway for energy exchange between rivers and riparian food webs.