Stable isotope values and trophic analysis of invasive three-spined stickleback in Upper Lake Constance points to significant piscivory

The three-spined stickleback Gasterosteus aculeatus was introduced into Lake Constance in the 1940s and occupied a limited range until late 2012. Since then the species has expanded from a solely littoral habitat in Upper Lake Constance, but now makes seasonal migrations into the pelagic zone. This behavioral change has been accompanied by a drastic increase in stickleback abundance. In order to integrate information about feeding of sticklebacks in Upper Lake Constance over two consecutive years, stomach content analysis was combined with seasonal stable isotope analysis on two types of tissue (muscle and liver). Isotope values were also obtained for zooplankton, whitefish larvae and eggs. We calculated the contribution of potential food sources for sticklebacks' diet using a Bayesian mixing model (SIMMR). Furthermore, we determined stickleback trophic position, and delta N-15 and delta C-13 values were compared with those of other fish species of Lake Constance. The results of the Bayesian model as well as the stomach content analysis showed clear evidence of stickleback predation on fish eggs and larvae. Stickleback delta N-15 values were elevated during winter and comparable to those of piscivorous pike, while delta N-15 values of zooplankton were reduced, and those of whitefish larvae were similar to those of sticklebacks after accounting trophic fractionation of N isotopes. Trophic position calculations further identified sticklebacks as piscivorous, while the delta C-13 values of the liver and stomach content analysis suggests that a benthic-pelagic species pair may exist in Lake Constance. These findings support the hypotheses that sticklebacks in Lake Constance can display piscivorous feeding behaviour on sympatric fish species, most likely whitefish larvae and eggs.