Assessing the direct and indirect effects of bigheaded carp (<i>Hypophthalmichthys</i> spp.) on freshwater food webs: A meta‐analysis

Abstract The rapid growth of aquaculture over the last century has led to the widespread introduction of aquatic species to freshwater environments. Bighead carp (Cyprinidae: Hypophthalmichthys nobilis ) and silver carp (Cyprinidae: Hypophthalmichthys molitrix ), collectively referred to as bigheaded carp, are among the top finfishes produced in aquaculture. These filter feeders possess feeding structures that allow them to efficiently remove plankton and other particles from the water. Unfortunately, when introduced into the wild and outside their historical range, their feeding traits generally result in the exploitation and suppression of phytoplankton and zooplankton, reducing food resources for native invertebrates and vertebrates. We conducted a meta‐analysis of published experiments to examine how bigheaded carp directly and indirectly affect organisms in aquatic food webs to better understand their ecological role within ecosystems. Specifically, we addressed the following questions: (1) What is the overall magnitude and direction of bigheaded carp effects on different aquatic organisms? (2) Do bigheaded carp affect organisms in planktonic and benthic habitats? (3) Are effects similar between bighead carp and silver carp? (4) Are effects consistent among experimental arenas (i.e., mesocosm, ponds)? We detected direct and indirect effects of bigheaded carp across multiple trophic levels. Large zooplankton often exhibited the strongest negative effects, with taxa‐specific differences between carp species. The suppression of zooplankton cascaded down to primary producers, resulting in an increase in phytoplankton (i.e., indirect effect). However, we observed no compensatory increases in rotifers when large zooplankton were reduced. Growth rates of planktivorous native fishes were the slowest when stocked with bigheaded carp. Our results further indicated that egested waste from bigheaded carp appeared to subsidise (i.e., direct effect) some benthic invertebrates such as Chironomidae midges. Effects of carp on zooplankton were generally similar between mesocosms and ponds. Invasive bigheaded carp greatly restructure their surrounding food web, both positively and negatively, through a complex suite of direct and indirect effects. These changes manifested across multiple trophic levels and differing habitats. Our analysis provides a more complete understanding of how these invasive carps change their surroundings by exploiting food resources and transferring materials from planktonic to benthic habitats. Taken together, the suite of direct and indirect effects shows how an invasive species can create imbalances within aquatic ecosystems, often to the detriment of native taxa. Bigheaded carp are invasive in many countries across the globe because of their popularity in aquaculture. Our findings reaffirm the threat these planktivores pose to native freshwater organisms.