Zooplankton and forage fish species off Peru: Large-scale bottom-up forcing and local-scale depletion

The Humboldt Current System, like all upwelling systems, has dramatic quantities of plankton-feeding fish, which suggested that their population dynamics may ‘drive’ or ‘control’ ecosystem dynamics. With this in mind we analysed the relationship between forage fish populations and their main prey, zooplankton populations. Our study combined a zooplankton sampling program (1961–2005) with simultaneous acoustic observations on fish from 40 pelagic surveys (1983–2005) conducted by the Peruvian Marine Research Institute (IMARPE) and landing statistics for anchoveta (Engraulis ringens) and sardine (Sardinops sagax) along the Peruvian coast from 1961 to 2005. The multi-year trend of anchoveta population abundance varied consistently with zooplankton biovolume trend, suggesting bottom-up control on anchovy at the population scale (since oceanographic conditions and phytoplankton production support the changes in zooplankton abundance). For a finer-scale analysis (km) we statistically modelled zooplankton biovolume as a function of geographical (latitude and distance from the 200-m isobath), environmental (sea surface temperature), temporal (year, month and time-of-day) and biological (acoustic anchovy and sardine biomass within 5 km of each zooplankton sample) covariates over all survey using both classification and regression trees (CART) and generalized additive models (GAM). CART showed local anchoveta density to have the strongest effect on zooplankton biovolume, with significantly reduced levels of biovolume for higher neighbourhood anchoveta biomass. Additionally, zooplankton biovolume was higher offshore than on the shelf. GAM results corroborated the CART findings, also showing a clear diel effect on zooplankton biovolume, probably due to diel migration or daytime net avoidance. Apparently, the observed multi-year population scale bottom-up control is not inconsistent with local depletion of zooplankton when anchoveta are locally abundant, since the latter effect was observed over a wide range of overall anchoveta abundance.