Progress in Oceanography Zooplankton and the oceanography of the eastern tropical Pacific : A review q Marı́a

We review the spatial and temporal patterns of zooplankton in the eastern tropical Pacific Ocean and relationships with oceanographic factors that affect zooplankton distribution, abundance and trophic relationships. Large-scale spatial patterns of some zooplankton groups show broad coincidence with surface water masses, circulation, and upwelling regions, in agreement with an ecological and dynamic partitioning of the pelagic ecosystem. The papers reviewed and a new compilation of zooplankton volume data at large-scale show that abundance patterns of zooplankton biomass have their highest values in the upwelling regions, including the Gulf of Tehuantepec, the Costa Rica Dome, the equatorial cold tongue, and the coast of Peru. Some of the first studies of zooplankton vertical distribution were done in this region, and a general review of the topic is presented. The possible physiological implications of vertical migration in zooplankton and the main hypotheses are described, with remarks on the importance of the oxygen minimum zone (OMZ) as a barrier to both the vertical distribution and migration of zooplankton in the region. Recent results, using multiple-net gear, show that vertical distribution is more complex than previously thought. There are some well-adapted species that do live and migrate within the OMZ. Temporal patterns are reviewed and summarized with historical data. Seasonal variations in zooplankton biomass follow productivity cycles in upwelling areas. No zooplankton time series exist to resolve ENSO effects in oceanic regions, but some El Niño events have had effects in the Peru Current ecosystem. Multidecadal periods of up to 50 years show a shift from a warm sardine regime with a low zooplankton biomass to a cool anchovy regime in the eastern Pacific with higher zooplankton biomasses. However, zooplankton volume off Peru has remained at low values since the 1972 El Niño, a trend opposite to that of anchoveta biomass since 1984. Studies of trophic relations emphasize the difference in the productivity cycle in the eastern tropical Pacific compared to temperate or polar ecosystems, with no particular peaks in the stocks of either zooplankton or phytoplankton. Productivity is more dependent on local events like coastal upwelling or water circulation, especially in the equatorial countercurrent and around the equatorial cool-tongue. Micrograzers are very important in the tropics as are predatory mesozooplankton. 0079-6611/$ see front matter 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.pocean. 2006.03. 003 q The order of authorship was determined by the flip of a coin. Sections 1–2.2 by M.A.F. and Sections 2.3–4.4 by J.F. * Corresponding author. E-mail addresses: mafa@hp.fciencias. unam.mx (M.A. Fernández-Álamo), jfarber@cicese.mx (J. Färber-Lorda). M.A. Fernández-Álamo, J. Färber-Lorda / Progress in Oceanography 69 (2006) 318–359 319 Up to 70% of the daily primary productivity is consumed by microzooplankton, which thus regulates the phytoplankton stocks. Micrograzers are an important link between primary producers, including bacteria, and mesozooplankton, constituting up to 80% of mesozooplankton food. Oceanography affects zooplankton trophic relationships through spatial–temporal effects on primary productivity and on the distributions of metabolic factors, food organisms, and predators. This paper is part of a comprehensive review of the oceanography of the eastern tropical Pacific. 2006 Elsevier Ltd. All rights reserved.