Planktonic copepods are primary consumers in the ocean and are perhaps the most numerous metazoans on earth. Secondary production by these zooplankton supports most food webs of the open sea, directly affecting pelagic fish populations and the biological pump of carbon into the deep ocean. Models of marine ecosystems are quite sensitive to the formulation of the term for zooplankton mortality, although there are few data available to constrain mortality rates in such models. Here we present the first evidence for nonlinear, density-dependent mortality rates of open-ocean zooplankton. A high-frequency time series reveals that per capita mortality rates of eggs of Calanus finmarchicus Gunnerus are a function of the abundance of adult females and juveniles. The temporal dynamics of zooplankton populations can be influenced as much by time-dependent mortality rates as by variations in 'bottom up' forcing. The functional form and rates chosen for zooplankton mortality in ecosystem models can alter the balance of pelagic ecosystems, modify elemental fluxes into the ocean's interior, and modulate interannual variability in pelagic ecosystems.

We compare the patterns of stage-specific mortality of Calanus finmarchicus at five localities across the North Atlantic Ocean during the spring-summer period of active population growth: Georges Bank, a continental shelf locality in the NW Atlantic, based on 30 broadscale survey cruises in the US GLOBEC program; the northern North Sea, studied during the historic FLEX program with sampling four times daily for 73 days; Ocean Station M in the central Norwegian Sea, based on an 80-day daily time-series; and Lurefjorden (sampled weekly in late winter-early summer) and Sørfjorden (sampled monthly), two fjords in southwestern Norway characterized by markedly different guilds of predators. The mortality estimation methods included Wood's Population Surface Method, the Vertical Life Table (VLT) method, and a modified VLT, according to the study site and copepod recruitment schedules. Contrary to assumptions implicit in many simulation models and indirect methods for estimating zooplankton mortality, both rates and stage-specific patterns of mortality of C. finmarchicus vary appreciably across the North Atlantic. Characteristics of local environments, including the predator field in particular, appear to strongly influence mortality schedules in different regions. In at least two sites (Georges Bank and Ocean Station M), mortality rates of early stages of C. finmarchicus are density-dependent. We attribute this density-dependent mortality to egg cannibalism, which introduces non-linear population responses to changing environmental conditions. Region-specific biological interactions can substantially modify the effects of physical climate variability and render simple linear relationships between climate and zooplankton abundance unlikely. © 2004 International Council for the Exploration of the Sea. Published by Elsevier Ltd. All rights reserved.