Modelers often need to quantify the rates at which zooplankton consume a variety of species, size classes and trophic types. Implicit in the equations used to describe the multiple resource functional response (i.e. how nutritional intake varies with resource densities) are assumptions that are not often stated, let alone tested. This is problematic because models are sensitive to the details of these formulations. Here, we enable modelers to make more informed decisions by providing them with a new framework for considering zooplankton feeding on multiple resources. We define a new classification of multiple resource responses that is based on preference, selection and switching, and we develop a set of mathematical diagnostics that elucidate model assumptions. We use these tools to evaluate the assumptions and biological dynamics inherent in published multiple resource responses. These models are shown to simulate different resource preferences, implied single resource responses, changes in intake with changing resource densities, nutritional benefits of generalism, and nutritional costs of selection. Certain formulations are further shown to exhibit anomalous dynamics such as negative switching and sub-optimal feeding. Such varied responses can have vastly different ecological consequences for both zooplankton and their resources; inappropriate choices may incorrectly quantify biologicallymediated fluxes and predict spurious dynamics. We discuss how our classes and diagnostics can help constrain parameters, interpret behaviors, and identify limitations to a formulation's applicability for both regional (e.g. HighNitrate-Low-Chlorophyll regions comprising large areas of the Pacific) and large-scale applications (e.g. global biogeochemical or climate change models). Strategies for assessing uncertainty and for using the mathematics to guide future experimental investigations are also discussed. r

Johnson, C. L., Leising, A. W., Runge, J. A., Head, E. J. H., Pepin, P., Plourde, S., and Durbin, E. G. 2008. Characteristics of Calanus finmarchicus dormancy patterns in the Northwest Atlantic. – ICES Journal of Marine Science, 65: 339–350. Demographic time-series from four fixed stations in the Northwest Atlantic Ocean demonstrate variable timing of entry into and emergence from dormancy in subpopulations of the planktonic copepod Calanus finmarchicus. A proxy for timing of entry was established as the date each year when the proportion of the fifth copepodid stage (CV) in the subpopulation rose to half its overall climatological maximum CV proportion at that station. The proxy for timing of emergence at each station was set as the first date when adults were more than 10% of the total abundance of copepodid stages. An alternate emergence proxy date was determined by back-calculating the spawning dates of the first early copepodid stages appearing in spring, using a stage-structured, individual-based model. No single environmental cue (photoperiod, surface temperature, or average surface-layer chlorophyll a concentration) consistently explained entry or emergence dates across all stations. Among hypotheses put forward to explain dormancy in Calanus species, we cannot eliminate the lipid accumulation window hypothesis for onset of dormancy or a lipid-modulated endogenous timer controlling dormancy duration. The fundamental premise of these hypotheses is that individuals can only enter dormancy if their food and temperature history allows them to accumulate sufficient lipid to endure overwintering, moult, and undergo early stages of gonad maturation.

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