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
An individual‐based life history and population dynamic model for the winter–spring dominant copepod of the subarctic North Atlantic, Calanus finmarchicus, is coupled with a regional model of advection for the Gulf of Maine and Georges Bank. Large numbers of vectors, each representing individual copepods with elements for age, stage, ovarian status and other population dynamic variables, are carried in a computation through hourly time steps. Each vector is updated at each time step according to development rate and reproductive functions derived from experimental data. Newly spawned eggs are each assigned new vectors as needed. All vectors are subject to random mortality. Thus, both life history progression and population dynamics of C. finmarchicus are represented for the temperatures in the Gulf of Maine–Georges Bank region in the active season. All vectors include elements representing depth, latitude and longitude. This allows coupling of the population dynamics to the tide‐ and wind‐driven Dartmouth model of New England regional circulation. Summary data from the physical model are used to advance vectors from resting‐stock locations in Gulf of Maine basins through two generations to sites of readiness for return to rest. Supply of Calanus stock to Georges Bank comes from all of the gulf and from the Scotian Shelf. The top of the bank is stocked from western gulf basins; the North‐east Peak is stocked from Georges Basin and the Scotian Shelf. All sources contribute to stock that accumulates in the SCOPEX gyre off the north‐west shoulder of Georges Bank, explaining the high abundance recurrently seen in that region. There is some return of resting stock to Wilkinson Basin in the western gulf, but other basins must mostly be restocked from upstream sources to the north‐east.
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