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

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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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A site-specific, coupled biologicaVphysica1 model is described for the dominant winterspring copepod in the Gulf of Maine. The biological portion describes temperature-and fooddependent progression through 13 life stages in an Eulerian (concentration-based) framework. The population is transported in a flow field depicting the climatological mean conditions in 2 mo 'seasons'. Behavioural assumptions account for depth selection and 2 limiting cases are contrasted: dispersal throughout the water colun~n, and aggregation in the surface layer. Simulations are inspired by MARMAP observat~ons, with an emphasis on the mid-winter initiation of the annual bloom by diapausing populations, and their role in supplying reproducing populations to Georges Bank during spring. Passive tracer sirnulations illustrate the role of the circulation. Georges Bank itself is an open system and depends on resupply from external sources. A l l 3 deep basins of the Gulf are capable of contributing populations to the Bank. The Scotian Shelf is capable of populating the Southern Flank. In the case where the organisms aggregate in the surface layer, the effect of convergence in downwelling zones is shown to be a significant contributor to population &stribution. Baseline population dynamics are initiated on January 1 by activating a diapausing population (Go) based on 10 yr mean abundance and distribution from the MARMAP program. The abundance and distribution of Go adults is reproduced with a 3-layer model, spatially vanable mortality, an extended period of activation, no food limitation, and a large, heretofore unobserved source of diapausing C5s near-bottom. Reproduction modeled in this system shows significant development of generation 1 (G,) over the Gulf of Maine in February-March, which is not observed. Delay of reproduction over the Gulf, and/or severe early-stage mortality, is required to conform with the data. The space-time pattern of this effect is consistent with the observed chlorophyll distribution and hming, and reasonable food-limitation thresholds. Inclusion of this effect initiates the spring Calanus bloom in the correct space-time pattern, with significant cohorts of Go females and G, nauplii over important cod and haddock spawning grounds on Georges Bank. The implication is that G, is locally spawned in food-rich waters over Georges Bank by females advected in the food-poor Gulf of Maine surface layer.

Gentleman, W. C., Neuheimer, A. B., and Campbell, R. G. 2008. Modelling copepod development: current limitations and a new realistic approach. – ICES Journal of Marine Science, 65: 399–413. To predict the influence of environmental variability on copepod dynamics and production, models must account for the effects of temperature and food on stage-dependent time-scales. Here, data for development-time means and variance of Calanus finmarchicus are used to quantify the limitations of existing models. Weight-based individual models are sensitive to uncertain parameters, such as moulting weights, assimilation efficiency, and environmental dependencies, making them highly difficult to calibrate. The accuracy of stage-based population models using ordinary differential equations depends on model structure, with some predicted generation times being incorrect by months. Even when large numbers of age classes are used to reduce modelled variability, it is not possible to make variability consistent with the data. Accuracy of mean times for stage-based population models using difference equations requires a small time-step, which results in large numbers of age classes and modelled variability that is underestimated by orders of magnitude, unless a probabilistic moult fraction is used. We present a new stage-based individual model that avoids the limitations of other models and successfully represents C. finmarchicus mean development timing and associated variability. This approach can be adapted easily for other species, as well as dynamic environmental conditions.

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