Placing biodiversity in ecosystem models without getting lost in translation

A key challenge to progressing our understanding of biodiversity’s role in the sustenance of ecosystem function is the extrapolation of the results of two decades of dedicated empirical research to regional, global and future landscapes. Ecosystem models provide a platform for this progression, potentially offering a holistic view of ecosystems where, guided by the mechanistic understanding of processes and their connection to the environment and biota, large-scale questions can be investigated. While the benefits of depicting biodiversity in such models are widely recognized, its application is limited by difficulties in the transfer of knowledge from small process oriented ecology into macro-scale modelling. Here, we build on previous work, breaking down key challenges of that knowledge transfer into a tangible framework, highlighting successful strategies that both modelling and ecology communities have developed to better interact with one another. We use a benthic and a pelagic case-study to illustrate how aspects of the links between biodiversity and ecosystem process have been depicted in marine ecosystem models (ERSEM and MIRO), from data, to conceptualisation and model development. We hope that this framework may help future interactions between biodiversity researchers and model developers by highlighting concrete solutions to common problems, and in this way contribute to the advance of the mechanistic understanding of the role of biodiversity in marine (and terrestrial) ecosystems.

Mechanisms shaping size structure and functional diversity of phytoplankton communities in the ocean

The factors regulating phytoplankton community composition play a crucial role in structuring aquatic food webs. However, consensus is still lacking about the mechanisms underlying the observed biogeographical differences in cell size composition of phytoplankton communities. Here we use a trait-based model to disentangle these mechanisms in two contrasting regions of the Atlantic Ocean. In our model, the phytoplankton community can self-assemble based on a trade-off emerging from relationships between cell size and (1) nutrient uptake, (2) zooplankton grazing, and (3) phytoplankton sinking. Grazing 'pushes' the community towards larger cell sizes, whereas nutrient uptake and sinking 'pull' the community towards smaller cell sizes. We find that the stable environmental conditions of the tropics strongly balance these forces leading to persistently small cell sizes and reduced size diversity. In contrast, the seasonality of the temperate region causes the community to regularly reorganize via shifts in species composition and to exhibit, on average, bigger cell sizes and higher size diversity than in the tropics. Our results raise the importance of environmental variability as a key structuring mechanism of plankton communities in the ocean and call for a reassessment of the current understanding of phytoplankton diversity patterns across latitudinal gradients.

A general framework for aquatic biogeochemical models

One of the most of challenging steps in the development of coupled hydrodynamic-biogeochemical models is the combination of multiple, often incompatible computer codes that describe individual physical, chemical, biological and geological processes. This “coupling” is time-consuming, error-prone, and demanding in terms of scientific and programming expertise. The open source, Fortran-based Framework for Aquatic Biogeochemical Models addresses these problems by providing a consistent set of programming interfaces through which hydrodynamic and biogeochemical models communicate. Models are coded once to connect to FABM, after which arbitrary combinations of hydrodynamic and biogeochemical models can be made. Thus, a biogeochemical model code works unmodified within models of a chemostat, a vertically structured water column, and a three-dimensional basin. Moreover, complex biogeochemistry can be distributed over many compact, self-contained modules, coupled at run-time. By enabling distributed development and user-controlled coupling of biogeochemical models, FABM enables optimal use of the expertise of scientists, programmers and end-users.

Trait-based representation of diatom functional diversity in a plankton functional type model of the eutrophied southern North Sea

We introduce a trait-based description of diatom functional diversity to an existing plankton functional type (PFT) model, implemented for the eutrophied coastal ecosystem in the Southern Bight of the North Sea. The trait-based description represents a continuum of diatom species, each characterized by a distinct cell volume, and includes size dependence of four diatom traits: the maximum growth rate, the half-saturation constants for nutrient uptake, the photosynthetic efficiency, and the relative affinity of copepods for diatoms. Through competition under seasonally varying forcing, the fitness of each diatom varies throughout time, and the outcome of competition results in a changing community structure. The predicted seasonal change in mean cell volume of the community is supported by field observations: smaller diatoms, which are more competitive in terms of resource acquisition, prevail during the first spring bloom, whereas the summer bloom is dominated by larger species which better resist grazing. The size-based model is used to determine the ecological niche of diatoms in the area and identifies a range of viable sizes that matches observations. The general trade-off between small, competitive diatoms and large, grazing-resistant species is a convenient framework to study patterns in diatom functional diversity. PFT models and trait-based approaches constitute promising complementary tools to study community structure in marine ecosystems.

ERSEM 15.06: a generic model for marine biogeochemistry and the ecosystem dynamics of the lower trophic levels

The ERSEM model is one of the most established ecosystem models for the lower trophic levels of the marine food-web in the scientific literature. Since its original development in the early nineties it has evolved significantly from a coastal ecosystem model for the North-Sea to a generic tool for ecosystem simulations from shelf seas to the global ocean. The current model release contains all essential elements for the pelagic and benthic part of the marine ecosystem, including the microbial food-web, the carbonate system and calcification. Its distribution is accompanied by a testing framework enabling the analysis of individual parts of the model. Here we provide a detailed mathematical description of all ERSEM components along with case-studies of mesocosm type simulations, water column implementations and a brief example of a full-scale application for the North-West European shelf. Validation against in situ data demonstrates the capability of the model to represent the marine ecosystem in contrasting environments.

ERSEM 15.06: a generic model for marine biogeochemistry and the ecosystem dynamics of the lower trophic levels

The European Regional Seas Ecosystem Model (ERSEM) is one of the most established ecosystem models for the lower trophic levels of the marine food web in the scientific literature. Since its original development in the early nineties it has evolved significantly from a coastal ecosystem model for the North Sea to a generic tool for ecosystem simulations from shelf seas to the global ocean. The current model release contains all essential elements for the pelagic and benthic parts of the marine ecosystem, including the microbial food web, the carbonate system, and calcification. Its distribution is accompanied by a testing framework enabling the analysis of individual parts of the model. Here we provide a detailed mathematical description of all ERSEM components along with case studies of mesocosm-type simulations, water column implementations, and a brief example of a full-scale application for the north-western European shelf. Validation against in situ data demonstrates the capability of the model to represent the marine ecosystem in contrasting environments.