Food Web Model Research Articles

Native richness and species level trophic traits predict establishment of alien freshwater fishes

Community level ecological traits are thought to affect invasibility as more diverse communities with complex trophic interactions may be associated with greater biotic resistance. Elucidation of the nature of this relationship is often hampered by difficulties in characterising food webs, particularly where field data are lacking. We attempted to overcome this by coupling food web modelling with information-theoretic analysis of the modelled webs. In addition, we also investigated the possibility that species level trends in trophic traits of established aliens might reflect exploitation of empty niches. We constructed hypothetical food webs of 26 natural and artificial lentic habitats from a data set consisting of 370 fish species representing 71 families. Using these food webs, we investigated associations at the community level between food web traits and network topology and number of alien fish species using an information-theoretic approach based on a set of competing a priori hypotheses. At the species level, we similarly tested for trends in trophic traits of established alien fishes using the information-theoretic approach in addition to nMDS of diet data. We found that native species richness in a community was the most important determinant of the number of alien fish taxa, displaying an inverse relationship. Our data also show that alien fish generally feed lower down the food web. Our findings suggest that the biotic resistance hypothesis, though scale dependent, can result in observable effects in animal communities. Moreover, we also found that the ability to exploit low energy yield food sources could favour the establishment of alien species via avoidance of resistive forces from native taxa.

Does structural sensitivity alter complexity–stability relationships?

Structural sensitivity, namely the sensitivity of a model dynamics to slight changes in its mathematical formulation, has already been studied in some models with a small number of state variables. The aim of this study is to investigate the impact of structural sensitivity in a food web model. Especially, the importance of structural sensitivity is compared to that of trophic complexity (number of species, connectance), which is known to strongly influence food web dynamics. Food web structures are built using the niche model. Then food web dynamics are modeled using several type II functional responses parameterized to fit the same predation fluxes. Food web persistence was found to be mostly determined by trophic complexity. At the opposite, even if food web connectance promotes equilibrium dynamics, their occurrence is mainly driven by the choice of the functional response. These conclusions are robust to changes in some parameter values, the fitting method and some model assumptions. In a one-prey/one-predator system, it was shown that the possibility that multiple stable states coexist can be highly structural sensitive. Quantifying this type of uncertainty at the scale of ecosystem models will be both a natural extension to this work and a challenging issue.

Top-down consumptive and trait-mediated control do affect soil food webs: It’s time for a new model

Soil food webs are complex and exhibit many of the features that are indicative of bottom-up or resource control. The perceived dominance of bottom-up control has resulted in food web models that have underexplored mechanisms of top-down or predator control. Developing soil food web models that include more mechanisms of top-down control may improve our ability to predict changes in food web configurations and ecosystem processes. Specifically, models could be improved by considering more detailed density-mediated and trait-mediated top-down effects via the inclusion of non-linear feeding relationships and context-specific variation in physiology, morphology, and behavior.

Carbon cycling in the deep eastern North Pacific benthic food web: Investigating the effect of organic carbon input

The deep ocean benthic environment plays a role in long-term carbon sequestration. Understanding carbon cycling in the deep ocean floor is critical to evaluate the impact of changing climate on the oceanic systems. Linear inverse modeling was used to quantify carbon transfer between compartments in the benthic food web at a long time-series study site in the abyssal northeastern Pacific (Station M). Linear inverse food web models were constructed for three separate years in the time-series when particulate organic carbon (POC) flux was relatively high (1990: 0.63 mean mmol C m?2 d?1), intermediate (1995: 0.24) and low (1996: 0.12). Carbon cycling in all years was dominated by the flows involved in the microbial loop; dissolved organic carbon uptake by microbes (0.80–0.95 mean mmol C m?2 d?1), microbial respiration (0.52–0.61), microbial biomass dissolution (0.09–0.18) and the dissolution of refractory detritus (0.46–0.65). Moreover, the magnitude of carbon flows involved in the microbial loop changed in relation to POC input, with a decline in contribution during the high POC influxes, such as those recently experienced at Station M. Results indicate that during high POC episodic pulses the role of faunal mediated carbon cycling would increase. Semi-labile detritus dominates benthic faunal diets and the role of labile detritus declined with increased total POC input. Linear inverse modeling represents an effective framework to analyze high-resolution time-series data and demonstrate the impact of climate change on the deep ocean carbon cycle in a coastal upwelling system.

A cautionary tale: The Goldilocks' Effect in measuring ecological complication in food webs

The purpose of this paper is to apply the Goldilocks' Effect to an experimental marine plankton community to explore the ‘just right’ level of ecological complication (EC) using loop analysis. First, the ‘just right’ level of variable aggregation (N) as well as the numbers and types of links need to be determined to then compute a minimal set of alternative descriptions of biologically reasonable food webs for a given ecosystem. A food web is biologically reasonable if it can structurally and functionally exist in nature, and encompass the main causal relationships and feedbacks that are necessary to explain how species densities change over time. An adequate food web model is more complicated that a single static representation. For example, there are several universes or sets of food webs that can be constructed. The macro universe is the set of mathematically possible networks of N variables, but relatively few are biologically reasonable. The micro universe represents all variables and links observed in a set of data fitted loop models and their combinations. It contains trillions of biologically-reasonable networks that can be summarized as Ecological Skeletons. The size of the micro universe is both an overestimate since some networks are unstable and include matrix redundancies, and it is an underestimate because there are inherent limitations on ecosystem observation in identifying all variables and links in a real world food web. The meso universe is the micro universe corrected for these over and under estimations or the ‘just right’ measure of EC. To date, its exact size remains unknown, but loop analysis provides a way to approach a realistic estimation. These results serve as a cautionary tale, however, that EC is more expansive than is commonly understood. The Goldilocks' Effect has been previously used with food webs. Loop analysis, however, contributes new insights to these discussions, and a useful methodology for identifying biologically reasonable food webs, and several associated structural and functional measures of EC.

Reticulated channels in soil food webs

Soil food web research has long recognized that real webs include a number of complexities often hidden in research focused on aboveground grazing webs. These complexities include nutrient recycling, omnivory (consuming more than one prey item) and multiple reticulated channels through which energy and resources flow. Increasing evidence in real and modeled food webs suggests a critical complexity of all webs is the presence of multi-channel feeding. Multi-channel feeding, or multi-channel omnivory, occurs when a consumer feeds across two energy channels: such as a consumer that eats both roots and detritus or bacteria and fungi. As growing empirical research has documented that multi-channel omnivores are the most common type of consumer in real webs, modeling studies have also highlighted that such omnivory can be stabilizing. Here I briefly review the evolving paradigm of multi-channel feeding in food webs and outline several major areas of research that could advance predictions of how common multi-channel feeding should be in soil food webs and its impacts on stability and trophic dynamics.

Simple assumptions predicts prey selection by piscivorous fishes

Studies on trophic interactions permits the use of community-wide network analyses to evaluate the consequences of human interventions in natural communities. In this paper, we aimed to get insights into the underlying mechanism of prey selection for four piscivorous species, and evaluate behavioral responses to prey selection after an impoundment. We assemble six food web models to search for the hypothesis that best predict observed prey selection pattern of piscivorous fishes combining the following assumptions: (i) predation window, defined as the size range of prey species consumed by a piscivorous fish; (ii) prey strategies to avoid predation (iii) and prey abundance. We tested the probability of each hypothesis to reproduce two empirical data, one before and one after an impoundment with minimum assumptions. Before impoundment, we found that predators presented switching behavior, preying preferably on abundant prey; while after impoundment, predators consumed prey within its predation window. Those results explained better than the null hypotesis and all other assumptions; and corroborate with both theoretical and empirical studies. We conclude that different assumptions drives piscivorous fish behavior in different environments; and modelling procedures can be used to assess gaps in trophic interactions of fish communities.

Eating from the same plate? Revisiting the role of labile carbon inputs in the soil food web

An increasing number of empirical studies are challenging the central fundamentals on which the classical soil food web model is built. This model assumes that bacteria consume labile substrates twice as fast as fungi, and that mycorrhizal fungi do not decompose organic matter. Here, we build on emerging evidence that points to significant consumption of labile C by fungi, and to the ability of ectomycorrhizal fungi to decompose organic matter, to show that labile C constitutes a major and presently underrated source of C for the soil food web. We use a simple model describing the dynamics of a recalcitrant and a labile C pool and their consumption by fungi and bacteria to show that fungal and bacterial populations can coexist in a stable state with large inputs into the labile C pool and a high fungal use of labile C. We propose a new conceptual model for the bottom trophic level of the soil food web, with organic C consisting of a continuous pool rather than two or three distinct pools, and saprotrophic fungi using substantial amounts of labile C. Incorporation of these concepts will increase our understanding of soil food web dynamics and functioning under changing conditions.

Lanice conchilega structures carbon flows in soft-bottom intertidal areas

Biogenic reefs constructed by the tube-building ecosystem engineer Lanice conchilega (Terrebilidae, Polychaeta) have profound structuring impacts on the benthic environment in that they alter the biogeochemical and physical properties of the sediment. This study provides new insights into the functioning and effects on food webs of L. conchilega reefs in intertidal sediments using linear inverse models to quantify the carbon flows in the food webs in the presence and absence of the tubeworm. The inverse food web models were based on an empirical dataset from 2 study sites, which provided biomass and stable isotope data, and information on general physiological constraints from the literature. Results of the model showed that the carbon input into reef food webs (mean ± SE; 191 ± 50 mmol C m -2 d -1 ) is ca. 40 times higher compared to bare sand areas (5 ± 2 mmol C m -2 d -1 ) and is mainly derived from organic matter (OM) in the water column. Most of the OM input towards these reefs is consumed by suspension-feeding macrofauna, particularly L. conchilega ; however, the worm is not an important source of carbon for other macrofaunal organisms. The ratio of OM input to primary production indicates that the OM needs to be produced in an area at least 15 times larger than the reef area, demonstrating significant OM ‘focussing’ within the reef food web. The reef structures created by L. conchilega act as a trap for OM, resulting in an overall higher macrofaunal biomass and much more diverse food webs than in the absence of the tubeworm.

The bacterial-fungal energy channel concept challenged by enormous functional versatility of soil protists

Protists (=protozoa) are commonly treated as bacterivores that control the bacterial energy channel in soil food webs. This ecologist’s perspective is, however, challenged by taxonomic studies showing that a range of protists feed on fungi, other protists and even nematodes. Recently, it was revealed that obligate and facultative mycophagous protists are common soil inhabitants, while others are facultative nematophagous. Furthermore, protists act as parasites and pathogens of plants and animals. This neglected functional diversity of protists, that is similarly prevalent for other groups of soil organisms, reveals that current food web model models are oversimplified.Facultative feeding of various protist taxa on bacteria and fungi, the source of both major energy channels, strongly implies that a clear split of the energy channels at lower trophic levels does not exist and that more complex energy flows prevail in soil food webs. Future efforts should therefore target ecological functioning of protists and other groups of soil organisms, on a species-specific level, to create more meaningful functional units that then need incorporation in modified soil food web models. Such efforts will help disentangling the structure, diversity and resulting functioning of complex soil systems, including energy flows through the soil food web.

Evolutionary food web models: effects of an additional resource

Many empirical food webs contain multiple resources, which can lead to the emergence of sub-communities—partitions—in a food web that are weakly connected with each other. These partitions interact and affect the complete food web. However, the fact that food webs can contain multiple resources is often neglected when describing food web assembly theoretically, by considering only a single resource. We present an allometric, evolutionary food web model and include two resources of different sizes. Simulations show that an additional resource can lead to the emergence of partitions, i.e. groups of species that specialise on different resources. For certain arrangements of these partitions, the interactions between them alter the food web properties. First, these interactions increase the variety of emerging network structures, since hierarchical bodysize relationships are weakened. Therefore, they could play an important role in explaining the variety of food web structures that is observed in empirical data. Second, interacting partitions can destabilise the population dynamics by introducing indirect interactions with a certain strength between predator and prey species, leading to biomass oscillations and evolutionary intermittence.

Reducing agricultural loss and food waste: how will nature fare?

Reducing agricultural loss and food waste: how will nature fare?

Developing a high taxonomic resolution food web model to assess the functional role of forage fish in the California Current ecosystem

Understanding the role of forage fish in marine food webs is an important part of ecosystem-based fisheries management. Food web models are a common tool used to account for important characteristics of forage fish and their trophodynamics. One primary limitation of many existing food web models is that the taxonomic resolution of forage fish and their predators is overly simplified. Here, we developed a food web model with high taxonomic resolution of forage fish and their predators in the California Current to more comprehensively describe trophic linkages involving forage fish and examine the ecological role of forage fish in this system. We parameterized a mass-balanced food web model (Ecopath) with 92 living functional groups, and used this to quantify diet dependency on forage fish, determine the main predators of forage fish, identify the topological position of forage fish in the food web, and calculate an index that identifies forage species or species aggregations that have key ecological roles (Supportive Role to Fishery ecosystem, SURF). Throughout, we characterized parameter uncertainty using a Monte Carlo approach. Though diets revealed some predators had high diet dependencies on individual forage fish species, most predators consumed multiple forage fish and also had notable diet overlap with forage fish. Consequently, no single forage fish appeared to act as a vital nexus species that is characteristic of “wasp-waisted” food webs in other upwelling regions. Additionally, no single forage fish was identified as “key” by the SURF index, but if predators and fisheries view certain pairs of forage fish as functionally equivalent, some plausible pairs would be identified as key assemblages. Specifically, sardine & anchovy (Sardinops sagax &Engraulis mordax) and herring & anchovy (Clupea pallasii &E. mordax) are key when predator populations depend on the aggregate availability of these species. This food web model can be used to support generalized equilibrium trade-off analysis or dynamic modeling to identify specific predators that would be of conservation concern under conditions of future forage fish depletion.

Structural sensitivity and resilience in a predator–prey model with density-dependent mortality

Numerous formulations with the same mathematical properties can be relevant to model a biological process. Different formulations can predict different model dynamics like equilibrium vs. oscillations even if they are quantitatively close (structural sensitivity). The question we address in this paper is: does the choice of a formulation affect predictions on the number of stable states? We focus on a predator–prey model with predator competition that exhibits multiple stable states. A bifurcation analysis is realized with respect to prey carrying capacity and species body mass ratio within range of values found in food web models. Bifurcation diagrams built for two type-II functional responses are different in two ways. First, the kind of stable state (equilibrium vs. oscillations) is different for 26.0–49.4% of the parameter values, depending on the parameter space investigated. Using generalized modelling, we highlight the role of functional response slope in this difference. Secondly, the number of stable states is higher with Ivlev's functional response for 0.1–14.3% of the parameter values. These two changes interact to create different model predictions if a parameter value or a state variable is altered. In these two examples of disturbance, Holling's disc equation predicts a higher system resilience. Indeed, Ivlev's functional response predicts that disturbance may trap the system into an alternative stable state that can be escaped from only by a larger alteration (hysteresis phenomena). Two questions arise from this work: (i) how much complex ecological models can be affected by this sensitivity to model formulation? and (ii) how to deal with these uncertainties in model predictions?

Intervality and coherence in complex networks.

Food webs-networks of predators and prey-have long been known to exhibit "intervality": species can generally be ordered along a single axis in such a way that the prey of any given predator tend to lie on unbroken compact intervals. Although the meaning of this axis-usually identified with a "niche" dimension-has remained a mystery, it is assumed to lie at the basis of the highly non-trivial structure of food webs. With this in mind, most trophic network modelling has for decades been based on assigning species a niche value by hand. However, we argue here that intervality should not be considered the cause but rather a consequence of food-web structure. First, analysing a set of 46 empirical food webs, we find that they also exhibit predator intervality: the predators of any given species are as likely to be contiguous as the prey are, but in a different ordering. Furthermore, this property is not exclusive of trophic networks: several networks of genes, neurons, metabolites, cellular machines, airports, and words are found to be approximately as interval as food webs. We go on to show that a simple model of food-web assembly which does not make use of a niche axis can nevertheless generate significant intervality. Therefore, the niche dimension (in the sense used for food-web modelling) could in fact be the consequence of other, more fundamental structural traits. We conclude that a new approach to food-web modelling is required for a deeper understanding of ecosystem assembly, structure, and function, and propose that certain topological features thought to be specific of food webs are in fact common to many complex networks.

Food habits of Arctic staghorn sculpin (Gymnocanthus tricuspis) and shorthorn sculpin (Myoxocephalus scorpius) in the northeastern Chukchi and western Beaufort Seas

Arctic staghorn sculpin (Gymnocanthus tricuspis) and shorthorn sculpin (Myoxocephalus scorpius) belong to Cottidae, the second most abundant fish family in the western Arctic. Although considered important in food webs, little is known about their food habits throughout this region. To address this knowledge gap, we examined and compared the diets of 515 Arctic staghorn sculpin and 422 shorthorn sculpin using stomachs collected over three summers in the northeastern Chukchi Sea (2010–2012) and one summer in the western Beaufort Sea (2011). We used permutational multivariate analysis of variance (PERMANOVA) and non-metric multidimensional scaling (nMDS) to compare sculpin diets between regions and selected size classes. Differences in mouth morphologies and predator size versus prey size relationships were examined using regression techniques. Arctic staghorn sculpin and shorthorn sculpin diet compositions differed greatly throughout the Chukchi and Beaufort Seas. Regardless of body size, the smaller-mouthed Arctic staghorn sculpin consumed mostly benthic amphipods and polychaetes, whereas the larger-mouthed shorthorn sculpin shifted from a diet composed of benthic and pelagic macroinvertebrates as smaller individuals to shrimps and fish prey as larger individuals. Within shared habitats, the sculpins appear to partition prey, either by taxa or size, in a manner that suggests no substantial overlap occurs between species. This study increases knowledge of sculpin feeding ecology in the western Arctic and offers regional, quantitative diet information that could support current and future food web modeling efforts.

Predator–prey mass ratio revisited: does preference of relative prey body size depend on individual predator size?

Summary Quantifying predator–prey body size relationships is key to understanding food webs. Food web models often assume that all individuals of predator species prefer the same relative body size of prey, using a single constant called preferred predator–prey mass ratio (preferred PPMR). In contrast, empirical studies have shown that relative prey body size in diet varies with individual predator size, challenging the food web models based on size‐invariant preferred PPMR and their predictions. We point out that this apparent inconsistency arises because empirical PPMR in those previous studies has been measured only through dietary data (i.e. realized PPMR rather than preferred PPMR) without considering the effects of environmental prey availability, suggesting the possibility that preferred PPMR may be in fact independent of individual predator size. Here, we present a new approach to revisit the assumption of size‐invariant preferred PPMR in food web models. The approach compares two measures of PPMR calculated from prey compositions in predator diet and environmental prey composition, respectively (i.e. realized PPMR vs. environmental PPMR). The deviations between realized and environmental PPMRs are considered as a proxy of individual variations in relative prey size preference (i.e. preferred PPMR). We apply this idea to long‐term dietary data of an omnivorous predatory fish species collected from a lake ecosystem over four decades. Our results showed that the preferred PPMR is independent of individual predator size when the foraging mode (i.e. the major prey type) of the predator is considered while the realized PPMR is size‐dependent regardless of the foraging mode, especially when analysed analogously to previous empirical studies. We suggest that the apparent inconsistency between theoretical assumption and empirical observation of PPMR is due to the conceptual and methodological confusion and could be resolved by distinguishing between preferred and realized PPMRs. Further, in contrast to the previous arguments based on realized PPMR, we provide the first empirical support for size‐invariant preferred PPMR. Future studies are encouraged to apply our ideas to other species/systems to test the robustness of size‐invariant preferred PPMR and to better describe food web models. A lay summary is available for this article.

Potential role of spiny dogfish in gray and harbor seal diets in the Gulf of Maine

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 550:249-270 (2016) - DOI: https://doi.org/10.3354/meps11718 Potential role of spiny dogfish in gray and harbor seal diets in the Gulf of Maine Carrie Byron*, Alexia Morgan Marine Science Center, University of New England, 11 Hills Beach Road, Biddeford, ME 04005, USA *Corresponding author: cbyron@une.edu ABSTRACT: Harbor seal Phoca vitulina and gray seal Halichoerus grypus populations have increased by 28.7% (1997-2001) and 8.6% (1999-present), respectively, in the Gulf of Maine. Here, we investigated what seals are eating and how their diet influences the food web in the Gulf of Maine. There is suggestion, but little evidence, that seals are eating spiny dogfish Squalus acanthias, currently an abundant species (standing stock biomass = 235000 t) in the Gulf of Maine and surrounding areas. We used an Ecopath food web model to explore the potential of dogfish being a primary food source for seals and how this may be impacting the ecosystem. We explored various plausible diets for seals and the implications for other non-prey species in the ecosystem. In addition, white sharks Carcharodon carcharias have increased in abundance regionally, and shark predation on seals is documented in the southern Gulf of Maine. We therefore also explored top-down impacts of white shark predation on seals and lower trophic level fish species. Our results suggest that (1) dogfish could be contributing to the recovery of seal populations by providing an alternative prey base, (2) there is a tipping point between 10 and 20% of dogfish in seal diet that elicits a linearly increasing trend in seal biomass over time, and (3) seals as prey will have a continued impact on the projected increase in white shark biomass. This work advances our understanding of top-down control and may help to inform fisheries and resource management. KEY WORDS: Food web · Ecopath with Ecosim · Phoca vitulina · Halichoerus grypus · Squalus acanthias · Cod · White shark · Carcharodon carcharias Full text in pdf format Supplementary material PreviousCite this article as: Byron C, Morgan A (2016) Potential role of spiny dogfish in gray and harbor seal diets in the Gulf of Maine. Mar Ecol Prog Ser 550:249-270. https://doi.org/10.3354/meps11718 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 550. Online publication date: May 25, 2016 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2016 Inter-Research.

Projecting changes in the distribution and productivity of living marine resources: A critical review of the suite of modelling approaches used in the large European project VECTORS

We review and compare four broad categories of spatially-explicit modelling approaches currently used to understand and project changes in the distribution and productivity of living marine resources including: 1) statistical species distribution models, 2) physiology-based, biophysical models of single life stages or the whole life cycle of species, 3) food web models, and 4) end-to-end models. Single pressures are rare and, in the future, models must be able to examine multiple factors affecting living marine resources such as interactions between: i) climate-driven changes in temperature regimes and acidification, ii) reductions in water quality due to eutrophication, iii) the introduction of alien invasive species, and/or iv) (over-)exploitation by fisheries. Statistical (correlative) approaches can be used to detect historical patterns which may not be relevant in the future. Advancing predictive capacity of changes in distribution and productivity of living marine resources requires explicit modelling of biological and physical mechanisms. New formulations are needed which (depending on the question) will need to strive for more realism in ecophysiology and behaviour of individuals, life history strategies of species, as well as trophodynamic interactions occurring at different spatial scales. Coupling existing models (e.g. physical, biological, economic) is one avenue that has proven successful. However, fundamental advancements are needed to address key issues such as the adaptive capacity of species/groups and ecosystems. The continued development of end-to-end models (e.g., physics to fish to human sectors) will be critical if we hope to assess how multiple pressures may interact to cause changes in living marine resources including the ecological and economic costs and trade-offs of different spatial management strategies. Given the strengths and weaknesses of the various types of models reviewed here, confidence in projections of changes in the distribution and productivity of living marine resources will be increased by assessing model structural uncertainty through biological ensemble modelling.

Plankton food-webs: to what extent can they be simplified?

<p class="p1">Plankton is a hugely diverse community including both unicellular and multicellular organisms, whose individual dimensions span over seven orders of magnitude. Plankton is a fundamental part of biogeochemical cycles and food-webs in aquatic systems. While knowledge has progressively accumulated at the level of single species and single trophic processes, the overwhelming biological diversity of plankton interactions is insufficiently known and a coherent and unifying trophic framework is virtually lacking. We performed an extensive review of the plankton literature to provide a compilation of data suitable for implementing food-web models including plankton trophic processes at high taxonomic resolution. We identified the components of the plankton community at the Long Term Ecological Research Station MareChiara in the Gulf of Naples. These components represented the sixty-three nodes of a plankton food-web. To each node we attributed biomass and vital rates, <em><span class="s1">i.e. </span></em>production, consumption, assimilation rates and ratio between autotrophy and heterotrophy in mixotrophic protists. Biomasses and rates values were defined for two opposite system’s conditions; relatively eutrophic and oligotrophic states. We finally identified 817 possible trophic links within the web and provided each of them with a relative weight, in order to define a diet-matrix, valid for both trophic states, which included all consumers, fromn anoflagellates to carnivorous plankton. Vital rates for plankton resulted, as expected, very wide; this strongly contrasts with the narrow ranges considered in plankton system models implemented so far. Moreover, the amount and variety of trophic links highlighted by our review is largely excluded by state-of-the-art biogeochemical and food-web models for aquatic systems. Plankton models could potentially benefit from the integration of the trophic diversity outlined in this paper: first, by using more realistic rates; second, by better defining trophic roles of consumers in the planktonic web. We suggest that most trophic habits present in planktonic organisms must be contemplated in new generation plankton models.</p>