Predator-prey Mass Ratios Research Articles

Zooplankton Are Not Fish: Improving Zooplankton Realism in Size-Spectrum Models Mediates Energy Transfer in Food Webs

The evidence for an equal distribution of biomass from bacteria to whales has led to development of size-spectrum models that represent the dynamics of the marine ecosystem using size rather than species identity. Recent advances have improved the realism of the fish component of the size-spectrum, but these often assume that small fish feed on an aggregated plankton size-spectrum, without any explicit representation of zooplankton dynamics. In these models, small zooplankton are grouped with phytoplankton as a resource for larval fish, and large zooplankton are parameterized as small fish. Here we investigate the impact of resolving zooplankton and their feeding traits in a dynamic size-spectrum model. First, we compare a base model, where zooplankton are parameterized as smaller fish, to a model that includes zooplankton-specific feeding parameters. Second, we evaluate how the parameterization of zooplankton feeding characteristics, specifically the predator-prey mass ratio (PPMR), assimilation efficiency and feeding kernel width, affects the productivity and stability of the fish community. Finally, we compare how feeding characteristics of different zooplankton functional groups mediate increases in primary production and fishing pressure. Incorporating zooplankton-specific feeding parameters increased productivity of the fish community, but also changed the dynamics of the entire system from a stable to an oscillating steady-state. The inclusion of zooplankton feeding characteristics mediated a trade-off between the productivity and resilience of the fish community, and its stability. Fish communities with increased productivity and lower stability were supported by zooplankton with a larger PPMR and a narrower feeding kernel – specialized herbivores. In contrast, fish communities that were stable had lower productivity, and were supported by zooplankton with a lower PPMR and a wider feeding kernel – generalist carnivores. Herbivorous zooplankton communities were more efficient at mediating increases in primary production, and supported fish communities more resilient to fishing. Our results illustrate that zooplankton are not just a static food source for larger organisms, nor can they be resolved as very small fish. The unique feeding characteristics of zooplankton have enormous implications for the dynamics of marine ecosystems, and their representation is of critical importance in size-spectrum models, and end-to-end ecosystem models more broadly.

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.

Trophic positions and predator–prey mass ratio of the pelagic food web in the East China Sea and Sea of Japan

Size-based food webs analysis is essential for understanding food web structure and evaluating the effects of human exploitation on food webs. We estimated the predator–prey mass ratio (PPMR) of the pelagic food web in the East China Sea and Sea of Japan by using the relationships between body mass and trophic position. Trophic position was calculated by additive and scaled models based on nitrogen stable isotope ratios (δ15N). The PPMRs based on additive and scaled models were 5032 (95% confidence interval (CI) 2066–15506) and 3430 (95% CI 1463–10083) respectively. The comparatively high PPMRs could reflect low ecosystem transfer efficiency and high metabolic rate.

Nitrogen and carbon stable isotope variation in northeast Atlantic fishes and squids

Nitrogen and carbon stable isotope data are frequently used to describe the origins and transformations of organic matter. Nitrogen stable isotopes (δ15N) in tissue are used to estimate species' trophic levels, the extent of omnivory, food chain length, and community‐wide relationships between body size and trophic level; the latter leading to estimates of predator–prey mass ratios for parameterization, calibration, and validation of food web models. Carbon stable isotopes (δ13C) are used to identify pathways linking producers and consumers and for studies of migration and movement. Collectively, δ15N and δ13C, often with other stable isotopes such as δ34S, may be used to define the contribution of different producers and pathways to consumer production, to assess the trophic impacts of invasive species and habitat modification, and to predict past habitat use, movements, and migrations. Stable isotope data often complement dietary data (e.g., from stomach contents) in food web studies, because stable isotope composition is indicative of assimilated diet over months to years, depending on species, size, environment, and tissue type. There are relatively few large‐scale compilations of δ15N and δ13C data for marine species from offshore habitats, but such data facilitate comparative analysis and research into food web structure and function. The data provided comprise 5535 records for individuals of 62 species of fish and squid weighing 0.3 g to 17 920 g and sampled from the northeast Atlantic shelf seas (Celtic Sea, North Sea, Irish Sea, Channel) from 2002 to 2010. For every sampled individual the record lists: species name, date of sampling, position of sampling, body mass, percentage nitrogen in muscle tissue, percentage carbon in muscle tissue, and δ15N and δ13C natural abundance in muscle tissue. Awareness of, and access to, these data should catalyze and facilitate new research with stable isotopes, to improve understanding of marine biology, food web ecology, and human impacts on the environment.

A coupled stable isotope-size spectrum approach to understanding pelagic food-web dynamics: A case study from the southwest sub-tropical Pacific

This study investigated the food web structure of the oligotrophic picophytoplankton-dominated pelagic ecosystem in the vicinity of New Caledonia, within the Archipelagic Deep Basin (ARCH) province of the southwest sub-tropical Pacific. Nitrogen stable isotope (δ15N) data were collected for mesozooplankton (0.2–2mm), macrozooplankton (2–20mm), micronekton (20–200mm) and nekton (>200mm) during 2002–2004 and 2011. Using a coupled δ15N size-spectrum approach, we estimated (1) organism trophic level (TL); (2) food chain length (FCL); (3) predator prey mass ratio (PPMR); and (4) transfer efficiency (TE). The role of phytoplankton size structure in determining these parameters was investigated. Applying a trophic enrichment factor (TEF) of 3.4, maximum TL was calculated at ~5. The number of TLs spanned by each length class was 1.97 for mesozooplankton, 2.07 for macrozooplankton, 2.75 for micronekton, and 2.21 for nekton. Estimated PPMR was 10,099:1 for mesozooplankton, 3683:1 for macrozooplankton/micronekton, and 2.44×105:1 for nekton, corresponding to TEs of 6.3%, 8.5% and 2.4%, respectively. PPMR and TE were strongly influenced by the TEF used, and TEF 3.4 likely over and underestimated PPMR and TE, respectively, for mesozooplankton and macrozooplankton/micronekton. Comparatively low PPMR for mesozooplankton and macrozooplankton/micronekton indicated longer food chains and higher connectivity within these groups than for the nekton. Conversely, the high PPMR yet high trophic niche width for the nekton indicated that they prey primarily on macrozooplankton/micronekton, with a relatively high degree of dietary specialisation. Our results are discussed in the context of other marine food webs. The ARCH food chain was found to be 1–1.5 trophic levels longer than the eutrophic micro-/nanophytoplankton-dominated Californian upwelling system, providing empirical support for the role of phytoplankton size in determining FCL. Group specific PPMR estimates demonstrated that it is changes in trophic pathways across the mesozooplankton/macrozooplankton/micronekton groups that are primarily responsible for higher FCL under oligotrophic conditions. Finally, we discuss consistently low δ15N values to the east of New Caledonia, and implications for the contribution of diazotroph nitrogen to the pelagic food web in this region.

Coexistence of structured populations with size-based prey selection

Species with a large adult–offspring size ratio and a preferred predator–prey mass ratio undergo ontogenetic trophic niche shift(s) throughout life. Trophic interactions between such species vary throughout life, resulting in different species-level interaction motifs depending on the maximum adult sizes and population size distributions. We explore the assembly and potential for coexistence of small communities where all species experience ontogenetic trophic niche shifts. The life-history of each species is described by a physiologically structured model and species identity is characterised by the trait: size at maturation. We show that a single species can exist in two different states: a ‘resource driven state’ and a ‘cannibalistic state’ with a large scope for emergent Allee effects and bistable states. Two species can coexist in two different configurations: in a ‘competitive coexistence’ state when the ratio between sizes at maturation of the two species is less than a predator–prey mass ratio and the resource level is low to intermediate, or in a ‘trophic ladder’ state if the ratio of sizes at maturation is larger than the predator–prey mass ratio at all resource levels. While there is a large scope for coexistence of two species, the scope for coexistence of three species is limited and we conclude that further trait differentiation is required for coexistence of more species-rich size-structured communities.

Predator–prey body size relationships when predators can consume prey larger than themselves

As predator-prey interactions are inherently size-dependent, predator and prey body sizes are key to understanding their feeding relationships. To describe predator-prey size relationships (PPSRs) when predators can consume prey larger than themselves, we conducted field observations targeting three aquatic hemipteran bugs, and assessed their body masses and those of their prey for each hunting event. The data revealed that their PPSR varied with predator size and species identity, although the use of the averaged sizes masked these effects. Specifically, two predators had slightly decreased predator-prey mass ratios (PPMRs) during growth, whereas the other predator specialized on particular sizes of prey, thereby showing a clear positive size-PPMR relationship. We discussed how these patterns could be different from fish predators swallowing smaller prey whole.

The Serengeti food web: empirical quantification and analysis of topological changes under increasing human impact

1. To address effects of land use and human overexploitation on wildlife populations, it is essential to better understand how human activities have changed species composition, diversity and functioning. Theoretical studies modelled how network properties change under human-induced, non-random species loss. However, we lack data on realistic species-loss sequences in threatened, real-world food webs to parameterize these models. 2. Here, we present a first size-structured topological food web of one of the most pristine terrestrial ecosystems in the world, the Serengeti ecosystem (Tanzania). The food web consists of 95 grouped nodes and includes both invertebrates and vertebrates ranging from body masses between 10(-7) and 10(4) kg. 3. We study the topological changes in this food web that result from the simulated IUCN-based species-loss sequence representing current species vulnerability to human disturbances in and around this savanna ecosystem. We then compare this realistic extinction scenario with other extinction sequences based on body size and connectance and perform an analysis of robustness of this savanna food web. 4. We demonstrate that real-world species loss in this case starts with the biggest (mega) herbivores and top predators, causing higher predator-prey mass ratios. However, unlike theoretically modelled linear species deletion sequences, this causes poor-connected species to be lost first, while more highly connected species become lost as human impact progresses. This food web shows high robustness to decreasing body size and increasing connectance deletion sequences compared with a high sensitivity to the decreasing connectance deletion scenario. 5. Furthermore, based on the current knowledge of the Serengeti ecosystem, we discuss how the focus on food web topology alone, disregarding nontrophic interactions, may lead to an underestimation of human impacts on wildlife communities, with the number of trophic links affected by a factor of two. 6. This study underlines the importance of integrative efforts between the development of food web theory and basic field work approaches in the quantification of the structure of interaction networks to sustain natural ecosystems in a changing world.

Food web framework for size-structured populations

We synthesise traditional unstructured food webs, allometric body size scaling, trait-based modelling, and physiologically structured modelling to provide a novel and ecologically relevant tool for size-structured food webs. The framework allows food web models to include ontogenetic growth and life-history omnivory at the individual level by resolving the population structure of each species as a size-spectrum. Each species is characterised by the trait ‘size at maturation’, and all model parameters are made species independent through scaling with individual body size and size at maturation. Parameter values are determined from cross-species analysis of fish communities as life-history omnivory is widespread in aquatic systems, but may be reparameterised for other systems. An ensemble of food webs is generated and the resulting communities are analysed at four levels of organisation: community level, species level, trait level, and individual level. The model may be solved analytically by assuming that the community spectrum follows a power law. The analytical solution provides a baseline expectation of the results of complex food web simulations, and agrees well with the predictions of the full model on biomass distribution as a function of individual size, biomass distribution as a function of size at maturation, and relation between predator–prey mass ratio of preferred and eaten food. The full model additionally predicts the diversity distribution as a function of size at maturation.

Global patterns in predator–prey size relationships reveal size dependency of trophic transfer efficiency

Predator-prey body size relationships influence food chain length, trophic structure, transfer efficiency, interaction strength, and the bioaccumulation of contaminants. Improved quantification of these relationships and their response to the environment is needed to parameterize food web models and describe food web structure and function. A compiled data set comprising 29582 records of individual prey eaten at 21 locations by individual predators that spanned 10 orders of magnitude in mass and lived in marine environments ranging from the poles to the tropics was used to investigate the influence of predator size and environment on predator and prey size relationships. Linear mixed effects models demonstrated that predator-prey mass ratios (PPMR) increased with predator mass. The amount of the increase varied among locations and predator species and individuals but was not significantly influenced by temperature, latitude, depth, or primary production. Increases in PPMR with predator mass implied nonlinear relationships between log body mass and trophic level and reductions in transfer efficiency with increasing body size. The results suggest that very general rules determine dominant trends in PPMR in diverse marine ecosystems, leading to the ubiquity of size-based trophic structuring and the consistency of observed relationships between the relative abundance of individuals and their body size.

Abundance–body mass relationships in size‐structured food webs

AbstractIn communities sharing a common energy source, the energetic equivalence hypothesis predicts that numerical abundance (N) scales with body mass (M) asM−0.75. However, in size‐structured food webs all individuals do not share a common energy source, and the energy available (E) to larger individuals is constrained by inefficient energy transfer through the food chains that support them. This is expected to lead to steeper scalings ofNwithM. Here, we formalize and test an existing model for predicting abundance–body mass scaling, where the decline inEwithMis calculated from the mean predator–prey body mass ratio (from size‐based nitrogen stable isotope analysis) and trophic transfer efficiency. We show that the steep predicted scalings of abundance and body mass (Nscales asM−1.2,Bscales asM−0.2) in a marine food web are consistent with empirical estimates and can be attributed to the small predator–prey body mass ratio (106 : 1). As a previous study has shown that environmental stability may favour low predator–prey mass ratios and long food chains, we predict that steeper abundance–body mass relationships will be found in more stable environments.

Smaller predator-prey body size ratios in longer food chains.

Maximum food-chain length has been correlated with resource availability, ecosystem size, environmental stability and colonization history. Some of these correlations may result from environmental effects on predator-prey body size ratios. We investigate relationships between maximum food-chain length, predator-prey mass ratios, primary production and environmental stability in marine food webs with a natural history of community assembly. Our analyses provide empirical evidence that smaller mean predator-prey body size ratios are characteristic of more stable environments and that food chains are longer when mean predator-prey body size ratios are small. We conclude that environmental effects on predator-prey body size ratios contribute to observed differences in maximum food-chain length.

Traumatic Injuries in Two Neotropical Frogs, Dendrobates auratus and Physalaemus pustulosus

served juvenile V bengalensis which contained an average of almost 12 items per lizard, with predatorprey mass ratios of 0.006:0.5. He also stated that body fat accounted for over 28% of body mass in juvenile V bengalensis from Pakistan, where the animals do not feed for several months at the age of nine months. In contrast, juvenile V niloticus contained only negligible amounts of body fat. Their relatively low rate of food intake is unlikely to be the result of local food shortages, because all specimens appeared in good condition. Lack of fat accumulation in these animals therefore suggests that they do not undergo extended fasting. Acknowledgments.-Thanks to G. A. Punguse, N. K. Ankudey, M. Komoah, and B.Basuglo of Ghana Wildlife Department, S. Yeboah of Cape Coast University, and P. Racey of Aberdeen University for their assistance. Anonymous referees gave valuable suggestions on earlier drafts of the manuscript.