Construction and evaluation of a robust trophic network model for the northern Gulf of Mexico ecosystem

A critical issue in understanding trophic connectivity in ecological systems is the lack in quality and quantity of information about feeding habits. In this work, we present a method for integrating a diversity of feeding habits data from published studies to evaluate the impact on indices that describe characteristics of individual taxa and their connectivity. We focus our study on feeding habits of the fishes of the northern Gulf of Mexico and seek to understand the importance of the forage fish Gulf Menhaden (Brevoortia patronus) in predator diets. We created a database of diet studies from the northern Gulf of Mexico that included six diet metrics: frequency of occurrence, wet weight, dry weight, number, volume, index of relative importance, and index of caloric importance. We then used this information to construct a set of traditional networks (all prey and predators were from a single taxonomic level and trophic connections were parameterized with a single diet metric). We also constructed a “robust” network where all taxa were identified to the lowest taxonomic level and trophic connections were parameterized using a resampling approach that included all available information. Linear regression and resampling methods were used to convert data reported in other diet metrics into the frequency of occurrence diet metric. For both traditional and robust networks, we used network indices to describe topological properties. With the robust network, we conducted removal simulations where the forage fish species Gulf Menhaden, and associated Clupeidae representatives, were removed from the network and the feeding effort of the predators was reallocated among their other prey items. We found that network and node-specific indices were sensitive to the choice of taxonomy and diet metric level. In the robust network, predator species with the greatest number of identified prey had the lowest precision in their connections and prey from the Arthropoda phyla had the lowest precision for connections. From the removal and reallocation simulations, we found that Actinopterygii and Arthropoda were the most impacted prey taxa with 1.2% to 4.3% increase in predation and approximately 23 taxa would receive 50% of the reallocated predation. Overall, the resampling methods we present provide a potential means for combining disparate diet data and enables a comprehensive understanding of trophic interactions within an ecosystem.     

Predator diversity and the functioning of ecosystems: the role of intraguild predation in dampening trophic cascades

Single trophic‐level studies of the relationship between biodiversity and ecosystem functioning highlight the importance of mechanisms such as resource partitioning, facilitation, and sampling effect. In a multi‐trophic context, trophic interactions such as intraguild predation may also be an important mediator of this relationship. Using a salt‐marsh food web, we investigated the interactive effects of predator species richness (one to three species) and trophic composition (strict predators, intraguild predators, or a mixture of the two) on ecosystem functions such as prey suppression and primary production via trophic cascades. We found that the trophic composition of the predator assemblage determined the impact of increasing predator species richness on the occurrence of trophic cascades. In addition, increasing the proportion of intraguild predator species present diminished herbivore suppression and reduced primary productivity. Therefore, trophic composition of the predator assemblage can play an important role in determining the nature of the relationship between predator diversity and ecosystem function.     

Intermediate predator impact on consumers weakens with increasing predator diversity in the presence of a top-predator

Adding or removing a top-predator is known to affect lower trophic levels with potentially large, indirect effects on primary production. However, little is known about how predator diversity may affect lower trophic levels, or how adding or removing a top-predator influences the effects of predator diversity. Using aquatic mesocosms containing three and four trophic levels, we tested whether intermediate predator diversity affected predation on consumers and if top-predator presence influenced such effects. We found that the presence of intermediate predators suppressed the consumer population and that this suppression tended to increase with increased intermediate predator diversity when the top-predator was absent. However, with the top-predator present, increased intermediate predator diversity showed the opposite effect on the consumers compared to without a top-predator, i.e. decreased suppression of consumers with increased diversity. Hence, in our study, the loss of intermediate predator species weakened or strengthened predator–prey interactions depending on if the top-predator was present or not, while loss of the top-predator only strengthened the predator–prey interactions. Therefore, the loss of a predator species may render different, but perhaps predictable effects on the functioning of a system depending on from which trophic level it is lost and on the initial number of species in that trophic level.     

长江口及邻近海域高营养层次生物群落功能群及其变化

通过对2006年6、8和10月在长江口及邻近海域3次调查采集样品的分析,对该水域的高营养层次生物群落的功能群组成及其变化进行了研究.结果表明：长江口及邻近海域高营养层次生物群落包括鱼食性、蟹食性、虾食性、底栖动物食性、浮游生物食性和广食性6个功能群.由于受海洋环境变化以及鱼类洄游活动的影响,各月份长江口及邻近海域高营养层次生物群落的组成及营养级都有较大的变化.6月高营养层次以鱼类、毛虾类和蟹类为主,以浮游生物食性功能群为主要功能群,营养级最低,为3.06;8月高营养层次以鱼类为主,虾食性功能群为主要功能群,营养级达到最高,为3.78;10月高营养层次虽仍以鱼类为主,虾蟹类比例增大,功能群以浮游动物食性和底栖动物食性功能群为主,营养级为3.58.     

Trophic cascades and trophic trickles in pelagic food webs

Prey-dependent models, with the predation rate (per predator) a function of prey numbers alone, predict the existence of a trophic cascade. In a trophic cascade, the addition of a top predator to a two-level food chain to make a three-level food chain will lead to increases in the population size of the primary producers, and the addition of nutrients to three-level chains will lead to increases in the population numbers at only the first and third trophic levels. In contrast, ratio-dependent models, with the predation rate (per predator) dependent on the ratio of predator numbers to prey, predict that additions of top predators will not increase the population sizes of the primary producers, and that the addition of nutrients to a three-level food chain will lead to increases in population numbers at all trophic levels. Surprisingly, recent meta-analyses show that freshwater pelagic food web patterns match neither prey-dependent models (in pelagic webs, ''prey'' are phytoplankton, and ''predators'' are zooplankton), nor ratio-dependent models. In this paper we use a modification of the prey-dependent model, incorporating strong interference within the zooplankton trophic level, that does yield patterns matching those found in nature. This zooplankton interference model corresponds to a more reticulate food web than in the linear, prey-dependent model, which lacks zooplankton interference. We thus reconcile data with a new model, and make the testable prediction that the strength of trophic cascades will depend on the degree of heterogeneity in the zooplankton level of the food chain.     

Predator interference emerging from trophotaxis in predator–prey systems: An individual-based approach

An individual-based model describing predator–prey interactions within a closed rectangular habitat was developed to study how different assumptions about the individual movements lead to the emergence at the population level of various kinds of prey- and predator-dependence in the spatially aggregated trophic function.In addition to random walk, both species are capable of directional movement, i.e., the model accounts for the predator prey-taxis and evasion of predators by prey individuals. The taxis stimulus of each species is the odour of the other species, which is distributed continuously in space. Spatial behaviour of individuals is determined by the specific response to the odour gradient and the tendency to maintain the taxis velocity.In order to facilitate the assessment of the trophic function, the model allows removing the effect of demographic density variations on the predator ration, keeping population sizes constant.Analyzing the dependence of the trophic function with the average predator density, we found that, depending on the intensity of taxis, the predator population exhibits various degrees of interference, from very low to very high values. In particular, a moderate taxis generates distinct levels of interference including the ratio-dependent case. The letter maximizes the average consumption rate.A new generalized function containing ratio-dependence and prey-dependence as special cases, at high and low population abundances, is suggested. This trophic function fits the simulated data better than the Hassell–Varley–Holling expression does.     

Prey size diversity hinders biomass trophic transfer and predator size diversity promotes it in planktonic communities

Body size exerts multiple effects on plankton food-web interactions. However, the influence of size structure on trophic transfer remains poorly quantified in the field. Here, we examine how the size diversity of prey (nano-microplankton) and predators (mesozooplankton) influence trophic transfer efficiency (using biomass ratio as a proxy) in natural marine ecosystems. Our results support previous studies on single trophic levels: transfer efficiency decreases with increasing prey size diversity and is enhanced with greater predator size diversity. We further show that communities with low nano-microplankton size diversity and high mesozooplankton size diversity tend to occur in warmer environments with low nutrient concentrations, thus promoting trophic transfer to higher trophic levels in those conditions. Moreover, we reveal an interactive effect of predator and prey size diversities: the positive effect of predator size diversity becomes influential when prey size diversity is high. Mechanistically, the negative effect of prey size diversity on trophic transfer may be explained by unicellular size-based metabolic constraints as well as trade-offs between growth and predation avoidance with size, whereas increasing predator size diversity may enhance diet niche partitioning and thus promote trophic transfer. These findings provide insights into size-based theories of ecosystem functioning, with implications for ecosystem predictive models.     

Latitude and protection affect decadal trends in reef trophic structure over a continental scale

The relative roles of top‐down (consumer‐driven) and bottom‐up (resource‐driven) forcing in exploited marine ecosystems have been much debated. Examples from a variety of marine systems of exploitation‐induced, top‐down trophic forcing have led to a general view that human‐induced predator perturbations can disrupt entire marine food webs, yet other studies that have found no such evidence provide a counterpoint. Though evidence continues to emerge, an unresolved debate exists regarding both the relative roles of top‐down versus bottom‐up forcing and the capacity of human exploitation to instigate top‐down, community‐level effects. Using time‐series data for 104 reef communities spanning tropical to temperate Australia from 1992 to 2013, we aimed to quantify relationships among long‐term trophic group population density trends, latitude, and exploitation status over a continental‐scale biogeographic range. Specifically, we amalgamated two long‐term monitoring databases of marine community dynamics to test for significant positive or negative trends in density of each of three key trophic levels (predators, herbivores, and algae) across the entire time series at each of the 104 locations. We found that trophic control tended toward bottom‐up driven in tropical systems and top‐down driven in temperate systems. Further, alternating long‐term population trends across multiple trophic levels (a method of identifying trophic cascades), presumably due to top‐down trophic forcing, occurred in roughly fifteen percent of locations where the prerequisite significant predator trends occurred. Such alternating trophic trends were significantly more likely to occur at locations with increasing predator densities over time. Within these locations, we found a marked latitudinal gradient in the prevalence of long‐term, alternating trophic group trends, from rare in the tropics (<5% of cases) to relatively common in temperate areas (~45%). Lastly, the strongest trends in predator and algal density occurred in older no‐take marine reserves; however, exploitation status did not affect the likelihood of alternating long‐term trophic group trends occurring. Our data suggest that the type and degree of trophic forcing in this system are likely related to one or more covariates of latitude, and that ecosystem resiliency to top‐down control does not universally vary in this system based on exploitation level.     

Temperature and prey density jointly influence trophic and non‐trophic interactions in multiple predator communities

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Different groups of ground-dwelling spiders share similar trophic niches in temperate forests

1. Generalistic interactions between predator and prey may vary with ecosystem type, predator traits, and prey traits, but the interplay of these factors has not been assessed in ground food webs.
2. We investigated trophic interactions of ground-dwelling spiders across eight forests in European Russia associated with body size, hunting strategy, microhabitat specialization, potential prey type, potential prey population density, and forest type (coniferous vs. broadleaved). We analyzed 128 individual spiders, including juveniles, all identified to the family level with two complementary methods: molecular gut content analysis, and stable isotope analysis of carbon and nitrogen.
3. The results suggest that feeding frequency of spiders is affected by predator body size and by selection of certain prey type. Stable isotope analysis showed similar trophic niches among spider families, varying moderately with forest type. Larger spiders had higher Δ¹³C values than smaller ones, but similar Δ¹⁵N values, suggesting that different size classes of spiders belong to different food chains. Results based on stable isotope and molecular gut content analyses were weakly linked, indicating them targeting different trophic niche dimensions.
4. At least for the group-level interactions, family identity and hunting strategy of predator has little predictive power while predator body size and prey traits affected trophic niche dimensions calling for future studies in this direction. Large spiders feed more and rely on different basal resources than small spiders, suggesting that including small species and juveniles provides a more comprehensive picture of food web organization.

     

Eco-evolutionary trophic dynamics: loss of top predators drives trophic evolution and ecology of prey

Ecosystems are being altered on a global scale by the extirpation of top predators. The ecological effects of predator removal have been investigated widely; however, predator removal can also change natural selection acting on prey, resulting in contemporary evolution. Here we tested the role of predator removal on the contemporary evolution of trophic traits in prey. We utilized a historical introduction experiment where Trinidadian guppies (Poecilia reticulata) were relocated from a site with predatory fishes to a site lacking predators. To assess the trophic consequences of predator release, we linked individual morphology (cranial, jaw, and body) to foraging performance. Our results show that predator release caused an increase in guppy density and a "sharpening" of guppy trophic traits, which enhanced food consumption rates. Predator release appears to have shifted natural selection away from predator escape ability and towards resource acquisition ability. Related diet and mesocosm studies suggest that this shift enhances the impact of guppies on lower trophic levels in a fashion nuanced by the omnivorous feeding ecology of the species. We conclude that extirpation of top predators may commonly select for enhanced feeding performance in prey, with important cascading consequences for communities and ecosystems.     

Predator diversity and environmental change modify the strengths of trophic and nontrophic interactions

Understanding the dependence of species interaction strengths on environmental factors and species diversity is crucial to predict community dynamics and persistence in a rapidly changing world. Nontrophic (e.g. predator interference) and trophic components together determine species interaction strengths, but the effects of environmental factors on these two components remain largely unknown. This impedes our ability to fully understand the links between environmental drivers and species interactions. Here, we used a dynamical modelling framework based on measured predator functional responses to investigate the effects of predator diversity, prey density, and temperature on trophic and nontrophic interaction strengths within a freshwater food web. We found that (i) species interaction strengths cannot be predicted from trophic interactions alone, (ii) nontrophic interaction strengths vary strongly among predator assemblages, (iii) temperature has opposite effects on trophic and nontrophic interaction strengths, and (iv) trophic interaction strengths decrease with prey density, whereas the dependence of nontrophic interaction strengths on prey density is concave up. Interestingly, the qualitative impacts of temperature and prey density on the strengths of trophic and nontrophic interactions were independent of predator identity, suggesting a general pattern. Our results indicate that taking multiple environmental factors and the nonlinearity of density‐dependent species interactions into account is an important step towards a better understanding of the effects of environmental variations on complex ecological communities. The functional response approach used in this study opens new avenues for (i) the quantification of the relative importance of the trophic and nontrophic components in species interactions and (ii) a better understanding how environmental factors affect these interactions and the dynamics of ecological communities.     

Stepping in Elton's footprints: a general scaling model for body masses and trophic levels across ecosystems

Despite growing awareness of the significance of body-size and predator-prey body-mass ratios for the stability of ecological networks, our understanding of their distribution within ecosystems is incomplete. Here, we study the relationships between predator and prey size, body-mass ratios and predator trophic levels using body-mass estimates of 1313 predators (invertebrates, ectotherm and endotherm vertebrates) from 35 food-webs (marine, stream, lake and terrestrial). Across all ecosystem and predator types, except for streams (which appear to have a different size structure in their predator-prey interactions), we find that (1) geometric mean prey mass increases with predator mass with a power-law exponent greater than unity and (2) predator size increases with trophic level. Consistent with our theoretical derivations, we show that the quantitative nature of these relationships implies systematic decreases in predator-prey body-mass ratios with the trophic level of the predator. Thus, predators are, on an average, more similar in size to their prey at the top of food-webs than that closer to the base. These findings contradict the traditional Eltonian paradigm and have implications for our understanding of body-mass constraints on food-web topology, community dynamics and stability.     

Predator personality structures prey communities and trophic cascades

Intraspecific variation is central to our understanding of evolution and population ecology, yet its consequences for community ecology are poorly understood. Animal personality – consistent individual differences in suites of behaviours – may be particularly important for trophic dynamics, where predator personality can determine activity rates and patterns of attack. We used mesocosms with aquatic food webs in which the top predator (dragonfly nymphs) varied in activity and subsequent attack rates on zooplankton, and tested the effects of predator personality. We found support for four hypotheses: (1) active predators disproportionately reduce the abundance of prey, (2) active predators select for predator‐resistant prey species, (3) active predators strengthen trophic cascades (increase phytoplankton abundance) and (4) active predators are more likely to cannibalise one another, weakening all other trends when at high densities. These results suggest that intraspecific variation in predator personality is an important determinant of prey abundance, community composition and trophic cascades.     

Predator decline leads to decreased stability in a coastal fish community

Fisheries exploitation has caused widespread declines in marine predators. Theory predicts that predator depletion will destabilise lower trophic levels, making natural communities more vulnerable to environmental perturbations. However, empirical evidence has been limited. Using a community matrix model, we empirically assessed trends in the stability of a multispecies coastal fish community over the course of predator depletion. Three indices of community stability (resistance, resilience and reactivity) revealed significantly decreasing stability concurrent with declining predator abundance. The trophically downgraded community exhibited weaker top‐down control, leading to predator‐release processes in lower trophic levels and increased susceptibility to perturbation. At the community level, our results suggest that high predator abundance acts as a stabilising force to the naturally stochastic and highly autocorrelated dynamics in low trophic species. These findings have important implications for the conservation and management of predators in marine ecosystems and provide empirical support for the theory of predatory control.     

Predator diversity enhances secondary production and decreases the likelihood of trophic cascades

We manipulated the diversity of top predators in a three trophic level marine food web. The food web included four top benthic marine fish predators (black goby, rock goby, sea scorpion and shore rockling), an intermediate trophic level of small fish, and a lower trophic level of benthic invertebrates. We kept predator density constant and monitored the response of the lower trophic levels. As top predator diversity increased, secondary production increased. We also observed that in the presence of the manipulated fish predators, the density of small gobiid fish (intermediate consumers) was suppressed, releasing certain groups of benthic invertebrates (caprellid amphipods, copepods, nematodes and spirorbid worms) from heavy intermediate predation pressure. We attribute the mechanism responsible for this trophic cascade to a trait-mediated indirect interaction, with the small gobiid fish changing their use of space in response to altered predator diversity. In the absence of top fish predators, a full-blown trophic cascade occurs. Therefore the diversity of predators reduces the likelihood of trophic cascades occurring and hence provides insurance against the loss of an important ecosystem function (i.e. secondary production).     

Environmental and plant genetic effects on tri‐trophic interactions

The effects of plant genotype and environmental factors on tri‐trophic interactions have usually been investigated separately, limiting our ability to compare the relative strength of these effects as well as their potential to interactively shape arthropod communities. We studied the interactions among the herb Ruellia nudiflora, a seed predator, and its parasitoids using 14 maternal plant families grown in a common garden. By fertilizing half of the plants of each family and subsequently recording fruit number, seed predator number, and parasitoid number per plant, we sought to compare the strength of plant genetic effects with those of soil fertility, and determine if these factors interactively shape tri‐trophic interactions. Furthermore, we evaluated if these bottom–up factors influenced higher trophic levels through changes in abundance across trophic levels (density‐mediated) or changes in the function of species interactions (trait‐mediated). Plant genetic effects on seed predators and parasitoids were stronger than fertilization effects. Moreover, we did not find plant genetic variation for fertilization effects on fruit, seed predator, or parasitoid abundance, showing that each factor acted independently on plant resources and higher trophic levels. Both bottom–up forces were transmitted via density‐mediated effects where increased fruit number from fertilization and plant genetic effects increased seed predator and parasitoid abundance; however, seed predator attack was density‐dependent, while parasitoid attack was density‐independent. Importantly, there was evidence (marginally significant in one case) that fertilization modified the function of plant‐seed predator and seed predator–parasitoid interactions by increasing the number of seed predators per fruit and decreasing the number of parasitoids per seed predator, respectively. These findings show that plant genetic and soil fertility effects cascaded up this simple food chain, that plant genetic effects were stronger across all trophic levels, and that these effects were transmitted independently and through contrasting mechanisms.     

Predator diversity strengthens trophic cascades in kelp forests by modifying herbivore behaviour

Although human-mediated extinctions disproportionately affect higher trophic levels, the ecosystem consequences of declining diversity are best known for plants and herbivores. We combined field surveys and experimental manipulations to examine the consequences of changing predator diversity for trophic cascades in kelp forests. In field surveys we found that predator diversity was negatively correlated with herbivore abundance and positively correlated with kelp abundance. To assess whether this relationship was causal, we manipulated predator richness in kelp mesocosms, and found that decreasing predator richness increased herbivore grazing, leading to a decrease in the biomass of the giant kelp Macrocystis. The presence of different predators caused different herbivores to alter their behaviour by reducing grazing, such that total grazing was lowest at highest predator diversity. Our results suggest that declining predator diversity can have cascading effects on community structure by reducing the abundance of key habitat-providing species.     

Pyramids and cascades: a synthesis of food chain functioning and stability

Food chain theory is one of the cornerstones of ecology, providing many of its basic predictions, such as biomass pyramids, trophic cascades and predator–prey oscillations. Yet, ninety years into this theory, the conditions under which these patterns may occur and persist in nature remain subject to debate. Rather than address each pattern in isolation, we propose that they must be understood together, calling for synthesis in a fragmented landscape of theoretical and empirical results. As a first step, we propose a minimal theory that combines the long‐standing energetic and dynamical approaches of food chains. We chart theoretical predictions on a concise map, where two main regimes emerge: across various functioning and stability metrics, one regime is characterised by pyramidal patterns and the other by cascade patterns. The axes of this map combine key physiological and ecological variables, such as metabolic rates and self‐regulation. A quantitative comparison with data sheds light on conflicting theoretical predictions and empirical puzzles, from size spectra to causes of trophic cascade strength. We conclude that drawing systematic connections between various existing approaches to food chains, and between their predictions on functioning and stability, is a crucial step in confronting this theory to real ecosystems.     

Cascading effects of predator diversity and omnivory in a marine food web

Over‐harvesting, habitat loss and exotic invasions have altered predator diversity and composition in a variety of communities which is predicted to affect other trophic levels and ecosystem functioning. We tested this hypothesis by manipulating predator identity and diversity in outdoor mesocosms that contained five species of macroalgae and a macroinvertebrate herbivore assemblage dominated by amphipods and isopods. We used five common predators including four carnivores (crabs, shrimp, blennies and killifish) and one omnivore (pinfish). Three carnivorous predators each induced a strong trophic cascade by reducing herbivore abundance and increasing algal biomass and diversity. Surprisingly, increasing predator diversity reversed these effects on macroalgae and altered algal composition, largely due to the inclusion and performance of omnivorous fish in diverse predator assemblages. Changes in predator diversity can cascade to lower trophic levels; the exact effects, however, will be difficult to predict due to the many complex interactions that occur in diverse food webs.