Ecological role and historical trends of large pelagic predators in a subtropical marine ecosystem of the South Atlantic

Large pelagic predators occupy high positions in food webs and could control lower trophic level species by direct and indirect ecological interactions. In this study we aimed to test the hypotheses: (1) pelagic predators are keystone species, and their removals could trigger impacts on the food chain; (2) higher landings of pelagic predators could trigger fishing impacts with time leading to a drop in the mean trophic level of catches; and (3) recovery in the pelagic predators populations, especially for sharks, could be achieved with fishing effort reduction. We performed a food web approach using an Ecopath with Ecosim model to represent the Southeastern and Southern Brazil, a subtropical marine ecosystem, in 2001. We then calibrated the baseline model using catch and fishing effort time series from 2001 to 2012. Afterwards, we simulated the impact of fishing effort changes on species and assessed the ecological impacts on the pelagic community from 2012 to 2025. Results showed that the model was well fitted to landing data for the majority of groups. The pelagic predators species were classified as keystone species impacting mainly on pelagic community. The ecosystem was resilient and fisheries seem sustainable at that time. However, the temporal simulation, from 2001 to 2012, revealed declines in the biomass of three sharks, tuna and billfish groups. It was possible observe declines in the mean trophic level of the catch and in the mean total length of landings. Longline fisheries particularly affected the sharks, billfish and swordfish, while hammerhead sharks were mostly impacted by gillnet fishery. Model simulations showed that large sharks’ biomasses could be recovered or maintained only after strong fishing effort reduction.     

Water temperature determines strength of top-down control in a stream food web

1. We examined effects of water temperature on the community structure of a three trophic level food chain (predatory fish, herbivorous caddisfly larvae and periphyton) in boreal streams. We used laboratory experiments to examine (i) the effects of water temperature on feeding activities of fish and caddisfly larvae and on periphyton productivity, to evaluate the thermal effects on each trophic level (species‐level experiment), and (ii) the effects of water temperature on predation pressure of fish on abundance of the lower trophic levels, to evaluate how temperature affects top‐down control by fish (community‐level experiment). 2. In the species‐level experiment, feeding activity of fish was high at 12 °C, which coincides with the mean summer temperature in forested streams of Hokkaido, Japan, but was depressed at 3 °C, which coincides with the mean winter temperature, and also above 18 °C, which coincides with the near maximum summer temperatures. Periphyton productivity increased over the range of water temperatures. 3. In the community‐level experiments, a top‐down effect of fish on the abundance of caddisfly larvae and periphyton was clear at 12 °C. This effect was not observed at 3 and 21 °C because of low predation pressure of fish at these temperatures. 4. These experiments revealed that trophic cascading effects may vary with temperature even in the presence of abundant predators. Physiological depression of predators because of thermal stress can alter top‐down control and lead to changes in community structure. 5. We suggest that thermal habitat alteration can change food web structure via combinations of direct and indirect trophic interactions.     

Linearity in the aggregate effects of multiple predators in a food web

Theory in community ecology often assumes that predator species have similar indirect effects and thus can be treated mathematically as a single functional unit (e.g. guild or trophic level). This assumption is questionable biologically because predator species typically differ in their effects, creating the potential for nonlinearities when they coexist. We evaluated the nature of indirect effects caused by three species of hunting spider predators, singly and in multiple species combinations, on grass and herb plants in experimental old-field food webs. Despite the potential for nonlinearity, indirect effects in different multiple predator combinations consistently did not differ significantly from the respective means of the single species effects. Thus, for this experimental system, the whole was simply the average of the parts. Consequently, models which abstract predator species as single trophic levels would successfully predict indirect effects in this system regardless of the composition of the predator fauna.     

Consequences of removing a keystone herbivore for the abundance and diversity of arthropods associated with a cruciferous shrub

Abstract. 1. The effect of the removal of Timarcha lugens (Chrysomelidae), one of the main herbivores of Hormathophylla spinosa (Cruciferae), on the abundance of co-occurring phytophagous insects, the abundance of non-phytophagous arthropods (detritivores, predators, and parasitoids), and the structure and diversity of the entire arthropod community, was studied for 3 years (1999–2001).
2. There was competition between T. lugens and co-occurring herbivores; the removal of T. lugens was correlated with an increase in the abundance of sap-suckers, flower-feeders, and, above all, folivores.
3. Timarcha lugens also had an indirect effect on arthropods belonging to other trophic levels; the abundance of predators increased significantly after the removal of T. lugens .
4. Community composition was affected by the experimental removal. In addition, the diversity of the overall community increased after removal of T. lugens .
5. The study demonstrated experimentally that T. lugens has a significant effect not only on other species belonging to the same trophic level, but also on the abundance of species belonging to higher trophic levels, and, consequently, on the entire structure and diversity of the complex community in which it is immersed.     

Top‐down and bottom‐up diversity cascades in detrital vs. living food webs

Apex predators and plant resources are both critical for maintaining diversity in biotic communities, but the indirect (‘cascading’) effects of top‐down and bottom‐up forces on diversity at different trophic levels are not well resolved in terrestrial systems. Manipulations of predators or resources can cause direct changes of diversity at one trophic level, which in turn can affect diversity at other trophic levels. The indirect diversity effects of resource and consumer variation should be strongest in aquatic systems, moderate in terrestrial systems, and weakest in decomposer food webs. We measured effects of top predators and plant resources on the diversity of endophytic animals in an understorey shrub Piper cenocladum (Piperaceae). Predators and resource availability had significant direct and indirect effects on the diversity of the endophytic animal community, but the effects were not interactive, nor were they consistent between living vs. detrital food webs. The addition of fourth trophic level beetle predators increased diversity of consumers supported by living plant tissue, whereas balanced plant resources (light and nutrients) increased the diversity of primary through tertiary consumers in the detrital resources food web. These results support the hypotheses that top‐down and bottom‐up diversity cascades occur in terrestrial systems, and that diversity is affected by different factors in living vs. detrital food webs.     

Predator-induced demographic shifts in coral reef fish assemblages

In recent years, it has become apparent that human impacts have altered community structure in coastal and marine ecosystems worldwide. Of these, fishing is one of the most pervasive, and a growing body of work suggests that fishing can have strong effects on the ecology of target species, especially top predators. However, the effects of removing top predators on lower trophic groups of prey fishes are less clear, particularly in highly diverse and trophically complex coral reef ecosystems. We examined patterns of abundance, size structure, and age-based demography through surveys and collection-based studies of five fish species from a variety of trophic levels at Kiritimati and Palmyra, two nearby atolls in the Northern Line Islands. These islands have similar biogeography and oceanography, and yet Kiritimati has ～10,000 people with extensive local fishing while Palmyra is a US National Wildlife Refuge with no permanent human population, no fishing, and an intact predator fauna. Surveys indicated that top predators were relatively larger and more abundant at unfished Palmyra, while prey functional groups were relatively smaller but showed no clear trends in abundance as would be expected from classic trophic cascades. Through detailed analyses of focal species, we found that size and longevity of a top predator were lower at fished Kiritimati than at unfished Palmyra. Demographic patterns also shifted dramatically for 4 of 5 fish species in lower trophic groups, opposite in direction to the top predator, including decreases in average size and longevity at Palmyra relative to Kiritimati. Overall, these results suggest that fishing may alter community structure in complex and non-intuitive ways, and that indirect demographic effects should be considered more broadly in ecosystem-based management.     

Predator diversity and density affect levels of predation upon strongly interactive species in temperate rocky reefs

Indirect effects of predators in the classic trophic cascade theory involve the effects of basal species (e.g. primary producers) mediated by predation upon strongly interactive consumers (e.g. grazers). The diversity and density of predators, and the way in which they interact, determine whether and how the effects of different predators on prey combine. Intraguild predation, for instance, was observed to dampen the effects of predators on prey in many ecosystems. In marine systems, species at high trophic levels are particularly susceptible to extinction (at least functionally). The loss of such species, which is mainly attributed to human activities (mostly fishing), is presently decreasing the diversity of marine predators in many areas of the world. Experimental studies that manipulate predator diversity and investigate the effects of this on strongly interactive consumers (i.e. those potentially capable of causing community-wide effects) in marine systems are scant, especially in the rocky sublittoral. I established an experiment that utilised cage enclosures to test whether the diversity and density of fish predators (two sea breams and two wrasses) would affect predation upon juvenile and adult sea urchins, the most important grazers in Mediterranean sublittoral rocky reefs. Changes in species identity (with sea breams producing major effects) and density of predators affected predation upon sea urchins more than changes in species richness per se. Predation upon adult sea urchins decreased in the presence of multiple predators, probably due to interference competition between sea breams and wrasses. This study suggests that factors that influence both fish predator diversity and density in Mediterranean rocky reefs (e.g. fishing and climate change) may have the potential to affect the predators' ability to control sea urchin population density, with possible repercussions for the whole benthic community structure.     

Disturbance-mediated trophic interactions and plant performance

Disturbances, such as flooding, play important roles in determining community structure. Most studies of disturbances focus on the direct effects and, hence, the indirect effects of disturbances are poorly understood. Within terrestrial riparian areas, annual flooding leads to differences in the arthropod community as compared to non-flooded areas. In turn, these differences are likely to alter the survival, growth, and reproduction of plant species via an indirect effect of flooding (i.e., changes in herbivory patterns). To test for such effects, an experiment was conducted wherein arthropod predators and herbivores were excluded from plots in flooded and non-flooded areas and the impact on a common riparian plant, Mimulus guttatus was examined. In general, the direct effect of flooding on M. guttatus was positive. The indirect effects, however, significantly decreased plant survival for both years of the experiment, regardless of predator presence, because of an increased exposure to grasshoppers, the most abundant herbivore in the non-flooded sites. Leafhoppers, which were more abundant in the flooded sites, had much weaker and varying effects. During 2000, when the leafhopper herbivory was high, arthropod predators did not significantly reduce damage to plants. In 2001, the mean herbivory damage was lower and predators were able to significantly reduce overall leafhopper damage. The effects of predators on leafhoppers, however, did not increase plant survival, final weight, or the reproduction potential and, thus, did not initiate a species-level trophic cascade. Overall, it was the differences in the herbivore community that led to a significant decrease in plant survival. While flooding certainly alters riparian plant survival through direct abiotic effects, it also indirectly affects riparian plants by changing the arthropod community, in particular herbivores, and hence trophic interactions.     

Roles of parasites in animal invasions

Biological invasions are global threats to biodiversity and parasites might play a role in determining invasion outcomes. Transmission of parasites from invading to native species can occur, aiding the invasion process, whilst the 'release' of invaders from parasites can also facilitate invasions. Parasites might also have indirect effects on the outcomes of invasions by mediating a range of competitive and predatory interactions among native and invading species. Although pathogen outbreaks can cause catastrophic species loss with knock-on effects for community structure, it is less clear what impact persistent, sub-lethal parasitism has on native-invader interactions and community structure. Here, we show that the influence of parasitism on the outcomes of animal invasions is more subtle and wide ranging than has been previously realized.     

Incorporating anthropogenic effects into trophic ecology: predator–prey interactions in a human-dominated landscape

Apex predators perform important functions that regulate ecosystems worldwide. However, little is known about how ecosystem regulation by predators is influenced by human activities. In particular, how important are top-down effects of predators relative to direct and indirect human-mediated bottom-up and top-down processes? Combining data on species'' occurrence from camera traps and hunting records, we aimed to quantify the relative effects of top-down and bottom-up processes in shaping predator and prey distributions in a human-dominated landscape in Transylvania, Romania. By global standards this system is diverse, including apex predators (brown bear and wolf), mesopredators (red fox) and large herbivores (roe and red deer). Humans and free-ranging dogs represent additional predators in the system. Using structural equation modelling, we found that apex predators suppress lower trophic levels, especially herbivores. However, direct and indirect top-down effects of humans affected the ecosystem more strongly, influencing species at all trophic levels. Our study highlights the need to explicitly embed humans and their influences within trophic cascade theory. This will greatly expand our understanding of species interactions in human-modified landscapes, which compose the majority of the Earth''s terrestrial surface.     

Predator Diversity Effects in an Exotic Freshwater Food Web

Cascading trophic interactions are often defined as the indirect effects of a predator on primary producers through the effect of the predator on herbivores. These effects can be both direct through removal of herbivores [density-mediated indirect interactions (DMIIs)] or indirect through changes in the behavior of the herbivores [trait-mediated indirect interactions (TMIIs)]. How the relative importance of these two indirect interactions varies with predator diversity remains poorly understood. We tested the effect of predator diversity on both TMIIs and DMIIs on phytoplankton using two competitive invasive dreissenid mussel species (zebra mussel and quagga mussel) as the herbivores and combinations of one, two or all three species of the predators pumpkinseed sunfish, round goby, and rusty crayfish. Predators had either direct access to mussels and induced both TMII and DMII, or no direct access and induced only TMII through the presence of risk cues. In both sets of treatments, the predators induced a trophic cascade which resulted in more phytoplankton remaining with predators present than with only mussels present. The trophic cascade was weaker in three-predator and two-predator treatments than in one-predator treatments when predators had direct access to dreissenids (DMIIs and TMIIs). Crayfish had higher cascading effects on phytoplankton than both pumpkinseed and round goby. Increased predator diversity decreased the strength of DMIIs but had no effect on the strength of TMIIs. The strength of TMIIs was higher with zebra than quagga mussels. Our study suggests that inter-specific interference among predators in multi-species treatments weakens the consumptive cascading effects of predation on lower trophic levels whereas the importance of predator diversity on trait mediated effects depends on predator identity.     

Trophic consequences of a biological invasion: do plant invasions increase predator abundance?

As invasive species become integrated into existing communities, they engage in a wide variety of trophic interactions with other community members. Many of these interactions are direct (e.g. predator–prey interactions or interference competition), but invasive species also can affect native community members indirectly, by influencing the abundances of intermediary species in trophic webs. Observational studies suggest that invasive plant species affect herbivorous arthropod communities and that these effects may flow up trophic webs to influence the abundance of predators. However, few studies have experimentally manipulated the presence of invasive plants to quantify the effects of plant invasion on higher trophic levels. Here, I use comparisons across sites that have or have not been invaded by the invasive plant Medicago polymorpha, combined with experimental removals of Medicago and insect herbivores, to investigate how a plant invasion affects the abundance of predators. Both manipulative and observational experiments showed that Medicago increased the abundance of the exotic herbivore Hypera and predatory spiders, suggesting positive bottom–up effects of plant invasions on higher trophic levels. Path analyses conducted on data from natural habitats revealed that Medicago primarily increased spider abundance through herbivore‐mediated indirect pathways. Specifically, Medicago density was positively correlated with the abundance of the dominant herbivore Hypera, and increased Hypera densities were correlated with increased spider abundance. Smaller‐scale experimental studies confirmed that Medicago may increase spider abundance through herbivore‐mediated indirect pathways, but also showed that the effects of Medicago varied across sites, including having no effect or having direct effects on spider abundance. If effects of invasive species commonly flow through trophic webs, then invasive species have the potential to affect numerous species throughout the community, especially those species whose dynamics are tightly connected to highly‐impacted community members through trophic linkages.     

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.     

Herbivore species richness, composition and community structure mediate predator richness effects and top-down control of herbivore biomass

Changes in predator species richness can have important consequences for ecosystem functioning at multiple trophic levels, but these effects are variable and depend on the ecological context in addition to the properties of predators themselves. Here, we report an experimental study to test how species identity, community attributes, and community structure at the herbivore level moderate the effects of predator richness on ecosystem functioning. Using mesocosms containing predatory insects and aphid prey, we independently manipulated species richness at both predator and herbivore trophic levels. Community structure was also manipulated by changing the distribution of herbivore species across two plant species. Predator species richness and herbivore species richness were found to negatively interact to influence predator biomass accumulation, an effect which is hypothesised to be due to the breakdown of functional complementarity among predators in species-rich herbivore assemblages. The strength of predator suppression of herbivore biomass decreased as herbivore species richness and distribution across host plants increased, and positive predator richness effects on herbivore biomass suppression were only observed in herbivore assemblages of relatively low productivity. In summary, the study shows that the species richness, productivity and host plant distribution of prey communities can all moderate the general influence of predators and the emergence of predator species richness effects on ecosystem functioning.     

Non-linear relationship between body size of terrestrial carnivores and their trophic niche breadth and overlap

Studying food partitioning of mammalian predators is important for understanding trophic structures and interactions between coexisting carnivore species. This is particularly pertinent in the light of expanding ranges of populations of generalist species whose habitat and diet overlap with more specialized species. Here, we tested the resource partitioning hypothesis in terrestrial carnivores, predicting that trophic niche breadth and overlap relate positively to body mass. We used dietary data from 18 terrestrial carnivore taxa in four families (Canidae, Mustelidae, Felidae and Ursidae; body mass 0.1–173.6 kg) in three regions in Central and Eastern Europe, i.e. deciduous forest and forest-steppe region (DFR), temperate deciduous and mixed forest region (MFR) and transitory mixed forest regions (TFR). We ranked carnivores along an axis of trophic niche (breadth and overlap), and analysed the relationship between trophic niche and body mass (or pair-wise difference in body mass). A hierarchical cluster analysis of diet composition divided carnivores into four ecological groups: wild ungulate predators; small-mammal predators; amphibians and small mammal predators and omnivores. The relationship between body mass of predators and both trophic niche breadth and trophic niche overlap were hump-shaped. The trophic niche breadth to body mass ratio was significantly lower in DFR than in TFR and trophic niche overlap was significantly higher in DFR than in MFR and TFR. The predominant food resource is small mammals whose abundance is related to local agricultural and forestry management practices. Modifications of management techniques can affect population dynamics and community composition of carnivore species, especially in the case of small-mammal predators.     

生态系统的组织理论：食物链动态论与互惠共生—控制论

本文主要介绍生态系统的组织理论,食物链动态理论和互惠共生－控制论,这两种理论代表了进化生态学家与系统生态学家两个学派。前者是建立在达尔文的“生存竞争”思想耻,强调初级生产力是关键变量,捕食作用和食物资源两者随初级生产力梯度的增加交替控制食物网结构,并预报相邻营养级的生物量和周转率为负相关关系,被称作“生态学的中心理论”后者是建立在控制论基础上,认为生态系统是正,负反馈联合构建的,并预报生物量和周转     

Community‐wide effects of below‐ground rhizobia on above‐ground arthropods

1. Plants take nutrients for their growth and reproduction from not only soil but also symbiotic microbes in the rhizosphere, and therefore below‐ground microbes may indirectly influence the above‐ground arthropod community through changes in the quality and quantity of plants. 2. Rhizobia are root‐nodulating bacteria that provide NH₄⁺ to legume plants. We examined bottom‐up effects of rhizobia on the community properties of the arthropods on host plants, using a root‐nodulating soybean strain (R+) and a non‐nodulating strain (R?) in a common garden. 3. R+ plants grew larger and produced a greater number of leaves than R? plants. We observed 28 species of herbivores and three taxonomic groups of predators on R+ and R? plants. The herbivorous species were classified into sap feeders (12 species) and chewers (16 species). 4. The species richness of overall herbivores, sap feeders, and chewers on R+ plants was greater than that on R? plants. Rhizobia positively affected the abundance of chewers. 5. The community composition of herbivores was significantly different between R? and R+ plants, although species diversity and evenness did not differ. 6. Rhizobia‐induced bottom‐up effects were transmitted to the third trophic level. The abundance, taxonomic richness, and diversity of the predators on R+ plants were greater but evenness was lower than those on R? plants. The community composition of predators was not affected by rhizobia. 7. These results indicate that the below‐ground microbes initiated bottom‐up effects on above‐ground herbivores and predators through trophic levels.     

Effects of trophic similarity on community composition

Understanding how ecological processes determine patterns among species coexisting within ecosystems is central to ecology. Here, we explore relationships between species’ local coexistence and their trophic niches in terms of their feeding relationships both as consumers and as resources. We build on recent concepts and methods from community phylogenetics to develop a framework for analysing mechanisms responsible for community composition using trophic similarity among species and null models of community assembly. We apply this framework to 50 food webs found in 50 Adirondack lakes and find that species composition in these communities appears to be driven by both bottom‐up effects by which the presence of prey species selects for predators of those prey, and top‐down effects by which prey more tolerant of predation out‐compete less tolerant prey of the same predators. This approach to community food webs is broadly applicable and shows how species interaction networks can inform an increasingly large array of theory central to community ecology.     

Relative strengths of top-down and bottom-up forces in a tropical forest community

We tested integrative bottom-up and top-down trophic cascade hypotheses with manipulative experiments in a tropical wet forest, using the ant-plant Piper cenocladum and its associated arthropod community. We examined enhanced nutrients and light along with predator and herbivore exclusions as sources of variation in the relative biomass of plants, their herbivores (via rates of herbivory), and resident predaceous ants. The combined manipulations of secondary consumers, primary consumers, and plant resources allowed us to examine some of the direct and indirect effects on each trophic level and to determine the relative contributions of bottom-up and top-down cascades to the structure of the community. We found that enhanced plant resources (nutrients and light) had direct positive effects on plant biomass. However, we found no evidence of indirect (cascading through the herbivores) effects of plant biomass on predators or top predators. In contrast, ants had indirect effects on plant biomass by decreasing herbivory on the plants. This top-down cascade occurred whether or not plant resources were enriched, conditions which are expected to modify top-down forces. Received: 9 August 1998 / Accepted: 1 December 1998     

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).