Significance of predation by protists in aquatic microbial food webs

Predation in aquatic microbial food webs is dominated by phagotrophic protists, yet these microorganisms are still understudied compared to bacteria and phytoplankton. In pelagic ecosystems, predaceous protists are ubiquitous, range in size from 2 μm flagellates to >100 μm ciliates and dinoflagellates, and exhibit a wide array of feeding strategies. Their trophic states run the gamut from strictly phagotrophic, to mixotrophic: partly autotrophic and partly phagotrophic, to primarily autotrophic but capable of phagotrophy. Protists are a major source of mortality for both heterotrophic and autotrophic bacteria. They compete with herbivorous meso- and macro-zooplankton for all size classes of phytoplankton. Protist grazing may affect the rate of organic sinking flux from the euphotic zone. Protist excretions are an important source of remineralized nutrients, and of colloidal and dissolved trace metals such as iron, in aquatic systems. Work on predation by protists is being facilitated by methodological advances, e.g., molecular genetic analysis of protistan diversity and application of flow cytometry to study population growth and feeding rates. Examples of new research areas are studies of impact of protistan predation on the community structure of prey assemblages and of chemical communication between predator and prey in microbial food webs. This revised version was published online in August 2006 with corrections to the Cover Date.     

Quantitative food webs of dipteran leafminers and their parasitoids in Argentina

The quantitative structure of two host–parasitoid communities based on leaf-mining flies (Diptera, Agromyzidae) in Argentina is described. The two communities consisted of 29 and 27 hosts, 46 and 40 parasitoids, and 193 and 179 recorded host–parasitoid associations. Also, food webs were constructed for one community based solely on samples taken in the wet and dry seasons. Data were expressed as quantitative food webs, and the manner in which food web properties, such as connectance and compartmentalization, were influenced by sampling intensity was explored. The potential importance of indirect effects between hosts mediated by parasitoids (e.g. apparent competition) was assessed using quantitative parasitoid overlap diagrams. The studys results suggest that indirect effects are likely to be important in these highly connected communities. The limitations of the studys analysis, and how the conclusions can be tested experimentally, are discussed.     

Ecological and evolutionary implications of food subsidies from humans

Human activities are the main current driver of global change. From hunter‐gatherers through to Neolithic societies–and particularly in contemporary industrialised countries–humans have (voluntarily or involuntarily) provided other animals with food, often with a high spatio‐temporal predictability. Nowadays, as much as 30–40% of all food produced in Earth is wasted. We argue here that predictable anthropogenic food subsidies (PAFS) provided historically by humans to animals has shaped many communities and ecosystems as we see them nowadays. PAFS improve individual fitness triggering population increases of opportunistic species, which may affect communities, food webs and ecosystems by altering processes such as competition, predator–prey interactions and nutrient transfer between biotopes and ecosystems. We also show that PAFS decrease temporal population variability, increase resilience of opportunistic species and reduce community diversity. Recent environmental policies, such as the regulation of dumps or the ban of fishing discards, constitute natural experiments that should improve our understanding of the role of food supply in a range of ecological and evolutionary processes at the ecosystem level. Comparison of subsidised and non‐subsidised ecosystems can help predict changes in diversity and the related ecosystem services that have suffered the impact of other global change agents.     

Functional links and robustness in food webs

The robustness of ecosystems to species losses is a central question in ecology, given the current pace of extinctions and the many species threatened by human impacts, including habitat destruction and climate change. Robustness from the perspective of secondary extinctions has been addressed in the context of food webs to consider the complex network of species interactions that underlie responses to perturbations. In-silico removal experiments have examined the structural properties of food webs that enhance or hamper the robustness of ecosystems to species losses, with a focus on the role of hubs, the most connected species. Here we take a different approach and focus on the role of the connections themselves. We show that trophic links can be divided into functional and redundant based on their contribution to robustness. The analysis of empirical webs shows that hubs are not necessarily the most important species as they may hold many redundant links. Furthermore, the fraction of functional connections is high and constant across systems regardless of size and interconnectedness. The main consequence of this scaling pattern is that ecosystem robustness can be considerably reduced by species extinctions even when these do not result in any secondary extinctions. This introduces the possibility of tipping points in the collapse of ecosystems.     

Precision of herbivore tolerance experiments with imposed and natural damage

Tiffin and Inouye (2000) discussed the use of natural and imposed (controlled) damage in experiments of herbivore tolerance. They constructed a statistical model of the effect of herbivory on plant fitness, including damage level and an environmental factor as the independent factors, in which tolerance is defined as a slope of the regression line when damage level is regressed with plant fitness. They claim that while experiments with imposed damage are more accurate (i.e., they give a more correct estimate of tolerance), experiments with natural damage are more precise under a wide range of parameter values (i.e., tolerance estimates explain a larger part of variation in fitness). I show, however, that experiments with imposed damage are less precise only when an experimenter uses an experimental design that has weaker statistical power than in experiments with natural herbivory. The experimenter can nevertheless control the damage levels to optimize the experimental designs. For instance, when half of the experimental plants are left undamaged and the other half treated with maximal relevant damage level, experiments with imposed damage are almost always much more precise than experiments with natural damage.     

Connecting ecology to daily life: how students and teachers relate food webs to the food they eat

This study used sociocultural learning theory to better understand how middle and high school environmental science and biology students and pre- and in-service science teachers connect the daily life activity of eating to the food web model learned in school. We sought to understand how student and teacher perceptions of the environment and their experiences influenced their responses to interview questions regarding this topic. Findings, based on transcribed interviews with 54 study participants, indicate that three quarters of teachers and students were unable to connect the food they eat with ecosystem food webs. Even so, many respondents particularly those from elite public schools, did not demonstrate common food web misconceptions identified by other researchers, instead showing a sophisticated understanding of food web interactions. These findings indicate that even though participants were proficient in their school science understanding of food web interactions, they did not readily think about how their everyday out of school activities, like eating, relate to those interactions. This may be representative of a more general disconnect between formal ecology instruction and daily life activities. We provide several recommendations for how this disconnect can be remedied in our classrooms.     

Biomass diversity and stability of food webs in aquatic ecosystems

The diversity–stability hypothesis in ecology asserts that biodiversity begets stability of ecological systems. This hypothesis has been supported by field studies on primary producers in grasslands, in which the interaction between species is mostly competition. As to ecosystems with multitrophic predatory interaction, however, no definite consensus has been arrived at for the relation between trophic diversity and ecosystem stability. The stability index suitable to ecosystems with predatory interaction is given by MacArthurs idea of stability and its formulation by Rutledge et al. More suitable indices of stability (relative conditional entropy) are proposed in this study for the comparison of different ecosystems, and the validity of the diversity–stability hypothesis for food webs (networks of predation) with many trophic compartments in natural aquatic ecosystems is examined. Results reveal that an increase in the biomass diversity of trophic compartments causes an increase in the whole systemic stability of food webs in aquatic ecosystems. Hence, evidence of the whole systemic validity of the diversity–stability hypothesis for natural aquatic ecosystems with ubiquitous multitrophic predatory interaction was obtained for the first time.     

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.     

Global change alters the stability of food webs

Recent research has generally shown that a small change in the number of species in a food web can have consequences both for community structure and ecosystem processes. However ‘change’ is not limited to just the number of species in a community, but might include an alteration to such properties as precipitation, nutrient cycling and temperature. How such changes might affect species interactions is important, not just through the presence or absence of interactions, but also because the patterning of interaction strengths among species is intimately associated with community stability. Interaction strengths encompass such properties as feeding rates and assimilation efficiencies, and encapsulate functionally important information with regard to ecosystem processes. Interaction strengths represent the pathways and transfer of energy through an ecosystem. We review the best empirical data available detailing the frequency distribution of interaction strengths in communities. We present the underlying (but consistent) pattern of species interactions and discuss the implications of this patterning. We then examine how such a basic pattern might be affected given various scenarios of ‘change’ and discuss the consequences for community stability and ecosystem functioning.     

Biodiversity lessens the risk of cascading extinction in model food webs

Due to the complex interactions between species in food webs, the extinction of one species could lead to a cascade of further extinctions and hence cause dramatic changes in species composition and ecosystem processes. We found that the risk of additional species extinction, following the loss of one species in model food webs, decreases with the number of species per functional group. For a given number of species per functional group, the risk of further extinctions is highest when an autotroph is removed and lowest when a top predator is removed. In addition, stability decreases when the distribution of interaction strengths in the webs is changed from equal to skew (few strong and many weak links). We also found that omnivory appears to stabilize model food webs. Our results indicate that high biodiversity may serve as an insurance against radical ecosystem changes.     

Communities at the extreme: Aquatic food webs in desert landscapes

Studying food webs across contrasting abiotic conditions is an important tool in understanding how environmental variability impacts community structure and ecosystem dynamics. The study of extreme environments provides insight into community‐wide level responses to environmental pressures with relevance to the future management of aquatic ecosystems. In the western Lake Eyre Basin of arid Australia, there are two characteristic and contrasting aquatic habitats: springs and rivers. Permanent isolated Great Artesian Basin springs represent hydrologically persistent environments in an arid desert landscape. In contrast, hydrologically variable river waterholes are ephemeral in space and time. We comprehensively sampled aquatic assemblages in contrasting ecosystem types to assess patterns in community composition and to quantify food web attributes with stable isotopes. Springs and rivers were found to have markedly different invertebrate communities, with rivers dominated by more dispersive species and springs associated with species that show high local endemism. Qualitative assessment of basal resources shows autochthonous carbon appears to be a key basal resource in both types of habitat, although the particular sources differed between habitats. Food‐web variables such as trophic length, trophic breadth, and community isotopic niche size were relatively similar in the two habitat types. The basis for the similarity in food‐web structure despite differences in community composition appears to be broader isotopic niches for predatory invertebrates and fish in springs as compared with rivers. In contrast to published theory, our findings suggest that the food webs of the hydrologically variable river sites may show less dietary generalization and more compact food‐web modules than in springs.     

A size-constrained feeding-niche model distinguishes predation patterns between aquatic and terrestrial food webs

Understanding the formation of feeding links provides insights into processes underlying food webs. Generally, predators feed on prey within a certain body-size range, but a systematic quantification of such feeding niches is lacking. We developed a size-constrained feeding-niche (SCFN) model and parameterized it with information on both realized and non-realized feeding links in 72 aquatic and 65 terrestrial food webs. Our analyses revealed profound differences in feeding niches between aquatic and terrestrial predators and variation along a temperature gradient. Specifically, the predator–prey body-size ratio and the range in prey sizes increase with the size of aquatic predators, whereas they are nearly constant across gradients in terrestrial predator size. Overall, our SCFN model well reproduces the feeding relationships and predation architecture across 137 natural food webs (including 3878 species and 136,839 realized links). Our results illuminate the organisation of natural food webs and enables novel trait-based and environment-explicit modelling approaches.     

Scaling from individuals to networks in food webs

1.  Food webs, the set of predator–prey interactions in an ecosystem, are a prototypical complex system. Much research to date has concentrated on the use of models to identify and explain the key structural features which characterize food webs.
2.  These models often fall into two general categories: (i) phenomenological models which are built upon a set of heuristic rules in order to explain some empirical observation and (ii) population-level models in which interactions between individuals result in emergent properties for the food web. Both types of models have helped to uncover how food-web structure is a product of factors such as foraging behaviour, prey selection and species' body sizes.
3.  Historically, the two types of models have followed rather different approaches to the problem. Despite the apparent differences, the overlap between the two styles of models is substantial. Examples are highlighted here.
4.  By paying greater attention to both the similarities and differences between the two, we will be better able to demonstrate the ecological insights offered by phenomenological models. This will help us, for example, design experiments which could validate or refute underlying assumptions of the models. By linking models to data, scaling from individuals to networks, we will be closer to understanding the true origins of food-web structure.     

食物网稳定性机制研究进展:一个基于等级系统的框架

食物网理论沟通了群落生态学和生态系统生态学,将生物多样性和生态系统功能的研究统一起来,是理解生态系统运作机制的关键。自从1973年Robert May的经典研究引发著名的"复杂性-稳定性"论辩之后,人们认识到食物网的稳定性是其结构维持、功能发挥和动态演化的一个重要前提,并开始了对食物网稳定性机制的探索。早期研究主要关注只包含拓扑关系的定性食物网,但后来人们逐渐认识到相互作用强度的重要性,并提出了诸如自限性、弱相互作用、适应性捕食等一系列机制。本文系统梳理了过往研究中模块层面的各类稳定性机制和全网层面对各模块的整合机制,从而清晰地展示了"模块-全网"双层框架的全貌。通过在其基础上的扩展,进而提出了一个基于等级系统的食物网稳定性框架,并从动力学和能量学角度,对各层级内部的稳定性机制以及层级之间的关系进行了探讨,以期为建立普适的食物网稳定性理论提供一些思路。未来的研究方向包括：①将稳定性机制的研究从食物网扩展到更一般的生态网络;②综合考虑生物物理要素、动力学稳定性、系统对能流功率的追求、环境的平稳程度、演化历史等影响因素,从而得到关于食物网结构和动态的更为深刻的认识。     

Cascading extinctions and community collapse in model food webs

Species loss in ecosystems can lead to secondary extinctions as a result of consumer–resource relationships and other species interactions. We compare levels of secondary extinctions in communities generated by four structural food-web models and a fifth null model in response to sequential primary species removals. We focus on various aspects of food-web structural integrity including robustness, community collapse and threshold periods, and how these features relate to assumptions underlying different models, different species loss sequences and simple measures of diversity and complexity. Hierarchical feeding, a fundamental characteristic of food-web structure, appears to impose a cost in terms of robustness and other aspects of structural integrity. However, exponential-type link distributions, also characteristic of more realistic models, generally confer greater structural robustness than the less skewed link distributions of less realistic models. In most cases for the more realistic models, increased robustness and decreased levels of web collapse are associated with increased diversity, measured as species richness S, and increased complexity, measured as connectance C. These and other results, including a surprising sensitivity of more realistic model food webs to loss of species with few links to other species, are compared with prior work based on empirical food-web data.     

Arthropod diversity and allochthonous-based food webs on tiny oceanic islands

Very small islands, on the order of a few hundred square metres in area, have rarely been the focus of ecological investigations. I sampled nine such islands in the central Exumas, Bahamas for arthropod species abundance and diversity using a combination of pitfall traps, pan traps and sticky traps. Three islands had no terrestrial vegetation, three islands contained only Sesuvium portulacastrum L., a salt‐tolerant perennial that had been experimentally introduced 10 years ago, and three islands supported one or two naturally occurring plant species. A relatively diverse arthropod assemblage was discovered, including representatives of 10 different orders of Crustacea and Insecta. Land hermit crabs were the most abundant crustaceans, and dipterans were the most abundant and speciose insects. Two of the most common insects were previously undescribed species. Measures of arthropod species abundance and diversity were not significantly different for vegetated vs. non‐vegetated islands. All 10 orders were present on bare islands, and nine of them were present on vegetated islands. Measures of arthropod species abundance and diversity were positively associated with island area, and negatively associated with distance from the nearest large island. Hypothesized food webs consist of several trophic levels and have strong allochthonous inputs. Tiny islands such as these hold insights into early successional processes and the base of insular food webs.     

A role for brain size and cognition in food webs

Predators tend to be large and mobile, enabling them to forage in spatially distinct food web compartments (e.g. littoral and pelagic aquatic macrohabitats). This feature can stabilise ecosystems when predators are capable of rapid behavioural response to changing resource conditions in distinct habitat compartments. However, what provides this ability to respond behaviourally has not been quantified. We hypothesised that predators require increased cognitive abilities to occupy their position in a food web, which puts pressure to increase brain size. Consistent with food web theory, we found that fish relative brain size increased with increased ability to forage across macrohabitats and increased relative trophic positions in a lacustrine food web, indicating that larger brains may afford the cognitive capacity to exploit various habitats flexibly, thus contributing to the stability of whole food webs.     

Resistance and tolerance to herbivory in Salix cordata are affected by different environmental factors

Abstract.  1. Effects of sand burial and nutrients on the ability of sand-dune willow ( Salix cordata ) to tolerate or resist herbivory by the beetle Altica subplicata were evaluated in field experiments.
2. To assess tolerance, all combinations of sand burial (none, 50%), nutrients (presence, absence), and beetles (presence, absence) were applied to caged plants and growth responses to herbivory were measured. Sand burial increased plant growth rate, but decreased S. cordata 's tolerance to herbivory. Although nutrients increased growth, tolerance to herbivory was unaffected.
3. To assess resistance, plants were exposed to all combinations of sand burial and nutrients, and then to natural beetle colonisation. The presence of nutrients, but not sand burial, significantly increased the percentage of plants with beetles, for both adults and larvae. This decreased resistance to beetles of plants grown with added nutrients occurred only in the absence of sand burial.
4. The performance and preference of beetles were examined in laboratory experiments. Larvae developed faster and had increased pupation success on plants with nutrients added. Beetles also showed a marginally significant feeding preference for leaves grown with added nutrients. Thus, S. cordata tolerance to herbivory was affected by sand burial, whereas resistance, preference, and performance were affected by nutrient level.