Inferring predator–prey interactions in food webs

Gradients in elevation are increasingly used to investigate how species respond to changes in local climatic conditions. Whilst many studies have shown elevational patterns in species richness and turnover, little is known about how food web structure is affected by elevation. Contrasting responses of predator and prey species to elevation may lead to changes in food web structure. We investigated how the quantitative structure of a herbivore-parasitoid food web changes with elevation in an Australian subtropical rain forest. On four occasions, spread over 1 year, we hand-collected leaf miners at twelve sites, along three elevational gradients (between 493 m and 1159 m a.s.l). A total of 5030 insects, including 603 parasitoids, were reared, and summary food webs were created for each site. We also carried out a replicated manipulative experiment by translocating an abundant leaf-mining weevil Platynotocis sp., which largely escaped parasitism at high elevations (≥ 900 m a.s.l.), to lower, warmer elevations, to test if it would experience higher parasitism pressure. We found strong evidence that the environmental change that occurs with increasing elevation affects food web structure. Quantitative measures of generality, vulnerability and interaction evenness decreased significantly with increasing elevation (and decreasing temperature), whilst elevation did not have a significant effect on connectance. Mined plant composition also had a significant effect on generality and vulnerability, but not on interaction evenness. Several relatively abundant species of leaf miner appeared to escape parasitism at higher elevations, but contrary to our prediction, Platynotocis sp. did not experience greater levels of parasitism when translocated to lower elevations. Our study indicates that leaf-mining herbivores and their parasitoids respond differently to environmental conditions imposed by elevation, thus producing structural changes in their food webs. Increasing temperatures and changes in vegetation communities that are likely to result from climate change may have a restructuring effect on host-parasitoid food webs. Our translocation experiment, however, indicated that leaf miners currently escaping parasitism at high elevations may not automatically experience higher parasitism under warmer conditions and future changes in food web structure may depend on the ability of parasitoids to adapt to novel hosts.

Consequent Effects of Parasitism on Population Dynamics, Food Webs, and Human Health Under Climate Change

To test the effects of temperature on food web structure and productivity, Mary O'Connor (above, checking temperatures) and colleagues placed five microcosms of food webs (shielded from full sunlight and UV) in eight independent water tables, each filled with a temperature-conditioned water bath.

Food webs from the Lerderderg River, an intermittent stream in Victoria, Australia, were compiled with the aim of examining changes in food web structure in a highly variable habitat. Emphasis was placed on a high degree of taxonomic precision. Spatial and temporal variation in the food webs was assessed by partitioning the study area into three sites, located ≈1.5 km apart along the river. Sites differed in overall stream width and the length of the low streamflow period during summer. Three separate webs for each site were compiled for four different times of the year. Relatively little spatial variation in community structure was observed. In contrast, temporal variation was considerable, with species composition and the number of species in the community changing considerably over the year. The number of species increased dramatically as the period of constant streamflow lengthened. Detritivores dominated the community, both in terms of species and individual numbers. The proportion of predators in the community increased slightly by the end of the year, suggesting that recolonization of the community by predators lags behind that of detritivores. The increase in the number of predator species also resulted in an increase in the mean food chain length through the year. Patterns observed in the food webs tended to fall within the range of values reported from several previous studies, suggesting that underlying constraints may structure certain aspects of food webs. However, the constancy of certain food web statistics was attributable either to methodological decisions made during compilation, or to an inherent property of the statistic itself. The potential sensitivity of several food web statistics to the methodology used to compile a food web render between—web comparisons difficult due to the confounding effects of methodology. This suggests that comparisons between food webs should be restricted to webs derived from similar habitats using a comparable methodology.

With an increasing reliance on nuclear power as a low-carbon energy source, understanding the pathways and dynamics of planned or unplanned radionuclide releases to the environment and subsequent transfer within food webs is essential for assessing environmental risks. This essay explores the mechanisms by which radionuclides accumulate in aquatic organisms and transfer through food webs. Factors influencing bioaccumulation, including environmental parameters, radionuclide properties, and organismal characteristics, are reviewed. Additionally, this essay investigates the role of food web structure in mediating the transfer of radionuclides, highlighting the influence of climate-induced changes in food web structure. The findings underline the complexity of radionuclide contaminant pathways in aquatic ecosystems and emphasize the need for research on the impact of shifting food web structures due to climate change on radionuclide transfer. This understanding is crucial for assessing and mitigating future environmental risks associated with nuclear energy production.

The influence of biotic and abiotic factors on species interactions and overall community structure has long interested ecologists. Despite a legacy of interest, there is still ambiguity into the role of biotic and abiotic factors due to highly nuanced, complex networks of interactions that are difficult to comprehend. Yet, understanding how such nuances is an essential goal to determine how they affect population and community structure. Thus, the goal of my dissertation was to understand how multiple biotic and abiotic mechanisms alter interactions among larval stages of two pond-breeding salamanders. Larval stages of pond-breeding salamanders represent an excellent system for understanding how species interactions vary along abiotic and biotic gradients. Intra-and interspecific interactions are frequently determined by size differences among individuals, where larger larvae are predators of smaller larvae and can out-compete them for shared resources. However, when size differences are minimized, only competition occurs. Such conjoined competition and predation is termed intraguild predation, and is a common interaction in many taxa. The factors that determine size differences among individuals (both within and between species) are critical towards to determining both the type of interaction, as well as the strength of such interactions. The focal species I used were the ringed salamander (Ambystoma annulatum) and spotted salamander (A. maculatum). The former breeds earlier than the latter, creating a larval size advantage which results in predation as the dominant interaction between species. However, factors that influence growth rates of ringed salamanders could result in minimized size differences, resulting in a change to the strength or type of interaction that occurs. For my dissertation, I experimentally investigated three different processes that were expected to affect the relative importance of predation and competition: density dependence, food web structure, and phenological shifts. In my first chapter, I tested whether the density of ringed salamanders influenced their growth rates to such a degree that the interaction type with spotted salamanders would switch from predation to competition. I found that increased intraspecific competition in ringed salamanders reduced their body size and increased their larval period length. However, intraspecific competition did not reduce their size to such a degree that predation on spotted salamanders was precluded. Spotted salamanders showed decreased survival and increased size at higher predator densities, indicative of thinning effects. The period of overlap in ponds also increased at higher predator densities, resulting in a larger temporal window for interactions to occur. In my second chapter, I tested how six different top predator food webs would influence intraguild predation between ringed and spotted salamanders. I also tested whether food web configuration would be simultaneously impacted by increased habitat complexity. I found that ringed salamander body size and survival were unaffected by habitat complexity, and that only certain combinations of predators affected these demographic rates. Spotted salamander body size and survival showed positive and negative relationships with ringed salamander survival, but the strength of these relationships varied depending on the predator and habitat complexity treatment. Thus, pairwise interactions may not exemplify typical interactions when embedded in more complex food webs with other predators. For my third chapter, I investigated whether phenological shifts in both the ringed and spotted salamanders, simultaneous to density dependence in the ringed salamander would influence the type and strength of their interactions. I found ringed salamander survival varied with phenological shifts but only when at high intraspecific densities. Spotted salamanders were relatively unaffected by phenological shifts, and that their interactions were, similar to the previous chapters, influenced primarily by survival of ringed salamanders. As phenological shifts are predicted for many species with climate change, this study highlights that not all species interactions will be subsequently affected, and that other underlying factors (e.g. density dependence) may be more important. Thus, the most important findings of my dissertation include 1) predator density can be a dominant factor in species interactions, 2) pairwise interactions may change when embedded in different habitats or food webs in non-intuitive ways, and 3) simultaneously testing multiple mechanisms can elicit a greater understanding of the relative importance of different ecological processes.

Assessing changes in food-web structure provides a useful monitoring tool for gauging the resilience of ecosystems in the face of climatic impacts. We consider the ecological resilience of a large estuarine lake (St Lucia Estuary, South Africa) in the wake of an extreme climatic event (prolonged drought). Using carbon and nitrogen stable isotopes, food-web structure was assessed at five sites across the estuary during the winter and spring of 2013. Sampling occurred approximately three years after heavy rains flooded the system and returned it to a relatively diluted state following an almost decade-long drought that decimated food webs in the upper parts of the estuary due to hypersalinity effects. Comparisons of niche width and variance of consumer food webs among sites revealed a general homogenisation of food webs across the entire system, contrasting with the spatial differentiation of food webs documented during the drought phase. Our results indicate that the estuary is able to maintain ecological resilience at the whole-system level in the face of an extreme drought. This is likely facilitated by source pools of species residing in the relatively stable lower estuary, which are able to rapidly recolonise areas denuded by drought in the upper estuary.

Simple SummaryIn terrestrial natural ecosystems, more complex and diverse networks of plant–insect primary consumers and their predators are often more productive, stable, and resilient. Plant diversity often positively correlates to the diversity of phytophagous insects and their natural enemies generating multitrophic interactions with changing outcomes (bottom-up effects). The use of cover crops can promote natural enemy populations and their temporal synchronization with a target pest, resulting in greater pest control. Therefore, changes in the habitat conditions can alter food webs. In agroecosystems, characteristics of the food trophic webs, as connectance, measured as the proportion of realized links in the network, could be linked to the efficiency of pest control. In this study, we evaluated how the use of oat cover crops affects composition and structure in the aphid–parasitoid–hyperparasitoid food webs of plum orchards with different habitat management contexts: plums with inter-rows of oats as a cover crop (OCC) and plums with inter-rows with spontaneous vegetation (SV). Quantitative food web metrics differed significantly among treatments showing a higher generality, vulnerability, interaction evenness, and linkage density in SV, while OCC presented a higher degree of specialization.By increasing plant diversity in agroecosystems, it has been proposed that one can enhance and stabilize ecosystem functioning by increasing natural enemies’ diversity. Food web structure determines ecosystem functioning as species at different trophic levels are linked in interacting networks. We compared the food web structure and composition of the aphid– parasitoid and aphid-hyperparasitoid networks in two differentially managed plum orchards: plums with inter-rows of oats as a cover crop (OCC) and plums with inter-rows of spontaneous vegetation (SV). We hypothesized that food web composition and structure vary between OCC and SV, with network specialization being higher in OCC and a more complex food web composition in SV treatment. We found a more complex food web composition with a higher species richness in SV compared to OCC. Quantitative food web metrics differed significantly among treatments showing a higher generality, vulnerability, interaction evenness, and linkage density in SV, while OCC presented a higher degree of specialization. Our results suggest that plant diversification can greatly influence the food web structure and composition, with bottom-up effects induced by plant and aphid hosts that might benefit parasitoids and provide a better understanding of the activity, abundance, and interactions between aphids, parasitoids, and hyperparasitoids in plum orchards.

In May 1993 and 1994, northern pike (Esox lucius) were added to eutrophic Lake 227 and oligotrophic Lake 110 at the Experimental Lakes Area in northwestern Ontario. Both lakes were previously dominated by cyprinids, and northern pike additions significantly decreased cyprinid densities in both lakes. Food web relationships were determined pre- and post-manipulation using stable carbon (δ13C) and nitrogen (δ15N) isotope analyses of zooplankton, benthic invertebrates, and fishes. In Lake 110, fathead minnow (Pimephales promelas) shifted from a zooplanktivorous to a zoobenthivorous diet as indicated by a shift in isotopic composition to more depleted δ15N and enriched δ13C values after northern pike additions. It was not possible to direct predator-induced shifts in cyprinid diets in Lake 227. Concentrations of mercury and organochlorines (sum of polychlorinated biphenyls (ΣPCB), dichlorodiphenyltrichloroethane (ΣDDT), and hexachlorocyclohexane (ΣHCH)) in these fish did not change markedly after northern pike introductions despite the shifts in diet for fathead minnow from Lake 110. However, concentrations of all contaminants were lower in biota from eutrophic Lake 227 when compared with Lake 110, which is consistent with previous studies. Our results indicate that nutrient availability, rather than short-term changes in food web structure, determined contaminant concentrations in fish and other biota from these lakes.

In recent years, sea ice cover along coasts of East Antarctica has tended to increase. To understand ecological implications of these environmental changes, we studied benthic food web structure on the coasts of Adélie Land during an event of unusually high sea ice cover (i.e. two successive austral summers without seasonal breakup). We used integrative trophic markers (stable isotope ratios of carbon, nitrogen and sulfur) to build ecological models and explored feeding habits of macroinvertebrates. In total, 28 taxa spanning most present animal groups and functional guilds were investigated. Our results indicate that the absence of seasonal sea ice breakup deeply influenced benthic food webs. Sympagic algae dominated the diet of many key consumers, and the trophic levels of invertebrates were low, suggesting omnivore consumers did not rely much on predation and/or scavenging. Our results provide insights about how Antarctic benthic consumers, which typically live in an extremely stable environment, might adapt their feeding habits in response to sudden changes in environmental conditions and trophic resource availability. They also show that local and/or global trends of sea ice increase in Antarctica have the potential to cause drastic changes in food web structure, and therefore to impact benthic communities.

1. Using a subtidal marine food web as a model system, we examined how food chain length (predators present or absent) and the prevalence of omnivory influenced temporal stability (and its components) of herbivores and plants. We held the density of top predators constant but manipulated their identity to generate a gradient in omnivory prevalence. 2. We measured temporal stability as the inverse of the coefficient of variation of abundance over time. Predators and omnivory could influence temporal stability through effects on abundance (the 'abundance' effect), summed variance across taxa (the 'portfolio effect') or summed covariances among taxa (the 'covariance effect'). 3. We found that increasing food chain length by predator addition destabilized aggregate herbivore abundance through their cascading effects on abundances. Thus, predators destabilized herbivores through the overyielding effect. We also found that the stability of herbivore abundance and microalgae declined with increasing prevalence of omnivory among top predators. Aggregate macroalgae was not affected, but the stability of one algal taxon increased with the prevalence of omnivory. 4. Our results suggest that herbivores are more sensitive than plants to changes in food web structure because of predator additions by invasion or deletions such as might occur via harvesting and habitat loss.

Mathematical models predict that species interactions such as competition and predation can generate chaos. However, experimental demonstrations of chaos in ecology are scarce, and have been limited to simple laboratory systems with a short duration and artificial species combinations. Here, we present the first experimental demonstration of chaos in a long-term experiment with a complex food web. Our food web was isolated from the Baltic Sea, and consisted of bacteria, several phytoplankton species, herbivorous and predatory zooplankton species, and detritivores. The food web was cultured in a laboratory mesocosm, and sampled twice a week for more than 2,300 days. Despite constant external conditions, the species abundances showed striking fluctuations over several orders of magnitude. These fluctuations displayed a variety of different periodicities, which could be attributed to different species interactions in the food web. The population dynamics were characterized by positive Lyapunov exponents of similar magnitude for each species. Predictability was limited to a time horizon of 15-30 days, only slightly longer than the local weather forecast. Hence, our results demonstrate that species interactions in food webs can generate chaos. This implies that stability is not required for the persistence of complex food webs, and that the long-term prediction of species abundances can be fundamentally impossible.

Kuroshio and Oyashio Currents

Current global change scenarios demand knowledge on how anthropogenic impacts affect ecosystem functioning through changes in food web structure. Frugivorous mesocarnivores are a key link in trophic cascades because, while their abundance and behaviour are usually controlled by apex predators, they can provide high‐quality seed‐dispersal services to plant communities. Thus, the recent rewilding of large carnivores worldwide can trigger cascading effects for plants. We investigated the top‐down effects of an apex predator (Iberian lynx Lynx pardinus) on seed‐dispersal services mediated by two mesocarnivore species (red fox Vulpes vulpes and stone marten Martes foina) at the plant community level by comparing areas with and without lynx in a Mediterranean mountain range in Southern Spain. We collected scats of mesocarnivores (n = 1575) to assess frugivory and seed dispersal of 15 plant species over two consecutive fruiting seasons and two habitat types (open and forest). Specifically, we assessed the effect of lynx presence on (i) seed occurrence and fleshy‐fruit biomass per scat, (ii) number of scats containing seeds and (iii) diversity of dispersed seeds. The quantity and diversity of dispersed seeds drastically decreased under predation risk for both mesocarnivore's species. Seed dispersal by stone martens was negatively affected by the presence of lynx, with a marked reduction in the number of scats with seeds (93%) and the diversity of dispersed seeds (46%). Foxes dispersed 68% fewer seeds in open habitats when coexisting with lynx, probably leading to differential contributions to seed‐dispersal effectiveness among habitats. Our study reveals a novel trophic cascade from apex predators to plant communities. The behavioural responses of frugivorous mesocarnivores to predation risk and the reduction in the intensity of their faecal deposition pattern are probably related to their lower abundance when co‐occurring with apex predators. While rewilding apex predators is a successful conservation tool, attention should be paid to cascading effects across food webs, particularly where frugivore megafauna is missing and mesocarnivores provide unique services to plants. Read the free Plain Language Summary for this article on the Journal blog.

Aquatic ecosystems support size structured food webs, wherein predator‐prey body sizes span orders of magnitude. As such, these food webs are replete with extremely generalized feeding strategies, especially among the larger bodied, higher trophic position taxa. The movement scale of aquatic organisms also generally increases with body size and trophic position. Together, these body size, mobility, and foraging relationships suggest that organisms lower in the food web generate relatively distinct energetic pathways by feeding over smaller spatial areas. Concurrently, the potential capacity for generalist foraging and spatial coupling of these pathways often increases, on average, moving up the food web toward higher trophic levels. We argue that these attributes make for a food web architecture that is inherently ‘adaptive’ in its response to environmental conditions. This is because variation in lower trophic level dynamics is dampened by the capacity of predators to flexibly alter their foraging behavior. We argue that empirical, theoretical, and applied research needs to embrace this inherently adaptive architecture if we are to understand the relationship between structure and function in the face of ongoing environmental change. Toward this goal, we discuss empirical patterns in the structure of lake food webs to suggest that ecosystems change consistently, from individual traits to the structure of whole food webs, under changing environmental conditions. We then explore an empirical example to reveal that explicitly unfolding the mechanisms that drive these adaptive responses offers insight into how human‐driven impacts, such as climate change, invasive species, and fisheries harvest, ought to influence ecosystem structure and function (e.g., stability, secondary productivity, maintenance of major energy pathways). We end by arguing that such a directed food web research program promises a powerful across‐scale framework for more effective ecosystem monitoring and management.