Trophic and non‐trophic interactions influence the mechanisms underlying biodiversity–ecosystem functioning relationships under different abiotic conditions

Plant diversity effects on ecosystem functioning usually have been studied from a plant perspective. However, the mechanisms underlying biodiversity–ecosystem functioning relationships may also depend on positive or negative interactions between plants and other biotic and abiotic factors, which remain poorly understood. Here we assessed whether plant–herbivore and/or plant–detritivore interactions modify the biodiversity–ecosystem functioning relationship and the mechanisms underlying biodiversity effects, including complementarity and selection effects, biomass allocation, vertical distribution of roots, and plant survival using a microcosm experiment. We also evaluated to what extent trophic and non‐trophic interactions are affected by abiotic conditions by studying drought effects. Our results show that biotic and abiotic conditions influence the shape of the biodiversity–ecosystem function relationship, varying from hump‐shaped to linear. For instance, total biomass increased linearly with plant richness in the presence of detritivores, but not in the absence of detritivores. Moreover, detritivore effects on belowground plant productivity were highly context dependent, varying in the presence of herbivores. Plant interactions with soil biota, especially with herbivores, influenced the mechanisms underlying diversity effects. Herbivores increased plant complementarity and modified biomass allocation and vertical distribution of roots. Furthermore, biotic–abiotic interactions influenced plant productivity differently across plant functional groups. Our findings emphasize the importance of complex biotic interactions underlying biodiversity effects, and that these biotic interactions may change with abiotic conditions. Despite minor changes in productivity in the short‐term, soil biota‐induced changes in plant–plant interactions and plant survival are likely to have significant long‐term consequences for ecosystem functioning. Considering the context‐dependency of multichannel interactions may contribute to reconciling differences among observed patterns in biodiversity studies. Further, abiotic conditions modified the effects of biotic interactions, suggesting that changes in environmental conditions may not only affect ecosystems directly, but also change the biotic composition of and dynamics within ecosystems.     

The influence of biotic interactions on soil biodiversity

Belowground communities usually support a much greater diversity of organisms than do corresponding aboveground ones, and while the factors that regulate their diversity are far less well understood, a growing number of recent studies have presented data relevant to understanding how these factors operate. This review considers how biotic factors influence community diversity within major groups of soil organisms across a broad spectrum of spatial scales, and addresses the mechanisms involved. At the most local scale, soil biodiversity may potentially be affected by interactions within trophic levels or by direct trophic interactions. Within the soil, larger bodied invertebrates can also influence diversity of smaller sized organisms by promoting dispersal and through modification of the soil habitat. At larger scales, individual plant species effects, vegetation composition, plant species diversity, mixing of plant litter types, and aboveground trophic interactions, all impact on soil biodiversity. Further, at the landscape scale, soil diversity also responds to vegetation change and succession. This review also considers how a conceptual understanding of the biotic drivers of soil biodiversity may assist our knowledge of key topics in community and ecosystem ecology, such as aboveground–belowground interactions, and the relationship between biodiversity and ecosystem functioning. It is concluded that an improved understanding of what drives the diversity of life in the soil, incorporated within appropriate conceptual frameworks, should significantly aid our understanding of the structure and functioning of terrestrial communities.     

Biodiversity, ecosystem functioning and food webs in fresh waters: assembling the jigsaw puzzle

1. Dramatic advances have been made recently in the study of biodiversity–ecosystem functioning (B-EF) relations and food web ecology. These fields are now starting to converge, and this fusion has the potential to improve our understanding of how environmental stressors modulate ecosystem processes and the supply of 'goods and services'.
2. Food web structure and dynamics can exert particularly strong influences on B-EF relations in fresh waters, as consumer–resource interactions (e.g. trophic cascades) are often more important than horizontal interactions within trophic levels. For instance, many freshwater food webs are size structured, with large organisms tending to occupy the higher trophic levels and often exerting powerful effects on ecosystem processes. However, because they are also vulnerable to perturbations, non-random losses of these large taxa can alter both food web structure and ecosystem functioning profoundly.
3. Recently, the focus of food web research has shifted away from exploring patterns, towards developing an understanding of processes (e.g. quantifying fluxes of individuals, biomass, energy, nutrients) and how the two interact. Many of the best-characterized food webs are from fresh waters, and these ecosystems are now being used to address some of the shortcomings of earlier B-EF studies. I have identified several key gaps in our current knowledge and highlighted potentially fruitful avenues of future B-EF and food web research.
4. A major challenge for this newly emerging research is to place it within a unified theoretical framework. The application of metabolic theory and ecological stoichiometry may help to achieve this goal by considering biological systems within the constraints imposed upon them by physical and chemical laws.     

A synthesis of subdisciplines: predator–prey interactions, and biodiversity and ecosystem functioning

The last 15 years has seen parallel surges of interest in two research areas that have rarely intersected: biodiversity and ecosystem functioning (BEF), and multispecies predator–prey interactions (PPI). Research addressing role of biodiversity in ecosystem functioning has focused primarily on single trophic‐level systems, emphasizing additive effects of diversity that manifest through resource partitioning and the sampling effect. Conversely, research addressing predator–prey interactions has focused on two trophic‐level systems, emphasizing indirect and non‐additive interactions among species. Here, we use a suite of consumer‐resource models to organize and synthesize the ways in which consumer species diversity affects the densities of both resources and consumer species. Specifically, we consider sampling effects, resource partitioning, indirect effects caused by intraguild interactions and non‐additive effects. We show that the relationship between consumer diversity and the density of resources and consumer species are broadly similar for systems with one vs. two trophic levels, and that indirect and non‐additive interactions generally do little more than modify the impacts of diversity established by the sampling effect and resource partitioning. The broad similarities between systems with one vs. two trophic levels argue for greater communication between researchers studying BEF, and researchers studying multispecies PPI.     

Temperature and trophic structure are driving microbial productivity along a biogeographical gradient

Temperature is known to influence ecosystem processes through its direct effect on biological rates such as respiration and nutrient cycling. These changes can then indirectly affect ecologically processes by altering trophic dynamics, the persistence of a species in a given environment, and, consequently, its distribution. However, it is not known if this direct effect of temperature on biological rates is singularly the most important factor for the functioning of ecosystems, or if trophic structure and the adaptation of a species to the local environment also play an essential role. Understanding the relative importance of these factors is crucial for predicting the impact that climate change will have on species and ecosystems. To achieve a more complete understanding of the impact of changing temperatures, it is necessary to integrate perspectives from biogeography, such as the influences of species distribution and local adaptation, with ecosystem and community ecology. By using the microbial community inhabiting the water‐filled leaves of Sarracenia purpurea, we tested the importance of temperature, trophic structure, and local adaptation on ecosystem functioning. We accomplished this by collecting communities along a natural temperature gradient and maintaining these communities in a common garden, factorial experiment. To test for the importance of local adaptation and temperature, the origin of each community was crossed with the temperature from each site. Additionally, to test the importance of top‐down trophic regulation for ecosystem functioning, the presence of the mosquito larvae top predator was manipulated. We found that temperature has a greater effect on ecosystem functioning than origin, and that top‐down trophic regulation increased with temperature. Our results emphasize the synergistic effects of temperature and biotic interactions when predicting the consequences of global warming on ecosystem functioning.     

Community and ecosystem responses to recent climate change

There is ample evidence for ecological responses to recent climate change. Most studies to date have concentrated on the effects of climate change on individuals and species, with particular emphasis on the effects on phenology and physiology of organisms as well as changes in the distribution and range shifts of species. However, responses by individual species to climate change are not isolated; they are connected through interactions with others at the same or adjacent trophic levels. Also from this more complex perspective, recent case studies have emphasized evidence on the effects of climate change on biotic interactions and ecosystem services. This review highlights the ‘knowns’ but also ‘unknowns’ resulting from recent climate impact studies and reveals limitations of (linear) extrapolations from recent climate-induced responses of species to expected trends and magnitudes of future climate change. Hence, there is need not only to continue to focus on the impacts of climate change on the actors in ecological networks but also and more intensively to focus on the linkages between them, and to acknowledge that biotic interactions and feedback processes lead to highly complex, nonlinear and sometimes abrupt responses.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

气候变化与生物多样性之间的复杂关系和反馈机制

气候变化与生物多样性丧失是人类社会正在经历的两大变化。气候变化影响生物多样性的方方面面, 是导致生物多样性丧失的一个主要驱动因子; 反过来, 生物多样性丧失会加剧气候变化。因此, 阻止甚至扭转气候变化和生物多样性丧失是当前人类社会亟需解决的全球性问题,但我们对气候变化与生物多样性之间的复杂关系和反馈机制尚缺乏清晰认识。本文总结了近年气候变化与生物多样性变化的研究进展, 重点概述了不同组织层次、空间尺度和维度的生物多样性对气候变化的响应和反馈等相关领域的研究进展和存在的主要问题。结果发现多数研究关注气候变化对生物多样性的直接影响, 涉及到生物多样性的不同组织层次、维度和营养级, 但针对气候变化间接影响的研究仍然较少, 机理研究同样需要加强; 生物多样性对生态系统功能影响的环境依赖和尺度推演、生物多样性对生态系统多功能性的作用机理和量化方法是当前研究面临的挑战; 生物多样性对生态系统响应气候变化的作用机制尚无统一的认识; 生物多样性对气候变化的正、负反馈效应是国内外研究的盲点。最后, 本文展望了未来发展方向和需要解决的关键科学问题, 包括多因子气候变化对生物多样性的影响; 减缓和适应气候变化的措施如何惠益于生物多样性保护; 生物多样性与生态系统功能的理论如何应用到现实世界; 生物多样性保护对实现碳中和目标的贡献。     

Spatial stoichiometry: cross‐ecosystem material flows and their impact on recipient ecosystems and organisms

Cross‐ecosystem material flows, in the form of inorganic nutrients, detritus and organisms, spatially connect ecosystems and impact food web dynamics. To date research on material flows has focused on the impact of the quantity of these flows and largely ignored their elemental composition, or quality. However, the ratios of elements like carbon, nitrogen and phosphorus can influence the impact material flows have on food web interactions through stoichiometric mismatches between resources and consumers. The type and movement of materials likely vary in their ability to change stoichiometric constraints within the recipient ecosystem and materials may undergo changes in their own stoichiometry during transport. In this literature review we evaluate the importance of cross‐ecosystem material flows within the framework of ecological stoichiometry. We explore how movement in space and time impacts the stoichiometry of material flow, as these transformations are essential to consider when assessing the ability of these flows to impact food web productivity and ecosystem functioning. Our review suggests that stoichiometry of cross‐ecosystem material flows are highly dynamic and undergo changes during transport across the landscape or from human influence. These material flows can impact recipient organisms if they change stoichiometry of the abiotic medium, or provide resources that have a different stoichiometry to in situ resources. They might also alter consumer excretion rates, in turn altering the availability of nutrients in the recipient ecosystem. These alterations in stoichiometric constraints of recipient organisms can have cascading trophic effects and shape food web dynamics. We highlight significant gaps in the literature and suggest new avenues for research that explore how cross‐ecosystem material flows impact recipient ecosystems when considering differences in stoichiometric quality, their movement through the landscape and across ecosystem boundaries, and the nutritional constraints of the recipient organisms.     

Integrating ecosystem functions into restoration ecology—recent advances and future directions

Including ecosystem functions into restoration ecology has been repeatedly suggested, yet there is limited evidence that this is taking place without bias to certain habitats, species, or functions. We reviewed the inclusion of ecosystem functions in restoration and potential relations to habitats and species by extracting 224 publications from the literature (2004–2013). Most studies investigated forests, fewer grasslands or freshwaters, and fewest wetlands or marine habitats. Of all studies, 14% analyzed only ecosystem functions, 44% considered both biotic composition and functions, 42% exclusively studied the biotic component, mostly vascular plants, more rarely invertebrates or vertebrates, and least often microbes. Most studies investigating ecosystem functions focused on nutrient cycling (26%), whereas productivity (18%), water relations (16%), and geomorphological processes (14%) were less covered; carbon sequestration (10%), decomposition (6%), and trophic interactions (6%) were rarely studied. Monitoring of ecosystem functions was common in forests and grasslands, but the functions considered depended on the study organisms. These associations indicate research opportunities for certain habitats, species, and functions. Overall, the call to include ecosystem functions in restoration has been heard; however, a lack of clarity about the ecosystem functions to be included and deficits of feasible field methods are major obstacles for a functional approach. Restoration ecology should learn from recent advances in rapid assessment of ecosystem functions, and by a closer integration with biodiversity–ecosystem functioning research. Not all functions need to be measured in all ecosystems, but more functions than the few commonly addressed would improve the understanding of restored ecosystems.     

Interspecific behavioural interference and range dynamics: current insights and future directions

Novel biotic interactions in shifting communities play a key role in determining the ability of species' ranges to track suitable habitat. To date, the impact of biotic interactions on range dynamics have predominantly been studied in the context of interactions between different trophic levels or, to a lesser extent, exploitative competition between species of the same trophic level. Yet, both theory and a growing number of empirical studies show that interspecific behavioural interference, such as interspecific territorial and mating interactions, can slow down range expansions, preclude coexistence, or drive local extinction, even in the absence of resource competition. We conducted a systematic review of the current empirical research into the consequences of interspecific behavioural interference on range dynamics. Our findings demonstrate there is abundant evidence that behavioural interference by one species can impact the spatial distribution of another. Furthermore, we identify several gaps where more empirical work is needed to test predictions from theory robustly. Finally, we outline several avenues for future research, providing suggestions for how interspecific behavioural interference could be incorporated into existing scientific frameworks for understanding how biotic interactions influence range expansions, such as species distribution models, to build a stronger understanding of the potential consequences of behavioural interference on the outcome of future range dynamics.     

Asymmetrical food web responses in trophic-level richness, biomass, and function following lake acidification

We tested for disproportional changes in annual and seasonal species richness and biomass among five trophic levels (phytoplankton, herbivorous, omnivorous, and carnivorous zooplankton, and fish) as well as altered trophic structure and ecosystem function following the 5-year experimental acidification of Little Rock Lake (Wisconsin, USA) from pH 6.1 to 4.7. Abiotic and biotic controls of trophic level response during acidification were also identified. Asymmetric reductions of species richness among trophic levels, separated by life stage and feeding type, were evident and changes in trophic structure were most pronounced by the end of the acidification period. Relative declines in richness of fish and zooplankton were greater than phytoplankton, which were generally unaffected, leading to a reduction of upper trophic level diversity. Each of the lower four trophic levels responded to a distinct combination of abiotic and biotic variables during acidification. pH was identified as a direct driver of change for only carnivorous zooplankton, while all other trophic levels were affected more by indirect interactions caused by acidification. Fluctuations in ecosystem function (zooplankton biomass and primary production) were also evident, with losses at all trophic levels only detected during the last year of acidification. The acidified basin displayed a tendency for greater variation in biomass for upper trophic levels relative to reference conditions implying greater unpredictability in ecosystem function. Together, these results suggest that trophic asymmetry may be an important and recurring feature of ecosystem response to anthropogenic stress.     

Thermal acclimation modulates the impacts of temperature and enrichment on trophic interaction strengths and population dynamics

Global change affects individual phenotypes and biotic interactions, which can have cascading effects up to the ecosystem level. However, the role of environmentally induced phenotypic plasticity in species interactions is poorly understood, leaving a substantial gap in our knowledge of the impacts of global change on ecosystems. Using a cladoceran–dragonfly system, we experimentally investigated the effects of thermal acclimation, acute temperature change and enrichment on predator functional response and metabolic rate. Using our experimental data, we next parameterized a population dynamics model to determine the consequences of these effects on trophic interaction strength and food‐chain stability. We found that (1) predation and metabolic rates of the dragonfly larvae increase with acute warming, (2) warm‐acclimated larvae have a higher maximum predation rate than cold‐acclimated ones, and (3) long‐term interaction strength increases with enrichment but decreases with both acclimation and acute temperatures. Overall, our experimental results show that thermal acclimation can buffer negative impacts of environmental change on predators and increase food‐web stability and persistence. We conclude that the effect of acclimation and, more generally, phenotypic plasticity on trophic interactions should not be overlooked if we aim to understand the effects of climate change and enrichment on species interaction strength and food‐web stability.     

A novel framework for linking functional diversity of plants with other trophic levels for the quantification of ecosystem services

A novel conceptual framework is presented that proposes to apply trait‐based approaches to predicting the impact of environmental change on ecosystem service delivery by multi‐trophic systems. Development of the framework was based on an extension of the response–effect trait approach to capture functional relationships that drive trophic interactions. The framework was populated with worked examples to demonstrate its flexibility and value for linking disparate data sources, identifying knowledge gaps and generating hypotheses for quantitative models.     

The relationship between biodiversity and ecosystem functioning in food webs

Recent theoretical and experimental work provides clear evidence that biodiversity loss can have profound impacts on functioning of natural and managed ecosystems and the ability of ecosystems to deliver ecological services to human societies. Work on simplified ecosystems in which the diversity of a single trophic level is manipulated shows that diversity can enhance ecosystem processes such as primary productivity and nutrient retention. Theory also strongly suggests that biodiversity can act as biological insurance against potential disruptions caused by environmental changes. However, these studies generally concern a single trophic level, primary producers for the most part. Changes in biodiversity also affect ecosystem functioning through trophic interactions. Here we review, through the analysis of a simple ecosystem model, several key aspects inherent in multitrophic systems that may strongly affect the relationship between diversity and ecosystem processes. Our analysis shows that trophic interactions have a strong impact on the relationships between diversity and ecosystem functioning, whether the ecosystem property considered is total biomass or temporal variability of biomass at the various trophic levels. In both cases, food-web structure and trade-offs that affect interaction strength have major effects on these relationships. Multitrophic interactions are expected to make biodiversity–ecosystem functioning relationships more complex and non-linear, in contrast to the monotonic changes predicted for simplified systems with a single trophic level.     

Warming weakens facilitative interactions between decomposers and detritivores,and modifies freshwater ecosystem functioning

Warming is among the major drivers of changes in biotic interactions and, in turn, ecosystem functioning. The decomposition process occurs in a chain of facilitative interactions between detritivores and microorganisms. It remains unclear, however, what effect warming may have on the interrelations between detritivores and microorganisms, and the consequences for the functioning of natural freshwater ecosystems. To address these gaps, we performed a field experiment using tank bromeliads and their associated aquatic fauna. We manipulated the presence of bacteria and detritivorous macroinvertebrates (control, “bacteria,” and “bacteria + macroinvertebrates”) under ambient and warming scenarios, and analyzed the effects on the microorganisms and ecosystem functioning (detritus mass loss, colored dissolved organic matter, and nitrogen flux). We applied antibiotic solution to eliminate or reduce bacteria from control bromeliads. After 60 days incubation, bacterial density was higher in the presence than in the absence of macroinvertebrates. In the absence of macroinvertebrates, temperature did not influence bacterial density. However, in the presence of macroinvertebrates, bacterial density decreased by 54% with warming. The magnitude of the effects of organisms on ecosystem functioning was higher in the combined presence of bacteria and macroinvertebrates. However, warming reduced the overall positive effects of detritivores on bacterial density, which in turn, cascaded down to ecosystem functioning by decreasing decomposition and nitrogen flux. These results show the existence of facilitative mechanisms between bacteria and detritivores in the decomposition process, which might collapse due to warming. Detritivores seem to contribute to nutrient cycling as they facilitate bacterial populations, probably by increasing nutrient input (feces) in the ecosystem. However, increased temperature mitigated these beneficial effects. Our results add to a growing research body that shows that warming can affect the structure of aquatic communities, and highlight the importance of considering the interactive effects between facilitation and climatic drivers on the functioning of freshwater ecosystems.     

The importance of ecological memory for trophic rewilding as an ecosystem restoration approach

Increasing human pressure on strongly defaunated ecosystems is characteristic of the Anthropocene and calls for proactive restoration approaches that promote self‐sustaining, functioning ecosystems. However, the suitability of novel restoration concepts such as trophic rewilding is still under discussion given fragmentary empirical data and limited theory development. Here, we develop a theoretical framework that integrates the concept of ‘ecological memory’ into trophic rewilding. The ecological memory of an ecosystem is defined as an ecosystem's accumulated abiotic and biotic material and information legacies from past dynamics. By summarising existing knowledge about the ecological effects of megafauna extinction and rewilding across a large range of spatial and temporal scales, we identify two key drivers of ecosystem responses to trophic rewilding: (i) impact potential of (re)introduced megafauna, and (ii) ecological memory characterising the focal ecosystem. The impact potential of (re)introduced megafauna species can be estimated from species properties such as lifetime per capita engineering capacity, population density, home range size and niche overlap with resident species. The importance of ecological memory characterising the focal ecosystem depends on (i) the absolute time since megafauna loss, (ii) the speed of abiotic and biotic turnover, (iii) the strength of species interactions characterising the focal ecosystem, and (iv) the compensatory capacity of surrounding source ecosystems. These properties related to the focal and surrounding ecosystems mediate material and information legacies (its ecological memory) and modulate the net ecosystem impact of (re)introduced megafauna species. We provide practical advice about how to quantify all these properties while highlighting the strong link between ecological memory and historically contingent ecosystem trajectories. With this newly established ecological memory–rewilding framework, we hope to guide future empirical studies that investigate the ecological effects of trophic rewilding and other ecosystem‐restoration approaches. The proposed integrated conceptual framework should also assist managers and decision makers to anticipate the possible trajectories of ecosystem dynamics after restoration actions and to weigh plausible alternatives. This will help practitioners to develop adaptive management strategies for trophic rewilding that could facilitate sustainable management of functioning ecosystems in an increasingly human‐dominated world.     

Multiple stressors on biotic interactions: how climate change and alien species interact to affect pollination

Global change may substantially affect biodiversity and ecosystem functioning but little is known about its effects on essential biotic interactions. Since different environmental drivers rarely act in isolation it is important to consider interactive effects. Here, we focus on how two key drivers of anthropogenic environmental change, climate change and the introduction of alien species, affect plant–pollinator interactions. Based on a literature survey we identify climatically sensitive aspects of species interactions, assess potential effects of climate change on these mechanisms, and derive hypotheses that may form the basis of future research. We find that both climate change and alien species will ultimately lead to the creation of novel communities. In these communities certain interactions may no longer occur while there will also be potential for the emergence of new relationships. Alien species can both partly compensate for the often negative effects of climate change but also amplify them in some cases. Since potential positive effects are often restricted to generalist interactions among species, climate change and alien species in combination can result in significant threats to more specialist interactions involving native species.     

Territorial ants depress plant growth through cascading non‐trophic effects in an alpine meadow

Understanding non‐trophic interactions is critical to mechanistically linking community structure and ecosystem functioning. Despite the widespread occurrence of territoriality across animal taxa and ecosystems, the cascading ecological consequences of non‐trophic interactions between territorial animals and intruders have been poorly studied. We experimentally investigated the non‐trophic interaction between territorial ants and members of a dung decomposer community (i.e. predatory arthropods, maggots and coprophagous beetles) in an alpine meadow. We further examined how this non‐trophic interaction cascaded to influence ecosystem properties including dung removal rate, soil nutrient status and aboveground plant biomass surrounding dung pats. Results indicated that territorial interference of ants on key decomposers cascaded to affect plant growth. Specifically, ants significantly decreased the abundance of coprophagous beetles at the time of their peak‐abundance and hence decreased dung removal rates and soil nitrogen concentrations, ultimately decreasing aboveground plant biomass. The strength of this non‐trophic cascading effect was comparable to those reported in studies addressing trophic cascades triggered by predator–prey interactions. Our findings suggest that the non‐trophic interactions and associated cascading effects stemming from territorial behavior should be incorporated into ecological network modeling and research addressing biodiversity–ecosystem functioning relationships.