Species portfolio effects dominate seasonal zooplankton stabilization within a large temperate lake.

Portfolio effects (PEs) in ecology refer to the suite of phenomenon where the temporal variation of aggregate ecosystem properties (i.e., abundance) is lower than that of their ecosystem components. An example of this is where differential responses of species to environmental variation generate stability at higher levels of ecological organization (e.g., local community, metapopulation, metacommunity). Most of the research examining such PEs has focused on spatial or interannual variation of ecosystems; however, as global change continues to alter seasonality and ecosystem functioning, understanding the underlying food web structures that help maintain stability at multiple spatial and temporal scales is critical to managing ecological systems. Recent advances investigating diversity-stability relationships has led to the development of frameworks that incorporate a metacommunity perspective which allows for the partitioning of PEs across organizational scales (i.e., local community, metapopulation, cross-community, metacommunity) from local population dynamics (total). This partitioning yields insights into the mechanisms that generate observed PEs in nature. Here, we employed one of these recently developed frameworks on a temporally (1986-1999, 2008-2019) and spatially (five sampling stations, local communities) extensive data set of zooplankton abundance (e.g., density) within a large temperate lake to investigate how temporal (seasonal) and spatial (among site) PEs influence stability within the zooplankton metacommunity. We found that seasonal asynchrony of different zooplankton species within local communities and across communities generated the vast majority of stabilization, while spatial (i.e., metapopulation) dynamics were more synchronous and contributed little to overall system stability. Furthermore, significantly positive diversity-asynchrony relationships at the total, local- and cross-community scales were found as asynchrony was positively correlated with local Shannon diversity. Last, a comparison of PEs over the time periods, during which significant local and global changes (i.e., climate warming, invasive species) have occurred suggests that PEs may be eroding, as increasingly synchronous dynamics and declining diversity in recent years have led to a rise in metacommunity variability. We end by arguing for the critical importance of understanding seasonally driven stabilizing mechanisms as local and global changes threaten to fundamentally alter seasonal signals with potentially strong implications for the structures that lend stability to ecosystems.

Consumer trophic positions respond variably to seasonally fluctuating environments.

The effects of environmental seasonality on food web structure have been notoriously understudied in empirical ecology. Here, we focus on seasonal changes in one key attribute of a food web, consumer trophic position. We ask whether fishes inhabiting tropical river-floodplain ecosystems behave as seasonal omnivores, by shifting their trophic positions in relation to the annual flood pulse, or whether they feed at the same trophic position all year, as much empirical work implicitly assumes. Using dietary data from the Tonle Sap Lake, Cambodia, and a literature review, we find evidence that some fishes, especially small piscivores, increased consumption of invertebrates and/or plant material during the wet season, as predicted. However, nitrogen stable isotope (δ15 N) data for 26 Tonle Sap fishes, spanning a broader range of functional groups, uncovered high variation in seasonal trophic position responses among species (0 to ±0.52 trophic positions). Based on these findings, species respond to the flood pulse differently. Diverse behavioral responses to seasonality, underpinned by spatiotemporal variation at multiple scales, could be central for rerouting matter and energy flow in these dynamic ecosystems. Seasonally flexible foraging behaviors warrant further study given their potential influence on food web dynamics in a range of fluctuating environments.

When too much isn't enough: Does current food production meet global nutritional needs?

Sustainably feeding the next generation is often described as one of the most pressing "grand challenges" facing the 21st century. Generally, scholars propose addressing this problem by increasing agricultural production, investing in technology to boost yields, changing diets, or reducing food waste. In this paper, we explore whether global food production is nutritionally balanced by comparing the diet that nutritionists recommend versus global agricultural production statistics. Results show that the global agricultural system currently overproduces grains, fats, and sugars while production of fruits and vegetables and protein is not sufficient to meet the nutritional needs of the current population. Correcting this imbalance could reduce the amount of arable land used by agriculture by 51 million ha globally but would increase total land used for agriculture by 407 million ha and increase greenhouse gas emissions. For a growing population, our calculations suggest that the only way to eat a nutritionally balanced diet, save land and reduce greenhouse gas emissions is to consume and produce more fruits and vegetables as well as transition to diets higher in plant-based protein. Such a move will help protect habitats and help meet the Sustainable Development Goals.

Food Web Structure in Temporally-Forced Ecosystems.

Temporal variation characterizes many of Earth's ecosystems. Despite this, little is known about how food webs respond to regular variation in time, such as occurs broadly with season. We argue that season, and likely any periodicity, structures food webs along a temporal axis in an analogous way to that previously recognized in space; predators shift their diet as different resource compartments and trophic levels become available through time. These characteristics are likely (i) central to ecosystem function and stability based on theory, and (ii) widespread across ecosystem types based on empirical observations. The temporal food web perspective outlined here could provide new insight into the ecosystem-level consequences of altered abiotic and biotic processes that might accompany globally changing environments.

HSS revisited: multi-channel processes mediate trophic control across a productivity gradient.

Classical food web theory holds that energy channels are regulated by top-down control with increasing productivity, arising from within-channel processes. However, these hypotheses do not consider the existence of parallel energy channels linked by shared resource pools and which can fuel generalist predators, imposing trophic control arising from multi-channel processes. Using 23 large marine food webs, we show that food web responses to increasing productivity are consistent with the apparent trophic cascade hypothesis (ATCH) - with rising productivity predators derive an increasing fraction of their diet from increasingly productive bottom-up controlled detritus channels, thereby subsidising predator biomass, and in turn strengthening top-down control in parallel grazing channels. These results testify to a fundamental role of detritus channels specifically and multi-channel processes in general in mediating food web response to productivity and demonstrate that the ATCH provides an alternative explanation for classical predictions of top-down control.

Integrating food web diversity, structure and stability.

Given the unprecedented rate of species extinctions facing the planet, understanding the causes and consequences of species diversity in ecosystems is of paramount importance. Ecologists have investigated both the influence of environmental variables on species diversity and the influence of species diversity on ecosystem function and stability. These investigations have largely been carried out without taking into account the overarching stabilizing structures of food webs that arise from evolutionary and successional processes and that are maintained through species interactions. Here, we argue that the same large-scale structures that have been purported to convey stability to food webs can also help to understand both the distribution of species diversity in nature and the relationship between species diversity and food web stability. Specifically, the allocation of species diversity to slow energy channels within food webs results in the skewed distribution of interactions strengths that has been shown to confer stability to complex food webs. We end by discussing the processes that might generate and maintain the structured, stable and diverse food webs observed in nature.

The more food webs change, the more they stay the same.

Here, we synthesize a number of recent empirical and theoretical papers to argue that food-web dynamics are characterized by high amounts of spatial and temporal variability and that organisms respond predictably, via behaviour, to these changing conditions. Such behavioural responses on the landscape drive a highly adaptive food-web structure in space and time. Empirical evidence suggests that underlying attributes of food webs are potentially scale-invariant such that food webs are characterized by hump-shaped trophic structures with fast and slow pathways that repeat at different resolutions within the food web. We place these empirical patterns within the context of recent food-web theory to show that adaptable food-web structure confers stability to an assemblage of interacting organisms in a variable world. Finally, we show that recent food-web analyses agree with two of the major predictions of this theory. We argue that the next major frontier in food-web theory and applied food-web ecology must consider the influence of variability on food-web structure.

Parallel ecological networks in ecosystems.

In ecosystems, species interact with other species directly and through abiotic factors in multiple ways, often forming complex networks of various types of ecological interaction. Out of this suite of interactions, predator-prey interactions have received most attention. The resulting food webs, however, will always operate simultaneously with networks based on other types of ecological interaction, such as through the activities of ecosystem engineers or mutualistic interactions. Little is known about how to classify, organize and quantify these other ecological networks and their mutual interplay. The aim of this paper is to provide new and testable ideas on how to understand and model ecosystems in which many different types of ecological interaction operate simultaneously. We approach this problem by first identifying six main types of interaction that operate within ecosystems, of which food web interactions are one. Then, we propose that food webs are structured among two main axes of organization: a vertical (classic) axis representing trophic position and a new horizontal 'ecological stoichiometry' axis representing decreasing palatability of plant parts and detritus for herbivores and detrivores and slower turnover times. The usefulness of these new ideas is then explored with three very different ecosystems as test cases: temperate intertidal mudflats; temperate short grass prairie; and tropical savannah.

A landscape theory for food web architecture.

Ecologists have long searched for structures and processes that impart stability in nature. In particular, food web ecology has held promise in tackling this issue. Empirical patterns in food webs have consistently shown that the distributions of species and interactions in nature are more likely to be stable than randomly constructed systems with the same number of species and interactions. Food web ecology still faces two fundamental challenges, however. First, the quantity and quality of food web data required to document both the species richness and the interaction strengths among all species within food webs is largely prohibitive. Second, where food webs have been well documented, spatial and temporal variation in food web structure has been ignored. Conversely, research that has addressed spatial and temporal variation in ecosystems has generally ignored the full complexity of food web architecture. Here, we incorporate empirical patterns, largely from macroecology and behavioural ecology, into a spatially implicit food web structure to construct a simple landscape theory of food web architecture. Such an approach both captures important architectural features of food webs and allows for an exploration of food web structure across a range of spatial scales. Finally, we demonstrated that food webs are hierarchically organized along the spatial and temporal niche axes of species and their utilization of food resources in ways that stabilize ecosystems.

Structural asymmetry and the stability of diverse food webs.

Untangling the influence of human activities on food-web stability and persistence is complex given the large numbers of species and overwhelming number of interactions within ecosystems. Although biodiversity has been associated with stability, the actual structures and processes that confer stability to diverse food webs remain largely unknown. Here we show that real food webs are structured such that top predators act as couplers of distinct energy channels that differ in both productivity and turnover rate. Our theoretical analysis shows that coupled fast and slow channels convey both local and non-local stability to food webs. Alarmingly, the same human actions that have been implicated in the loss of biodiversity also directly erode the very structures and processes that we show to confer stability on food webs.