Neil Rooney

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.

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.

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.

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.

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.

From energetics to ecosystems : the dynamics and structure of ecological systems

SECTION I.- A Process-Oriented Approach to the Multispecies Functional Response.- Homage to Yodzis and Innes 1992: Scaling up Feeding-Based Population Dynamics to Complex Ecological Networks.- Food Webs, Body Size and the Curse of the Latin Binomial.- An Energetic Framework for Trophic Control.- SECTION II.- Experimental Studies of Food Webs: Causes and Consequences of Trophic Interactions.- Interplay Between Scale, Resolution, Life History and Food Web Properties.- Heteroclinic Cycles in the Rain Forest: Insights from Complex Dynamics.- Emergence in Ecological Systems.- Dynamic Signatures of Real and Model Ecosystems.- SECTION III.- Evolutionary Branching of Single Traits.- Feedback Effects Between the Food Chain and Induced Defense Strategies.- Evolutionary Demography: The Invasion Exponent and the Effective Population Density in Nonlinear Matrix Models.- Of Experimentalists, Empiricists, and Theoreticians.

Food Web Structure in Temporally-Forced Ecosystems

Temporal variation characterizes many of Earths 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.

Submerged Macrophyte-bed Effects on Water-Column Phosphorus, Chlorophyll a , and Bacterial Production

Submerged macrophytes are a major component of freshwater ecosystems, yet their net effect on water column phosphorus (P), algae, and bacterioplankton is not well understood. A 4-month mass-balance study during the summer quantified the net effect of a large (∼5.5 ha) undisturbed macrophyte bed on these water-column properties. The bed is located in a slow-flowing (0.05–0.1 cm s−1) channel between two lakes, allowing for the quantification of inputs and outputs. The P budget for the study period showed that, despite considerable short-term variation, the macrophyte bed was a negligible net sink for P (0.06 mg m−2 day−1, range from −0.76 to +0.79 mg m−2 day−1), demonstrating that loading and uptake processes in the weedbed roughly balance over the summer. Chlorophyll a was disproportionately retained relative to particulate organic carbon (POC), indicating that the algal component of the POC was preferentially trapped. However, the principal contribution of the weedbed to the open water was a consistent positive influence on bacterioplankton production over the summer. Conservative extrapolations based on measured August specific exports (m−2 day−1) of P and bacterial production exiting the weedbed applied to five regional lakes varying in lake morphometry and macrophyte cover suggest that even in the most macrophyte dominated of lakes (66% cover), P loading from submerged weedbeds never exceeds 1% day−1 of standing epilimnetic P levels, whereas subsidization of bacterioplankton production can reach upward of 20% day−1. The presence of submerged macrophytes therefore differentially modifies algae and bacteria in the water column, while modestly altering P dynamics over the summer.

Interactions among epilimnetic phosphorus, phytoplankton biomass and bacterioplankton metabolism in lakes of varying submerged macrophyte cover

The effect of submerged macrophytes on interactions among epilimnetic phosphorus, phytoplankton, and heterotrophic bacterioplankton has been acknowledged, but remains poorly understood. Here, we test the hypotheses that the mean summer phytoplankton biomass (chlorophyll a): phosphorus ratios decrease with increased macrophyte cover in a series of nine lakes. Further, we test that both planktonic respiration and bacterioplankton production increase with respect to phytoplankton biomass along the same gradient of increasing macrophyte cover. Increased macrophyte cover was associated with a lower fraction of particulate phosphorus in epilimnia, with total particulate phosphorus declining from over 80% of total phosphorus in a macrophyte free lake to less than 50% in a macrophyte rich lake. Phytoplankton biomass (chlorophyll a) too was lower in macrophyte dominated lakes, despite relatively high levels of total dissolved phosphorus. Planktonic respiration and bacterioplankton production were higher in macrophyte rich lakes than would be expected from phytoplankton biomass alone, pointing to a subsidy of bacterioplankton metabolism by macrophyte beds at the whole lake scale. The results suggest that the classical view of pelagic interactions, which proposes phosphorus determines phytoplankton abundance, which in turn determines bacterial abundance through the production of organic carbon, becomes less relevant as macrophyte cover increases.

The adaptive capacity of lake food webs: from individuals to ecosystems

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.