Individual energetics scale up to community coexistence: Movement, metabolism and biodiversity dynamics in fragmented landscapes.

Unravelling the intricate mechanisms that govern community coexistence remains a daunting challenge, particularly amidst ongoing environmental change. Individual physiology and metabolism are often studied to understand the response of individual animals to environmental change. However, this perspective is currently largely lacking in community ecology. We argue that the integration of individual metabolism into community theory can offer new insights into coexistence. We present the first individual-based metabolic community model for a terrestrial mammal community to simulate energy dynamics and home range behaviour in different environments. Using this model, we investigate how ecologically similar species coexist and maintain their energy balance under food competition. Only if individuals of different species are able to balance their incoming and outgoing energy over the long-term will they be able to coexist. After thoroughly testing and validating the model against real-world patterns such as of home range dynamics and field metabolic rates, we applied it as a case study to scenarios of habitat fragmentation - a widely discussed topic in biodiversity research. First, comparing single-species simulations with community simulations, we find that the effect of habitat fragmentation on populations is strongly context-dependent. While populations of species living alone in the landscape were mostly positively affected by fragmentation, the diversity of a community of species was highest under medium fragmentation scenarios. Under medium fragmentation, energy balance and reproductive investment were also most similar among species. We therefore suggest that similarity in energy balance among species promotes coexistence. We argue that energetics should be part of community ecology theory, as the relative energetic status and reproductive investment can reveal why and under what environmental conditions coexistence is likely to occur. As a result, landscapes can potentially be protected and designed to maximize coexistence. The metabolic community model presented here can be a promising tool to investigate other scenarios of environmental change or other species communities to further disentangle global change effects and preserve biodiversity.

Energy-mediated responses to changing prey size and distribution in marine top predator movements and population dynamics.

Climate change is modifying the structure of marine ecosystems, including that of fish communities. Alterations in abiotic and biotic conditions can decrease fish size and change community spatial arrangement, ultimately impacting predator species which rely on these communities. To conserve predators and understand the drivers of observed changes in their population dynamics, we must advance our understanding of how shifting environmental conditions can impact populations by limiting food available to individuals. To investigate the impacts of changing fish size and spatial aggregation on a top predator population, we applied an existing agent-based model parameterized for harbour porpoises Phocoena phocoena which represents animal energetics and movements in high detail. We used this framework to quantify the impacts of shifting prey size and spatial aggregation on porpoise movement, space use, energetics and population dynamics. Simulated individuals were more likely to switch from area-restricted search to transit behaviour with increasing prey size, particularly when starving, due to elevated resource competition. In simulations with highly aggregated prey, higher prey encounter rates counteracted resource competition, resulting in no impacts of prey spatial aggregation on movement behaviour. Reduced energy intake with decreasing prey size and aggregation level caused population decline, with a 15% decrease in fish length resulting in total population collapse Increasing prey consumption rates by 42.8 ± 4.5% could offset population declines; however, this increase was 21.3 ± 12.7% higher than needed to account for changes in total energy availability alone. This suggests that animals in realistic seascapes require additional energy to locate smaller prey which should be considered when assessing the impacts of decreased energy availability. Changes in prey size and aggregation influenced movements and population dynamics of simulated harbour porpoises, revealing that climate-induced changes in prey structure, not only prey abundance, may threaten predator populations. We demonstrate how a population model with realistic animal movements and process-based energetics can be used to investigate population consequences of shifting food availability, such as those mediated by climate change, and provide a mechanistic explanation for how changes in prey structure can impact energetics, behaviour and ultimately viability of predator populations.

Movement and Seasonal Energetics Mediate Vulnerability to Disturbance in Marine Mammal Populations.

AbstractIn marine environments, noise from human activities is increasing dramatically, causing animals to alter their behavior and forage less efficiently. These alterations incur energetic costs that can result in reproductive failure and death and may ultimately influence population viability, yet the link between population dynamics and individual energetics is poorly understood. We present an energy budget model for simulating effects of acoustic disturbance on populations. It accounts for environmental variability and individual state, while incorporating realistic animal movements. Using harbor porpoises (Phocoena phocoena) as a case study, we evaluated population consequences of disturbance from seismic surveys and investigated underlying drivers of vulnerability. The framework reproduced empirical estimates of population structure and seasonal variations in energetics. The largest effects predicted for seismic surveys were in late summer and fall and were unrelated to local abundance, but instead were related to lactation costs, water temperature, and body fat. Our results demonstrate that consideration of temporal variation in individual energetics and their link to costs associated with disturbances is imperative when predicting disturbance impacts. These mechanisms are general to animal species, and the framework presented here can be used for gaining new insights into the spatiotemporal variability of animal movements and energetics that control population dynamics.

Local buffer mechanisms for population persistence.

Assessing and predicting the persistence of populations is essential for the conservation and control of species. Here, we argue that local mechanisms require a better conceptual synthesis to facilitate a more holistic consideration along with regional mechanisms known from metapopulation theory. We summarise the evidence for local buffer mechanisms along with their capacities and emphasise the need to include multiple buffer mechanisms in studies of population persistence. We propose an accessible framework for local buffer mechanisms that distinguishes between damping (reducing fluctuations in population size) and repelling (reducing population declines) mechanisms. We highlight opportunities for empirical and modelling studies to investigate the interactions and capacities of buffer mechanisms to facilitate better ecological understanding in times of ecological upheaval.

Key questions in marine mammal bioenergetics.

Bioenergetic approaches are increasingly used to understand how marine mammal populations could be affected by a changing and disturbed aquatic environment. There remain considerable gaps in our knowledge of marine mammal bioenergetics, which hinder the application of bioenergetic studies to inform policy decisions. We conducted a priority-setting exercise to identify high-priority unanswered questions in marine mammal bioenergetics, with an emphasis on questions relevant to conservation and management. Electronic communication and a virtual workshop were used to solicit and collate potential research questions from the marine mammal bioenergetic community. From a final list of 39 questions, 11 were identified as 'key' questions because they received votes from at least 50% of survey participants. Key questions included those related to energy intake (prey landscapes, exposure to human activities) and expenditure (field metabolic rate, exposure to human activities, lactation, time-activity budgets), energy allocation priorities, metrics of body condition and relationships with survival and reproductive success and extrapolation of data from one species to another. Existing tools to address key questions include labelled water, animal-borne sensors, mark-resight data from long-term research programs, environmental DNA and unmanned vehicles. Further validation of existing approaches and development of new methodologies are needed to comprehensively address some key questions, particularly for cetaceans. The identification of these key questions can provide a guiding framework to set research priorities, which ultimately may yield more accurate information to inform policies and better conserve marine mammal populations.

From theory to practice in pattern-oriented modelling: identifying and using empirical patterns in predictive models.

To robustly predict the effects of disturbance and ecosystem changes on species, it is necessary to produce structurally realistic models with high predictive power and flexibility. To ensure that these models reflect the natural conditions necessary for reliable prediction, models must be informed and tested using relevant empirical observations. Pattern-oriented modelling (POM) offers a systematic framework for employing empirical patterns throughout the modelling process and has been coupled with complex systems modelling, such as in agent-based models (ABMs). However, while the production of ABMs has been rising rapidly, the explicit use of POM has not increased. Challenges with identifying patterns and an absence of specific guidelines on how to implement empirical observations may limit the accessibility of POM and lead to the production of models which lack a systematic consideration of reality. This review serves to provide guidance on how to identify and apply patterns following a POM approach in ABMs (POM-ABMs), specifically addressing: where in the ecological hierarchy can we find patterns; what kinds of patterns are useful; how should simulations and observations be compared; and when in the modelling cycle are patterns used? The guidance and examples provided herein are intended to encourage the application of POM and inspire efficient identification and implementation of patterns for both new and experienced modellers alike. Additionally, by generalising patterns found especially useful for POM-ABM development, these guidelines provide practical help for the identification of data gaps and guide the collection of observations useful for the development and verification of predictive models. Improving the accessibility and explicitness of POM could facilitate the production of robust and structurally realistic models in the ecological community, contributing to the advancement of predictive ecology at large.