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How the combination of positive and negative species interactions acts to drive community dynamics is a fundamental question in ecology. Here we explore one aspect of this question by expanding the theory of predator-mediated coexistence to include the potential role of facilitation between the predator and inferior competitor. To motivate and illustrate our simple model, we focus on sea-urchin--algae interactions in temperate rocky reef systems and incorporate recruitment facilitation, a common characteristic of marine systems. Specifically, the model represents sea urchin grazing on macroalgae, macroalgal competition with crustose coralline algae (CCA), and facilitation of sea urchin recruitment to CCA. These interactions generate alternative stable states, one dominated by macroalgae and the other by urchins, which do not occur when recruitment facilitation of urchins to CCA is ignored. Therefore, recruitment facilitation provides a possible mechanism for alternative kelp forest and urchin barren states in temperate marine systems, where storm events or harvesting of urchins or their predators can drive switches between states that are difficult to reverse. In systems with such dynamics, spatial management such as no-take marine reserves may play a crucial role in protecting community structure by increasing the resilience to shifts between states. Key words: alternative states; crustose coralline algae; herbivory; hysteresis; kelp forest; macroalgae; recruitment facilitation; resilience; urchin barren.

Human-caused global change produces biotic and abiotic conditions that increase the uncertainty and risk of failure of restoration efforts. A focus of managing for resiliency, that is, the ability of the system to respond to disturbance, has the potential to reduce this uncertainty and risk. However, identifying what drives resiliency might depend on how one measures it. An example of a system where identifying how the drivers of different aspects of resiliency can inform restoration under climate change is the northern coast of California, where kelp experienced a decline in coverage of over 95% due to the combination of an intense marine heat wave and the functional extinction of the primary predator of the kelp-grazing purple sea urchin, the sunflower sea star. Although restoration efforts focused on urchin removal and kelp reintroduction in this system are ongoing, the question of how to increase the resiliency of this system to future marine heat waves remains open. In this paper, we introduce a dynamical model that describes a tritrophic food chain of kelp, purple urchins, and a purple urchin predator such as the sunflower sea star. We run a global sensitivity analysis of three different resiliency metrics (recovery likelihood, recovery rate, and resistance to disturbance) of the kelp forest to identify their ecological drivers. We find that each metric depends the most on a unique set of drivers: Recovery likelihood depends the most on live and drift kelp production, recovery rate depends the most on urchin production and feedbacks that determine urchin grazing on live kelp, and resistance depends the most on feedbacks that determine predator consumption of urchins. Therefore, an understanding of the potential role of predator reintroduction or recovery in kelp systems relies on a comprehensive approach to measuring resiliency., (© 2024 The Author(s). Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Climate refugia are areas where species can persist through climate change with little to no movement. Among the factors associated with climate refugia are high spatial heterogeneity, such that there is only a short distance between current and future optimal climates, as well as biotic or abiotic environmental factors that buffer against variability in time. However, these types of climate refugia may be declining due to anthropogenic homogenization of environments and degradation of environmental buffers. To quantify the potential for restoration of refugia-like environmental conditions to increase population persistence under climate change, we simulated a population's capacity to track their temperature over space and time given different levels of spatial and temporal variability in temperature. To determine how species traits affected the efficacy of restoring heterogeneity, we explored an array of values for species' dispersal ability, thermal tolerance, and fecundity. We found that species were more likely to persist in environments with higher spatial heterogeneity and lower environmental stochasticity. When simulating a management action that increased the spatial heterogeneity of a previously homogenized environment, species were more likely to persist through climate change, and population sizes were generally higher, but there was little effect with mild temperature change. The benefits of heterogeneity restoration were greatest for species with limited dispersal ability. In contrast, species with longer dispersal but lower fecundity were more likely to benefit from a reduction in environmental stochasticity than an increase in spatial heterogeneity. Our results suggest that restoring environments to refugia-like conditions could promote species' persistence under the influence of climate change in addition to conservation strategies such as assisted migration, corridors, and increased protection., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

In restoration ecology, the Field of Dreams hypothesis posits that restoration efforts that create a suitable environment could lead to the eventual recovery of the remaining aspects of the ecosystem through natural processes. Natural processes following partial restoration has led to ecosystem recovery in both terrestrial and aquatic systems. However, understanding the efficacy of a "Field of Dreams" approach requires a comparison of different approaches to partial restoration in terms of spatial, temporal, and ecological scale with what would happen given more comprehensive restoration efforts. We explore the relative effect of partial restoration and ongoing recovery on restoration efficacy with a dynamical model based on temperate rocky reefs in Northern California. We analyze our model for both the ability and rate of bull kelp forest recovery under different restoration strategies. We compare the efficacy of a partial restoration approach with a more comprehensive restoration effort by exploring how kelp recovery likelihood and rate change with varying intensities of urchin removal and kelp outplanting over different time periods and spatial scales. We find that, in the case of bull kelp forests, setting more favorable initial conditions for kelp recovery by implementing both urchin harvesting and kelp outplanting at the start of the restoration project has a bigger impact on the kelp recovery rate than applying restoration efforts through a longer period of time. Therefore, partial restoration efforts, in terms of spatial and temporal scale, can be significantly more effective when applied across multiple ecological scales in terms of both the capacity and rate for achieving the target outcomes., (© 2023 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Outcomes of management efforts to recover or restore populations of harvested species can be highly dependent on environmental and community context. Predator-prey interactions can alter recovery trajectories, and the timing of management actions within multi-trophic level harvest scenarios may influence the dynamics of recovery and lead to management trade-offs. Recent work using a generalist predator-prey model suggests that management promoting synchronized recovery of predators and prey leads to faster and less variable recovery trajectories than sequential recovery (predator or prey first). However, more complex communities may require different management actions to minimize recovery time and variability. Here, we use a tri-trophic level rocky reef community dynamics model with size-structure and fisheries at multiple trophic levels to investigate the importance of three ecological processes to recovery of fished communities: (1) size-structured predation, (2) non-consumptive effects of predators on prey behavior, and (3) varying levels of recruitment. We also test the effects of initiating recovery from community states associated with varying degrees of fishery-induced degradation and develop a simulation in which the basal resource (kelp) is harvested. In this system, a predator-first closure generally leads to the least volatile and quickest recovery, whether from a kelp forest, urchin barren, or intermediate community state. The benefits gained by selecting this strategy are magnified when recovering from the degraded community, the urchin barren, because initial conditions in the degraded state lead to lengthy recovery times. However, the shape of the size-structured predation relationship can strongly affect recovery volatility, where the differences between alternate management strategies are negated with size-independent predation. External recruitment reduces return times by bolstering the predatory lobster population. These results show that in a tightly linked tri-trophic level food web with top-down control, a predator-first fishery closure can be the most effective strategy to reduce volatility and shorten recovery, particularly when the system is starting from the degraded community state. Given the ubiquity of top predator loss across many ecosystems, we highlight the value of incorporating insights from community ecology into ecosystem management., (© 2021 Ecological Society of America. This article has been contributed to by US Government employees and their work is in the public domain in the USA.)

Marine Protected Areas (MPAs) are increasingly established globally as a spatial management tool to aid in conservation and fisheries management objectives. Assessing whether MPAs are having the desired effects on populations requires effective monitoring programs. A cornerstone of an effective monitoring program is an assessment of the statistical power of sampling designs to detect changes when they occur. We present a novel approach to power assessment that combines spatial point process models, integral projection models (IPMs) and sampling simulations to assess the power of different sample designs across a network of MPAs. We focus on the use of remotely operated vehicle (ROV) video cameras as the sampling method, though the results could be extended to other sampling methods. We use empirical data from baseline surveys of an example indicator fish species across three MPAs in California, USA as a case study. Spatial models simulated time series of spatial distributions across sites that accounted for the effects of environmental covariates, while IPMs simulated expected trends over time in abundances and sizes of fish. We tested the power of different levels of sampling effort (i.e., the number of 500-m ROV transects) and temporal replication (every 1-3 yr) to detect expected post-MPA changes in fish abundance and biomass. We found that changes in biomass are detectable earlier than changes in abundance. We also found that detectability of MPA effects was higher in sites with higher initial densities. Increasing the sampling effort had a greater effect than increasing sampling frequency on the time taken to achieve high power. High power was best achieved by combining data from multiple sites. Our approach provides a powerful tool to explore the interaction between sampling effort, spatial distributions, population dynamics, and metrics for detecting change in previously fished populations., (© 2020 by the Ecological Society of America.)

Harvest mortality typically truncates the harvested species' size structure, thereby reducing phenotypic complexity, which can lead to reduced population productivity, increased population variability, and selection on an array of life history traits that can further alter these demographic processes. Marine protected areas (MPAs) are a potential tool to protect older, larger individuals and therefore mitigate such ecological and evolutionary effects of harvest, depending on the degree of connectivity among areas. Such MPA protection relies on a shift in size-dependent mortality, the measurement of which can therefore serve as an early indicator of whether MPAs might achieve the desired longer-term ecological and evolutionary responses. We directly measured MPA effects on size-selective mortality and associated size structure using mark-recapture data on European lobster (Homarus gammarus) collected at three MPA-control area pairs in southern Norway during one decade (n = 5,943). Mark-recapture modeling, accounting for variation in recapture probabilities, revealed (1) that annual mean survival was higher inside MPAs (0.592) vs. control areas (0.298) and (2) that significant negative relationships between survival and body size occurred at the control areas but not in the MPAs, where the effect of body size was predominantly positive. Additionally, we found (3) that mean and maximum body size increased over time inside MPAs but not in control areas. Overall, our results suggest that MPAs can rebuild phenotypic complexity (i.e., size structure) and provide protection from harvest selection., (© 2020 The Authors. Ecological Applications published by Wiley Periodicals, Inc. on behalf of Ecological Society of America.)

Whether ecosystems recover from disturbance depends on the presence of alternative stable states, which are theoretically possible in simple models of many systems. However, definitive empirical evidence for this phenomenon remains limited to demographically closed ecosystems such as lakes. In more interconnected systems such as temperate rocky reefs, the local relevance of alternative stable states might erode as immigration overwhelms local feedbacks and produces a single stable state. At larger spatial scales, dispersal might counter localized disturbance and feedbacks to synchronize states throughout a region. Here, we quantify how interconnectedness affects the relevance of alternative stable states using dynamical models of California rocky reef communities that incorporate observed environmental stochasticity and feedback loops in kelp-urchin-predator interactions. Our models demonstrate the potential for localized alternative states despite high interconnectedness likely due to feedbacks affecting dispersers as they settle into local communities. Regionally, such feedbacks affecting settlement can produce a mosaic of alternative stable states that span local (10-20 km) scales despite the synchronizing effect of long-distance dispersal. The specific spatial scale and duration of each state predominantly depend on the scales of environmental variation and on local dynamics (here, fishing). Model predictions reflect observed scales of community states in California rocky reefs and suggest how alternative states co-occur in the wide array of marine and terrestrial systems with settlement feedbacks., (© 2019 by the Ecological Society of America.)

Disturbance plays a key role in structuring community dynamics and is central to conservation and natural resource management. However, ecologists continue to debate the importance of disturbance for species coexistence and biodiversity. Such disagreements may arise in part because few studies have examined variation across multiple dimensions of disturbance (e.g., size, frequency) and how the effects of disturbance may depend on species attributes (e.g., competitiveness, dispersal ability). In light of this gap in understanding and accelerating changes to disturbance regimes worldwide, we used spatial population models to explore how disturbance size and frequency interact with species attributes to affect coexistence between seagrass (Zostera marina) and colonial burrowing shrimp (Neotrypaea californiensis) that compete for benthic space in estuaries throughout the west coast of North America. By simulating population dynamics under a range of ecologically relevant disturbance regimes, we discovered that intermediate disturbance (approximately 9-23% of landscape area per year) to short-dispersing, competitively dominant seagrass can foster long-term stable coexistence with broad-dispersing, competitively inferior burrowing shrimp via the spatial storage effect. When holding the total extent of disturbance constant, the individual size and annual frequency of disturbance altered landscape spatial patterns and mediated the dominance and evenness of competitors. Many small disturbances favored short-dispersing seagrass by hastening recolonization, whereas fewer large disturbances benefited rapidly colonizing burrowing shrimp by creating temporary refugia from competition. As a result, large, infrequent disturbances generally improved the strength and stability of coexistence relative to small, frequent disturbances. Regardless of disturbance size or frequency, the dispersal ability of the superior competitor (seagrass), the competitive ability of the inferior competitor (burrowing shrimp), and the reproduction and survival of both species strongly influenced population abundances and coexistence. Our results show that disturbance size and frequency can promote or constrain coexistence by altering the duration of time over which inferior competitors can escape competitive exclusion, particularly when colonization depends on the spatial pattern of disturbance due to dispersal traits. For coastal managers and conservation practitioners, our findings indicate that reducing particularly large disturbances may help conserve globally imperiled seagrass meadows and control burrowing shrimp colonies that can threaten the viability of oyster aquaculture., (© 2019 by the Ecological Society of America.)

Adaptive management of marine protected areas (MPAs) requires developing methods to evaluate whether monitoring data indicate that they are performing as expected. Modeling the expected responses of targeted species to an MPA network, with a clear timeline for those expectations, can aid in the development of a monitoring program that efficiently evaluates expectations over appropriate time frames. Here, we describe the expected trajectories in abundance and biomass following MPA implementation for populations of 19 nearshore fishery species in California. To capture the process of filling in the age structure truncated by fishing, we used age-structured population models with stochastic larval recruitment to predict responses to MPA implementation. We implemented both demographically open (high larval immigration) and closed (high self-recruitment) populations to model the range of possible trajectories as they depend on recruitment dynamics. From these simulations, we quantified the time scales over which anticipated increases in abundance and biomass inside MPAs would become statistically detectable. Predicted population biomass responses range from little change, for species with low fishing rates, to increasing by a factor of nearly seven, for species with high fishing rates before MPA establishment. Increases in biomass following MPA implementation are usually greater in both magnitude and statistical detectability than increases in abundance. For most species, increases in abundance would not begin to become detectable for at least 10 years after implementation. Overall, these results inform potential indicator metrics (biomass), potential indicator species (those with a high fishing : natural mortality ratio), and time frame (>10 yr) for MPA monitoring assessment as part of the adaptive management process., (© 2019 by the Ecological Society of America.)

A major goal of ecosystem-based fisheries management is to prevent fishery-induced shifts in community states. This requires an understanding of ecological resilience: the ability of an ecosystem to return to the same state following a perturbation, which can strongly depend on species interactions across trophic levels. We use a structured model of a temperate rocky reef to explore how multi-trophic level fisheries impact ecological resilience. Increasing fishing mortality of prey (urchins) has a minor effect on equilibrium biomass of kelp, urchins, and spiny lobster predators, but increases resilience by reducing the range of predator harvest rates at which alternative stable states are possible. Size-structured predation on urchins acts as the feedback maintaining each state. Our results demonstrate that the resilience of ecosystems strongly depends on the interactive effects of predator and prey harvest in multi-trophic level fisheries, which are common in marine ecosystems but are unaccounted for by traditional management., (© 2017 by the Ecological Society of America.)

Integral projection models (IPMs) have a number of advantages over matrix-model approaches for analyzing size-structured population dynamics, because the latter require parameter estimates for each age or stage transition. However, IPMs still require appropriate data. Typically they are parameterized using individual-scale relationships between body size and demographic rates, but these are not always available. We present an alternative approach for estimating demographic parameters from time series of size-structured survey data using a Bayesian state-space IPM (SSIPM). By fitting an IPM in a state-space framework, we estimate unknown parameters and explicitly account for process and measurement error in a dataset to estimate the underlying process model dynamics. We tested our method by fitting SSIPMs to simulated data; the model fit the simulated size distributions well and estimated unknown demographic parameters accurately. We then illustrated our method using nine years of annual surveys of the density and size distribution of two fish species (blue rockfish, Sebastes mystinus, and gopher rockfish, S. carnatus) at seven kelp forest sites in California. The SSIPM produced reasonable fits to the data, and estimated fishing rates for both species that were higher than our Bayesian prior estimates based on coast-wide stock assessment estimates of harvest. That improvement reinforces the value of being able to estimate demographic parameters from local-scale monitoring data. We highlight a number of key decision points in SSIPM development (e.g., open vs. closed demography, number of particles in the state-space filter) so that users can apply the method to their own datasets., (© 2016 by the Ecological Society of America.)

Extreme events, which have profound ecological consequences, are changing in both frequency and magnitude with climate change. Because extreme temperatures induce coral bleaching, we can explore the relative impacts of changes in frequency and magnitude of high temperature events on coral reefs. Here, we combined climate projections and a dynamic population model to determine how changing bleaching regimes influence coral persistence. We additionally explored how coral traits and competition with macroalgae mediate changes in bleaching regimes. Our results predict that severe bleaching events reduce coral persistence more than frequent bleaching. Corals with low adult mortality and high growth rates are successful when bleaching is mild, but bleaching resistance is necessary to persist when bleaching is severe, regardless of frequency. The existence of macroalgae-dominated stable states reduces coral persistence and changes the relative importance of coral traits. Building on previous studies, our results predict that management efforts may need to prioritize protection of "weaker" corals with high adult mortality when bleaching is mild, and protection of "stronger" corals with high bleaching resistance when bleaching is severe. In summary, future reef projections and conservation targets depend on both local bleaching regimes and biodiversity.

Ecological theory predicts that interactions between antagonistic ecosystem engineers can lead to local competitive exclusion, but disturbance can facilitate broader coexistence. However, few empirical studies have tested the potential for disturbance to mediate competition between engineers. We examined the capacity for disturbance and habitat modification to explain the disjunct distributions of two benthic ecosystem engineers, eelgrass Zostera marina and the burrowing ghost shrimp Neotrypaea californiensis, in two California estuaries. Sediment sampling in eelgrass and ghost shrimp patches revealed that ghost shrimp change benthic biogeochemistry over small scales (centimeters) but not patch scales (meters to tens of meters), suggesting a limited capacity for sediment modification to explain species distributions. To determine the relative competitive abilities of engineers, we conducted reciprocal transplantations of ghost shrimp and eelgrass. Local ghost shrimp densities declined rapidly following the addition of eelgrass, and transplanted eelgrass expanded laterally into the surrounding ghost shrimp-dominated areas. When transplanted into eelgrass patches, ghost shrimp failed to persist. Ghost shrimp were also displaced from plots with structural mimics of eelgrass rhizomes and roots, suggesting that autogenic habitat modification by eelgrass is an important mechanism determining ghost shrimp distributions. However, ghost shrimp were able to rapidly colonize experimental disturbances to eelgrass patch edges, which are common in shallow estuaries. We conclude that coexistence in this system is maintained by spatiotemporally asynchronous disturbances and a competition-colonization trade-off: eelgrass is a competitively superior ecosystem engineer, but benthic disturbances permit the coexistence of ghost shrimp at the landscape scale by modulating the availability of space.

If a species is bycatch in a fishery targeted at its competitor or predator, it experiences both direct anthropogenic mortality and indirect positive effects through species interactions. If the species involved interact strongly, the release from competition or predation can counteract or exceed the negative effects of bycatch. We used a set of two- and three-species community modules to analyze the relative importance of species interactions when modeling the overall effect of harvest with bycatch on a nontarget species. To measure the trade-off between direct mortality and indirect positive effects, we developed a "bycatch transition point" metric to determine, for different scenarios, what levels of bycatch shift overall harvest impact from positive to negative. Under strong direct competition with a targeted competitor, release from competition due to harvest leads to a net increase in abundance even under moderate levels of bycatch. For a three-species model with a shared obligate predator, the release from apparent competition exceeds direct competitive release and outweighs the decrease from bycatch mortality under a wide range of parameters. Therefore, in communities where a shared predator forms a strong link between the target and nontarget species, the effects of indirect interactions on populations can be larger than those of direct interactions. The bycatch transition point metric can be used for tightly linked species to evaluate the relative strengths of positive indirect effects and negative anthropogenic impacts such as bycatch, habitat degradation, and introduction of invasive species.

Given climate change, thermal stress-related mass coral-bleaching events present one of the greatest anthropogenic threats to coral reefs. While corals and their symbiotic algae may respond to future temperatures through genetic adaptation and shifts in community compositions, the climate may change too rapidly for coral response. To test this potential for response, here we develop a model of coral and symbiont ecological dynamics and symbiont evolutionary dynamics. Model results without variation in symbiont thermal tolerance predict coral reef collapse within decades under multiple future climate scenarios, consistent with previous threshold-based predictions. However, model results with genetic or community-level variation in symbiont thermal tolerance can predict coral reef persistence into the next century, provided low enough greenhouse gas emissions occur. Therefore, the level of greenhouse gas emissions will have a significant effect on the future of coral reefs, and accounting for biodiversity and biological dynamics is vital to estimating the size of this effect.