Metamicrobiome diversity promotes the evolution of host-microbial mutualisms.

Ecological theory suggests that a host organism's internal spatial structure can promote the persistence of mutualistic microbes by allowing for the turnover of tissue occupied by non-beneficial or cheating microbes. This type of regulation, whereby a host preferentially rewards tissue occupied by beneficial members of its microbiome but sanctions tissue occupied by non-beneficial cheaters, is expected to generate a competition-extinction trade-off by allowing beneficial microbes to experience a lower extinction rate than competitively dominant cheaters. Using an adaptive dynamics approach, we demonstrate that although ecologically stable, microbial regulation via sanctioning is not stable in any evolutionary sense, as each individual host will be under pressure to reduce the costs incurred from cheater suppression in order to maximize its own fitness at the expense of the rest of the host population. However, increasing the diversity of non-beneficial cheaters in the host population metamicrobiome can lead to an increase in the relative fitness of hosts that actively sanction non-performing tissue, thus facilitating the evolutionary emergence and persistence of such strategies in host-microbial systems. These counter-intuitive results demonstrate how diversity at multiple levels of biological organization and spatiotemporal scales can interact to facilitate the establishment and maintenance of mutualistic relationships.

Host-microbial systems as glass cannons: Explaining microbiome stability in corals exposed to extrinsic perturbations.

Although stability is relatively well understood in macro-organisms, much less is known about its drivers in host-microbial systems where processes operating at multiple levels of biological organisation jointly regulate the microbiome. We conducted an experiment to examine the microbiome stability of three Caribbean corals (Acropora cervicornis, Pseudodiploria strigosa and Porites astreoides) by placing them in aquaria and exposing them to a pulse perturbation consisting of a large dose of broad-spectrum antibiotics before transplanting them into the field. We found that coral hosts harboured persistent, species-specific microbiomes. Stability was generally high but variable across coral species, with A. cervicornis microbiomes displaying the lowest community turnover in both the non-perturbed and the perturbed field transplants. Interestingly, the microbiome of P. astreoides was stable in the non-perturbed field transplants, but unstable in the perturbed field transplants. A mathematical model of host-microbial dynamics helped resolve this paradox by showing that when microbiome regulation is driven by host sanctioning, both resistance and resilience to invasion are low and can lead to instability despite the high direct costs bourne by corals. Conversely, when microbiome regulation is mainly associated with microbial processes, both resistance and resilience to invasion are high and promote stability at no direct cost to corals. We suggest that corals that are mainly regulated by microbial processes can be likened to 'glass cannons' because the high stability they exhibit in the field is due to their microbiome's potent suppression of invasive microbes. However, these corals are susceptible to destabilisation when exposed to perturbations that target the vulnerable members of their microbiomes who are responsible for mounting such powerful attacks against invasive microbes. The differential patterns of stability exhibited by P. astreoides across perturbed and non-perturbed field transplants suggest it is a 'glass cannon' whose microbiome is regulated by microbial processes, whereas A. cervicornis' consistent patterns of stability suggest that its microbiome is mainly regulated by host-level processes. Our results show that understanding how processes that operate at multiple levels of biological organisation interact to regulate microbiomes is critical for predicting the effects of environmental perturbations on host-microbial systems.

Intensification and spatial homogenization of coastal upwelling under climate change.

The timing and strength of wind-driven coastal upwelling along the eastern margins of major ocean basins regulate the productivity of critical fisheries and marine ecosystems by bringing deep and nutrient-rich waters to the sunlit surface, where photosynthesis can occur. How coastal upwelling regimes might change in a warming climate is therefore a question of vital importance. Although enhanced land-ocean differential heating due to greenhouse warming has been proposed to intensify coastal upwelling by strengthening alongshore winds, analyses of observations and previous climate models have provided little consensus on historical and projected trends in coastal upwelling. Here we show that there are strong and consistent changes in the timing, intensity and spatial heterogeneity of coastal upwelling in response to future warming in most Eastern Boundary Upwelling Systems (EBUSs). An ensemble of climate models shows that by the end of the twenty-first century the upwelling season will start earlier, end later and become more intense at high but not low latitudes. This projected increase in upwelling intensity and duration at high latitudes will result in a substantial reduction of the existing latitudinal variation in coastal upwelling. These patterns are consistent across three of the four EBUSs (Canary, Benguela and Humboldt, but not California). The lack of upwelling intensification and greater uncertainty associated with the California EBUS may reflect regional controls associated with the atmospheric response to climate change. Given the strong linkages between upwelling and marine ecosystems, the projected changes in the intensity, timing and spatial structure of coastal upwelling may influence the geographical distribution of marine biodiversity.

Temperature dependency of intraguild predation between native and invasive crabs.

Environmental factors such as temperature can affect the geographical distribution of species directly by exceeding physiological tolerances, or indirectly by altering physiological rates that dictate the sign and strength of species interactions. Although the direct effects of environmental conditions are relatively well studied, the effects of environmentally mediated species interactions have garnered less attention. In this study, we examined the temperature dependency of size-structured intraguild predation (IGP) between native blue crabs (Callinectes sapidus, the IG predator) and invasive green crabs (Carcinus maenas, the IG prey) to evaluate how the effect of temperature on competitive and predatory rates may influence the latitudinal distribution of these species. In outdoor mesocosm experiments, we quantified interactions between blue crabs, green crabs, and shared prey (mussels) at three temperatures reflective of those across their range, using two size classes of blue crab. At low temperatures, green crabs had a competitive advantage and IGP by blue crabs on green crabs was low. At high temperatures, size-matched blue and green crabs were competitively similar, large blue crabs had a competitive advantage, and IGP on green crabs was high. We then used parameter values generated from these experiments (temperature- and size-dependent attack rates and handling times) in a size-structured IGP model in which we varied IGP attack rate, maturation rate of the blue crab from the non-predatory to predatory size class, and resource carrying capacity at each of the three temperatures. In the model, green crabs were likely to competitively exclude blue crabs at low temperature, whereas blue crabs were likely to competitively and consumptively exclude green crabs at higher temperatures, particularly when resource productivities and rates of IGP were high. While many factors may play a role in delimiting species ranges, our results suggest that temperature-dependent interactions can influence local coexistence and are worth considering when developing mechanistic species distribution models and evaluating responses to environmental change.

Predicting shifting sustainability trade-offs in marine finfish aquaculture under climate change.

Defining sustainability goals is a crucial but difficult task because it often involves the quantification of multiple interrelated and sometimes conflicting components. This complexity may be exacerbated by climate change, which will increase environmental vulnerability in aquaculture and potentially compromise the ability to meet the needs of a growing human population. Here, we developed an approach to inform sustainable aquaculture by quantifying spatio-temporal shifts in critical trade-offs between environmental costs and benefits using the time to reach the commercial size as a possible proxy of economic implications of aquaculture under climate change. Our results indicate that optimizing aquaculture practices by minimizing impact (this study considers as impact a benthic carbon deposition ≥ 1 g C m-2  day-1 ) will become increasingly difficult under climate change. Moreover, an increasing temperature will produce a poleward shift in sustainability trade-offs. These findings suggest that future sustainable management strategies and plans will need to account for the effects of climate change across scales. Overall, our results highlight the importance of integrating environmental factors in order to sustainably manage critical natural resources under shifting climatic conditions.

The cryptic role of biodiversity in the emergence of host-microbial mutualisms.

The persistence of mutualisms in host-microbial - or holobiont - systems is difficult to explain because microbial mutualists, who bear the costs of providing benefits to their host, are always prone to being competitively displaced by non-mutualist 'cheater' species. This disruptive effect of competition is expected to be particularly strong when the benefits provided by the mutualists entail costs such as reduced competitive ability. Using a metacommunity model, we show that competition between multiple cheaters within the host's microbiome, when combined with the spatial structure of host-microbial interactions, can have a constructive rather than a disruptive effect by allowing the emergence and maintenance of mutualistic microorganisms within the host. These results indicate that many of the microorganisms inhabiting a host's microbiome, including those that would otherwise be considered opportunistic or even potential pathogens, play a cryptic yet critical role in promoting the health and persistence of the holobiont across spatial scales.

Designing effective reserve networks for nonequilibrium metacommunities.

The proliferation of efficient fishing practices has promoted the depletion of commercial stocks around the world and caused significant collateral damage to marine habitats. Recent empirical studies have shown that marine reserves can play an important role in reversing these effects. Equilibrium metapopulation models predict that networks of marine reserves can provide similar benefits so long as individual reserves are sufficiently large to achieve self-sustainability, or spaced based on the extent of dispersal of the target species in order to maintain connectivity between neighboring reserves. However, these guidelines have not been tested in nonequilibrium metacommunity models that exhibit the kinds of complex spatiotemporal dynamics typically seen in natural marine communities. Here, we used a spatially explicit predator-prey model whose predictions have been validated in a marine system to show that current guidelines are not optimal for metacommunities. In equilibrium metacommunities, there is a community-level trade-off for designing effective reserves: Networks whose size and spacing are smaller than the extent of dispersal maximize global predator abundance but minimize global prey abundance because of trophic cascades, whereas the converse is true for reserve networks whose size and spacing are larger than the extent of dispersal. In nonequilibrium metacommunities, reserves whose size and spacing match the extent of spatial autocorrelation in adult abundance (i.e., the extent of patchiness) escape this community-level trade-off by maximizing global abundance and persistence of both the prey and the predator. Overall, these results suggest that using the extent of adult patchiness instead of the extent of larval dispersal as the size and spacing of reserve networks is critical for designing community-based management strategies. By emphasizing patchiness over dispersal distance, our results show how the apparent complexity of nonequilibrium communities can actually simplify management guidelines and reduce uncertainty associated with the assessment of dispersal in marine environments.

Ecological processes can synchronize marine population dynamics over continental scales.

Determining the relative importance of local and regional processes for the distribution of population abundance is a fundamental but contentious issue in ecology. In marine systems, classical theory holds that the influence of demographic processes and dispersal is confined to local populations whereas the environment controls regional patterns of abundance. Here, we use spatial synchrony to compare the distribution of population abundance of the dominant mussel Mytilus californianus observed along the West Coast of the United States to that predicted by dynamical models undergoing different dispersal and environmental treatments to infer the relative influence of local and regional processes. We reveal synchronized fluctuations in the abundance of mussel populations across a whole continent despite limited larval dispersal and strong environmental forcing. We show that dispersal among neighboring populations interacts with local demographic processes to generate characteristic patterns of spatial synchrony that can govern the dynamic distribution of mussel abundance over 1,800 km of coastline. Our study emphasizes the importance of dispersal and local dynamics for the distribution of abundance at the continental scale. It further highlights potential limits to the use of "climate envelope" models for predicting the response of large-scale ecosystems to global climate change.

Network-based restoration strategies maximize ecosystem recovery.

Redressing global patterns of biodiversity loss requires quantitative frameworks that can predict ecosystem collapse and inform restoration strategies. By applying a network-based dynamical approach to synthetic and real-world mutualistic ecosystems, we show that biodiversity recovery following collapse is maximized when extirpated species are reintroduced based solely on their total number of connections in the original interaction network. More complex network-based strategies that prioritize the reintroduction of species that improve 'higher order' topological features such as compartmentalization do not provide meaningful performance improvements. These results suggest that it is possible to design nearly optimal restoration strategies that maximize biodiversity recovery for data-poor ecosystems in order to ensure the delivery of critical natural services that fuel economic development, food security, and human health around the globe.

Coastal upwelling generates cryptic temperature refugia.

Understanding the effects of climate-mediated environmental variation on the distribution of organisms is critically important in an era of global change. We used wavelet analysis to quantify the spatiotemporal (co)variation in daily water temperature for predicting the distribution of cryptic refugia across 16 intertidal sites that were characterized as 'no', 'weak' or 'strong' upwelling and spanned 2000 km of the European Atlantic Coast. Sites experiencing weak upwelling exhibited high synchrony in temperature but low levels of co-variability at monthly to weekly timescales, whereas the opposite was true for sites experiencing strong upwelling. This suggests upwelling generates temporal thermal refugia that can promote organismal performance by both supplying colder water that mitigates thermal stress during hot Summer months and ensuring high levels of fine-scale variation in temperature that reduce the duration of thermal extremes. Additionally, pairwise correlograms based on the Pearson-product moment correlation coefficient and wavelet coherence revealed scale dependent trends in temperature fluctuations across space, with a rapid decay in strong upwelling sites at monthly and weekly timescales. This suggests upwelling also generates spatial thermal refugia that can 'rescue' populations from unfavorable conditions at local and regional scales. Overall, this study highlights the importance of identifying cryptic spatiotemporal refugia that emerge from fine-scale environmental variation to map potential patterns of organismal performance in a rapidly changing world.

Recruitment facilitation can promote coexistence and buffer population growth in metacommunities.

Although positive species interactions are ubiquitous in nature, theory has generally focused on the role of negative interactions to explain patterns of species diversity. Here, we incorporate recruitment facilitation, a positive interaction prevalent in marine and terrestrial systems, into a metacommunity framework to assess how the interplay between colonisation, competition and facilitation mediates coexistence. We show that when subordinate species facilitate the recruitment of dominant species, multi-species metacommunities can persist stably even if the colonisation rate of the dominant species is greater than that of the subordinate species. In addition, recruitment facilitation can buffer population growth from changes in colonisation rates, and thus explain the paradoxical mismatch between patterns of abundance and recruitment in marine systems. Overall, our results demonstrate that recruitment facilitation can have profound effects on the assembly, dissolution and regulation of metacommunities by mediating the relative influence of local and regional processes on population abundance and species diversity.

Synchrony and stability of food webs in metacommunities.

Synchrony has fundamental but conflicting implications for the persistence and stability of food webs at local and regional scales. In a constant environment, compensatory dynamics between species can maintain food web stability, but factors that synchronize population fluctuations within and between communities are expected to be destabilizing. We studied the dynamics of a food web in a metacommunity to determine how environmental variability and dispersal affect stability by altering compensatory dynamics and average species abundance. When dispersal rate is high, weak correlated environmental fluctuations promote food web stability by reducing the amplitude of compensatory dynamics. However, when dispersal rate is low, weak environmental fluctuations reduce food web stability by inducing intraspecific synchrony across communities. Irrespective of dispersal rate, strong environmental fluctuations disrupt compensatory dynamics and decrease stability by inducing intermittent correlated fluctuations between consumers in local food webs, which reduce both total consumer abundance and predator abundance. Strong correlated environmental fluctuations lead to (i) spatially asynchronous and highly correlated local consumer dynamics when dispersal is low and (ii) spatially synchronous but intermediate local consumer correlation when dispersal is high. By controlling intraspecific synchrony, dispersal mediates the capacity of strong environmental fluctuations to disrupt compensatory dynamics at both local and metacommunity scales.

Predicting the stability of multitrophic communities in a variable world.

Identifying the factors that destabilize communities is critical for predicting and mitigating the ecological impacts of environmental change. Although theory has shown that local ecosystem size and regional dispersal can determine biodiversity, less is known about the direct and indirect effects of these factors on community stability. Here we show that multitrophic community instability of invertebrates and fishes in coastal ponds is negatively related to local pond size and positively related to distance to the ocean, a proxy for dispersal limitation. Importantly, the effects of pond size and distance on instability were direct rather than indirectly mediated by species richness. This suggests that the diversity-stability relationship is an epiphenomenon whose resolution is neither necessary nor sufficient to understand the stability of these multitrophic communities. Instead, well-established and easily measured local and regional factors historically linked to species richness can be used to predict multitrophic community stability in a variable world.

Biogeography of ocean acidification: Differential field performance of transplanted mussels to upwelling-driven variation in carbonate chemistry.

Ocean acidification (OA) represents a serious challenge to marine ecosystems. Laboratory studies addressing OA indicate broadly negative effects for marine organisms, particularly those relying on calcification processes. Growing evidence also suggests OA combined with other environmental stressors may be even more deleterious. Scaling these laboratory studies to ecological performance in the field, where environmental heterogeneity may mediate responses, is a critical next step toward understanding OA impacts on natural communities. We leveraged an upwelling-driven pH mosaic along the California Current System to deconstruct the relative influences of pH, ocean temperature, and food availability on seasonal growth, condition and shell thickness of the ecologically dominant intertidal mussel Mytilus californianus. In 2011 and 2012, ecological performance of adult mussels from local and commonly sourced populations was measured at 8 rocky intertidal sites between central Oregon and southern California. Sites coincided with a large-scale network of intertidal pH sensors, allowing comparisons among pH and other environmental stressors. Adult California mussel growth and size varied latitudinally among sites and inter-annually, and mean shell thickness index and shell weight growth were reduced with low pH. Surprisingly, shell length growth and the ratio of tissue to shell weight were enhanced, not diminished as expected, by low pH. In contrast, and as expected, shell weight growth and shell thickness were both diminished by low pH, consistent with the idea that OA exposure can compromise shell-dependent defenses against predators or wave forces. We also found that adult mussel shell weight growth and relative tissue mass were negatively associated with increased pH variability. Including local pH conditions with previously documented influences of ocean temperature, food availability, aerial exposure, and origin site enhanced the explanatory power of models describing observed performance differences. Responses of local mussel populations differed from those of a common source population suggesting mussel performance partially depended on genetic or persistent phenotypic differences. In light of prior research showing deleterious effects of low pH on larval mussels, our results suggest a life history transition leading to greater resilience in at least some performance metrics to ocean acidification by adult California mussels. Our data also demonstrate "hot" (more extreme) and "cold" (less extreme) spots in both mussel responses and environmental conditions, a pattern that may enable mitigation approaches in response to future changes in climate.

Regional environmental variation and local species interactions influence biogeographic structure on oyster reefs.

Although species interactions are often assumed to be strongest at small spatial scales, they can interact with regional environmental factors to modify food web dynamics across biogeographic scales. The eastern oyster (Crassostrea virginica) is a widespread foundational species of both ecological and economic importance. The oyster and its associated assemblage of fish and macroinvertebrates is an ideal system to investigate how regional differences in environmental variables influence trophic interactions and food web structure. We quantified multiple environmental factors, oyster reef properties, associated species, and trophic guilds on intertidal oyster reefs within 10 estuaries along 900 km of the southeastern United States. Geographical gradients in fall water temperature and mean water depth likely influenced regional (i.e., the northern, central and southern sections of the SAB) variation in oyster reef food web structure. Variation in the biomass of mud crabs, an intermediate predator, was mostly (84.1%) explained by reefs within each site, and did not differ substantially among regions; however, regional variation in the biomass of top predators and of juvenile oysters also contributed to biogeographic variation in food web structure. In particular, region explained almost half (40.2%) of the variation in biomass of predators of blue crab, a top predator that was prevalent only in the central region where water depth was greater. Field experiments revealed that oyster mortality due to predation was greatest in the central region, suggesting spatial variation in the importance of trophic cascades. However, high oyster recruitment in the middle region probably compensates for this enhanced predation, potentially explaining why relatively less variation (17.9%) in oyster cluster biomass was explained by region. Region also explained over half of the variation in biomass of mud crab predators (55.2%), with the southern region containing almost an order of magnitude more biomass than the other two regions. In this region, higher water temperatures in the fall corresponded with higher biomass of fish that consume mud crabs and of fish that consume juvenile and forage fish, whereas biomas of their prey (mud crabs and juvenile and forage fish, respectively) was generally low in the southern region. Collectively, these results show how environmental gradients interact with trophic cascades to structure food webs associated with foundation species across biogeographic regions.

Not even wrong: the spurious measurement of biodiversity's effects on ecosystem functioning.

Understanding how biodiversity influences ecosystem functioning is one of the central goals of modern ecology. The early and often acrimonious debates about the relationship between biodiversity and ecosystem functioning were largely resolved following the advent of a statistical partitioning scheme that decomposed the net effect of biodiversity on ecosystem functioning into a "selection" effect and a "complementarity" effect. Here we show that both the biodiversity effect and its statistical decomposition into selection and complementarity are fundamentally flawed because these methods use a naïve null expectation based on neutrality, likely leading to an overestimate of the net biodiversity effect, and because they fail to account for the nonlinear abundance-ecosystem-functioning relationships widely observed in nature. Furthermore, under nonlinearity no such statistical scheme can be devised to partition the biodiversity effect. We also present an alternative approach that provides a more reasonable starting point for estimating biodiversity effects. Overall, our results suggest that all studies conducted since the early 1990s are likely to have overestimated the positive effects of biodiversity on ecosystem functioning.

Structure and stability of the coral microbiome in space and time.

Although it is well established that the microbial communities inhabiting corals perform key functions that promote the health and persistence of their hosts, little is known about their spatial structure and temporal stability. We examined the natural variability of microbial communities associated with six Caribbean coral species from three genera at four reef sites over one year. We identified differences in microbial community composition between coral genera and species that persisted across space and time, suggesting that local host identity likely plays a dominant role in structuring the microbiome. However, we found that microbial community dissimilarity increased with geographical distance, which indicates that regional processes such as dispersal limitation and spatiotemporal environmental heterogeneity also influence microbial community composition. In addition, network analysis revealed that the strength of host identity varied across coral host genera, with species from the genus Acropora having the most influence over their microbial community. Overall, our results demonstrate that despite high levels of microbial diversity, coral species are characterized by signature microbiomes that are stable in both space and time.

Regional processes are stronger determinants of rocky intertidal community dynamics than local biotic interactions.

Understanding the relative roles of species interactions and environmental factors in structuring communities has historically focused on local scales where manipulative experiments are possible. However, recent interest in predicting the effects of climate change and species invasions has spurred increasing attention to processes occurring at larger spatial and temporal scales. The "meta-ecosystem" approach is an ideal framework for integrating processes operating at multiple scales as it explicitly considers the influence of local biotic interactions and regional flows of energy, materials, and organisms on community structure. Using a comparative-experimental design, we asked (1) what is the relative importance of local biotic interactions and oceanic processes in determining rocky intertidal community structure in the low zone within the Northern California Current System, and (2) what factors are most important in regulating this structure and why? We focused on functional group interactions between macrophytes and sessile invertebrates and their consumers (grazers, predators), how these varied across spatial scales, and with ocean-driven conditions (upwelling, temperature) and ecological subsidies (nutrients, phytoplankton, sessile invertebrate recruits). Experiments were conducted at 13 sites divided across four capes in Oregon and northern California. Results showed that biotic interactions were variable in space and time but overall, sessile invertebrates had no effect on macrophytes while macrophytes had weakly negative effects on sessile invertebrates. Consumers, particularly predators, also had weakly negative effects on both functional groups. Overall, we found that 40-49% of the variance in community structure at the local scale was explained by external factors (e.g., spatial scale, time, upwelling, temperature, ecological subsidies) vs. 19-39% explained by functional group interactions. When individual functional group interaction strengths were used, only 2-3% of the variation was explained by any one functional group while 28-54% of the variation was explained by external factors. We conclude that community structure in the low intertidal zone is driven primarily by external factors at the regional scale with local biotic interactions playing a secondary role.

Increased spatial and temporal autocorrelation of temperature under climate change.

Understanding spatiotemporal variation in environmental conditions is important to determine how climate change will impact ecological communities. The spatial and temporal autocorrelation of temperature can have strong impacts on community structure and persistence by increasing the duration and the magnitude of unfavorable conditions in sink populations and disrupting spatial rescue effects by synchronizing spatially segregated populations. Although increases in spatial and temporal autocorrelation of temperature have been documented in historical data, little is known about how climate change will impact these trends. We examined daily air temperature data from 21 General Circulation Models under the business-as-usual carbon emission scenario to quantify patterns of spatial and temporal autocorrelation between 1871 and 2099. Although both spatial and temporal autocorrelation increased over time, there was significant regional variation in the temporal autocorrelation trends. Additionally, we found a consistent breakpoint in the relationship between spatial autocorrelation and time around the year 2030, indicating an acceleration in the rate of increase of the spatial autocorrelation over the second half of the 21st century. Overall, our results suggest that ecological populations might experience elevated extinction risk under climate change because increased spatial and temporal autocorrelation of temperature is expected to erode both spatial and temporal refugia.

Local disturbance cycles and the maintenance of heterogeneity across scales in marine metapopulations.

In marine systems, the occurrence and implications of disturbance-recovery cycles have been revealed at the landscape level, but only in demographically open or closed systems where landscape-level dynamics are assumed to have no feedback effect on regional dynamics. We present a mussel metapopulation model to elucidate the role of landscape-level disturbance cycles for regional response of mussel populations to onshore productivity and larval transport. Landscape dynamics are generated through spatially explicit rules, and each landscape is connected to its neighbor through unidirectional larval dispersal. The role of landscape disturbance cycles in the regional system behavior is elucidated (1) in demographically open vs. demographically coupled systems, in relation to (2) onshore reproductive output and (3) the temporal scale of landscape disturbance dynamics. By controlling for spatial structure at the landscape and metapopulation levels, we first demonstrate the interaction between landscape and oceanographic connectivity. The temporal scale of disturbance cycles, as controlled by mussel colonization rate, plays a critical role in the regional behavior of the system. Indeed, fast disturbance cycles are responsible for regional synchrony in relation to onshore reproductive output. Slow disturbance cycles, however, lead to increased robustness to changes in productivity and to demographic coupling. These testable predictions indicate that the occurrence and temporal scale of local disturbance-recovery dynamics can drive large-scale variability in demographically open systems, and the response of metapopulations to changes in nearshore productivity.