Kyle C. Cavanaugh

Poleward expansion of mangroves is a threshold response to decreased frequency of extreme cold events

Significance Coastal mangrove forests support a diverse array of associated species and provide ecosystem services to human communities. Mangroves cannot tolerate extreme freezing temperatures and so are generally limited to tropical environments. However, climate change in the form of increasing temperatures has the potential to facilitate increases in mangrove abundance near tropical–temperate transition zones. Here, we use 28 y of satellite imagery to demonstrate that increases in mangrove area have already occurred along the northeast coast of Florida. These increases correspond to decreases in the frequency of extreme cold events in this region. We also identify a temperature-related ecological threshold of −4°C. These results suggest that landscape-scale increases in mangrove area may occur in other regions where this threshold is crossed. Regional warming associated with climate change is linked with altered range and abundance of species and ecosystems worldwide. However, the ecological impacts of changes in the frequency of extreme events have not been as well documented, especially for coastal and marine environments. We used 28 y of satellite imagery to demonstrate that the area of mangrove forests has doubled at the northern end of their historic range on the east coast of Florida. This expansion is associated with a reduction in the frequency of “extreme” cold events (days colder than −4 °C), but uncorrelated with changes in mean annual temperature, mean annual precipitation, and land use. Our analyses provide evidence for a threshold response, with declining frequency of severe cold winter events allowing for poleward expansion of mangroves. Future warming may result in increases in mangrove cover beyond current latitudinal limits of mangrove forests, thereby altering the structure and function of these important coastal ecosystems.

Operationalizing the social-ecological systems framework to assess sustainability

Significance Meeting human needs while sustaining ecosystems and the benefits they provide is a global challenge. Coastal marine systems present a particularly important case, given that >50% of the world’s population lives within 100 km of the coast and fisheries are the primary source of protein for >1 billion people worldwide. Our integrative analysis here yields an understanding of the sustainability of coupled social-ecological systems that is quite distinct from that provided by either the biophysical or the social sciences alone and that illustrates the feasibility and value of operationalizing the social-ecological systems framework for comparative analyses of coupled systems, particularly in data-poor and developing nation settings. Environmental governance is more effective when the scales of ecological processes are well matched with the human institutions charged with managing human–environment interactions. The social-ecological systems (SESs) framework provides guidance on how to assess the social and ecological dimensions that contribute to sustainable resource use and management, but rarely if ever has been operationalized for multiple localities in a spatially explicit, quantitative manner. Here, we use the case of small-scale fisheries in Baja California Sur, Mexico, to identify distinct SES regions and test key aspects of coupled SESs theory. Regions that exhibit greater potential for social-ecological sustainability in one dimension do not necessarily exhibit it in others, highlighting the importance of integrative, coupled system analyses when implementing spatial planning and other ecosystem-based strategies.

Wave disturbance overwhelms top-down and bottom-up control of primary production in California kelp forests.

We took advantage of regional differences in environmental forcing and consumer abundance to examine the relative importance of nutrient availability (bottom-up), grazing pressure (top-down), and storm waves (disturbance) in determining the standing biomass and net primary production (NPP) of the giant kelp Macrocystis pyrifera in central and southern California. Using a nine-year data set collected from 17 sites we show that, despite high densities of sea urchin grazers and prolonged periods of low nutrient availability in southern California, NPP by giant kelp was twice that of central California where nutrient concentrations were consistently high and sea urchins were nearly absent due to predation by sea otters. Waves associated with winter storms were consistently higher in central California, and the loss of kelp biomass to winter wave disturbance was on average twice that of southern California. These observations suggest that the more intense wave disturbance in central California limited NPP by giant kelp under otherwise favorable conditions. Regional patterns of interannual variation in NPP were similar to those of wave disturbance in that year-to-year variation in disturbance and NPP were both greater in southern California. Our findings provide strong evidence that regional differences in wave disturbance overwhelmed those of nutrient supply and grazing intensity to determine NPP by giant kelp. The important role of disturbance in controlling NPP revealed by our study is likely not unique to giant kelp forests, as vegetation dynamics in many systems are dominated by post-disturbance succession with climax communities being relatively uncommon. The effects of disturbance frequency may be easier to detect in giant kelp because it is fast growing and relatively short lived, with cycles of disturbance and recovery occurring on time scales of years. Much longer data sets (decades to centuries) will likely be needed to properly evaluate the role of disturbance relative to other processes in determining patterns of NPP in other systems.

Global patterns of kelp forest change over the past half-century

Significance Kelp forests support diverse and productive ecological communities throughout temperate and arctic regions worldwide, providing numerous ecosystem services to humans. Literature suggests that kelp forests are increasingly threatened by a variety of human impacts, including climate change, overfishing, and direct harvest. We provide the first globally comprehensive analysis of kelp forest change over the past 50 y, identifying a high degree of variation in the magnitude and direction of change across the geographic range of kelps. These results suggest region-specific responses to global change, with local drivers playing an important role in driving patterns of kelp abundance. Increased monitoring aimed at understanding regional kelp forest dynamics is likely to prove most effective for the adaptive management of these important ecosystems. Kelp forests (Order Laminariales) form key biogenic habitats in coastal regions of temperate and Arctic seas worldwide, providing ecosystem services valued in the range of billions of dollars annually. Although local evidence suggests that kelp forests are increasingly threatened by a variety of stressors, no comprehensive global analysis of change in kelp abundances currently exists. Here, we build and analyze a global database of kelp time series spanning the past half-century to assess regional and global trends in kelp abundances. We detected a high degree of geographic variation in trends, with regional variability in the direction and magnitude of change far exceeding a small global average decline (instantaneous rate of change = −0.018 y−1). Our analysis identified declines in 38% of ecoregions for which there are data (−0.015 to −0.18 y−1), increases in 27% of ecoregions (0.015 to 0.11 y−1), and no detectable change in 35% of ecoregions. These spatially variable trajectories reflected regional differences in the drivers of change, uncertainty in some regions owing to poor spatial and temporal data coverage, and the dynamic nature of kelp populations. We conclude that although global drivers could be affecting kelp forests at multiple scales, local stressors and regional variation in the effects of these drivers dominate kelp dynamics, in contrast to many other marine and terrestrial foundation species.

Integrating physiological threshold experiments with climate modeling to project mangrove species’ range expansion

Predictions of climate-related shifts in species ranges have largely been based on correlative models. Due to limitations of these models, there is a need for more integration of experimental approaches when studying impacts of climate change on species distributions. Here, we used controlled experiments to identify physiological thresholds that control poleward range limits of three species of mangroves found in North America. We found that all three species exhibited a threshold response to extreme cold, but freeze tolerance thresholds varied among species. From these experiments, we developed a climate metric, freeze degree days (FDD), which incorporates both the intensity and the frequency of freezes. When included in distribution models, FDD accurately predicted mangrove presence/absence. Using 28 years of satellite imagery, we linked FDD to observed changes in mangrove abundance in Florida, further exemplifying the importance of extreme cold. We then used downscaled climate projections of FDD to project that these range limits will move northward by 2.2-3.2 km yr(-1) over the next 50 years.

Review of the ecosystem service implications of mangrove encroachment into salt marshes

Salt marsh and mangrove have been recognized as being among the most valuable ecosystem types globally in terms of their supply of ecosystem services and support for human livelihoods. These coastal ecosystems are also susceptible to the impacts of climate change and rising sea levels, with evidence of global shifts in the distribution of mangroves, including encroachment into salt marshes. The encroachment of woody mangrove shrubs and trees into herbaceous salt marshes may represent a substantial change in ecosystem structure, although resulting impacts on ecosystem functions and service provisions are largely unknown. In this review, we assess changes in ecosystem services associated with mangrove encroachment. While there is quantitative evidence to suggest that mangrove encroachment may enhance carbon storage and the capacity of a wetland to increase surface elevation in response to sea-level rise, for most services there has been no direct assessment of encroachment impact. On the basis of current understanding of ecosystem structure and function, we theorize that mangrove encroachment may increase nutrient storage and improve storm protection, but cause declines in habitat availability for fauna requiring open vegetation structure (such as migratory birds and foraging bats) as well as the recreational and cultural activities associated with this fauna (e.g., birdwatching and/or hunting). Changes to provisional services such as fisheries productivity and cultural services are likely to be site specific and dependent on the species involved. We discuss the need for explicit experimental testing of the effects of encroachment on ecosystem services in order to address key knowledge gaps, and present an overview of the options available to coastal resource managers during a time of environmental change.

Synchrony in dynamics of giant kelp forests is driven by both local recruitment and regional environmental controls

Populations of many species display spatially synchronous fluctuations in abundance. Synchrony is most commonly attributed to three processes: factors that influence recruitment (e.g., dispersal, early survival), large-scale environmental variability, and spatially autocorrelated trophic interactions. However it is often difficult to link population synchrony to a specific dominant process, particularly when multiple synchronizing forces are operating. We utilized a new satellite-based data set of giant kelp (Macrocystis pyrifera) canopy biomass to examine population synchrony in southern California kelp forests on spatial scales ranging from 50 m to 300 km and temporal scales ranging from 1 to 11 years. We examined the relationship between synchrony and distance for adult kelp populations, kelp recruits, sea urchin abundance (a major grazer of kelp), and environmental variables known to influence kelp population dynamics. Population synchrony in giant kelp decreased with distance between populations: an initial rapid exponential decrease between 50 m and 1.3 km was followed by a second, large-scale decrease between distances of 1.3 km and 172 km. The 50-m to 1.3-km spatial scale corresponded to the scales of synchrony in the abundance of sea urchins and young kelp recruits, suggesting that local drivers of predation and recruitment influence small-scale synchrony in kelp populations. The spatial correlation patterns of environmental variables, particularly wave height, were similar to the synchrony-distance relationship of kelp populations from 1.3 km to 172 km, suggesting that regional environmental variability, i.e., the Moran effect, was the dominant process affecting synchrony at larger spatial scales. This two-step pattern in the relationship between kelp biomass synchrony and distance was apparent in each of the 11 years of our study. Our results highlight the potential for synthesizing approaches from both landscape and population ecology in order to identify the multiple processes that generate synchrony in population dynamics.

Connectivity structures local population dynamics: a long-term empirical test in a large metapopulation system

Ecological theory predicts that demographic connectivity structures the dynamics of local populations within metapopulation systems, but empirical support has been constrained by major limitations in data and methodology. We tested this prediction for giant kelp Macrocystis pyrifera, a key habitat-forming species in temperate coastal ecosystems worldwide, in southern California, USA. We combined a long-term (22 years), large-scale (~500 km coastline), high-resolution census of abundance with novel patch delineation methods and an innovative connectivity measure incorporating oceanographic transport and source fecundity. Connectivity strongly predicted local dynamics (well-connected patches had lower probabilities of extinction and higher probabilities of colonization, leading to greater likelihoods of occupancy) but this relationship was mediated by patch size. Moreover, the relationship between connectivity and local population dynamics varied over time, possibly due to temporal variation in oceanographic transport processes. Surprisingly, connectivity had a smaller influence on colonization relative to extinction, possibly because local ecological factors differ greatly between extinct and extant patches. Our results provide the first comprehensive evidence that southern California giant kelp populations function as a metapopulation system, challenging the view that populations of this important foundation species are governed exclusively by self-replenishment.

Fluctuations in population fecundity drive variation in demographic connectivity and metapopulation dynamics.

Demographic connectivity is vital to sustaining metapopulations yet often changes dramatically through time due to variation in the production and dispersal of offspring. However, the relative importance of variation in fecundity and dispersal in determining the connectivity and dynamics of metapopulations is poorly understood due to the paucity of comprehensive spatio-temporal data on these processes for most species. We quantified connectivity in metapopulations of a marine foundation species (giant kelp Macrocystis pyrifera) across 11 years and approximately 900 km of coastline by estimating population fecundity with satellite imagery and propagule dispersal using a high-resolution ocean circulation model. By varying the temporal complexity of different connectivity measures and comparing their ability to explain observed extinction–colonization dynamics, we discovered that fluctuations in population fecundity, rather than fluctuations in dispersal, are the dominant driver of variation in connectivity and contribute substantially to metapopulation recovery and persistence. Thus, for species with high variability in reproductive output and modest variability in dispersal (most plants, many animals), connectivity measures ignoring fluctuations in fecundity may overestimate connectivity and likelihoods of persistence, limiting their value for understanding and conserving metapopulations. However, we demonstrate how connectivity measures can be simplified while retaining utility, validating a practical solution for data-limited systems.

Impacts of mangrove encroachment and mosquito impoundment management on coastal protection services

The ecosystem services afforded by coastal wetlands are threatened by climate change and other anthropogenic stressors. The Kennedy Space Center and Merritt Island National Wildlife Refuge in east central Florida offer a representative site for investigating how changes to vegetation distribution interact with management to impact coastal protection. Here, salt marshes are converting to mangroves, and mosquito impoundment structures are being modified. The resulting changes to vegetation composition and topography influence coastal protection services in wetlands. We used a model-based assessment of wave attenuation and erosion to compare vegetation (mangrove, salt marsh) and impoundment state (intact, graded). Our findings suggest that the habitat needed to attenuate 90% of wave height is significantly larger for salt marshes than mangroves. Erosion prevention was significantly higher (470%) in scenarios with mangroves than in salt marshes. Intact berms attenuated waves over shorter distances, but did not significantly reduce erosion. Differences in coastal protection were driven more by vegetation than by impoundment state. Overall, our findings reveal that mangroves provide more coastal protection services, and therefore more coastal protection value, than salt marshes in east central Florida. Other coastal regions undergoing similar habitat conversion may also benefit from increased coastal protection in the future.