Spatial synchrony cascades across ecosystem boundaries and up food webs via resource subsidies.

Cross-ecosystem subsidies are critical to ecosystem structure and function, especially in recipient ecosystems where they are the primary source of organic matter to the food web. Subsidies are indicative of processes connecting ecosystems and can couple ecological dynamics across system boundaries. However, the degree to which such flows can induce cross-ecosystem cascades of spatial synchrony, the tendency for system fluctuations to be correlated across locations, is not well understood. Synchrony has destabilizing effects on ecosystems, adding to the importance of understanding spatiotemporal patterns of synchrony transmission. In order to understand whether and how spatial synchrony cascades across the marine-terrestrial boundary via resource subsidies, we studied the relationship between giant kelp forests on rocky nearshore reefs and sandy beach ecosystems that receive resource subsidies in the form of kelp wrack (detritus). We found that synchrony cascades from rocky reefs to sandy beaches, with spatiotemporal patterns mediated by fluctuations in live kelp biomass, wave action, and beach width. Moreover, wrack deposition synchronized local abundances of shorebirds that move among beaches seeking to forage on wrack-associated invertebrates, demonstrating that synchrony due to subsidies propagates across trophic levels in the recipient ecosystem. Synchronizing resource subsidies likely play an underappreciated role in the spatiotemporal structure, functioning, and stability of ecosystems.

Nutrient availability and senescence spatially structure the dynamics of a foundation species.

Disentangling the roles of the external environment and internal biotic drivers of plant population dynamics is challenging due to the absence of relevant physiological and abundance information over appropriate space and time scales. Remote observations of giant kelp biomass and photosynthetic pigment concentrations are used to show that spatiotemporal patterns of physiological condition, and thus growth and production, are regulated by different processes depending on the scale of observation. Nutrient supply was linked to regional scale (>1 km) physiological condition dynamics, and kelp forest stands were more persistent where nutrient levels were consistently high. However, on local scales (<1 km), internal senescence processes related to canopy age demographics determined patterns of biomass loss across individual kelp forests despite uniform nutrient conditions. Repeat measurements of physiology over continuous spatial fields can provide insights into complex dynamics that are unexplained by the environmental drivers thought to regulate abundance. Emerging remote sensing technologies that provide simultaneous estimates of abundance and physiology can quantify the roles of environmental change and demographics governing plant population dynamics for a wide range of aquatic and terrestrial ecosystems.

Tail-dependent spatial synchrony arises from nonlinear driver-response relationships.

Spatial synchrony may be tail-dependent, that is, stronger when populations are abundant than scarce, or vice-versa. Here, 'tail-dependent' follows from distributions having a lower tail consisting of relatively low values and an upper tail of relatively high values. We present a general theory of how the distribution and correlation structure of an environmental driver translates into tail-dependent spatial synchrony through a non-linear response, and examine empirical evidence for theoretical predictions in giant kelp along the California coastline. In sheltered areas, kelp declines synchronously (lower-tail dependence) when waves are relatively intense, because waves below a certain height do little damage to kelp. Conversely, in exposed areas, kelp is synchronised primarily by periods of calmness that cause shared recovery (upper-tail dependence). We find evidence for geographies of tail dependence in synchrony, which helps structure regional population resilience: areas where population declines are asynchronous may be more resilient to disturbance because remnant populations facilitate reestablishment.

Disturbance and nutrients synchronise kelp forests across scales through interacting Moran effects.

Spatial synchrony is a ubiquitous and important feature of population dynamics, but many aspects of this phenomenon are not well understood. In particular, it is largely unknown how multiple environmental drivers interact to determine synchrony via Moran effects, and how these impacts vary across spatial and temporal scales. Using new wavelet statistical techniques, we characterised synchrony in populations of giant kelp Macrocystis pyrifera, a widely distributed marine foundation species, and related synchrony to variation in oceanographic conditions across 33 years (1987-2019) and >900 km of coastline in California, USA. We discovered that disturbance (storm-driven waves) and resources (seawater nutrients)-underpinned by climatic variability-act individually and interactively to produce synchrony in giant kelp across geography and timescales. Our findings demonstrate that understanding and predicting synchrony, and thus the regional stability of populations, relies on resolving the synergistic and antagonistic Moran effects of multiple environmental drivers acting on different timescales.

Scale-specific drivers of kelp forest communities.

Identifying spatial scales of variation in natural communities and the processes driving them is critical for obtaining a predictive understanding of biodiversity. In this study, we focused on diverse communities inhabiting productive kelp forests on shallow subtidal rocky reefs in southern California, USA. We combined long-term community surveys from 86 sites with detailed environmental data to determine what structures assemblages of fishes, invertebrates and algae at multiple spatial scales. We identified the spatial scales of variation in species composition using a hierarchical analysis based on eigenfunctions, and assessed how sea surface temperature (SST), water column chlorophyll, giant kelp biomass, wave exposure and potential propagule delivery strength contributed to community variation at each scale. Spatial effects occurring at multiple scales explained 60% of the variation in fish assemblages and 52% of the variation in the assemblages of invertebrates and algae. Most variation occurred over broad spatial scales (> 200 km) consistent with spatial heterogeneity in SST and potential propagule delivery strength, while the latter also explained community variation at medium scales (65-200 km). Small scale (1-65 km) community variation was substantial but not linked to any of the measured drivers. Conclusions were consistent for both reef fishes and benthic invertebrates and algae, despite sharp differences in their adult mobility. Our results demonstrate the scale dependence of environmental drivers on kelp forest communities, showing that most species were strongly sorted along oceanographic conditions over various spatial scales. Such spatial effects must be integrated into models assessing the response of marine ecosystems to climate change.

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.

CO2 exchange and evapotranspiration across dryland ecosystems of southwestern North America.

Global-scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO2 sink. However, such analyses are poorly constrained by measured CO2 exchange in drylands. Here we address this observation gap with eddy covariance data from 25 sites in the water-limited Southwest region of North America with observed ranges in annual precipitation of 100-1000 mm, annual temperatures of 2-25°C, and records of 3-10 years (150 site-years in total). Annual fluxes were integrated using site-specific ecohydrologic years to group precipitation with resulting ecosystem exchanges. We found a wide range of carbon sink/source function, with mean annual net ecosystem production (NEP) varying from -350 to +330 gCm-2 across sites with diverse vegetation types, contrasting with the more constant sink typically measured in mesic ecosystems. In this region, only forest-dominated sites were consistent carbon sinks. Interannual variability of NEP, gross ecosystem production (GEP), and ecosystem respiration (Reco ) was larger than for mesic regions, and half the sites switched between functioning as C sinks/C sources in wet/dry years. The sites demonstrated coherent responses of GEP and NEP to anomalies in annual evapotranspiration (ET), used here as a proxy for annually available water after hydrologic losses. Notably, GEP and Reco were negatively related to temperature, both interannually within site and spatially across sites, in contrast to positive temperature effects commonly reported for mesic ecosystems. Models based on MODIS satellite observations matched the cross-site spatial pattern in mean annual GEP but consistently underestimated mean annual ET by ~50%. Importantly, the MODIS-based models captured only 20-30% of interannual variation magnitude. These results suggest the contribution of this dryland region to variability of regional to global CO2 exchange may be up to 3-5 times larger than current estimates.

Drivers of redistribution of fishing and non-fishing effort after the implementation of a marine protected area network.

Marine spatial planning (MSP) is increasingly utilized to sustainably manage ocean uses. Marine protected areas (MPAs), a form of spatial management in which parts of the ocean are regulated to fishing, are now a common tool in MSP for conserving marine biodiversity and managing fisheries. However, the use of MPAs in MSP often neglects, or simplifies, the redistribution of fishing and non-fishing activities inside and outside of MPAs following their implementation. This redistribution of effort can have important implications for effective MSP. Using long-term (14 yr) aerial surveys of boats at the California Channel Islands, we examined the spatial redistribution of fishing and non-fishing activities and their drivers following MPA establishment. Our data represent 6 yr of information before the implementation of an MPA network and 8 yr after implementation. Different types of boats responded in different ways to the closures, ranging from behaviors by commercial dive boats that support the hypothesis of fishing-the-line, to behaviors by urchin, sport fishing, and recreational boats that support the theory of ideal free distribution. Additionally, we found that boats engaged in recreational activities targeted areas that are sheltered from large waves and located near their home ports, while boats engaged in fishing activities also avoided high wave areas but were not constrained by the distance to their home ports. We did not observe the expected pattern of effort concentration near MPA borders for some boat types; this can be explained by the habitat preference of certain activities (for some activities, the desired habitat attributes are not inside the MPAs), species' biology (species such as urchins where the MPA benefit would likely come from larval export rather than adult spillover), or policy-infraction avoidance. The diversity of boat responses reveals variance from the usual simplified assumption that all extractive boats respond similarly to MPA establishment. Our work is the first empirical study to analyze the response of both commercial and recreational boats to closure. Our results will inform MSP in better accounting for effort redistribution by ocean users in response to the implementation of MPAs and other closures.

Terrestrial carbon balance in a drier world: the effects of water availability in southwestern North America.

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO2 exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO2 exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site-years of annual eddy covariance measurements of net and gross CO2 exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 - 1000 mm in annual precipitation and records of 4-9 years each. In addition to evaluating spatial relationships among CO2 and water fluxes across sites, we separately quantified site-level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis-ET relationship was linear, suggesting ET was a better proxy for water available to drive CO2 exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site-level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long-term mean CO2 exchanges with climatic ET. Consequently, a hypothetical 100-mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm(-2) yr(-1). Most of the unexplained NEP variability was related to persistent, site-specific function, suggesting prioritization of research on slow-changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site-level responses to interannual weather can be extrapolated for prediction of CO2 exchanges over decadal and longer timescales relevant to societal response to climate change.

Seascape drivers of Macrocystis pyrifera population genetic structure in the northeast Pacific.

At small spatial and temporal scales, genetic differentiation is largely controlled by constraints on gene flow, while genetic diversity across a species' distribution is shaped on longer temporal and spatial scales. We assess the hypothesis that oceanographic transport and other seascape features explain different scales of genetic structure of giant kelp, Macrocystis pyrifera. We followed a hierarchical approach to perform a microsatellite-based analysis of genetic differentiation in Macrocystis across its distribution in the northeast Pacific. We used seascape genetic approaches to identify large-scale biogeographic population clusters and investigate whether they could be explained by oceanographic transport and other environmental drivers. We then modelled population genetic differentiation within clusters as a function of oceanographic transport and other environmental factors. Five geographic clusters were identified: Alaska/Canada, central California, continental Santa Barbara, California Channel Islands and mainland southern California/Baja California peninsula. The strongest break occurred between central and southern California, with mainland Santa Barbara sites forming a transition zone between the two. Breaks between clusters corresponded approximately to previously identified biogeographic breaks, but were not solely explained by oceanographic transport. An isolation-by-environment (IBE) pattern was observed where the northern and southern Channel Islands clustered together, but not with closer mainland sites, despite the greater distance between them. The strongest environmental association with this IBE pattern was observed with light extinction coefficient, which extends suitable habitat to deeper areas. Within clusters, we found support for previous results showing that oceanographic connectivity plays an important role in the population genetic structure of Macrocystis in the Northern hemisphere.

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.

Dispersal synchronizes giant kelp forests.

Spatial synchrony is the tendency for population fluctuations to be correlated among different locations. This phenomenon is a ubiquitous feature of population dynamics and is important for ecosystem stability, but several aspects of synchrony remain unresolved. In particular, the extent to which any particular mechanism, such as dispersal, contributes to observed synchrony in natural populations has been difficult to determine. To address this gap, we leveraged recent methodological improvements to determine how dispersal structures synchrony in giant kelp (Macrocystis pyrifera), a global marine foundation species that has served as a useful system for understanding synchrony. We quantified population synchrony and fecundity with satellite imagery across 11 years and 880 km of coastline in southern California, USA, and estimated propagule dispersal probabilities using a high-resolution ocean circulation model. Using matrix regression models that control for the influence of geographic distance, resources (seawater nitrate), and disturbance (destructive waves), we discovered that dispersal was an important driver of synchrony. Our findings were robust to assumptions about propagule mortality during dispersal and consistent between two metrics of dispersal: (1) the individual probability of dispersal and (2) estimates of demographic connectivity that incorporate fecundity (the number of propagules dispersing). We also found that dispersal and environmental conditions resulted in geographic clusters with distinct patterns of synchrony. This study is among the few to statistically associate synchrony with dispersal in a natural population and the first to do so in a marine organism. The synchronizing effects of dispersal and environmental conditions on foundation species, such as giant kelp, likely have cascading effects on the spatial stability of biodiversity and ecosystem function.

A marine heatwave changes the stabilizing effects of biodiversity in kelp forests.

Biodiversity can stabilize ecological communities through biological insurance, but climate and other environmental changes may disrupt this process via simultaneous ecosystem destabilization and biodiversity loss. While changes to diversity-stability relationships (DSRs) and the underlying mechanisms have been extensively explored in terrestrial plant communities, this topic remains largely unexplored in benthic marine ecosystems that comprise diverse assemblages of producers and consumers. By analyzing two decades of kelp forest biodiversity survey data, we discovered changes in diversity, stability, and their relationships at multiple scales (biological organizational levels, spatial scales, and functional groups) that were linked with the most severe marine heatwave ever documented in the North Pacific Ocean. Moreover, changes in the strength of DSRs during/after the heatwave were more apparent among functional groups than both biological organizational levels (population vs. ecosystem levels) and spatial scales (local vs. broad scales). Specifically, the strength of DSRs decreased for fishes, increased for mobile invertebrates and understory algae, and were unchanged for sessile invertebrates during/after the heatwave. Our findings suggest that biodiversity plays a key role in stabilizing marine ecosystems, but the resilience of DSRs to adverse climate impacts primarily depends on the functional identities of ecological communities.

Kelpwatch: A new visualization and analysis tool to explore kelp canopy dynamics reveals variable response to and recovery from marine heatwaves.

Giant kelp and bull kelp forests are increasingly at risk from marine heatwave events, herbivore outbreaks, and the loss or alterations in the behavior of key herbivore predators. The dynamic floating canopy of these kelps is well-suited to study via satellite imagery, which provides high temporal and spatial resolution data of floating kelp canopy across the western United States and Mexico. However, the size and complexity of the satellite image dataset has made ecological analysis difficult for scientists and managers. To increase accessibility of this rich dataset, we created Kelpwatch, a web-based visualization and analysis tool. This tool allows researchers and managers to quantify kelp forest change in response to disturbances, assess historical trends, and allow for effective and actionable kelp forest management. Here, we demonstrate how Kelpwatch can be used to analyze long-term trends in kelp canopy across regions, quantify spatial variability in the response to and recovery from the 2014 to 2016 marine heatwave events, and provide a local analysis of kelp canopy status around the Monterey Peninsula, California. We found that 18.6% of regional sites displayed a significant trend in kelp canopy area over the past 38 years and that there was a latitudinal response to heatwave events for each kelp species. The recovery from heatwave events was more variable across space, with some local areas like Bahía Tortugas in Baja California Sur showing high recovery while kelp canopies around the Monterey Peninsula continued a slow decline and patchy recovery compared to the rest of the Central California region. Kelpwatch provides near real time spatial data and analysis support and makes complex earth observation data actionable for scientists and managers, which can help identify areas for research, monitoring, and management efforts.

Automated satellite remote sensing of giant kelp at the Falkland Islands (Islas Malvinas).

Giant kelp populations that support productive and diverse coastal ecosystems at temperate and subpolar latitudes of both hemispheres are vulnerable to changing climate conditions as well as direct human impacts. Observations of giant kelp forests are spatially and temporally uneven, with disproportionate coverage in the northern hemisphere, despite the size and comparable density of southern hemisphere kelp forests. Satellite imagery enables the mapping of existing and historical giant kelp populations in understudied regions, but automating the detection of giant kelp using satellite imagery requires approaches that are robust to the optical complexity of the shallow, nearshore environment. We present and compare two approaches for automating the detection of giant kelp in satellite datasets: one based on crowd sourcing of satellite imagery classifications and another based on a decision tree paired with a spectral unmixing algorithm (automated using Google Earth Engine). Both approaches are applied to satellite imagery (Landsat) of the Falkland Islands or Islas Malvinas (FLK), an archipelago in the southern Atlantic Ocean that supports expansive giant kelp ecosystems. The performance of each method is evaluated by comparing the automated classifications with a subset of expert-annotated imagery (8 images spanning the majority of our continuous timeseries, cumulatively covering over 2,700 km of coastline, and including all relevant sensors). Using the remote sensing approaches evaluated herein, we present the first continuous timeseries of giant kelp observations in the FLK region using Landsat imagery spanning over three decades. We do not detect evidence of long-term change in the FLK region, although we observe a recent decline in total canopy area from 2017-2021. Using a nitrate model based on nearby ocean state measurements obtained from ships and incorporating satellite sea surface temperature products, we find that the area of giant kelp forests in the FLK region is positively correlated with the nitrate content observed during the prior year. Our results indicate that giant kelp classifications using citizen science are approximately consistent with classifications based on a state-of-the-art automated spectral approach. Despite differences in accuracy and sensitivity, both approaches find high interannual variability that impedes the detection of potential long-term changes in giant kelp canopy area, although recent canopy area declines are notable and should continue to be monitored carefully.

Large-scale shift in the structure of a kelp forest ecosystem co-occurs with an epizootic and marine heatwave.

Climate change is responsible for increased frequency, intensity, and duration of extreme events, such as marine heatwaves (MHWs). Within eastern boundary current systems, MHWs have profound impacts on temperature-nutrient dynamics that drive primary productivity. Bull kelp (Nereocystis luetkeana) forests, a vital nearshore habitat, experienced unprecedented losses along 350 km of coastline in northern California beginning in 2014 and continuing through 2019. These losses have had devastating consequences to northern California communities, economies, and fisheries. Using a suite of in situ and satellite-derived data, we demonstrate that the abrupt ecosystem shift initiated by a multi-year MHW was preceded by declines in keystone predator population densities. We show strong evidence that northern California kelp forests, while temporally dynamic, were historically resilient to fluctuating environmental conditions, even in the absence of key top predators, but that a series of coupled environmental and biological shifts between 2014 and 2016 resulted in the formation of a persistent, altered ecosystem state with low primary productivity. Based on our findings, we recommend the implementation of ecosystem-based and adaptive management strategies, such as (1) monitoring the status of key ecosystem attributes: kelp distribution and abundance, and densities of sea urchins and their predators, (2) developing management responses to threshold levels of these attributes, and (3) creating quantitative restoration suitability indices for informing kelp restoration efforts.

Variation in disturbance to a foundation species structures the dynamics of a benthic reef community.

Disturbance and foundation species can both have strong impacts on ecosystem structure and function, but studies of their interacting effects are hindered by the long life spans and slow growth of most foundation species. Here, we investigated the extent to which foundation species may mediate the impacts of disturbance on ecological communities, using the kelp forest ecosystem as a study system. Giant kelp (Macrocystis pyrifera) grows rapidly and experiences wave disturbance from winter storms. We developed and analyzed a model of the effects of variable storm regimes on giant kelp population dynamics and of the cascading effects on kelp-mediated competition between benthic community members in kelp forests. Simulations of severe storm regimes resulted in a greater abundance of understory macroalgae and a lower abundance of sessile invertebrates than did milder regimes. Both the cascading effects of periodic loss of giant kelp as well as the degree to which storms directly impacted the benthos (in the form of scouring) influenced the outcome of competition between benthic community members. The model's qualitative predictions were consistent with empirical data from a 20-yr time series of community dynamics, suggesting that interannual variability in disturbance that affects giant kelp abundance can have strong consequences for benthic community structure. Our findings point to the value of long-term studies in elucidating the interacting effects of disturbance and foundation species.

Remote sensing: generation of long-term kelp bed data sets for evaluation of impacts of climatic variation.

A critical tool in assessing ecosystem change is the analysis of long-term data sets, yet such information is generally sparse and often unavailable for many habitats. Kelp forests are an example of rapidly changing ecosystems that are in most cases data poor. Because kelp forests are highly dynamic and have high intrinsic interannual variability, understanding how regional-scale drivers are driving kelp populations-and particularly how kelp populations are responding to climate change-requires long-term data sets. However, much of the work on kelp responses to climate change has focused on just a few, relatively long-lived, perennial, canopy-forming species. To understand how kelp populations with different life history traits are responding to climate-related variability, we leverage 35 yr of Landsat satellite imagery to track the population size of an annual, ruderal kelp, Nereocystis luetkeana, across Oregon. We found high levels of interannual variability in Nereocystis canopy area and varying population trajectories over the last 35 yr. Surprisingly, Oregon Nereocystis population sizes were unresponsive to a 2014 marine heat wave accompanied by increases in urchin densities that decimated northern California Nereocystis populations. Some Oregon Nereocystis populations have even increased in area relative to pre-2014 levels. Analysis of environmental drivers found that Nereocystis population size was negatively correlated with estimated nitrate levels and positively correlated with winter wave height. This pattern is the inverse of the predicted relationship based on extensive prior work on the perennial kelp Macrocystis pyrifera and may be related to the annual life cycle of Nereocystis. This article demonstrates (1) the value of novel remote sensing tools to create long-term data sets that may challenge our understanding of nearshore marine species and (2) the need to incorporate life history traits into our theory of how climate change will shape the ocean of the future.

Kelp forests at the end of the earth: 45 years later.

The kelp forests of southern South America are some of the least disturbed on the planet. The remoteness of this region has, until recently, spared it from many of the direct anthropogenic stressors that have negatively affected these ecosystems elsewhere. Re-surveys of 11 locations at the easternmost extent of Tierra del Fuego originally conducted in 1973 showed no significant differences in the densities of adult and juvenile Macrocystis pyrifera kelp or kelp holdfast diameter between the two survey periods. Additionally, sea urchin assemblage structure at the same sites were not significantly different between the two time periods, with the dominant species Loxechinus albus accounting for 66.3% of total sea urchin abundance in 2018 and 61.1% in 1973. Time series of Landsat imagery of the region from 1998 to 2018 showed no long-term trends in kelp canopy over the past 20 years. However, ~ 4-year oscillations in canopy fraction were observed and were strongly and negatively correlated with the NOAA Multivariate ENSO index and sea surface temperature. More extensive surveying in 2018 showed significant differences in benthic community structure between exposed and sheltered locations. Fish species endemic to the Magellanic Province accounted for 73% of all nearshore species observed and from 98-100% of the numerical abundance enumerated at sites. Fish assemblage structure varied significantly among locations and wave exposures. The recent creation of the Yaganes Marine Park is an important step in protecting this unique and biologically rich region; however, the nearshore waters of the region are currently not included in this protection. There is a general lack of information on changes in kelp forests over long time periods, making a global assessment difficult. A complete picture of how these ecosystems are responding to human pressures must also include remote locations and locations with little to no impact.

Marine biodiversity at the end of the world: Cape Horn and Diego Ramírez islands.

The vast and complex coast of the Magellan Region of extreme southern Chile possesses a diversity of habitats including fjords, deep channels, and extensive kelp forests, with a unique mix of temperate and sub-Antarctic species. The Cape Horn and Diego Ramírez archipelagos are the most southerly locations in the Americas, with the southernmost kelp forests, and some of the least explored places on earth. The giant kelp Macrocystis pyrifera plays a key role in structuring the ecological communities of the entire region, with the large brown seaweed Lessonia spp. forming dense understories. Kelp densities were highest around Cape Horn, followed by Diego Ramírez, and lowest within the fjord region of Francisco Coloane Marine Park (mean canopy densities of 2.51 kg m-2, 2.29 kg m-2, and 2.14 kg m-2, respectively). There were clear differences in marine communities among these sub-regions, with the lowest diversity in the fjords. We observed 18 species of nearshore fishes, with average species richness nearly 50% higher at Diego Ramírez compared with Cape Horn and Francisco Coloane. The number of individual fishes was nearly 10 times higher at Diego Ramírez and 4 times higher at Cape Horn compared with the fjords. Dropcam surveys of mesophotic depths (53-105 m) identified 30 taxa from 25 families, 15 classes, and 7 phyla. While much of these deeper habitats consisted of soft sediment and cobble, in rocky habitats, echinoderms, mollusks, bryozoans, and sponges were common. The southern hagfish (Myxine australis) was the most frequently encountered of the deep-sea fishes (50% of deployments), and while the Fueguian sprat (Sprattus fuegensis) was the most abundant fish species, its distribution was patchy. The Cape Horn and Diego Ramírez archipelagos represent some of the last intact sub-Antarctic ecosystems remaining and a recently declared large protected area will help ensure the health of this unique region.