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Model simulations show that a network of marine reserves can enhance yield in depleted fisheries by protecting populations, particularly large, old spawners that supply larvae for interspersed fishing grounds. The ability of marine reserves to enhance sustainable fisheries is much less evident. We report empirical evidence of a marine reserve network improving yield for a sustainable spiny lobster fishery, apparently through the spillover of adult lobsters and behavioral adaptation by the fishing fleet. Results of a Before-After, Control-Impact analysis found catch, effort, and Catch-Per-Unit Effort increased after the establishment of marine reserves in the northern region of the fishery where fishers responded by fishing intensively at reserve borders, but declined in the southern region where they vacated once productive fishing grounds. Our results highlight the value of collaborative research and education programs involving diverse stakeholders for preparing fisheries to operate productively within a seascape that includes a large marine reserve network.

We conducted a population genetic analysis of the stalked kelp, Pterygophora californica, in the Santa Barbara Channel, California, USA. The results were compared with previous work on the genetic differentiation of giant kelp, Macrocystis pyrifera, in the same region. These two sympatric kelps not only share many life history and dispersal characteristics but also differ in that dislodged P. californica does not produce floating rafts with buoyant fertile sporophytes, commonly observed for M. pyrifera. We used a comparative population genetic approach with these two species to test the hypothesis that the ability to produce floating rafts increases the genetic connectivity among kelp patches in the Santa Barbara Channel. We quantified the association of habitat continuity and oceanographic distance with the genetic differentiation observed in stalked kelp, like previously conducted for giant kelp. We compared both overall (across all patches) and pairwise (between patches) genetic differentiation. We found that oceanographic transit time, habitat continuity, and geographic distance were all associated with genetic connectivity in P. californica, supporting similar previous findings for M. pyrifera. Controlling for differences in heterozygosity between kelp species using Jost's DEST , we showed that global differentiation and pairwise differentiation were similar among patches between the two kelp species, indicating that they have similar dispersal capabilities despite their differences in rafting ability. These results suggest that rafting sporophytes do not play a significant role in effective dispersal of M. pyrifera at ecologically relevant spatial and temporal scales.

Herbivores can reach extraordinary abundances in many ecosystems. When herbivore abundance is high, heavy grazing can severely defoliate primary producers and, in some cases, even drive ecosystem to undergo regime shifts from a high productivity state to a denuded, low productivity state. While the phenomenon of herbivore-driven regime shifts is well documented, we only partially understand the mechanisms underlying these events. Here, we combine herbivory experiments with 21 years of long-term monitoring data of kelp forest ecosystems to test the hypothesis that herbivores drive regime shifts when herbivory exceeds primary production. To test this hypothesis, we quantified how the foraging habits of an important group of marine herbivores-sea urchins-change with increases in sea urchin biomass and trigger regime shifts to a foundation species, giant kelp (Macrocystis pyrifera). Using experiments, we quantified how the grazing capacity of urchins increases as urchin biomass increases, then we combined these estimates of urchin grazing capacity with estimates of kelp production to predict when and where urchin grazing capacity exceeded kelp production. When grazing capacity exceeded kelp production, sea urchins caused a 50-fold reduction in giant kelp biomass. Our findings support the hypothesis that the balance between herbivory and production underlies herbivore-driven regime shifts in Southern California kelp forests and provides insight into when and where urchins are likely to force regime shifts in kelp forest ecosystems.

Ecosystems across the United States are changing in complex and surprising ways. Ongoing demand for critical ecosystem services requires an understanding of the populations and communities in these ecosystems in the future. This paper represents a synthesis effort of the U.S. National Science Foundation-funded Long-Term Ecological Research (LTER) network addressing the core research area of "populations and communities." The objective of this effort was to show the importance of long-term data collection and experiments for addressing the hardest questions in scientific ecology that have significant implications for environmental policy and management. Each LTER site developed at least one compelling case study about what their site could look like in 50-100 yr as human and environmental drivers influencing specific ecosystems change. As the case studies were prepared, five themes emerged, and the studies were grouped into papers in this LTER Futures Special Feature addressing state change, connectivity, resilience, time lags, and cascading effects. This paper addresses the "connectivity" theme and has examples from the Phoenix (urban), Niwot Ridge (alpine tundra), McMurdo Dry Valleys (polar desert), Plum Island (coastal), Santa Barbara Coastal (coastal), and Jornada (arid grassland and shrubland) sites. Connectivity has multiple dimensions, ranging from multi-scalar interactions in space to complex interactions over time that govern the transport of materials and the distribution and movement of organisms. The case studies presented here range widely, showing how land-use legacies interact with climate to alter the structure and function of arid ecosystems and flows of resources and organisms in Antarctic polar desert, alpine, urban, and coastal marine ecosystems. Long-term ecological research demonstrates that connectivity can, in some circumstances, sustain valuable ecosystem functions, such as the persistence of foundation species and their associated biodiversity or, it can be an agent of state change, as when it increases wind and water erosion. Increased connectivity due to warming can also lead to species range expansions or contractions and the introduction of undesirable species. Continued long-term studies are essential for addressing the complexities of connectivity. The diversity of ecosystems within the LTER network is a strong platform for these studies. National Science Foundation Long-Term Ecological Research programNational Science Foundation (NSF) [DEB-1637590, 1832016, DEB-0213767, 0816963, 1354494, 1902712, NSF OCE-0423565, 1058747, 1637630, OPP-1637708, OCE-1232779, DEB-1832194, DEB-1637686] Published version This research was supported by the National Science Foundation Long-Term Ecological Research program grants to the Central Arizona-Phoenix (DEB-1637590, 1832016), Plum Island (DEB-0213767, 0816963, 1354494, 1902712, NSF OCE-0423565, 1058747, 1637630), McMurdo (OPP-1637708), Santa Barbara (OCE-1232779), Jornada (DEB-1832194), and Niwot (DEB-1637686) LTER sites.

Disturbances often cause the disproportionate loss of foundation species but understanding how the frequency and severity of disturbance to such organisms influence biological communities remains unresolved. This gap in knowledge exists in part because of the rarity of ecologically meaningful studies capable of disentangling different elements of disturbance. Hence, we carried out a long-term (9 yr), large-scale (2,000 m2 plots), spatially replicated (4 sites) field experiment in which we manipulated disturbance to a globally distributed marine foundation species, the giant kelp Macrocystis pyrifera, and tracked community responses over time. To distinguish the effects of disturbance frequency and severity on the biodiversity and composition of temperate rocky reef communities, we simulated the repeated loss of giant kelp from destructive winter waves across a background of natural variation in disturbance. By following the response of over 200 taxa from the surrounding community, we discovered that the frequency of disturbance to giant kelp changed the biomass, diversity, and composition of community guilds in a manner commensurate with their dependence on the physical (i.e., benthic light and space), trophic (i.e., living and detrital biomass), and habitat (i.e., biogenic structure) resources mediated by this foundation species. Annual winter disturbance to giant kelp reduced living and detrital giant kelp biomass by 57% and 40%, respectively, enhanced bottom light by 22%, and halved the seafloor area covered by giant kelp holdfasts. Concomitantly, the biomass of understory algae and epilithic sessile invertebrates more than doubled, while the biomass of rock-boring clams, mobile invertebrates, and fishes decreased 30-61%. Frequent loss of giant kelp boosted understory algal richness by 82% and lowered sessile invertebrate richness by 13% but did not affect the biodiversity of mobile fauna. In contrast to changes driven by disturbance frequency, interannual variation in the severity of disturbance to giant kelp had weaker, less consistent effects, causing only modest changes in assemblages of sessile invertebrates, mobile invertebrate herbivores, and fishes. Our results broaden the foundation species concept by demonstrating that repeated disturbance to a dominant habitat-forming species can outweigh the influence of less frequent but severe disturbances for the surrounding community.

Distinguishing between human impacts and natural variation in abundance remains difficult because most species exhibit complex patterns of variation in space and time. When ecological monitoring data are available, a before-after-control-impact (BACI) analysis can control natural spatial and temporal variation to better identify an impact and estimate its magnitude. However, populations with limited distributions and confounding spatial-temporal dynamics can violate core assumptions of BACI-type designs. In this study, we assessed how such properties affect the potential to identify impacts. Specifically, we quantified the conditions under which BACI analyses correctly (or incorrectly) identified simulated anthropogenic impacts in a spatially and temporally replicated data set of fish, macroalgal, and invertebrate species found on nearshore subtidal reefs in southern California, USA. We found BACI failed to assess very localized impacts, and had low power but high precision when assessing region-wide impacts. Power was highest for severe impacts of moderate spatial scale, and impacts were most easily detected in species with stable, widely distributed populations. Serial autocorrelation in the data greatly inflated false impact detection rates, and could be partly controlled for statistically, while spatial synchrony in dynamics had no consistent effect on power or false detection rates. Unfortunately, species that offer high power to detect real impacts were also more likely to detect impacts where none had occurred. However, considering power and false detection rates together can identify promising indicator species, and collectively analyzing data for similar species improved the net ability to assess impacts. These insights set expectations for the sizes and severities of impacts that BACI analyses can detect in real systems, point to the importance of serial autocorrelation (but not of spatial synchrony), and indicate how to choose the species, and groups of species, that can best identify impacts.

Disturbances often disproportionately impact different vegetation layers in forests and other vertically stratified ecosystems, shaping community structure and ecosystem function. However, disturbance‐driven changes may be mediated by environmental conditions that affect habitat quality and species interactions. In a decade‐long field experiment, we tested how kelp forest net primary productivity (NPP) responds to repeated canopy loss along a gradient in grazing and substrate suitability. We discovered that habitat quality can mediate the effects of intensified disturbance on canopy and understory NPP. Experimental annual and quarterly disturbances suppressed total macroalgal NPP, but effects were strongest in high‐quality habitats that supported dense kelp canopies that were removed by disturbance. Understory macroalgae partly compensated for canopy NPP losses and this effect magnified with increasing habitat quality. Disturbance‐driven increases in understory NPP were still rising after 5–10 years of disturbance, demonstrating the value of long‐term experimentation for understanding ecosystem responses to changing disturbance regimes., In a decade‐long field experiment, we show that habitat quality can mediate the effects of intensified disturbance on canopy and understory net primary productivity (NPP) in kelp forests. Experimental annual and quarterly disturbances suppressed total macroalgal NPP; however, understory macroalgae partly compensated for canopy NPP losses and this effect magnified with increasing habitat quality. Disturbance‐driven increases in understory NPP were still rising after 5–10 years of disturbance, demonstrating the value of long‐term experimentation for understanding ecosystem responses to changing disturbance regimes.

Marine Protected Areas (MPAs) are designed to enhance biodiversity and ecosystem services. Some MPAs are also established to benefit fisheries through increased egg and larval production, or the spillover of mobile juveniles and adults. Whether spillover influences fishery landings depend on the population status and movement patterns of target species both inside and outside of MPAs, as well as the status of the fishery and behavior of the fleet. We tested whether an increase in the lobster population inside two newly established MPAs influenced local catch, fishing effort, and catch-per-unit-effort (CPUE) within the sustainable California spiny lobster fishery. We found greater build-up of lobsters within MPAs relative to unprotected areas, and greater increases in fishing effort and total lobster catch, but not CPUE, in fishing zones containing MPAs vs. those without MPAs. Our results show that a 35% reduction in fishing area resulting from MPA designation was compensated for by a 225% increase in total catch after 6-years, thus indicating at a local scale that the trade-off of fishing ground for no-fishing zones benefitted the fishery.

Foundation species structure communities, promote biodiversity, and stabilize ecosystem processes by creating locally stable environmental conditions. Despite their critical importance, the role of foundation species in stabilizing natural communities has seldom been quantified. In theory, the stability of a foundation species should promote community stability by enhancing species richness, altering the population fluctuations of individual species, or both. Here we tested the hypothesis that the stability of a marine foundation species, the giant kelp Macrocystis pyrifera, increased the stability of the aggregate biomass of a phylogenetically diverse assemblage of understory algae and sessile invertebrates that compete for space beneath the giant kelp canopy. To achieve this goal, we analyzed an 18-yr time series of the biomass of giant kelp and its associated benthic community collected from 32 plots distributed among nine shallow reefs in the Santa Barbara Channel, USA. We showed that the stability of understory algae and sessile invertebrates was positively and indirectly related to the stability of giant kelp, which primarily resulted from giant kelp's direct positive association with species richness. The stability of all community types was positively related to species richness via increased species stability and species asynchrony. The stabilizing effects of richness were three to four times stronger when algae and invertebrates were considered separately rather than in combination. Our finding that diversity-stability relationships were stronger in communities consisting of species with similar resource requirements suggests that competition for shared resources rather than differential responses to environmental conditions played a more important role in stabilizing the community. Increasing threats to structure-forming foundation species worldwide necessitates a detailed understanding of how they influence their associated community. This study is among the first to show that dampened temporal fluctuations in the biomass of a foundation species is an important determinant of the stability of the complex communities it supports.

© 2018 The US and Mexico share a common history in many areas, including language and culture. They face ecological changes due to the increased frequency and severity of droughts and rising energy demands; trends that entail economic costs for both nations and major implications for human wellbeing. We describe an ongoing effort by the Environment Working Group (EWG), created by The University of California's UC-Mexico initiative in 2015, to promote binational research, teaching, and outreach collaborations on the implications of climate change for Mexico and California. We synthesize current knowledge about the most pressing issues related to climate change in the US-Mexico border region and provide examples of cross-border discoveries and research initiatives, highlighting the need to move forward in six broad rubrics. This and similar binational cooperation efforts can lead to improved living standards, generate a collaborative mindset among participating universities, and create an international network to address urgent sustainability challenges affecting both countries.

Photoautotrophs vary the concentration of photosynthetic pigments in response to changing environmental conditions. In the ocean, the chlorophyll a to carbon ratio (Chl:C) has been used as a proxy for the physiological condition of phytoplankton, and there is laboratory evidence that the growth rate of juvenile giant kelp (Macrocystis pyrifera), a coastal foundation species, is positively related to Chl:C under nutrient-limited conditions. We examined the relative roles of nutrients and light in determining Chl:C dynamics and if fluctuations in Chl:C were related to changes in canopy biomass and net primary production (NPP) from sites spanning 750 km of the California coast. Seventy percent of the variability in canopy Chl:C was explained by a combination of photosynthetically active radiation and seawater nitrate concentration. In the periodically nutrient-limited waters of the Southern California Bight, changes in Chl:C positively resembled changes in available nitrate, whereas Chl:C negatively tracked changes in light for the more nutrient-replete central California coastline. Values of Chl:C were positively related to residuals from an autoregressive model of kelp biomass at the southern California sites indicating that Chl:C is a proxy for physiological state of the kelp canopy. NPP estimated through correlations with kelp biomass and lagged Chl:C compared well to established field-based estimates. These results open the possibility of assessing giant kelp physiological condition from estimates of Chl:C modeled from sea surface irradiance and nitrate concentration, which in turn can be used to estimate giant kelp primary production over large spatial and temporal scales using future remote sensing technologies.

Seasonal or chronic nutrient limitations in the photic zone limit large-scale cultivation of seaweed (macroalgae) in much of the world's oceans, hindering the development of macroalgae as a biofuel feedstock. One possible solution is to supply nutrients using a diel depth-cycling approach, physically moving the macroalgae between deep nutrient-rich water at night and shallow depths within the photic zone during the day. This study tested the effects of depth-cycling on the growth, morphology, and chemical composition of the giant kelp Macrocystis pyrifera, a target species for renewable biomass production. Giant kelp grown under depth-cycling conditions had an average growth rate of 5% per day and produced four times more biomass (wet weight) than individuals grown in a kelp bed without depth-cycling. Analysis of tissue from the depth-cycled kelp showed elevated levels of protein, lower C:N ratios, and distinct δ15N and δ13C values suggesting that the depth-cycled kelp were not nitrogen-deficient and assimilated nutrients from deep water. Depth-cycled kelp also exhibited smaller and thicker-walled pneumatocysts and larger blades. Overall, this study supports further investigation of depth-cycling as a macroalgal farming strategy.

Because natural ecosystems are complex, it is difficult to predict how their variability scales across space and levels of organization. The species-insurance hypothesis predicts that asynchronous dynamics among species should reduce variability when biomass is aggregated either from local species populations to local multispecies communities, or from metapopulations to metacommunities. Similarly, the spatial-insurance hypothesis predicts that asynchronous spatial dynamics among either local populations or local communities should stabilize metapopulation biomass and metacommunity biomass, respectively. In combination, both species and spatial insurance reduce variation in metacommunity biomass over time, yet these insurances are rarely considered together in natural systems. We partitioned the extent that species insurance and spatial insurance reduced the annual variation in macroalgal biomass in a southern California kelp forest. We quantified variability and synchrony at two levels of organization (population and community) and two spatial scales (local plots and region) and quantified the strength of species and spatial insurance by comparing observed variability and synchrony in aggregate biomass to null models of independent species or spatial dynamics based on cyclic-shift permutation. Spatial insurance was weak, presumably because large-scale oceanographic processes in the study region led to high spatial synchrony at both population- and community-level biomass. Species insurance was stronger due to asynchronous dynamics among the metapopulations of a few common species. In particular, a regional decline in the dominant understory kelp species Pterygophora californica was compensated for by the rise of three subdominant species. These compensatory dynamics were associated with positive values of the Pacific Decadal Oscillation, indicating that differential species tolerances to warmer temperature and nutrient-poor conditions may underlie species insurance in this system. Our results illustrate how species insurance can stabilize aggregate community properties in natural ecosystems where environmental conditions vary over broad spatial scales.

Globally, anthropogenic pressures are reducing the abundances of marine species and altering ecosystems through modification of trophic interactions. Yet, consumer declines also disrupt important bottom-up processes, like nutrient recycling, which are critical for ecosystem functioning. Consumer-mediated nutrient dynamics (CND) is now considered a major biogeochemical component of most ecosystems, but lacking long-term studies, it is difficult to predict how CND will respond to accelerating disturbances in the wake of global change. To aid such predictions, we coupled empirical ammonium excretion rates with an 18-year time series of the standing biomass of common benthic macroinvertebrates in southern California kelp forests. This time series of excretion rates encompassed an extended period of extreme ocean warming, disease outbreaks, and the abolishment of fishing at two of our study sites, allowing us to assess kelp forest CND across a wide range of environmental conditions. At their peak, reef invertebrates supplied an average of 18.3 ± 3.0 µmol NH4+ m-2 hr-1 to kelp forests when sea stars were regionally abundant, but dropped to 3.5 ± 1.0 µmol NH4+ m-2 hr-1 following their mass mortality due to disease during a prolonged period of extreme warming. However, a coincident increase in the abundance of the California spiny lobster, Palinurus interupptus (Randall, 1840), likely in response to both reduced fishing and a warmer ocean, compensated for much of the recycled ammonium lost to sea star mortality. Both lobsters and sea stars are widely recognized as key predators that can profoundly influence community structure in benthic marine systems. Our study is the first to demonstrate their importance in nutrient cycling, thus expanding their roles in the ecosystem. Climate change is increasing the frequency and severity of warming events, and rising human populations are intensifying fishing pressure in coastal ecosystems worldwide. Our study documents how these projected global changes can drive regime shifts in CND and fundamentally alter a critical ecosystem function.

Enhanced release of alkalinity from the seafloor, principally driven by anaerobic degradation of organic matter under low-oxygen conditions and associated secondary redox reactions, can increase the carbon dioxide (CO2) buffering capacity of seawater and therefore oceanic CO2 uptake. The Baltic Sea has undergone severe changes in oxygenation state and total alkalinity (TA) over the past decades. The link between these concurrent changes has not yet been investigated in detail. A recent system-wide TA budget constructed for the past 50 years using BALTSEM, a coupled physical-biogeochemical model for the whole Baltic Sea area, revealed an unknown TA source. Here we use BALTSEM in combination with observational data and one-dimensional reactive transport modelling of sedimentary processes in the Fårö Deep, a deep Baltic Sea basin, to test whether sulfate reduction coupled to iron (Fe) sulfide burial can explain the missing TA source in the Baltic Proper. We calculated that this burial can account for 26 % of the missing source in this basin, with the remaining TA possibly originating from unknown river inputs or submarine groundwater discharge. We also show that temporal variability in the input of Fe to the sediments since the 1970s drives changes in sulfur burial in the Fårö Deep, suggesting that Fe availability is the ultimate limiting factor for TA generation under anoxic conditions. The implementation of projected climate change and two nutrient load scenarios for the 21st century in BALTSEM shows that reducing nutrient loads will improve deep water oxygen conditions, but at the expense of lower surface water TA concentrations, CO2 buffering capacities and faster acidification. When these changes additionally lead to a decrease in Fe inputs to the sediment of the deep basins, anaerobic TA generation will be reduced even further, thus exacerbating acidification. This work highlights that Fe dynamics play a key role in the release of TA from sediments where Fe sulfide formation is limited by Fe availability, as exemplified for the Baltic Sea. Moreover, it demonstrates that burial of Fe sulfides should be included in TA budgets of low oxygen basins.

Data from five sites of the International Long Term Ecological Research (ILTER) network in the North-Eastern Pacific, Western Arctic Ocean, Northern Baltic Sea, South-Eastern North Sea and in the Western Mediterranean Sea were analyzed by dynamic factor analysis (DFA) to trace common multi-year trends in abundance and composition of phytoplankton, benthic fauna and temperate reef fish. Multiannual trends were related to climate and environmental variables to study interactions. Two common trends in biological responses were detected, with temperature and climate indices as explanatory variables in four of the five LTER sites considered. Only one trend was observed at the fifth site, the Northern Baltic Sea, where no explanatory variables were identified. Our findings revealed quasi-synchronous biological shifts in the different marine ecosystems coincident with the 2000 climatic regime shift and provided evidence on a possible further biological shift around 2010. The observed biological modifications were coupled with abrupt or continuous increase in sea water and air temperature confirming the key-role of temperature in structuring marine communities.

Recent decades have witnessed an exponential spread of low-oxygen regions in the coastal ocean due at least in-part to enhanced terrestrial nutrient inputs. As oxygen deprivation is a major stressor on marine ecosystems, there is a great need to quantitatively link shifts in nutrient loading with changes in oxygen concentrations. To this end, we have developed and here describe, evaluate, and apply the Coastal Ocean Oxygen Linked to Benthic Exchange And Nutrient Supply (COOLBEANS) model, a first-of-its-kind, spatially explicit (with 152 coastal segments) model, global model of coastal oxygen and nutrient dynamics. In COOLBEANS, benthic oxygen demand (BOD) is calculated using empirical models for aerobic respiration, iron reduction, and sulfate reduction, while oxygen supply is represented by a simple parameterization of exchange between surface and bottom waters. A nutrient cycling component translates shifts in riverine nutrient inputs into changes in organic matter delivery to sediments and, ultimately, oxygen uptake. Modeled BOD reproduces observations reasonably well (Nash-Sutcliffe efficiency = 0.71), and estimates of exchange between surface and bottom waters correlate with stratification. The model examines sensitivity of bottom water oxygen to changes in nutrient inputs and vertical exchange between surface and bottom waters, highlighting the importance of this vertical exchange in defining the susceptibility of a system to oxygen depletion. These sensitivities along with estimated maximum hypoxic areas that are supported by present day nutrient loads are consistent with existing hypoxic regions. Sensitivities are put into context by applying historic changes in nitrogen loading observed in the Gulf of Mexico to the global coastal ocean, demonstrating that such loads would drive many systems anoxic or even sulfidic.

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.

Sea urchins are voracious herbivores that influence the ecological structure and function of nearshore ecosystems throughout the world. Like many species that produce planktonic larvae, their recruitment is thought to be particularly sensitive to climatic fluctuations in temperature that directly or indirectly affect adult reproduction and larval transport and survival. Yet how climate alters sea urchin populations in space and time by modifying larval recruitment and year-class strength on the time-scales that regulate populations remains understudied. Using a, spatially replicated weekly-biweekly dataset spanning 27 years and 1100 km of coastline, we characterized seasonal, interannual, and spatial patterns of larval settlement of the purple sea urchin (Strongylocentrotus purpuratus). We show that large spatial differences in temporal patterns of larval settlement were associated with different responses to fluctuations in ocean temperature and climate. Importantly, we found a strong correlation between larval settlement and regional year class strength suggesting that such temporal and spatial variation in settlement plays an important role in controlling population dynamics. These results provide strong evidence over extensive temporal and spatial domains that climatic fluctuations shape broad-scale patterns of larval settlement and subsequent population structure of an important marine herbivore known to control the productivity, community state and provisioning services of marine ecosystems.

Motivation The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. Main types of variables included The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. Spatial location and grain BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). Time period and grain BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. Major taxa and level of measurement BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. Software format .csv and .SQL., Global Ecology and Biogeography, 27 (7), ISSN:1466-822X, ISSN:1466-8238

Foundation species define the ecosystems they live in, but ecologists have often characterized dominant plants as foundational without supporting evidence. Giant kelp has long been considered a marine foundation species due to its complex structure and high productivity; however, there is little quantitative evidence to evaluate this. Here, we apply structural equation modelling to a 15-year time series of reef community data to evaluate how giant kelp affects the reef community. Although species richness was positively associated with giant kelp biomass, most direct paths did not involve giant kelp. Instead, the foundational qualities of giant kelp were driven mostly by indirect effects attributed to its dominant physical structure and associated engineering influence on the ecosystem, rather than by its use as food by invertebrates and fishes. Giant kelp structure has indirect effects because it shades out understorey algae that compete with sessile invertebrates. When released from competition, sessile species in turn increase the diversity of mobile predators. Sea urchin grazing effects could have been misinterpreted as kelp effects, because sea urchins can overgraze giant kelp, understorey algae and sessile invertebrates alike. Our results confirm the high diversity and biomass associated with kelp forests, but highlight how species interactions and habitat attributes can be misconstrued as direct consequences of a foundation species like giant kelp.

We investigated the patterns and controls of dissolved organic carbon (DOC) production by the giant kelp (Macrocystis pyrifera) using data from short-term in situ incubations of entire blades and portions of stipes. These data were incorporated into an empirical model of reef-scale net primary production (NPP) at Mohawk Reef in southern California, U.S.A. for an 8-yr period. Rates of DOC release of incubated blades varied unpredictably with time of year, but were significantly related to the irradiance at the sea surface during the incubations. The growth stage, C/N ratio, and epiphyte load of the blades and the temperature of the ocean during the incubations had no discernable effect on rates of DOC release. Blades produced on average 2–3 times more DOC than stipes, and stipes and blades produced on average 30% and 80% more DOC respectively during the day compared to the night. Modeled DOC NPP at the reef scale was on average highest in summer and spring (∼0.5 g C m−2 d−1) and lowest in winter and autumn (∼0.31 g C m−2 d−1), but it varied greatly among years for any given season as large oscillations in standing biomass led to corresponding fluctuations in reef-scale DOC NPP. The fraction of NPP released as DOC was highly variable when examined at the monthly time scale, but became much more stable at seasonal and annual time scales averaging 14% of total NPP.

Marine reserves have become increasingly valuable tools with which to manage eco- systems. These reserves consistently restore populations of top predators, often reducing avail- ability of their favored prey. We hypothesized that such prey reduction in reserves causes pro- tected predators to alter their attack behavior to include less palatable prey, potentially amplifying top-down effects on community structure. To test this hypothesis, we presented the relatively unpalatable sea hare Aplysia californica to freely foraging spiny lobsters Panulirus interruptus in 4 marine no-take reserves, each paired with an adjacent fished area. We found that lobsters only attacked sea hares inside reserves, where lobster density was significantly greater than that of the adjacent fished areas. Attacks on otherwise unpalatable prey exclusively in no-take reserves was likely caused by increased hunger, since in the laboratory only food-deprived lobsters attacked sea hares. These findings are the first to suggest that management involving no-take reserves may have unintended consequences on community structure that result from behavioral changes in key predators in the face of increased competition for food. We suspect that these effects may become more widely detected as reserves across the globe grow older and are researched further.

Global climate change is likely to constrain low latitude range edges across many taxa and habitats. Such is the case for NE Atlantic marine macroalgal forests, important ecosystems whose main structuring species is the annual kelp Saccorhiza polyschides. We coupled ecological niche modelling with simulations of potential dispersal and delayed development stages to infer the major forces shaping range edges and to predict their dynamics. Models indicated that the southern limit is set by high winter temperatures above the physiological tolerance of overwintering microscopic stages and reduced upwelling during recruitment. The best range predictions were achieved assuming low spatial dispersal (5 km) and delayed stages up to two years (temporal dispersal). Reconstructing distributions through time indicated losses of similar to 30% from 1986 to 2014, restricting S. polyschides to upwelling regions at the southern edge. Future predictions further restrict populations to a unique refugium in northwestern Iberia. Losses were dependent on the emissions scenario, with the most drastic one shifting similar to 38% of the current distribution by 2100. Such distributional changes might not be rescued by dispersal in space or time (as shown for the recent past) and are expected to drive major biodiversity loss and changes in ecosystem functioning. Electricity of Portugal (Fundo EDP para a Biodiversidade); FCT - Portuguese Science Foundation [PTDC/MAR-EST/6053/2014, EXTANT-EXCL/AAG-GLO/0661/2012, SFRH/BPD/111003/2015]

Background Molecular markers are revealing a much more diverse and evolutionarily complex picture of marine biodiversity than previously anticipated. Cryptic and/or endemic marine species are continually being found throughout the world oceans, predominantly in inconspicuous tropical groups but also in larger, canopy-forming taxa from well studied temperate regions. Interspecific hybridization has also been found to be prevalent in many marine groups, for instance within dense congeneric assemblages, with introgressive gene-flow being the most common outcome. Here, using a congeneric phylogeographic approach, we investigated two monotypic and geographically complementary sister genera of north-east Pacific intertidal seaweeds (Hesperophycus and Pelvetiopsis), for which preliminary molecular tests revealed unexpected conflicts consistent with unrecognized cryptic diversity and hybridization. Results The three recovered mtDNA clades did not match a priori species delimitations. H. californicus was congruent, whereas widespread P. limitata encompassed two additional narrow-endemic species from California - P. arborescens (here genetically confirmed) and P. hybrida sp. nov. The congruence between the genotypic clusters and the mtDNA clades was absolute. Fixed heterozygosity was apparent in a high proportion of loci in P. limitata and P. hybrida, with genetic analyses showing that the latter was composed of both H. californicus and P. arborescens genomes. All four inferred species could be distinguished based on their general morphology. Conclusions This study confirmed additional diversity and reticulation within NE Pacific Hesperophycus/Pelvetiopsis, including the validity of the much endangered, modern climatic relict P. arborescens, and the identification of a new, stable allopolyploid species (P. hybrida) with clearly discernable ancestry (♀ H. californicus x ♂ P. arborescens), morphology, and geographical distribution. Allopolyploid speciation is otherwise completely unknown in brown seaweeds, and its unique occurrence within this genus (P. limitata possibly representing a second example) remains enigmatic. The taxonomic separation of Hesperophycus and Pelvetiopsis is not supported and the genera should be synonymized; we retain only the latter. The transitional coastline between Point Conception and Monterey Bay represented a diversity hotspot for the genus and the likely sites of extraordinary evolutionary events of allopolyploid speciation at sympatric range contact zones. This study pinpoints how much diversity (and evolutionary processes) potentially remains undiscovered even on a conspicuous seaweed genus from the well-studied Californian intertidal shores let alone in other, less studied marine groups and regions/depths. info:eu-repo/semantics/publishedVersion