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Changes in body size have been documented across taxa in response to human activities and climate change. Body size influences many aspects of an individual's physiology, behavior, and ecology, ultimately affecting life history performance and resilience to stressors. In this study, we developed an analytical approach to model individual growth patterns using aerial imagery collected via drones, which can be used to investigate shifts in body size in a population and the associated drivers. We applied the method to a large morphological dataset of gray whales (Eschrichtius robustus) using a distinct foraging ground along the NE Pacific coast, and found that the asymptotic length of these whales has declined since around the year 2000 at an average rate of 0.05-0.12 m/y. The decline has been stronger in females, which are estimated to be now comparable in size to males, minimizing sexual dimorphism. We show that the decline in asymptotic length is correlated with two oceanographic metrics acting as proxies of habitat quality at different scales: the mean Pacific Decadal Oscillation index, and the mean ratio between upwelling intensity in a season and the number of relaxation events. These results suggest that the decline in gray whale body size may represent a plastic response to changing environmental conditions. Decreasing body size could have cascading effects on the population's demography, ability to adjust to environmental changes, and ecological influence on the structure of their community. This finding adds to the mounting evidence that body size is shrinking in several marine populations in association with climate change and other anthropogenic stressors. Our modeling approach is broadly applicable across multiple systems where morphological data on megafauna are collected using drones.

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Environmental changes such as seasonality, decadal oscillation, and anthropogenic forcing may shape the dynamics of lower trophic-level organisms. In this study, 9-years (2010-2018) of monitoring data on microscopic protists such as diatoms and dinoflagellates, and environmental variables were analyzed to clarify the relationships between plankton and local/synoptic environmental changes. We found that time-series temperature increased in May, whereas it decreased in August and November. Nutrients (e.g., phosphate) decreased in May, remained unchanged in August, and increased in November from 2010 to 2018. The partial pressure of CO2 increased in May, August, and November over time. It is notable that the change in seawater temperature (-0.54 to 0.32 °C per year) and CO2 levels (3.6-5.7 µatm CO2 per year) in the latest decade in the eastern Tsugaru Strait were highly dynamic than the projected anthropogenic climate change. Protist abundance generally increased or stayed unchanged during the examined period. In August and November, when cooling and decreases in pH occurred, diatoms such as Chaetoceros subgenus Hyalochaete spp. and Rhizosoleniaceae temporally increased from 2010 to 2018. During the study period, we found that locally aquacultured scallops elevated soft tissue mass relative to the total weight as diatom abundance increased, and the relative scallop soft tissue mass was positively related to the Pacific Decadal Oscillation index. These results indicate that decadal climatic forcing in the ocean modifies the local physical and chemical environment, which strongly affects phytoplankton dynamics rather than the effect of anthropogenic climate change in the eastern Tsugaru Strait.

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The North Yellow Sea (NYS) is a productive marginal sea of the western North Pacific. In summer and autumn, CaCO3 saturation states beneath the seasonal thermocline in the NYS have frequently fallen below critical levels, indicating that marine calcifying organisms are under threat. To explore the long-term evolution of the acidification of the NYS, we reconstructed seasonal variations in subsurface aragonite saturation state (Ωarag) and pH during 1976-2017, using wintertime and summertime temperature, salinity, dissolved oxygen and pH data mainly from a quality-controlled oceanographic database. Over the past 40 years, the wintertime warming rate in the NYS was twice the rate of global ocean surface warming. Warming-induced decrease in CO2 solubility canceled out a part of the wintertime Ωarag decrease caused by atmospheric CO2 increase, and also had minor effect on pH changes in winter. Although the NYS is a semi-enclosed marginal sea, its interannual variations of wintertime temperature, salinity, pH and Ωarag were correlated to Pacific Decadal Oscillation with a lag of 2-3 years. Due to the eutrophication-induced enhancement of net community respiration beneath the seasonal thermocline, long-term declines of bottom-water Ωarag and pH in summer were substantially faster than the declines of assumed air-equilibrated Ωarag and pH in spring. Over the past 40 years, the amplitudes of seasonal variations of bottom-water Ωarag and pH from spring to summer/autumn have increased by 4-7 times. This amplification has pushed the NYS towards the critical threshold of net community CaCO3 dissolution at a pace faster than that forecast under scenarios of atmospheric CO2 increase. In summary, our results provide insights into the combined effects of ocean warming, eutrophication, atmospheric CO2 rise and climate variability on coastal hydrochemistry, explaining how the environmental stresses on local marine calcifying organisms and the benthic ecosystem increased over the past 40 years.

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'The Blob', a mass of anomalously warm water in the Northeast Pacific Ocean peaking from 2014 to 2016, caused a decrease in primary productivity with cascading effects on the marine ecosystem. Among the more obvious manifestations of the event were seabird breeding failures and mass mortality events. Here, we used corticosterone in breast feathers (fCort), grown in the winter period during migration, as an indicator of nutritional stress to investigate the impact of the Blob on two sentinel Pacific auk species (family Alcidae). Feathers were collected from breeding females over 8 years from 2010 to 2017, encompassing the Blob period. Since Pacific auks replace body feathers at sea during the migratory period, measures of fCort provide an accumulated measure of nutritional stress or allostatic load during this time. Changes in diet were also measured using δ15N and δ13C values from feathers. Relative to years prior to the Blob, the primarily zooplanktivorous Cassin's auklets (Ptychoramphus aleuticus) had elevated fCort in 2014-2017, which correlated with the occurrence of the Blob and a recovery period afterwards, with relatively stable feather isotope values. In contrast, generalist rhinoceros auklets (Cerorhinca monocerata) displayed stable fCort values across years and increased δ15N values during the Blob. As marine heatwaves increase in intensity and frequency due to climate change, this study provides insight into the variable response of Pacific auks to such phenomena and suggests a means for monitoring population-level responses to climatological variation.

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Dendroclimatic reconstructions, which are a well-known tool for extending records of climatic variability, have recently been expanded by using wood anatomical parameters. However, the relationships between wood cellular structures and large-scale climatic patterns, such as El Niño-Southern Oscillation (ENSO) and the Pacific Decadal Oscillation (PDO), are still not completely understood, hindering the potential for wood anatomy as a paleoclimatic proxy. To better understand the teleconnection between regional and local climate processes in the western United States, our main objective was to assess the value of these emerging tree-ring parameters for reconstructing climate dynamics. Using Confocal Laser Scanning Microscopy, we measured cell lumen diameter and cell wall thickness (CWT) for the period 1966 to 2015 in five Douglas-firs [Pseudotsuga menziesii (Mirb.) Franco] from two sites in eastern Arizona (United States). Dendroclimatic analysis was performed using chronologies developed for 10 equally distributed sectors of the ring and daily climatic records to identify the strongest climatic signal for each sector. We found that lumen diameter in the first ring sector was sensitive to previous fall-winter temperature (September 25th to January 23rd), while a precipitation signal (October 27th to February 13th) persisted for the entire first half of the ring. The lack of synchronous patterns between trees for CWT prevented conducting meaningful climate-response analysis for that anatomical parameter. Time series of lumen diameter showed an anti-phase relationship with the Southern Oscillation Index (a proxy for ENSO) at 10 to 14year periodicity and particularly in 1980-2005, suggesting that chronologies of wood anatomical parameters respond to multidecadal variability of regional climatic modes. Our findings demonstrate the potential of cell structural characteristics of southwestern United States conifers for reconstructing past climatic variability, while also improving our understanding of how large-scale ocean-atmosphere interactions impact local hydroclimatic patterns.

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Climate change is likely to change the relationships between commonly used climate indices and underlying patterns of climate variability, but this complexity is rarely considered in studies using climate indices. Here, we show that the physical and ecological conditions mapping onto the Pacific Decadal Oscillation (PDO) index and North Pacific Gyre Oscillation (NPGO) index have changed over multidecadal timescales. These changes apparently began around a 1988/1989 North Pacific climate shift that was marked by abrupt northeast Pacific warming, declining temporal variance in the Aleutian Low (a leading atmospheric driver of the PDO), and increasing correlation between the PDO and NPGO patterns. Sea level pressure and surface temperature patterns associated with each climate index changed after 1988/1989, indicating that identical index values reflect different states of basin-scale climate over time. The PDO and NPGO also show time-dependent skill as indices of regional northeast Pacific ecosystem variability. Since the late 1980s, both indices have become less relevant to physical-ecological variability in regional ecosystems from the Bering Sea to the southern California Current. Users of these climate indices should be aware of nonstationary relationships with underlying climate variability within the historical record, and the potential for further nonstationarity with ongoing climate change.

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Rangeland ecosystems worldwide are characterized by a high degree of uncertainty in precipitation, both within and across years. Such uncertainty creates challenges for livestock managers seeking to match herbivore numbers with forage availability to prevent vegetation degradation and optimize livestock production. Here, we assess variation in annual large herbivore production (LHP, kg/ha) across multiple herbivore densities over a 78-yr period (1940-2018) in a semiarid rangeland ecosystem (shortgrass steppe of eastern Colorado, USA) that has experienced several phase changes in global-level sea surface temperature (SST) anomalies, as measured by the Pacific Decadal Oscillation (PDO) and the El Niño-Southern Oscillation (ENSO). We examined the influence of prevailing PDO phase, magnitude of late winter (February-April) ENSO, prior growing-season precipitation (prior April to prior September) and precipitation during the six months (prior October to current April) preceding the growing season on LHP. All of these are known prior to the start of the growing season in the shortgrass steppe and could potentially be used by livestock managers to adjust herbivore densities. Annual LHP was greater during warm PDO irrespective of herbivore density, while variance in LHP increased by 69% (moderate density) and 91% (high density) under cold-phase compared to warm-phase PDO. No differences in LHP attributed to PDO phase were observed with low herbivore density. ENSO effects on LHP, specifically La Niña, were more pronounced during cold-phase PDO years. High herbivore density increased LHP at a greater rate than at moderate and low densities with increasing fall and winter precipitation. Differential gain, a weighted measure of LHP under higher relative to lower herbivore densities, was sensitive to prevailing PDO phase, ENSO magnitude, and precipitation amounts from the prior growing season and current fall-winter season. Temporal hierarchical approaches using PDO, ENSO, and local-scale precipitation can enhance decision-making for flexible herbivore densities. Herbivore densities could be increased above recommended levels with lowered risk of negative returns for managers during warm-phase PDO to result in greater LHP and less variability. Conversely, during cold-phase PDO, managers should be cognizant of the additional influences of ENSO and prior fall-winter precipitation, which can help predict when to reduce herbivore densities and minimize risk of forage shortages.

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Tree growth at northern treelines is generally temperature-limited due to cold and short growing seasons. However, temperature-induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree-ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell-wall thickness, cell number) and TRW were correlated with the drought-sensitive standardized precipitation-evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925-1946), cool/wet (1947-1976) and again warm/dry (1977-1998) climate regimes. Xylem anatomical traits revealed water-limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture-driven shift in growth-limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture-driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity.

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The supply of nutrients is a fundamental regulator of ocean productivity and carbon sequestration. Nutrient sources, sinks, residence times, and elemental ratios vary over broad scales, including those resulting from climate-driven changes in upper water column stratification, advection, and the deposition of atmospheric dust. These changes can alter the proximate elemental control of ecosystem productivity with cascading ecological effects and impacts on carbon sequestration. Here, we report multidecadal observations revealing that the ecosystem in the eastern region of the North Pacific Subtropical Gyre (NPSG) oscillates on subdecadal scales between inorganic phosphorus (P i ) sufficiency and limitation, when P i concentration in surface waters decreases below 50-60 nmol⋅kg-1 In situ observations and model simulations suggest that sea-level pressure changes over the northwest Pacific may induce basin-scale variations in the atmospheric transport and deposition of Asian dust-associated iron (Fe), causing the eastern portion of the NPSG ecosystem to shift between states of Fe and P i limitation. Our results highlight the critical need to include both atmospheric and ocean circulation variability when modeling the response of open ocean pelagic ecosystems under future climate change scenarios.

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Common approaches for summarizing multivariate environmental or community data assume that relationships among variables are stationary over time, and this assumption is often not tested. Here we test the hypothesis that relationships among environmental and community time series are nonstationary in the Gulf of Alaska ecosystem (North Pacific Ocean) over multidecadal time scales. Dynamic factor analysis (DFA) is applied to environmental and community data from before and after 1988/1989, corresponding to the timing of an abrupt decline in temporal variance of the Aleutian Low atmospheric pattern, a leading driver of Gulf of Alaska climate. Results show that covariance among local atmosphere and ocean environmental variables weakened simultaneous to the decline in Aleutian Low variance. At the same time, community-wide responses of 14 fish and crustacean populations to physical forcing weakened, as indicated by nonstationary environment-biology regression coefficients. In line with theoretical predictions, this loss of a shared response to environmental variability was accompanied by weakening community covariance. Individual populations also showed nonstationary relationships with shared trends of community variability. We conclude that assumptions of fixed environmental and community relationships are likely to produce mistaken inference in this ecosystem. Similar concerns may apply in other ecosystems subject to changing climate patterns.

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Data from decadal hindcast experiments conducted under CMIP5 were used to assess the ability of CM2.1, HadCM3, MIROC5, and CCSM4 Earth System Models (ESMs) to hindcast sea-surface temperature (SST) indices of the Pacific Decadal Oscillation (PDO), the tropical Atlantic SST gradient (TAG) variability, and the West Pacific Warm Pool (WPWP) SST variability from 1961 to 2010. The ESMs were initialized at specific times with observed data to make 10- and 30-year hindcasts/forecasts. Deterministic and probabilistic skill estimates show predictability of detrended WPWP index to 5 years' lead time and of non-detrended WPWP index to 10 years' lead time. These estimates also show atypical skill dependence of PDO and TAG indices on lead times, with increasing skill in the middle to end of 10-year hindcasts. The skill of ESMs to hindcast an observed DCV index (signal skill) is surprisingly greater than the skill to hindcast their own DCV index (noise skill) at some lead times. All ESMs hindcast occurrence frequencies of positive and negative phases of the indices, and probabilities of same-phase transitions from one year to the next reasonably well. Four, major, low-latitude volcanic eruptions are associated with phase transitions of all observed and some of the ensemble-average hindcast indices. All ESMs' WPWP index hindcasts respond correctly to all eruptions as do three observed PDO phase transitions. No one of the ESMs' hindcasts of the TAG index responds correctly to these eruptions. Some of the ESMs hindcast correct phase transitions in the absence of eruptions also, implying that initializations with observed data are beneficial in predicting phase transitions. The skills of DCV indices' phase prediction up to at least two years in advance can be used to inform societal impacts adaptation decisions in water resources management and agriculture. The Atlantic region's responses in these ESMs appear to be fundamentally incorrect.

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Anomalously cold winters with extreme storms strain natural gas (NG) markets due to heightened demand for heating and electricity generation. While extended weather forecasting has become an indicator for NG management, seasonal (2-3 month) prediction could mitigate the impact of extreme winters on the NG market for consumers and industry. Interrelated climate patterns of ocean and atmospheric circulation anomalies exhibit characteristics useful for developing effective seasonal outlooks of NG storage and consumption due to their influence on the persistence and intensity of extreme winter weather in North America. This study explores the connection between the Pacific-North American climate systems and the NG market in the U.S., connecting macro-scale oceanic and atmospheric processes to regional NG storage and consumption. Western Pacific sea surface temperatures and atmospheric pressure patterns describe significant variation in seasonal NG storage and consumption. Prediction of these coupled climate processes is useful for estimating NG storage and consumption; this could facilitate economic adaptation toward extreme winter weather conditions. Understanding the implicated impact of climate variability on NG is a crucial step toward economic adaptation to climate change.

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Studies of climate effects on ecology often account for non-stationarity in individual physical and biological variables, but rarely allow for non-stationary relationships among variables. Here, we show that non-stationary relationships among physical and biological variables are central to understanding climate effects on salmon (Onchorynchus spp.) in the Gulf of Alaska during 1965-2012. The relative importance of two leading patterns in North Pacific climate, the Pacific Decadal Oscillation (PDO) and North Pacific Gyre Oscillation (NPGO), changed around 1988/1989 as reflected by changing correlations with leading axes of sea surface temperature variability. Simultaneously, relationships between the PDO and Gulf of Alaska environmental variables weakened, and long-standing temperature-salmon and PDO-salmon covariance declined to zero. We propose a mechanistic explanation for changing climate-salmon relationships in terms of non-stationary atmosphere-ocean interactions coinciding with changing PDO-NPGO relative importance. We also show that regression models assuming stationary climate-salmon relationships are inappropriate over the multidecadal time scale we consider. Relaxing assumptions of stationary relationships markedly improved modelling of climate effects on salmon catches and productivity. Attempts to understand the implications of changing climate patterns in other ecosystems might also be aided by the application of models that allow associations among environmental and biological variables to change over time.

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BACKGROUND: Spatial scale is important when studying ecological processes. The Greater sage-grouse (Centrocercus urophasianus) is a large sexually dimorphic tetraonid that is endemic to the sagebrush biome of western North America. The impacts of oil and gas (OAG) development at individual leks has been well-documented. However, no previous studies have quantified the population-level response. METHODS: Hierarchical models were used to estimate the effects of the areal disturbance due to well pads as well as climatic variation on individual lek counts and Greater sage-grouse populations (management units) over 32 years. The lek counts were analyzed using generalized linear mixed models while the management units were analyzed using Gompertz population dynamic models. The models were fitted using frequentist and Bayesian methods. An information-theoretic approach was used to identify the most important spatial scale and time lags. The relative importance of OAG and climate at the local and population-level scales was assessed using information-theoretic (Akaike's weights) and estimation (effect size) statistics. RESULTS: At the local scale, OAG was an important negative predictor of the lek count. At the population scale, there was only weak support for OAG as a predictor of density changes but the estimated impacts on the long-term carrying capacity were consistent with summation of the local impacts. Regional climatic variation, as indexed by the Pacific Decadal Oscillation, was an important positive predictor of density changes at both the local and population level (particularly in the most recent part of the time series). CONCLUSIONS: Additional studies to reduce the uncertainty in the range of possible effects of OAG at the population scale are required. Wildlife agencies need to account for the effects of regional climatic variation when managing sage-grouse populations.

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Understanding and modeling population change is urgently needed to predict effects of climate change on biodiversity. High trophic-level organisms are influenced by fluctuations of prey quality and abundance, which themselves may depend on climate oscillations. Modeling effects of such fluctuations is challenging because prey populations may vary with multiple climate oscillations occurring at different time scales. The analysis of a 28-yr time series of capture-recapture data of a tropical seabird, the Nazca Booby (Sula granti), in the Galápagos, Ecuador, allowed us to test for demographic effects of two major ocean oscillations occurring at distinct time-scales: the inter-annual El Niño Southern Oscillation (ENSO) and inter-decadal oscillations. As expected for a tropical seabird, survival of fledgling birds was highly affected by extreme ENSO events; by contrast, neither recruitment nor breeding participation were affected by either ENSO or decadal oscillations. More interesting, adult survival, a demographic trait that canalizes response to environmental variations, was unaffected by inter-annual ENSO oscillations yet was shaped by the Pacific Decadal Oscillation and small pelagic fish regime. Adult survival decreased during oceanic conditions associated with higher breeding success, an association probably mediated in this species by costs of reproduction that reduce survival when breeding attempts end later. To our knowledge, this is the first study suggesting that survival of a vertebrate can be vulnerable to a natural multidecadal oscillation.

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The degree to which ecosystems are regulated through bottom-up, top-down, or direct physical processes represents a long-standing issue in ecology, with important consequences for resource management and conservation. In marine ecosystems, the role of bottom-up and top-down forcing has been shown to vary over spatio-temporal scales, often linked to highly variable and heterogeneously distributed environmental conditions. Ecosystem dynamics in the Northeast Pacific have been suggested to be predominately bottom-up regulated. However, it remains unknown to what extent top-down regulation occurs, or whether the relative importance of bottom-up and top-down forcing may shift in response to climate change. In this study, we investigate the effects and relative importance of bottom-up, top-down, and physical forcing during changing climate conditions on ecosystem regulation in the Southern California Current System (SCCS) using a generalized food web model. This statistical approach is based on nonlinear threshold models and a long-term data set (~60 years) covering multiple trophic levels from phytoplankton to predatory fish. We found bottom-up control to be the primary mode of ecosystem regulation. However, our results also demonstrate an alternative mode of regulation represented by interacting bottom-up and top-down forcing, analogous to wasp-waist dynamics, but occurring across multiple trophic levels and only during periods of reduced bottom-up forcing (i.e., weak upwelling, low nutrient concentrations, and primary production). The shifts in ecosystem regulation are caused by changes in ocean-atmosphere forcing and triggered by highly variable climate conditions associated with El Niño. Furthermore, we show that biota respond differently to major El Niño events during positive or negative phases of the Pacific Decadal Oscillation (PDO), as well as highlight potential concerns for marine and fisheries management by demonstrating increased sensitivity of pelagic fish to exploitation during El Niño.

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The Humboldt Current System (HCS) has the highest production of forage fish in the world, although it is highly variable and the future of the primary component, anchovy, is uncertain in the context of global warming. Paradigms based on late 20th century observations suggest that large-scale forcing controls decadal-scale fluctuations of anchovy and sardine across different boundary currents of the Pacific. We develop records of anchovy and sardine fluctuations since 1860 AD using fish scales from multiple sites containing laminated sediments and compare them with Pacific basin-scale and regional indices of ocean climate variability. Our records reveal two main anchovy and sardine phases with a timescale that is not consistent with previously proposed periodicities. Rather, the regime shifts in the HCS are related to 3D habitat changes driven by changes in upwelling intensity from both regional and large-scale forcing. Moreover, we show that a long-term increase in coastal upwelling translates via a bottom-up mechanism to top predators suggesting that the warming climate, at least up to the start of the 21st century, was favorable for fishery productivity in the HCS.

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Primary producers respond to climate directly and indirectly due to effects on their consumers. In the temperate coastal ocean, the highly productive brown algae known as kelp have both strong climate and grazer linkages. We analyzed the demographic response of the kelp Pleurophycus gardneri over a 25-year span to determine the interaction between ocean climate indicators and invertebrate infestation rates. Pleurophycus hosts amphipod species that burrow in the stipe, increasing mortality. Although kelp performance is generally greater with more negative values of the Pacific Decadal Oscillation (PDO) and colder seawater temperatures, Pleurophycus showed the opposite pattern. When we compared the 1990s, a period of positive values for the PDO and warmer sea surface temperatures, with the following decade, a period characterized by negative PDO values, we documented a contradictory outcome for proxies of kelp fitness. In the 1990s, Pleurophycus unexpectedly showed greater longevity, faster growth, greater reproductive effort, and a trend toward decreased amphipod infestation compared with the 2006-2012 period. In contrast, the period from 2006 to 2012 showed opposite kelp performance patterns and with a trend toward greater amphipod infestation. Pleurophycus performance metrics suggest that some coastal primary producers will respond differently to climate drivers, particularly if they interact strongly with grazers.

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Domoic acid is a potent neurotoxin produced by certain marine microalgae that can accumulate in the foodweb, posing a health threat to human seafood consumers and wildlife in coastal regions worldwide. Evidence of climatic regulation of domoic acid in shellfish over the past 20 y in the Northern California Current regime is shown. The timing of elevated domoic acid is strongly related to warm phases of the Pacific Decadal Oscillation and the Oceanic Niño Index, an indicator of El Niño events. Ocean conditions in the northeast Pacific that are associated with warm phases of these indices, including changes in prevailing currents and advection of anomalously warm water masses onto the continental shelf, are hypothesized to contribute to increases in this toxin. We present an applied domoic acid risk assessment model for the US West Coast based on combined climatic and local variables. Evidence of regional- to basin-scale controls on domoic acid has not previously been presented. Our findings have implications in coastal zones worldwide that are affected by this toxin and are particularly relevant given the increased frequency of anomalously warm ocean conditions.

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Novel forest decline is increasing due to global environmental change, yet the causal factors and their interactions remain poorly understood. Using tree ring analyses, we show how climate and multiple biotic factors caused the decline of whitebark pine (Pinus albicaulis) in 16 stands in the southern Canadian Rockies. In our study area, 72% of whitebark pines were dead and 18% had partially dead crowns. Tree mortality peaked in the 1970s; however, the annual basal area increment of disturbed trees began to decline significantly in the late 1940s. Growth decline persisted up to 30 years before trees died from mountain pine beetle (Dendroctonus ponderosae), Ips spp. bark beetles or non-native blister rust pathogen (Cronartium ribicola). Climate-growth relations varied over time and differed among the healthy and disturbed subpopulations of whitebark pine. Prior to the 1940s, cool temperatures limited the growth of all subpopulations. Growth of live, healthy trees became limited by drought during the cool phase (1947 -1976) of the Pacific Decadal Oscillation (PDO) and then reverted to positive correlations with temperature during the subsequent warm PDO phase. In the 1940s, the climate-growth relations of the disturbed subpopulations diverged from the live, healthy trees with trees ultimately killed by mountain pine beetle diverging the most. We propose that multiple factors interacted over several decades to cause unprecedented rates of whitebark pine mortality. Climatic variation during the cool PDO phase caused drought stress that may have predisposed trees to blister rust. Subsequent decline in snowpack and warming temperatures likely incited further climatic stress and with blister rust reduced tree resistance to bark beetles. Ultimately, bark beetles and blister rust contributed to tree death. Our findings suggest the complexity of whitebark pine decline and the importance of considering multiway drought-disease-insect interactions over various timescales when interpreting forest decline.

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