William J. Sydeman

The Pace of Shifting Climate in Marine and Terrestrial Ecosystems

Ecologically relevant measures of contemporary global climate change can predict species distributions and vulnerabilities. Climate change challenges organisms to adapt or move to track changes in environments in space and time. We used two measures of thermal shifts from analyses of global temperatures over the past 50 years to describe the pace of climate change that species should track: the velocity of climate change (geographic shifts of isotherms over time) and the shift in seasonal timing of temperatures. Both measures are higher in the ocean than on land at some latitudes, despite slower ocean warming. These indices give a complex mosaic of predicted range shifts and phenology changes that deviate from simple poleward migration and earlier springs or later falls. They also emphasize potential conservation concerns, because areas of high marine biodiversity often have greater velocities of climate change and seasonal shifts.

Global Seabird Response to Forage Fish Depletion—One-Third for the Birds

One-third of maximum fish biomass must be available for seabirds to sustain high breeding success. Determining the form of key predator-prey relationships is critical for understanding marine ecosystem dynamics. Using a comprehensive global database, we quantified the effect of fluctuations in food abundance on seabird breeding success. We identified a threshold in prey (fish and krill, termed “forage fish”) abundance below which seabirds experience consistently reduced and more variable productivity. This response was common to all seven ecosystems and 14 bird species examined within the Atlantic, Pacific, and Southern Oceans. The threshold approximated one-third of the maximum prey biomass observed in long-term studies. This provides an indicator of the minimal forage fish biomass needed to sustain seabird productivity over the long term.

Geographical limits to species-range shifts are suggested by climate velocity

The reorganization of patterns of species diversity driven by anthropogenic climate change, and the consequences for humans, are not yet fully understood or appreciated. Nevertheless, changes in climate conditions are useful for predicting shifts in species distributions at global and local scales. Here we use the velocity of climate change to derive spatial trajectories for climatic niches from 1960 to 2009 (ref. 7) and from 2006 to 2100, and use the properties of these trajectories to infer changes in species distributions. Coastlines act as barriers and locally cooler areas act as attractors for trajectories, creating source and sink areas for local climatic conditions. Climate source areas indicate where locally novel conditions are not connected to areas where similar climates previously occurred, and are thereby inaccessible to climate migrants tracking isotherms: 16% of global surface area for 1960 to 2009, and 34% of ocean for the ‘business as usual’ climate scenario (representative concentration pathway (RCP) 8.5) representing continued use of fossil fuels without mitigation. Climate sink areas are where climate conditions locally disappear, potentially blocking the movement of climate migrants. Sink areas comprise 1.0% of ocean area and 3.6% of land and are prevalent on coasts and high ground. Using this approach to infer shifts in species distributions gives global and regional maps of the expected direction and rate of shifts of climate migrants, and suggests areas of potential loss of species richness.

Climate change and wind intensification in coastal upwelling ecosystems

Strong winds, upwelling, and teeming shores Climate warming has produced stronger winds along some coasts, a result of growing differences in temperature and pressure between land and sea. These winds cause cold nutrient-rich seawater to rise to the surface, affecting climate and fueling marine productivity. Sydeman et al. examined data from the five major world regions where upwelling is occurring. Particularly in the California, Humboldt, and Benguela upwelling systems, winds have become stronger over the past 60 years. These regions represent up to a fifth of wild marine fish catches and are hot spots of biodiversity. Science, this issue p. 77 Increasing greenhouse gas concentrations have caused windier conditions in most major coastal upwelling regions. In 1990, Andrew Bakun proposed that increasing greenhouse gas concentrations would force intensification of upwelling-favorable winds in eastern boundary current systems that contribute substantial services to society. Because there is considerable disagreement about whether contemporary wind trends support Bakun’s hypothesis, we performed a meta-analysis of the literature on upwelling-favorable wind intensification. The preponderance of published analyses suggests that winds have intensified in the California, Benguela, and Humboldt upwelling systems and weakened in the Iberian system over time scales ranging up to 60 years; wind change is equivocal in the Canary system. Stronger intensification signals are observed at higher latitudes, consistent with the warming pattern associated with climate change. Overall, reported changes in coastal winds, although subtle and spatially variable, support Bakun’s hypothesis of upwelling intensification in eastern boundary current systems.

Breeding Performance in the Western Gull: Effects of Parental Age, Timing of Breeding and Year in Relation to Food Availability

(1) We studied age-specific breeding success of western gulls (Larus occidentalis) on south-east Farallon Island, California in relation to annual variation in food abundance, 1983-89. We assessed yearly reproductive performance on the basis of clutch initiation date, clutch size, hatching and fledging success, and the number of chicks raised to independence. We categorized age of parent into three groups: young, middle-aged and old. (2) Mature birds initiated breeding earlier and laid larger clutches; they fledged more chicks than younger birds as a result of hatching a greater proportion of their eggs and fledging a greater proportion of their hatched chicks. The oldest birds generally did not differ from middle-aged in breeding performance. The difference in reproductive performance between young and mature birds was most apparent during years when food availability was reduced. (3) Timing of breeding influenced reproductive performance independently of parental age; reproductive performance decreased throughout the breeding season. Within-season reductions in reproductive performance were most apparent for young breeders and individuals with small clutches or broods. (4) Breeding performance varied significantly among years independently of age of the study animals. Food availability accounted for part of the yearly variation in breeding success. (5) We suggest that clutch/brood size reflects individual quality. Age-specific breeding success in western gulls is best explained by previous breeding experience and individual quality.

Trophic relationships among seabirds in central California : Combined stable isotope and conventional dietary approach

We used stable isotope analysis (SIA) and conventional techniques of diet assessment to determine marine trophic relationships in the Gulf of the Farallones, California, with an emphasis on marine birds. Stable-carbon (δ 13 C) and nitrogen (δ 15 N) isotopes were obtained from 98 tissue samples of 16 species representing primary and secondary consumers in 1993-1994. The values of δ 13 C ranged from -20.1‰ in whole euphausiids (krill) to -15.0‰ in muscle of northern sea lions. Values of δ 15 N showed step-wise trophic enrichment and ranged from 11.2‰ in euphausiids to 19.8‰ in sea lions. SIA of egg albumen from birds indicated reliance on zooplankton by Cassins Auklet, Common Murre, and Western Gull, and on fish by Brandts and Pelagic Cormorants, Rhinoceros Auklets, and Pigeon Guillemots during egg formation (April-May). However, analysis of prey brought to chicks during summer indicated the prevalence of fish in the diet of most seabirds, except Cassins Auklet which fed primarily on krill. Results suggest a shift in trophic level and diet between spring and summer from krill to fish for Common Murres. δ 13 C analysis confirmed that Brandts Cormorants and northern sea lions feed in neritic habitats, whereas Cassins and Rhinoceros Auklets foraged in epipelagic offshore waters. Our approach demonstrates the utility of combining both SIA and conventional dietary assessments to understand trophic relationships in dynamic marine ecosystems.

Quantitative approaches in climate change ecology

Contemporary impacts of anthropogenic climate change on ecosystems are increasingly being recognized. Documenting the extent of these impacts requires quantitative tools for analyses of ecological observations to distinguish climate impacts in noisy data and to understand interactions between climate variability and other drivers of change. To assist the development of reliable statistical approaches, we review the marine climate change literature and provide suggestions for quantitative approaches in climate change ecology. We compiled 267 peer-reviewed articles that examined relationships between climate change and marine ecological variables. Of the articles with time series data (n = 186), 75% used statistics to test for a dependency of ecological variables on climate variables. We identified several common weaknesses in statistical approaches, including marginalizing other important non-climate drivers of change, ignoring temporal and spatial autocorrelation, averaging across spatial patterns and not reporting key metrics. We provide a list of issues that need to be addressed to make inferences more defensible, including the consideration of (i) data limitations and the comparability of data sets; (ii) alternative mechanisms for change; (iii) appropriate response variables; (iv) a suitable model for the process under study; (v) temporal autocorrelation; (vi) spatial autocorrelation and patterns; and (vii) the reporting of rates of change. While the focus of our review was marine studies, these suggestions are equally applicable to terrestrial studies. Consideration of these suggestions will help advance global knowledge of climate impacts and understanding of the processes driving ecological change.

Timing is everything: flexible phenology and shifting selection in a colonial seabird

1. In order to reproduce successfully in a temporally varying environment, iteroparous animals must exhibit considerable behavioural flexibility across their lifetimes. By adjusting timing of breeding each year, parents can ensure optimal overlap between the energy intensive period of offspring production and the seasonal peak in favourable environmental conditions, thereby increasing their chances of successfully rearing young. 2. Few studies investigate variation among individuals in how they respond to fluctuating conditions, or how selection acts on these individual differences, but this information is essential for understanding how populations will cope with rapid environmental change. 3. We explored inter-annual trends in breeding time and individual responses to environmental variability in common guillemots Uria aalge, an important marine top predator in the highly variable California Current System. Complex, nonlinear relationships between phenology and oceanic and climate variables were found at the population level. Using a novel application of a statistical technique called random regression, we showed that individual females responded in a nonlinear fashion to environmental variability, and that reaction norm shape differed among females. 4. The pattern and strength of selection varied substantially over a 34-year period, but in general, earlier laying was favoured. Females deviating significantly from the population mean laying date each year also suffered reduced breeding success, with the strength of nonlinear selection varying in relation to environmental conditions. 5. We discuss our results in the wider context of an emerging literature on the evolutionary ecology of individual-level plasticity in the wild. Better understanding of how species-specific factors and local habitat features affect the timing and success of breeding will improve our ability to predict how populations will respond to climate change.

Anticipated Effects of Climate Change on Coastal Upwelling Ecosystems

Ecosystem productivity in coastal ocean upwelling systems is threatened by climate change. Increases in spring and summer upwelling intensity, and associated increases in the rate of offshore advection, are expected. While this could counter effects of habitat warming, it could also lead to more frequent hypoxic events and lower densities of suitable-sized food particles for fish larvae. With upwelling intensification, ocean acidity will rise, affecting organisms with carbonate structures. Regardless of changes in upwelling, near-surface stratification, turbulent diffusion rates, source water origins, and perhaps thermocline depths associated with large-scale climate episodes (ENSO) maybe affected. Major impacts on pelagic fish resources appear unlikely unless couples with overfishing, although changes toward more subtropical community composition are likely. Marine mammals and seabirds that are tied to sparsely distributed nesting or resting grounds could experience difficulties in obtaining prey resources, or adaptively respond by moving to more favorable biogeographic provinces.

Responses of marine organisms to climate change across oceans

Climate change is driving changes in the physical and chemical properties of the ocean that have consequences for marine ecosystems. Here, we review evidence for the responses of marine life to recent climate change across ocean regions, from tropical seas to polar oceans. We consider observed changes in calcification rates, demography, abundance, distribution and phenology of marine species. We draw on a database of observed climate change impacts on marine species, supplemented with evidence in the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. We discuss factors that limit or facilitate species’ responses, such as fishing pressure, the availability of prey, habitat, light and other resources, and dispersal by ocean currents. We find that general trends in species responses are consistent with expectations from climate change, including poleward and deeper distributional shifts, advances in spring phenology, declines in calcification and increases in the abundance of warm-water species. The volume and type of evidence of species responses to climate change is variable across ocean regions and taxonomic groups, with much evidence derived from the heavily-studied north Atlantic Ocean. Most investigations of marine biological impacts of climate change are of the impacts of changing temperature, with few observations of effects of changing oxygen, wave climate, precipitation (coastal waters) or ocean acidification. Observations of species responses that have been linked to anthropogenic climate change are widespread, but are still lacking for some taxonomic groups (e.g., phytoplankton, benthic invertebrates, marine mammals).