Niches, models, and climate change: assessing the assumptions and uncertainties.

As the rate and magnitude of climate change accelerate, understanding the consequences becomes increasingly important. Species distribution models (SDMs) based on current ecological niche constraints are used to project future species distributions. These models contain assumptions that add to the uncertainty in model projections stemming from the structure of the models, the algorithms used to translate niche associations into distributional probabilities, the quality and quantity of data, and mismatches between the scales of modeling and data. We illustrate the application of SDMs using two climate models and two distributional algorithms, together with information on distributional shifts in vegetation types, to project fine-scale future distributions of 60 California landbird species. Most species are projected to decrease in distribution by 2070. Changes in total species richness vary over the state, with large losses of species in some "hotspots" of vulnerability. Differences in distributional shifts among species will change species co-occurrences, creating spatial variation in similarities between current and future assemblages. We use these analyses to consider how assumptions can be addressed and uncertainties reduced. SDMs can provide a useful way to incorporate future conditions into conservation and management practices and decisions, but the uncertainties of model projections must be balanced with the risks of taking the wrong actions or the costs of inaction. Doing this will require that the sources and magnitudes of uncertainty are documented, and that conservationists and resource managers be willing to act despite the uncertainties. The alternative, of ignoring the future, is not an option.

Where the wild things are: predicting hotspots of seabird aggregations in the California Current System.

Marine protected areas (MPAs) provide an important tool for conservation of marine ecosystems. To be most effective, these areas should be strategically located in a manner that supports ecosystem function. To inform marine spatial planning and support strategic establishment of MPAs within the California Current System, we identified areas predicted to support multispecies aggregations of seabirds ("hotspots"). We developed habitat-association models for 16 species using information from at-sea observations collected over an 11-year period (1997-2008), bathymetric data, and remotely sensed oceanographic data for an area from north of Vancouver Island, Canada, to the USA/Mexico border and seaward 600 km from the coast. This approach enabled us to predict distribution and abundance of seabirds even in areas of few or no surveys. We developed single-species predictive models using a machine-learning algorithm: bagged decision trees. Single-species predictions were then combined to identify potential hotspots of seabird aggregation, using three criteria: (1) overall abundance among species, (2) importance of specific areas ("core areas") to individual species, and (3) predicted persistence of hotspots across years. Model predictions were applied to the entire California Current for four seasons (represented by February, May, July, and October) in each of 11 years. Overall, bathymetric variables were often important predictive variables, whereas oceanographic variables derived from remotely sensed data were generally less important. Predicted hotspots often aligned with currently protected areas (e.g., National Marine Sanctuaries), but we also identified potential hotspots in Northern California/Southern Oregon (from Cape Mendocino to Heceta Bank), Southern California (adjacent to the Channel Islands), and adjacent to Vancouver Island, British Columbia, that are not currently included in protected areas. Prioritization and identification of multispecies hotspots will depend on which group of species is of highest management priority. Modeling hotspots at a broad spatial scale can contribute to MPA site selection, particularly if complemented by fine-scale information for focal areas.

Matching the multiple scales of conservation with the multiple scales of climate change.

To anticipate the rapidly changing world resulting from global climate change, the projections of climate models must be incorporated into conservation. This requires that the scales of conservation be aligned with the scales of climate-change projections. We considered how conservation has incorporated spatial scale into protecting biodiversity, how the projections of climate-change models vary with scale, and how the two do or do not align. Conservation planners use information about past and current ecological conditions at multiple scales to identify conservation targets and threats and guide conservation actions. Projections of climate change are also made at multiple scales, from global and regional circulation models to projections downscaled to local scales. These downscaled projections carry with them the uncertainties associated with the broad-scale models from which they are derived; thus, their high resolution may be more apparent than real. Conservation at regional or global scales is about establishing priorities and influencing policy. At these scales, the coarseness and uncertainties of global and regional climate models may be less important than what they reveal about possible futures. At the ecoregional scale, the uncertainties associated with downscaling climate models become more critical because the distributions of conservation targets on which plans are founded may shift under future climates. At a local scale, variations in topography and land cover influence local climate, often overriding the projections of broad-scale climate models and increasing uncertainty. Despite the uncertainties, ecologists and conservationists must work with climate-change modelers to focus on the most likely projections. The future will be different from the past and full of surprises; judicious use of model projections at appropriate scales may help us prepare.

Assessing Ecological Impact Assessment: Lessons from Mono Lake, California.

Because of its high salinity and alkalinity, Mono Lake, in eastern California (USA), is a relatively simple ecosystem. It has become the focus of an environmental controversy over the effects of 50 yr of diversions of water from tributary streams to supply water to Los Angeles. Diversions lowered the lake level, increased the salinity, changed the availability of aquatic habitats, and altered the configuration of the shoreline and of islands that support breeding colonies of gulls. We consider (1) how two independent panels of experts synthesized scientific information on the lake ecosystem to assess the environmental consequences of these changes, and (2) how the findings of these groups influenced policy decisions and how well subsequent changes in the lake matched expectations. Despite differences in composition and approach, the two panels reached generally similar conclusions. These conclusions have been a major component of legal activities and the development of management plans for the lake and basin ecosystem. Both panels concluded that, because of the simplicity of the lake ecosystem, ecological consequences of changes in lake level and salinity associated with continuing diversions were likely to be unusually clear-cut. At certain lake levels these changes would be expected to alter algal and invertebrate populations and the populations of aquatic birds that feed upon them or to disrupt breeding activities in gull colonies. Projections about when critical lake levels might be reached, however, have not been met. This is largely because stream flows into the lake have been altered from recent historic patterns by the cessation of water diversions due to governmental and legal actions (prompted in part by the panels' findings) and by a prolonged drought. These events illustrate the difficulty of projecting a timetable for environmental changes, even in simple and well-studied ecosystems.

Model estimation of energy flow in North American grassland bird communities.

The energy demands and general food consumption rates of bird populations breeding in North American grasslands are estimated using a simulation model which employs information on population natural history and individual metabolism gathered from several study locations. The total breeding season energy demand of the grassland/shrub-steppe avifaunas ranged from 0.89 kcal m-2 season-1 in arid shrub-steppe to 2.92 kcal m-2 season-1 in a mesic tallgrass prairie. There was substantial variation between years and between census plots, however, and in general the average avian community energy demands did not differ significantly over the range of locations. Production accounted for 0.9 to 1.5% of the total seasonal energy demand. Roughly 11 to 18% of the seasonal energy flow was required in the production of eggs and maintenance and growth of nestlings and fledglings.On the average, between 209 and 386 kg dry wt km-2 of prey were consumed by the bird communities breeding in the grassland locations. Seeds contributed more to the total biomass consumed at the drier plots, but in general, animal prey types comprised roughly 80% of the total biomass eaten. Phytophagous insects were the major component of the animal prey.These low magnitudes of energy flow and biomass consumption attest to the relatively minor role of birds in the processing of energy and biomass in grassland ecosystems. If these populations do play an 'importnat' role in the functioning of grassland ecosystems, it must be quite subtle and indirect.

Temporal variation in habitat structure and shrubsteppe bird dynamics.

We investigate the role of temporal variation in habitat physiognomy in influencing the dynamics of shrubsteppe bird populations and communities. During a 3-y (1977-1979) study of 14 sites in the northern Great Basin of North America, annual precipitation varied substantially, with one of the driest years on record followed by one of the wettest. This resulted in significant physiognomic variation (increasing height and coverage of vegetation, decreasing horizontal patchiness), mediated largely by changes in the annual elements of the flora, particularly forbs and grasses. Shrub species coverage values, on the other hand, demonstrated no statistically detectable year-to-year changes, nor were they correlated with any physiognomic variation. Despite large scale physiognomic changes, no bird species' abundance varied in a statistically significant fashion; neither could variation in bird abundances be correlated with variation in either physiognomy or shrub species coverages.Multivariate analyses revealed essentially the same patterns as the univariate analyses: substantial changes in physiognomy, few changes in shrub species coverages or bird species abundances, and little correlation of temporal variation among the three data sets. Calculation of the Euclidean distances that sites "moved" in multivariate physiognomic, bird species, or shrub species hyperspaces yields synthetic gradients of annual "turnover" of sites with respect to those data sets. Sites identified as demonstrating high physiognomic turnover were characterized by high coverage of grass and forbs, while low turnover sites had greater coverage of shrubs and higher shrub species diversity. Relatively high bird turnover sites had greater numbers of Western Meadowlarks and Black-throated Sparrows, while more stable sites had high numbers of Brewer's Sparrows. Physiognomically, high bird turnover sites were grassier and had greater total vegetation coverage, while low bird turnover sites had more bare ground and higher horizontal patchiness. A site's position on the avifauna turnover axis, however, was uncorrelated with its position on the physiognomic turnover axis. Shrub species showed virtually no annual turnover.Reanalysis of a previous Principal Components Analysis (PCA) of these same data sets that was applied without regard to year of sampling revealed that the first physiognomic component (41% of the total physiognomic variability) did in fact have a strong temporal element, and that this element was consistent with the changes in univariate characters noted above. No other physiognomic component could be associated with annual variation, nor could any components of parallel bird abundance or shrub species coverage PCA's.Regional patterns indicate that sites tended to be very consistent from year to year in their relationships to one another as defined by their relative locations in either physiognomic or shrub species hyperspaces, but varied independently of one another with respect to their bird species abundances and composition. Examination of the temporal consistency of site relationships between "bird space" and "vegetation space" reveals that bird communities are to a large degree independent of a site's physiognomic position, but instead are strongly associated with its position in shrub species hyperspace.The overall patterns that emerge from these analyses are consistent with the so-called "checkerboard effect" that results from the apparently random annual redistribution of individual birds, and leads to the conclusion that populations of shrubsteppe birds are not existing at maximum density or "carrying capacity." Such observations are consistent with contentions that these populations lack close biological coupling with coexisting species and that interactions among these species (e.g. competition) are likely to play little if any role in the organization of their communities.

Diet niche relationships among North American grassland and shrubsteppe birds.

We consider the dietary relationships of the numerically dominant breeding bird species in four North American grassland/shrubsteppe habitats, sampled over 2-3 consecutive years. Overall, the diets of these species contained primarily insects: orthopterans comprised 29% of the diet biomass, coleopterans 24%, and lepidopteran larvae 23%, while seeds contributed 15% of the average diet. These diets varied substantially, however, and we evaluated several aspects of this variation. Intersexual differences in diets within a species were few, despite the occurrence of significant sexual size dimorphism in several species. For many species, however, there were substantial shifts in dietary composition between years at a given location; overall, the average between-year similarity of species' dietary composition was 70%. Different species exhibited rather different diet patterns. Horned Larks were relatively omnivorous, had broad diet composition niches, and varied considerably in diets between different locations. Meadowlarks were also broad-niched and geographically variable in their diets, but were the most highly carnivorous of the species we considered. Dietary niche breadths of Grasshopper Sparrows were intermediate, but diet composition was rather stable, both between years and between locations. Chestnut-collared Longspurs exhibited narrow diet niches, but substantial annual variation: each year this species apparently exploited a different but limited set of prey types rather heavily. Larger avian predators generally consumed a broader array of functional groups of prey, but did not differ in the taxonomic variety of their diets from small birds. Variation in diet composition between individuals within local populations was considerable; in most species, an individual contained on the average 30-40% of the prey taxa represented in entire population smaples.Patterns of dietary overlap among species were quite inconsistent from year to year at most locations, although at the shrubsteppe site overlap among all species present was consistently quite high. Relatively few cooccurring species pairs exhibited low diet overlap. The degree of diet niche overlap was unrelated to body size differences of the birds, despite as much as six-fold differences in weight among some coexisting species. Relationships of the bird species on another dimension of the trophic niche, prey size, also differed substantially between sites and years. The ranking of co-occurring species by the mean sizes of the prey they consumed generally did not parallel their rankings by body sizes, and in some cases the smallest and the largest species present ate prey of similar sizes. At the shrubsteppe site, all the breeding species exhibited quite similar frequency distributions of prey sizes in their diets.As species number and diversity increased in the breeding avifaunas, diet niche breadths generally decreased, species packing by prey size decreased, and diet composition niche overlap remained relatively unchanged. These trends are in at least partial agreement with predictions of diffuse competition theory, but the patterns were derived from broad inter-site comparisons of overall site averages, and the relationships generally did not hold within local assemblages of species. In general, our attempts to match values of dietary niche features with site characteristics failed to demonstrate close agreement with the predictions of prevailing ecological theory based upon assumptions of resource limitation and competition. Instead, our findings seem generally most consistent with the suggestion that food is not normally limiting to bird populations in these systems, and individuals and populations are exploiting the food resources in an opportunistic fashion, which leads to considerable individual, between-year, and between-location variation in diet compositions and interspecific overlaps.Our attempts to discern clear relationships that accord with theoretical expectations in these avian assemblages are thwarted by our lack of detailed information on the resource base and by the lack of clear tests that will separate alternative hypotheses of community organization and structuring. We suggest that these complications may compromise the findings of many community studies.

Quantitative Assessment of Current Risks to Harlequin Ducks in Prince William Sound, Alaska, from the Exxon Valdez Oil Spill.

Harlequin Ducks (Histrionicus histrionicus) were adversely affected by the Exxon Valdez oil spill (EVOS) in Prince William Sound (PWS), Alaska, and some have suggested effects continue two decades later. We present an ecological risk assessment evaluating quantitatively whether PWS seaducks continue to be at-risk from polycyclic aromatic hydrocarbons (PAHs) in residual Exxon Valdez oil. Potential pathways for PAH exposures are identified for initially oiled and never-oiled reference sites. Some potential pathways are implausible (e.g., a seaduck excavating subsurface oil residues), whereas other pathways warrant quantification. We used data on PAH concentrations in PWS prey species, sediments, and seawater collected during 2001-2008 to develop a stochastic individual-based model projecting assimilated doses to seaducks. We simulated exposures to 500,000 individuals in each of eight age/gender classes, capturing the variability within a population of seaducks living in PWS. Doses to the maximum-exposed individuals are ∼400-4,000 times lower than chronic toxicity reference values established using USEPA protocols for seaducks. These exposures are so low that no individual-level effects are plausible, even within a simulated population that is orders-of-magnitude larger than exists in PWS. We conclude that toxicological risks to PWS seaducks from residual Exxon Valdez oil two decades later are essentially non-existent.

Re-shuffling of species with climate disruption: a no-analog future for California birds?

By facilitating independent shifts in species' distributions, climate disruption may result in the rapid development of novel species assemblages that challenge the capacity of species to co-exist and adapt. We used a multivariate approach borrowed from paleoecology to quantify the potential change in California terrestrial breeding bird communities based on current and future species-distribution models for 60 focal species. Projections of future no-analog communities based on two climate models and two species-distribution-model algorithms indicate that by 2070 over half of California could be occupied by novel assemblages of bird species, implying the potential for dramatic community reshuffling and altered patterns of species interactions. The expected percentage of no-analog bird communities was dependent on the community scale examined, but consistent geographic patterns indicated several locations that are particularly likely to host novel bird communities in the future. These no-analog areas did not always coincide with areas of greatest projected species turnover. Efforts to conserve and manage biodiversity could be substantially improved by considering not just future changes in the distribution of individual species, but including the potential for unprecedented changes in community composition and unanticipated consequences of novel species assemblages.