Relationship of size at return with environmental variation,hatchery production,and productivity of wild pink salmon in Prince William Sound,Alaska: does size matter?

Pink salmon (Oncorhynchus gorbuscha) returning to Prince William Sound (PWS), Alaska, have increased to historically high levels of abundance in recent years, but average body size at return has declined. We examined how body size at return of PWS pink salmon was related to 10 biophysical factors, including the scale of hatchery production. We also examined the effect of body size at return on productivity of wild pink salmon in PWS. For the 1975–1999 brood years, we found that an index of total abundance of pink salmon in the Gulf of Alaska and sea surface temperature during the year of return best explained the variation in pink salmon body size over time. Body size at return was significantly correlated with productivity of wild pink salmon. We used stepwise-regression to fit a generalized linear version of the Ricker spawner-recruit model to determine if body size would explain significant variation in wild-stock productivity in context with other environmental variation, including hatchery production. The results indicate that variability in wild-stock productivity is primarily driven by density-independent factors in the marine environment, but that body size of wild spawners also significantly affects productivity of wild PWS pink salmon. We conclude that the success of large-scale enhancement increasing the total run in PWS may have contributed to the decline in body size because of density-dependent growth in the Gulf of Alaska. We used a simulation model to estimate the impact of hatchery-induced changes in adult body size on wild-stock production in PWS. We estimated an annual wild-stock yield loss of 1.03 million pink salmon, less than 5% of the annual hatchery return of 24.2 million adult pink salmon for brood years 1990–1999.     

Asynchrony in population dynamics of sockeye salmon in southwest Alaska

Ecologists have examined the synchronization of population dynamics across space as a means to understand how populations respond to climate variation. However, response diversity may reflect important variation among local population dynamics driven by population‐specific responses to regional environmental change. We used long‐term data on sockeye salmon Oncorhynchus nerka from pristine watersheds of southwestern Alaska to show that populations spawning in close proximity (<40 km) to one another have a limited degree of synchrony in their dynamics, even after accounting for density‐dependent processes. In fact, the dynamics of local populations of stream‐spawning sockeye salmon were no more coherent than those of stocks at a much coarser resolution across this region of Alaska. We examined four hypotheses to explain the observed patterns of asynchrony among stream‐spawning populations, and found that populations spawning in dissimilar habitats, and using different nursery lakes were less synchronized in their productivity. Similarity in the age structure of spawning adults was less correlated with synchrony in productivity. These results emphasize the importance of maintaining diverse spawning and rearing habitat for the conservation of Pacific salmon, and should guide conservation planning for Pacific salmon populations in regions where natural dynamics have been altered by habitat loss, hatchery practices, and over‐fishing.     

Variability in primary productivity determines metapopulation dynamics

Temporal variability in primary productivity can change habitat quality for consumer species by affecting the energy levels available as food resources. However, it remains unclear how habitat-quality fluctuations may determine the dynamics of spatially structured populations, where the effects of habitat size, quality and isolation have been customarily assessed assuming static habitats. We present the first empirical evaluation on the effects of stochastic fluctuations in primary productivity—a major outcome of ecosystem functions—on the metapopulation dynamics of a primary consumer. A unique 13-year dataset from an herbivore rodent was used to test the hypothesis that inter-annual variations in primary productivity determine spatiotemporal habitat occupancy patterns and colonization and extinction processes. Inter-annual variability in productivity and in the growing season phenology significantly influenced habitat colonization patterns and occupancy dynamics. These effects lead to changes in connectivity to other potentially occupied habitat patches, which then feed back into occupancy dynamics. According to the results, the dynamics of primary productivity accounted for more than 50% of the variation in occupancy probability, depending on patch size and landscape configuration. Evidence connecting primary productivity dynamics and spatiotemporal population processes has broad implications for metapopulation persistence in fluctuating and changing environments.     

Serologic survey for Brucella spp., phocid herpesvirus-1, phocid herpesvirus-2, and phocine distemper virus in harbor seals from Alaska, 1976-1999

Harbor seals (Phoca vitulina richardsi) were captured in the coastal regions of Southeast Alaska, Gulf of Alaska, Prince William Sound (PWS), and Kodiak Island during 1976-1999. Blood was collected from 286 seals. Sera were tested for evidence of exposure to Brucella spp., phocid herpesvirus-1 (PhoHV-1), phocid herpesvirus-2 (PhHV-2), and phocine distemper virus (PDV). Antibody prevalence rates were 46% (46/100) for Brucella spp., 93% (225/243) for PhoHV-1, 0% (0/286) for PhHV-2, and 1% (2/160) for PDV. Antibody prevalence for Brucella spp. was directly related to host age. Antibody prevalence for PhoHV-1 was higher in PWS as compared to the other three regions. No evidence of mortality attributable to these four agents was observed during the course of this study. Based on the results of this survey, none of these agents is considered a significant mortality factor in harbor seals from the four regions of coastal Alaska included in the study.     

Linkages between large‐scale climate patterns and the dynamics of Arctic caribou populations

Recent research has linked climate warming to global declines in caribou and reindeer (both Rangifer tarandus) populations. We hypothesize large‐scale climate patterns are a contributing factor explaining why these declines are not universal. To test our hypothesis for such relationships among Alaska caribou herds, we calculated the population growth rate and percent change of four arctic herds using existing population estimates, and explored associations with indices of the Arctic Oscillation (AO) and the Pacific Decadal Oscillation (PDO). The AO, which more strongly affects eastern Alaska, was negatively associated with the population trends of the Porcupine Caribou Herd and Central Arctic Herd, the easternmost of the herds. We hypothesize that either increased snowfall or suboptimal growing conditions for summer forage plants could explain this negative relationship. Intensity of the PDO, which has greatest effects in western Alaska, was negatively associated with the growth rate of the Teshekpuk Caribou Herd in northwestern Alaska, but the Western Arctic Herd in western Alaska displayed the opposite trend. We suggest that the contrasting patterns of association relate to the spatial variability of the effects of the PDO on western and northwestern Alaska. Although predation and winter range quality have often been considered the primary causes of population variation, our results show that large‐scale climate patterns may play an important role in caribou population dynamics in arctic Alaska. Our findings reveal that climate warming has not acted uniformly to reduce caribou populations globally. Further research should focus on the relative importance of mechanisms by which climate indices influence caribou population dynamics.     

Source-Sink Estimates of Genetic Introgression Show Influence of Hatchery Strays on Wild Chum Salmon Populations in Prince William Sound,Alaska

The extent to which stray, hatchery-reared salmon affect wild populations is much debated. Although experiments show that artificial breeding and culture influence the genetics of hatchery salmon, little is known about the interaction between hatchery and wild salmon in a natural setting. Here, we estimated historical and contemporary genetic population structures of chum salmon (Oncorhynchus keta) in Prince William Sound (PWS), Alaska, with 135 single nucleotide polymorphism (SNP) markers. Historical population structure was inferred from the analysis of DNA from fish scales, which had been archived since the late 1960’s for several populations in PWS. Parallel analyses with microsatellites and a test based on Hardy-Weinberg proportions showed that about 50% of the fish-scale DNA was cross-contaminated with DNA from other fish. These samples were removed from the analysis. We used a novel application of the classical source-sink model to compare SNP allele frequencies in these archived fish-scales (1964–1982) with frequencies in contemporary samples (2008–2010) and found a temporal shift toward hatchery allele frequencies in some wild populations. Other populations showed markedly less introgression, despite moderate amounts of hatchery straying. The extent of introgression may reflect similarities in spawning time and life-history traits between hatchery and wild fish, or the degree that hybrids return to a natal spawning area. The source-sink model is a powerful means of detecting low levels of introgression over several generations.     

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 sea- ducks. 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.     

Straying of hatchery salmon in Prince William Sound,Alaska

The straying of hatchery salmon may harm wild salmon populations through a variety of ecological and genetic mechanisms. Surveys of pink (Oncorhynchus gorbuscha), chum (O. keta) and sockeye (O. nerka) salmon in wild salmon spawning locations in Prince William Sound (PWS), Alaska since 1997 show a wide range of hatchery straying. The analysis of thermally marked otoliths collected from carcasses indicate that 0–98% of pink salmon, 0–63% of chum salmon and 0–93% of sockeye salmon in spawning areas are hatchery fish, producing an unknown number of hatchery-wild hybrids. Most spawning locations sampled (77%) had hatchery pink salmon from three or more hatcheries, and 51% had annual escapements consisting of more than 10% hatchery pink salmon during at least one of the years surveyed. An exponential decay model of the percentage of hatchery pink salmon strays with distance from hatcheries indicated that streams throughout PWS contain more than 10% hatchery pink salmon. The prevalence of hatchery pink salmon strays in streams increased throughout the spawning season, while the prevalence of hatchery chum salmon decreased. The level of hatchery salmon strays in many areas of PWS are beyond all proposed thresholds (2–10%), which confounds wild salmon escapement goals and may harm the productivity, genetic diversity and fitness of wild salmon in this region     

Polar ecosystem dynamics: recovery of communities from organic enrichment in McMurdo Sound, Antarctica

Community structure and diversity are influenced by patterns of disturbance and input of food. In Antarctica, the marine ecosystem undergoes highly seasonal changes in availability of light and in primary production. Near research stations, organic input from human activities can disturb the regular productivity regime with a consistent input of sewage. McMurdo Sound has both high-productivity and low-productivity habitats, thereby providing an ideal test bed for community recovery dynamics under polar conditions. We used experimental manipulations of the subtidal communities to test the hypotheses that (1) benthic communities respond differently to disturbance from organic enrichment versus burial and (2) community response also varies in areas with different natural patterns of food supply. Both in low- and high-food habitats, the strongest community response was to organic enrichment and resulted in dominance of typical organic-enrichment specialists. In habitats with highly seasonal productivity, community response was predictable and recovery was rapid. In habitats with low productivity, community variability was high and caging treatments suggested that inconsistencies were due to patchy impacts by scavengers. In areas normally subject to regular organic enrichment, either from primary production or from further up the food web (defecation by marine mammals), recovery of benthic communities takes only years even in a polar system. However, a low-productivity regime is as common in near shore habitats around the continent; under these conditions, recovery of benthic communities from disturbance is likely to be much slower and follow a variable ecological trajectory.     

Evidence for competition at sea between Norton Sound chum salmon and Asian hatchery chum salmon

Increasing production of hatchery salmon over the past four decades has led to concerns about possible density-dependent effects on wild Pacific salmon populations in the North Pacific Ocean. The concern arises because salmon from distant regions overlap in the ocean, and wild salmon populations having low productivity may compete for food with abundant hatchery populations. We tested the hypothesis that adult length-at-age, age-at-maturation, productivity, and abundance of a Norton Sound, Alaska, chum salmon population were influenced by Asian hatchery chum salmon, which have become exceptionally abundant and surpassed the abundance of wild chum salmon in the North Pacific beginning in the early 1980s. We found that smaller adult length-at-age, delayed age-at-maturation, and reduced productivity and abundance of the Norton Sound salmon population were associated with greater production of Asian hatchery chum salmon since 1965. Modeling of the density-dependent relationship, while controlling for other influential variables, indicated that an increase in adult hatchery chum salmon abundance from 10 million to 80 million adult fish led to a 72% reduction in the abundance of the wild chum salmon population. These findings indicate that competition with hatchery chum salmon contributed to the low productivity and abundance of Norton Sound chum salmon, which includes several stocks that are classified as Stocks of Concern by the State of Alaska. This study provides new evidence indicating that large-scale hatchery production may influence body size, age-at-maturation, productivity and abundance of a distant wild salmon population.     

The Influence of Climate, Soils, Weather, and Land Use on Primary Production and Biomass Seasonality in the US Great Plains

Identifying the conditions and mechanisms that control ecosystem processes, such as net primary production, is a central goal of ecosystem ecology. Ideas have ranged from single limiting-resource theories to colimitation by nutrients and climate, to simulation models with edaphic, climatic, and competitive controls. Although some investigators have begun to consider the influence of land-use practices, especially cropping, few studies have quantified the impact of cropping at large scales relative to other known controls over ecosystem processes. We used a 9-year record of productivity, biomass seasonality, climate, weather, soil conditions, and cropping in the US Great Plains to quantify the controls over spatial and temporal patterns of net primary production and to estimate sensitivity to specific driving variables. We considered climate, soil conditions, and long-term average cropping as controls over spatial patterns, while weather and interannual cropping variations were used as controls over temporal variability. We found that variation in primary production is primarily spatial, whereas variation in seasonality is more evenly split between spatial and temporal components. Our statistical (multiple linear regression) models explained more of the variation in the amount of primary production than in its seasonality, and more of the spatial than the temporal patterns. Our results indicate that although climate is the most important variable for explaining spatial patterns, cropping explains a substantial amount of the residual variability. Soil texture and depth contributed very little to our models of spatial variability. Weather and cropping deviation both made modest contributions to the models of temporal variability. These results suggest that the controls over seasonality and temporal variation are not well understood. Our sensitivity analysis indicates that production is more sensitive to climate than to weather and that it is very sensitive to cropping intensity. In addition to identifying potential gaps in out knowledge, these results provide insight into the probable long- and short-term ecosystem response to changes in climate, weather, and cropping.     

Concordant population dynamics of Lepidoptera herbivores in a forest ecosystem

It is frequently assumed that population fluctuations are largely independent within a community of trophically‐similar species, but this need not be so. If population fluctuations are partly synchronized or concordant, this will produce interannual variability in the community's aggregate abundance and generate temporal variance in ecosystem structure. We studied the community of Lepidoptera inhabiting northern hardwood forests in New Hampshire, USA, to evaluate the hypothesis that fluctuations in consumer communities can arise from concordant dynamics of constituent populations. Interannual comparisons of moth abundances for >75 species sampled at three sites over four years revealed that concordant dynamics contribute strongly to interannual variability in the abundance of consumers. A conspicuous decline in community abundance from 2004 to 2005 was the result of predominantly negative population growth rates of the component species, while an increase in community abundance from 2006 to 2007 was the result of predominantly positive population growth rates. Population dynamics most strongly linked species that feed in the early season (perhaps due to shared responses to climatic effects), but not species that might share natural enemies or host plants. The observed concordant dynamics introduced conspicuous temporal variation in the abundance of primary consumers relative to plants and secondary consumers, thereby altering the forest's trophic structure. Such variance in the aggregate abundance of forest primary consumers could generate time‐lagged fluctuations in abundances of secondary consumers and will generally have important consequences for ecosystem properties and processes that are nonlinear functions of consumer abundance, such as plant community structure and nutrient cycling.     

Bottom‐up processes drive reproductive success in an apex predator

One of the central goals of the field of population ecology is to identify the drivers of population dynamics, particularly in the context of predator–prey relationships. Understanding the relative role of top‐down versus bottom‐up drivers is of particular interest in understanding ecosystem dynamics. Our goal was to explore predator–prey relationships in a boreal ecosystem in interior Alaska through the use of multispecies long‐term monitoring data. We used 29 years of field data and a dynamic multistate site occupancy modeling approach to explore the trophic relationships between an apex predator, the golden eagle, and cyclic populations of the two primary prey species available to eagles early in the breeding season, snowshoe hare and willow ptarmigan. We found that golden eagle reproductive success was reliant on prey numbers, but also responded prior to changes in the phase of the snowshoe hare population cycle and failed to respond to variation in hare cycle amplitude. There was no lagged response to ptarmigan populations, and ptarmigan populations recovered quickly from the low phase. Together, these results suggested that eagle reproduction is largely driven by bottom‐up processes, with little evidence of top‐down control of either ptarmigan or hare populations. Although the relationship between golden eagle reproductive success and prey abundance had been previously established, here we established prey populations are likely driving eagle dynamics through bottom‐up processes. The key to this insight was our focus on golden eagle reproductive parameters rather than overall abundance. Although our inference is limited to the golden eagle–hare–ptarmigan relationships we studied, our results suggest caution in interpreting predator–prey abundance patterns among other species as strong evidence for top‐down control.     

Molecular status of the dusky Canada goose (Branta canadensis occidentalis): A genetic assessment of a translocation effort

Until recently, the dusky Canada goose (Branta canadensis occidentalis) was managedas one breeding population from the CopperRiver Delta (CRD), Alaska. Population numberson the CRD have declined precipitously over thelast three decades, due in part to changes inhabitat. In 1981, a pair of Canada geese,presumably B.c. occidentalis, wasreported nesting on Middleton Island (MID), inthe Gulf of Alaska. Numbers of Canada geese onthe island increased in the decade subsequentto a translocation of geese from CRD to MID,but it is unclear whether the increase isattributable to the translocation effort. Weused genetic data derived from three classes ofgenetic markers to clarify relationships ofCanada geese breeding in south-coastal Alaska. Geese were sampled from 5 populations: CRD,MID, Anchorage (ANC), Admiralty Island (ADM) insoutheastern Alaska, and Green Island (GRN) inPrince William Sound (PWS). Mitochondrial DNAanalyses demonstrate Canada geese from MID arenearly monomorphic for a unique haplotype fixedon GRN but not found in CRD or any otherbreeding population. Furthermore, nuclearmarkers consistently cluster MID with GRN tothe exclusion of CRD. We suggest the currentpopulation on MID is not derived from birdstranslocated from CRD, but rather that MID wasmost likely colonised by birds inhabiting otherisland habitats within the PWS. Furthermore,since geese from the CRD share mtDNA haplotypeswith geese from other breeding locales, theyapparently share recent common ancestry and/orgene flow with populations representing othersubspecies. Our genetic data raise questionsabout the validity of current management unitsof Canada geese.     

Revealing environmentally driven population dynamics of an Arctic diatom using a novel microsatellite PoolSeq barcoding approach

Ecological stability under environmental change is determined by both interspecific and intraspecific processes. Particularly for planktonic microorganisms, it is challenging to follow intraspecific dynamics over space and time. We propose a new method, microsatellite PoolSeq barcoding (MPB), for tracing allele frequency changes in protist populations. We successfully applied this method to experimental community incubations and field samples of the diatom Thalassiosira hyalina from the Arctic, a rapidly changing ecosystem. Validation of the method found compelling accuracy in comparison with established genotyping approaches within different diversity contexts. In experimental and environmental samples, we show that MPB can detect meaningful patterns of population dynamics, resolving allelic stability and shifts within a key diatom species in response to experimental treatments as well as different bloom phases and years. Through our novel MPB approach, we produced a large dataset of populations at different time-points and locations with comparably little effort. Results like this can add insights into the roles of selection and plasticity in natural protist populations under stable experimental but also variable field conditions. Especially for organisms where genotype sampling remains challenging, MPB holds great potential to efficiently resolve eco-evolutionary dynamics and to assess the mechanisms and limits of resilience to environmental stressors.     

The evolution of ecosystem processes: growth rate and elemental stoichiometry of a key herbivore in temperate and arctic habitats

Understanding the reciprocal interactions between the evolved characteristics of species and the environment in which each species is embedded is a major priority for evolutionary ecology. Here we use the perspective of ecological stoichiometry to test the hypothesis that natural selection on body growth rate affects consumer body stoichiometry. As body elemental composition (nitrogen, phosphorus) of consumers influences nutrient cycling and trophic dynamics in food webs, such differences should also affect biogeochemical processes and trophic dynamics. Consistent with the growth rate hypothesis, body growth rate and phosphorus content of individuals of the Daphnia pulex species complex were lower in Wisconsin compared to Alaska, where the brevity of the growing season places a premium on growth rate. Consistent with stoichiometric theory, we also show that, relative to animals sampled in Wisconsin, animals sampled in Alaska were poor recyclers of P and suffered greater declines in growth when fed low‐quality, P‐deficient food. These results highlight the importance of evolutionary context in establishing the reciprocal relationships between single species and ecosystem processes such as trophic dynamics and consumer‐driven nutrient recycling.     

Regional processes in mangrove ecosystems: spatial scaling relationships, biomass, and turnover rates following catastrophic disturbance

Physiological processes and local-scale structural dynamics of mangroves are relatively well studied. Regional-scale processes, however, are not as well understood. Here we provide long-term data on trends in structure and forest turnover at a large scale, following hurricane damage in mangrove ecosystems of South Florida, U.S.A. Twelve mangrove vegetation plots were monitored at periodic intervals, between October 1992 and March 2005. Mangrove forests of this region are defined by a −1.5 scaling relationship between mean stem diameter and stem density, mirroring self-thinning theory for mono-specific stands. This relationship is reflected in tree size frequency scaling exponents which, through time, have exhibited trends toward a community average that is indicative of full spatial resource utilization. These trends, together with an asymptotic standing biomass accumulation, indicate that coastal mangrove ecosystems do adhere to size-structured organizing principles as described for upland tree communities. Regenerative dynamics are different between areas inside and outside of the primary wind-path of Hurricane Andrew which occurred in 1992. Forest dynamic turnover rates, however, are steady through time. This suggests that ecological, more-so than structural factors, control forest productivity. In agreement, the relative mean rate of biomass growth exhibits an inverse relationship with the seasonal range of porewater salinities. The ecosystem average in forest scaling relationships may provide a useful investigative tool of mangrove community biomass relationships, as well as offer a robust indicator of general ecosystem health for use in mangrove forest ecosystem management and restoration.     

The importance of neutral over niche processes in structuring Ediacaran early animal communities

The relative influence of niche vs. neutral processes in ecosystem dynamics is an on‐going debate, but the extent to which they structured the earliest animal communities is unknown. Some of the oldest known metazoan‐dominated paleocommunities occur in Ediacaran age (~ 565 million years old) strata in Newfoundland, Canada and Charnwood Forest, UK. These comprise large and diverse populations of sessile organisms that are amenable to spatial point process analyses, enabling inference of the most likely underlying niche or neutral processes governing community structure. We mapped seven Ediacaran paleocommunities using LiDAR, photogrammetry and a laser line probe. We found that neutral processes dominate these paleocommunities, with niche processes exerting limited influence, in contrast with the niche‐dominated dynamics of modern marine ecosystems. The dominance of neutral processes suggests that early metazoan diversification may not have been driven by systematic adaptations to the local environment, but instead may have resulted from stochastic demographic differences.     

Population density of North American elk: effects on plant diversity

Large, herbivorous mammals have profound effects on ecosystem structure and function and often act as keystone species in ecosystems they inhabit. Density-dependent processes associated with population structure of large mammals may interact with ecosystem functioning to increase or decrease biodiversity, depending on the relationship of herbivore populations relative to the carrying capacity (K) of the ecosystem. We tested for indirect effects of population density of large herbivores on plant species richness and diversity in a montane ecosystem, where increased net aboveground primary productivity (NAPP) in response to low levels of herbivory has been reported. We documented a positive, linear relationship between plant-species diversity and richness with NAPP. Structural equation modeling revealed significant indirect relationships between population density of herbivores, NAPP, and species diversity. We observed an indirect effect of density-dependent processes in large, herbivorous mammals and species diversity of plants through changes in NAPP in this montane ecosystem. Changes in species diversity of plants in response to herbivory may be more indirect in ecosystems with long histories of herbivory. Those subtle or indirect effects of herbivory may have strong effects on ecosystem functioning, but may be overlooked in plant communities that are relatively resilient to herbivory.