New hypothesis helps explain elasmobranch "outburst" on Georges Bank in the 1980s.

Regime shifts are a feature of many ecosystems. During the last 40 years, intensive commercial exploitation and environmental changes have driven substantial shifts in ecosystem structure and function in the northwest Atlantic. In the Georges Bank-southern New England region, commercially important species have declined, and the ecosystem shifted to one dominated by economically undesirable species such as skates and dogfish. Aggregated abundance indices indicate a large increase of small and medium-sized elasmobranchs in the early 1980s following the decline of many commercial species. It has been hypothesized that ecological interactions such as competition and predation within the Georges Bank region were responsible for and are maintaining the "elasmobranch outburst" at the heart of the observed ecosystem shift. We offer an alternative hypothesis invoking population connectivity among winter skate populations such that the observed abundance increase is a result of migratory dynamics, perhaps with the Scotian Shelf (i.e., it is an open population). Here we critically evaluate the survey data for winter skate, the species principally responsible for the increase in total skate abundance during the 1980s on Georges Bank, to assess support for both hypotheses. We show that time series from different surveys within the Georges Bank region exhibit low coherence, indicating that a widespread population increase was not consistently shown by all surveys. Further, we argue that observed length-frequency data for Georges Bank indicate biologically unrealistic population fluctuations if the population is closed. Neither finding supports the elasmobranch outburst hypothesis. In contrast, survey time series for Georges Bank and the Scotian Shelf are negatively correlated, in support of the population connectivity hypothesis. Further, we argue that understanding the mechanisms of ecosystem state changes and population connectivity are needed to make inferences about both the causes and appropriate management responses to large-scale system change.     

Fish production and the marine ecosystems of the Scotian Shelf,eastern Canada

Fishery production and food webs have been studied on the Scotian Shelf and upper continental slope along a transect running 270 km SE of Halifax, Canada. The area (TCNAF Division 4W) supports a fishery of roughly 0.15x10⁶ metric tons. Overall primary production of the shelf waters is 102 g C m^-2 year^-1 and of the slope wacers about 128 g C m^-2 year^-1. Demersal fish production (average 4.1 kcal m^-2 year^-1) is highest over the shelf and declines in an offshore direction, while pelagic fish production (average 16 kcal m^-2 year^-1) is highest over the slope and declines in an inshore direction. Hypothetical food webs of these two intergrading ecosystems have been constructed, based on data for primary production, fish catches, and the biomasses of zooplankton and macrobenthos. These lead us to suggest that there are basic differences in food chains and efficiencies between the two ecosystems that account for their differences in production. Although primary production is 17% higher on an average on the Nova Scotian transect than in the North Sea, the apparent zooplankton and macrobenthos production is 31% lower and macrobenthos production may also be lower. Overall fish catch from the Scotian Shelf and slope is about 47% lower per unit area than the catch in the North Sea, despite the fact that the demersal catches are identical. This is accounted for by a much lower overall pelagic catch from the Nova Scotian area, centered in a region that is small compared to the total area. Fish production in different regions cannot be predicted merely on the basis of differences in level of primary production, but must take into account differences in the structure of the ecosystems.     

Advancing objectives-based,integrated ocean management through marine spatial planning: current and future directions on the Scotian Shelf off Nova Scotia,Canada

The national legislative and policy context for integrated ocean management in Canada is provided by the Oceans Act (1996) and the supporting policy statement, Canada’s Oceans Strategy. Under the Oceans Act, Fisheries and Oceans Canada (DFO) is the lead federal authority for ocean affairs and is charged with leading and facilitating the development and implementation of integrated management plans for all marine waters. Integrated management efforts in Canada are being undertaken through an area-based approach that enables marine planning, management and decision making to occur at appropriate spatial scales, from regional to site-specific. This article focuses on the Eastern Scotian Shelf Integrated Management (ESSIM) process, an offshore-focused effort to develop an integrated ocean management plan for a large portion of the Scotian Shelf, off Nova Scotia. The resulting Eastern Scotian Shelf Integrated Ocean Management Plan (the ESSIM plan) has been developed through a collaborative process involving all interested and affected government departments and ocean stakeholders, and provides an objectives-based approach to ocean management. The ESSIM plan contains a set of long-term, overarching goals for collaborative governance and integrated management, sustainable human use, and healthy ecosystems. These goals are supported by more specific objectives that express desired outcomes and conditions for the marine region. The objectives-based approach seeks to ensure that interrelationships among ecosystem and human use objectives are recognized and reflected in the identification of management strategies and supporting actions. This article considers the role of marine spatial planning within the context of the integrated ocean management process underway for the Scotian Shelf. The policy and management context for integrated ocean management in Canada is briefly described and a summary of the ESSIM plan is provided. The current and potential role for marine spatial planning in implementing the objectives and strategies of the ESSIM plan is highlighted using examples related to multiple ocean use and marine conservation and protected area planning. The article concludes by drawing out key lessons learned to date through the ESSIM process for marine spatial planning and looks to the future in terms of the development of tools and approaches for this integral aspect of integrated ocean management.     

Multistate modeling of habitat dynamics: factors affecting Florida scrub transition probabilities

Many ecosystems are influenced by disturbances that create specific successional states and habitat structures that species need to persist. Estimating transition probabilities between habitat states and modeling the factors that influence such transitions have many applications for investigating and managing disturbance-prone ecosystems. We identify the correspondence between multistate capture-recapture models and Markov models of habitat dynamics. We exploit this correspondence by fitting and comparing competing models of different ecological covariates affecting habitat transition probabilities in Florida scrub and flatwoods, a habitat important to many unique plants and animals. We subdivided a large scrub and flatwoods ecosystem along central Florida's Atlantic coast into 10-ha grid cells, which approximated average territory size of the threatened Florida Scrub-Jay (Aphelocoma coerulescens), a management indicator species. We used 1.0-m resolution aerial imagery for 1994, 1999, and 2004 to classify grid cells into four habitat quality states that were directly related to Florida Scrub-Jay source-sink dynamics and management decision making. Results showed that static site features related to fire propagation (vegetation type, edges) and temporally varying disturbances (fires, mechanical cutting) best explained transition probabilities. Results indicated that much of the scrub and flatwoods ecosystem was resistant to moving from a degraded state to a desired state without mechanical cutting, an expensive restoration tool. We used habitat models parameterized with the estimated transition probabilities to investigate the consequences of alternative management scenarios on future habitat dynamics. We recommend this multistate modeling approach as being broadly applicable for studying ecosystem, land cover, or habitat dynamics. The approach provides maximum-likelihood estimates of transition parameters, including precision measures, and can be used to assess evidence among competing ecological models that describe system dynamics.     

Fish reintroductions reveal smooth transitions between lake community states

Whether communities respond smoothly or discontinuously to changing environmental conditions has important consequences for the preservation and restoration of ecosystems. Theory shows that communities may exhibit a variety of responses to environmental change, including abrupt transitions due to the existence of alternate states. However, there have been few opportunities to look for such transitions in nature. Here, we examine the impact of a two-orders-of-magnitude decrease and then increase in planktivore abundance in Wintergreen lake (Michigan, USA), caused by the extinction and reintroduction of two dominant fish species (largemouth bass, Micropterus salmoides, and bluegill, Lepomis macrochirus). Over a 16 + yr period of slow change from high planktivory to low planktivory back to high planktivory, the zooplankton community changed smoothly and predictably between states. In years of low planktivory, the zooplankton assemblage was dominated by a single, large, cladoceran species, Daphnia pulicaria, whereas in years of high planktivory, D. pulicaria disappeared and was replaced by a suite of small-bodied cladocerans. We quantified the multivariate change in zooplankton community dissimilarity and found that community state smoothly tracked changes in planktivore density in both a forward and backward direction. Thus, there was little evidence of discontinuity in this system where transitions are strongly driven by planktivory.     

Effects of temperature on morphological landmarks critical to growth and survival in larval Atlantic cod (Gadus morhua)

The morphology and function of structures important to energy acquisition were studied from spawning to the stage of transformation of larva to pelagic juvenile in Atlantic cod, Gadus morhua L., from December 1991 to July 1992. Fertilized eggs produced by adult fish from two genetically discrete populations (Newfoundland and Scotian Shelf) were raised under similar conditions in the laboratory at temperatures of 5 and 10°C. Subsamples of larvae were removed from cultures daily for 10 d, and then less frequently, and fixed for light microscopy and scanning electron microscopy. Nine functional morphological landmarks important to feeding, respiration and locomotion were chosen from observation of 280 ind. These landmarks defined 12 major developmental stages, from hatching to the pelagic juvenile stage. One of the feeding landmarks, intestinal stage, varied as a function of age and size and the variance in development was higher at 10°C than at 5°C; Newfoundland larvae developed more complex intestines than did Scotian Shelf larvae. In addition, Newfoundland larvae had significantly higher growth rates than those of Scotian Shelf larvae. Despite the higher growth rates and greater structural complexity of the intestine in Newfoundland larvae, the rate of yolk utilization was not significantly different between Newfoundland and Scotian Shelf larvae. Staging of respiratory landmarks showed that the gill arches were probably used preferentially in feeding while respiration was cutaneous. The gills, operculum and gill rakers developed late in larval life and accompanied the transition from cutaneous to branchial respiration. In the yolk-sac period, development of feeding and respiratory structures may be largely genetically controlled. During exogenous feeding, extrinsic factors also become important, as shown by the size and age-independent variation in intestinal development of larval cod raised at different temperatures.     

Indicators of ecosystem function identify alternate states in the sagebrush steppe

Models of ecosystem change that incorporate nonlinear dynamics and thresholds, such as state-and-transition models (STMs), are increasingly popular tools for land management decision-making. However, few models are based on systematic collection and documentation of ecological data, and of these, most rely solely on structural indicators (species composition) to identify states and transitions. As STMs are adopted as an assessment framework throughout the United States, finding effective and efficient ways to create data-driven models that integrate ecosystem function and structure is vital. This study aims to (1) evaluate the utility of functional indicators (indicators of rangeland health, IRH) as proxies for more difficult ecosystem function measurements and (2) create a data-driven STM for the sagebrush steppe of Colorado, USA, that incorporates both ecosystem structure and function. We sampled soils, plant communities, and IRH at 41 plots with similar clayey soils but different site histories to identify potential states and infer the effects of management practices and disturbances on transitions. We found that many IRH were correlated with quantitative measures of functional indicators, suggesting that the IRH can be used to approximate ecosystem function. In addition to a reference state that functions as expected for this soil type, we identified four biotically and functionally distinct potential states, consistent with the theoretical concept of alternate states. Three potential states were related to management practices (chemical and mechanical shrub treatments and seeding history) while one was related only to ecosystem processes (erosion). IRH and potential states were also related to environmental variation (slope, soil texture), suggesting that there are environmental factors within areas with similar soils that affect ecosystem dynamics and should be noted within STMs. Our approach generated an objective, data-driven model of ecosystem dynamics for rangeland management. Our findings suggest that the IRH approximate ecosystem processes and can distinguish between alternate states and communities and identify transitions when building data-driven STMs. Functional indicators are a simple, efficient way to create data-driven models that are consistent with alternate state theory. Managers can use them to improve current model-building methods and thus apply state-and-transition models more broadly for land management decision-making.     

Marine Protected Area Design and the Spatial and Temporal Distribution of Cetaceans in a Submarine Canyon

Abstract: The Gully, the largest submarine canyon off the coast of eastern Canada, is currently under consideration as a marine conservation area, primarily because of the increasing interest in oil and gas production on the Scotian Shelf. Cetaceans, as a guild of abundant, large organisms that are relatively sensitive to such threats, provide a reliable means to determine the boundaries for a conservation area in this region. We compared the abundance of cetaceans between the Gully and other parts of the Scotian Shelf and Slope and found that abundance was higher in the Gully. We also assessed cetacean distribution and relative abundance within the Gully relative to search effort for several spatial and temporal parameters: depth, slope, sea surface temperature, and month. Distribution within the Gully was most strongly correlated with depth, but was also significantly correlated with sea surface temperature and month. Five of the 11 cetacean species commonly found in the Gully, and all those for which the Gully formed significant habitat on the Scotian Shelf, were concentrated in the deep (200–2000 m) mouth of the canyon. We suggest that a year-round marine protected area is necessary for the Gully. A core protection zone should be defined in the Gully based on depth and bounded by the 200-m isobath. A buffer zone around the core zone should be defined to provide protection from activities with further-reaching effects, such as noise, dredging, and chemical pollution.     

Robustness of variance and autocorrelation as indicators of critical slowing down

Ecosystems close to a critical threshold lose resilience, in the sense that perturbations can more easily push them into an alternative state. Recently, it has been proposed that such loss of resilience may be detected from elevated autocorrelation and variance in the fluctuations of the state of an ecosystem due to critical slowing down; the underlying generic phenomenon that occurs at critical thresholds. Here we explore the robustness of autocorrelation and variance as indicators of imminent critical transitions. We show both analytically and in simulations that variance may sometimes decrease close to a transition. This can happen when environmental factors fluctuate stochastically and the ecosystem becomes less sensitive to these factors near the threshold, or when critical slowing down reduces the ecosystem's capacity to follow high-frequency fluctuations in the environment. In addition, when available data is limited, variance can be systematically underestimated due to the prevalence of low frequencies close to a transition. By contrast, autocorrelation always increases toward critical transitions in our analyses. To exemplify this point, we provide cases of rising autocorrelation and increasing or decreasing variance in time series prior to past climate transitions.     

Movements,environmental associations,and presumed spawning locations of Atlantic halibut (Hippoglossus hippoglossus) in the northwest Atlantic determined using archival satellite pop-up tags

Large Atlantic halibut (Hippoglossus hippoglossus) off the eastern coast of Canada were tagged with pop-up satellite archival transmission tags (N = 17) to track movements, determine ambient depth and temperature, and infer spawning activity. Many halibut showed seasonal movements from deepwater slope areas in fall and winter to shallower feeding grounds on the Scotian Shelf and Grand Banks in summer. Halibut depths ranged between 0 and 1,640 m. Mean temperature of occupation was 4.7 °C. Multiple short-term vertical ascents from a consistent baseline depth, characterized as spawning rises, were identified in seven of the tagged halibut south of the Grand Banks. All presumed spawning rises occurred in multiples of 2–6 events at 2- to 9-day intervals between October and January, spanning an average vertical extent of 50–100 m at depths of about 800–1,000 m. Given the direction and velocity of the slope water currents and the duration of the pelagic stage, the calculated 300–500 km drift of the eggs and larvae would take them onto the Scotian Shelf, as well as into the Gulf of St. Lawrence. Therefore, the location of the presumed spawning grounds is consistent with expectations based on migration compensation theory, the northeasterly migratory patterns of the juveniles, the relatively static distribution of the adults off southern Newfoundland, and the prevailing currents at depth.     

Ecosystem states: Creating a data-derived, ecosystem-scale ecological response model that is explicit in space and time

Increasing difficulties associated with balancing consumptive demands for water and achieving ecological benefits in aquatic ecosystems provide opportunities for new ecosystem-scale ecological response models to assist managers. Using an Australian estuary as a case study, we developed a novel approach to create a data-derived state-and-transition model. The model identifies suites of co-occurring birds, fish, benthic invertebrates and aquatic macrophytes (as ‘states’) and the changing physico-chemical conditions that are associated with each (‘transitions’). The approach first used cluster analysis to identify sets of co-occurring biota. Differences in the physico-chemical data associated with each state were identified using classification trees, with the biotic distinctness of the resultant statistical model tested using analysis of similarities. The predictive capacity of the model was tested using new cases. Two models were created using different time-steps (annual and quarterly) and then combined to capture both longer-term trends and more-recent declines in ecological condition. We identified eight ecosystem states that were differentiated by a mix of water-quantity and water-quality variables. Each ecosystem state represented a distinct biotic assemblage under well-defined physico-chemical conditions. Two ‘basins of attraction’ were identified, with four tidally-influenced states, and another four independent of tidal influence. Within each basin, states described a continuum of relative health, manifest through declining taxonomic diversity and abundances. The main threshold determining relative health was whether freshwater flows had occurred in the region during the previous 339 days. Canonical analyses of principal coordinates tested the predictive capacity of the model and demonstrated that the variance in the environmental data set was well captured (87%) with 52% of the variance in the biological data set also captured. The latter increased to >80% when long- and short-term biological data were analysed separately, indicating that the model described the available data for the Coorong well. This approach thus created a data-derived, multivariate model, where neither states nor transitions were determined a priori. The approach did not over-fit the data, was robust to patchy or missing data, the choice of initial clustering technique and random errors in the biological data set, and was well-received by local natural resource managers. However, the model did not capture causal relationships and requires additional testing, particularly during future episodes of ecological recovery. The approach shows significant promise for simplifying management definitions of ecological condition and, via scenario analyses, can be used to assist in manager decision-making of large, complex aquatic ecosystems in the future.     

Plague bacterium as a transformer species in prairie dogs and the grasslands of western North America

Invasive transformer species change the character, condition, form, or nature of ecosystems and deserve considerable attention from conservation scientists. We applied the transformer species concept to the plague bacterium Yersinia pestis in western North America, where the pathogen was introduced around 1900. Y. pestis transforms grassland ecosystems by severely depleting the abundance of prairie dogs (Cynomys spp.) and thereby causing declines in native species abundance and diversity, including threatened and endangered species; altering food web connections; altering the import and export of nutrients; causing a loss of ecosystem resilience to encroaching invasive plants; and modifying prairie dog burrows. Y. pestis poses an important challenge to conservation biologists because it causes trophic‐level perturbations that affect the stability of ecosystems. Unfortunately, understanding of the effects of Y. pestis on ecosystems is rudimentary, highlighting an acute need for continued research.     

A stochastic weather generation model for Alaska

A specific problem encountered in ecosystem-level simulation of Arctic ecosystems is the depth and extent of the driving variable record. Often, climate records are of short duration, gathered at locations different from the area to be simulated, or do not contain all the variables required by a given model. This paper addresses this problem for ecosystem simulation in Alaska with the development of a weather generator. The generator, called WGENAL, is based on the WGEN climate generator developed and validated in the 48 conterminous states. Because of the extreme variability of weather in Alaska that is not accommodated by the statistical metrics in the earlier model, a new climate generator was developed. WGENAL generates daily values of precipitation, maximum temperature, minimum temperature, solar radiation, and wind run. Precipitation is generated using a Markov chain-gamma model. A two-parameter gamma distribution is used to generate wind run. Temperatures and solar radiation are generated using procedures developed in the earlier study. Validation of the generator shows it provides adequate diurnal and seasonal weather records for Fairbanks. Other comparisons of synthetic weather with observed weather for sites north of the Brooks Range in Alaska are also within the error of the original data.     

Age-structured modeling reveals long-term declines in the natality of western Steller sea lions.

Since the mid-1970s, the western Steller sea lion (Eumetopias jubatus), inhabiting Alaskan waters from Prince William Sound west through the Aleutian Islands, has declined by over 80%. Changing oceanographic conditions, competition from fishing operations, direct human-related mortality, and predators have been suggested as factors driving the decline, but the indirect and interactive nature of their effects on sea lions have made it difficult to attribute changes in abundance to specific factors. In part, this is because only changes in abundance, not changes in vital rates, are known. To determine how vital rates of the western Steller sea lion have changed during its 28-year decline, we first estimated the changes in Steller sea lion age structure using measurements of animals in aerial photographs taken during population surveys since 1985 in the central Gulf of Alaska (CGOA). We then fit an age-structured model with temporally varying vital rates to the age-structure data and to total population and pup counts. The model fits indicate that birth rate in the CGOA steadily declined from 1976 to 2004. Over the same period, survivorship first dropped severely in the early 1980s, when the population collapsed, and then survivorship steadily recovered. The best-fitting model indicates that in 2004, the birth rate in the central Gulf of Alaska was 36% lower than in the 1970s, while adult and juvenile survivorship were close to or slightly above 1970s levels. These predictions and other model predictions concerning population structure match independent field data from mark-recapture studies and photometric analyses. The dominant eigenvalue for the estimated 2004 Leslie matrix is 1.0014, indicating a stable population. The stability, however, depends on very high adult survival, and the shift in vital rates results in a population that is more sensitive to changes in adult survivorship. Although our modeling analysis focused exclusively on the central Gulf of Alaska, the western Gulf of Alaska and eastern Aleutians show a similar pattern of declining pup fraction with no increase in the juvenile, or pre-breeding, fraction. This suggests that declining birth rate may be a problem for western Steller sea lions across the Gulf of Alaska and into the Aleutian Islands.     

A 700-year paleoecological record of boreal ecosystem responses to climatic variation from Alaska

Recent observations and model simulations have highlighted the sensitivity of the forest-tundra ecotone to climatic forcing. In contrast, paleoecological studies have not provided evidence of tree-line fluctuations in response to Holocene climatic changes in Alaska, suggesting that the forest-tundra boundary in certain areas may be relatively stable at multicentennial to millennial time scales. We conducted a multiproxy study of sediment cores from an Alaskan lake near the altitudinal limits of key boreal-forest species. Paleoecological data were compared with independent climatic reconstructions to assess ecosystem responses of the forest tundra boundary to Little Ice Age (LIA) climatic fluctuations. Pollen, diatom, charcoal, macrofossil, and magnetic analyses provide the first continuous record of vegetation fire-climate interactions at decadal to centennial time scales during the past 700 years from southern Alaska. Boreal-forest diebacks characterized by declines of Picea mariana, P. glauca, and tree Betula occurred during the LIA (AD 1500-1800), whereas shrubs (Alnus viridis, Betula glandulosa/nana) and herbaceous taxa (Epilobium, Aconitum) expanded. Marked increases in charcoal abundance and changes in magnetic properties suggest increases in fire importance and soil erosion during the same period. In addition, the conspicuous reduction or disappearance of certain aquatic (e.g., Isoetes, Nuphar, Pediastrum) and wetland (Sphagnum) plants and major shifts in diatom assemblages suggest pronounced lake-level fluctuations and rapid ecosystem reorganization in response to LIA climatic deterioration. Our results imply that temperature shifts of 1-2 degrees C, when accompanied by major changes in moisture balance, can greatly alter high-altitudinal terrestrial, wetland, and aquatic ecosystems, including conversion between boreal-forest tree line and tundra. The climatic and ecosystem variations in our study area appear to be coherent with changes in solar irradiance, suggesting that changes in solar activity contributed to the environmental instability of the past 700 years.     

Multi-decadal drought and amplified moisture variability drove rapid forest community change in a humid region

Climate variability, particularly the frequency of extreme events, is likely to increase in the coming decades, with poorly understood consequences for terrestrial ecosystems. Hydroclimatic variations of the Medieval Climate Anomaly (MCA) provide a setting for studying ecological responses to recent climate variability at magnitudes and timescales comparable to expectations of coming centuries. We examined forest response to the MCA in the humid western Great Lakes region of North America, using proxy records of vegetation, fire, and hydroclimate. Multi-decadal moisture variability during the MCA was associated with a widespread, episodic decline in Fagus grandifolia (beech) populations. Spatial patterns of drought and forest changes were coherent, with beech declining only in areas where proxy-climate records indicate that severe MCA droughts occurred. The occurrence of widespread, drought-induced ecological changes in the Great Lakes region indicates that ecosystems in humid regions are vulnerable to rapid changes in drought magnitude and frequency.     

Interspecific variation in thermal denaturation of proteins in the congeneric musselsMytilus trossulus andM. galloprovincialis: evidence from the heat-shock response and protein ubiquitination

The genetic population structure of the precominant zooplankter, the copepod Calanus finmarchicus (Gunnerus), was examined to determine whether genetically distinct populations exist in the Gulf of Maine. C. finmarchicus was sampled in three regions of the Gulf of Maine (Great South Channel, spring 1989; northern Gulf of Maine, winter 1990; Great South Channel and Georges Bank, spring 1990). Copepods from seven locations in the Great South Channel, five in the northern Gulf of Maine and four on or near Georges Bank were assayed for allozyme variation and mitochondrial DNA variation of amplified 16S rRNA and cytochrome b genes. Restriction fragment length polymorphism (RFLP) analyses of both mitochondrial DNA genes revealed no variation among any of the individuals assayed. Analysis of five polymorphic allozyme loci revealed that genetic variation among the three geographic regions was low, and genetic identities were high between all locations (I>0.97). Most of the genetic variation was among locations regardless of region. Chi-square tests were used to examine genetic similarity between specific pairs of locations within and between regions. In the northern Gulf of Maine, genetic homogeneity occurred over larger spatial scales (hundreds of km) than in either the Great South Channel or Georges Bank (tens of km). Only copepods from the Bay of Fundy and Nova Scotian Shelf locations were genetically distinct from Wilkinson Basin copepods at two loci. Copepod populations from the northern locations may have been partially isolated or they may represent immigrant populations (e.g., from the Gulf of St. Lawrence). Several pairs of locations were genetically distinct at one or more loci in the two southern regions. Differences between locations in these regions may represent distinct populations advected into the areas at different times or from different sources (e.g., genetic variation may represent a mixture of genetically distinct northern and southern copepod populations). These results suggest extensive gene flow among populations of C. finmarchicus in the Gulf of Maine with some evidence of genetic population subdivision near the Gulf's northeastern and southern boundaries.     

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups. ‘Resident’-type (fish-eating) killer whales were more abundant than the ‘transient’-type (mammal-eating). Abundance estimates of resident killer whales (991 [95% CI = 379–2,585] [IGS] and 1,587 [95% CI = 608–4,140] [PEGS]), were at least four times greater than those of the transient killer whales (200 [95% CI = 81–488] [IGS] and 251 [95% CI = 97–644] whales [PEGS]). The IGS estimate of abundance is preferred for resident killer whales because the estimate based on PEGS data may show an upward bias. The PEGS estimate of abundance is likely more accurate for transients. Residents were most abundant near Kodiak Island in the northern Gulf of Alaska, around Umnak and Unalaska Islands in the eastern Aleutians, and in Seguam Pass in the central Aleutians. This ecotype was not observed between 156 and 164°W, south of the Alaska Peninsula. In contrast, transient killer whale sightings were found at higher densities south of the Alaska Peninsula between the Shumagin Islands and the eastern Aleutians. Only two sightings of ‘offshore’-type killer whales were recorded during the surveys, one northeast of Unalaska Island and the other south of Kodiak Island. These are the first estimates of abundance of killer whale ecotypes in the Aleutian Islands and Alaska Peninsula area and provide a baseline for quantifying the role of these top predators in their ecosystem. Electronic Supplementary Material  Supplementary material is available in the online version of this article at and is accessible for authorized users.

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Spatial and social connectivity of fish-eating “Resident” killer whales (Orcinus orca) in the northern North Pacific

The productive North Pacific waters of the Gulf of Alaska, Aleutian Islands and Bering Sea support a high density of fish-eating “Resident” type killer whales (Orcinus orca), which overlap in distribution with commercial fisheries, producing both direct and indirect interactions. To provide a spatial context for these interactions, we analyzed a 10-year dataset of 3,058 whale photo-identifications from 331 encounters within a large (linear ~4,000 km) coastal study area to investigate the ranging and social patterns of 532 individually identifiable whales photographed in more than one encounter. Although capable of large-scale movements (maximum 1,443 km), we documented ranges generally <200 km, with high site fidelity across summer sampling intervals and also re-sightings during a winter survey. Bayesian analysis of pair-wise associations identified four defined clusters, likely representing groupings of stable matrilines, with distinct ranging patterns, that combined to form a large network of associated whales that ranged across most of the study area. This provides evidence of structure within the Alaska stock of Resident killer whales, important for evaluating ecosystem and fisheries impacts. This network included whales known to depredate groundfish from longline fisheries, and we suggest that such large-scale connectivity has facilitated the spread of depredation.     

Hydroclimatic variability drives episodic expansion of a floating peat mat in a North American kettlehole basin

The coming century is predicted to feature enhanced climatic variability, including increased frequency, intensity, and duration of extreme climatic events. Ecologists are faced with the critical challenge of anticipating potentially nonlinear ecosystem responses to these changes. High-resolution paleoecological data sets that capture past ecosystem responses to climate variability provide valuable long-term perspectives on the sensitivity of ecosystems to climate-forced state shifts. We used a suite of paleoecological analyses at Titus Bog in northwestern Pennsylvania, USA, to test the hypothesis that the development and expansion of floating peatlands in kettlehole basins represents a threshold response to hydroclimate variability. In contrast with expectations of gradual autogenic peat mat expansion, our results indicate that peat mat expansion at Titus Bog was highly episodic and occurred in three distinct pulses centered on 800, 650, and 400 cal yr BP. Each of these expansion events coincided with or immediately followed decadal-to-mutlidecadal droughts recorded in regional paleoclimate reconstructions. These patterns indicate that peatland development in kettlehole basins can follow nonlinear trajectories, with episodes of rapid advancement triggered by climatic forcing. Future climate changes may increase the likelihood of peatland expansion in kettlehole basins, potentially leading to abrupt changes in adjacent lake ecosystems.