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John F. Piatt

Publications and source records attributed to John F. Piatt.

At least 73 records · Page 4Linked to original sources

Prey density and the behavioral flexibility of a marine predator: The common murre ( Uria aalge )

Flexible time budgets allow individual animals to buffer the effects of variable food availability by allocating more time to foraging when food density decreases. This trait should be especially important for marine predators that forage on patchy and ephemeral food resources. We examined flexible time allocation by a long-lived marine predator, the Common Murre (Uria aalge), using data collected in a five-year study at three colonies in Alaska (USA) with contrasting environmental conditions. Annual hydroacoustic surveys revealed an order-of-magnitude variation in food density among the 15 colony-years of study. We used data on parental time budgets and local prey density to test predictions from two hypotheses: Hypothesis A, the colony attendance of seabirds varies nonlinearly with food density; and Hypothesis B, flexible time allocation of parent murres buffers chicks against variable food availability. Hypothesis A was supported; colony attendance by murres was positively correlated with food over a limited range of poor-to-moderate food densities, but independent of food over a broader range of higher densities. This is the first empirical evidence for a nonlinear response of a marine predator's time budget to changes in prey density. Predictions from Hypothesis B were largely supported: (1) chick-feeding rates were fairly constant over a wide range of densities and only dropped below 3.5 meals per day at the low end of prey density, and (2) there was a nonlinear relationship between chick-feeding rates and time spent at the colony, with chick-feeding rates only declining after time at the colony by the nonbrooding parent was reduced to a minimum. The ability of parents to adjust their foraging time by more than 2 h/d explains why they were able to maintain chick-feeding rates of more than 3.5 meals/d across a 10-fold range in local food density. ?? 2007 by the Ecological Society of America.

Ecology

Status review of the Marbled Murrelet (Brachyramphus marmoratus) in Alaska and British Columbia

The Marbled Murrelet ( Brachyramphus marmoratus ) is a small, diving seabird inhabiting inshore waters of the Northeastern Pacific Ocean. This species feeds on small, schooling fishes and zooplankton, and nests primarily on the moss-covered branches of large, old-growth conifers, and also, in some parts of its range, on the ground. We reviewed existing information on this species to evaluate its current status in the northern part of its range—Alaska (U.S.) and British Columbia (Canada). Within the southern part of its range (Washington, Oregon, and California, U.S.), the Marbled Murrelet was listed as a threatened species under the Endangered Species Act (ESA) in 1993, and the U.S. Fish and Wildlife Service (USFWS) needed information on the species throughout its range for ESA deliberations. We compiled published information on the conservation status, population biology, foraging ecology, population genetics, population status and trends, demography, marine and nesting habitat characteristics, threats, and ongoing conservation efforts for Marbled Murrelets in Alaska and British Columbia. We conducted a new genetic study using samples from a segment of the range that had not been included in previous studies (Washington, Oregon) and additional nuclear intron and microsatellite markers. We also analyzed available at-sea survey data from several locations for trend. To understand the reasonableness of the empirical trend data, we developed demographic models incorporating stochasticity to discern what population trends were possible by chance. The genetic studies substantially confirmed previous findings on population structure in the Marbled Murrelet. Our present work finds three populations: (1) one comprising birds in the central and western Aleutian Islands; (2) one comprising birds in central California; and (3) one comprising birds within the center of the range from the eastern Aleutians to northern California. Our knowledge of genetic structure within this central population is limited and it requires additional study. Compiling available abundance information, we estimated that in the recent past, Marbled Murrelets in Alaska numbered on the order of 1 million birds. We were unable to generate a similar estimate for historical population size in British Columbia. Using trend information from at-sea surveys spanning a wide geographic range in Alaska, murrelet numbers declined significantly at five of eight trend sites at annual rates of -5.4 to -12.7 percent since the early 1990s. Applying these rates of decline to the historical population estimate, the current murrelet population in Alaska is projected to be on the order of 270,000 birds. This represents an overall population decline of about 70 percent during the past 25 years. In British Columbia, available trend data indicate that murrelet populations there have experienced similar declines. We updated a recent (2002) population estimate for British Columbia, concluding that there are now between 54,000 and 92,000 murrelets in British Columbia. The rates of decline we observed are within, but at the high end of, a range of rates expected by chance. Given that declines were estimated for sites over essentially the entire northern range of the species, there is cause for concern about the species’ status. In their marine habitats, Marbled Murrelets overlap with salmon (Oncorhynchus sp.) gillnetting operations in British Columbia and in Alaska (especially in Prince William Sound and Southeast Alaska), and annual bycatch mortality is likely in the low thousands per year, although bycatch rates are difficult to measure. The species’ inshore distribution coincides with high levels of vessel traffic and makes them especially vulnerable to both chronic oil pollution and to catastrophic spills (e.g., the 1989 Exxon Valdez oil spill [EVOS] in south-central Alaska, which is estimated to have killed 12,000 to 15,000 murrelets). In their forested nesting habitats, Marbled Murrelets have lost about 15 percent of their suitable nesting habitat in Southeast Alaska, and 33 to 49 percent in British Columbia, from industrial-scale logging within the past half century. Increased predation also may be a threat to murrelet populations, related to fragmentation and edge effects from logging and development, and recent population increases observed for some important murrelet predators, including Bald Eagles ( Haliaeetus leucocephalus ), Common Ravens ( Corvus corax ), and Steller’s Jays ( Cyanocitta stelleri ). Nesting habitat losses cannot explain the declines observed in areas where industrial logging has not occurred on a large scale (e.g., Prince William Sound) or at all (Glacier Bay). The apparent change in population size and rates of decline reported for the Marbled Murrelet are large, and we therefore considered alternative explanations and precedents for changes of similar magnitude in other marine wildlife populations in the Northeastern Pacific Ocean. The declines are likely real, and related to combined and cumulative effects from climate-related changes in the marine ecosystem (most likely the 1977 regime shift) and human activities (logging, gillnet bycatch, oil pollution). Much uncertainty about the decline could be alleviated by continuing to repeat boat surveys in Prince William Sound and lower Cook Inlet, and by repeating the boat survey of Southeast Alaska that was conducted in 1994. This survey used a statistically sound design and covered the region that has been and likely remains the center of the species’ abundance. Important questions remain to be addressed about methods for measuring population status and change, adult mortality (major sources, density dependence, seasonal concordance), and the movements of wintering populations.

Alaska, British Columbia

Proceedings of the Fourth Glacier Bay Science Symposium

Foreword Glacier Bay was established as a National Monument in 1925, in part to protect its unique character and natural beauty, but also to create a natural laboratory to examine evolution of the glacial landscape. Today, Glacier Bay National Park and Preserve is still a place of profound natural beauty and dynamic landscapes. It also remains a focal point for scientific research and includes continuing observations begun decades ago of glacial processes and terrestrial ecosystems. In recent years, research has focused on glacial-marine interactions and ecosystem processes that occur below the surface of the bay. In October 2004, Glacier Bay National Park convened the fourth in a series of science symposiums to provide an opportunity for researchers, managers, interpreters, educators, students and the general public to share knowledge about Glacier Bay. The Fourth Glacier Bay Science Symposium was held in Juneau, Alaska, rather than at the Park, reflecting a desire to maximize attendance and communication among a growing and diverse number of stakeholders interested in science in the park. More than 400 people attended the symposium. Participants provided 46 oral presentations and 41 posters covering a wide array of disciplines including geology, glaciology, oceanography, wildlife and fisheries biology, terrestrial and marine ecology, socio-cultural research and management issues. A panel discussion focused on the importance of connectivity in Glacier Bay research, and keynote speakers (Gary Davis and Terry Chapin) spoke of long-term monitoring and ecological processes. These proceedings include 56 papers from the symposium. A summary of the Glacier Bay Science Plan-itself a subject of a meeting during the symposium and the result of ongoing discussions between scientists and resource managers-also is provided. We hope these proceedings illustrate the diversity of completed and ongoing scientific studies, conducted within the Park. To this end, we invited all presenters to submit brief technical summaries of their work, to capture the gist of their study and its main findings without an overload of details and methodology. We also asked authors to include a few words on the management implications of their work to help bridge the gap between scientists and managers in understanding how specific research questions may translate to management practice. Papers in this volume are laid out by subject matter, from terrestrial and freshwater subjects to glacial-marine geology, to the ecology of marine animals and ending with risk assessment, human impacts and science-management considerations. In summary, we hope the proceedings will serve as a useful reference to completed and ongoing studies in Glacier Bay National Park, and thereby provide park enthusiasts, scientists, and managers with a road map of scientific progress.

Scientific Investigations Report

Distribution of ground-nesting marine birds along shorelines in Glacier Bay, southeastern Alaska: An assessment related to potential disturbance by back-country users

With the exception of a few large colonies, the distribution of ground-nesting marine birds in Glacier Bay National Park in southeastern Alaska is largely unknown. As visitor use increases in back-country areas of the park, there is growing concern over the potential impact of human activities on breeding birds. During the 2003–05 breeding seasons, the shoreline of Glacier Bay was surveyed to locate ground-nesting marine birds and their nesting areas, including wildlife closures and historical sites for egg collection by Alaska Native peoples. The nesting distribution of four common ground-nesting marine bird species was determined: Arctic Tern ( Sterna paradisaea ), Black Oystercatcher ( Haematopus bachmani ), Mew Gull ( Larus canus ), and Glaucous-winged Gull ( Larus glaucescens ). Observations of less abundant species also were recorded, including Herring Gull ( Larus argentatus ), Red-throated Loon ( Gavia stellata ), Canada Goose ( Branta canadensis ), Willow Ptarmigan ( Lagopus lagopus ), Semipalmated Plover ( Charadrius semipalmatus ), Spotted Sandpiper ( Actitis macularia ), Least Sandpiper ( Calidris minutilla ), Parasitic Jaeger ( Stercorarius parasiticus ), and Aleutian Tern ( Sterna aleutica ). Nesting distribution for Arctic Terns was largely restricted to the upper arms of the bay and a few treeless islets in the lower bay, whereas Black Oystercatchers were more widely distributed along shorelines in the park. Mew Gulls nested throughout the upper bay in Geikie Inlet and in Fingers and Berg Bays, and most Glaucous-winged Gull nests were found at wildlife closures in the central and lower bays. Several areas were identified where human disturbance could affect breeding birds. This study comprises the first bay-wide survey for the breeding distribution of ground-nesting marine birds in Glacier Bay National Park, providing a minimum estimate of their numbers and distribution within the park. This information can be used to assess future human disturbance and track natural changes in nesting bird distribution over time.

Alaska

Seabirds as indicators of marine ecosystems: Introduction: A modern role for seabirds as indicators

A key requirement for implementing ecosystem-based management is to obtain timely information on significant fluctuations in the ecosystem (Botsford et al. 1997). However, obtaining all necessary information about physical and biological changes at appropriate temporal and spatial scales is a daunting task. Intuitively, one might assume that physical data are more important for the interpretation of ecosystem changes than biological data, but analyses of time series data suggest otherwise: physical data are more erratic and often confusing over the short term compared to biological data, which tend to fluctuate less on annual time scales (Hare & Mantua 2000). Even so, biological time-series may also be confusing when coexisting marine species respond differently to ecosystem variability. For example, while warming temperatures in the Gulf of Alaska following the 1976 to 1977 regime shift favored an increase in gadoids and flatfish, a variety of forage fish and pandalid shrimp species virtually disappeared (Anderson & Piatt 1999). Zooplankton communities in the Gulf of Alaska also demonstrated similar patterns of response (Francis et al. 1998). At the basin scale, favorable conditions for salmon in Alaska following the regime shift were matched inversely by poor conditions in the California Current (Francis et al. 1998). In marine birds, subtropical species increased, while subarctic ones decreased during a warming phase in the southern California Bight. Clearly, no single index can tell the whole story accurately. Multi-species, multi-region, and multi-trophic level approaches are needed to quantify fluctuations in marine ecosystem processes and in the distribution and abundance of its inhabitants, to determine critical parameter thresholds and to use this information in management and marine conservation.

Marine Ecology Progress Series

Seabird behavior as an indicator of food supplies: Sensitivity across the breeding season

We used empirical data on the time allocation of common murres Uria aalge in relation to measures of local prey density to examine whether adults provisioning chicks are more sensitive to changes in prey density than birds that are incubating eggs. We hypothesized that seasonal differences in food requirements of incubating and chick-rearing parents would affect the form of the relationship between time spent at the colony and local food density. We found that the relationship did differ between the incubation and chick-rearing period in 3 important ways: (1) there was a strong non-linear relationship between food density and colony attendance during chick-rearing and a weaker relationship during incubation; (2) incubating birds were able to maintain relatively constant rates of attendance over a wider range of food densities than chick-rearing birds and only reduced colony attendance under extremely poor feeding conditions, if at all; and (3) incubating birds spent more time attending nest sites at the colony than provisioning birds. These differences confirmed that chick-rearing parents are more sensitive to changes in food density than incubating parents, and that measurements of time allocation during the incubation period would have limited value as an indicator of ecosystem change. ?? Inter-Research 2007.

Marine Ecology Progress Series

Stress hormones link food availability and population processes in seabirds

Catastrophic population declines in marine top predators in the northern Pacific have been hypothesized to result from nutritional stress affecting reproduction and survival of individuals. However, empirical evidence for food-related stress in wild animals is frequently lacking or inconclusive. We used a field endocrinology approach to measure stress, identify its causes, and examine a link between stress and population processes in the common murre Uria aalge. We tested the empirical relationship between variations in the stress hormone corticosterone (CORT) and food abundance, reproduction, and persistence of individuals at declining and increasing colonies in Cook Inlet, Alaska, from 1996 to 2001. We found that CORT secretion in murres is independent of colony, reproductive stage effects, and gender of individuals, but is directly negatively correlated with abundance of their food. Baseline CORT reflected current food abundance, whereas acute stress-induced CORT reflected food abundance in the previous month. As food supply diminished, increased CORT secretion predicted a decrease in reproductive performance. At a declining colony, increased baseline levels of CORT during reproduction predicted disappearance of individuals from the population. Persistence of individuals in a growing colony was independent of CORT during reproduction. The obtained results support the hypothesis that nutritional stress during reproduction affects reproduction and survival in seabirds. This study provides the first unequivocal evidence for CORT secretion as a mechanistic link between fluctuations in food abundance and population processes in seabirds. ?? Inter-Research 2007.

Marine Ecology Progress Series

Seabirds as indicators of marine food supplies: Cairns revisited

In his seminal paper about using seabirds as indicators of marine food supplies, Cairns (1987, Biol Oceanogr 5:261&ndash;271) predicted that (1) parameters of seabird biology and behavior would vary in curvilinear fashion with changes in food supply, (2) the threshold of prey density over which birds responded would be different for each parameter, and (3) different seabird species would respond differently to variation in food availability depending on foraging behavior and ability to adjust time budgets. We tested these predictions using data collected at colonies of common murre Uria aalge and black-legged kittiwake Rissa tridactyla in Cook Inlet, Alaska. (1) Of 22 seabird responses fitted with linear and non-linear functions, 16 responses exhibited significant curvilinear shapes, and Akaike&rsquo;s information criterion (AIC) analysis indicated that curvilinear functions provided the best-fitting model for 12 of those. (2) However, there were few differences among parameters in their threshold to prey density, presumably because most responses ultimately depend upon a single threshold for prey acquisition at sea. (3) There were similarities and some differences in how species responded to variability in prey density. Both murres and kittiwakes minimized variability (CV < 15%) in their own body condition and growth of chicks in the face of high annual variability (CV = 69%) in local prey density. Whereas kittiwake breeding success (CV = 63%, r 2 = 0.89) reflected prey variability, murre breeding success did not (CV = 29%, r 2 < 0.00). It appears that murres were able to buffer breeding success by reallocating discretionary &lsquo;loafing&rsquo; time to foraging effort in response (r 2 = 0.64) to declining prey density. Kittiwakes had little or no discretionary time, so fledging success was a more direct function of local prey density. Implications of these results for using &lsquo;seabirds as indicators&rsquo; are discussed.

Marine Ecology Progress Series

Investigating trophic relationships of pinnipeds in Alaska and Washington using stable isotope ratios of nitrogen and carbon

We measured stable-nitrogen (δ 15 N) and stable-carbon (δ 13 C) isotope ratios in muscle and hair from 7 northern fur seals ( Callorhinus ursinus ) from the Pribilof Islands, Alaska, and 27 Steller sea lions ( Eumetopias jubatus ), and 14 harbor seals ( Phoca vitulina ) from the Gulf of Alaska and coast of Washington State, in order to contrast dietary information derived from isotopic vs. available conventional dietary studies. Stable-nitrogen-isotope analysis of muscle revealed that harbor seals were enriched over sea lions (mean δ 15 N = 18.6‰ vs. 17.5‰) which were in turn enriched over northern fur seals (mean δ 15 N = 16.6‰). Trophic segregation among these species likely results primarily from differential reliance on herring ( Clupea harengus ), Atka mackerel ( Pleurogrammus monopterygius ), and large vs. small walleye pollock ( Theregra chalcogramma ). According to their δ 15 N values, adult male Steller sea lions showed a higher trophic position than adult females (mean δ 15 N: 18.0‰ vs. 17.2‰), whereas adult female northern fur seals were trophically higher than juvenile male fur seals (mean δ 15 N: 16.5‰ vs. 15.0‰). Each of these observed differences likely resulted from differential reliance on squid or differences in the size range of pollock consumed. Three northern fur seal pups showed higher δ 15 N enrichment over adults (mean 17.7‰ vs. 15.8‰) due to their reliance on their mother's milk. Stable-carbon isotope measurements of hair revealed a cline toward more negative values with latitude. Segregation in hair δ 13 C between Steller sea lions and harbor seals off the coast of Washington (mean δ 13 C: -13.6‰ vs. - 15.0‰) reflected the greater association of harbor seals with freshwater input from the Columbia River. Our study demonstrates the utility of the stable isotope approach to augment conventional dietary analyses of pinnipeds and other marine mammals.

Alaska, Washington

Predictable hotspots and foraging habitat of the endangered short-tailed albatross (Phoebastria albatrus) in the North Pacific: Implications for conservation

The short-tailed albatross ( Phoebastria albatrus ) is a rare and endangered seabird that ranges widely over the northern North Pacific. Populations are slowly recovering but birds face several threats at sea, in particular the incidental capture of birds in long-line fisheries. Conservation efforts are hampered by a lack of information about the at-sea distribution of this species, especially knowledge of where it may predictably co-occur with long-line fishing effort. During 18 years of transiting the Aleutian Islands Unit of the Alaska Maritime National Wildlife Refuge on a research vessel, we observed short-tailed albatross on 65 occasions. They were consistently observed near Ingenstrem Rocks (Buldir Pass) in the western Aleutians and near Seguam Pass in the central Aleutians. Based on the oceanographic characteristics of the locations where we saw most of the birds, we hypothesized that short-tailed albatross &ldquo;hotspots&rdquo; were located where tidal currents and steep bottom topography generate strong vertical mixing along the Aleutian Archipelago. As a test of this hypothesis, we analyzed a database containing 1432 opportunistic observations of 2463 short-tailed albatross at sea in the North Pacific. These data showed that short-tailed albatross were closely associated with shelf-edge habitats throughout the northern Gulf of Alaska and Bering Sea. In addition to Ingenstrem Rocks and Seguam Pass, important hotspots for short-tailed albatross in the Aleutians included Near Strait, Samalga Pass, and the shelf-edge south of Umnak/Unalaska islands. In the Bering Sea, hotspots were located along margins of Zhemchug, St. Matthews and Pervenets canyons. Because these short-tailed albatross hotspots are predictable, they are also protectable by regulation of threatening activities at local spatial scales.

Deep-Sea Research Part II: Topical Studies in Ocea

Balancing predation and egg harvest in a colonial seabird: A simulation model

We developed an individual-based model to study the effects of different regimes of harvesting eggs and natural predation on reproductive success in a colony of the glaucous-winged gull ( Larus glaucescens ) in Glacier Bay National Park, Alaska. The model incorporates the sequence of egg laying, relaying, and incubation to hatching for individual nests and calculates hatching success, incubation length, and the total number of eggs laid (as a result of re-nesting and relaying) in all nests in the colony. Stochasticity is incorporated in the distribution of nest lay dates, predation rates, and nests attacked during predation and harvest events. We estimated parameter values by fitting the model to data collected at a small colony during 1999 and 2000 using maximum likelihood. We then simulated harvests and analyzed model predictions. Model outputs indicate that harvesting early, and at one time, provides a predictable take of eggs with the least impact to gulls.

Ecological Modelling

Testing the junk-food hypothesis on marine birds: Effects of prey type on growth and development

The junk-food hypothesis attributes declines in productivity of marine birds and mammals to changes in the species of prey they consume and corresponding differences in nutritional quality of those prey. To test this hypothesis nestling Black-legged Kittiwakes (Rissa tridactyla) and Tufted Puffins (Fratercula cirrhata) were raised in captivity under controlled conditions to determine whether the type and quality of fish consumed by young seabirds constrains their growth and development. Some nestlings were fed rations of Capelin (Mallotus villosus), Herring (Clupea pallasi) or Sand Lance (Ammodytes hexapterus) and their growth was compared with nestlings raised on equal biomass rations of Walleye Pollock (Theragra chalcograma). Nestlings fed rations of herring, sand lance, or capelin experienced higher growth increments than nestlings fed pollock. The energy density of forage fish fed to nestlings had a marked effect on growth increments and could be expected to have an effect on pre- and post-fledging survival of nestlings in the wild. These results provide empirical support for the junk-food hypothesis.

Waterbirds

Population genetic structure and conservation of marbled murrelets ( Brachyramphus marmoratus )

Marbled murrelets ( Brachyramphus marmoratus ) are coastal seabirds that nest from California to the Aleutian Islands. They are declining and considered threatened in several regions. We compared variation in the mitochondrial control region, four nuclear introns and three microsatellite loci among 194 murrelets from throughout their range except Washington and Oregon. Significant population genetic structure was found: nine private control region haplotypes and three private intron alleles occurred at high frequency in the Aleutians and California; global estimates of FST or ??ST and most pairwise estimates involving the Aleutians and/or California were significant; and marked isolation-by-distance was found. Given the available samples, murrelets appear to comprise five genetic management units: (1) western Aleutian Islands, (2) central Aleutian Islands, (3) mainland Alaska and British Columbia, (4) northern California, and (5) central California.

Conservation Genetics

A mechanistic link between chick diet and decline in seabirds?

A climatic regime shift during the mid-1970s in the North Pacific resulted in decreased availability of lipidrich fish to seabirds and was followed by a dramatic decline in number of kittiwakes breeding on the Pribilof Islands. Although production of chicks in the mid-1970s was adequate to sustain kittiwake populations in the early 1980s, the disappearance of birds from breeding colonies apparently exceeded recruitment. No mechanism has been proposed to explain why recruitment would differ among fledglings fed lipid-rich or lipid-poor fish during development. Here we show that diets low in lipids induce nutritional stress and impair cognitive abilities in young red-legged kittiwakes, Rissa brevirostris. Specifically, growth retardation, increased secretion of stress hormones and inferior ability to associate food distribution with visual cues were observed in individuals fed lipid-poor diets. We conclude that lipid-poor diets during development affect the quality of young seabirds, which is likely to result in their increased mortality and low recruitment. ?? 2005 The Royal Society.

Proceedings of the Royal Society B: Biological Sci

Prey consumption and energy transfer by marine birds in the Gulf of Alaska

We investigated prey consumption by marine birds and their contribution to cross-shelf fluxes in the northern Gulf of Alaska. We utilized data from the North Pacific Pelagic Seabird Database for modeling energy demand and prey consumption. We found that prey consumption by marine birds was much greater over the continental shelf than it was over the basin. Over the shelf, subsurface-foraging marine birds dominated food consumption, whereas over the basin, surface-foraging birds took the most prey biomass. Daily consumption by marine birds during the non-breeding season ("winter") from September through April was greater than daily consumption during the breeding season, between May and August. Over the shelf, shearwaters, murres and, in winter, sea ducks, were the most important consumers. Over the basin, northern fulmars, gulls and kittiwakes predominated in winter and storm-petrels dominated in May to August. Our results suggest that marine birds contribute little to cross-shelf fluxes of energy or matter, but they do remove energy from the marine system through consumption, respiration and migration. ?? 2005 Elsevier Ltd. All rights reserved.

Conference Paper

Parallel structure among environmental gradients and three trophic levels in a subarctic estuary

We assessed spatial and temporal variability in the physical environment of a subarctic estuary, and examined concurrent patterns of chlorophyll &alpha; abundance (fluorescence), and zooplankton and forage fish community structure. Surveys were conducted in lower Cook Inlet, Alaska, during late July and early August from 1997 through 1999. Principle components analysis (PCA) revealed that spatial heterogeneity in the physical oceanographic environment of lower Cook Inlet could be modeled as three marine-estuarine gradients characterized by temperature, salinity, bottom depth, and turbidity. The gradients persisted from 1997 through 1999, and PCA explained 68% to 92% of the variance in physical oceanography for each gradient-year combination. Correlations between chlorophyll &alpha; abundance and distribution and the PCA axes were weak. Chlorophyll was reduced by turbidity, and low levels occurred in areas with high levels of suspended sediments. Detrended correspondence analysis (DCA) was used to order the sample sites based on species composition and to order the zooplankton and forage fish taxa based on similarities among sample sites for each gradient-year. Correlations between the structure of the physical environment (PCA axis 1) and zooplankton community structure (DCA axis 1) were strong (r = 0.43-0.86) in all years for the three marine-estuarine gradients, suggesting that zooplankton community composition was structured by the physical environment. The physical environment (PCA) and forage fish community structure (DCA) were weakly correlated in all years along Gradient 2, defined by halocline intensity and surface temperature and salinity, even though these physical variables were more important for defining zooplankton habitats. However, the physical environment (PCA) and forage fish community structure (DCA) were strongly correlated along the primary marine-estuarine gradient (#1) in 1997 (r = 0.87) and 1998 (r = 0.82). The correlation was poor (r = 0.32) in 1999, when fish community structure changed markedly in lower Cook Inlet. Capelin (Mallotus villosus), walleye pollock ( Theragra chalcogramma ), and arrowtooth flounder ( Atheresthes stomias ) were caught farther north than in previous years. Waters were significantly colder and more saline in 1999, a La Nina year, than in other years of the study. Interannual fluctuations in environmental conditions in lower Cook Inlet did not have substantial effects on zooplankton community structure, although abundance of individual taxa varied significantly. The abundance and distribution of chlorophyll &alpha;, zooplankton and forage fish were affected much more by spatial variability in physical oceanography than by interannual variability. Our examination of physical-biological linkages in lower Cook Inlet supports the concept of "bottom-up control," i.e., that variability in the physical environment structures higher trophic-level communities by influencing their distribution and abundance across space.

Progress in Oceanography