Seabirds as indicators

In his seminal paper about using seabirds as indicators of marine food supplies, Cairns (1987, Biol Oceanogr 5:261-271) 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'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 'loafing' 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 'seabirds as indicators' are discussed.

In his seminal paper about using seabirds as indicators of marine food supplies, Cairns (1987, Biol Oceanogr 5:261-271) 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'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 'loafing' 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 'seabirds as indicators' are discussed.

The growth rate of chicks is a parameter often used to examine the effects of changes in food availability on reproductive performance of seabirds. We analyzed the relationship of growth rates in Black-legged Kittiwake Rissa tridactyla chicks at three colonies in Prince William Sound, Alaska, and three colonies in Lower Cook Inlet, Alaska, to two components of nestling meals that represent diet quantity (meal delivery rate and meal size) and one that represents diet quality (energy density). During this study, kittiwakes appeared to rely on an array of relatively high-lipid prey. Growth rates of alpha and singleton chicks were positively correlated with annual productivity and were best explained by meal delivery rate and meal size; growth rates of beta chicks were not correlated with productivity and were best explained by meal delivery rate alone. The lack of a positive effect of energy density on growth rates was in contrast to results from a companion study that found a moderately positive effect of energy density on kittiwake productivity. Taken together, results from these two studies demonstrate the need to measure both diet quantity and quality to reliably assess the relationships between diet composition, provisioning rates, growth rates, and productivity in seabirds.

Elevated seawater temperatures are linked to the development of harmful algal blooms (HABs), which pose a growing threat to marine birds and other wildlife. During late 2015 and early 2016, a massive die-off of Common Murres (Uria aalge; hereafter, murres) was observed in the Gulf of Alaska coincident with a strong marine heat wave. Previous studies have documented illness and death among seabirds resulting from exposure to the HAB neurotoxins saxitoxin (STX) and domoic acid (DA). Given the unusual mortality event, corresponding warm water anomalies, and recent detection of STX and DA throughout coastal Alaskan waters, HABs were identified as a possible factor of concern. To evaluate whether algal toxins may have contributed to murre deaths, we tested for STX and DA in a suite of tissues obtained from beach-cast murre carcasses associated with the die-off as well as from apparently healthy murres and Black-legged Kittiwakes (Rissa tridactyla; hereafter, kittiwakes) sampled in the preceding and following summers. We also tested forage fish and marine invertebrates collected in the Gulf of Alaska in 2015-2017 to evaluate potential sources of HAB toxin exposure for seabirds. Saxitoxin was present in multiple tissue types of both die-off (36.4 %) and healthy (41.7 %) murres and healthy kittiwakes (54.2 %). Among birds, we detected the highest concentrations of STX in liver tissues (range 1.4-10.8 μg 100 g −1) of die-off murres. Saxitoxin was relatively common in forage fish (20.3 %) and invertebrates (53.8 %). No established toxicity limits currently exist for seabirds, but concentrations of STX in birds and forage fish in our study were lower than values reported from most other bird die-offs in which STX intoxication was causally linked. We detected low concentrations of DA in a single bird sample and in 33.3 % of invertebrates and 4.0 % of forage fish samples. Although these results do not support the hypothesis that acute exposure to STX or DA was a primary factor in the 2015-2016 mortality event, additional information about the sensitivity of murres to these toxins is needed before we can discount their potential role in the die-off. The widespread occurrence of STX in seabirds, forage fish, and invertebrates in the Gulf of Alaska indicates that algal toxins should be considered in future assessments of seabird health, especially given the potential for greater occurrence of HABs in the future.

During breeding, long-lived species face important time and energy constraints that can lead to breeding failure when food becomes scarce. Despite the potential implications of intra-season dynamics in breeding failure for individual behavior, carry-over effects, dispersal decisions and population dynamics, little information is currently available on these dynamics at fine temporal scales. Here, we monitored the foraging behavior and the proportion of successful black-legged kittiwake pairs from nest construction to chick fledging in a colony of the southern Barents Sea, to relate foraging effort to the dynamics of breeding failure over an entire breeding season, and to infer the environmental conditions leading to this failure. Specifically, we tracked kittiwakes with GPS and satellite tags during incubation and early chick-rearing to document nest attendance, foraging range, time budgets and daily energy expenditures (DEE). We also monitored diet changes over time. We predicted that breeding failure would follow a non-linear trend characterized by a break point after which breeding success would drop abruptly and would be related to a substantial increase in foraging effort. Kittiwakes showed contrasting foraging patterns between incubation and chick-rearing: they extended their foraging range from 20 km during incubation to more than 450 km during chick-rearing and switched diet. They also increased their DEE and readjusted their time budgets by increasing time spent at sea. These changes corresponded to a break point in breeding dynamics beyond which the proportion of successful pairs abruptly dropped. At the end of the season, less than 10% of kittiwake pairs raised chicks in the monitored plots. This integrative study confirms that breeding failure is a non-linear process characterized by a threshold beyond which individuals face an energetic trade-off and cannot simultaneously sustain high reproductive and selfmaintenance efforts. In this way, the occurrence of sudden environmental changes complicates our ability to predict population dynamics and poses conservation challenges.

Some seabird populations damaged by the Exxon Valdez oil spill have not recovered, and populations will need to be monitored for many years to assess both recovery and ecological conditions affecting recovery. Detailed studies of individual seabird colonies and marine ecosystems in the Gulf of Alaska (GOA) have been conducted by the USGS and USFWS under the auspices of damage assessment and restoration programs of the EVOSTC. Much has been learned about factors influencing seabird populations and their capacity to recover from the spill in the GOA. As we move towards long-term monitoring of populations, however, we need to develop protocols and long-term monitoring strategies that focus on key parameters of interest and that are inexpensive, practical and applicable over a large geographic area.

We have been conducting detailed studies of the biology of seabirds in relation to oceanography and forage fish ecology in lower Cook Inlet, Alaska, since 1995. This fortuitously allowed us to document biological effects of the 1997/98 ENSO in this region. Anomalously warm sea surface temperatures (SST’s) were observed in the Gulf of Alaska (GOA) beginning in June of 1997, but not in Cook Inlet until September, 1997. Warm temperature anomalies at the surface and at depth persisted until May of 1998, when temperatures returned to average in the GOA and Cook Inlet. Thus, temperature anomalies occurred outside the core window of productivity (June-August) for forage fish and seabirds in both 1997 and 1998. Abundance or production of phytoplankton, zooplankton, fish, and seabirds in lower Cook Inlet varied among years, and overall appeared to be depressed in 1998. We observed a few biological anomalies that might be attributed to ENSO effects: 1) a significant die-off of Common Murres o...

Some seabird populations damaged by the Exxon Valdez oil spill have not recovered, and populations will need to be monitored for many years to assess both recovery and ecological conditions affecting recovery. Detailed studies of individual seabird colonies and marine ecosystems in the Gulf of Alaska (GOA) have been conducted by the USGS and USFWS under the auspices of damage assessment and restoration programs of the EVOSTC. Much has been learned about factors influencing seabird populations and their capacity to recover from the spill in the GOA. As we move towards long-term monitoring of populations, however, we need to develop protocols and long-term monitoring strategies that focus on key parameters of interest and that are inexpensive, practical and applicable over a large geographic area.

Reproductive success at seabird colonies is often quite variable, even within a single region. Although exhaustive investigations of reproductive success drivers are not logistically feasible, many long-term seabird colony monitoring datasets may be combined with ancillary environmental datasets to reveal key relationships that may inform management and future investigations. Here we explored biological and environmental influences on reproductive success of two surface-feeding seabird species (Black-legged Kittiwakes Rissa tridactyla and Common Terns Sterna hirundo) that inhabit different oceans and climate zones and experience different management approaches; the terns’ predators were controlled while the kittiwakes were not managed. Both systems showed evidence of negative density dependence between colony size and reproductive success, contrasting with positive associations between region-wide population density and reproductive success. Direct environmental influences were also...

Reproductive success is one of the most easily-measured and widely studied demographic parameters of colonial nesting seabirds. Nevertheless, factors affecting the sequential stages (egg laying, incubation, chickrearing) of reproductive success are less understood. We investigated the separate sequential stages of reproductive success in piscivorous black-legged kittiwakes (Rissa tridactyla) and thick-billed murres (Uria lomvia) using a 36-year dataset (1975-2010) on the major Pribilof Islands (St. Paul and St. George), which have recently had contrasting population trajectories. Our objectives were to evaluate how the proportion of successful nests varied among stages, and to quantify factors influencing the probability of nest success at each stage in each island. We modeled the probability of nest success at each stage using General Linear Mixed Models incorporating broad-scale and local climate variables, and diet as covariates as well as other measures of reproduction such as timing of breeding and reproductive output in the previous year and previous stage. For both species we found: (1) Success in previous stages of the breeding cycle and success in the prior year better explained overall success than any environmental variables. Phenology was also an important predictor of laying success for kittiwakes. (2) Fledging success was lower when chick diets contained oceanic fish found farther from the colonies and small invertebrates, rather than coastal fish species. (3) Differences in reproductive variables at St. Paul and St. George islands did not correspond to population trends between the two islands. Our results highlight the potential importance of adult condition and annual survival to kittiwake and murre productivity and ultimately, populations. Adult condition carrying over from the previous year ultimately seems to drive annual breeding success in a cascade effect. Furthermore, condition and survival appear to be important contributors to population dynamics at each island. Therefore, adult condition and survival prior to breeding, and factors that influence these parameters such as foraging conditions in the non-breeding season, may be important datasets for understanding drivers of seabird demography at the Pribilof Islands.

Forage fishes were sampled with a mid-water trawl in lower Cook Inlet, Alaska, USA, from late July to early August 1996 to 1999. We sampled 3 oceanographically distinct areas of lower Cook Inlet: waters adjacent to Chisik Island, in Kachemak Bay, and near the Barren Islands. In 163 tows using a mid-water trawl, 229 437 fishes with fork length < 200 mm were captured. More than 39 species were captured in lower Cook Inlet, but Pacific sand lance Ammodytes hexapterus, juvenile Pacific herring Clupea pallasi, and juvenile walleye pollock Theragra chalcogramma comprised 97.5% of the total individuals. Both species richness and species diversity were highest in warm, lowsalinity, weakly stratified waters near Chisik Island. Kachemak Bay, which had thermohaline values between those found near Chisik Island and the Barren Islands, had an intermediate value of species richness. Species richness was lowest at the Barren Islands, an exposed region that regularly receives oceanic, upwelled water from the Gulf of Alaska. Non-metric multidimensional scaling (NMDS) was used to compute axes of species composition based on an ordination of pairwise site dissimilarities. Each axis was strongly rank-correlated with unique groups of species and examined separately as a function of environmental parameters (temperature, salinity, depth), area, and year. Oceanographic parameters accounted for 41 and 12% of the variability among forage fishes indicated by Axis 1 and Axis 2, respectively. Axis 1 also captured the spatial variability in the upwelled area of lower Cook Inlet and essentially contrasted the distribution of species among shallow, nearshore (sand lance, herring) and deep, offshore (walleye pollock) habitats. Axis 2 captured the spatial variability in forage fish communities from the north (Chisik Island) to the south (Barren Islands) of lower Cook Inlet and essentially contrasted a highly diverse community dominated by salmonids and osmerids (warmer, less saline) with a fish community dominated by Pacific sand lance (colder, more saline). Axis 3 reflected the negative spatial association of capelin Mallotus villosus and Pacific cod Gadus macrocephalus. Correlations of year with Axes 1 and 3 indicate that from 1996 to 1999 the forage fish community significantly decreased in lipid-poor gadids (walleye pollock and Pacific cod), and significantly increased in lipid-rich species such as Pacific sand lance, Pacific herring, and capelin. KEY WORDS: Community structure • Forage fish • Lipid • Mid-water distribution • Capelin • Gadids • Cook Inlet • Gulf of Alaska Resale or republication not permitted without written consent of the publisher 1 The term 'forage fishes' applies to mid-water, schooling fishes that are prey to marine mammals, seabirds, and larger fishes during some phase of their life-history (Springer & Speckman 1997)

Food availability comprises a complex interaction of factors that integrates abundance, taxonomic composition, accessibility, and quality of the prey base. The relationship between food availability and reproductive performance can be assessed via the nutritional stress (NSH) and junkfood (JFH) hypotheses. With respect to reproductive success, NSH posits that a deficiency in any of the aforementioned metrics can have a deleterious effect on a population via poor reproductive success. JFH, a component of NSH, posits specifically that it is a decline in the quality of food (i.e. energy density and lipid content) that leads to poor reproductive success. We assessed each in relation to reproductive success in a piscivorous seabird, the black-legged kittiwake Rissa tridactyla. We measured productivity, taxonomic composition, frequency, size, and quality of meals delivered to nestlings from 1996 to 1999 at 6 colonies in Alaska, USA, 3 each in Prince William Sound and Lower Cook Inlet. Productivity varied widely among colony-years. Pacific herring Clupea pallasi, sand lance Ammodytes hexapterus, and capelin Mallotus villosus comprised ca. 80% of the diet among colonyyears, and each was characterized by relatively high energy density. Diet quality for kittiwakes in this region therefore remained uniformly high during this study. Meal delivery rate and meal size were quite variable among colony-years, however, and best explained the variability in productivity. Parent kittiwakes appeared to select prey that were energy dense and that maximized the biomass provisioned to broods. While these results fail to support JFH, they do provide substantial support for NSH.

To better understand how fluctuations in prey abundance may impact seabird reproductive success, we studied short-term changes in prey populations and their effect on prey selection and brood-rearing in the black-legged kittiwake Rissa tridactyla, a predator of near-surface-schooling forage fishes. Our fine-scale approach involved a weekly assessment of forage fish abundance and brood-rearing conditions during 4 consecutive years (1996 to 1999) at the Shoup Bay kittiwake colony in Prince William Sound, Alaska. We conducted forage fish surveys from a fixed-wing aircraft to determine weekly prey abundance throughout the known foraging range of breeding kittiwakes. Our results provide clear evidence that short-term fluctuations in prey availability are responsible for dramatic, within-season changes in the breeding conditions of black-legged kittiwakes. Adult kittiwakes often showed immediate response to changes in the prey base by altering prey selection; however, there were instances when kittiwakes selected prey species disproportionate to their availability (typically selecting for Pacific herring Clupea pallasi and against Pacific sand lance Ammodytes hexapterus). Changes in prey selection often resulted in striking differences in the amount of time required to obtain a load of food. The cascading effects of longer foraging trips was translated into reduced nestling growth and survival. Of the 3 components of energy provisioning to nestlings (meal delivery rate, meal size, and energy density), meal delivery rate had the strongest and most consistent positive effect on nestling growth and survival. Overall, these results demonstrate that complex foraging conditions limit the reproductive success of a central place-foraging species relying on an ephemeral food source. Moreover, we demonstrated that feeding conditions during the first 2 wk of brood-rearing were most critical for survival of the brood. Given the potential for such marked within-season variation in breeding conditions, it is critical that investigators adequately sample throughout the brood-rearing period, or, alternatively, select that portion that is germane to their study.

Variability in forage fish abundance strongly affects seabird behavior and reproductive success, although details of this relationship are unclear. During 1997 and 1998, we measured (1) daily energy expenditure (DEE) of 80 parent black-legged kittiwakes Rissa tridactyla at 2 colonies in Prince William Sound, Alaska (North Icy Bay and Shoup Bay), (2) abundance of surface-schooling forage fishes within the foraging range of each colony, and (3) diet composition, energy delivery rates to nestlings, and reproductive success of kittiwakes at these same colonies. Female DEE was highest at North Icy Bay in 1998, while male DEE did not differ by colony year. Abundances of Pacific herring Clupea pallasi and sand lance Ammodytes hexapterus were highest near North Icy Bay in 1998 and nearly equal in density, although Age 1+ herring comprised the majority of the diet there. Energy delivery rates to nestlings, nestling growth rates, and productivity were also highest at North Icy Bay in 1998. We suggest that female kittiwakes responded to the increased abundance of Age 1+ herring near North Icy Bay in 1998 by increasing their DEE, which in turn positively affected reproductive success. Given that adult kittiwakes have been shown to suffer decreased survival as a response to increased energy expenditure during brood rearing, the positive correlation we observed between increased abundance of a high quality food source, parental effort, and productivity is consistent with maximizing lifetime reproductive success. The lack of a response in male DEE suggests that broodrearing roles in kittiwakes differ between genders.

We used a supplemental-feeding experiment, the doubly labeled water technique, and a model-selection approach based upon the Akaike Information Criterion to examine effects of food availability on energy expenditure rates of Black-legged Kittiwakes (Rissa tridactyla) raising young. Energy expenditure rates of supplementally fed females (n = 14) and males (n = 16) were 34 and 20% lower than those of control females (n = 14) and males (n = 18), respectively. Energy expenditure rates of females were more responsive to fluctuations in food availability than those of males. Fed males likely expended more energy while off the nest than fed females, possibly because of nest defense. Energy expenditure rates of fed kittiwakes were similar to values reported for kittiwakes that were either not raising young or not foraging. Parent kittiwakes, therefore, adjusted parental effort in response to variation in breeding conditions due to changes in food availability. Adjustments in reproductive effort in response to variable foraging conditions may have significant effects on the survival and productivity of individuals, and thus provide substantial fitness benefits for long-lived seabirds such as Black-legged Kittiwakes. Résumé : Nous avons utilisé une expérience d'addition de nourriture, la technique de l'eau doublement marquée et un modèle de sélection basé sur le critère d'information d'Akaike pour étudier les effets de la disponibilité de la nourriture sur les taux de dépense énergétique chez des mouettes tridactyles (Rissa tridactyla) pendant l'élevage des jeunes. Les taux de dépense énergétique des femelles (n = 14) et des mâles (n = 16) qui ont reçu des suppléments de nourriture sont respectivement de 34 % et de 20 % moins élevés que ceux de femelles (n = 14) et de mâles (n = 18) témoins. Les taux de dépense énergétique des femelles sont plus sensibles aux fluctuations de la disponibilité de la nourriture que ceux des mâles. Les mâles nourris dépensent probablement plus d'énergie hors du nid que les femelles nourries, peut-être parce qu'ils assurent la défense du nid. Les taux de dépense énergétique des mouettes nourries sont semblables aux valeurs rapportées dans la littérature dans le cas de mouettes qui n'élèvent pas de jeunes ou qui ne font pas de quête de nourriture. Les mouettes ajustent donc leur effort parental en fonction des variations des conditions de la reproduction, elles-mêmes dépendantes de la disponibilité de la nourriture. L'ajustement de l'effort reproducteur aux conditions variables de quête de nourriture peut avoir un effet significatif sur la survie et la productivité individuelles, ce qui a pour conséquence d'améliorer le fitness de façon importante chez les oiseaux marins, tels que la mouette tridactyle.

Forage fishes were sampled with a mid-water trawl in lower Cook Inlet, Alaska, USA, from late July to early August 1996 to 1999. We sampled 3 oceanographically distinct areas of lower Cook Inlet: waters adjacent to Chisik Island, in Kachemak Bay, and near the Barren Islands. In 163 tows using a mid-water trawl, 229 437 fishes with fork length < 200 mm were captured. More than 39 species were captured in lower Cook Inlet, but Pacific sand lance Ammodytes hexapterus, juvenile Pacific herring Clupea pallasi, and juvenile walleye pollock Theragra chalcogramma comprised 97.5% of the total individuals. Both species richness and species diversity were highest in warm, lowsalinity, weakly stratified waters near Chisik Island. Kachemak Bay, which had thermohaline values between those found near Chisik Island and the Barren Islands, had an intermediate value of species richness. Species richness was lowest at the Barren Islands, an exposed region that regularly receives oceanic, upwelled water from the Gulf of Alaska. Non-metric multidimensional scaling (NMDS) was used to compute axes of species composition based on an ordination of pairwise site dissimilarities. Each axis was strongly rank-correlated with unique groups of species and examined separately as a function of environmental parameters (temperature, salinity, depth), area, and year. Oceanographic parameters accounted for 41 and 12% of the variability among forage fishes indicated by Axis 1 and Axis 2, respectively. Axis 1 also captured the spatial variability in the upwelled area of lower Cook Inlet and essentially contrasted the distribution of species among shallow, nearshore (sand lance, herring) and deep, offshore (walleye pollock) habitats. Axis 2 captured the spatial variability in forage fish communities from the north (Chisik Island) to the south (Barren Islands) of lower Cook Inlet and essentially contrasted a highly diverse community dominated by salmonids and osmerids (warmer, less saline) with a fish community dominated by Pacific sand lance (colder, more saline). Axis 3 reflected the negative spatial association of capelin Mallotus villosus and Pacific cod Gadus macrocephalus. Correlations of year with Axes 1 and 3 indicate that from 1996 to 1999 the forage fish community significantly decreased in lipid-poor gadids (walleye pollock and Pacific cod), and significantly increased in lipid-rich species such as Pacific sand lance, Pacific herring, and capelin. KEY WORDS: Community structure • Forage fish • Lipid • Mid-water distribution • Capelin • Gadids • Cook Inlet • Gulf of Alaska Resale or republication not permitted without written consent of the publisher 1 The term 'forage fishes' applies to mid-water, schooling fishes that are prey to marine mammals, seabirds, and larger fishes during some phase of their life-history (Springer & Speckman 1997)

We have been conducting detailed studies of the biology of seabirds in relation to oceanography and forage fish ecology in lower Cook Inlet, Alaska, since 1995. This fortuitously allowed us to document biological effects of the 1997/98 ENSO in this region. Anomalously warm sea surface temperatures (SST's) were observed in the Gulf of Alaska (GOA) beginning in June of 1997, but not in Cook Inlet until September, 1997. Warm temperature anomalies at the surface and at depth persisted until May of 1998, when temperatures returned to average in the GOA and Cook Inlet. Thus, temperature anomalies occurred outside the core window of productivity (June-August) for forage fish and seabirds in both 1997 and 1998. Abundance or production of phytoplankton, zooplankton, fish, and seabirds in lower Cook Inlet varied among years, and overall appeared to be depressed in 1998. We observed a few biological anomalies that might be attributed to ENSO effects: 1) a significant die-off of Common Murres occurred in March-May of 1998, 2) murres and Black-legged Kittiwakes were physiologically stressed during the 1998 breeding season, 3) murres failed to reproduce at one colony in 1998, 4) kittiwake breeding success was lower than usual at colonies in 1998, 5) phenology of breeding was later in 1998 for both murres and kittiwakes, and 6) peaks of forage fish abundance were later in 1998. We presume that seabird anomolies were linked to a reduction or delay in food availability, but the mechanism by which anomalously warm water temperatures in winter affects forage fish availability during the summer breeding season for seabirds is not known.

Dramatic changes in seabird and marine mammal populations in the Gulf of Alaska have been linked to shifts in abundance and composition of forage fish stocks over the past 20 years (Piatt and Anderson, 1996; Anderson et al. 1). Coincident with cyclical fluctuations in seawater temperatures, abundance of several key forage species, including capelin (Mallotus villosus), pricklebacks (mostly Lumpenella longirostris), and Pacific sandfish (Trichodon trichodon) declined precipitously in the late 1970s. Meanwhile populations of large predatory fish, including walleye pollock (Theragra chalcogramma), Pacific cod (Gadus macrocephalus), and several pleuronectids increased dramatically. Correspondingly, seabird diets shifted from mostly capelin in the 1970s to mostly Pacific sand lance (Ammodytes hexapterus) and juvenile pollock by the late 1980s (Piatt and Anderson, 1996). Furthermore, a

This project initiated the process of updating an existing long-term monitoring protocol for black-legged kittiwakes (Rissa tridactyla, henceforth kittiwakes) as part of a larger monitoring effort, the marine predators vital sign program, in Glacier Bay National Park and Preserve (GLBA). Production of the original monitoring protocol for kittiwakes was spearheaded by Hooge et al. (1998) during the 1990s. The purpose of this earlier work was to develop an efficient survey methodology to annually census/monitor kittiwake populations in GLBA at their breeding colonies. Hooge et al. (1998) compared the accuracy of direct visual counts versus counts from photographs. The authors used a repeat survey design and examined the minimum number of replicate surveys necessary for various levels of precision. Additionally, they presented process-based information on the equipment, frames, slide counting, and data management components of the monitoring. In 2011, the Southeast Alaska Inventory and...

Ethnographic studies show that seabirds were regularly eaten in the Western and Northern Isles of Scotland until recent time. Archaeology has shown that seabird were most often exploited between the 4th millennium BC and the17th century AD. Remains of domestic chicken (Gallus gallus) are rare, even on the latest sites. The gannet (Sula bassana) was the species most often captured for food. The great auk (Alca impennis) was important on prehistoric sites.

Reproductive success is one of the most easily-measured and widely studied demographic parameters of colonial nesting seabirds. Nevertheless, factors affecting the sequential stages (egg laying, incubation, chickrearing) of reproductive success are less understood. We investigated the separate sequential stages of reproductive success in piscivorous black-legged kittiwakes (Rissa tridactyla) and thick-billed murres (Uria lomvia) using a 36-year dataset (1975-2010) on the major Pribilof Islands (St. Paul and St. George), which have recently had contrasting population trajectories. Our objectives were to evaluate how the proportion of successful nests varied among stages, and to quantify factors influencing the probability of nest success at each stage in each island. We modeled the probability of nest success at each stage using General Linear Mixed Models incorporating broad-scale and local climate variables, and diet as covariates as well as other measures of reproduction such as timing of breeding and reproductive output in the previous year and previous stage. For both species we found: (1) Success in previous stages of the breeding cycle and success in the prior year better explained overall success than any environmental variables. Phenology was also an important predictor of laying success for kittiwakes. (2) Fledging success was lower when chick diets contained oceanic fish found farther from the colonies and small invertebrates, rather than coastal fish species. (3) Differences in reproductive variables at St. Paul and St. George islands did not correspond to population trends between the two islands. Our results highlight the potential importance of adult condition and annual survival to kittiwake and murre productivity and ultimately, populations. Adult condition carrying over from the previous year ultimately seems to drive annual breeding success in a cascade effect. Furthermore, condition and survival appear to be important contributors to population dynamics at each island. Therefore, adult condition and survival prior to breeding, and factors that influence these parameters such as foraging conditions in the non-breeding season, may be important datasets for understanding drivers of seabird demography at the Pribilof Islands.

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.

As apex predators in marine ecosystems, seabirds may primarily experience climate change impacts indirectly, via changes to their food webs. Observed seabird population declines have been linked to climate-driven oceanographic and food web changes. However, relationships have often been derived from relatively few colonies and consider only sea surface temperature (SST), so important drivers, and spatial variation in drivers, could remain undetected. Further, explicit climate change projections have rarely been made, so longer-term risks remain unclear. Here, we use tracking data to estimate foraging areas for 11 black-legged kittiwake Rissa tridacty la colonies in the UK and Ireland, thus reducing reliance on single colonies and allowing calculation of colony-specific oceanographic conditions. We use mixed models to consider how SST, the potential energy anomaly (indicating density stratification strength) and the timing of seasonal stratification influence kittiwake productivity. Across all colonies, higher breeding success was associated with weaker stratification before breeding and lower SSTs during the breeding season. Eight colonies with sufficient data were modelled individually: higher productivity was associated with later stratification at 3 colonies, weaker stratification at 2, and lower SSTs at one, whilst 2 colonies showed no significant relationships. Hence, key drivers of productivity varied among colonies. Climate change projections, made using fitted models, indicated that breeding success could decline by 21 to 43% between 1961−90 and 2070−99. Climate change therefore poses a longer-term threat to kittiwakes, but as this will be mediated via availability of key prey species, other marine apex predators could also face similar threats.

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.

Food availability comprises a complex interaction of factors that integrates abundance, taxonomic composition, accessibility, and quality of the prey base. The relationship between food availability and reproductive performance can be assessed via the nutritional stress (NSH) and junkfood (JFH) hypotheses. With respect to reproductive success, NSH posits that a deficiency in any of the aforementioned metrics can have a deleterious effect on a population via poor reproductive success. JFH, a component of NSH, posits specifically that it is a decline in the quality of food (i.e. energy density and lipid content) that leads to poor reproductive success. We assessed each in relation to reproductive success in a piscivorous seabird, the black-legged kittiwake Rissa tridactyla. We measured productivity, taxonomic composition, frequency, size, and quality of meals delivered to nestlings from 1996 to 1999 at 6 colonies in Alaska, USA, 3 each in Prince William Sound and Lower Cook Inlet. Productivity varied widely among colony-years. Pacific herring Clupea pallasi, sand lance Ammodytes hexapterus, and capelin Mallotus villosus comprised ca. 80% of the diet among colonyyears, and each was characterized by relatively high energy density. Diet quality for kittiwakes in this region therefore remained uniformly high during this study. Meal delivery rate and meal size were quite variable among colony-years, however, and best explained the variability in productivity. Parent kittiwakes appeared to select prey that were energy dense and that maximized the biomass provisioned to broods. While these results fail to support JFH, they do provide substantial support for NSH.

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.

Both within and among seabird species, different aspects of breeding biology may respond to changes in prey availability in distinct ways, and the identification of species-specific breeding parameters that are sensitive to food availability is useful for monitoring purposes. We present data from a 5-year study (1995)(1996)(1997)(1998)(1999) of the breeding ecology of Horned Puffins (Fratercula corniculata) in Alaska. The El Niño -Southern Oscillation event of 1997-1998 provided an opportunity to examine the sensitivity of various breeding parameters to a reduction in prey availability caused by the anomalous oceanographic conditions of 1998. Horned Puffins were able to maintain high fledging success (83-97%) over the 5 years of the study, despite the poor local feeding conditions in 1998. The rate of increase in chick mass was lowest in 1998, and evidence suggests that chicks also fledged at the youngest ages in that year. The impacts of reduced food availability on growth varied among body structures, suggesting differential allocation of energy and nutrients. There was no variation among years in either chick diet or the mass of food loads delivered by adults. We suggest that rates of chick growth, specifically mass increase, may be a good parameter to measure for use in monitoring Horned Puffins.

We used a supplemental-feeding experiment, the doubly labeled water technique, and a model-selection approach based upon the Akaike Information Criterion to examine effects of food availability on energy expenditure rates of Black-legged Kittiwakes (Rissa tridactyla) raising young. Energy expenditure rates of supplementally fed females (n = 14) and males (n = 16) were 34 and 20% lower than those of control females (n = 14) and males (n = 18), respectively. Energy expenditure rates of females were more responsive to fluctuations in food availability than those of males. Fed males likely expended more energy while off the nest than fed females, possibly because of nest defense. Energy expenditure rates of fed kittiwakes were similar to values reported for kittiwakes that were either not raising young or not foraging. Parent kittiwakes, therefore, adjusted parental effort in response to variation in breeding conditions due to changes in food availability. Adjustments in reproductive effort in response to variable foraging conditions may have significant effects on the survival and productivity of individuals, and thus provide substantial fitness benefits for long-lived seabirds such as Black-legged Kittiwakes.

We fed Herring Clupea harengus to pairs of Black-legged Kittiwakes Rissa tridactyla throughout the breeding season in two years at a colony in the northern Gulf of Alaska. We measured responses to supplemental feeding in a wide array of breeding parameters to gauge their relative sensitivity to food supply, and thus their potential as indicators of natural foraging conditions. Conventional measures of success (hatching, fledging and overall productivity) were more effective as indicators of food supply than behavioural attributes such as courtship feeding, chick provisioning rates and sibling aggression. However, behaviour such as nest relief during incubation and adult attendance with older chicks were also highly responsive to supplemental food and may be useful for monitoring environmental conditions in studies of shorter duration. On average, the chick-rearing stage contained more sensitive indicators of food availability than prelaying or incubation stages. Overall, rates of hatching and fledging success, and the mean duration of incubation shifts were the most food-sensitive parameters studied.

Seabirds are suggested as useful indicators of the abundance and distribution of various of their prey species as well as indicators of changes in oceanic conditions. Additionally, it is shown that seabirds with varying foraging methods are affected differently, with respect to breeding success and by changes in prey numbers and availability.  Diets of three seabirds changed dramatically over a two-year period with respect to numbers species, frequency of occurrence, and length of prey. A concomitant reduction n breeding success, by up to 90%, was noted for surface-feeding Black legged Kittiwakes and Glaucous-winged Gulls, but not for diving Tufted Puffins.  Concurrent sampling of fish prey populations revealed stratification and depression of stocks in the second year of study of the predominant prey eaten in the first year of study. Possible explanations of changes in fish stocks are lower sea temperatures the second year and seawater dilution in the first year.

Seabirds are among the best biological indicator species of the health of the ocean. They are colonial; therefore they are visible. They  usually return to the same sites each year to nest, and in this way are not geographically or temporally patchy as are other oceanic organisms. At seabird colonies, long-term studies can easily be conducted. Annual numbers of breeders and measures of reproductive success such as laying, hatching, or fledging success, or chick growth are good indices of population trends and stability, and can model population parameters of their prey. Other indicator measurements are  weights of adults, of eggs, and fledglings. Various measures of seabird prey can be made and changes in prey numbers, frequency, and weight will give a good indication what might be occurring in the prey base and thus in the surrounding oceanic environment. In this way, seabirds can be used as proxies for oceanic conditions, and can, for instance, predict events such as El Niňo or a collapse of a fishery long before they are verified by others.

Seabirds are central place foragers during the breeding season and, as marine food resources are often patchily distributed, flexibility in foraging behaviour may be important in maintaining prey delivery rates to chicks. We developed a methodological approach using a combination of GPS data loggers and temperature-depth recorders that allowed us to describe the behaviour of surface-feeding seabirds. Specifically, we tested whether differences in foraging behaviour of black-legged kittiwakes Rissa tridactyla could be linked with reproductive success by comparing 2 consecutive years at 2 sites. At Rathlin Island (Northern Ireland) during 2010, foraging differed markedly from that during 2009 and from that at Lambay Island (Republic of Ireland) during both years. Birds exhibited foraging trips of greater duration, travelled a greater total distance, spent more time in transit and spent longer recuperating on the surface of the water. This notable shift was associated with a decline in breeding success, with greater loss of eggs to predation and lower prey delivery rates, resulting in the starvation of 15% of chicks. We suggest that food resources were reduced or geographically less accessible during 2010, with suitable foraging areas located further from the colony. Birds did not invest greater amounts of time attempting to catch prey. Thus, our results indicate that kittiwakes at Rathlin modulated their foraging behaviour not by increasing foraging effort through feeding more intensively within prey patches but by extending their range to increase the probability of encountering more profitable prey patches.

AяѠѡџюѐѡ.-In spite of their putative importance in the evolution of certain traits (e.g., nocturnality, coloniality, cliff nesting), the eff ects of aerial predators on behavior of adult seabirds at colonies have been poorly investigated. We hypothesized that TuĞ ed Puffi ns (Fratercula cirrhata) respond to danger posed by aerial predators by modifying their behavior to mitigate danger. We observed TuĞ ed Puffi ns making repeated colony fl y-ins and departures and characterized (1) the timing of this behavior, (2) the activity rate (number of birds arriving or departing), and (3) the risk-level of activity, with respect to predation danger posed by Bald Eagles (Haliaeetus leucocephalus) and Peregrine Falcons (Falco peregrinus). As we predicted, we found that TuĞ ed Puffi ns (1) dilute danger by synchronizing their fl y-in and departure activities, (2) reduce fl y-in and departure activity rates when predators are present, and (3) switch to lower-risk fl y-in activities (e.g., staying over water where they have an escape route from an aerial aĴ ack) when predators are present.

Background: Windscapes affect energy costs for flying animals, but animals can adjust their behavior to accommodate wind-induced energy costs. Theory predicts that flying animals should decrease air speed to compensate for increased tailwind speed and increase air speed to compensate for increased crosswind speed. In addition, animals are expected to vary their foraging effort in time and space to maximize energy efficiency across variable windscapes. Results: We examined the influence of wind on seabird (thick-billed murre Uria lomvia and black-legged kittiwake Rissa tridactyla) foraging behavior. Airspeed and mechanical flight costs (dynamic body acceleration and wing beat frequency) increased with headwind speed during commuting flights. As predicted, birds adjusted their airspeed to compensate for crosswinds and to reduce the effect of a headwind, but they could not completely compensate for the latter. As we were able to account for the effect of sampling frequency and wind speed, we accurately estimated commuting flight speed with no wind as 16.6 ms −1 (murres) and 10.6 ms −1 (kittiwakes). High winds decreased delivery rates of schooling fish (murres), energy (murres) and food (kittiwakes) but did not impact daily energy expenditure or chick growth rates. During high winds, murres switched from feeding their offspring with schooling fish, which required substantial above-water searching, to amphipods, which required less above-water searching. Conclusions: Adults buffered the adverse effect of high winds on chick growth rates by switching to other food sources during windy days or increasing food delivery rates when weather improved.

In his seminal paper about using seabirds as indicators of marine food supplies, Cairns (1987, Biol Oceanogr 5:261-271) 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'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 'loafing' 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 'seabirds as indicators' are discussed.

Overfishing of predatory fish has contributed to an increase in forage-fish stocks. At the same time, a rising demand for forage fish to supply fish meal markets, in combination with climate change, has put strong pressure on these stocks, and this, in turn, has had an impact on marine top predators. We examined how inter-annual variation in food quality (sprat Sprattus sprattus weight-at-age) and quantity (sprat abundance) influenced Baltic Sea common murres Uria aalge during chick-rearing. Fledging success, i.e. survival from hatching to fledging, showed a positive relationship with food quality, but we found no effect of food quantity. We found no relationship between food quality and parental behaviour or chick feeding parameters, but a negative relationship between food quantity and trip duration. Our data indicate that there was room for parental birds to increase their effort to compensate for reduced food quality, but we found no signs of such compensation. We analysed different types of fish and seabird life-history data to separate effects of food quantity and quality on a top predator. Understanding such effects can contribute to clarifying causes and consequences for observed changes in life-history parameters and population dynamics of top predators.