John F. Piatt

Using stable isotopes to determine seabird trophic relationships

1. The stable isotopes of nitrogen (δ 15 N) and carbon (δ 13 C) were analysed in 22 species of marine birds from coastal waters of the northeast Pacific Ocean. Analyses confirm that stable nitrogen isotopes can predict seabird trophic positions. 2. Based on δ 15 N analyses, seabird trophic-level inferences generally agree with those of conventional dietary studies, but suggest that lower trophic-level organisms are more important to several seabirds than was recognized previously. 3. Stable-carbon isotope analysis may be a good indicator of inshore vs. offshore feeding preference

Global Seabird Response to Forage Fish Depletion—One-Third for the Birds

One-third of maximum fish biomass must be available for seabirds to sustain high breeding success. Determining the form of key predator-prey relationships is critical for understanding marine ecosystem dynamics. Using a comprehensive global database, we quantified the effect of fluctuations in food abundance on seabird breeding success. We identified a threshold in prey (fish and krill, termed “forage fish”) abundance below which seabirds experience consistently reduced and more variable productivity. This response was common to all seven ecosystems and 14 bird species examined within the Atlantic, Pacific, and Southern Oceans. The threshold approximated one-third of the maximum prey biomass observed in long-term studies. This provides an indicator of the minimal forage fish biomass needed to sustain seabird productivity over the long term.

Benefits and costs of increased levels of corticosterone in seabird chicks.

Seabird chicks respond to food shortages by increasing corticosterone (cort) secretion, which is probably associated with fitness benefits and costs. To examine this, we experimentally increased levels of circulating cort in captive black-legged kittiwake chicks fed ad libitum. We found that cort-implanted chicks begged more frequently and were more aggressive compared to controls. These behavioral modifications must be beneficial to chicks as they facilitate acquisition of food from the parents and might trigger brood reduction and reduced competition for food. Cort-implanted chicks also increased food intake; however, their growth rates were similar to controls. To examine the costs of chronically increased circulating levels of cort, we removed cort implants and, after a 10-day recovery period, tested cognitive abilities of young kittiwakes. We found that the ability of kittiwakes to associate a visual cue with the presence of food in a choice situation was compromised by the experimental elevation of cort during development. To examine the long-term costs of increased levels of cort, 8 months later we tested the performance of the same individuals in a spatial task requiring them to make a detour around a barrier in order to escape from an enclosure. Individuals treated with cort during development took significantly more time to solve this task compared to controls. The results of this study suggest that the adrenocortical response of a developing bird to environmental stressors is associated with both benefits (increased food intake, foraging behavior, and aggression) and costs (low growth efficiency and compromised cognitive abilities later in life). This provides an evolutionary framework for relating juvenile physiological traits to fitness of birds in subsequent life-history stages.

The adrenocortical stress-response of Black-legged Kittiwake chicks in relation to dietary restrictions

Abstract In this study we examined hormonal responses of Black-legged Kittiwake (Rissatridactyla) chicks to experimental variations in energy content and nutritional quality (low or high lipid to protein ratio, LPR) of their food. Starting at the age of 10 days, chicks were fed either high or low LPR fish at 30, 50, 70 and 100% of ad libitum energy intake. After 20 days of treatment, chicks were exposed to a standardized acute handling and restraint stress protocol, where a baseline sample was taken immediately after taking a chick from the nest, and three additional blood samples were taken at intervals up to 50 min. Testosterone and corticosterone titres in plasma were measured via radioimmunoassay. We found that baseline testosterone levels were not significantly affected by the experimental treatments. Food-restricted chicks had elevated baseline and acute stress-induced levels of corticosterone compared to chicks fed ad libitum. An elevation of circulating levels of corticosterone in energetically stressed individuals was further magnified by low nutritional quality of food. Baseline and acute stress-induced corticosterone levels of chicks were negatively correlated with their fat reserves. We conclude that the physiological condition of Black-legged Kittiwake chicks can be assessed reliably by measuring circulating levels of corticosterone. We discuss short- and long-term effects of elevated corticosterone secretion in food-stressed nest-bound chicks.

Proximate composition and energy density of some north pacific forage fishes

Abstract Mature pelagic forage fish species (capelin, sand lance, squid) had greater lipid concentrations than juvenile age-classes of large demersal and pelagic fish species (walleye pollock, Pacific cod, Atka mackerel, greenling, prowfish, rockfish, sablefish). Myctophids preyed on by puffins have at least twice as much lipid per gram compared to mature capelin, sand lance and squid, and an order of magnitude greater lipid concentrations than juvenile forage fish. Energy density of forage fishes was positively correlated with lipid content, and negatively correlated with water, ash-free lean dry mass (mostly protein), and ash contents.

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 lipid-rich 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.

Increasing frequency of plastic particles ingested by seabirds in the subarctic North Pacific

Abstract We examined gut contents of 1799 seabirds comprising 24 species collected in 1988–1990 to assess the types and quantities of plastic particles ingested by seabirds in the subarctic waters of Alaska. Of the 15 species found to ingest plastic, most were surface-feeders (shearwaters, petrels, gulls) or plankton-feeding divers (auklets, puffins). Of 4417 plastic particles examined, 76% were industrial pellets and 21% were fragments of ‘user’ plastic. Ingestion rates varied geographically, but no trends were evident and rates of plastic ingestion varied far more among species within areas than within species among areas. Comparison with similar data from 1968 seabirds comprising 37 species collected in 1969–1977 revealed that plastic ingestion by seabirds has increased significantly during the 10–15-year interval between studies. This was demonstrated by: (i) an increase in the total number of species ingesting plastic; (ii) an increase in the frequency of occurrence of plastic particles within species that ingested plastic; and, (iii) an increase in the mean number of plastic particles ingested by individuals of those species.

MECHANISMS OF POPULATION DIFFERENTIATION IN MARBLED MURRELETS: HISTORICAL VERSUS CONTEMPORARY PROCESSES

Abstract.— Mechanisms of population differentiation in highly vagile species such as seabirds are poorly understood. Previous studies of marbled murrelets (Brachyramphus marmoratus; Charadriiformes: Alcidae) found significant population genetic structure, but could not determine whether this structure is due to historical vicariance (e.g., due to Pleistocene glaciers), isolation by distance, drift or selection in peripheral populations, or nesting habitat selection. To discriminate among these possibilities, we analyzed sequence variation in nine nuclear introns from 120 marbled murrelets sampled from British Columbia to the western Aleutian Islands. Mismatch distributions indicated that murrelets underwent at least one population expansion during the Pleistocene and probably are not in genetic equilibrium. Maximum‐likelihood analysis of allele frequencies suggested that murrelets from “mainland” sites (from the Alaskan Peninsula east) are genetically different from those in the Aleutians and that these two lineages diverged prior to the last glaciation. Analyses of molecular variance, as well as estimates of gene flow derived using coalescent theory, indicate that population genetic structure is best explained by peripheral isolation of murrelets in the Aleutian Islands, rather than by selection associated with different nesting habitats. No isolation‐by‐distance effects could be detected. Our results are consistent with a rapid expansion of murrelets from a single refugium during the early–mid Pleistocene, subsequent isolation and divergence in two or more refugia during the final Pleistocene glacial advance, and secondary contact following retreat of the ice sheets. Population genetic structure now appears to be maintained by distance effects combined with small populations and a highly fragmented habitat in the Aleutian Islands.

Changes in proximate composition and somatic energy content for Pacific sand lance (Ammodytes hexapterus) from Kachemak Bay, Alaska relative to maturity and season

Mean dry-weight energy values of adult Pacific sand lance (Ammodytes hexapterus) peaked in spring and early summer (20.91 kJg−1 for males, 21.08 kJg−1 for females), then declined by about 25% during late summer and fall (15.91 kJg−1 for males, 15.74 kJg−1 for females). Late summer declines in energy density paralleled gonadal development. Gender differences in energy density (males<females) were only apparent from August to October. Adult sand lance spawn in October, entering the winter with close to their minimum whole body energy content. Juvenile sand lance exhibited a relatively constant protein to lipid ratio until they reached 80 mm fork length. Thereafter, relative proportions of protein remained constant while lipid proportions increased significantly. Dry weight energy densities of juveniles increased from a minimum 16.67 kJg−1 to a maximum of 19.68 kJg−1, and (per g) are higher than adults in late summer. The seasonal food value of adult sand lance to predators varies markedly, but maximum energetic value coincides with important feeding periods for marine mammals, fish, and seabirds.

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 predators 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.