Small tube-nosed seabirds fledge on the full moon and throughout the lunar cycle.

Many seabirds are attracted to anthropogenic light, and the risk is greater for recent fledglings. Moon phase predicts the probability of stranding (fewer birds strand on the full moon), but it remains uncertain whether moon phase is associated with when young seabirds fledge. Fledging behaviour of nocturnal, burrowing seabirds can be difficult to monitor using traditional methods but can provide insight into environmental factors that influence the risk of stranding. We used passive integrated transponder tags to monitor the fledging dates and times of Leach's storm-petrel (Hydrobates leucorhous) chicks across four breeding seasons (2017, 2018, 2021, 2022) at a major colony in Newfoundland and Labrador, Canada. We also assessed whether moon phase and incident illumination related to fledging date and time. The median fledge time was 1.6 h after sunset (0.6-11.7 h). The median fledge date was 10 October, and fledging dates ranged from 13 September to 13 November. Most importantly, moon phase was not associated with the time and date that Leach's storm-petrel chicks fledged. These results suggest that recently fledged storm-petrels are less attracted to anthropogenic light during high levels of natural illumination, which could indicate periods of higher stranding risk and help concentrate conservation efforts.

Female and male Leach's Storm Petrels (Hydrobates leucorhous) pursue different foraging strategies during the incubation period.

Reproduction in procellariiform birds is characterized by a single egg clutch, slow development, a long breeding season and obligate biparental care. Female Leach's Storm Petrels Hydrobates leucorhous, nearly monomorphic members of this order, produce eggs that are between 20 and 25% of adult body weight. We tested whether female foraging behaviour differs from male foraging behaviour during the ~ 44-day incubation period across seven breeding colonies in the Northwest Atlantic. Over six breeding seasons, we used a combination of Global Positioning System and Global Location Sensor devices to measure characteristics of individual foraging trips during the incubation period. Females travelled significantly greater distances and went farther from the breeding colony than did males on individual foraging trips. For both sexes, the longer the foraging trip, the greater the distance. Independent of trip duration, females travelled farther, and spent a greater proportion of their foraging trips prospecting widely as defined by behavioural categories derived from a Hidden Markov Model. For both sexes, trip duration decreased with date. Sex differences in these foraging metrics were apparently not a consequence of morphological differences or spatial segregation. Our data are consistent with the idea that female foraging strategies differed from male foraging strategies during incubation in ways that would be expected if females were still compensating for egg formation.

Climate oscillations drive millennial-scale changes in seabird colony size.

Seabird population size is intimately linked to the physical, chemical, and biological processes of the oceans. Yet, the overall effects of long-term changes in ocean dynamics on seabird colonies are difficult to quantify. Here, we used dated lake sediments to reconstruct ~10,000-years of seabird dynamics in the Northwest Atlantic to determine the influences of Holocene-scale climatic oscillations on colony size. On Baccalieu Island (Newfoundland and Labrador, Canada)-where the world's largest colony of Leach's storm-petrel (Hydrobates leucorhous Vieillot 1818) currently breeds-our data track seabird colony growth in response to warming during the Holocene Thermal Maximum (ca. 9000 to 6000 BP). From ca. 5200 BP to the onset of the Little Ice Age (ca. 550 BP), changes in colony size were correlated to variations in the North Atlantic Oscillation (NAO). By contrasting the seabird trends from Baccalieu Island to millennial-scale changes of storm-petrel populations from Grand Colombier Island (an island in the Northwest Atlantic that is subjected a to different ocean climate), we infer that changes in NAO influenced the ocean circulation, which translated into, among many things, changes in pycnocline depth across the Northwest Atlantic basin where the storm-petrels feed. We hypothesize that the depth of the pycnocline is likely a strong bottom-up control on surface-feeding storm-petrels through its influence on prey accessibility. Since the Little Ice Age (LIA), the effects of ocean dynamics on seabird colony size have been altered by anthropogenic impacts. Subsequently, the colony on Baccalieu Island grew at an unprecedented rate to become the world's largest resulting from favorable conditions linked to climate warming, increased vegetation (thereby nesting habitat), and attraction of recruits from other colonies that are now in decline. We show that although ocean dynamics were an important driver of seabird colony dynamics, its recent influence has been modified by human interference.

Influence of offshore oil and gas structures on seascape ecological connectivity.

Offshore platforms, subsea pipelines, wells and related fixed structures supporting the oil and gas (O&G) industry are prevalent in oceans across the globe, with many approaching the end of their operational life and requiring decommissioning. Although structures can possess high ecological diversity and productivity, information on how they interact with broader ecological processes remains unclear. Here, we review the current state of knowledge on the role of O&G infrastructure in maintaining, altering or enhancing ecological connectivity with natural marine habitats. There is a paucity of studies on the subject with only 33 papers specifically targeting connectivity and O&G structures, although other studies provide important related information. Evidence for O&G structures facilitating vertical and horizontal seascape connectivity exists for larvae and mobile adult invertebrates, fish and megafauna; including threatened and commercially important species. The degree to which these structures represent a beneficial or detrimental net impact remains unclear, is complex and ultimately needs more research to determine the extent to which natural connectivity networks are conserved, enhanced or disrupted. We discuss the potential impacts of different decommissioning approaches on seascape connectivity and identify, through expert elicitation, critical knowledge gaps that, if addressed, may further inform decision making for the life cycle of O&G infrastructure, with relevance for other industries (e.g. renewables). The most highly ranked critical knowledge gap was a need to understand how O&G structures modify and influence the movement patterns of mobile species and dispersal stages of sessile marine species. Understanding how different decommissioning options affect species survival and movement was also highly ranked, as was understanding the extent to which O&G structures contribute to extending species distributions by providing rest stops, foraging habitat, and stepping stones. These questions could be addressed with further dedicated studies of animal movement in relation to structures using telemetry, molecular techniques and movement models. Our review and these priority questions provide a roadmap for advancing research needed to support evidence-based decision making for decommissioning O&G infrastructure.

Striking centennial-scale changes in the population size of a threatened seabird.

Many animal populations are under stress and declining. For numerous marine bird species, only recent or sparse monitoring data are available, lacking the appropriate temporal perspective needed to consider natural, long-term population dynamics when developing conservation strategies. Here, we use a combination of established palaeoenvironmental approaches to examine the centennial-scale dynamics of the world's largest colony (representing approx. 50% of the global population) of the declining and vulnerable Leach's Storm-petrel (Hydrobates leucorhous). By reconstructing the last approximately 1700 years of the colony's population trends, we corroborate recent surveys indicating rapid declines since the 1980s. More surprisingly, however, was that the colony size was smaller and has changed strikingly in the past, even prior to the introduction of human stressors. Our results challenge notions that very large colonies are generally stable in the absence of anthropogenic pressures and speak to an increasingly pressing need to better understand inter-colony movement and recruitment when inferring range- and species-wide trends. While the recently documented decline in storm-petrels clearly warrants conservation concern, we show that colony size was consistently much lower in the past and changed markedly in the absence of major anthropogenic activity. In response, we emphasize the need for enlarged protected area networks to maintain natural population cycles, coupled with continued research to identify the driver(s) of the current global seabird decline.

Circumpolar dynamics of a marine top-predator track ocean warming rates.

Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long-lived, wide-ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black-legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea-surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large-scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom-up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large-scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.

Feather corticosterone during non-breeding correlates with multiple measures of physiology during subsequent breeding in a migratory seabird.

Carry-over effects in migratory birds are likely mediated by physiological processes that are activated in response to environmental variation. Such processes affect body condition and/or reproductive success, and can include corticosterone (CORT) because this hormone responds to environmental stressors and influences energy balance. Few studies have considered how CORT levels during non-breeding relate to a broader physiological profile during subsequent breeding, and fewer still have considered measures other than body condition. To explore CORT's potential role in carry-over effects, we investigated the relationship between CORT and foraging ecology of northern gannets (Morus bassanus) during the non-breeding period, and tested for associations between these factors and variation in a suite of physiological and biochemical metrics during subsequent breeding. Northern gannets are the largest seabird top predator in the North Atlantic and were among the hardest hit by the Deepwater Horizon oil blowout in the Gulf of Mexico in 2010. We used light-level geolocators to confirm winter origins of individuals in our study. No interrelationships were found among levels of CORT from feathers grown during non-breeding (CORTf) and variation in foraging ecology, measured by stable isotopes of carbon (δ13C) and nitrogen (δ15N) from the same feathers. CORTf was correlated negatively with hematocrit and positively with triglyceride measured during subsequent incubation, and explained more variation in these variables than did body mass during incubation. These findings provide support for the hypothesis that energy management, measured using CORTf, during non-breeding carries over to influence physiological measures other than body condition. Gannets that previously wintered within the Gulf of Mexico in the years following the Deepwater Horizon oil blowout had higher levels of CORTf compared to birds that wintered along the Atlantic coast, suggesting an increased energetic cost associated with visiting the Gulf of Mexico. Our results indicate that CORT during non-breeding is associated with a broader physiological profile during subsequent breeding than previously reported in birds.

Challenges to Oil Spill Assessment for Seabirds in the Deep Ocean.

We synthesize impediments for evaluating effects to seabirds from open ocean hydrocarbon releases. Effects on seabirds from ship discharges, spills, and well blowouts often are poorly detected and monitored far from land. Regulatory regimes for ocean spills can result in monitoring efforts that are not entirely transparent. We illustrate how interdisciplinary technologies address deficits that hamper individual or population level assessments for seabirds, and we demonstrate where emerging technologies might be engaged to bridge gaps in oil spill monitoring. Although acute mortality from direct oil exposure poses the greatest risk to seabirds, other hazards from light-attraction, flaring, collisions, chronic pollution, and hydrocarbon inhalation around oil infrastructure also may induce bird mortality in the deep ocean.

Reduction of coastal lighting decreases seabird strandings.

Artificial light at night (ALAN) is negatively impacting numerous species of nocturnally active birds. Nocturnal positive phototaxis, the movement towards ALAN, is exhibited by many marine birds and can result in stranding on land. Seabird species facing major population declines may be most at risk. Leach's Storm-Petrels (Hydrobates leucorhous) are small, threatened seabirds with an extensive breeding range in the North Atlantic and North Pacific Oceans. The Atlantic population, which represents approximately 40-48% of the global population, is declining sharply. Nocturnal positive phototaxis is considered to be a key contributing factor. The Leach's Storm-Petrel is the seabird species most often found stranded around ALAN in the North Atlantic, though there is little experimental evidence that reduction of ALAN decreases the occurrence of stranded storm-petrels. During a two-year study at a large, brightly illuminated seafood processing plant adjacent to the Leach's Storm-Petrel's largest colony, we compared the number of birds that stranded when the lights at the plant were on versus significantly reduced. We recorded survival, counted carcasses of adults and juveniles, and released any rescued individuals. Daily morning surveys revealed that reducing ALAN decreased strandings by 57.11% (95% CI: 39.29% - 83.01%) per night and night surveys revealed that reducing ALAN decreased stranding of adult birds by 11.94% (95% CI: 3.47% - 41.13%) per night. The peak stranding period occurred from 25 September to 28 October, and 94.9% of the birds found during this period were fledglings. These results provide evidence to support the implementation of widespread reduction and modification of coastal artificial light as a conservation strategy, especially during avian fledging and migration periods.

Quantifying inter-annual variability on the space-use of parental Northern Gannets (Morus bassanus) in pursuit of different prey types.

Spatial planning for marine areas of multi-species conservation concern requires in-depth assessment of the distribution of predators and their prey. Northern Gannets Morus bassanus are generalist predators that predate several different forage fishes depending on their availability. In the western North Atlantic, gannets employ different dive tactics while in pursuit of different prey types, performing deep, prolonged U-shaped dives when foraging on capelin (Mallotus villosus), and rapid, shallow, V-shaped dives when foraging on larger pelagic fishes. Therefore, much can be inferred about the distribution and abundance of key forage fishes by assessing the foraging behaviour and space-use of gannets. In this study, we aimed to quantify space-use and to determine areas of suitable foraging habitat for gannets in pursuit of different prey types using habitat suitability models and kernel density utilization distributions. We deployed 25 GPS/Time-depth recorder devices on parental Northern Gannets at Cape St. Mary's, Newfoundland, Canada from 2019 to 2021. To assess the influence of environmental variables on gannets foraging for different prey types, we constructed three different habitat suitability models: a U-shaped dive model, and two V-shaped dive models (early and late chick-rearing). Suitable foraging habitat for capelin, deduced by the U-shaped dive model, was defined by coastal, shallow waters with flat relief and sea surface temperatures (SST) of 11-15° C. Suitable habitat for early V-shaped dives was defined by shallow and coastal waters with steep slope and SST of 12-15°C and ~18°C, likely reflecting the variability in environmental preferences of different prey species captured when performing V-shaped dives. Suitable habitat for late V-shaped dives was defined by shallow coastal waters (<100m depth), as well as waters deeper than 200 m, and by SST greater than 16°C. We show that space-use by gannets can vary both within and between years depending on environmental conditions and the prey they are searching for, with consequences for the extent of potential interaction with anthropogenic activities. Further, we suggest regions defined as suitable for U-shaped dives are likely to be critical habitat of multi-species conservation concern, as these regions are likely to represent consistent capelin spawning habitat.

Factors influencing mercury levels in Leach's storm-petrels at northwest Atlantic colonies.

Mercury (Hg) is a globally distributed heavy metal, with negative effects on wildlife. Its most toxic form, methylmercury (MeHg), predominates in aquatic systems. Levels of MeHg in marine predators can vary widely among individuals and populations. Leach's storm-petrels (Hydrobates leucorhous) have elevated levels of Hg but the role of Hg in storm-petrel population declines is unknown. In this study, we used egg and blood samples to study variation in Hg exposure among several northwest Atlantic colonies during breeding seasons, thereby evaluating relative toxicity risk within and among colonies. Total mercury (THg) concentrations were higher with increasing colony latitude, and were more pronounced in blood than in eggs. THg concentrations in blood were mostly associated with low toxicity risk in birds from the southern colonies and moderate risks in birds from the northern colonies; however, those values did not affect hatching or fledging success. THg concentrations in both eggs and blood were positively correlated with δ34S, emphasizing the role of sulfate-reducing bacteria in methylation of THg acquired through marine food webs, which is consistent with enriched δ34S profiles. By associating tracking data from foraging trips with THg from blood, we determined that blood THg levels were higher when storm-petrel's intensive search locations were over deeper waters. We conclude that spatial variation in THg concentrations in Leach's storm-petrels is attributable to differences in ocean depth at foraging locations, both at individual and colony levels. Differences in diet among colonies observed previously are the most likely cause for observed blood THg differences. As one of the few pelagic seabird species breeding in Atlantic Canada, with limited overlap in core foraging areas among colonies, Leach's storm-petrels can be used as biomonitors for less sampled offshore pelagic regions. The global trend in Hg emissions combined with legacy levels warrant continued monitoring for toxicity effects in seabirds.

Foraging range scales with colony size in high-latitude seabirds.

Density-dependent prey depletion around breeding colonies has long been considered an important factor controlling the population dynamics of colonial animals.1-4 Ashmole proposed that as seabird colony size increases, intraspecific competition leads to declines in reproductive success, as breeding adults must spend more time and energy to find prey farther from the colony.1 Seabird colony size often varies over several orders of magnitude within the same species and can include millions of individuals per colony.5,6 As such, colony size likely plays an important role in determining the individual behavior of its members and how the colony interacts with the surrounding environment.6 Using tracking data from murres (Uria spp.), the world's most densely breeding seabirds, we show that the distribution of foraging-trip distances scales to colony size0.33 during the chick-rearing stage, consistent with Ashmole's halo theory.1,2 This pattern occurred across colonies varying in size over three orders of magnitude and distributed throughout the North Atlantic region. The strong relationship between colony size and foraging range means that the foraging areas of some colonial species can be estimated from colony sizes, which is more practical to measure over a large geographic scale. Two-thirds of the North Atlantic murre population breed at the 16 largest colonies; by extrapolating the predicted foraging ranges to sites without tracking data, we show that only two of these large colonies have significant coverage as marine protected areas. Our results are an important example of how theoretical models, in this case, Ashmole's version of central-place-foraging theory, can be applied to inform conservation and management in colonial breeding species.

Global political responsibility for the conservation of albatrosses and large petrels.

Migratory marine species cross political borders and enter the high seas, where the lack of an effective global management framework for biodiversity leaves them vulnerable to threats. Here, we combine 10,108 tracks from 5775 individual birds at 87 sites with data on breeding population sizes to estimate the relative year-round importance of national jurisdictions and high seas areas for 39 species of albatrosses and large petrels. Populations from every country made extensive use of the high seas, indicating the stake each country has in the management of biodiversity in international waters. We quantified the links among national populations of these threatened seabirds and the regional fisheries management organizations (RFMOs) which regulate fishing in the high seas. This work makes explicit the relative responsibilities that each country and RFMO has for the management of shared biodiversity, providing invaluable information for the conservation and management of migratory species in the marine realm.

An investigation of physiological effects of the Deepwater Horizon oil spill on a long-distance migratory seabird, the northern gannet.

Exposure to oil can have long-term impacts on migratory birds. Following the 2010 Deepwater Horizon blowout in the Gulf of Mexico (GOM), we investigated potential impacts of oil exposure on a population of northern gannets (Morus bassanus) that breed on Bonaventure Island (Québec, Canada) and winter in GOM and along the U.S. Atlantic coast (AC). Blood and feather samples were collected from adults previously equipped with geolocators to determine wintering locations. Parent and alkylated polycyclic aromatic hydrocarbons (PAHs); trace metals; stable isotopes of carbon, nitrogen, and hydrogen; and immune, thyroid, steroid, retinoid, and genetic endpoints were measured. PAH and trace metal concentrations did not differ between gannets using different wintering sites. Feather stable isotope values varied significantly between birds from different wintering locations. Gannets wintering in GOM showed higher feather corticosterone and plasma thyroid hormone levels, which may indicate increased energetic demands and/or greater exposure to environmental stressors.

Author Correction: Important At-Sea Areas of Colonial Breeding Marine Predators on the Southern Patagonian Shelf.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Important At-Sea Areas of Colonial Breeding Marine Predators on the Southern Patagonian Shelf.

The Patagonian Shelf Large Marine Ecosystem supports high levels of biodiversity and endemism and is one of the most productive marine ecosystems in the world. Despite the important role marine predators play in structuring the ecosystems, areas of high diversity where multiple predators congregate remains poorly known on the Patagonian Shelf. Here, we used biotelemetry and biologging tags to track the movements of six seabird species and three pinniped species breeding at the Falkland Islands. Using Generalized Additive Models, we then modelled these animals' use of space as functions of dynamic and static environmental indices that described their habitat. Based on these models, we mapped the predicted distribution of animals from both sampled and unsampled colonies and thereby identified areas where multiple species were likely to overlap at sea. Maximum foraging trip distance ranged from 79 to 1,325 km. However, most of the 1,891 foraging trips by 686 animals were restricted to the Patagonian Shelf and shelf slope, which highlighted a preference for these habitats. Of the seven candidate explanatory covariates used to predict distribution, distance from the colony was retained in models for all species and negatively affected the probability of occurrence. Predicted overlap among species was highest on the Patagonian Shelf around the Falkland Islands and the Burdwood Bank. The predicted area of overlap is consistent with areas that are also important habitat for marine predators migrating from distant breeding locations. Our findings provide comprehensive multi-species predictions for some of the largest marine predator populations on the Patagonian Shelf, which will contribute to future marine spatial planning initiatives. Crucially, our findings highlight that spatially explicit conservation measures are likely to benefit multiple species, while threats are likely to impact multiple species.

Foraging areas, offshore habitat use, and colony overlap by incubating Leach's storm-petrels Oceanodroma leucorhoa in the Northwest Atlantic.

Despite their importance in marine food webs, much has yet to be learned about the spatial ecology of small seabirds. This includes the Leach's storm-petrel Oceanodroma leucorhoa, a species that is declining throughout its Northwest Atlantic breeding range. In 2013 and 2014, we used global location sensors to track foraging movements of incubating storm-petrels from 7 eastern Canadian breeding colonies. We determined and compared the foraging trip and at-sea habitat characteristics, analysed spatial overlap among colonies, and determined whether colony foraging ranges intersected with offshore oil and gas operations. Individuals tracked during the incubation period made 4.0 ± 1.4 day foraging trips, travelling to highly pelagic waters over and beyond continental slopes which ranged, on average, 400 to 830 km from colonies. Cumulative travel distances ranged from ~900 to 2,100 km among colonies. While colony size did not influence foraging trip characteristics or the size of areas used at sea, foraging distances tended to be shorter for individuals breeding at the southern end of the range. Core areas did not overlap considerably among colonies, and individuals from all sites except Kent Island in the Bay of Fundy foraged over waters with median depths > 1,950 m and average chlorophyll a concentrations ≤ 0.6 mg/m3. Sea surface temperatures within colony core areas varied considerably (11-23°C), coincident with the birds' use of cold waters of the Labrador Current or warmer waters of the Gulf Stream Current. Offshore oil and gas operations intersected with the foraging ranges of 5 of 7 colonies. Three of these, including Baccalieu Island, Newfoundland, which supports the species' largest population, have experienced substantial declines in the last few decades. Future work should prioritize modelling efforts to incorporate information on relative predation risk at colonies, spatially explicit risks at-sea on the breeding and wintering grounds, effects of climate and marine ecosystem change, as well as lethal and sub-lethal effects of environmental contaminants, to better understand drivers of Leach's storm-petrel populations trends in Atlantic Canada.

Seasonal Variation in Parental Care Drives Sex-Specific Foraging by a Monomorphic Seabird.

Evidence of sex-specific foraging in monomorphic seabirds is increasing though the underlying mechanisms remain poorly understood. We investigate differential parental care as a mechanism for sex-specific foraging in monomorphic Common Murres (Uria aalge), where the male parent alone provisions the chick after colony departure. Using a combination of geolocation-immersion loggers and stable isotopes, we assess two hypotheses: the reproductive role specialization hypothesis and the energetic constraint hypothesis. We compare the foraging behavior of females (n = 15) and males (n = 9) during bi-parental at the colony, post-fledging male-only parental care and winter when parental care is absent. As predicted by the reproductive role specialization hypothesis, we found evidence of sex-specific foraging during post-fledging only, the stage with the greatest divergence in parental care roles. Single-parenting males spent almost twice as much time diving per day and foraged at lower quality prey patches relative to independent females. This implies a potential energetic constraint for males during the estimated 62.8 ± 8.9 days of offspring dependence at sea. Contrary to the predictions of the energetic constraint hypothesis, we found no evidence of sex-specific foraging during biparental care, suggesting that male parents did not forage for their own benefit before colony departure in anticipation of post-fledging energy constraints. We hypothesize that unpredictable prey conditions at Newfoundland colonies in recent years may limit male parental ability to allocate additional time and energy to self-feeding during biparental care, without compromising chick survival. Our findings support differential parental care as a mechanism for sex-specific foraging in monomorphic murres, and highlight the need to consider ecological context in the interpretation of sex-specific foraging behavior.

Constraints on nest site selection: a comparison of predator and flood avoidance in four species of marsh-nesting birds (genera: Catoptrophorus, Larus, Rallus, and Sterna).

Nest site characteristics associated with flood and predator avoidance were compared for four nonpasserine species of marsh-nesting birds: clapper rails, willets, laughing gulls, and common terns. Species with short nests, willets and terns, minimized flood damage by nesting on higher ground than did gulls and rails that build tall nests. Species with dispersed, cryptic nests had taller surrounding grass than did open-nesting colonial species. Total nest height was similar for species with tall nests and for terns whose short nests were elevated by placement on mats of dead Spartina grass. Willets had lower nest heights than the other species, probably because the inverse relation between grass height and ground height in the salt marsh makes it difficult for willets to find sites with high enough ground for flood avoidance while still retaining high enough grass for nest crypticity. Ground height for common terns and grass height for gulls and rails appear to be cues used in nest site selection. Nests of each species in which these characteristics were maximized were more successful in a major tidal flood. Laughing gulls and clapper rails appear to be more specialized salt marsh nesters than the other two species.

Stable isotopic determinations of trophic relationships of great auks.

Stable isotopic ratios of animal tissues are related to those of their foods and can be used in palaeoecological reconstructions, including those of extinct animals. Nitrogen isotopic analyses of marine organisms from coastal Newfoundland and Georges Bank were used to construct a model predicting collagen δ15N values for seabirds feeding at various trophic levels (TL). This model was tested by measuring bone collagen δ15N values of extant alcids from the northwest Atlantic and high Arctic. Isotopic analysis of bone collagen of the extinct great auk (Pinguinus impennis), the last flightless seabird in the northern hemisphere, indicate that this species occupied a trophic continuum from TL3 (crustacean diets) to TL5 (diets of piscivorous fish). We suggest that (a) great auk chicks and juveniles occupied lower trophic levels and probably consumed euphausiids, and (b) great auks fed offspring via regurgitation, as do dovekies (Alle alle), the only extant fully planktivorous alcid in the Atlantic, and unrelated penguins of the southern hemisphere.