The incredible shrinking puffin: Decreasing size and increasing proportional bill size of Atlantic puffins nesting at Machias Seal Island.

Climate change imposes physiological constraints on organisms particularly through changing thermoregulatory requirements. Bergmann's and Allen's rules suggest that body size and the size of thermoregulatory structures differ between warm and cold locations, where body size decreases with temperature and thermoregulatory structures increase. However, phenotypic plastic responses to malnutrition during development can result in the same patterns while lacking fitness benefits. The Gulf of Maine (GOM), located at the southern end of the Labrador current, is warming faster than most of the world's oceans, and many of the marine species that occupy these waters exist at the southern edge of their distributions including Atlantic puffins (Fratercula arctica; hereafter "puffin"). Monitoring of puffins in the GOM, at Machias Seal Island (MSI), has continued annually since 1995. We asked whether changes in adult puffin body size and the proportional size of bill to body have changed with observed rapid ocean warming. We found that the size of fledgling puffins is negatively related to sea surface temperature anomalies (warm conditions = small fledgers), adult puffin size is related to fledgling size (small fledgers = small adults), and adult puffins have decreased in size in recent years in response to malnutrition during development. We found an increase in the proportional size of bill to wing chord, likely in response to some mix of malnutrition during development and increasing air temperatures. Although studies have assessed clinal variation in seabird morphology with temperature, this is the first study addressing changes in seabird morphology in relation to ocean warming. Our results suggest that puffins nesting in the GOM have morphological plasticity that may help them acclimate to ocean warming.

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.

Foot web pentosidine does not covary strongly with age in four species of wild seabirds.

Age is an important parameter for a variety of ecological applications, including population viability analyses, contaminants monitoring and targeting of individuals for conservation. While many organisms can be aged by annual rings, dentition and other techniques (i.e., fish otoliths, clam growth rings, mammal tooth wear), there are no minimally invasive biomarkers for accurately aging birds in the wild. For the past century, banding has been the only way to identify a bird of known age, which requires continuous effort on a large scale with possibly low return rates. Recent studies have identified pentosidine as a potential biomarker of chronological aging in several bird species. To test this idea in four species of long-lived seabirds, we collected skin biopsies from the foot webs of previously banded, known-age seabirds: black-legged kittiwakes (Rissa tridactyla; 0-19 y old), Atlantic puffins (Fratercula arctica; 5-26 y old), razorbills (Alca torda; 0-15 d old) and thick-billed murres (Uria lomvia; 0-35 y old). Foot web samples were specifically chosen because this was the least invasive site for substantial skin biopsy. Samples were analysed with high performance liquid chromatography to quantify pentosidine levels. Collagen levels were estimated through hydroxyproline assays to normalize pentosidine content across individuals. Kittiwakes displayed a weak correlation (r2 = 0.20) between age and pentosidine/collagen. Puffins (adults only, r2 = 0.02), razorbills (chicks only, r2 = 0.08), and murres (adults, r2 = 0.04) did not show any associations with age. We concluded that pentosidine content in the foot web does not appear to be a reliable method for aging seabirds in the wild. An absence of change in pentosidine in the foot web with age is further evidence that long-lived seabirds may maintain physiological performance into old age.

It's about time: A synthesis of changing phenology in the Gulf of Maine ecosystem.

The timing of recurring biological and seasonal environmental events is changing on a global scale relative to temperature and other climate drivers. This study considers the Gulf of Maine ecosystem, a region of high social and ecological importance in the Northwest Atlantic Ocean and synthesizes current knowledge of (a) key seasonal processes, patterns, and events; (b) direct evidence for shifts in timing; (c) implications of phenological responses for linked ecological-human systems; and (d) potential phenology-focused adaptation strategies and actions. Twenty studies demonstrated shifts in timing of regional marine organisms and seasonal environmental events. The most common response was earlier timing, observed in spring onset, spring and winter hydrology, zooplankton abundance, occurrence of several larval fishes, and diadromous fish migrations. Later timing was documented for fall onset, reproduction and fledging in Atlantic puffins, spring and fall phytoplankton blooms, and occurrence of additional larval fishes. Changes in event duration generally increased and were detected in zooplankton peak abundance, early life history periods of macro-invertebrates, and lobster fishery landings. Reduced duration was observed in winter-spring ice-affected stream flows. Two studies projected phenological changes, both finding diapause duration would decrease in zooplankton under future climate scenarios. Phenological responses were species-specific and varied depending on the environmental driver, spatial, and temporal scales evaluated. Overall, a wide range of baseline phenology and relevant modeling studies exist, yet surprisingly few document long-term shifts. Results reveal a need for increased emphasis on phenological shifts in the Gulf of Maine and identify opportunities for future research and consideration of phenological changes in adaptation efforts.

Ocean-wide Drivers of Migration Strategies and Their Influence on Population Breeding Performance in a Declining Seabird.

Which factors shape animals' migration movements across large geographical scales, how different migratory strategies emerge between populations, and how these may affect population dynamics are central questions in the field of animal migration [1] that only large-scale studies of migration patterns across a species' range can answer [2]. To address these questions, we track the migration of 270 Atlantic puffins Fratercula arctica, a red-listed, declining seabird, across their entire breeding range. We investigate the role of demographic, geographical, and environmental variables in driving spatial and behavioral differences on an ocean-basin scale by measuring puffins' among-colony differences in migratory routes and day-to-day behavior (estimated with individual daily activity budgets and energy expenditure). We show that competition and local winter resource availability are important drivers of migratory movements, with birds from larger colonies or with poorer local winter conditions migrating further and visiting less-productive waters; this in turn led to differences in flight activity and energy expenditure. Other behavioral differences emerge with latitude, with foraging effort and energy expenditure increasing when birds winter further north in colder waters. Importantly, these ocean-wide migration patterns can ultimately be linked with breeding performance: colony productivity is negatively associated with wintering latitude, population size, and migration distance, which demonstrates the cost of competition and migration on future breeding and the link between non-breeding and breeding periods. Our results help us to understand the drivers of animal migration and have important implications for population dynamics and the conservation of migratory species.

From puffins to plankton: a DNA-based analysis of a seabird food chain in the northern Gulf of Maine.

The predator-prey interactions within food chains are used to both characterize and understand ecosystems. Conventional methods of constructing food chains from visual identification of prey in predator diet can suffer from poor taxonomic resolution, misidentification, and bias against small or completely digestible prey. Next-generation sequencing (NGS) technology has become a powerful tool for diet reconstruction through barcoding of DNA in stomach content or fecal samples. Here we use multi-locus (16S and CO1) next-generation sequencing of DNA barcodes on the feces of Atlantic puffin (Fratercula arctica) chicks (n=65) and adults (n=64) and the stomach contents of their main prey, Atlantic herring (Clupea harengus, n=44) to investigate a previously studied food chain. We compared conventional and molecular-derived chick diet, tested the similarity between the diets of puffin adults and chicks, and determined whether herring prey can be detected in puffin diet samples. There was high variability in the coverage of prey groups between 16S and CO1 markers. We identified more unique prey with our 16S compared to CO1 barcoding markers (51 and 39 taxa respectively) with only 12 taxa identified by both genes. We found no significant difference between the 16S-identified diets of puffin adults (n=17) and chicks (n=41). Our molecular method is more taxonomically resolved and detected chick prey at higher frequencies than conventional field observations. Many likely planktonic prey of herring were detected in feces from puffin adults and chicks, highlighting the impact secondary consumption may have on the interpretation of molecular dietary analysis. This study represents the first simultaneous molecular investigation into the diet of multiple components of a food chain and highlights the utility of a multi-locus approach to diet reconstruction that is broadly applicable to food web analysis.

Contaminants in magnificent frigatebird eggs from Barbuda, West Indies.

We examined persistent organic pollutant and trace element concentrations in eggs of magnificent frigatebirds (Fregata magnificens), a species of conservation concern breeding in the West Indies. Despite that frigatebirds feed at high positions in tropical marine food chains, we detected low levels of most contaminants, suggesting limited contamination of their prey.

Recent Bayesian stable-isotope mixing models are highly sensitive to variation in discrimination factors.

Stable isotopes are now used widely in ecological studies, including diet reconstruction, where quantitative inferences about diet composition are derived from the use of mixing models. Recent Bayesian models (MixSIR, SIAR) allow users to incorporate variability in discrimination factors (delta13C or delta15N), or the amount of change in either delta13C or delta15N between prey and consumer, but to date there has been no systematic assessment of the effect of variation in delta13C or delta15N on model outputs. We used whole blood from Common Terns (Sterna hirundo) and muscle from their common prey items (fish and euphausiids) to build a series of mixing models in SIAR (stable isotope analysis in R) using various discrimination factors from the published literature for marine birds. The estimated proportion of each diet component was affected significantly by delta13C or delta15N. We also use recently published stable-isotope data on the reliance of critically endangered Balearic Shearwaters (Puffinus mauretanicus) on fisheries discards to show that discrimination factor choice can have profound implications for conservation and management actions. It is therefore crucial for researchers wishing to use mixing models to have an accurate estimate of delta13C and delta15N, because quantitative diet estimates can help to direct future research or prioritize conservation and management actions.

Mercury concentrations in seabird tissues from Machias Seal Island, New Brunswick, Canada.

Mercury is a pervasive environmental contaminant, the anthropogenic portion of which is increasing globally, and in northeastern North America in particular. Seabirds frequently are used as indicators of the marine environment, including mercury contamination. We analysed paired samples for total mercury (Hg) concentrations in feathers and blood from adult and chick, albumen, and lipid-free yolk of seven seabirds breeding on Machias Seal Island, New Brunswick, Canada - Arctic Tern (Sterna paradisaea), Atlantic Puffin (Fratercula arctica), Common Eider (Somateria mollissima), Common Murre (Uria aalge), Common Tern (Sterna hirundo), Leach's Storm-petrel (Oceanodroma leucorhoa), and Razorbill (Alca torda). We also used stable-isotope ratios of carbon (delta(13)C), and nitrogen (delta(15)N) to evaluate the relationship between carbon source and trophic position and mercury. We found high Hg concentrations across tissue types in Leach's Storm-petrels, and Razorbills, with lower concentrations in other species, the lowest being in Common Eiders. Storm-petrels prey on mesopelagic fish that accumulate mercury, and Razorbills feed on larger, older fish that bioaccumulate heavy metals. Biomagnification of Hg, or the increase in Hg concentration with trophic position as measured by delta(15)N, was significant and greater in albumen than other tissues, whereas in other tissues, delta(15)N explained little of the overall variation in Hg concentration. Hg concentrations in egg components are higher on Machias Seal Island than other sites globally and in the Gulf of Maine region, but only for some species. Further detailed investigations are required to determine the cause of this trend.

Total and methyl mercury concentrations in seabird feathers and eggs.

Seabirds are used frequently as indicators of mercury contamination in marine ecosystems, but few studies have examined the forms of mercury found in seabird tissues. Here we compare concentrations of total and organic mercury in feathers (n=5) of six sympatric nesting seabirds and in egg components of Leach's storm-petrels from Machias Seal Island, New Brunswick, Canada, during the 2006 breeding season. Essentially all (82-133%) mercury found in seabird feathers and egg components was methyl mercury, with no interspecific differences in percentage methyl mercury. This pattern is consistent with the hypothesis that feather-molt and egg production eliminate toxic methyl mercury, while inorganic forms from demethylation in the liver remain in internal tissues. Additional studies across more species, and comparisons with percentage methyl mercury in internal tissues, are required to validate this theory.

High within-individual variation in total mercury concentration in seabird feathers.

To our knowledge, no rigorous assessment of the variation in mercury concentrations within individual seabirds has been made using multiple body feathers. We analyzed five feathers from individual Arctic terns (Sterna paradisaea Pontoppidan), common terns (Sterna hirundo L.), and Leach's storm-petrels (Oceanodroma leucorhoa Veillot) and found levels of within-individual variability higher than population or time-series variation. Using a randomization procedure, we found a large range of possible mercury concentrations if only one feather per individual had been sampled. Researchers should report within-individual variability in future studies.

Thresholds in songbird occurrence in relation to landscape structure.

Theory predicts the occurrence of threshold levels of habitat in landscapes, below which ecological processes change abruptly. Simulation models indicate that below critical thresholds, fragmentation of habitat influences patch occupancy by decreasing colonization rates and increasing rates of local extinction. Uncovering such putative relationships is important for understanding the demography of species and in developing sound conservation strategies. Using segmented logistic regression, we tested for thresholds in occurrence of 15 bird species as a function of the amount of suitable habitat at multiple scales (150-2000-m radii). Suitable habitat was defined quantitatively based on previously derived, spatially explicit distribution models for each species. The occurrence of 10 out of 15 species was influenced by the amount of habitat at a landscape scale (>or=500-m radius). Of these species all but one were best predicted by threshold models. Six out of nine species exhibited asymptotic thresholds; the effects of habitat loss intensified at low amounts of habitat in a landscape. Landscape thresholds ranged from 8.6% habitat to 28.7% (x= 18.5 +/- 2.6%[95% CI]). For two species landscape thresholds coincided with sensitivity to fragmentation; both species were more likely to occur in large patches, but only when the amount of habitat in a landscape was low. This supports the fragmentation threshold hypothesis. Nevertheless, the occurrence of most species appeared to be unaffected by fragmentation, regardless of the amount of habitat present at landscape extents. The thresholds we identified may be useful to managers in establishing conservation targets. Our results indicate that findings of landscape-scale studies conducted in regions with relatively high proportions of habitat and low fragmentation may not be applicable in regions with low habitat proportions and high fragmentation.

Time and recruitment costs as currencies in manipulation studies on the costs of reproduction.

Life history theory predicts that parents will have lower Darwinian fitness if they tend clutches that are above or below the size they naturally produce. We experimentally tested for relationships between fitness and clutch size in Tree Swallow (Tachycineta bicolor) offspring and parents. Over 130 trios of nests initiated on the same day were randomly divided among reduce (-3 eggs), control (3 eggs picked up and replaced), or add (+3 eggs) manipulations. Pre-manipulation modal clutch size was six eggs (range before manipulations was 1-10; afterwards, it was 1-11). Hatching took longer in larger clutches, but the proportion of eggs hatching and fledging was similar for clutches from 4 to 10, so that clutches of 10 produced the maximum number of fledgling. Parental feeding rates were higher for larger broods, but per capita feeds to nestlings were fewer, and nestlings were smaller. Nonetheless, survival of both young and adults, based on recaptures in subsequent years, was not significantly affected by manipulations. Manipulations also had no significant effect on subsequent reproduction, including the number of fledglings produced by either local recruits or returning breeders. Collectively, our results failed to detect fitness costs associated with tending larger clutches for either parents or the offspring reared and suggested directional selection for larger clutch size. However, because clutches that hatch later produce fewer recruits, the extra days required to lay more eggs and to fledge extra young may eliminate a large part of the advantage that would accrue to parents producing enlarged clutches. For example, our data suggest that there may be less than a 16% benefit to producing nine instead of six eggs, rather than 50%, as is suggested by experimentally manipulated egg numbers alone. Thus, time, rather than costs of reproduction, may be the crucial constraint selecting against Tree Swallows laying larger clutches.

Independent effects of fragmentation on forest songbirds: an organism-based approach.

The degree to which spatial patterns influence the dynamics and distribution of populations is a central question in ecology. This question is even more pressing in the context of rapid habitat loss and fragmentation, which threaten global biodiversity. However, the relative influence of habitat loss and landscape fragmentation, the spatial patterning of remaining habitat, remains unclear. If landscape pattern affects population size, managers may be able to design landscapes that mitigate habitat loss. We present the results of a mensurative experiment designed to test four habitat loss vs. fragmentation hypotheses. Unlike previous studies, we measured landscape structure using quantitative, spatially explicit habitat distribution models previously developed for two species: Blackburnian Warbler (Dendroica fusca) and Ovenbird (Seiurus aurocapilla). We used a stratified sampling design that reduced the confounding of habitat amount and fragmentation variables. Occurrence and reoccurrence of both species were strongly influenced by characteristics at scales greater than the individual territory, indicating little support for the random-sample hypothesis. However, the type and spatial extent of landscape influence differed. Both occurrence and reoccurrence of Blackburnian Warblers were influenced by the amount of poor-quality matrix at 300- and 2000-m spatial extents. The occurrence and reoccurrence of Ovenbirds depended on a landscape pattern variable, patch size, but only in cases when patches were isolated. These results support the hypothesis that landscape pattern is important for some species only when the amount of suitable habitat is low. Although theoretical models have predicted such an interaction between landscape fragmentation and composition, to our knowledge this is the first study to report empirical evidence of such nonlinear fragmentation effects. Defining landscapes quantitatively from an organism-based perspective may increase power to detect fragmentation effects, particularly in forest mosaics where boundaries between patches and matrix are ambiguous. Our results indicate that manipulating landscape pattern may reduce negative impacts of habitat loss for Ovenbird, but not Blackburnian Warbler. We emphasize that most variance in the occurrence of both species was explained by local scale or landscape composition variables rather than variables reflecting landscape pattern.