Spatially explicit network analysis reveals multi-species annual cycle movement patterns of sea ducks.

Conservation of long-distance migratory species poses unique challenges. Migratory connectivity, that is, the extent to which groupings of individuals at breeding sites are maintained in wintering areas, is frequently used to evaluate population structure and assess use of key habitat areas. However, for species with complex or variable annual cycle movements, this traditional bimodal framework of migratory connectivity may be overly simplistic. Like many other waterfowl, sea ducks often travel to specific pre- and post-breeding sites outside their nesting and wintering areas to prepare for migration by feeding extensively and, in some cases, molting their flight feathers. These additional migrations may play a key role in population structure, but are not included in traditional models of migratory connectivity. Network analysis, which applies graph theory to assess linkages between discrete locations or entities, offers a powerful tool for quantitatively assessing the contributions of different sites used throughout the annual cycle to complex spatial networks. We collected satellite telemetry data on annual cycle movements of 672 individual sea ducks of five species from throughout eastern North America and the Great Lakes. From these data, we constructed a multi-species network model of migratory patterns and site use over the course of breeding, molting, wintering, and migratory staging. Our results highlight inter- and intra-specific differences in the patterns and complexity of annual cycle movement patterns, including the central importance of staging and molting sites in James Bay, the St. Lawrence River, and southern New England to multi-species annual cycle habitat linkages, and highlight the value of Long-tailed Ducks (Calengula haemalis) as an umbrella species to represent the movement patterns of multiple sea duck species. We also discuss potential applications of network migration models to conservation prioritization, identification of population units, and integrating different data streams.

Rapid loss of maternal immunity and increase in environmentally mediated antibody generation in urban gulls.

Monitoring pathogen circulation in wildlife sentinel populations can help to understand and predict the spread of disease at the wildlife-livestock-human interface. Immobile young provide a useful target population for disease surveillance, since they can be easily captured for sampling and their levels of antibodies against infectious agents can provide an index of localized circulation. However, early-life immune responses include both maternally-derived antibodies and antibodies resulting from exposure to pathogens, and disentangling these two processes requires understanding their individual dynamics. We conducted an egg-swapping experiment in an urban-nesting sentinel seabird, the yellow-legged gull, and measured antibody levels against three pathogens of interest (avian influenza virus AIV, Toxoplasma gondii TOX, and infectious bronchitis virus IBV) across various life stages, throughout chick growth, and between nestlings raised by biological or non-biological parents. We found that levels of background circulation differed among pathogens, with AIV antibodies widely present across all life stages, TOX antibodies rarer, and IBV antibodies absent. Antibody titers declined steadily from adult through egg, nestling, and chick stages. For the two circulating pathogens, maternal antibodies declined exponentially after hatching at similar rates, but the rate of linear increase due to environmental exposure was significantly higher in the more prevalent pathogen (AIV). Differences in nestling antibody levels due to parental effects also persisted longer for AIV (25 days, vs. 14 days for TOX). Our results suggest that yellow-legged gulls can be a useful sentinel population of locally transmitted infectious agents, provided that chicks are sampled at ages when environmental exposure outweighs maternal effects.

Distributing transmitters to maximize population-level representativeness in automated radio telemetry studies of animal movement.

Telemetry is a powerful and indispensable tool for evaluating wildlife movement and distribution patterns, particularly in systems where opportunities for direct observation are limited. However, the effort and expense required to track individuals often results in small sample sizes, which can lead to biased results if the sample of tracked individuals does not fully capture spatial, temporal, and individual variability within the target population. To better understand the influence of sampling design on results of automated radio telemetry studies, we conducted a retrospective power analysis of very high frequency (VHF) radio telemetry data from the Motus Wildlife Tracking System for two species of birds along the United States Atlantic coast: a shorebird, the piping plover (Charadrius melodus), and a nearshore seabird, the common tern (Sterna hirundo). We found that ~ 100-150 tracked individuals were required to identify 90% of locations known to be used by the tracked population, with 40-50 additional individuals required to include 95% of used locations. For any number of individuals, the percentage of stations included in the sample was higher for common terns than for piping plovers when tags were deployed within a single site and year. Percentages of stations included increased for piping plovers when birds were tagged over multiple sites and, to a lesser extent, years, and increased with average length of the tracking period. The probability that any given receiver station used by the population would be included in a subsample increased with the number of birds tracked, station proximity to a migratory stopover or staging site, number of receiving antennas per station, and percentage of the tracked population present. Our results provide general guidance for the number and distribution of tagged birds required to obtain representative VHF telemetry data, while also highlighting the importance of accounting for station network configuration and species-specific differences in behavior when designing automated radio telemetry studies to address specific research questions. Our results have broad applications to remotely track movements of small-bodied migratory wildlife in inaccessible habitats, including predicting and monitoring effects of offshore wind energy development.

Blood biochemistry and hematology of adult and chick brown pelicans in the northern Gulf of Mexico: baseline health values and ecological relationships.

The northern Gulf of Mexico supports a diverse community of nearshore seabirds during both breeding and nonbreeding periods of the annual cycle and is also a highly industrialized marine ecosystem with substantial levels of oil and gas development particularly in the west and central regions. Stakeholders in the region often assess risk to species of interest based on these differing levels of development. We collected blood samples from 81 adult and 35 chick eastern brown pelicans (Pelecanus occidentalis carolinensis) from 10 colonies across the northern Gulf of Mexico and used these to establish baseline values for hematology and blood biochemistry. We assessed the potential influence of body condition, sex and home range size on hematology and blood biochemistry. We also assessed potential influences of oil and gas activity by considering differing levels of oil and gas development that occur regionally throughout the study area. Although blood analyte concentrations of adults and chicks were often associated with these regional differences, the pattern we observed was not entirely consistent with the differing levels of oil and gas activity across the Gulf, suggesting that regional levels of oil and gas activity around breeding sites may not be the primary drivers of hematology and blood biochemistry. We note that baseline values or reference intervals are not available for other nearshore seabirds that breed in the northern Gulf. Given that exposure and risk may differ among this suite of species based on diet, foraging strategies and life history strategies, similar assessments and monitoring may be warranted.

Tropical cyclones alter short-term activity patterns of a coastal seabird.

BACKGROUND: Mobile organisms in marine environments are expected to modify their behavior in response to external stressors. Among environmental drivers of animal movement are long-term climatic indices influencing organism distribution and short-term meteorological events anticipated to alter acute movement behavior. However, few studies exist documenting the response of vagile species to meteorological anomalies in coastal and marine systems. METHODS: Here we examined the movements of Eastern brown pelicans (Pelecanus occidentalis carolinensis) in the South Atlantic Bight in response to the passage of three separate hurricane events in 2 years. Pelicans (n = 32) were tracked with GPS satellite transmitters from four colonies in coastal South Carolina, USA, for the entirety of at least one storm event. An Expectation Maximization binary Clustering algorithm was used to discretize pelican behavioral states, which were pooled into 'active' versus 'inactive' states. Multinomial logistic regression was used to assess behavioral state probabilities in relation to changes in barometric pressure and wind velocity. RESULTS: Individual pelicans were more likely to remain inactive during tropical cyclone passage compared to baseline conditions generally, although responses varied by hurricane. When inactive, pelicans tended to seek shelter using local geomorphological features along the coastline such as barrier islands and estuarine systems. CONCLUSIONS: Our telemetry data showed that large subtropical seabirds such as pelicans may mitigate risk associated with spatially-extensive meteorological events by decreasing daily movements. Sheltering may be related to changes in barometric pressure and wind velocity, and represents a strategy common to several other classes of marine vertebrate predators for increasing survival probabilities.

Modeling spatiotemporal abundance of mobile wildlife in highly variable environments using boosted GAMLSS hurdle models.

Modeling organism distributions from survey data involves numerous statistical challenges, including accounting for zero-inflation, overdispersion, and selection and incorporation of environmental covariates. In environments with high spatial and temporal variability, addressing these challenges often requires numerous assumptions regarding organism distributions and their relationships to biophysical features. These assumptions may limit the resolution or accuracy of predictions resulting from survey-based distribution models. We propose an iterative modeling approach that incorporates a negative binomial hurdle, followed by modeling of the relationship of organism distribution and abundance to environmental covariates using generalized additive models (GAM) and generalized additive models for location, scale, and shape (GAMLSS). Our approach accounts for key features of survey data by separating binary (presence-absence) from count (abundance) data, separately modeling the mean and dispersion of count data, and incorporating selection of appropriate covariates and response functions from a suite of potential covariates while avoiding overfitting. We apply our modeling approach to surveys of sea duck abundance and distribution in Nantucket Sound (Massachusetts, USA), which has been proposed as a location for offshore wind energy development. Our model results highlight the importance of spatiotemporal variation in this system, as well as identifying key habitat features including distance to shore, sediment grain size, and seafloor topographic variation. Our work provides a powerful, flexible, and highly repeatable modeling framework with minimal assumptions that can be broadly applied to the modeling of survey data with high spatiotemporal variability. Applying GAMLSS models to the count portion of survey data allows us to incorporate potential overdispersion, which can dramatically affect model results in highly dynamic systems. Our approach is particularly relevant to systems in which little a priori knowledge is available regarding relationships between organism distributions and biophysical features, since it incorporates simultaneous selection of covariates and their functional relationships with organism responses.

Influence of density-dependent competition on foraging and migratory behavior of a subtropical colonial seabird.

Density-dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central-place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density-dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density-dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density-dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density-dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density-dependent competition on breeding success and population regulation remain uncertain in this system.

Physical condition and stress levels during early development reflect feeding rates and predict pre- and post-fledging survival in a nearshore seabird.

The effects of acute environmental stressors on reproduction in wildlife are often difficult to measure because of the labour and disturbance involved in collecting accurate reproductive data. Stress hormones represent a promising option for assessing the effects of environmental perturbations on altricial young; however, it is necessary first to establish how stress levels are affected by environmental conditions during development and whether elevated stress results in reduced survival and recruitment rates. In birds, the stress hormone corticosterone is deposited in feathers during the entire period of feather growth, making it an integrated measure of background stress levels during development. We tested the utility of feather corticosterone levels in 3- to 4-week-old nestling brown pelicans (Pelecanus occidentalis) for predicting survival rates at both the individual and colony levels. We also assessed the relationship of feather corticosterone to nestling body condition and rates of energy delivery to nestlings. Chicks with higher body condition and lower corticosterone levels were more likely to fledge and to be resighted after fledging, whereas those with lower body condition and higher corticosterone levels were less likely to fledge or be resighted after fledging. Feather corticosterone was also associated with intracolony differences in survival between ground and elevated nest sites. Colony-wide, mean feather corticosterone predicted nest productivity, chick survival and post-fledging dispersal more effectively than did body condition, although these relationships were strongest before fledglings dispersed away from the colony. Both reproductive success and nestling corticosterone were strongly related to nutritional conditions, particularly meal delivery rates. We conclude that feather corticosterone is a powerful predictor of reproductive success and could provide a useful metric for rapidly assessing the effects of changes in environmental conditions, provided pre-existing baseline variation is monitored and understood.