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.
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.
Studies of the effects of transmitters on wildlife often focus on survival. However, sublethal behavioral changes resulting from radio-marking have the potential to affect inferences from telemetry data and may vary based on individual and environmental characteristics. We used a long-term, multi-species tracking study of sea ducks to assess behavioral patterns at multiple temporal scales following implantation of intracoelomic satellite transmitters. We applied state-space models to assess short-term behavioral patterns in 476 individuals with implanted satellite transmitters, as well as comparing breeding site attendance and migratory phenology across multiple years after capture. In the short term, our results suggest an increase in dispersive behavior immediately following capture and transmitter implantation; however, behavior returned to seasonally average patterns within ~5 days after release. Over multiple years, we found that breeding site attendance by both males and females was depressed during the first breeding season after radio-marking relative to subsequent years, with larger relative decreases in breeding site attendance among males than females. We also found that spring and breeding migrations occurred later in the first year after radio-marking than in subsequent years. Across all behavioral effects, the severity of behavioral change often varied by species, sex, age, and capture season. We conclude that, although individuals appear to adjust relatively quickly (i.e. within 1 week) to implanted satellite transmitters, changes in breeding phenology may occur over the longer term and should be considered when analyzing and reporting telemetry data.
Located at the shortest overland route between the Gulf of Mexico and the Pacific Ocean, Mexico's Tehuantepec Isthmus is a globally important migratory corridor for many terrestrial bird species. The Pacific coast of the Isthmus also contains a significant wetland complex that supports large multi-species aggregations of nonbreeding waterbirds during the boreal winter. In recent years, extensive wind energy development has occurred in the plains bordering these wetlands, directly along the migratory flyway. Using recent studies of movement patterns of three marineassociated bird species -reddish egrets Egretta rufescens, brown pelicans Pelecanus occidentalis, and red knots Calidris canutus -from the northern Gulf of Mexico, we assess the use of the isthmus as a migratory corridor. Our data provide evidence that marine birds from the Gulf region regularly overwinter along the Pacific coast of Mexico and use the isthmus as a migratory corridor, creating the potential for interaction with terrestrial wind farms during non-breeding. This study is the first to describe migration by marine-associated bird species between the Gulf of Mexico and Pacific coast. These data contribute new information toward ongoing efforts to understand the complex migration patterns of mobile marine species, with the goal of informing integrated conservation efforts for species whose year-round habitat needs cross ecoregional and geopolitical boundaries.
Although the use of bird-borne data loggers has become widespread in avian field research, the effects of capture and transmitter attachment on behavior and demographic rates are not often measured. Tag-and capture-induced effects on individual behavior, survival and reproduction may limit extrapolation of transmitter data to wider populations. However, measuring individual responses to capture and tagging is a necessary step in developing research techniques that minimize negative effects. We measured the short-term behavioral effects of handling and GPS transmitter attachment on Brown Pelicans under both captive and field conditions, and followed tagged individuals through a full breeding season to assess whether capture and transmitter attachment increased rates of nest abandonment or breeding failure. We observed slight increases in preening among tagged individuals 0-2 h after capture relative to controls that had not been captured or tagged, with a corresponding reduction in time spent resting. One to three days post-capture, nesting behavior of tagged pelicans resembled that of neighbors that had not been captured or tagged. Eighty-eight percent of tagged breeders remained at the same nest location for more than 48 h after capture, attending nests and chicks for an average of 49 days, and 51% were assumed to successfully fledge young. Breeding success was driven primarily by variation in location; however, sex and handling time also influenced the probability of successful breeding in tagged pelicans, suggesting that individual characteristics and the capture process itself can confound the effects of capture and transmitter attachment. We conclude that pelicans fitted with GPS transmitters exhibit comparable behaviors to untagged individuals within a day of capture and that GPS tracking is a viable technique for studying behavior and demography in this species. We also identify measures to minimize post-capture nest abandonment rates in tracking studies, including minimizing handling time and covering nests during processing.
Snapshot nestling health measures can reveal reproductive effects of environmental stress while minimizing handling, disturbance and researcher effort. We tested two short-term measurements, body condition and feather corticosterone, as predictors of brown pelican nestling survival. Both measurements predicted nestling survival to fledge, and feather corticosterone also predicted post-fledging survival.
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.
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