Thomas W. Bodey

Space Partitioning Without Territoriality in Gannets

This Is the Place Bats, bees, seals, and many seabirds practice central-place foraging, leaving a central home site, such as a hive or a rookery, to forage in a specific territory. Such species also share the challenge of competing for local resources with individuals from separate colonies. Using satellite tags, Wakefield et al. (p. 68, published online 6 June; see the Perspective by Weimerskirch) followed over 180 northern gannets to determine potential drivers of foraging territory division. Boundaries among colonial territories arose as a result of competition with individuals from other territories. Individuals from the same colony appeared to share information about foraging sites, presumably contributing to the establishment and maintenance of specific, long-term colonial territories. Gannets establish foraging territory boundaries in colonies through local competition and information exchange. [Also see Perspective by Weimerskirch] Colonial breeding is widespread among animals. Some, such as eusocial insects, may use agonistic behavior to partition available foraging habitat into mutually exclusive territories; others, such as breeding seabirds, do not. We found that northern gannets, satellite-tracked from 12 neighboring colonies, nonetheless forage in largely mutually exclusive areas and that these colony-specific home ranges are determined by density-dependent competition. This segregation may be enhanced by individual-level public information transfer, leading to cultural evolution and divergence among colonies.

Long‐term individual foraging site fidelity—why some gannets don't change their spots

Many established models of animal foraging assume that individuals are ecologically equivalent. However, it is increasingly recognized that populations may comprise individuals who differ consistently in their diets and foraging behaviors. For example, recent studies have shown that individual foraging site fidelity (IFSF, when individuals consistently forage in only a small part of their populations home range) occurs in some colonial breeders. Short-term IFSF could result from animals using a win-stay, lose-shift foraging strategy. Alternatively, it may be a consequence of individual specialization. Pelagic seabirds are colonial central-place foragers, classically assumed to use flexible foraging strategies to target widely dispersed, spatiotemporally patchy prey. However, tracking has shown that IFSF occurs in many seabirds, although it is not known whether this persists across years. To test for long-term IFSF and to examine alternative hypotheses concerning its cause, we repeatedly tracked 55 Northern Gannets (Morus bassanus) from a large colony in the North Sea within and across three successive breeding seasons. Gannets foraged in neritic waters, predictably structured by tidal mixing and thermal stratification, but subject to stochastic, wind-induced overturning. Both within and across years, coarse to mesoscale (tens of kilometers) IFSF was significant but not absolute, and foraging birds departed the colony in individually consistent directions. Carbon stable isotope ratios in gannet blood tissues were repeatable within years and nitrogen ratios were also repeatable across years, suggesting long-term individual dietary specialization. Individuals were also consistent across years in habitat use with respect to relative sea surface temperature and in some dive metrics, yet none of these factors accounted for IFSF. Moreover, at the scale of weeks, IFSF did not decay over time and the magnitude of IFSF across years was similar to that within years, suggesting that IFSF is not primarily the result of win-stay, lose-shift foraging. Rather, we hypothesize that site familiarity, accrued early in-life, causes IFSF by canalizing subsequent foraging decisions. Evidence from this and other studies suggests that IFSF may be common in colonial central-place foragers, with far-reaching consequences for our attempts to understand and conserve these animals in a rapidly changing environment.

Localised control of an introduced predator: creating problems for the future?

Introduced mammalian predators have had significant impacts on many native prey species. Although control of such predators for conservation management is becoming increasingly commonplace, it is often undertaken at a relatively small scale in relation to the overall predator population. Processes such as immigration mean that it remains difficult to determine the effectiveness of control measures. We investigated the impacts of feral ferret Mustela furo removal on the entire feral ferret population on Rathlin Island, UK. Removal of ferrets prior to breeding led to a substantial increase in the post-dispersal population through the enhanced survival of juveniles. Despite increased numbers, overwinter survival remained high, potentially aided by the reduced territoriality shown by this feral species compared to wild carnivores. The response of this ferret population to control is a further illustration of the complex ecological processes and outcomes arising from the anthropogenic disruption of wildlife populations. It highlights how partial or localised management may prove ineffective, and at worst might exacerbate the problems that management was designed to avert.

Mesopredators constrain a top predator: competitive release of ravens after culling crows

Although predator control programmes rarely consider complex competitive interactions among predators, it is becoming clear that removal of larger ‘superior’ competitors often releases the ‘inferior’ ones and can precipitate trophic cascades. In contrast, our study indicates that culling hooded crows Corvus cornix appears to release a larger competitor, the common raven Corvus corax. Ravens ranged more widely, and the predation of artificial nests was significantly faster (although total predation was similar), after the removal of crows. Our study provides evidence of a novel reversal of competitive release where a larger species was freed from constraints imposed on its distribution and behaviour by a smaller species, and emphasizes the importance of considering community and ecosystem effects of predator manipulations when undertaken for conservation or game management.

Effects of age and reproductive status on individual foraging site fidelity in a long-lived marine predator

Individual foraging specializations, where individuals use a small component of the population niche width, are widespread in nature with important ecological and evolutionary implications. In long-lived animals, foraging ability develops with age, but we know little about the ontogeny of individuality in foraging. Here we use precision global positioning system (GPS) loggers to examine how individual foraging site fidelity (IFSF), a common component of foraging specialization, varies between breeders, failed breeders and immatures in a long-lived marine predator—the northern gannet Morus bassanus. Breeders (aged 5+) showed strong IFSF: they had similar routes and were faithful to distal points during successive trips. However, centrally placed immatures (aged 2–3) were far more exploratory and lacked route or foraging site fidelity. Failed breeders were intermediate: some with strong fidelity, others being more exploratory. Individual foraging specializations were previously thought to arise as a function of heritable phenotypic differences or via social transmission. Our results instead suggest a third alternative—in long-lived species foraging sites are learned during exploratory behaviours early in life, which become canalized with age and experience, and refined where possible—the exploration-refinement foraging hypothesis. We speculate similar patterns may be present in other long-lived species and moreover that long periods of immaturity may be a consequence of such memory-based individual foraging strategies.

Species versus guild level differentiation revealed across the annual cycle by isotopic niche examination

Interspecific competitive interactions typically result in niche differentiation to alleviate competition through mechanisms including character displacement. However, competition is not the sole constraint on resource partitioning, and its effects are mediated by factors including the environmental context in which species coexist. Colonial seabirds provide an excellent opportunity to investigate the importance of competition in shaping realized niche widths because their life histories lead to variation in intra- and interspecific competition across the annual cycle. Dense breeding aggregations result in intense competition for prey in surrounding waters, whereas non-breeding dispersal to larger geographical areas produces lower densities of competitors. Bayesian hierarchical models of the isotopic niche, closely aligned to the trophic niche, reveal the degree of segregation between species and functional groups during both time periods. Surprisingly, species explained far more of the variance in the isotopic niche during the non-breeding than the breeding period. Our results underline the key role of non-breeding dynamics in alleviating competition and promoting distinctions between species through the facilitation of resource partitioning. Such situations may be common in a diverse range of communities sustained by ephemeral but abundant food items. This highlights how consideration of the hierarchical grouping of competitive interactions alongside consideration of abiotic constraints across the complete annual cycle allows a full understanding of the role of competition in driving patterns of character displacement.

The diet of an invasive nonnative predator, the feral ferret Mustela furo, and implications for the conservation of ground-nesting birds

Introduced carnivores have had a significant impact on the fauna of a number of countries, particularly on islands. In the British Isles, several offshore islands holding internationally important aggregations of seabirds and shorebirds support self-sustaining feral ferret Mustela furo populations, often as the top terrestrial predator. However, little is known about the interactions between ferrets and both native and nonnative prey in these locations. We examined the diet of feral ferrets on Rathlin Island, Northern Ireland. We determined the frequency of occurrence of prey items and constructed energetic models to determine their potential impact on both native and introduced prey. Rabbits Oryctolagus cuniculus occurred in 75% of scats, while birds, carrion, and brown rats Rattus norvegicus were important secondary items. There was little difference between the diets of males and females. Estimates of the energy requirements of the population at current, and with hypothetically reduced, rabbit availability revealed the potential for carrion to maintain the ferret population over winter. Management options could thus focus on reducing anthropogenic food sources as an immediate way of mitigating the threat to ground-nesting birds, while other strategies, including eradication, are considered.

A phylogenetically controlled meta-analysis of biologging device effects on birds: Deleterious effects and a call for more standardized reporting of study data

1.The use of biologging devices continues to increase, with technological advances yielding remarkable ecological insights and generating new research questions. However, as devices develop and are deployed more widely, there is a need to update our knowledge of the potential ethical impacts to allow scientists to balance these against the knowledge gained. 2.We employed a suite of phylogenetically controlled meta-analyses on a dataset comprising more than 450 published effect sizes across 214 different studies to examine the effects of biologger tagging on five key traits in birds. 3.Overall, we found small but significant negative effects of tagging on survival, reproduction, parental care. In addition, tagging was positively associated with foraging trip duration, but had no effect on body mass. Meta-regressions revealed that flying style, migration distance and proportional tag mass were significant influences producing these deleterious effects, with attachment type and position additionally important covariates influencing survival- and reproduction-based effect sizes. 4.There was a positive correlation between the effects of tagging on survival and reproduction, highlighting that effects may be cumulative, with the full effects of tagging not necessarily apparent in studies focused on single traits. We discuss the tradeoff between these negative effects and the advances gained through the use of biologgers. 5.Finally, given the number of studies from our initial literature search that lacked sufficient data for inclusion in analyses, we provide recommendations on the essential information that all biologging studies should report in order to facilitate future assessments of impacts on animals. This article is protected by copyright. All rights reserved.

Search and foraging behaviors from movement data: A comparison of methods.

Abstract Search behavior is often used as a proxy for foraging effort within studies of animal movement, despite it being only one part of the foraging process, which also includes prey capture. While methods for validating prey capture exist, many studies rely solely on behavioral annotation of animal movement data to identify search and infer prey capture attempts. However, the degree to which search correlates with prey capture is largely untested. This study applied seven behavioral annotation methods to identify search behavior from GPS tracks of northern gannets (Morus bassanus), and compared outputs to the occurrence of dives recorded by simultaneously deployed time–depth recorders. We tested how behavioral annotation methods vary in their ability to identify search behavior leading to dive events. There was considerable variation in the number of dives occurring within search areas across methods. Hidden Markov models proved to be the most successful, with 81% of all dives occurring within areas identified as search. k‐Means clustering and first passage time had the highest rates of dives occurring outside identified search behavior. First passage time and hidden Markov models had the lowest rates of false positives, identifying fewer search areas with no dives. All behavioral annotation methods had advantages and drawbacks in terms of the complexity of analysis and ability to reflect prey capture events while minimizing the number of false positives and false negatives. We used these results, with consideration of analytical difficulty, to provide advice on the most appropriate methods for use where prey capture behavior is not available. This study highlights a need to critically assess and carefully choose a behavioral annotation method suitable for the research question being addressed, or resulting species management frameworks established.

Climatic conditions produce contrasting influences on demographic traits in a long‐distance Arctic migrant

The manner in which patterns of variation and interactions among demographic rates contribute to population growth rate (λ) is key to understanding how animal populations will respond to changing climatic conditions. Migratory species are likely to be particularly sensitive to climatic conditions as they experience a range of different environments throughout their annual cycle. However, few studies have provided fully integrated demographic analyses of migratory populations in response to changing climatic conditions. Here, we employed integrated population models to demonstrate that the environmental conditions experienced during a short but critical period play a central role in the demography of a long-distance migrant, the light-bellied Brent goose (Branta bernicla hrota). Female survival was positively associated with June North Atlantic Oscillation (NAO) values, whereas male survival was not. In contrast, breeding productivity was negatively associated with June NAO, suggesting a trade-off between female survival and reproductive success. Both adult female and adult male survival showed low temporal variation, whereas there was high temporal variation in recruitment and breeding productivity. In addition, while annual population growth was positively correlated with annual breeding productivity, a sensitivity analysis revealed that population growth was most sensitive to changes in adult survival. Our results demonstrate that the environmental conditions experienced during a relatively short-time window at the start of the breeding season play a critical role in shaping the demography of a long-distant Arctic migrant. Crucially, different demographic rates responded in opposing directions to climatic variation, emphasising the need for integrated analysis of multiple demographic traits when understanding population dynamics.