Effects of windscape on three-dimensional foraging behaviour in a wide-ranging marine predator, the northern gannet

Marine birds are strongly exposed to weather conditions at sea but to date, few studies have investigated the influences of wind or rainfall on their time-activity budgets or foraging routines. Here, we used data from GPS and pressure loggers to investigate the effects of wind speed and direction and rainfall on the 3-dimensional foraging behaviour of gannets Morus bassanus breeding at Bass Rock, Scotland. We found that birds spent more time actively foraging during stronger winds, but there was no subsequent increase in overall trip duration because individuals compensated by decreasing the time they spent on the water during stronger winds. Birds returned more quickly from distant foraging grounds, and those encountering head winds spent less time on the water and so were able to compensate to some extent for an adverse effect of head winds on speed of travel over the return leg. These data strongly suggest that by reducing time spent on the water, birds were able to buffer trip durations against adverse effects of strong winds encountered during both commuting and active foraging. Birds also commuted at greater heights with increasing tail wind speed and at lower heights with increasing head wind speed, potentially providing an additional behavioural buffer against the adverse effects of strong head winds during foraging trips. There was no discernible effect of rain on foraging, but the behavioural flexibility recorded here is likely to be critical to maintaining nest attendance patterns and food provisioning rates of chicks across variable environmental conditions encountered at sea

Incorporating intraspecific trait variation into functional diversity: Impacts of selective logging on birds in Borneo

1. As conservation increasingly recognises the importance of species’ functional roles in ecosystem processes, studies are shifting away from measuring species richness towards measures that account for the functional differences between species in a community. These functional diversity (FD) indices have received much recent attention and refinement, but their greatest limitation remains their inability to incorporate information about intraspecific trait variation (ITV). 2. We use an individual-based model to account for ITV when calculating the functional diversity of two avian communities in Borneo; one in primary (unlogged) forest and one in selectively logged forest. We deal with the scarcity of trait data for individual species by developing a simulation approach, taking data from the literature where necessary. Using a bootstrapping procedure, we produce a range of ecologically feasible FD values taking account of ITV for five commonly-used FD indices, and we quantify the confidence that can be placed in these values using a newly-developed bootstrapping method: btFD. 3. We found that incorporating ITV significantly altered the FD values of all indices used in our models. The rank order of FD for the two communities, indicating whether diversity was higher in primary or selectively logged forest, was largely unchanged by the inclusion of ITV. However, by accounting for ITV, we were able to reveal previously unrecognized impacts of selective logging on avian functional diversity through a narrower dispersion of individuals in functional trait space in logged forest. 4. Our results highlight the importance of incorporating ITV into measures of functional diversity, whilst our simulation approach addresses the frequently encountered difficulty of working with sparse trait data and quantifies the confidence that should be placed in such findings

Vulnerability of northern gannets to offshore wind farms; seasonal and sex-specific collision risk and demographic consequences

There is a pressing need to quantify the risks of renewable energy developments such as offshore wind farms for protected populations. However, assessments are often based on incomplete data, or fail to consider variation in risk between sexes and at different times of year. We tracked northern gannets foraging from the world's largest colony (Bass Rock, Scotland) across five consecutive breeding seasons. We examine how seasonal and sex differences in behaviour affect the collision risk from planned and operational wind farms within their foraging range and assess the likely consequences for long-term population viability. Both sexes made shorter trips during chick-rearing than prior to chick-hatching, spent a greater proportion of time within wind farm sites and had an eight times greater potential collision risk during chick-rearing. Females made longer trips than males at both these times of year, flew higher and spent more time within wind farm sites, leading to three times greater collision risk for females. After accounting for the potential additional mortality from collisions, and assuming that the death of a parent also led to the loss of its offspring, the breeding population was projected to increase by 3.57% (95% CI: 2.16–5.15%) per year, compared with 6.56% (95% CI: 4.59–8.73%) in the absence of turbines, suggesting a negligible effect on population viability. However, additional mortality could result in greater immigration from neighbouring colonies, potentially affecting their viability and highlighting a need for research within a metapopulation framework to assess the impacts of offshore wind developments on vulnerable species across multiple connected sites

Using time-series similarity measures to compare animal movement trajectories in ecology

Identifying and understanding patterns in movement data are amongst the principal aims of movement ecology. By quantifying the similarity of movement trajectories, inferences can be made about diverse processes, ranging from individual specialisation to the ontogeny of foraging strategies. Movement analysis is not unique to ecology however, and methods for estimating the similarity of movement trajectories have been developed in other fields but are currently under-utilised by ecologists. Here, we introduce five commonly used measures of trajectory similarity: dynamic time warping (DTW), longest common subsequence (LCSS), edit distance for real sequences (EDR), Fréchet distance and nearest neighbour distance (NND), of which only NND is routinely used by ecologists. We investigate the performance of each of these measures by simulating movement trajectories using an Ornstein-Uhlenbeck (OU) model in which we varied the following parameters: (1) the point of attraction, (2) the strength of attraction to this point and (3) the noise or volatility added to the movement process in order to determine which measures were most responsive to such changes. In addition, we demonstrate how these measures can be applied using movement trajectories of breeding northern gannets (Morus bassanus) by performing trajectory clustering on a large ecological dataset. Simulations showed that DTW and Fréchet distance were most responsive to changes in movement parameters and were able to distinguish between all the different parameter combinations we trialled. In contrast, NND was the least sensitive measure trialled. When applied to our gannet dataset, the five similarity measures were highly correlated despite differences in their underlying calculation. Clustering of trajectories within and across individuals allowed us to easily visualise and compare patterns of space use over time across a large dataset. Trajectory clusters reflected the bearing on which birds departed the colony and highlighted the use of well-known bathymetric features. As both the volume of movement data and the need to quantify similarity amongst animal trajectories grow, the measures described here and the bridge they provide to other fields of research will become increasingly useful in ecology

Tropical land-use change alters trait-based community assembly rules for dung beetles and birds

Tropical rainforest disturbance and conversion are critical drivers of biodiversity loss. A key knowledge gap is understanding the impacts of habitat modification on mechanisms of community assembly, which are predicted to respond differently between taxa and across spatial scales. We use a null model approach to detect trait assembly of species at local- and landscape-scales, and then subdivide communities with different habitat associations and foraging guilds to investigate whether the detection of assembly mechanisms varies between groups. We focus on two indicator taxa, dung beetles and birds, across a disturbance gradient of primary rainforest, selectively logged rainforest, and oil palm plantations in Borneo, Southeast Asia. Random community assembly was predominant for dung beetles across habitats, whereas trait convergence, indicative of environmental filtering, occurred across the disturbance gradient for birds. Assembly patterns at the two spatial scales were similar. Subdividing for habitat association and foraging guild revealed patterns hidden when focusing on the overall community. Dung beetle forest specialists and habitat generalists showed opposing assembly mechanisms in primary forest, community assembly of habitat generalists for both taxa differed with disturbance intensity, and insectivorous birds strongly influenced overall community assembly relative to other guilds. Our study reveals the sensitivity of community assembly mechanisms to anthropogenic disturbance via a shift in the relative contribution of stochastic and deterministic processes. This highlights the need for greater understanding of how habitat modification alters species interactions and the importance of incorporating species’ traits within assessments

Avian blood parasite infection during the non-breeding season: an overlooked issue in declining populations?

Background Pathogens and parasites can have major impacts on host population dynamics, both through direct mortality and via indirect effects. Both types of effect may be stronger in species whose populations are already under pressure. We investigated the potential for blood parasites to impact upon their hosts at the immunological, physiological and population level during the non-breeding season using a declining population of yellowhammers Emberiza citrinella as a model. Results Yellowhammers infected by Haemoproteus spp. showed both a reduced heterophil to lymphocyte (H:L) ratio, and an elevated standardised white blood cell (WBC) count compared to uninfected birds, indicating an immunological response to infection. Infected birds had shorter wings during the first winter of sampling but not during the second, colder, winter; survival analysis of 321 birds sampled across four winters indicated that increased wing length conferred a survival advantage. Conclusions We suggest that the potential impacts of blood parasite infections on over-wintering birds may have been underestimated. Further research should consider the potential impacts of sub-clinical parasite infections on the dynamics of vulnerable populations, and we suggest using declining populations as model systems within which to investigate these relationships as well as examining interactions between sub-clinical disease and other environmental stressors

Seabird movement reveals the ecological footprint of fishing vessels.

Exploitation of the seas is currently unsustainable, with increasing demand for marine resources placing intense pressure on the Earth's largest ecosystem [1]. The scale of anthropogenic effects varies from local to entire ocean basins [1-3]. For example, discards of commercial capture fisheries can have both positive and negative impacts on scavengers at the population and community-level [2-6], although this is driven by individual foraging behaviour [3,7]. Currently, we have little understanding of the scale at which individual animals initiate such behaviours. We use the known interaction between fisheries and a wide-ranging seabird, the Northern gannet Morus bassanus[3], to investigate how fishing vessels affect individual birds' behaviours in near real-time. We document the footprint of fishing vessels' (≥15 m length) influence on foraging decisions (≤11 km), and a potential underlying behavioural mechanism, by revealing how birds respond differently to vessels depending on gear type and activity. Such influences have important implications for fisheries, including the proposed discard ban [8]), and wider marine management

Post-fledging movements, mortality and migration of juvenile northern gannets

Studying the at-sea movements and behaviour of juvenile seabirds is logistically challenging, but new technologies now allow tracking birds on their first migration, giving a more complete picture of population-level spatial ecology. We investigated the post-fledging migration of juvenile northern gannets Morus bassanus from the world's largest colony, at Bass Rock, Scotland. We first examined the movements and survival of 38 juveniles over their initial days at sea with GPS precision for up to 53 d post-fledging. We then compared their migration journeys with those of 35 adults tracked with geolocators. Almost one-third of juveniles died within 2 mo of leaving the colony, and this mortality was often associated with apparent uncertainties in their direction of migration, including marked, abrupt and often repeated changes in bearing within the North Sea. Both juveniles and adults then migrated as far as the Canary Current Large Marine Ecosystem (CCLME) off the Atlantic coast of West Africa, initially taking both clockwise and counter-clockwise routes around the UK. Juveniles covered a distance each day similar to that of adults, but they reached the CCLME much more quickly, mainly because they travelled more directly, staying close to the coast throughout most of their migration, whereas adults additionally spent long periods over relatively restricted areas of ocean further offshore. The CCLME is a hotspot of unregulated fishing activity, and our findings highlight the importance of this region across different age-classes of birds, echoing previous calls that the regional strengthening of marine conservation should be a high priority

Risks to different populations and age classes of gannets from impacts of offshore wind farms in the southern North Sea

The southern North Sea holds the world's highest concentration of offshore wind farms (OWFs). Northern gannets (Morus bassanus), a species considered at high risk from OWF impacts, show a strong seasonal peak there in November, but it is unclear which populations and age classes are most at risk of collision with wind turbines. We tagged adult and juvenile gannets at the world's largest colony (Bass Rock) and reviewed two sources of survey data for different age classes to study their movements through southern North Sea waters. Tracked birds showed peak numbers in the southern North Sea in mid-October, with much smaller numbers there during November. Adults were distributed throughout the area, including waters close to OWFs, whereas juveniles were confined to the coast. Survey data indicated high proportions of immature gannets in southern North Sea waters, suggesting higher collision risk than for adults. Gannets present in November may be predominantly from colonies further north than Bass Rock

Frequency and consequences of individual dietary specialisation in a wide-ranging marine predator, the northern gannet

Individual specialisations in animals are important contributors to a wide range of ecological and evolutionary processes, and have been particularly documented in relation to multiple aspects of foraging behaviours. Central-place foragers, such as seabirds, frequently exhibit pronounced specialisations and individual differences in a variety of foraging traits. In particular, the availability of fisheries discards alongside natural prey resources provides additional potential for differentiation and specialisation for opportunistically scavenging seabird species. However, the consequences of such specialisations for at-sea distributions and intraspecific interactions are not well known. Here, we investigated the links between the degree of dietary specialisation on natural or discarded prey and the foraging movements and spatial occupancy of northern gannets Morus bassanus in relation to differing intraspecific competition at 6 colonies of differing sizes. We found that, at most colonies, individuals with different dietary strategies concentrated foraging at differing levels of intraspecific competition. In addition, individuals pursuing different strategies were frequently, but not consistently, spatially separated, distinctions that were most acutely seen in females. However, this variation in individual strategy had no significant impact on current body condition. These analyses demonstrate how foraging-associated metrics need not covary within an unconstrained system. They also reveal that specialisation can have important consequences for the competitive regimes individuals experience, highlighting the complexity of examining interacting consequences at large spatial scales