The hidden cost of disturbance: Eurasian Oystercatchers (Haematopus ostralegus) avoid a disturbed roost site during the tourist season

Disturbance may impact individual birds and ultimately bird populations. If animals avoid disturbed sites this may prevent them from being disturbed directly but may also negatively impact their movement patterns and energy budgets. Avoidance is, however, challenging to study, because it requires following individuals over large spatial scales in order to compare their movement rates between sites in relation to spatiotemporal variation in disturbance intensity. We studied how 48 GPS-tracked non-breeding Eurasian Oystercatchers Haematopus ostralegus used two neighbouring roost sites in the Wadden Sea. One roost site is highly influenced by seasonal recreational disturbance whereas the other is an undisturbed sandbar. We analysed roost choice and the probability of moving away from the disturbed roost site with regard to a seasonal recreation activity index, weekends and night-time. Oystercatchers often chose to roost on the undisturbed site, even if they were foraging closer to the disturbed roost. The probability that Oystercatchers chose to roost on the disturbed site was negatively correlated with the recreation activity index and was lowest in the tourist season (summer and early autumn), indicating that birds used the site less often when recreation levels were high. Furthermore, the probability that birds moved away from the disturbed site during high tide was positively correlated with the recreation activity index. The choice to roost on the undisturbed site implies that birds must fly an additional 8 km during one high-tide period, which equates to 3.4% of daily energy expenditure of an average Oystercatcher. Our study tentatively suggests that the costs of avoidance may outweigh the energetic cost of direct flight responses and hence that avoidance of disturbed sites requires more attention in future disturbance impact studies. Nature managers should evaluate whether high-quality undisturbed roosting sites are available near foraging sites, and in our case closing of a section of the disturbed site during high tides in the tourist season may mitigate much disturbance impact

Balancing food and density-dependence in the spatial distribution of an interference-prone forager

Foraging distributions are thought to be density-dependent, because animals not only select for a high availability and quality of resources, but also avoid conspecific interference. Since these processes are confounded, their relative importance in shaping foraging distributions remains poorly understood. Here we aimed to rank the contribution of density-dependent and density-independent effects on the spatio-temporal foraging patterns of eurasian oystercatchers. In our intertidal study area, tides caused continuous variation in oystercatcher density, providing an opportunity to disentangle conspecific interference and density-independent interactions with the food landscape. Spatial distributions were quantified using high-resolution individual tracking of foraging activity and location. In a model environment that included a realistic reconstruction of both the tides and the benthic food, we tested a family of behaviour-based optimality models against these tracking data. Density-independent interactions affected spatial distributions much more strongly than conspecific interference, even in an interference-prone species like oystercatchers. Spatial distributions were governed by avoidance of bill injury costs, selection for high interference-free intake rates and a decreasing availability of benthic bivalve prey after their exposure. These density-independent interactions outweighed interference competition in terms of effect size. We suggest that the bottleneck in our mechanistic understanding of foraging distributions may be primarily the role of density-independent prey attributes unrelated to intake rates, like damage costs in the case of oystercatchers foraging on perilous prey. At a landscape scale, above the finest inter-individual distances, effects of conspecific interaction on spatial distributions may have been overemphasised

Spatiotemporal variation in disturbance impacts derived from simultaneous tracking of aircraft and shorebirds

Assessing the impacts of disturbance over large areas and long time periods is crucial for nature management, but also challenging since impacts depend on both wildlife responses to disturbance and on the spatiotemporal distribution of disturbance sources. Combined tracking of animals and disturbance sources enables quantification of wildlife responses as a function of the distance to a disturbance source. We provide a framework to derive such distance–response curves and combine those with disturbance source presence data to quantify energetic costs of disturbance at a landscape scale. We tracked 90 Eurasian Oystercatchers Haematopus ostralegus and all aircraft in a military training area in the Dutch Wadden Sea. We quantified distance–response curves estimating flight probability and additional displacement for five types of aircraft activities, by comparing bird movement prior to aircraft presence with movement during aircraft presence. We then used the distance–response curves to map mean and variation in additional daily energy expenditure due to cumulative aircraft disturbance across the landscape for a 700-day period. Flight probability and displacement responses differed strongly among aircraft activities and decreased from transport aeroplanes, through bombing jets, helicopters, jets to small civil aeroplanes. Since the most disturbing aircraft activities were also the rarest ones, mean additional daily energy expenditure did not exceed 0.25%. However, days with substantial (>1%) additional expenditure occurred between 0.1% and 3.7% of all days across high-tide roosts in the tidal basin. Notably, expenditure particularly spiked on days with transport aeroplane activity (up to 8.5%). Synthesis and applications. We quantified cumulative energetic flight costs due to aircraft disturbance and found that these were low and unlikely to impact survival of oystercatchers in our study area. Our results provide evidence that the legal minimum flight height of 450 m for small civil aeroplanes effectively limits the disturbance of oystercatchers. Mitigation should focus on limiting the number of days when disturbance has a high impact by reducing rare but highly disturbing activities, especially transport aeroplanes. Our approach can be applied to other species and disturbance sources that are automatically tracked, for example boats and walkers, ultimately to quantify the entire anthropogenic disturbance landscape

The demographic causes of population change vary across four decades in a long-lived shorebird

Understanding which factors cause populations to decline begins with identifying which parts of the life cycle, and which vital rates, have changed over time. However, in a world where humans are altering the environment both rapidly and in different ways, the demographic causes of decline likely vary over time. Identifying temporal variation in demographic causes of decline is crucial to assure that conservation actions target current and not past threats. However, this has rarely been studied as it requires long time series. Here we investigate how the demography of a long-lived shorebird (the Eurasian Oystercatcher Haematopus ostralegus) has changed in the past four decades, resulting in a shift from stable dynamics to strong declines (−9% per year), and recently back to a modest decline. Since individuals of this species are likely to respond differently to environmental change, we captured individual heterogeneity through three state variables: age, breeding status, and lay date (using integral projection models). Timing of egg-laying explained significant levels of variation in reproduction, with a parabolic relationship of maximal productivity near the average lay date. Reproduction explained most variation in population growth rates, largely due to poor nest success and hatchling survival. However, the demographic causes of decline have also been in flux over the last three decades: hatchling survival was low in the 2000s but improved in the 2010s, while adult survival declined in the 2000s and remains low today. Overall, the joint action of several key demographic variables explain the decline of the oystercatcher, and improvements in a single vital rate cannot halt the decline. Conservations actions will thus need to address threats occurring at different stages of the oystercatcher's life cycle. The dynamic nature of the threat landscape is further supported by the finding that the average individual no longer has the highest performance in the population, and emphasizes how individual heterogeneity in vital rates can play an important role in modulating population growth rates. Our results indicate that understanding population decline in the current era requires disentangling demographic mechanisms, individual variability, and their changes over time

Disturbance increases high tide travel distance of a roosting shorebird but only marginally affects daily energy expenditure

Background: Anthropogenic disturbance can negatively affect an animal's energy budget by evoking movement responses. Existing research focuses mainly on immediate displacement as a disturbance effect, since this can be easily observed in the field. However, effects on movement over longer timescales are poorly examined and it is largely unknown if and to what extent they reflect immediate responses. Longer-term responses could for example be larger than immediate responses if birds, after disturbance, return to the original location and thereby travel twice the immediate disturbed distance. Methods: We combined GPS tracking data with observational data to quantify the effects of anthropogenic (air force and walkers) and non-anthropogenic disturbances on distances travelled by roosting Eurasian Oystercatchers (Haematopus ostralegus) during the non-breeding season. We compared immediate displacement after a disturbance with distance travelled during the entire high tide period (longer-term response), while accounting for environmental factors. Additionally, we calculated energy expenditure due to disturbance based on observed disturbance frequencies. Results: Disturbance resulted in an immediate displacement response of ~ 200 m (median). Air force disturbances tended to yield larger immediate responses than walker and, especially, than non-anthropogenic disturbances. Longer-term responses and immediate responses were approximately similar, suggesting that, over longer timescales, spatial disturbance effects in the study area remain confined to immediate effects. However, disturbances were infrequent (0.17 disturbances per bird per hour) and most disturbances were of natural origin (62%). Consequently, anthropogenic disturbance of roosting oystercatchers in the study area on average costs 0.08% of the daily energy expenditure. Conclusions: Our results suggest that immediate spatial responses to disturbance can be a useful proxy for spatial responses over longer timescales. Over the non-exhaustive range of conditions investigated, energetic consequences of spatial disturbance responses for an oystercatcher in the study area are marginal due to low disturbance levels

In this Issue

Aim: Molluscivorous shorebirds supposedly developed their present wintering distribution after the last ice age. Currently, molluscivorous shorebirds are abundant on almost all shores of the world, except for those in the Indo-West Pacific (IWP). Long before shorebirds arrived on the scene, molluscan prey in the IWP evolved strong anti-predation traits in a prolonged evolutionary arms race with durophagous predators including brachyuran crabs. Here, we investigate whether the absence of molluscivorous shorebirds from a site in Oman can be explained by the molluscan community being too well-defended. Location: The intertidal mudflats of Barr Al Hikman, Oman. Methods: Based on samples from 282 locations across the intertidal area the standing stock of the macrozoobenthic community was investigated. By measuring anti-predation traits (burrowing depth, size and strength of armour), the fraction of molluscs available to molluscivorous shorebirds was calculated. Results: Molluscs dominated the macrozoobenthic community at Barr Al Hikman. However, less than 17% of the total molluscan biomass was available to shorebirds. Most molluscs were unavailable either because of their hard-to-crush shells, or because they lived too deeply in the sediment. Repair scars and direct observations confirmed crab predation on molluscs. Although standing stock densities of the Barr Al Hikman molluscs were of the same order of magnitude as at intertidal mudflat areas where molluscivorous shorebirds are abundant, the molluscan biomass available to shorebirds was distinctly lower at Barr Al Hikman. Main conclusions: The established strong molluscan anti-predation traits against crabs precludes molluscan exploitation by shorebirds at Barr Al Hikman. This study exemplifies that dispersal of "novel" predators is hampered in areas where native predators and prey exhibit strongly developed attack and defence mechanisms, and highlights that evolutionary arms races can have consequences for the global distribution of species