Biologging or biotelemetry, such as GPS tracking, is a common, modern method to study movement ecology in birds and other animals (i.e. 1, 2). Tracking studies have enormously improved our understanding of animal distribution and behaviour at locations and times where human observations are difficult or impossible. Tracking technology has provided thereby valuable insight into migration strategies, foraging ecology and energetics of the full annual cycle (i.e. 3, 4–9). However, to generalize behaviour assessed from animal tracking, it is important to critically examine whether the tracking device and its attachment affects the behaviour of the study species (10, 11). Using a meta-analysis including more than 450 studies, Body et al. (12) showed that in general, tagging birds had a negative effect on survival and reproduction, and was positively associated with foraging trip duration; however the impact varied per species, device, and attachment method. These negative effects of tracking devices on birds may be caused by their added mass (13), discomfort (14) or increased drag during flight or during dives (15, 16). Additionally, tracking devices fitted for longer-term deployment, for instance with a wing harness (17), may cause feather damage, feather loss, or abrasion of the skin underneath the device or harness (14). In Greater Sage-Grouse (Centrocercus urophasianus) skin abrasions were found underneath the harness depending on the material used (18), in Black-legged Kittiwake (Rissa tridactyla) the skin underneath the tracking device was inflamed when recaptured (19), and Lesser Black-backed Gulls (Larus fuscus) which were recovered two years after tagging showed feather damage (17).
In our own studies with Lesser Black-backed Gulls, with multi-year GPS deployments using a wing harness (i.e. 20, 21, 22), we observed bare patches lacking down and contour feathers under the GPS tag in re-trapped individuals after at least one cycle of postnuptial moult (see photo’s in Additional File 1). A lack of contour feathers results in a permeable plumage and may compromise insulation, thereby affecting thermoregulatory costs of birds that frequently forage on or dive in (sea)water (23). An increase in thermoregulatory costs may lead to longer foraging trips (24) or mortality if this energy loss cannot be compensated for, but it could also lead to shifts away from aquatic habitats. In an experiment to test the effects of tracking devices fitted with a wing harness on captive Mallards (Anas platyrhynchos), those with a device were less likely to be in the water than the control group (25), which was explained by a disruption of the waterproof layer by the device and harness. Obviously, such adaptive behaviour would only be feasible for species which have the option to choose between aquatic or terrestrial habitats.
Lesser Black-backed Gulls are among the most commonly tracked seabirds, and previous studies found no negative effects of GPS tags on offspring development within the year of tagging (26) or on return rates in the following year (17). Studies on longer-term effects of tracking devices on behaviour or habitat choice are lacking. Lesser Black-backed Gulls are omnivores and use a range foraging habitats, including marine, subtidal, and terrestrial habitats, such as agricultural fields and urban areas (6, 21, 27–30). Tracking studies revealed that there is a considerable variation in habitat use between individuals, in part influenced by breeding status and sex (27, 30, 31). In our studies GPS tags are attached using permanent harnesses, thereby affecting feather growth underneath the device. If insulation is disrupted, especially in cold water, gulls could shift their foraging behaviour from sea to land in years following deployment.
In this analysis, we utilized a long-term dataset of 110 GPS-tagged Lesser Black-backed Gulls breeding in three colonies to investigate whether the devices affected habitat choice. Given individual variability in habitat choice, we examined whether birds that spent a relatively high amount of time at sea during the year of tag deployment were less likely to return to the colony in subsequent years. If no relationship was found, we could rule out selective disappearance. As we could not compare the time spent at sea between tagged and untagged individual birds, we investigated whether the time spent at sea decreased over years within individuals. Since a within-individual reduction in time spent at sea could potentially result from gradual environmental changes affecting the entire population, we also compared newly tagged gulls with those tagged in previous years. We hypothesized that if GPS tags caused a shift from sea to land in years following deployment, newly tagged gulls would spent more time at sea than previously tagged ones. Because breeders and non-breeders differ in the time they spend at sea (27), we focused our study on breeding individuals. We expected that if there were an effect, the strongest reduction would occur in the first year after deployment, assuming a lack of regrowth of mantle feathers during post-nuptial moult in the first autumn after tagging.