Behavioral responses of nesting birds to human disturbance along recreational trails

ABSTRACT The presence of recreational trails can impact breeding birds either indirectly by altering habitat or the movement patterns of predators or directly if the presence of humans disturbs birds. We examined the behavioral responses of nesting female Northern Cardinals (Cardinalis cardinalis) to human disturbance using both experimental and observational approaches. From April to August 2006 and 2007, we monitored Northern Cardinal nests in 18 riparian forests in Ohio, USA. Two experimental trials were conducted at each nest (N= 63), with Flight Initiation Distance (FID, the distance at which a bird flushed from the nest) recorded as we approached nests by walking directly toward them and by walking along trails located variable distances from nests. We also measured flight initiation distance (FID) when nests were approached during routine nest checks (N= 160). Cardinals were six times more likely to flush when nests were approached directly, and females on higher nests were less likely to flush regardless of distance to trail. FID was not significantly influenced by the distance of nests from trails. We found no association between nest survival and the tendency of birds to flush. Rather, nest survival was best explained by nest height. Thus, our findings suggest that the responses of birds to human use of recreational trails have only short‐term effects, with no apparent effects of on nest survival. Because the reaction of birds to humans in our study depended on how nests were approached, studies where FID is used as an indicator of sensitivity to human disturbance and is determined by direct approaches may overestimate the potential impact of trails on nesting birds.     

Trophy hunting and perceived risk in closed ecosystems: Flight behaviour of three gregarious African ungulates in a semi‐arid tropical savanna

Although being an important conservation tool in Africa, trophy hunting is known to influence risk perception in wildlife species, thus affecting the behaviour and fitness of most targeted species. We studied the effects of trophy hunting on the flight behaviour of impala (Aepyceros melampus), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in two closed ecosystems, Cawston Ranch (hunting area) and Stanley and Livingstone Private Game Reserve (tourist area), western Zimbabwe. Using standardized field procedures, we assessed the flight behavioural responses of the three species in two seasons: non‐hunting (December–March) and hunting (April–November) between March 2013 and November 2014. We tested the effect of habitat, group size, sex, season, start distance and alert distance on flight initiation distance using linear mixed models. Habitat, group size sex and alert distance did not have any effect on flight initiation distance for the three species. The three species were more alert and displayed longer flight initiation distances in the hunting area compared with the tourist area. Flight initiation distances for the three species were higher during the hunting season for the hunting area and low during the non‐hunting season. Flight distances of the three species did not differ between the hunting area and the tourist area. We concluded that trophy hunting increased perceived risk of wild ungulates in closed hunting areas, whereas ungulates in non‐hunting areas are less responsive and somehow habituated to human presence. Management plans should include minimum approach distances by tourists as well as establishing seasonal restrictions on special zones to promote species viability. Research aimed at integrating behavioural responses with physiological aspects of target species should be promoted to ensure that managers are able to deal with the behavioural trade‐offs of trophy hunting at local and regional scale.     

Determinants of uncertainty in wildlife responses to human disturbance

Outdoor recreation is increasing in intensity and space. Areas previously inaccessible are now being visited by ever‐growing numbers of people, which increases human–wildlife encounters across habitats. This has raised concern among researchers and conservationists as, even in non‐aggressive encounters, animals often perceive humans as predators and mount physiological and behavioural responses that can have negative consequences. However, despite all the research in recent decades, not many general patterns have emerged, especially at the level of populations, and many studies have yielded seemingly contradictory or inconclusive results. We argue that this is partly due to incomplete knowledge of the number and complexity of factors that may modulate the responses of animals. Thus, we aim to provide a conceptual approach intended to highlight the reasons that make it difficult to detect general patterns. We present a comprehensive compilation of factors modulating animal responses to humans at increasing levels (from sensory detection and immediate behavioural and physiological reactions, to changes in fitness and population trends), which may help understanding the uncertainty in the patterns. We observed that there are many modulating factors, which can be categorized as reflecting characteristics of the recreational activity itself (e.g. intensity of human presence), of the animals concerned (e.g. age or antipredatory strategy), and of the spatio‐temporal context (e.g. habitat or timing of the encounter). Some factors appear to have non‐linear and complex effects, which, if not considered, may lead to erroneous conclusions. Finally, we conclude that the difficulty in finding general patterns will be amplified at higher levels (i.e. at the level of populations), since as we proceed from one level to the next, the number of potential modulating factors accumulates, adding noise and obscuring direct associations between recreation and wildlife. More comprehensive knowledge about which (and how) factors affect animal responses across levels will certainly improve future research design and interpretation, and thus, our understanding of human recreational impacts on wildlife.     

Island tameness: living on islands reduces flight initiation distance

One of Darwin''s most widely known conjectures is that prey are tame on remote islands, where mammalian predators are absent. Many species appear to permit close approach on such islands, but no comparative studies have demonstrated reduced wariness quantified as flight initiation distance (FID; i.e. predator–prey distance when the prey begins to flee) in comparison with mainland relatives. We used the phylogenetic comparative method to assess influence of distance from the mainland and island area on FID of 66 lizard species. Because body size and predator approach speed affect predation risk, we included these as independent variables. Multiple regression showed that FID decreases as distance from mainland increases and is shorter in island than mainland populations. Although FID increased as area increased in some models, collinearity made it difficult to separate effects of area from distance and island occupancy. FID increases as SVL increases and approach speed increases; these effects are statistically independent of effects of distance to mainland and island occupancy. Ordinary least-squares models fit the data better than phylogenetic regressions, indicating little or no phylogenetic signal in residual FID after accounting for the independent variables. Our results demonstrate that island tameness is a real phenomenon in lizards.     

Flight initiation distance and escape behavior in the black redstart (Phoenicurus ochruros)

There are many anti‐predatory escape strategies in animals. A well‐established method to assess escape behavior is the flight initiation distance (FID), which is the distance between prey and predator at which an animal flees. Previous studies in various species throughout the animal kingdom have shown that group size, urbanization, and distance to refuge and body mass affect FID. In most species, FID increases if body mass, group size or distance to refuge decreases. However, how age and sexual dimorphism affect FID is rather unknown. Here, we assess the escape behavior and FID of the black redstart (Phoenicurus ochruros), a small turdid passerine. When approached by a human, males initiated flights later, that is allowing a closer approach than females. Males of this species are more conspicuous, and therefore, may exhibit aposematism to deter potential predators or are less fearful than females. Additionally, juveniles fled at shorter distances and fled to lower heights than adults. Lastly, concerning escape strategy, black redstarts, unless other passerine birds, fled less often into cover, but rather onto open or elevated spots. Black redstarts are especially prone to predation by ambushing predators that might hide in cover. Hence, this species most likely has a higher chance of escaping by fleeing to an open spot rather than to a potentially risky cover.     

At home with the birds: Kalahari tree skinks associate with sociable weaver nests despite African pygmy falcon presence

The way in which animals use habitat can affect their access to key resources or how they are buffered from environmental variables such as the extreme temperatures of deserts. One strategy of animals is to modify the environment or take advantage of structures constructed by other species. The sociable weaver bird (Philetairus socius) constructs enormous colonial nests in trees. These nests are frequented by Kalahari tree skinks (Trachylepis spilogaster) and the two species coexist over a large portion of the Kalahari Desert in South Africa. We tested whether skinks were more abundant in trees containing sociable weaver nests and asked whether the physical features of trees were important predictors of skink abundance. We then focused on potential costs of this association by examining the relationship between skink abundance and the presence of a potential predator, the pygmy falcon (Polihierax semitorquatus), which exclusively uses weaver nest colonies for roosting and nesting. Finally, we simulated a predatory threat to determine if skinks assess risk differently if a weaver nest is present. We found a significant positive association between the presence of weaver nests and skink abundance. In the absence of nests, the type of tree did not influence skink abundance. Skinks used weaver nests and were more likely to perch on the nest than the tree. When threatened with predation, skinks preferred to take refuge in nests. Surprisingly, the presence of nesting pygmy falcons in nests did not influence skink abundance, perhaps because of the abundance of nearby refuges within nests, tree crevices, or in debris at the tree base. We suggest that sociable weaver nests provide multiple benefits to skinks including lowered predation risk, thermal refuges and greater prey availability, although this requires experimental testing. In the current era of global climate change, sociable weaver nests may become a crucial resource for skinks seeking refuge as the Kalahari climate warms.     

Consistent avoidance of human disturbance over large geographical distances by a migratory bird

Recent work on animal personalities has demonstrated that individuals may show consistent behaviour across situations and contexts. These studies were often carried out in one location and/or during short time intervals. Many animals, however, migrate and spend their life in several geographically distinct locations, and they may either adopt behaviours specific to the local environment or keep consistent behaviours over ecologically distinct locations. Long-distance migratory species offer excellent opportunities to test whether the animals maintain their personalities over large geographical scale, although the practical difficulties associated with these studies have hampered such tests. Here, we demonstrate for the first time consistency in disturbance tolerance behaviour in a long-distance migratory bird, using the common crane Grus grus as an ecological model species. Cranes that hatched in undisturbed habitats in Finland choose undisturbed migratory stop-over sites in Hungary, 1300-2000 km away from their breeding ground. This is remarkable, because these sites are not only separated by large distances, they also differ ecologically: the breeding sites are wooded bogs and subarctic tundra, whereas the migratory stop-over sites are temperate zone alkaline grasslands. The significance of our study goes beyond evolutionary biology and behavioural ecology: local effects on behaviour may carry over large distances, and this hitherto hidden implication of habitat selection needs to be incorporated into conservation planning.     

Effects of risk assessment, predator behavior, and habitat on escape behavior in Columbian black-tailed deer

The relationship between preflight risk assessment by prey andthe escape behaviors they perform while fleeing from predatorsis relatively unexplored. To examine this relationship, a humanobserver approached groups of Columbian black-tailed deer (Odocoileushemionus columbianus), varying his behavior to simulate moreor less threatening behavior. We measured the focal deer's angleof escape, distance moved during flight, duration of trottingand stotting behavior, and change in elevation during flight.Analyses revealed positive relationships between the distancemoved during flight and the distance at which they fled. Whenflight was initiated when the approacher was close, deer fledrelatively shorter distances and took flight paths at more acuteangles, a property that would force a real predator to changedirection suddenly. Our results indicate that deer do not compensatefor allowing the observer to approach more closely by fleeinggreater distances. Rather, distance moved and flight initiationdistance are linked by level of reactivity and habituation:more reactive or less habituated deer both flee at a greaterdistance and move away to a greater distance during flight.More threatening behavior by the approacher led to longer durationsof rapid flight behavior (e.g., trotting and stotting), anddeer tended to flee uphill and into taller vegetation, usingthese landscape features as refuge from danger. Finally, weprovide the first evidence for Pitcher's untested "antiambush"hypothesis for the function of stotting and discuss its significance.In general, both preflight predator behavior and habitat featuresinfluence both duration and direction of escape.     

Effects of human disturbance on liana community diversity and structure in a tropical rainforest,Malaysia: implication for conservation

Aim Due to the important role of lianas in the functioning of forest ecosystem, knowledge of the factors that affect them are important in the management of forests. Currently, there are conflicting reports on the response of liana communities to disturbance, calling for more research in the area. The present study was carried out to investigate the response of liana diversity and structure to human disturbance within two major forests in the Penang National Park, Malaysia. The study also looked at the implication of the findings for conservation.Methods A total of 15 40 × 40-m 2 (or 40-m × 40-m) plots each were randomly located across a range of habitats in a primary forest and disturbed secondary forest. Trees with diameter at breast height ≥10 cm were examined for lianas with diameter ≥2 cm. Both lianas and trees were enumerated and compared between the two forests. Diversity and structural variables of lianas were compared between the two forests using the t -test analysis. Tree abundance was also compared between the two forests with t -test, while linear regression analysis was run to determine the effects of tree abundance on liana abundance.Important findings A total of 46 liana species belonging to 27 genera and 15 families were identified in the study. Human disturbance significantly reduced liana species richness and species diversity in the secondary forest. Liana abundance remained the same in both forests whereas liana basal area was significantly higher in the primary forest. Twiners and hook climbers were significantly more abundant in the primary and secondary forest, respectively. Large diameter lianas were more abundant in the primary forest compared with the secondary forest. The diameter distribution of most families in the primary forest followed the inverted J-shaped curve whereas only a few of the families in the secondary forest did so. Tree abundance was significantly higher in the primary forest. The abundance of lianas significantly depended on tree abundance in all the forests. The study has provided evidence of negative effects of human disturbance on liana diversity and structure that does not auger well for biodiversity in the forest. In view of the critical role of lianas in maintaining biodiversity in the forest ecosystem, lianas in the national park should be protected from further exploitation.     

A meta‐analysis of human disturbance impacts on Antarctic wildlife

Evidence‐based assessments are increasingly recognized as the best‐practice approach to determine appropriate conservation interventions, but such assessments of the impact of human disturbance on wildlife are rare. Human disturbance comprises anthropogenic activities that are typically non‐lethal, but may cause short‐ and/or longer‐term stress and fitness responses in wildlife. Expanding human activity in the Antarctic region is of particular concern because it increases the scope and potential for increased human disturbance to wildlife in a region that is often thought of as relatively untouched by anthropogenic influences. Here, we use a meta‐analytical approach to synthesise research on human disturbance to wildlife over the last three decades in the Antarctic and sub‐Antarctic region. We combine data from 62 studies across 21 species on the behavioural, physiological and population responses of wildlife to pedestrian, vehicle and research disturbances. The overall effect size indicated a small, albeit statistically significant negative effect of disturbance (?0.39; 95% CI: ?0.60 to ?0.18). Negative effects were found for both physiological and population responses, but no evidence was found for a significant impact on wildlife behavioural responses. Negative effects were found across pedestrian, vehicle and research disturbances. Significant and high among‐study heterogeneity was found in both disturbance and response sub‐groups. Among species, it remains unclear to what extent different forms of disturbance translate into negative population responses. Most current guidelines to limit wildlife disturbance impacts in Antarctica recommend that approaches be tailored to animal behavioural cues, but our work demonstrates that behavioural changes do not necessarily reflect more cryptic, and more deleterious impacts, such as changes in physiology. In consequence, we recommend that pedestrian approach guidelines in the Antarctic region be revisited. Due to the high heterogeneity in effects, management guidelines for different sites and species will need to be developed on a case‐by‐case basis, ideally in conjunction with carefully designed experiments. Guidelines to reduce the impact of research activities per se require development to reduce the potential impacts of conducting research. We identify research questions that, if answered, will further improve the evidence base for guidelines to manage human disturbance in Antarctica.     

Effects of Recreation and Hunting on Flushing Distance of Capercaillie

Abstract: The capercaillie (Tetrao urogallus) in Central Europe is an endangered species of grouse that is thought to be highly susceptible to human disturbance, possibly causing local populations to decline. We investigated the behavioral response of capercaillie in the Black Forest, Germany, and the French Pyrenees to an off-trail hiker by measuring flushing distances. Flushing distance varied with capercaillie sex, visibility of the hiker, intensity of winter tourism, and hunting pressure. Independent of the study area, males flushed at consistently longer distances than females, and lower visual blocking between bird and hiker resulted in longer flushing distances. Capercaillie flew at longer distances from an approaching hiker in areas with high intensity of winter tourism or hunting pressure than in undisturbed areas. We recommend the establishment of regulations requiring hikers to stay on trails and to close trails where intertrail distances fall below 100 m (90% of all flushing events appeared within 50 m). Furthermore, planting or preserving evergreen conifer trees in dense rows along critical parts of disturbance sources, reducing the degree of visibility between capercaillie and recreationists would increase habitat for capercaillie in forests with predictable recreation activities.     

Population genetics reveals high connectivity of giant panda populations across human disturbance features in key nature reserve

The giant panda is an example of a species that has faced extensive historical habitat fragmentation, and anthropogenic disturbance and is assumed to be isolated in numerous subpopulations with limited gene flow between them. To investigate the population size, health, and connectivity of pandas in a key habitat area, we noninvasively collected a total of 539 fresh wild giant panda fecal samples for DNA extraction within Wolong Nature Reserve, Sichuan, China. Seven validated tetra‐microsatellite markers were used to analyze each sample, and a total of 142 unique genotypes were identified. Nonspatial and spatial capture–recapture models estimated the population size of the reserve at 164 and 137 individuals (95% confidence intervals 153–175 and 115–163), respectively. Relatively high levels of genetic variation and low levels of inbreeding were estimated, indicating adequate genetic diversity. Surprisingly, no significant genetic boundaries were found within the population despite the national road G350 that bisects the reserve, which is also bordered with patches of development and agricultural land. We attribute this to high rates of migration, with four giant panda road‐crossing events confirmed within a year based on repeated captures of individuals. This likely means that giant panda populations within mountain ranges are better connected than previously thought. Increased development and tourism traffic in the area and throughout the current panda distribution pose a threat of increasing population isolation, however. Maintaining and restoring adequate habitat corridors for dispersal is thus a vital step for preserving the levels of gene flow seen in our analysis and the continued conservation of the giant panda meta‐population in both Wolong and throughout their current range.     

Antipredator behaviour of African ungulates around human settlements

As human populations grow and come into more frequent contact with wildlife, it is important to understand how anthropogenic disturbance alters wildlife behaviour. Using fine‐scale spatial analyses, we examined how proximity to human settlements affects antipredator responses of ungulates. We studied seven common ungulate species (Kirk's dik‐dik, Thomson's gazelle, impala, common warthog, common wildebeest, common zebra and Masai giraffe) in the Tarangire–Manyara ecosystem in northern Tanzania. In zebra and giraffe, flight responses to humans were significantly more likely when closer to settlements; however, there was a weak relationship between flight responses and distance to settlement in all other species. While there was largely a weak relationship between proximity to human settlements, the distribution of settlements in the landscape appears to affect wildlife behaviour, suggesting that animals perceive and respond to spatial variation in risk exerted by humans.     

Myrmecochory in Australia's seasonal tropics: Effects of disturbance on distance dispersal

Abstract Successful ecosystem restoration requires the re-establishment of fundamental ecological processes, many of which involve plant-animal interactions. Myrmecochory (seed dispersal by ants) is a particularly important plant-animal mutualism in Australia, but little is known about its response to either disturbance or restoration following disturbance. Here we investigate the effects of disturbance on seed dispersal by ants, and the extent to which the ant-seed relationship has re-established at sites undergoing rehabilitation, at Ranger uranium mine in the seasonal tropics of Australia's Northern Territory. We focused on the composition of seed-dispersing ant assemblages, rates of seed removal by ants, and the dispersal curves generated by ants, as determined by observations of removal from seed depots. Ten sites were studied, comprising four ‘natural’ (undisturbed) sites representing a range of savanna habitats occurring in the region, four disturbed sites representing a range of habitat disturbance but with intact soil, and two waste rock sites subject to preliminary revegetation trials. A total of 22 ant species from 10 genera were observed during 154 observations of seed removal, most commonly Rhytidoponera aurata (53 records), Monomorium (rothsteini gp) sp. 1 (14), Iridomyrmex sanguineus (13), Iridomyrmex pallidus (12) and Pleidole sp. 3 (10). Removal rates (over 3 h) averaged 29% across all sites and time periods, varying markedly both between and within sites. However, mean rates of removal were similar between natural, disturbed and waste rock sites (29%, 28% and 31%, respectively). A high incidence (62% of all depots) of'aril robbing’ by ants (primarily Monomorium spp.) eating arils in situ, without removal, was observed. Dispersal distances varied markedly between ant species, with Iridomyrmex sanguineus having both the highest mean (7.25m) and maximum (13.08 m) dispersal distances. Species of Pheidole typically dispersed seeds less than 0.5 m, and Meranoplus, Monomorium and Tetramorium spp. only ever moved seeds a few centimetres, usually dropping and abandoning them before reaching the nests. The dispersal curves characteristic of each site varied markedly due to the different composition of seed-dispersing ants. The mean dispersal distance at disturbed sites (3.91 m) was significantly higher than at natural sites (2.19 m), and the curves were strongly skewed in the former, but relatively uniform in the latter. The implications of these differences for recovery following disturbance are unclear. At rehabilitated waste rock sites, all observed removals involved distances less than 0.5 m, with a mean of 17 cm. This lack of effective ant-seed relationships might represent a barrier to further vegetation development at rehabilitated sites.     

Fifty years of chasing lizards: new insights advance optimal escape theory

Systematic reviews and meta‐analyses often examine data from diverse taxa to identify general patterns of effect sizes. Meta‐analyses that focus on identifying generalisations in a single taxon are also valuable because species in a taxon are more likely to share similar unique constraints. We conducted a comprehensive phylogenetic meta‐analysis of flight initiation distance in lizards. Flight initiation distance (FID) is a common metric used to quantify risk‐taking and has previously been shown to reflect adaptive decision‐making. The past decade has seen an explosion of studies focused on quantifying FID in lizards, and, because lizards occur in a wide range of habitats, are ecologically diverse, and are typically smaller and differ physiologically from the better studied mammals and birds, they are worthy of detailed examination. We found that variables that reflect the costs or benefits of flight (being engaged in social interactions, having food available) as well as certain predator effects (predator size and approach speed) had large effects on FID in the directions predicted by optimal escape theory. Variables that were associated with morphology (with the exception of crypsis) and physiology had relatively small effects, whereas habitat selection factors typically had moderate to large effect sizes. Lizards, like other taxa, are very sensitive to the costs of flight.     

Life history,predation and flight initiation distance in a migratory bird

Life‐history trade‐offs occur as a consequence of the compromise between maximization of different components such as the size and the number of clutches. Flight initiation distance (FID) potentially constitutes a general proximate factor influencing such trade‐offs reflecting the risks that individuals take. Therefore, greater investment in reproduction occurs at a higher risk of death, resulting in selection for efficient flight morphology. I analysed long‐term data on FID in a population of barn swallows Hirundo rustica during 1984–2013 with 2196 records of FID for 1789 individuals. FID had a repeatability of 0.62 (SE = 0.04) and a heritability of 0.48 (SE = 0.07). FID varied between individuals and sites, and it increased over time as climate ameliorated. FID showed a U‐shaped relationship with age, with young and very old individuals having the longest FIDs. Barn swallows that arrived early from spring migration, started to breed early and produced many fledglings had the longest FID. Individuals with the longest tails had the longest FID, and individuals with the shortest aspect ratios and wing loadings had the longest FID. Individuals that died from predation had shorter FID than survivors. These findings are consistent with the hypothesis that FID relates directly to life history, with longer FIDs being associated with smaller levels of risk‐taking.     

Magnitude of food reward affects escape behavior and acceptable risk in Balearic lizards, Podarcis lilfordi

During encounters with predators, prey must balance the degreeof risk against the loss of fitness-enhancing benefits suchas feeding and social activities. Most studies of tradeoffsbetween risk and cost of escaping have measured flight initiationdistance and time to emerge from refuge, for which theory providesrobustly supported predictions. Tradeoffs involving other aspectsof encounters, including distance fled and time between escapeand return to a food source, have received little theoreticalor empirical attention. By adapting models of flight initiationdistance and time between entry into refuge and emergence, wepredict effects of predation risk and cost on distance fledand time to return to a source of benefit after fleeing. Actingas simulated predators that approached at a fixed speed, weconducted an experimental field study to test the hypothesesthat flight initiation distance, distance fled, and time toreturn to food by Balearic lizards (Podarcis lilfordi) decreasewith the presence and amount of insect food. Predictions ofthe models were strongly supported, including those for distancefled and return time, but predictions for other cost factorsand predation risk factors remain to be tested.     

Effect of Approaching Boats on Nesting Black Skimmers: Using Response Distances to Establish Protective Buffer Zones

ABSTRACT Understanding how birds respond to the activities of people is an important component of conserving wildlife. We measured responses of nesting black skimmers (Rynchops niger) to an approaching boat in Barnegat Bay, New Jersey, USA, by examining distance to first respond, distance to flush, and time to return to the colony. Our objective was to determine if response distances of skimmers changed as a function of year, reproductive stage, direction of approach (direct or tangential), or number of birds nesting in the colony. Generally, reproductive stage had the greatest effect on all responses, followed by direction of approach, number of adults present at the colony, number of nests, and year, which also explained variation in behavioral responses. The distance at which skimmers first flew when a boat approached decreased from the pre-egg-laying period to hatching, and then increased slightly later in the season. Time (x̄± SE) for skimmers to return to the nesting colony varied seasonally, with birds taking longer to return during the pre-egg period (9.5 ± min) than during hatching (0.7 ± 0.1 min). The decision process for determining set-back distances to protect nesting skimmers should involve selecting 1) behavioral response of highest concern, 2) reproductive stage of highest concern, and 3) an appropriate level of response at which to establish the buffer area. We recommend that managers use a set-back distance of ± 118 m from the perimeter of the colony for black skimmers, which is the 95% percentile of the distance that skimmers first flew in response to approaching boats. Managers can use these data to set buffer distances for skimmers and other colonial birds.