Research Paper

INTRODUCTION

Mining and other forms of resource extraction are important economic drivers in Canada’s North because they provide well-paying, stable jobs for people living in rural northern communities (Cameron and Levitan 2014, Belayneh et al. 2018). In 2014, 18% of the gross domestic product of Nunavut, Canada, was associated with resource extraction (AMAP 2017). Mineral, oil, and gas exploration are predicted to grow across the Arctic landscape (Kondratiev 2020), leading to land-use changes and disturbance of wildlife (Wilson et al. 2013). Although resource extraction plays an important role in northern economies, balancing economic growth with conservation and respect for Indigenous rights will be a challenge (Arbo et al. 2012, Bernauer 2019, Tolvanen et al. 2019).

Because of the limited human footprint, Arctic ecosystems are often considered secure from human-induced change. However, the effects of resource extraction on habitat loss can be substantial at a local or regional scale. For example, Pirie et al. (2009) predicted an 8 to 30% decrease in nesting habitat for Whimbrel (Numenius phaeopus) while running scenario-based analyses of subsidence-induced flooding related to natural gas extraction and development within the Kendall Island Bird Sanctuary (623 km²) and Fish Island (7560 km²) in the outer Mackenzie Delta, Northwest Territories. An increase in resource extraction in northern landscapes means a greater likelihood of loss of nesting habitat (Gajera et al. 2013, Bernath-Plaisted and Koper 2016). More subtle effects are also possible, mines may have a relatively limited effect on northern species (Smith et al. 2005, Merondun et al. 2024; see also https://www.nwtgeoscience.ca/forum/session/long-term-monitoring-finds-no-effect-Arctic-mine-migratory-birds ), but mines and associated infrastructure may alter behavior of breeding birds and, as a result, nest success (Male and Nol 2005, Van Wilgenburg et al. 2013, Ludlow and Davis 2018).

Some resource extraction activities can be timed to minimize disturbance to nesting birds, but other activities must be carried out at times that conflict with the timing of breeding of birds. For example, large-scale dewatering of lakes for mining pit development often occurs during the spring and early summer, adding to run-off on the tundra because water is directed elsewhere. This additional run-off during the breeding season can lead to destruction of nesting areas, or direct loss of nests, through flooding (Pirie et al. 2009, Gajera et al. 2013).

The Migratory Birds Convention Act (MBCA; Parliament of Canada 1994) prohibits disturbance, or destruction of migratory birds and their nests, including unintentional harm through incidental take, which is defined as the inadvertent harming, killing, disturbance, or destruction of migratory birds, nests, and eggs (CWS 2014). Nest losses due to mining development are considered incidental take (Van Wilgenburg et al. 2013) and are a concern in many activities related to resource development, including vegetation clearing, road or pipeline maintenance, and flooding from dams used to control water on the landscape. Although incidental take of migratory birds or their nests is sometimes unavoidable when carrying out otherwise permitted activities, industry has a legal requirement to understand and mitigate the impacts.

Deterrents can be used to discourage birds from occupying mining sites that may be harmful to them (Boag and Lewin 1980, Ronconi and St. Clair 2006). Deterrents are used to convey to birds, through perceived predation risk, that prospective nesting, feeding, or roosting areas are not suitable, thus discouraging birds from settling in or remaining in areas where human activities might cause them harm (Ball 2000, Schlichting et al. 2017) or where humans identify birds as pests. For example, deterrents such as robotic peregrine falcon effigies, audio calls of predators, high-intensity strobe lights, and propane cannons are used at tailing ponds in the oil sands region of Alberta, Canada, to prevent birds from using these contaminated waters (Ronconi et al. 2004). Although deterrents have been used in various situations, there is limited research on the effectiveness of various deterrent technologies (Marcus et al. 2007, Kruk et al. 1997), particularly in Arctic regions (Umlah 1996, Racca 2005).

Agnico Eagle Mines Ltd. proposed and built the Whale Tail Project in central Nunavut, Canada as an expansion to their gold mining activities, the Meadowbank Complex, which had first become commercially productive in 2010 (Agnico Eagle Mines Ltd. 2024). This project included the construction of two dikes within the northern portion of Whale Tail Lake that diverted water (dewatering) from the Whale Tail mining pit into surrounding lakes and tributaries (Fig. 1). One dike resulted in flooding that elevated water levels by 4 m (pre-flood water elevation: 152 m.a.s.l., post-flood 156 m.a.s.l.) from May 2019 to August 2020, causing approximately 157 ha of tundra adjacent to the lakes to flood. This flooding coincided with timing of bird nesting over two breeding seasons. We were asked by mine personnel to test potential solutions that would deter birds from nesting within the predicted flood area during dewatering, to minimize the number of bird nests lost (i.e., incidental take) due to flooding.

Our objective was to test the effectiveness of visual and audio deterrents to prevent birds from nesting in tundra areas exposed to mining-induced flooding. We accomplished this objective with the use of an experimental design applied to plots located outside the proposed flood area. We hypothesized that deterrent use would dissuade Arctic-nesting birds from establishing nests. We also assessed the hypothesis that, if birds failed to be deterred and nested in plots with deterrents that they would exhibit a change in behavior from disturbance by deterrents, and that these behavioral changes could potentially impact daily nest survival. Changes in behaviors due to deterrents could include limiting time on nests or outright abandoning active nests. We compared (1) territory densities, (2) daily nest survival of passerines and apparent survival of shorebirds before and after deterrent deployment, and (3) incubation behavior of Lapland Longspur (Calcarius lapponicus) between plots with and without deterrents. We predicted that density of territorial males would be reduced from pre-treatment levels in plots where we applied deterrents, whereas there should be no such reduction in density in control plots. We predicted that daily nest survival of passerines and apparent survival of shorebirds would be higher in the pre-treatment compared to treatment plots but would remain similar before and after treatments in control plots. We also predicted that incubating female Lapland Longspur would show more frequent and longer incubation off-bouts within deterrent plots than in control plots. We report on the time used in implementing deterrents and their costs to assist others in determining whether the effort and cost are warranted.

METHODS

Study area and design

The study was conducted in the Kivalliq Region of Nunavut, 130 km north of Baker Lake (N 65° 24′ 23.98″, W 96° 40′ 32.68″) in June and July of 2018 and 2019. Situated in the Northern Arctic ecozone, the landscape is primarily tundra heath, lichen, and rock in the uplands and graminoid-dominated wetlands in the lowlands, with scattered lakes and ponds (Campbell et al. 2012). June temperatures in the study area averaged 4.0 °C (min: -0.8, max: 8.7) and 6.7 °C (min: 2.5, max: 10.6) in 2018 and 2019, respectively, and July temperatures averaged 13.5 °C (min: 7.9, max: 18.9) and 10.5 °C (min: 6.7, max: 14.8) in 2018 and 2019, respectively. Precipitation totals for June and July were 132 mm and 139 mm in 2018 and 2019, respectively (Agnico Eagle Mine weather station).

In 2018, we established 15 6-ha plots (300 x 200 m) along Amaruq Road (Fig. 1), 5–17 km south of Amaruq Mine for convenience of access. Plots were placed > 200 m from Amaruq Road to avoid potential impacts of road dust, which can cause early snowmelt and in turn, impact nest timing and success (Smith et al. 2010, Grabowski et al. 2013). These plots were located 13–18 km southeast from the area that was slated for flooding. The vegetation and bird communities within these study plots were primarily lowland wet sedge meadows, like those in nearby flooded areas (Holmes 2022).

We used a before-after control-impact study design (BACI; Smokorowski and Randall 2017). Study plots were grouped into five clusters of three plots, with each cluster consisting of one control plot and two deterrent plots. In each cluster, plots were randomly assigned to control or treatments. The same plots were used in the study in both years, with no deterrents deployed in 2018, and the deterrent treatments applied in 2019. The distance between clusters of plots was 1–9 km (Fig. 1). The location of plots was constrained by the need to select for lowland habitats.

We deployed deterrents from 6–17 June 2019, before most nests were initiated. The two deterrent treatments differed in their intensity (Table 1). The high-intensity treatment included a hawk-kite effigy, an audio deterrent system with four speakers, and ribbons of flash tape, distributed evenly throughout the 6-ha plot. The low-intensity treatment contained only a hawk-kite effigy and an audio deterrent system with four speakers (Table 1).

We erected hawk-kite effigies using 5.2 m tall fiberglass pole in the middle of each treatment plot, near to the central unit of the audio deterrent system. The hawk-kite effigies were attached to the top of the pole with a kite string, where they soared in the wind. We used two different brands of hawk-kites: Peregrine Falcon kite (Margo Supplies, High Water, Alberta) placed in the high-intensity treatment and Birds of Prey Falcon kite (Sutton Agricultural Enterprises, Salinas, California), placed in the low-intensity treatment (Table 1). Hawk-kite effigies were similar in size and structure, but with different materials and coloration.

Audio deterrents (Super BirdXpeller® PRO, Bird-X, Inc., Elmhurst, Illinois) consisted of a central unit with four speakers (sound pressure: 110–115 dB at 1 m). We placed the main unit in the middle of the 6-ha plot, with each speaker attached to a wooden stake, hammered into the tundra at a height of 1 m, at each cardinal direction, 25 m from the main unit. The Bird X unit played a mix of distress calls from American Crow (Corvus brachyrhynchos) and generalist predator calls (Ring-billed Gull, Larus delawarensis; Common Raven, Corvus corax; Blue Jay, Cyanocitta cristata; Black-billed Magpie, Pica pica; Peregrine Falcon, Falco peregrinus). Among these, only Common Raven and Peregrine Falcon occur in the area. Recorded calls of locally relevant predators (e.g., Arctic fox, Vulpes lagopus; Arctic ground squirrel, Urocitellus parryii; Rough-legged Hawk, Buteo lagopus; Parasitic Jaeger, Stercorarius parasiticus) are not available on these units. Although we acknowledge this large list of potential predator calls as a limitation of our design, other work has suggested that animals should respond to generalized predator calls especially if the calls are from taxonomically similar species (Ferrari et al. 2007). In the high-intensity plots, the four speakers in each plot played all at once, broadcasting one of the six predator calls at a time in random order continuously for 24 hr at 10–30-min intervals. In the low-intensity plots, one of the four speakers in each plot played one predator call at a time with the predator calls broadcasting sequentially (e.g., speaker 1 played Blue Jay, speaker 2 played Common Raven, etc.) continuously for 24 h at 5–10-min intervals. Both patterns of audio deterrent were broadcast from 10 June to 16 July. To maintain electrical power for the audio deterrents throughout the experiment, speakers were powered with 40-Ah 12-v car batteries and 25-W solar panels (Fig. 2).

A grid was erected by stringing monofilament fishing line (100-lb test, Hercules PE braided fishing line, 4 Strands) between 1.5 m long, narrow (1.5 x 1.5 mm) aluminum angle stakes that we hammered 0.5 m into the tundra every 60 m across the 300 m length of the study plot, and every 20 m along the 200 m length of the plot (Fig. 2). Every 20 m along the monofilament lines, we affixed a piece of flash tape (Birdscare Flash tape, Sutton Agricultural Enterprises, Salinas, California): a metallic tape with red and silver on opposite sides, 30 mm wide and 0.025 mm thick. Each flash tape piece was 5 m long, with one end knotted to the fishing line and the other end free to sway with the wind.

Deterrents were maintained as needed every four days during nest monitoring by tightening the fishing line, replacing flash tape, ensuring that the hawk-kite effigy was still intact, and checking electrical connections and power supply for audio deterrents. Because effort required to use deterrents is an important practical consideration, we recorded the number of person-hours spent erecting and maintaining the deterrents and recorded the full cost to purchase the equipment and supplies used.

Species composition, territory mapping, nest searching, and monitoring

The breeding bird population in the study area consisted primarily of Arctic-nesting passerines, with lower densities of shorebirds and ducks. The four most common species were Lapland Longspur, Semipalmated Sandpiper (Calidris pusilla), Least Sandpiper (Calidris minutilla), and Horned Lark (Eremophila alpestris). Lapland Longspur and the two sandpiper species nest in low elevation sedge meadows, close to water bodies. Horned Lark nest at slightly higher elevations in adjacent upland, dry habitats (Camfield et al. 2010).

Nest searching, monitoring, and territory mapping occurred 4 June to 15 July. Territory mapping was most intensive at the beginning of each breeding season (4–25 June), after male birds arrived and began to sing and display. Territory mapping continued throughout the breeding season when males were observed singing and or in display. Once nesting began, there were fewer breeding displays. Mapping was done by following displaying males and recording the bird’s location with a waypoint (± 3 m) using a Garmin GPS. We aimed to obtain a minimum of five waypoints for each territorial bird, every four days. While following males, we attempted to avoid disturbing them by staying low to the ground and moving only after the bird moved to a new position to sing. Because the habitat was very open, we were able to watch individual males move to new singing locations. When uncertain about the territory to which a singing bird belonged, we omitted the observations from calculations of territory size.

Nest searching was conducted by systematically walking plots and observing behavioral cues of breeding adults (e.g., flushing, mate courtship, alarm calls). Nests were marked with a tongue depressor driven partially into the ground within 5 m of the nest, in a random direction. The observer recorded nest coordinates using the average waypoint function in the Garmin GPS unit, as well as species, number of eggs present, and date found. The average waypoint function with Garmin is a method of averaging multi-sample waypoints to obtain a more accurate GPS position, facilitating nest relocation. For nests found outside of the laying period, we floated eggs to determine nest age and to estimate initiation date and hatch date (adapted from Liebezeit et al. 2007).

Nests were monitored on a 4-day schedule until fates were determined. Nests of shorebirds with precocial young were considered successful if at least one egg hatched, whereas passerine nests with altricial young were considered successful if at least one chick fledged. Signs of predation (loss of a whole clutch, nest disturbance, large eggshell fragments, or yolk) or abandonment (no sign of adults or cold eggs; Mabee et al. 2006) indicated a failed nesting attempt.

Scientific permits were issued by Environment and Climate Change Canada under Migratory Birds Regulations Sections 4, 12, and 19. All animals used in this research paper were cared for in accordance with the guidelines set out by the Canadian Council on Animal Care and all methods regarding use of animals for this project were reviewed and approved by the Trent University Animal Care Committee.

Incubation behavior of Lapland Longspur, nest concealment, and weather

In 2019, we deployed Tinytag© (Tiny Tag Plus 2 Logger, TGP-4020, Gemini Data Loggers) temperature probes in nests of Lapland Longspurs located within treatment plots of either intensity that were < 20 m from flash tape or < 150 m from a deterrent audio speaker, and in nests within control plots to determine on-off bouts of incubating females. We placed 20 temperature probes within Lapland Longspur nests in 2019: 10 nests in control plots and 10 in treatment plots. Incubation data were obtained from 21–30 June and 3–4 July. We deployed one temperature probe in a Lapland Longspur renesting attempt from 10–18 July.

We inserted the temperature probe of the Tinytag© into the nest so that it was in contact with the brood patch of the incubating female. The data logger unit was secured under peat near the nest so as not to attract predators or rouse suspicion of incubating birds. Probes recorded a temperature every minute, providing a high-resolution dataset with which to evaluate incubation behavior (Joyce et al. 2001, Schneider and McWilliams 2007).

At each nest, we estimated vertical concealment as percentage of the nest concealed by vegetation when viewed from directly above. Ambient temperature data were collected from a weather station located at Amaruq Mine, Nunavut, operated by Agnico Eagle Mines Limited (N 65° 24′ 23.98″, W 96° 40′ 32.68″).

Statistical analysis

Territory densities

Territory maps that included a minimum of five waypoint locations of a singing male were delineated using minimum convex polygons (MCPs) in the R Studio package adehabitatHR (version 1.2.1335; Calenge 2006, RStudio Team 2020). We used MCP’s to estimate relative rather than absolute territory size between treatments because it is a simple and commonly used estimator (Haenel et al. 2003). Nests in plots that were not associated within a delineated territory were given an assumed territory size by using the st_buffer function in sf package to assign a circular spatial buffer around the nest (Pebesma 2018). This buffer size was based on the average relative territory size that we documented for each species in the study, based on the MCP approach, i.e., average territory size (± SD) of Lapland Longspur = 3988 ± 4750 m²; Horned Lark = 7258 ± 9012 m²; Semipalmated Sandpiper and Least Sandpiper = 4295 ± 3646 m². We used the st_intersection function in the sf package (Pebesma 2018) to determine the proportion of each territory (MCP or buffer based) that occurred within the boundaries of the plots.

We tested for significance of differences in territory densities in our BACI design, by examining the interaction term between year (before/after) and treatment (Smokorowski and Randall 2017). We summed the count of territories in a plot (including the fraction within the plot for those that straddled the border) and converted them into a density (territories per km²) in each 6-ha plot, separately for each year (2018, 2019). We used a generalized linear mixed-effects model using the lme4 and lmerTest packages in R Studio (Bates et al. 2015, Kuznetsova et al. 2016) with density of territories as the response variable and treatments (high-intensity treatment, low-intensity treatment, and control, Table 1), year (2018 or 2019) and their interaction as fixed effects. We included an offset of observer-hours within the model to account for the number of hours spent nest searching and territory mapping in each plot. This was because we spent more time in treatment plots due to the need for deterrent maintenance. We included cluster as a random effect to account for the geographic association of plots within clusters.

This statistical model was first applied to assess potential changes in territory densities for all species combined, then for two shorebirds, and individually for Lapland Longspur and Horned Lark. Additionally, we assessed whether there was a difference in relative territory size of Lapland Longspur between treatment and control plots using a linear model (lm). We assessed the differences in relative territory size only for territories that had at least 50% of their area within a plot.

Daily survival rate of nests

We calculated daily nest survival rates (Mayfield 1961, Dinsmore and Dinsmore 2007) using the RMark package (Laake 2013) in R Studio for passerines because they had a sufficient sample size for such analyses. We started with 90 nests but excluded 13 nests with unknown fate. We converted ordinal dates into nest-specific dates (hereafter season-date) in which the first initiation date was season-date 0 (on 10 June, ordinal date 161, for passerines) and the final nest check was season-date 36 (16 July, ordinal date 197).

In addition to examining effects of treatment on daily nest survival, we assessed the effect of nest age (i.e., the age of a nest in days, where age 0 = the day that the first egg was laid) because this predictor variable is often important in explaining daily nest survival (Grant et al. 2005). We eliminated three nests from these analyses in which nest initiation date (and therefore nest age) was unknown. In the first stage of modeling, we ran six candidate models (including an intercept-only model) consisting of temporal variables: nest age, season-date, quadratic season-date (date²), cubic season-date (date³), and year. We found that nest age was not a good predictor of daily nest survival because it was not in a top or competitive model as judged by Akaike’s information criterion adjusted for small sample size (AIC_c; i.e., ΔAIC_c > 6; Akaike 1973, Burnham et al. 2011). Therefore, we removed nest age from the model, which allowed us to add back the three nests lacking estimates of nest age (n = 77).

We then re-ran models consisting of temporal variables: season-date, quadratic season-date (date²), cubic season-date (date³), and year. We carried the top temporal model (date + date²) from this stage of modeling forward, as a foundation for testing the impacts of treatment on daily nest survival, by considering variables: treatment, year, and the interaction of year and treatment. Lastly, we used AIC_c (Hurvich and Tsai 1991) to assess relative model support. We considered models with ΔAIC_c*** < 2 as competitive. In addition, only variables with 95% confidence intervals not overlapping zero were considered important for explaining nest daily survival rate (Fromberger et al. 2020).

We had too few nests to statistically compare daily nest survival rates between treatments for Semipalmated Sandpiper (n = 23) and Least Sandpiper. Therefore, we report apparent nest survival (proportion of successful nests out of total nests) among years and treatments for these species.

Incubation behavior

We assumed that when Tiny Tags registered temperatures within 3 °C of 39 °C that incubating Lapland Longspur females were present on the nest (“on bouts”) and when nest temperature dropped to the ambient surrounding temperature that incubating birds had left the nest for an incubation recess (“off-bouts”). We used the program RHYTHM (1.0; Cooper and Mills 2005) in combination with Raven Lite (2.0) to automate measurements of length of off-bouts. We considered a drop in temperature greater than 3 °C within 1 minute to signal the start of an off-bout. This criterion differs slightly from that used in similar studies of Horned Lark (Camfield and Martin 2009, MacDonald et al. 2013), which considered a recess as a drop of > 3 °C in the nest and a decline in temperature that lasted more than 3 minutes. We chose our criteria to account for swift off-bouts that may last for less than three minutes, in situations where birds are potentially disturbed frequently by deterrents such as the flash tape. Results of the recesses estimated by Raven Lite (2.0) were manually examined for quality control. We manually added off-bouts in accordance with the above criterion where they were missed.

In a preliminary inspection of data, we noticed a greater number of recesses and longer recesses during daytime hours (6:00–18:00 CST) than during typical nighttime hours (18:00–6:00 CST), despite an average 20–21 hr of daylight in the study area during June and July (Government of Canada 2020). Therefore, to account for the effect of time of day (Camfield and Martin 2009), we separated data into periods of daytime and nighttime hours. We removed the first and last time-of-day period of the temperature recordings (i.e., immediately post-deployment and pre-retrieval) to account for the lack of an entire 12-hr period of incubation data. We calculated the number of off-bouts and proportion of time spent off the nest during each 12-hr period of night and day, for each incubation day of a nest. To evaluate whether incubation rhythm was impacted by treatment type, we used linear mixed-effects models using a maximum likelihood method of parameter estimation in the statistical package lme4 (Bates et al. 2015) in R Studio. We also removed data recorded on 1–2 July 2019 from the analysis due to two days of inclement weather that kept birds off their nests for over 12 hr. We focused analyses on incubation days 3–10, because sample size was insufficient for comparison of treatment versus control on days 1, 2, 11, and 12.

We modeled two response variables separately: proportion of time off the nest and number of off-bouts within 12-hr periods and included treatment type and nest fate as fixed effects and nest ID as a random factor to account for repeated sampling of individual nests. Because nest incubation patterns can potentially vary as a function of ambient temperature, time of day (day/night), and stage in the incubation cycle (nest age; Wiebe and Martin 1998, Camfield and Martin 2009, Ricklefs and Brawn 2012), we included these variables as fixed effects. Only one of the 20 nests that we monitored was known to be a second nesting attempt (i.e., a re-nest). Therefore, we did not include renesting as a factor in the analysis. We used AIC_c (Hurvich and Tsai 1991) to determine the strongest model.

We used an alpha of 0.05 to assess statistical significance for all analyses including for relative territory densities, daily nest survival, and incubation behavior.

RESULTS

Relative territory densities

Over the 2 years of study, we mapped a total of 118 territories of the focal species that had a portion of their area within the study plots. Of these 118 territories, 60 were not associated with a nest that we located, and 58 were associated with at least 1 nest. We found an additional 66 (53%) nests that did not meet the criteria for spot-mapping (5 waypoints per territory) and assigned the average territory size to these territories. The average start date of territory determination for passerines was 18 June and 9 June in 2018 and 2019, respectively, and for shorebirds 19 June and 15 June in 2018 and 2019, respectively.

Densities of territorial males for the four study species combined (Table 2) ranged among plots from 56.1–104.5 territories/km². Lapland Longspur were the most abundant species in the plots, followed by the two shorebird species, and then Horned Lark. Deterrents did not reduce the density of territorial birds within the plots. There was no significant interaction between year and treatment for the model including all study species together (F_2,20 = 0.54, p = 0.59). However, there was a significant increase in the densities of territorial males between 2018 and 2019 for all treatments (F_1,20 = 4.58, p = 0.04), with the cold and late snowmelt year of 2018 having a lower density of birds than in 2019 (average air temperature, 1 June to 10 June, in 2018: -1.35 °C; in 2019: 1.99 °C).

Densities of shorebirds averaged between 8.2–23.8 territories per km² (Table 2). Although average shorebird densities increased by 65% between 2018 and 2019 in the high-intensity treatment, and decreased in the controls by 8%, there was no significant interaction between year and treatment for densities of shorebirds (Table 2). Lapland Longspur territory densities ranged from 41.4–76.7 territories per km², also exhibiting an 42% increase in the high-intensity treatment and a 13% increase in the control but with no significant interaction between year and treatment (Table 2). There was no significant difference in the territory size of Lapland Longspur between treatment and control plots within the experimental year (t = 0.81, p = 0.42; mean relative territory size (± SD) of Lapland Longspurs was 3935 ± 3772 m² in the control plots and 5358 ± 7501 m²in the treatment plots.

Horned Lark ranged from 3.2–15.2 territories per km² among plots (Table 2). For this species, there was a significant interaction between year and treatment (Table 2), with a significant reduction in densities of territorial male Horned Larks in high-intensity treatment plots between years, after deterrents were applied (t = -2.98, p = 0.007). Relative territory density of Horned Lark males was reduced by 74% in 2019 relative to the pre-treatment year 2018 (Table 2). However, average densities were low for Horned Lark with about one territory per plot.

Nest success

During the two years of the study, we found and monitored a total of 145 nests (2018 = 72; 2019 = 73), of which 125 were from our 4 focal species. The average nest initiation date for the four focal species was 18 June in 2018 and 15 June in 2019, whereas the average date for deterrent deployment was 12 June in 2019, with full deployment completed over the period from 6 June to 17 June. At the plot level, our estimates of timing of nest initiation and deterrent deployment suggested that 79% of nests in low-intensity treatment plots and 92% of nests in high-intensity treatment plots were initiated after the deterrents were deployed in respective plots.

Analyses of daily nest survival rates included data from 77 passerine nests (Lapland Longspur, n = 64; Horned Lark, n = 13). Of these, 45 nests fledged (58% passerine apparent nest success), with 4 known abandonments (12% of failed passerine nests) and 16 known cases of predation (50% of failed passerine nests). For shorebirds, 20 out of 27 nests (Semipalmated Sandpiper, n = 23; and Least Sandpiper, n = 4) hatched (74% shorebird apparent nest success), with no known abandonments and 7 known cases of predation. In control plots, 1 of 3 shorebird nests in 2018 (33.3%) and 0 of 4 shorebird nests (0%) in 2019 failed. In comparison, in treatment plots, 2 of 7 nests in 2018 (28%) and 4 of 13 nests in 2019 (15%) failed (Fisher’s exact test: p = 0.63).

Neither treatment, nor their interaction predicted daily nest survival rates of passerines. However, year, season-date (date) and quadratic season-date (date²) were included in the top models explaining daily nest survival rates (Table 3). Only date and date² parameter estimates had confidence intervals that did not overlap 0 (Table 4). Passerine daily nest survival rates in both years had a similar pattern, with nests experiencing the lowest daily survival rates across years on day 23 (Fig. 3).

Incubation behavior

We ran 11 model variations for both response variables: proportion of time spent off the nest or the number of off-bouts per 12-hr period. Our top model for proportion of time spent off the nest per 12-hr period included time of day (Table 5), with time of day and nest age (stage in the incubation cycle) in the second-best model (ΔAIC_c*** < 2; Table 5). The top model for number of off-bouts per 12-hr period included time of day and average temperature (Table 5). Therefore, we conclude that treatment did not significantly influence either response variable based on our top models (Table 5). There was a significant effect of time of day (day vs. night) on both metrics (Fig. 4; the proportion of time spent off the nest per 12-hr period, t = -18.20, p = < 0.0000000000000002, number of off-bouts per 12 hr period, t = -9.67, p = 0.000000000000002) with birds spending more time off the nest during the day. Nest age also had a significant negative influence on the proportion of time spent off the nest per 12-hr period (t = -3.52, p = 0.00053), but not on the number off-bouts per 12-hr period (Fig. 4). Additionally, average temperature had a significant positive influence on the number of off-bouts per 12-hr period (t = 4.25, p = < 0.00004).

Costs of deterrent deployment and maintenance

Deterrents took a total of 200 person hours to deploy in the 10 6-ha treatment plots, plus an additional 120 person hours to assemble and troubleshoot prior to deployment (Table 6). For example, a crew of 6–8 people spent 4 hr deploying the flash tape grid within a single high-intensity treatment, 6-ha plot. Deterrent maintenance was done every 4 days, ranging from 30 min to 4 hr per plot, depending on damage and needs. Examples of maintenance included ensuring that the hawk-kite effigy poles were erect and that the kite was still intact, ensuring the fishing line holding together the flash tape grid was taut, and ensuring flash tape was not tangled around hummocks or vegetation. In some cases, deterrents were destroyed, taking hours to fix or were unable to be repaired. For example, in one instance the flash tape grid was dismantled likely due to disturbance by barren-ground caribou (Rangifer tarandus groenlandicus) or muskoxen (Ovibos moschatus), causing the entire grid to collapse and requiring a full, new deployment. This re-deployment took hours and demonstrated a possible risk to mid-size to large Arctic mammals that may have become entangled or injured in the flash tape grid.

Financial costs (all reported in 2019 in Canadian dollars) for audio and visual deterrents and accessories (Table 6), included the Bird-X Super Bird X-peller Pro audio units costing $510 each, with 10 purchased in 2019. This cost included audio chips for each audio unit ($60 each). To keep batteries charged so the audio deterrents would run 24 hr a day for 6 weeks, we purchased 10 solar panels, $90 each. The 12-v batteries used to run audio deterrents and hold the solar panel charge cost $55 per battery (10 batteries, $550.00). Visual deterrent costs came to a total of $5131, with Hawk-Kite effigies costing $525 for 12 Hawk-Kites, with fiber glass poles (10) totaling $517 and replacement strings (7) $42. Flash tape rolls were $5.40 per roll, totaling $1917 for 355 rolls. Fishing line used to string the flash tape grid together, cost $850 for 34 rolls of 100-lb Hercules PE braided fishing line 4 strands. Finally, the aluminum angle stakes used to erect the flash tape grid, with 640 pieces of 1-m long angles, cost $1280. The complete cost of purchasing deterrents and their accessories (without labor) was $11,731.

DISCUSSION

We found little evidence that deterrents were effective at dissuading the most abundant species of Arctic-nesting birds in our study plots. The only exception was Horned Lark for which there was a reduction in densities in the plots with deterrents between years. Whether examining factors operating during settlement and territory establishment, once nesting had been initiated (e.g., nest behavior), or reproductive success, we found few responses to our auditory and visual deterrent combinations. Lapland Longspurs that nested within the deterrent treatment plots did not exhibit a greater frequency of incubation off-bouts than birds who were not exposed to deterrents, and there were no impacts of deterrents on daily nest survival rates.

Other studies have demonstrated measurable effects of deterrents (Marcus et al. 2007, McGowan et al. 2019). However, the study species in these experiments (Common Tern, Sterna hirundo and Least Tern, Sternula antillarum) rely on active nest defense (Coulson 2002). In contrast, our study species more often rely on concealment and cryptic behavior to avoid predation, using their plumage as camouflage to avoid being detected (Caro 2005, Ruiz-Rodríguez et al. 2013). For example, Lapland Longspur sit on their nests until human observers are within 5 m before flushing (G. Holmes, personal observation). Semipalmated and Least Sandpipers both rely on vegetation over their nests to conceal their presence (Cunningham et al. 2016), although once flushed, they both sometimes exhibit conspicuous “rodent-run” behaviors to distract potential predators away from the nest site (Ashkenazie and Safriel 1979). It is possible that more aggressive species that use active forms of nest defense, such as terns and gulls, may be more responsive to predators, and therefore more likely to respond to artificially elevated predation risk mimicked by deterrents. These interspecific differences in anti-predator behaviors may explain why we saw so little response to deterrents in our study species.

Deterrent impact on relative territory densities

Arctic-nesting species have meaningful nesting-site fidelity, returning to the same territories year after year (Gratto et al. 1985, Craig et al. 2015), and sometimes even the same nest cups (Herzog et al. 2018), potentially making it difficult to deter birds from occupying a territory that is habitually theirs. In a separate study in the same study area (S. Bonnett, personal communication), we found 7 out of 54 individuals of 4 species that we banded in 2018 returned to nest in the study area in 2019 (Lapland Longspur: 1 male, Semipalmated Sandpiper: 5 unknown sex, Least Sandpiper: 1 unknown sex). In four of six of the cases for shorebird species, birds nested in the same nest cup in both years. The single resighted Lapland Longspur male had also returned to the same territory in both years. Although weather effects in 2018 versus 2019 complicate interpretation of site fidelity in our small sample of resighting data, high site fidelity could potentially limit birds’ responsiveness to deterrents. Whatever the causes, we directly observed birds’ unwillingness to abandon their territories and nest sites because of deterrents. As additional evidence for this, we observed five Lapland Longspur nests where flash tape extended directly over the nest cups (Fig. 5). Horned Larks showed a significant reduction in relative territory density between years because of treatment use. However, Horned Larks are the least common passerine with an average of less than one pair per plot. Deterrents in theory should apply to the most common species to be effective in minimizing incidental take. Despite this apparent unresponsiveness to deterrents by shorebirds and Lapland Longspurs, we acknowledge that our study on territory densities, which treated the plot as the experimental unit (n = 15), may have lacked statistical power to detect differences. Variation in territory density across plots and years could have obscured the effects of treatment if they were subtle.

Deterrent impact on daily nest survival

Deterrents did not impact daily nest survival rates or apparent survival of either passerines or shorebirds, indicating that despite the deterrents, birds that bred were still successful. As found in many other studies of factors influencing nest survival in passerines (Grant et al. 2005), date was a predictor of daily survival rate of the nests of passerines. Daily nest survival rates for passerines were highest in the early and late-breeding season, and lowest mid-season. The low daily nest survival rate for passerines mid-season aligned with egg hatching and the beginning of feeding of nestlings, a period that often coincides with the highest rates of predation (Grant et al. 2005). As in other studies of nest survival in Arctic-nesting birds (Lecomte et al. 2008, Flemming et al. 2019), we also found that predation was the major cause of nest failure.

Temporal constraints on nesting could also limit birds’ responsiveness to deterrents. Arctic-nesting birds must contend with a short breeding season by synchronizing their arrival and nest initiation with the relevant ecological processes of the Arctic if they are to breed successfully (Smith et al. 2010). Due to a short breeding season, there are few opportunities to re-nest and therefore birds must compensate by increasing investment in a single brood (Badyaev and Ghalambor 2001, Klaassen et al. 2006, Camfield 2008). Furthermore, breeding must be timed such that hatch coincides with insect emergence, to maximize chick growth (Liebezeit et al. 2014). Such temporal and ecological constraints may also explain the general lack of response from Arctic-nesting species to deterrents.

Deterrent impact on incubation behavior

Incubation behavior of female Lapland Longspur nesting in treatment plots was not different from those nesting in control plots. This indicates that audio and visual deterrents did not cause significant disturbance to incubation patterns of Lapland Longspur females. Other factors can influence length or duration of incubation recesses, but we controlled for these factors in our analysis. For example, we found that proportion of time off the nest for Lapland Longspurs decreased with incubation day, while the number of off-bouts stayed the same, in both treatment and control plots. A decrease in proportion of time off nest as incubation period advances is consistent with incubation behavior of other ground-nesting birds such as Horned Lark (Camfield and Martin 2009) and White-tailed Ptarmigan (Lagopus leucura; Wiebe and Martin 1997) and could be used as a tactic to avoid being detected by predators (Meyer et al. 2020) or to avoid chilling of the eggs (Olson et al. 2006). Time of day had the greatest influence on incubation behavior for both length and duration of incubation recesses, with fewer and short lengths of off-bouts occurring at night compared to during the day, despite the 24-h daylight in our study area. This pattern has been observed in other studies of Arctic birds, in which high activity of predators and low availability of surface-active arthropods at night make it less profitable to leave the nest at night versus during the day (Smith et al. 2012). Additionally, fewer off-bouts and more time on the nest in the colder hours of the Arctic day (i.e., night) could reduce chilling of developing embryos and is consistent with pronounced circadian cycles in the incubation scheduling of several other uniparentally incubating Arctic-breeding birds (Cartar and Montgomerie 1985, Steiger et al. 2013).

Deterrent outcomes and recommendations

We deployed deterrents as early in the season as was practical. Although our deterrents were in place before nests were established, territories may have already been established in some cases, given that we observed territorial males singing during deployment. Deploying deterrents in Arctic environments before territories are established may be more effective but is logistically challenging. This would require erecting deterrents during late winter and early spring, when the ground is still frozen and snow is present on the landscape, because most Arctic-nesting birds start to breed in available snow-free places as the land emerges (Liebezeit et al. 2014). These wintery conditions made it difficult to travel with equipment on foot, and frozen ground would make it difficult to hammer stakes into the ground and ensure that audio deterrent speakers stay upright. The use of snowmobiles could facilitate travel but would require additional costs and bring additional disturbance. Although sturdier stakes could have been employed (e.g., rebar), use of this much heavier material would have greatly increased both costs of transport and physical effort.

During the experiment, there were a few instances in which deterrents were damaged or destroyed, probably by mammals. In some plots where we found Arctic ground squirrels (Urocitellus parryii) or Arctic hare (Lepus arcticus), we anticipated there might be some damage to the wires associated with the audio deterrent units. Damage by Arctic ground squirrels occurred on one occasion when a speaker cord was chewed but quickly replaced thereafter. Of greater concern may be the impacts of deterrents on large ungulate species, such as ground-barren caribou and muskoxen, where deterrents could have caused harm through entanglement. There were multiple occasions during which visual deterrents were destroyed by caribou or muskoxen walking through the treatment plots, causing aluminum poles to be ripped out of the ground and carried away.

The cost of deterrents was on the low end of the scale compared to deterrents in other studies (Ronconi et al. 2004). Options that might have been more effective, include trained domestic dogs (Ball 2000) or radar-activated on-demand deterrence systems (Ronconi et al. 2004). However, deployment and maintenance of deterrents used here were time-consuming and required many person hours. Ultimately, the cost and labor of deterrents were not justifiable given the outcome of our experiment because three of four of our study species of Arctic-nesting birds were not successfully deterred from nesting, and the fourth (Horned Lark) only moderately.

Although incidental take is often accepted as an avoidable environmental cost for resource extraction (CWS 2014), this is nevertheless prohibited under the Migratory Birds Convention Act (MCBA 1994), and industry must make every effort to minimize, track, and report these losses. We demonstrated that traditional deterrent methods vary in their utility, depending on both species and context. Through further research efforts such as this, it is imperative that industry devises ways to reduce their impact on the natural world, including impacts on the countless populations of migratory birds already facing staggering losses across North America (Rosenberg et al. 2019).

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