Males of many polygynous species compete for access to fertile females without providing them with resources other than sperm and without investing in care for the offspring (male dominance polygyny)1,2,3,4. In such systems, local competition for access to females is intense and typically only a few males obtain matings, leading to strong sexual selection2,4,5. Sampling multiple breeding areas could then provide a mechanism for males to increase their chances to reproduce. However, little is known about such sampling behaviour and about the spatial scale at which males compete6,7,8. Here we show that most males of a migratory, polygynous shorebird, the pectoral sandpiper (Calidris melanotos)9,10, that arrived at a known breeding location in northern Alaska9 subsequently moved through a considerable part of the entire species’ breeding range (up to 13,045 km in a four-week period), sampling as many as 23 additional potential breeding sites. Our data suggest that males do not have a final breeding destination after migration from their wintering quarters, but make nomadic movements that are probably not a consequence of breeding failure. Tenure, the duration of stay at a site, correlated strongly with the number of breeding females at the site, suggesting that decisions to leave are dependent on local mating opportunities. Nomadic movements may allow males to display and sire offspring at multiple sites within a single breeding season. Sexual selection may then favour high-performance males that are able to reduce sleep to compete locally9 and to fly long distances between breeding sites, leading to a population with unrestricted interbreeding and without local adaptation and speciation.
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We thank R. Barth, J. Conklin, W. Forstmeier, A. Jacot, K. Kapetanopoulos, S. Kuhn, L. Langlois, P. Loës, C. Muck, A. Mutzel, M. Oltrogge, E. Penning, H. Schielzeth, X. Schleuning, S. Steiger, K. Teltscher, K. Temnow, A. Türk, D. Werner, A. Wittenzellner and L. Zimmer for help in the field, S. Kuhn and K. Teltscher for genotyping, and W. Goymann, I. Schwabl and M. Trappschuh for hormone analyses. We thank P. Mombaerts for photos and for encouraging us to conduct the tracking study on a large scale, J. Conklin for discussion, W. Forstmeier for comments on the manuscript, and E. Schlicht for discussion, literature search, and comments on the manuscript. We thank R. Lanctot from USFWS in Anchorage and the Barrow Arctic Science Consortium and UMIAQ Barrow Science Support for logistical support. This work was funded by the Max Planck Society.
The authors declare no competing financial interests.
Reviewer Information: Nature thanks J. Gill, L. Oring and the other anonymous reviewer(s) for their contribution to the peer review of this work.
Extended data figures and tables
Extended Data Figure 1 Spatio-temporal illustration of the site sampling behaviour of male pectoral sandpipers during two breeding seasons.
a, The cumulative number of potential breeding sites visited by each male. b, The cumulative distance travelled by each male within the Arctic breeding region. Horizontal grey bar, range of clutch initiation dates (n = 746) from six sites across Alaska (this study, refs 41,42). Lines connect successive points (no step function).
Extended Data Figure 2 Increase in expected male reproductive success with the number of females breeding in Barrow in each year.
The expected breeding success was estimated as best linear unbiased predictors (BLUPs) from a generalized linear mixed-effects model with Poisson error distribution where the number of young sired was fitted as dependent variable, the number of breeding females as predictor, tenure (ln transformed) as covariate, and year and male identity as random intercepts (see Extended Data Table 5). Tenure was fitted as a random slope for each year. The correlation between the expected breeding success and the number of breeding females was r = 0.95, n = 5, P = 0.016.
Extended Data Figure 3 Spatio-temporal patterns of snow cover within the entire known breeding area of the pectoral sandpiper during the 2012 and 2014 breeding seasons.
a, Seasonal changes in snow cover (percentage of the breeding range covered with snow). b, Spatial autocorrelation of snow cover in each year. Each line represents a single day during the breeding season (day 1 = 1 June) and shows how spatial autocorrelation (as measured by Moran’s I) decreases with distance (data for 10 km intervals). The graphs illustrate that similarity in snow cover between two locations exists only on a small scale and quickly declines with increasing distance, even early in the season.
Extended Data Figure 4 The opportunity for sexual selection in male pectoral sandpipers under two hypothetical mating scenarios.
In Barrow, mating success, that is, the number of females with whom a male sired at least one offspring was determined by how long the male was present on the study site (Fig. 3d and Extended Data Table 5). Assuming that a similar relationship between tenure and mating success exists elsewhere, mating success of each male at each of the sites he visited in suitable breeding habitat in the Arctic can be predicted from his tenure at each of these sites. We calculated male mating success under two scenarios: (1) assuming that males could sire offspring at all visited sites, (2) assuming that males only sired offspring at the site where they had their longest tenure. The predicted mating success of each male was then used to compute Is (the opportunity for sexual selection43) as the variance in mating success divided by the squared mean mating success. Only males recorded for at least 30 days (n = 44) were included, and data from Barrow (the catching site) were excluded. Shown are Is with non-parametric bootstrapped 95% confidence intervals. The value computed for scenario 1 (Is = 0.376) is almost 20 times higher than the value computed for scenario 2 (Is = 0.019), suggesting that the effect of male breeding site sampling on sexual selection strongly depends on whether males actually compete for matings at each of the areas they visited. This is because there is substantial variation among males in both the number of sites they visited and in their tenure at these sites. Assuming that mating success depends on tenure as observed in Barrow, then, under scenario 1, the most successful males are those that visit the most sites, not those that stay longer at fewer sites.
a, 5 g solar PTT-100 transmitter glued directly on the back of a male pectoral sandpiper (tail is on the left). b, Flying male with transmitter in Barrow, Alaska. Note the colour bands (red–white–red) used for identification during field observations. c, Displaying male with transmitter and colour bands (red–yellow–red) on the tundra in Barrow. Credits: a, P. Mombaerts; b, D. Werner, Max Planck Institute for Ornithology; c, K. Kapetanopoulos, Max Planck Institute for Ornithology.
This file contains maps of tracks flown and areas visited for each individual male pectoral sandpiper captured in Barrow, Alaska, at the beginning of the breeding season in 2014 (N = 60) and 2012 (N = 60). Staging areas are shown as minimum convex polygons scaled with a factor of 15% (referenced to the map scale: lower right corner) to increase visibility. Only staging areas within the suitable breeding habitat (blue area) are shown. Darker blue: known breeding range of the species. Black dots: raw ARGOS locations; red lines: predicted track of the individual. Below each map the following information is provided: ID, a combination of individual identity and year; Overall tenure, the sum of the tenures at each residency area; Total time recorded, number of days between the first recorded signal in the first residency area (Barrow) and the last recorded signal in the last visited area (excluding the last area when tenure cannot be estimated due to the loss of the satellite tag); Total distance, sum of the distances flown between each visited area. For individuals that were followed until they left the breeding area, the total distance covered is marked as complete. (ZIP 4458 kb)
Birds were caught in Barrow, Alaska in 2012 (N = 60, red) and in 2014 (N = 60, blue). The current residency areas are shown as coloured rhombi; they disappear when the bird moves on or when the satellite transmitter stops sending data. Pale green area: suitable breeding habitat of the pectoral sandpiper; dark green area: known breeding range of the species (see Extended Data Fig. 1 for references). Map projection: Polar Lambert azimuthal equal-area with longitude origin 156.65° W (Barrow). Note the running date and time, and the scale bar on the bottom right. In 2014, blizzard conditions (frozen tundra covered with fresh snow) in late May caused most males to leave Barrow and fly to the SW; only one male later returned to the study site for some time. (MOV 11582 kb)
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Kempenaers, B., Valcu, M. Breeding site sampling across the Arctic by individual males of a polygynous shorebird. Nature 541, 528–531 (2017). https://doi.org/10.1038/nature20813
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