In our study of ruff, a lekking shorebird, we found that the “lek ratio” — the proportion of the total population of males occurring on leks was low, averaging 12% over the breeding season (Fig. 1D). Off-lek males spent approximately the same proportion of time as lek males in displaying to females (Fig. 4). We defined three spacing tacties that male ruffs use to position themselves to court females: “Following” — directly pursuing females, and two types of lekking behavior, “Intercepting” — waiting for females at resource-rich sites, and “True Lekking” — waiting for females at places without any resources other than the males themselves. Males switched among these tactics, causing the lek ratio to vary over the season (Fig. 1 D). Lek ratio increased when the number of females present in the study area plummeted at the end of May, and was positively correlated throughout the season with the copulation rate of the preceding day, suggesting that males were tracking the behavior, as well as the number, of females available (Table 1, Fig. 1). Early in the season, males off leks spent most of their time feeding (Fig. 4), and lek ratio was positively correlated with temperature (Table 1), suggesting that some males may have been unable to lek during cold weather. Males on leks mated at significantly higher rates than Followers (Table 4). On average, males at our interception lek were less site faithful and less peristent than males at true leks, and the interception lek itself disappeared after females stopped coming to use its adjacent resource (Table 2, Fig. 5). The most successful individuals in our population were the True Lekking males, rather than the Interceptors.
In addition to the conditinal lekking tactics described here, ruff display a dimorphism in behaviorat leks. “Independent” males defend small courts on leks, while “Satellite” males share courts and mutually display with independents Both independents and satellites may use all three conditional tactics. We propose that satellites evolved as specialized. Followers, adept at tracking the movements of females among leks.