Low levels of relatedness on black grouse leks despite male philopatry

In lekking species, males cluster on specific areas for display (the leks) and females generally prefer to copulate with males on large aggregations. The maintenance of leks in which only a few males reproduce might be explained if subordinate males gain indirect fitness benefits. By joining a lek on which relatives are displaying, subordinates might attract more females to the lek thereby increasing the mating opportunities of their kin. In black grouse, a genetic structure among leks has previously been found suggesting that relatives could display together. Using 11 microsatellite loci, we extended this result by testing for the presence of kin structures in nine black grouse leks (101 males). The genetic differentiation among flocks was higher in males than in females, suggesting female-biased dispersal and male philopatry. Because of this genetic structure, males were more related within than among leks. However, the mean relatedness within each lek hardly differed from zero. The lekking males were not more related than random assortments of males from the winter flocks and there were no kin clusters within leks. Thus, black grouse males do not choose to display with and close to relatives. Male philopatry alone was not sufficient to induce elevated levels of relatedness on the leks either because of male partial dispersal or a rapid turnover of the successful males. The indirect fitness benefits associated with males' settlement decision are probably limited compared to the direct benefits of joining large aggregations such as increased current and future mating opportunities.     

Genetic structure among closely spaced leks in a peripheral population of lesser prairie-chickens

We evaluated the genetic structure of birds from four closely spaced leks in a peripheral population of lesser prairie-chickens (Tympanuchus pallidicinctus). Analyses of molecular variance revealed significant genetic structuring among birds from different leks for six microsatellite loci (FST = 0.036; P = 0.002), but we found no genetic differentiation at the mtDNA control region. Significant deviations from Hardy-Weinberg revealed an excess of homozygote genotypes within each of the leks studied (FIS = 0.190-0.307), indicative of increased inbreeding. Estimates of relatedness using microsatellite data suggest that the genetic structuring among lesser prairie-chicken leks occurs in part because of a lek mating system in which males at some leks are related. Structuring may also be caused by stochastic effects associated with a historical decline in population size leading to small, semi-isolated leks and high site fidelity by reproductive males. Results from this study suggest that microspatial genetic structuring may occur in lek-mating bird species with low levels of dispersal.     

Evaluating the spatiotemporal indicators of the population decline of a threatened large forest grouse

Many wildlife populations are declining due to anthropologic and environmental pressures. Indicators can facilitate detection of and reaction to population declines. We tested spatial population pattern variables as indicators for the future fate of capercaillie populations in Slovenia. This population was surveyed from 1979 to 1983, 1998 to 2000, and 2008 to 2011. We tested the following indicators of spatial population patterns: the proportion of occupied leks in the neighbourhood of each lek, the distance to the nth nearest occupied lek, the distance to the centre of the cluster and the distance to the closest neighbouring cluster. The AUC and conditional and marginal R2 were used to evaluate the strength of the indicators. There was a steep decline in the number of leks in the Dinaric Mountains during the period 1980–2010, while the number of leks in the Eastern Slovenian Alps and Julian Alps decreased until 2000 and then stabilized. There was an increase in the distance to the nearest occupied leks over the years, but there were differences in this indicator between the clusters. A combination of the distance to the fourth nearest occupied lek and the occupancy of the first nine leks best predicted the future of a lek, showing that the indicators used in the study can be used to predict the future fate of a lek. Our results can help guide future conservation and management decisions to conserve capercaille populations.     

Distribution and size of capercaillie leks in relation to old forest fragmentation

Summary Distribution and size of 38 capercaillie Tetrao urogallus leks were related to amount and configuration of old forest patches in two south-east Norwegian coniferous forests. The smallest occupied patch was 48 ha containing a solitary displaying cock. All patches larger than 1 km² contained leks. Number of cocks per lek increased with increasing patch size. Number of leks per patch increased in a step-wise manner with one lek added for each 2.5–3 km² increase in patch size. In large patches there was one lek per 3–5 km² old forest, and density of lekking cocks was 2–2.5 per km². In small patches density of cocks varied considerably. Density of cocks was not related to patch isolation or patch shape. However, among leks surrounded by 50–60% old forest within a 1 km radius, number of cocks increased with increasing old forest fine-graininess. We argue that when old forests cover more than 50%, a fine-grained mosaic may support higher densities of lekking cocks than a coarse-grained mosaic. Conversely, when old forests cover less than 50%, a fine-grained mosaic is unfavourable, because each old forest patch becomes too small and isolated. Finally, we present a predictive model of how old forest fragmentation influences density of leks, number of cocks per lek, and total density of cocks.     

Genotyping faeces reveals facultative kin association on capercaillie’s leks

The role that kin selection might play in the evolution of lekking in birds remains controversial. Recent molecular data suggest that males displaying on leks are related. Here we investigated the genetic structure and pattern of relatedness on leks of a declining population of capercaillie (Tetrao urogallus) using microsatellite genetic markers. Since the species is highly sensitive to disturbance, we adopted a non-invasive method by using faecal samples collected in the field. Based on a dataset of 50 males distributed in 6 sub-populations, we found significant genetic structuring among sub-populations, and a significant pattern of isolation by distance among leks. Estimates of relatedness showed that males displaying on the same lek were related, even when controlling for the effects of genetical differentiation among sub-populations. In addition, the frequency distribution of relatedness values indicated that leks contain a mixture of close kin and unrelated individuals (34 and 66%, respectively). This pattern is consistent with the hypothesis that leks often contain kin associations, which might be due to very restricted dispersal of some of the males or to joint dispersal of kin. The results are discussed with respect to their implication for the conservation of endangered populations.     

Destination‐based seed dispersal homogenizes genetic structure of a tropical palm

As the dominant seed dispersal agents in many ecosystems, frugivorous animals profoundly impact gene movement and fine‐scale genetic structure of plants. Most frugivores engage in some form of destination‐based dispersal, in that they move seeds towards specific destinations, resulting in clumped distributions of seeds away from the source tree. Molecular analyses of dispersed seeds and seedlings suggest that destination‐based dispersal may often yield clusters of maternal genotypes and lead to pronounced local genetic structure. The long‐wattled umbrellabird Cephalopterus penduliger is a frugivorous bird whose lek mating system creates a species‐specific pattern of seed dispersal that can potentially be distinguished from background dispersal processes. We used this system to test how destination‐based dispersal by umbrellabirds into the lek affects gene movement and genetic structure of one of their preferred food sources Oenocarpus bataua, a canopy palm tree. Relative to background dispersal processes, umbrellabird mating behaviour yielded more diverse seed pools in leks that included on average five times more seed sources and a higher incidence of long‐distance dispersal events. This resulted in markedly lower fine‐scale spatial genetic structure among established seedlings in leks than background areas. These species‐specific impacts of destination‐based dispersal illustrate how detailed knowledge of disperser behaviour can elucidate the mechanistic link driving observed patterns of seed movement and genetic structure.     

To lek or not to lek: mating strategies of male fallow deer

We studied the mating system of fallow deer (Dama dama) for6 years in central Italy. Males in this population could defendterritories that were either single, clumped in leks, or satelliteto leks. The most highly successful males in our study werein leks. When we considered all males, there were no significantdifferences in average copulatory success according to territorytype because many lek males did not achieve any copulations,which were seen in only a few lek territories. The variancein copulatory success, however, was much greater for leks thanelsewhere. Single territories were occupied for shorter timesduring the rut than lek territories. Fighting among males wasmore frequent in the lek, even when we excluded highly successfullek males from the analysis. Chases of nonterritorial malesand harem size were correlated with the number of copulationsachieved by individual males, but did not vary according toterritory type. Copulatory success of some individuals increasedwith age, but there were no age differences among males holdingdifferent types of territories. Satellite males switched tolek territoriality in the course of one rut, but switches fromsingle territory to lek territory were rare. We suggest thatmales in single territories are inferior competitors that selecta low-risk, lowbenefit strategy, whereas those in lek territorieswhere no copulations were seen may be attempting to establishthemselves on the lek to increase their copulatory success infuture years.     

Detectability of lesser prairie-chicken leks: A comparison of surveys from aircraft

Lesser prairie-chickens (Tympanuchus pallidicinctus) are traditionally monitored by spring road-based lek surveys and counts of males attending leks. Several weaknesses exist with ground-based monitoring methods such as the bias of restricting surveys to roads, unknown probability of lek detection, and man-hours required to survey large tracts of habitat. We evaluated aerial surveys to locate lesser prairie-chicken leks in Texas and New Mexico using a Cessna 172 airplane (C172), R-22 Beta II helicopter (R-22), and R-44 Raven II helicopter (R-44) during spring 2007–2008. We determined lek activity during surveys with remote cameras placed on leks and cross-referenced time on the photo frame to time on our Global Positioning System flight log. From remote cameras we found that 305 leks were available for detection during survey flights. We determined lek detectability was 32.7% (95% CI = 20.3–47.1%) in the C172, 72.3% (64.50–79.14%) in the R-22, and 89.8% (82.0–95.0%) in the R-44. We created 16 a priori logistic regression models incorporating aircraft platform, distance to lek, survey date, lek size, and lek type to explain lek detection from aerial surveys. Our top ranked model included platform, distance, and lek type (model weight; w_i = 0.288). We had four competitive models and model averaged to draw inferences. Model averaging showed that detectability was generally greatest with the R-44, followed by the R-22, and lowest with the C172, with a slight deviation from this ranking at increased distances. Within our transect width, model averaging also suggested that detectability decreased as distance from the transect to the lek increased during helicopter surveys, and detectability increased as distance from the transect to the lek increased during C172 surveys. Furthermore, man-made leks were more likely to be detected than natural leks and large leks were more likely to be detected than medium or small leks. Aerial surveys effectively locate new leks and monitor lek density, and alleviate weaknesses associated with ground-based monitoring. We recommend using the R-44 to conduct lek surveys while flying at an altitude of 15 m at a speed of 60 km/hr on sunny mornings. © 2011 The Wildlife Society.     

Local genetic structure in a white-bearded manakin population

Local genetic structure was studied in lekking white-bearded manakins in a study area on northern Trinidad, West Indies. The study population consisted of nine leks, at which a total of 238 birds were caught. By genotyping the individuals at eight polymorphic microsatellite loci we inferred some males on leks to be related (r = 0.25) as we found an average number of 14.8 half-sib relationships and two full-sib relationships per lek. We found that the sampled birds belonged to one genetic population that was slightly inbred (FIS and FIT = 0.02). Kinship coefficients decreased with increasing geographical distance, indicating that related birds displayed at the same or nearby leks. However, leks did not consist of only one family group because the average genetic distance (aij) between males within leks was higher than when comparing males on leks within close proximity. These patterns suggest limited male dispersal, that some type of kin recognition process between individuals may exist in this species and that males on leks may be more likely to establish themselves as territory-holding birds if a relative is already present.     

Both habitat change and local lek structure influence patterns of spatial loss and recovery in a black grouse population

Land use change is a major driver of declines in wildlife populations. Where human economic or recreational interests and wildlife share landscapes this problem is exacerbated. Changes in UK black grouse Tetrao tetrix populations are thought to have been strongly influenced by upland land use change. In a long-studied population within Perthshire, lek persistence is positively correlated with lek size, and remaining leks clustered most strongly within the landscape when the population is lowest, suggesting that there may be a demographic and/or spatial context to the reaction of the population to habitat changes. Hierarchical cluster analysis of lek locations revealed that patterns of lek occupancy when the population was declining were different to those during the later recovery period. Response curves from lek-habitat models developed using MaxEnt for periods with a declining population, low population, and recovering population were consistent across years for most habitat measures. We found evidence linking lek persistence with habitat quality changes and more leks which appeared between 1994 and 2008 were in improving habitat than those which disappeared during the same period. Generalised additive models identified changes in woodland and starting lek size as being important indicators of lek survival between declining and low/recovery periods. There may also have been a role for local densities in explaining recovery since the population low point. Persistence of black grouse leks was influenced by habitat, but changes in this alone did not fully account for black grouse declines. Even when surrounded by good quality habitat, leks can be susceptible to extirpation due to isolation.     

Acoustic Divergence with Gene Flow in a Lekking Hummingbird with Complex Songs

Hummingbirds have developed a remarkable diversity of learned vocalizations, from single-note songs to phonologically and syntactically complex songs. In this study we evaluated if geographic song variation of wedge-tailed sabrewings (Campylopterus curvipennis) is correlated with genetic divergence, and examined processes that explain best the origin of intraspecific song variation. We contrasted estimates of genetic differentiation, genetic structure, and gene flow across leks from microsatellite loci of wedge-tailed sabrewings with measures for acoustic signals involved in mating derived from recordings of males singing at leks throughout eastern Mexico. We found a strong acoustic structure across leks and geography, where lek members had an exclusive assemblage of syllable types, differed in spectral and temporal measurements of song, and song sharing decreased with geographic distance. However, neutral genetic and song divergence were not correlated, and measures of genetic differentiation and migration estimates indicated gene flow across leks. The persistence of acoustic structuring in wedge-tailed sabrewings may thus best be explained by stochastic processes across leks, in which intraspecific vocal variation is maintained in the absence of genetic differentiation by postdispersal learning and social conditions, and by geographical isolation due to the accumulation of small differences, producing most dramatic changes between populations further apart.     

Do male moor frogs (Rana arvalis) lek with kin?

Many amphibian species are known to form leks during breeding season, yet it has seldom been tested which evolutionary forces are likely to act on lek formation in this taxon. We tested the kin selection hypothesis for lek formation by using eight variable microsatellite loci to compare the genetic relationship of 203 males in seven Rana arvalis leks. The results indicate that moor frog males do not lek with kin: their relatedness within leks was not higher than expected by chance. Furthermore, spatially distinct leks within same water bodies could not be distinguished from each other as separate units. These results are not expected if kin selection underlie lek formation. On the basis of these results and general knowledge of anuran breeding biology, we suggest that lek formation in explosively breeding amphibians might have evolved by female choice for breeding aggregations, combined with female choice of habitat. Future work should aim at predicting aggregations based on rules of phonotaxis over different spatial scales, and empirical work should document visitation rates not only for leks of a specific size, but also for different travel distances that visiting females may have had to cover.     

Aural and Visual Detection of Greater Sage-Grouse Leks: Implications for Population Trend Estimates

Counts of greater sage-grouse (Centrocercus urophasianus) at leks have been used in harvest management, Endangered Species Act listing decisions, and land management policies for over half a century. Lek count sampling methods focus on counting male sage-grouse at known leks, primarily those observed visually from roads or vantage points, but leks are likely missed that are unknown prior to the survey and are difficult to detect while driving between known lek sites. One way to ameliorate this shortfall may be to conduct short point-count surveys at multiple stops along lek-survey routes or while driving between lek counts, thereby detecting newly established or unknown leks. To evaluate the feasibility of this approach, we estimated aural and visual detection probability of active sage-grouse leks during 1-minute point-count surveys at known distances and examined the effects of environmental factors on aural lek detection in southern Idaho, USA, 2016–2017. Our results demonstrate that field observers can aurally detect sage-grouse leks at approximately 3 times greater distances compared to detecting leks visually. The probability of hearing an active lek was highest near the peak of male and female attendance (8 Apr), within an hour of sunrise, on relatively calm and cold days, when the observer was at a higher elevation relative to the lek, and during conditions with no background noise. Detection probability declined with distance and the probability of aural detection was 0.59 at 1 km from a lek when other variables were held at their means. Hence, conducting ≥3 1-minute surveys along a lek route would be expected to detect ≥93% of all leks within 1.5 km of each survey under the average environmental conditions in our study. Our results suggest that surveys could greatly improve detection of unknown or newly established leks and can facilitate a more accurate assessment of sage-grouse population trends through lek counts. Moreover, our results demonstrate how environmental factors influence the detection of leks during surveys, and therefore which variables should be considered for inclusion in any future revisions of lek count protocols or in analyses of lek count data. © 2021 The Wildlife Society.     

Female mate choice across spatial scales: influence of lek and male attributes on mating success of blue-crowned manakins

Lekking males compete for females within and among leks, yet female choice is expected to work differently at each of these spatial scales. We used paternity analyses to examine how lek versus male attributes influence mate choice in the blue-crowned manakin Lepidothrix coronata. We tested the hypotheses that females prefer (i) to mate at larger leks where a larger number of potential mates can be assessed, (ii) to mate with unrelated or highly heterozygous males expected to produce high-quality offspring, (iii) to mate with males that display at higher rates, and that (iv) display honestly reflects male genetic quality. Our results show that (i) males at larger leks are not more likely to sire young, although females nesting close to small leks travel further to reach larger leks, (ii) siring males are not less related to females or more heterozygous than expected, (iii) within a lek, high-display males are more likely to sire young, and (iv) both male heterozygosity and display rate increased with lek size, and as a result display does not reliably reflect male genetic quality across leks. We suggest that female mate choice in this species is probably driven by a Fisherian process rather than adaptive genetic benefits.     

The anatomy of population change in a black grouse population 1992–2008

We interrogate an 18-year-long dataset containing counts of displaying male black grouse Tetrao tetrix and incidental counts of females within an 800-km² region of Perthshire, Scotland. We examine the trends in the population and investigate how different components of the population might act as signposts of different stages of overall population change. We found statistical evidence for a decline in black grouse numbers between 1992 and 2000, and then a recovery from 2002 to 2008, but little evidence for a link between population change and weather during the decline phase. There was some evidence for a positive relationship between male and female counts. The two main components of male population size, lek size and lek frequency followed the overall population trend while it was increasing, but during the earlier decline, the two became uncoupled, to expose a complex structure within the data. During the decline, when black grouse numbers were approaching their minimum, mean lek size was actually increasing. Small leks lost proportionally more birds than did large leks, and lek longevity was positively correlated with lek size, indicating that maintenance of large leks is crucial in buffering the population against serious declines. During the decline, the spatial arrangement of leks changed, with remnant leks showing tight clustering at larger spatial scales, before expanding out to fill the large areas of unoccupied landscape during the population increase. We discuss these findings in terms of species monitoring and suggest that counts of young males may add much useful demographic information with little extra effort.     

Distribution dynamics of a great bustard metapopulation throughout a decade: influence of conspecific attraction and recruitment

Dispersing individuals can use conspecifics as indicators of habitat quality and aggregate at traditionally occupied sites, leaving other favourable patches unoccupied. Here we test the predictions of the conspecific-based habitat selection hypothesis on a Spanish great bustard (Otis tarda) metapopulation, currently fragmented due to recent human-induced habitat changes. The number of birds had increased by 23% between 1988 and 1998, but not consistently among leks. Leks that were large in 1988 increased, while those that were small decreased, which suggests that dispersing individuals used the numbers of conspecifics as cues for breeding-site selection. Moreover, leks with high productivity increased, while those with low productivity decreased. Finally, lek distribution was markedly stable throughout the decade, with no establishment of new leks, and suitable habitat patches remained unoccupied, as predicted by the conspecific attraction hypothesis. These results were corroborated by a simulation model which incorporated natal dispersal rates between leks as obtained through radio-tracking of 15 birds that survived throughout their 4-year dispersal period. In conclusion, in spite of the apparent increase in total numbers throughout the decade, both conspecific attraction and local differences in reproductive success contributed to a more aggregated distribution, increasing the species' vulnerability to local catastrophes, and the risks of reduced genetic diversity and extinction of small leks.     

Lek fidelity and movement among leks by male Greater Sage‐grouse Centrocercus urophasianus: a capture‐mark‐recapture approach

Males in lek mating systems tend to exhibit high fidelity to breeding leks despite substantial evidence of skewed mating success among males. Although movements between leks are often reported to be rare, such movements provide a mechanism for an individual to improve lifetime fitness in response to heterogeneity in reproductive conditions. Additionally, estimates of apparent movements among leks are potentially biased due to unaccounted variation in detection probability across time and space. We monitored breeding male Greater Sage‐grouse Centrocercus urophasianus on 13 leks in eastern Nevada over a 10‐year period, and estimated movement rates among leks using capture‐mark‐recapture methods. We expected that male movement rates among leks would be low, despite predictions of low breeding success for most males, and that detection rates would be highly variable among leks and years. We used a robust design multistate analysis in Program mark to estimate probability of movements among leks, while accounting for imperfect detection of males. Male Sage‐grouse were extremely faithful to their leks; the annual probability of a male moving away from its original lek of capture was approximately 3% (se = 0.01). Detection probabilities varied substantially among leks (range = 0.21–0.95), and among years (range = 0.30–0.76), but remained relatively constant within years at each lek. These results suggest that male Sage‐grouse dispersal is either rare, or consists primarily of dispersal of sub‐adults from their natal areas prior to the breeding season. The study highlights the benefits of robust design multistate models over standard ‘live‐encounter’ analyses, as they not only permit estimation of additional parameters, such as movement rates, but also allow for more precise parameter estimates that are less sensitive to heterogeneity in detection rates. Additionally, as these data were collected using capture‐mark‐recapture methods, our approach to estimating movement rates would be beneficial in systems where radiotagging is detrimental to the study organism.     

Thresholds and Time Lags in Effects of Energy Development on Greater Sage-Grouse Populations

ABSTRACT Rapid expansion of energy development in some portions of the Intermountain West, USA, has prompted concern regarding impacts to declining greater sage-grouse (Centrocercus urophasianus) populations. We used retrospective analyses of public data to explicitly investigate potential thresholds in the relationship between lek attendance by male greater sage-grouse, the presence of oil or gas wells near leks (surface occupancy), and landscape-level density of well pads. We used generalized linear models and generalized estimating equations to analyze data on peak male attendance at 704 leks over 12 years in Wyoming, USA. Within this framework we also tested for time-lag effects between development activity and changes in lek attendance. Surface occupancy of oil or gas wells adjacent to leks was negatively associated with male lek attendance in 5 of 7 study areas. For example, leks that had ≥ 1 oil or gas well within a 0.4-km (0.25-mile) radius encircling the lek had 35–91% fewer attending males than leks with no well within this radius. In 2 of these 5 study areas, negative effects of well surface occupancy were present out to 4.8 km, the largest radius we investigated. Declining lek attendance was also associated with a higher landscape-level density of well pads; lek attendance at well-pad densities of 1.54 well pads/km² (4 well pads/mile²) ranged from 13% to 74% lower than attendance at unimpacted leks (leks with zero well pads within 8.5 km). Lek attendance at a well-pad density of 3.09 well pads/km² (8 well pads/mile²) ranged from 77% to 79% lower than attendance at leks with no well pad within 8.5 km. Further, our analysis of time-lag effects suggested that there is a delay of 2–10 years between activity associated with energy development and its measurable effects on lek attendance. These results offer new information for consideration by land managers on spatial and temporal associations between human activity and lek attendance in sage-grouse, and suggest that regional variation is an important consideration in refining existing management strategies.     

Mating Behavior Drives Seed Dispersal by the Long‐wattled Umbrellabird Cephalopterus penduliger

Frugivores exhibit considerable variation in the seed dispersal services they provide. Understanding what drives these differences is a key goal for ecologists because of the central role seed dispersal plays in shaping ecological and genetic diversity in plant populations. The lek‐mating system of the Long‐wattled Umbrellabird (Cephalopterus penduliger) provides a powerful lens to examine how mating behavior may impact seed dispersal outcomes. As in all lek‐breeding species, male Umbrellabirds congregate in traditional sites (leks) to display, whereas females are solitary and visit leks only rarely. This study demonstrates how differences in mating behavior between the sexes drive distinctive seed movement and deposition patterns by male vs. female Umbrellabirds. Using radio tracking and gut retention trials, we documented divergent movement patterns between the sexes that are directly attributable to mating behavior differences. These movement differences led males to disperse seeds long distances from source trees and to deposit the majority of seeds they ingested within the lek; females dispersed seeds shorter distances and more evenly across the landscape. We empirically confirmed that the density of dispersed seeds was higher in leks than in control areas outside the lek, yet found no evidence that this higher density of seeds in leks reduced probability of seedling establishment. This research not only provides a mechanistic explanation for long dispersal distances and high levels of genetic diversity previously reported for seeds in Umbrellabird leks, but also highlights the importance of explicitly considering behavior in studies of animal‐mediated seed dispersal.     

Responses of Male and Female Black Grouse to Male Vocal Display

The evolution of leks may be explained by several hypotheses. The ‘female preference’ hypothesis, which states that females favour males that have aggregated, has recently gained some empirical support. Low-quality, unattractive males may, however, settle near attractive males, as predicted by the ‘hotshot’ hypothesis. We tested whether black grouse Tetrao tetrix females use auditory cues to find the preferred leks, and whether males respond to vocal display emitted on leks. We conducted a playback experiment with male vocal display (rookooing) on leks, where the visiting females and displaying males were counted. The number of males tended to increase more on playback leks. Specifically, the number of 1-year-old males was greater on playback sites than on control sites. Also, the number of females, in relation to the lek size before the start of the experiment tended to increase. In addition, we used aviary playback trials to test whether females distinguish between single-male and multi-male displays. In a choice test, females showed greater preference for the ‘multi-male’ tape. The tendency for increased male numbers on playback leks resulted from increased visits of young, mobile males which were attracted to leks that they perceived to be large. This suggests a ‘hotshot’-type mechanism in the settlement of young males. Because females also responded to the supplemented auditory advertisement, or directly to the increased number of males, the ‘female preference’ hypothesis is also supported. Females may, at least in part, base their decision of which lek to visit on auditory cues, but visual contact to males may be also needed.