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.     

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.     

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.     

The importance of within-year repeated counts and the influence of scale on long-term monitoring of sage-grouse

Long-term population monitoring is the cornerstone of animal conservation and management. The accuracy and precision of models developed using monitoring data can be influenced by the protocols guiding data collection. The greater sage-grouse (Centrocercus urophasianus) is a species of concern that has been monitored over decades, primarily, by counting the number of males that attend lek (breeding) sites. These lek count data have been used to assess long-term population trends and for multiple mechanistic studies. However, some studies have questioned the efficacy of lek counts to accurately identify population trends. In response, monitoring protocols were changed to have a goal of counting lek sites multiple times within a season. We assessed the influence of this change in monitoring protocols on model accuracy and precision applying generalized additive models to describe trends over time. We found that at large spatial scales including >50 leks, the absence of repeated counts within a year did not significantly alter population trend estimates or interpretation. Increasing sample size decreased the model confidence intervals. We developed a population trend model for Wyoming greater sage-grouse from 1965 to 2008, identifying significant changes in the population indices and capturing the cyclic nature of this species. Most sage-grouse declines in Wyoming occurred between 1965 and the 1990s and lek count numbers generally increased from the mid-1990s to 2008. Our results validate the combination of monitoring data collected under different protocols in past and future studies—provided those studies are addressing large-scale questions. We suggest that a larger sample of individual leks is preferable to multiple counts of a smaller sample of leks. © 2011 The Wildlife Society.     

Assessing lek attendance of male greater sage-grouse using fine-resolution GPS data: Implications for population monitoring of lek mating grouse

Counts of males displaying on breeding grounds are the primary management tool used to assess population trends in lekking grouse species. Despite the importance of male lek attendance (i.e., proportion of males on leks available for detection) influencing lek counts, patterns of within season and between season variability in attendance rates are not well understood. We used high-frequency global positioning system (GPS) telemetry data from male greater sage-grouse (Centrocercus urophasianus; n = 67) over five lekking seasons (2013–2017) at eight study sites in Nevada to estimate lek attendance rates. Specifically, we recorded daily locations of sage-grouse in relation to mapped lek boundaries and used generalized additive models to assess temporal variation in attendance rates by age class (subadult vs. adult). Average timing of peak attendance occurred on 16 April but varied from March 16, 2014 to April 21 , 2016. Overall, adult males attended leks at higher rates (0.683 at peak) and earlier in the season (19 March) than subadults (0.421 at peak on April 19). Peak attendance probability was positively related to cumulative winter precipitation. Daily probabilities of lek switching differed between adults (0.019 at peak on March 3) and subadults (0.046 at peak on March 22), and lek switching was negatively related to distance to nearest lek. Our results indicate variable patterns in lek attendance through time, and that lek switching may occur at higher rates than previously thought. We demonstrate the use of generalizable daily attendance curves to date-correct lek counts and derive estimates of male abundance, although such an approach will likely require the incorporation of information on age structure to produce robust results that are useful for population monitoring.     

Greater Sage-Grouse Population Response to Energy Development and Habitat Loss

ABSTRACT Modification of landscapes due to energy development may alter both habitat use and vital rates of sensitive wildlife species. Greater sage-grouse (Centrocercus urophasianus) in the Powder River Basin (PRB) of Wyoming and Montana, USA, have experienced rapid, widespread changes to their habitat due to recent coal-bed natural gas (CBNG) development. We analyzed lek-count, habitat, and infrastructure data to assess how CBNG development and other landscape features influenced trends in the numbers of male sage-grouse observed and persistence of leks in the PRB. From 2001 to 2005, the number of males observed on leks in CBNG fields declined more rapidly than leks outside of CBNG. Of leks active in 1997 or later, only 38% of 26 leks in CBNG fields remained active by 2004–2005, compared to 84% of 250 leks outside CBNG fields. By 2005, leks in CBNG fields had 46% fewer males per active lek than leks outside of CBNG. Persistence of 110 leks was positively influenced by the proportion of sagebrush habitat within 6.4 km of the lek. After controlling for habitat, we found support for negative effects of CBNG development within 0.8 km and 3.2 km of the lek and for a time lag between CBNG development and lek disappearance. Current lease stipulations that prohibit development within 0.4 km of sage-grouse leks on federal lands are inadequate to ensure lek persistence and may result in impacts to breeding populations over larger areas. Seasonal restrictions on drilling and construction do not address impacts caused by loss of sagebrush and incursion of infrastructure that can affect populations over long periods of time. Regulatory agencies may need to increase spatial restrictions on development, industry may need to rapidly implement more effective mitigation measures, or both, to reduce impacts of CBNG development on sage-grouse populations in the PRB.     

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.     

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.     

Lek formation by female choice: a simulation study

We used computer simulation to study the formation of matingaggregations under the influence of female choice. Female movementswere not constrained by home range size. Males formed clumpeddispersions if females expressed strong preferences either forlarger leks or for male quality. High travel costs for femalesvisiting leks produced male dispersions that matched the initialdistribution of females. High costs to females for waiting formatings produced evenly dispersed males. Depending on the relativeweights of these factors, a continuous range of outcomes froma single lek per population to evenly dispersed males was produced.The results suggest that effects of female choice on lek evolutionare more likely to be detected by studying the processes bywhich leks form than by characterizing the spatial distributionof displaying males.     

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.     

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.     

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.     

Disturbance factors influencing greater sage-grouse lek abandonment in north-central Wyoming

Detecting the disappearance of active leks is the most efficient way to determine large declines in greater sage-grouse (Centrocercus urophasianus) populations; thus, understanding factors that influence lek abandonment is critical. We evaluated factors that may have influenced the probability of sage-grouse lek abandonment in the Bighorn Basin (BHB) of north-central Wyoming from 1980 to 2009. Our objective was to examine lek abandonment based on landscape characteristics that explain differences between occupied and unoccupied leks. We evaluated lek abandonment from 144 occupied and 39 unoccupied leks from the Wyoming Game and Fish Department lek database with sufficient data for our 30-year analysis. We conducted our analysis with binary logistic regression using landscape predictor variables obtained from geographic coverages at 5 scales (1.0-, 3.2-, 4.0-, 5.0-, and 6.4-km radii around leks) to evaluate how these disturbances have influenced lek abandonment. Coverages included anthropogenic characteristics such as agricultural development, oil and gas development, prescribed burned treatments, and roads; and environmental characteristics such as vegetation attributes and wildfire. Our combined model included the number of oil and gas wells in a 1.0-km radius, percent area of wildfire in a 1.0-km radius, and variability in shrub height in a 1.0-km radius around sage-grouse leks. Abandoned (unoccupied) leks had 1.1-times the variability of shrub height in a 1.0-km radius, 3.1-times the percentage of wildfire in a 1.0-km radius, and 10.3-times the number of oil and gas wells in a 1.0-km radius compared to occupied leks. The model-averaged odds of lek persistence with every 1 unit increase in oil and gas wells within a 1.0-km radius was 0.66 (90% CI: 0.37–0.94), odds with every 1% increase in wildfire in a 1.0-km radius was 0.99 (90% CI: 0.85–1.12), and odds with every 1 unit increase in the standard deviation of shrub height within a 1.0-km radius around a lek was 0.77 (90% CI: 0.45–1.08). Because the 90% confidence intervals around the odds ratios of wells did not overlap 1.0, we suggest this predictor variable was most influential in our model-averaged estimates. The BHB has lower developed reserves of oil and gas than many other regions; however, our study supports findings from other studies that demonstrate energy development increases lek abandonment. Our findings indicate conservation efforts should be focused on minimizing well development and implementing wildfire suppression tactics near active sage-grouse leks. © 2012 The Wildlife Society.     

Kin groups in closely spaced capercaillie leks

We quantified the genetic structure among 58 capercaillie (Tetrao urogallus) from three closely spaced leks in boreal Russia using ten microsatellite loci. We did not find significant structuring with respect to lek membership in an overall analysis. However, all but one individual could be strongly assigned to one of two genetic clusters, which we interpret as reflecting different local lineages or extended family groups. There was evidence for structuring among some clusters within leks, and one smaller lek consisted of nine out of ten males from one cluster, which could reflect kin biasing of lek membership. One cluster was apparently outperforming the other, since nine out of ten juveniles sampled belonged to it.     

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.     

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.     

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.     

Lekking, resource defense, and harassment in two subspecies of lechwe antelope

For lek-breeding in ungulate populations to continue, benefitsto males defending lek territories and to females visiting leksmust outweigh the costs. In this study, Kafue lechwe males onleks gained higher mating rates than nonlekking males, a resultof sexually receptive females leaving herds and aggregatingon leks. When the numbers of females on leks were experimentallyreduced, benefits to males decreased, resulting in males graduallyabandoning lek territories. A comparison with a population ofnonlekking, resource-defending black lechwe showed that matingattempts by estrous females in herds were disrupted by harassingmales eight times more frequently in a population of lek-breedingKafue lechwe than in the nonlekking black lechwe. Despite thefact that there were fewer Kafue lechwe females on single territories,harassment of estrous females by males was greater on singleterritories of Kafue lechwe than on leks and greater than onblack lechwe resource territories. Females were also absenton Kafue lechwe single territories for long periods becauseof erratic, widespread movements of compact herds resultingfrom unpredictable distributions of resources. In contrast,black lechwe females were more evenly dispersed over homogeneousresources and for a given territory, females were likely tobe present most of the time. Therefore, unlike black lechwe,male Kafue lechwe find it uneconomical to defend resource territories.Thus, costs to estrous females mating off leks and the absenceof benefits to males attempting to defend resource-based territoriesmay be important cofactors in the appearance of lek-breedingin some ungulate populations     

Evolution of black grouse leks: female preferences benefit males in larger leks

Leks, communal display grounds in which males display and femalesonly attend to mate, represent one of the most remarkable outcomesof sexual selection. There have been no detailed studies thatcompare the behavior of males and females between leks of differentsizes to test if there is any benefit for male clumping andto test the many hypotheses suggested to explain lekking. Inthe black grouse, Tetrao tetrix, larger leks have many morefemale visits and copulations, leading to a higher average malemating success. Females visiting larger leks are also more likelyto mate, indicating that female preferences of males are importantfor the evolution of leks. Yearling males seldom copulate, buttheir presence on the lek increases the mating success of adultmales, suggesting that lek size per se and not only male qualityaffects female preferences for larger leks. The distributionof males over lek sizes agrees with a game theory model of idealfree distribution with unequal competitors, in which less competitivemales go to smaller leks.     

Lek mating systems: a case study in the Little Bustard Tetrax tetrax

Leks have recently been defined as male display aggregations that females attend primarily for the purpose of mating. This is an extended version of previous definitions, as a clear-cut definition of leks is difficult to obtain. Four criteria should be verified to identify a lekking species: (i) there is no male parental investment beyond the sperm; (ii) males aggregate at specific sites for display; (iii) the only resource females find on the lek is the male, i.e. the male genes; (iv) females can select her mate(s), although the necessity of this latter condition for lekking species has been highly debated. We applied these criteria to the endangered little bustard Tetrax tetrax, a species that is claimed to show an exploded lek mating system, but for which this has never been fully investigated. We monitored a population of little bustards in western France during 2 years to investigate the two central criteria in the assessment of their mating system: male aggregation in arenas and lack of consistent resources in male territories. We analysed the spatial distribution of little bustard male territories, the individual variation in size, and the land use characteristics of male territories, with particular attention to the habitats that may be considered as defensible resources. Displaying males showed an aggregated spatial distribution over the study area during the 2 years of survey. Male territories were rather large (19+/-16 ha), but a large among-male variability in territory size was observed. Land use within the territories included mainly permanent and semi-permanent crops. The variability in land use among territories suggests also that resources found within male territories were selected according to male needs (food and display) rather than to female needs (permanent crops that are more appropriate for reproduction). The mating system of the little bustard seems to match the general (and extended) definition of leks, at least in some populations. However, limits between resource defence polygyny and extreme exploded or resource-based leks are thin and unclear, and the little bustard is a good example of how lek definitions may be difficult to apply in non clear-cut empirical situations.