Restrictive mating by females on black grouse leks

In bird species with pair bonds, extra-pair matings could allow females to choose genetically superior males. This is not needed in lekking species because female choice is not constrained by pairing opportunities. However, polyandry has been reported in most lekking species studied so far. Using 12 microsatellite loci, we determined the paternity of 135 broods of black grouse sampled between 2001 and 2005 (970 hatchlings and 811 adult birds genotyped). The paternity assignments were combined to lek observations to investigate the mating behaviour of black grouse females. About 10% of the matings seemed to take place with males displaying solitarily. Forty per cent of the copulations between males displaying on the studied leks and radio-tagged females were not recorded. This was due to difficulties in identifying the females and because our observations did not cover all the possible time for matings. However, females of the undetected copulations had chosen males that were already known to be successful on the leks. There was a strong consistency between the observations and true paternity, even when the copulation was disturbed by a neighbouring male. Multiple mating and multiple paternities were rare. We can now confidently ascertain that most females mate only once with one male for the whole clutch. This mating behaviour requires that a single insemination is sufficient to fertilize a clutch and that females can determine whether the sperm has been successfully transferred. Grouse Tetraoninae with many lekking species may be the only bird taxon that has evolved these traits.     

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.     

Testing predictions for the evolution of lekking in the sandfly, Lutzomyia longipalpis

We carried out a combination of field and laboratory experiments to investigate two possible mechanisms promoting the evolution of lekking in the sandfly, Lutzomyia longipalpis. Males in this species lek on or near hosts that females visit to mate and feed. To test whether the distribution of resources, lek size and female number were positively correlated (a prediction of hotspot models) we manipulated the availability of potential lek sites by varying the numbers of chickens, Gallus gallus domesticus, held in field cages. We then investigated the distribution of males and females across leks formed within these cages and measured males to assess the relationship between male size and lek size. The number of males attracted to a lek increased with the number of hosts present and on average males on larger leks had longer abdomens. The number of females was positively correlated with the number of males present, but the average number of females per male decreased with increasing lek size. In a second experiment, we introduced single females to laboratory leks of different sizes, and compared female latency to mate. Contrary to an explicit prediction of the black hole model, lek size had a direct effect on a female's latency to mate. Females at larger leks encountered and copulated with males sooner than those at smaller aggregations, but the length of time a pair spent courting did not vary with lek size. Our results suggest that the black hole model is unlikely to explain the evolution of lekking in this species. The data instead support the hotspot model as a possible mechanism promoting lekking in L. longipalpis but suggest that female preference for larger leks may need to be invoked to obtain the degree of aggregation observed.     

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.     

Active formation of mixed-species grouse leks: a role for predation in lek evolution?

Behavioural ecologists have interpreted avian leks as products of sexual selection, in which males display socially to increase their opportunities to mate. However, without invoking reproductive queuing or kin selection, this paradigm does not necessarily explain why many males that fail to mate participate in leks. An alternative solution, that males also aggregate to reduce predation, has previously lacked compelling support. We show that mixed-species leks, comprising two congeneric grouse, form when single males or small groups of one species, the greater prairie chicken Tympanuchus cupido, join leks of another, the sharp-tailed grouse T. phasianellus. We documented the process by observing lek dynamics and comparing group sizes between mixed- and single-species leks. Joining implies that prairie chickens benefit from displaying with sharp-tailed grouse. The numbers of females of each species attending a lek increased with the number of conspecific, but not heterospecific, males. This suggests that the joining of heterospecifics is unlikely to increase mating opportunities, and leaves lowered predation risk as the most likely benefit of associating with heterospecifics. Active formation of mixed-species leks therefore suggests that predation may be sufficient to drive lek formation. The benefits of participation in mixed leks may be asymmetrical because prairie chickens display more and are less vigilant than sharp-tailed grouse.     

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.     

Long-range visibility of greater sage grouse leks: a GIS-based analysis

We investigated whether male greater sage grouse, Centrocercus urophasianus, select lek locations on the basis of topographic features that affect their visibility to both conspecific females and a major avian predator, the golden eagle, Aquila chrysaetos. We mapped locations of displaying males at all leks in a local population and used a Geographic Information System (GIS) and digital elevation model (DEM) to generate ‘viewsheds’ around male locations within a boundary set by the estimated maximal visual acuity of the viewer. Areas visible around leks were compared to those visible around random sites with the same conformation of displaying males. Male sage grouse displayed at sites where surrounding topography both diminished long-range visibility (>1000 m) and enhanced short-range visibility (<500 m) to ground observers. These characteristics could (1) force eagles to monitor lek activity from the air, where they may be more visible to their prey, (2) make displaying males more visible to females and (3) allow males to monitor predators approaching the lek more easily. These results suggest that, in open habitats, visually signalling animals may exploit local topography to control both their visibility to receivers and the visibility of their immediate surroundings.     

Female-female aggression and female mate choice on black grouse leks

We studied female-female aggression in relation to female mate choice in black grouse, Tetrao tetrix, in central Finland, in 1994-1998. Aggression occurred on average every other minute when there was more than one female on a territory, and aggressive behaviour was most prominent when several females attended the lek. Interactions tended to be proportionally most frequent on the territories of the highest-ranking males, although not significantly so. Females that were chased by other females did not mate with lower-ranking males than their aggressors did. Furthermore, chased females were only rarely (6% of cases) forced to move off the territory by agonistic interactions and copulations were disrupted by other females even less often (3% of cases). The choice of a mating territory did not depend on the outcome of aggression even though the aggressors were more likely to mate on the territory where aggression occurred than elsewhere. There was a marginally significant tendency for aggressors to mate earlier in the season. Females placed themselves further away from other females on the territory when soliciting a copulation than just before aggression. Our results suggest that aggression between females does not effectively constrain female choice in black grouse. Its function may be to aid females to secure undisturbed mating opportunities for themselves rather than to prevent others from mating with a particular male. Copyright 2000 The Association for the Study of Animal Behaviour.     

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.     

Manipulating lek size and composition using decoys: an experimental investigation of lek evolution models

Four theoretical models have been proposed to account for the origin and maintenance of leks: hotspot, female preference, hotshot, and black hole models. Each has been validated in particular cases, and most are not mutually exclusive; therefore, it has been difficult to contrast and separate them, empirically and experimentally. By using decoys to mimic natural leks in the little bustard, artificial leks attracted wild birds. Then, by manipulating artificial lek size and structure (sex ratio, male phenotype), the study of responses of wild males and females allowed us to test specific predictions derived from the four classical models of lek evolution. The hotspot model was not supported because female decoys did not attract wild males. Conversely, hotshot males do exist in this species (attracting both wild females and males), as does a female preference for a particular lek size (four males). Finally, males aggressive toward decoys attracted fewer females, consistent with one of the mechanisms by which the black hole model may work. Therefore, three models of lek evolution were partly or fully supported by our experimental results: hotshot, female preference, and black hole models. We suggest that these models actually fit within each other, ensuring the evolution, functioning, and long-term maintenance of leks.     

Limited indirect fitness benefits of male group membership in a lekking species

In group living species, individuals may gain the indirect fitness benefits characterizing kin selection when groups contain close relatives. However, tests of kin selection have primarily focused on cooperatively breeding and eusocial species, whereas its importance in other forms of group living remains to be fully understood. Lekking is a form of grouping where males display on small aggregated territories, which females then visit to mate. As females prefer larger aggregations, territorial males might gain indirect fitness benefits if their presence increases the fitness of close relatives. Previous studies have tested specific predictions of kin selection models using measures such as group‐level relatedness. However, a full understanding of the contribution of kin selection in the evolution of group living requires estimating individuals' indirect fitness benefits across multiple sites and years. Using behavioural and genetic data from the black grouse (Tetrao tetrix), we show that the indirect fitness benefits of group membership were very small because newcomers joined leks containing few close relatives who had limited mating success. Males' indirect fitness benefits were higher in yearlings during increasing population density but marginally changed the variation in male mating success. Kin selection acting through increasing group size is therefore unlikely to contribute substantially to the evolution and maintenance of lekking in this black grouse population.     

The flexible lek: Phymatopus hecta the gold swift demonstrates the evolution of leking and male swarming via a hotspot (Lepidoptera: Hepialidae)

Leking moths present an evolutionary problem in their apparently simultaneous reversal of male–female attraction. The mating system of Phymatopus (or Hepialus) hecta embraces an unusually wide range of procedures. Both males and females use medium‐range olfactory attractants (also probably visual signals), and both sexes will lure, and both will approach, the other, either when flying or perched. This produces an ‘infinite variety’ which includes the classic moth mating procedure (males fly to sessile female); a typical lek procedure (female flies to sessile male); a mating swarm (hovering male follows passing female); and intermediates such as a mutual courtship dance. Male behaviour includes a flying display, two sessile displays, and an escalating war of attrition. The system is versatile, persistent, and probably evolutionarily stable. The lek site has a high density of perches suitable for copulation which facilitate predator‐escape by means of a dead drop. The whole supports a model for the evolution of resource‐based leks which commence with a concentration of females on a hotspot, leading to a concentration of males; and then an escalating process of sexual selection as males become increasingly attractive over a distance to females, and females use the males as a way of locating the resource. The main stabilizing pressure may be selection for efficient mate acquisition, and as with grouse leking systems, the precondition for evolution was probably having travelling females that actively sought a reproductive resource or a predator free space. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 184–201.     

Microsatellite analysis of female mating behaviour in lek-breeding sage grouse

We used microsatellite DNA markers to genotype chicks in 10 broods of lek-breeding sage grouse, Centrocercus urophasianus, whose mothers' behaviour was studied by radio-tracking and observing leks. Previous behavioural studies suggested that almost all matings are performed by territorial males on leks and that multiple mating is rare. Two broods (20%) were sired by more than one male. Genetic analyses of the broods of eight females that visited an intensively studied lek were consistent with behavioural observations. Four females observed mating produced singly sired broods and males other than the individual observed copulating were excluded as sires for most or all of their chicks. Territorial males at the study lek were excluded as sires of broods of four other females that visited the lek but were not observed mating there. Radio-tracking suggested that two of these females mated at other leks. Our results confirm the reliability of mating observations at leks, but do not rule out a possible unseen component of the mating system.     

Indirect Effects of an Existing Wind Energy Facility on Lekking Behavior of Greater Prairie‐Chickens

Rapid expansion of the wind energy industry has raised concerns about the potential effects of anthropogenic disturbance on prairie grouse. While efforts have been made to address the effects of wind energy facilities on measures of fitness, their effect on the behaviors of prairie grouse has been largely neglected. To address these concerns, we investigated the effects of an existing wind energy facility in Nebraska that became operational in 2005 on the lekking behavior of male greater prairie‐chickens Tympanuchus cupido pinnatus between March and May 2013. Given the potential for disturbance caused by wind turbine noise to disrupt acoustic communication and thus behavior, we predicted that males at leks close to, compared to far from, the wind energy facility would spend more time in agonistic behaviors, and less in booming displays. Given the potential for wind turbine noise to reduce the number of females attending leks (hereafter ‘female lek attendance’), we also predicted that males at leks close to the wind energy facility would spend more time in non‐breeding behaviors and less time in breeding behaviors than males farther from the facility. Although we found no effect of the wind energy facility on female lek attendance, males at leks closer to the wind energy facility spent less time in non‐breeding behaviors than those at leks farther away. However, distance from the wind energy facility had no effect on time spent performing booming displays, flutter jumps, or in agonistic behaviors. Given that lekking behaviors of males influence mating success, our results may have consequences for the fitness of prairie grouse breeding in the vicinity of wind energy facilities.     

Context-dependent effects of tail-ornament damage on mating success in black grouse

Male black grouse (Tetrao tetrix) may receive damage to theirtail ornaments, the lyre, during goshawk predation attemptsand during fights with other males. In this study we confirma previous observation that black grouse males with damagedtail ornaments suffer reduced mating success. In males thatheld territories on the edge of the leks, tail damage was unrelatedto mating success, whereas in central males damage was negativelycorrelated with mating success. We tested experimentally whetherabsence of damage is used by females in mate choice. In maleswith edge territories, intact, control males had higher matingsuccess than males with cut tails, but in males with centralterritories, lyre cutting had no effect on mating success. Theseresults suggest two interpretations. First, female choice alsodepends on factors other than tail damage such as position onthe lek and dominance. Second, the effect of tail damage iscontext dependent; in males that otherwise meet females standards(e.g., dominant males), the effect of tail damage is negligible,but in less dominant males, tail damage could be used by femalesin mate selection. The second interpretation provides an explanationfor why the data on unmanipulated and manipulated birds differ.In experimental central birds, factors other than tail damageprobably determine male mating success, whereas in experimentaledge birds such factors are probably absent and therefore taildamage is relatively more important. In central unmanipulatedbirds, however, males with natural damage are probably not chosenbecause tail damage and absence of other attractive traits arecorrelated. The absence of an effect on peripheral unmanipulatedbirds may be explained by their overall low mating success     

Female choice selects for lifetime lekking performance in black grouse males

''Good genes'' models assume that females can use a signal such as mating effort to assess a male''s lifetime fitness. Inferring long-term performance from short-term behavioural observations can be unreliable, and repeated sampling may be needed for more accurate assessment of males. Additionally, if sexual advertisement is viewed as a life-history trait subject to trade-offs, reliable comparison of mates should yield information on all life-history components rather than on one trait value in one season. We show that in the lekking black grouse (Tetrao tetrix), a male''s success is best explained by assuming that females are informed of the past history of males up to the beginning of the study (eight years). Much of this extremely lasting ''memory'' can be attributed to females observing long-term outcomes of male–male competition: current territory position is the only momentarily observable variable that has high power in predicting female choice, and it correlates to a male''s past lekking effort on a cumulative lifetime scale. We conclude that females can use territory position as a signal that conveys information of a male''s lifetime performance that combines lekking effort and longevity. Females may thus overcome the problem of male allocations varying in time, without the need to pay costs associated with repeated sampling.     

Mate choice for offspring performance: major benefits or minor costs?

There is considerable disagreement over whether or not gaining viability benefits to offspring could be substantial enough to overcome the costs of female choosiness. A recent review suggested that the ''lek paradox'' might be resolved by large indirect benefits as indicated by highly heritable ornamental traits. We selected males of a wolf spider Hygrolycosa rubrofasciata in relation to their sexual signalling rate (audible drumming). The estimated correlated response in offspring viability was rather small (0.12 s.d.). However, it may be large enough if the costs of being choosy are small. In fact, females mate with better-than-average males just by responding passively to a random drumming signal, and the active choice by females seemed to increase this benefit only slightly. In many mating systems, females obtain better-than-average males as a consequence of intense male–male competition or because of the extraordinary variance in male signalling. The costs of any additional choice may be so minor that female choice for honestly signalling males may evolve even with minute benefits in offspring viability. This may be the general solution to the lek paradox, as most studies report no apparent fitness benefits. Publication bias favouring statistically significant results may have led to an overemphasis on the few studies with large effects.     

Sharp-tailed grouse lek attendance and fidelity in upper Michigan

To assess and improve existing monitoring protocols for sharp-tailed grouse (Tympanuchus phasianellus) in the eastern Upper Peninsula of Michigan, we used data from 58 radio-collared grouse (46 M, 12 F) monitored within 3 openland landscape types: a xeric, conifer-dominated site, a wetland-dominated site, and a site dominated by low-intensity agriculture. We used lek counts and radio telemetry to determine lek attendance rates, factors affecting lek attendance rates, lek fidelity, and inter-sexual variation in these parameters. Our analysis indicated lek attendance varied with respect to sex of bird, day of year, time after sunrise, and wind speed. Peak male lek attendance rates exceeded those of females by up to 40%, and peak lekking activity for both males and females occurred during the second and third weeks of April. Male lekking activity occurred earlier and was sustained longer than that of females. Lekking activity was negatively related to time of day and wind speed. We observed strong lek fidelity as radio-collared birds attended a primary lek 94% of the time, indicating a low probability of multiple counting of individual birds. We also proposed a method to adjust lek count data for the probability that birds are on a lek during lek counts. Our proposed method can be used by researchers and managers to improve estimates of the number of birds attending a lek by reducing the negative bias associated with observed counts. © 2011 The Wildlife Society.     

Combs and sexual selection in black grouse (Tetrao tetrix)

We studied supra-orbital combs in lekking black grouse (Tetrao tetrix) in relation to sexual selection at five leks in Finland1991-1998 and four leks in Sweden 1992-1995. Comb size wasestimated in two ways: by observing its natural size in thefield at different behaviors ("observed comb size"), and bymeasuring the comb size from captured birds ("measured combsize"). The size of combs is highly variable, and individualscan change it within seconds. Males express their larger combsduring display, as compared to other behaviors. Observed mean comb sizes were larger on leks with a higher number of malesand a higher number of copulations. Measured and observed combsizes and copulatory success did not significantly correlatewhen all males where analyzed, but a positive and significantrelationship between observed comb size and copulatory success was found within males that achieved copulations. Measured comblength correlated positively with the amount of testosterone.While females were present on the lek, displaying and successfulmales showed the largest observed comb size. When we comparedobserved comb size during fighting between successful and unsuccessfulmales and correlated comb size of pairs of fighting males withtheir fighting activity, no significant differences in combsize were found. The result that comb size correlated significantlywith an increase in testosterone level and that larger combsize, within successful males, predicted higher copulatorysuccess suggests that combs may be a cue for females to assessmale quality. The lack of a significant relationship betweenobserved comb size and fighting behavior suggests that combsize either has minor importance in male-male signaling onthe lek or that males may express similar-sized combs duringfighting to avoid serious fights and thus risk of comb injuries.     

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.