Lek centre attracts black grouse females

In several lek mating systems, centrally located males enjoy higher mating success than peripheral males. The mechanism behind this pattern, however, has been controversial and a rigorous test of the different alternatives is missing. Here we report that in black grouse Tetrao tetrix , a lekking bird species, central males achieved many more copulations than males in the periphery. Generally, central territories were smaller than peripheral ones, and males were thus more clumped in the centre. A choice experiment in an aviary provided experimental support for females being attracted to densely clustered males, so that males in larger clusters have higher expected average mating success. Thu…

Male coloration and species recognition in sympatric flycatchers

Currently favoured views for explaining ornaments in males emphasize female preference such that females benefit from increased offspring production, good genes of the offspring, or the attractiveness of sons. Results from long-term studies in the pied flycatcher Ficedula hypoleuca suggest that factors associated with species recognition may also be important for male coloration. In sympatry the collared flycatcher F. albicollis is dominant in competition for nesting sites over the pied flycatcher. Bright pied flycatcher males resemble collared flycatcher males and suffer from interspecific interference, whereas dull and female-like males can acquire nesting sites close to those of the coll…

The effects of parasites on male ornaments and female choice in the lek-breeding black grouse (Tetrao tetrix)

We describe the results of two studies of parasitic infection in the black grouse (Tetrao tetrix). The first deals with our own observations of lekking black grouse in which the parasite levels of two blood parasites, the protozoan Leucocytozoon lovati and microfilaria, probably produced by a nematode worm Splendidofilaria tuvensis, were scored. We also obtained measures of age, survival, number of copulations, body mass and length of the ornamental tail feathers (the lyre) of the lekking males. The second study analysed the data from Lund (1954) which involved eight gut parasites obtained from birds which were killed. In the first study we found higher levels of infection of Leucocytozoon …

Extra-pair paternity and male characteristics in the pied flycatcher

The pied flycatcher (Ficedula hypoleuca) is sexually dichromatic with extreme variation in male plumage coloration. The benefit for males of having black plumage is controversial, and few studies have found evidence for a sexual selection benefit of being black rather than brown. However, blacker males may be better able to achieve extra-pair fertilizations (EPFs), which may be an important component of sexual selection. We studied the role of EPFs in sexual selection in the pied flycatcher by establishing a set-up where two males with different back coloration (blacker vs browner) bred simultaneously near each other. DNA fingerprinting analysis revealed that 11% of offspring resulted from …

Paternity, copulation disturbance and female choice in lekking black grouse

Female copulation behaviour in the black grouse, Tetrao tetrix, was studied by detailed observations of individually marked birds together with DNA fingerprinting analyses for paternity assessment. For each breeding occasion females typically mated only once with one male, and did not mate outside the lek arena. They re-mated with the same or another male usually only if the initial copulation was disturbed and was probably unsuccessful in sperm transfer. The willingness of females to copulate only once with one male suggests that multiple mating with several males incurs a cost that more than outweighs any possible fertility or sperm competition benefits. Attempts by neighbouring males to …

Why Do Black Grouse Males Perform on Lek Sites outside the Breeding Season?

Studies of sexual selection have focused mainly on explanations of how male characteristics influence mating success during the breeding season. By contrast, few studies have investigated how behaviours prior to the breeding season influence mate choice d

Assortative mating and female clutch investment in black grouse

Variation in female behaviour has only recently received attention in studies of sexual selection. It has been suggested that females may invest differentially in their offspring in relation to the quality of their mate. This may lead to females that mate with high-quality and/or attractive males laying larger clutches. Females may also differ in their ability to choose between males. For example, females in good physical condition may make better choices. If physical condition and clutch size are positively correlated, this hypothesis could also produce a relationship between male attractiveness and female clutch size. We found, in lekking black grouse, Tetrao tetrix, that females mated to…

The search cost in mate choice of the pied flycatcher

Variation in clutch size in relation to nest size in birds

© 2014 The Authors. Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nes…

Mate sampling behaviour of black grouse females (Tetrao tetrix)

We studied female mate sampling behaviour in lekking black grouse (Tetrao tetrix). Females mainly visited males occupying territories in the centre of the lek with relatively large territories. They were also more likely to visit males that had high attendance. The same factors were also correlated with male mating success. A multiple regression model including these factors explained more of the variance in female visits per male (53%) than in mating success (33%). The pattern of female sampling conformed with a pool comparison (best-of-n) tactic. Such a tactic is expected if the costs of sampling are low. Females of high body mass visited more males than lighter females, however, which in…

Can female preference explain sexual dichromatism in the pied flycatcher, Ficedula hypoleuca?

How important female choice is for the evolution of male secondary sexual characteristics is controversial. Two field and one laboratory experiment, using the pied flycatcher, were performed to test the female choice aspect of sexual selection. In addition, non-manipulative data from 5 years are presented. The observational data suggest a slight preference for dark males by females but in field experiments in which males had territories at random sites (i.e. they did not choose a territory) or the colour of concurrently arriving males was altered, there was no preference for darker ones. Similarly, oestradiol-treated females did not prefer black or brown males in the laboratory. Thus, there…

Copying the Mate Choice of Others? Observations On Female Black Grouse

Abstract It has been suggested that the non-random mating often observed in lekking species is a consequence of either male-male competition or active female mate choice. Here we show that the highly skewed mating distributions observed in a black grouse lek in three years were indeed different from random expectations. We suggest that females copying the mate choice of others enhance this skew. Observations in favour of copying are: females pay multiple visits to the lek during several days; females arrive and move in bands which makes it possible to observe the visits to male territories and matings of other females; in the main lek in the study area, males often mated in sequence indica…

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

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

Polyterritoriality and deception

Testosterone and male mating success on the black grouse leks

On black grouse (Tetrao tetrix) leks females prefer dominant, viable males that have managed to acquire relatively central territories. The immunocompetence hypothesis predicts that, because high levels of testosterone are costly to the immune system, male sexual traits that are controlled by testosterone are likely to serve as reliable indicators of male health. Indeed, testosterone concentrations of black grouse males were highly variable, and strongly correlated with male mating success. This is related to the fact that males with high testosterone levels had most central territories. However, the association of testostestorone level with male mating success was not solely related to cen…

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, t…

Singing rate and female attraction in the pied flycatcher: an experiment

«Attirer les femelles» est une des principales fonctions du chant des oiseaux. Beaucoup d'etudes ont mis en evidence l'effet de variations du repertoire sonore sur le succes d'accouplement mais peu d'entre elles sont consacrees a l'intensite du chant. Dans une telle optique, cette etude examine les effets de manipulations experimentales de chants sur la selection du partenaire par les femelles

Clutch-size variation in Western Palaearctic secondary hole-nesting passerine birds in relation to nest box design.

Møller, A.P. [et al.]

Inbreeding depression and male fitness in black grouse

The male lifetime lekking performance was studied, and related to inbreeding-outbreeding in a wild population of black grouse (Tetrao tetrix) in central Finland between 1989 and 1995. Inbreeding was measured as the mean heterozygosity and mean d(2) of 15 microsatellite loci. We found a significantly positive relationship between mean d(2) and lifetime copulation success (LCS), while the relationship between heterozygosity and LCS was close to significant. We also found that males that never obtained a lek territory had significantly lower mean heterozygosity than males that were observed on a territory at least during one mating season in their life. Furthermore, among males that were succe…

Polygyny and Breeding Success of Pied Flycatchers Nesting in Natural Cavities

Many hole-nesting passerine birds accept, or even prefer, nest boxes. Hence most of our knowledge about the breeding biology, population dynamics, life-history evolution and so forth of such species comes from nest box studies. This, in some cases, might lead to erroneous conclusions. For example, breeding density in nest box areas may often be much higher than in the natural situation (e.g. von Haartman, 1971), possibly leading to unnatural density dependent effects. Furthermore, the use of nest boxes may reduce predation risks (e.g. Nilsson 1975, 1984 a,b, Moller 1989), while the routine procedure of cleaning boxes after each breeding season may reduce the number of parasites, thus affect…

Interspecific variation in the relationship between clutch size, laying date and intensity of urbanization in four species of hole-nesting birds

Marie Vaugoyeau [et al.]

Lekking in the black grouse— a test of male viability

LEKS, where males congregate to display and females attend only to mate, present one of the most remarkable outcomes of sexual selection1. It is a common but untested belief that females mate with the most vigorous males2. In leks of the black grouse Tetrao tetrix, males dominant in winter flocks were most successful in mating, as were males winning fights over female dummies placed at territory boundaries. Males tear feathers from each others' tail ornaments in combats, and attractive males always had undamaged tails. We report here that by choosing victorious males, females mate with males that are most likely to survive the following six months. There is a strong association between fema…

Fluctuating asymmetry and copulation success in lekking black grouse

Abstract In lekking black grouse, Tetrao tetrix males at the centre of the leks obtain more copulations than males at the edges. We found that males with territories at the edge of the lek obtained fewer matings and also have the most asymmetric tarsi. However, when considering the tail ornament (the lyre) no correlation between asymmetry and mating success was found. Although females are unlikely to select males on the basis of tarsus symmetry, asymmetry in this trait may reflect male condition. We suggest that males in poor condition, as reflected by higher levels of fluctuating asymmetry in the tarsi, do not achieve central positions on the lek arena. This would account for the lower suc…

Combs and sexual selection in black grouse (Tetrao tetrix)

We studied supra-orbital combs in lekking black grouse (Telrao tetrix) in relation to sexual selection at five leks in Finland 1991-1998 and four leks in Sweden 1992-1995. Comb size was estimated in two ways: by observing its natural size in the field at

Patterns of variation in tail ornament size in birds

In recent years several different kinds of sexual selection models have been developed, and tail ornaments in birds have frequently been used as an example of a sexually selected character where the models might apply. However, very little is known about intra- and interpopulation variation in ornament size. We have studied the elongated tail ornaments in four species of whydahs Vidua, the forktailed flycatcher Tyrannus savana and the Asian paradise flycatcher Terpsiphone paradisi. Ornaments were relatively longer in males with the longest tarsi (‘heterogony’ with positive allometry). Also, tail lengths were remarkably variable within each geographical area, the coefficient of variation (av…