Lek Formation Research Articles

Male Aggregation in Dwarf Tyrant-Manakins and What It Tells Us about the Origin of Leks.

Despite considerable research on lek-breeding birds, many aspects of the evolution of the lek social system, including the origin of male aggregation, remain unresolved. Originally, clusters may have formed passively as a by-product of independent but concurrent responses by males to some external stimulus (e.g., a concentration of display sites, prominent food source, or a travel corridor commonly used by females) that drew them at a particular time to a site where they might increase their probabilities of encountering females. In contrast, male aggregation may have been active, with a male purposely seeking to associate with another male (or males) because proximity to that male allowed him to enhance his own reproductive success by improving his ability to attract females (i.e., learning from his associate) or to intercept females attracted to that male. To identify factors important in the formation of male aggregations, I studied these alternatives in the Dwarf Tyrant-Manakin (Tyranneutes stolzmanni), a species in which some breeding males occupy solitary courts where they attempt to attract females for reproduction, whereas other males occupy courts clustered into leks, where they interact to attract females. I compared habitat characteristics and resource availability at courts of grouped and solitary males as well as certain traits of the males in an attempt to distinguish between passive or active lek formation in this species. Little evidence suggests that Dwarf Tyrant-Manakins are passively clustered as a result of common attraction to or use of a limited or unevenly distributed resource. Several observations suggest the contrary. Evidence for the active aggregation of males is more promising, suggesting that males may associate with other males on the basis of their vocal behavior. The issue is confounded, however, by the observation that male call behavior appears to change with a change in social status. A definitive answer for T. stolzmanni will require additional analyses of vocal behavior as well as of the characteristics of the vocalizations themselves. Finally, the behavior and social organization of this species may reflect characteristics of the ancestral manakin line that gave rise to both the Tyrant and Core clades of modern manakins.

Biology of a putative male aggregation-sex pheromone in Sirex noctilio (Hymenoptera: Siricidae).

A putative male-produced pheromone has recently been described for the global pest of pines, Sirex noctilio, but field-activity has not been demonstrated. This study aimed to investigate the pheromone biology of S. noctilio in more detail. Specifically, we i) analysed effluvia and extracts for additional compounds by gas chromatography coupled with electro-antennographic detection (GC-EAD), mass spectrometry (GC-MS) and two dimensional time of flight mass spectrometry (GC X GC TOF MS), ii) conducted dose-response experiments for putative pheromone components, iii) determined the site of synthesis/ storage of the putative pheromone and iv) determined the release rate of the putative pheromone from males and three types of lures. A blend of four compounds was identified, including the previously described (Z)-3-decenol and (Z)-4-decenol, and two new compounds (Z)-3-octenol and (Z)-3-dodecenol. All compounds elicited a response from both male and female antennae, but the strength of the response varied according to sex, compound and dose tested. (Z)-3-Decenol and (Z)-3-octenol at lower and higher doses, respectively, elicited larger responses in males and females than the other two compounds. (Z)-3-Octenol and (Z)-4-decenol generally elicited larger female than male antennal responses. The site of synthesis and/or storage in males was determined to be the hind legs, likely in the leg-tendon gland. The relative release rate of the major compound by male wasps was shown to be 90 ± 12.4 ng/min, which is between 4 and 15 times greater than that observed from typical lures used previously. These observations are consistent with the hypothesis that these compounds may mediate lek formation in S. noctilio males and lek location in females.

Male relatedness, lekking behavior patterns, and the potential for kin selection in a Neotropical hummingbird

AbstractSeveral models have been proposed to explain the evolution of leks, both in terms of direct or indirect fitness benefits, and in survival. According to kin selection theory, male skewed reproductive success leads unsuccessful males to join successful relatives to increase their inclusive fitness, because their genes would be transmitted indirectly to the next generation. Wedge-tailed Sabrewing (Campylopterus curvipennis) is a hummingbird species whose males congregate at leks, in which spatially clustered males sing a particular song with marked differences among neighboring males (song neighborhoods). The maintenance of song neighborhoods presumably depends on juvenile newcomers copying the song type of their neighbors when they establish within a lek, and their acceptance could be more likely if a relative has already settled down in a territory, which in turn could offer fitness benefits explained by kin selection theory. To investigate the potential for kin selection in this species, we genotyped 126 hummingbirds at 10 microsatellite loci and estimated pairwise relatedness among males at 6 leks and in 4 song neighborhoods within 1 focal lek. Within leks, most males were unrelated and only a few were relatives. Moreover, even though relatedness within leks was higher than between leks, it was not higher than 0, which is likely due to isolation by distance. Our results do not support the idea of kin selection as an important force acting on the formation of leks in this species. Additionally, we found no evidence for kin clustering within song neighborhoods, suggesting that juveniles attempting to settle in a lek have to learn the song of the neighborhood (regardless of their kinship) to gain access to territories.

Effect of mass rearing and gamma irradiation on the mating behaviour of Eldana saccharina

AbstractAt low doses of radiation, F1 sterility of the sugarcane stalk borer, Eldana saccharina Walker (Lepidoptera: Pyralidae), can be attained. This bodes well for progression of the sterile insect technique (SIT) as part of an area‐wide integrated pest management strategy against this pest of economic importance. However, success of the SIT is dependent on mating ability and performance of the mass‐reared sterile insects once released into the field. Mating behaviour of non‐irradiated laboratory‐reared male E. saccharina moths and those subjected to radiation (150, 200, or 250 Gy) was investigated in still‐air cage laboratory trials. Video technology was used to determine whether mass rearing and irradiation of adult moths altered their performance in lek formation and mating success, compared to their untreated wild counterparts. Mating frequency, mating duration, onset of mating, and calling during the first two scotophase periods following adult eclosion were used to make these assessments. Results indicated that both irradiated and non‐irradiated laboratory‐reared males are able to form leks and mate with wild females, and are therefore as effective as their wild counterparts with regards to mating performance. However, a quantitative departure of male mating behaviour away from that exhibited by wild males was observed in the non‐irradiated and irradiated laboratory males. These commenced calling and mating earlier than wild males in both scotophases, except for those irradiated at 250 Gy in the case of mating onset. The mating duration of treated males was similar to that of the untreated wild males, except for those irradiated at 150 Gy, which was significantly higher. Our study provides evidence that increasing radiation dose negatively impacts on performance of laboratory‐produced moths relative to their wild counterparts, which in turn could affect the efficacy of the SIT. At 150 and 200 Gy, these parameters are not affected significantly when compared to untreated wild males, and thus these radiation doses can be used safely. However, at higher doses negative effects do become evident.

The kin selection hypothesis in a lekking mole cricket: assessing nested patterns of relatedness

Loosely defined, a lek is a male mating aggregation visited by females primarily for the purpose of fertilization. No consensus has been forged that successfully explains how and why leks have evolved across the full breadth of lekking taxa. The two major conceptual explanations (cooperation and competition) are both intricately intertwined in any given system and exhibit varying levels of plasticity based on an organism's environment and life history. The kin selection hypothesis suggests that if females prefer larger leks (as is often the case), unattractive males may aggregate with their attractive relatives in order to boost the latter's reproductive success, while effectively sacrificing any opportunities of their own. Here we develop microsatellite markers to genetically evaluate the kin selection hypothesis by measuring relatedness and precise spatial locations of males in a population of the lekking prairie mole cricket (Gryllotalpa major Saussure). Results indicate that neighbouring males are often highly related, suggesting that kin selection may play some role in this system. However, because leks are comprised of multiple kin groups, if kin selection is operating it is likely doing so at a smaller scale than predicted by the kin selection hypothesis of lek formation. The high levels of subgroup relatedness within this species likely occurs as a passive process due to male viscosity, but the functional implications of this interesting genetic organization remain unknown.

Reproductive Behavior and Basic Biology of the Oriental Bamboo-Inhabiting Anoplomus rufipes and a Comparison with Frugivorous Dacinae Fruit Flies.

The reproductive behaviors and mating systems of the fruit-infesting species of the Dacinae tribes Ceratitidini and Dacini are increasingly well understood, while in the non-frugivorous tribe Gastrozonini, data are lacking. In the present study, the reproductive behavior of Anoplomus rufipes from North Thailand was studied in the field, other behaviors also in the laboratory. A. rufipes mated on young bamboo plants growing in areas destroyed by fire. Exudates of extrafloral nectaries produced by the young bamboo plants provided food for the females. Factors affecting the choice of the mating site were favorable microclimatic conditions and food. Courtship behavior was performed on the upper sides of bamboo leaves and included pheromone calling (abdominal elevation, anal pouch eversion, abdominal pleural distention), anal dabbing, looping flights and a specific lofting/body swaying behavior. The males searched individually for females or formed leks containing up to four males. The reproductive behaviors and lek formation of A. rufipes are compared to other Dacinae (Ceratitis, Bactrocera), and their functions are discussed. Hitherto unknown data on the general biology of A. rufipes are also included. A. rufipes larvae infested living bamboo shoots of Cephalostachyum pergracile, and the observed behaviors of the adults included locomotion, grooming, feeding, oral droplet deposition, bubbling and agonistic behavior.

Population structure and mating system of the Red-crested Korhaan (Lophotis ruficrista) in South Africa

BackgroundThe mating system of the korhaans and bustards in southern Africa is either based on polygyny or monogamy. The Red-crested Korhaan (Lophotis ruficrista) has been described as polygynous but otherwise very little is known about its breeding biology. The aims were to describe the population structure and male behavior during breeding.MethodsThe data collected for this paper was mainly based on field transect surveys carried out in 2009, 2010 and 2011. The population structure of the Red-crested Korhaan in the temperate Borakalalo Game Reserve (BGR) was compared with that in the arid Molopo Nature Reserve (MNR) (both situated in the North West province of South Africa). The study was mainly conducted in the BGR but additional work was conducted in the MNR for comparison.ResultsThe difference in mean group sizes between MNR (1.03) and BGR (1.07) was statistically not significant. Group sizes were not affected by climate despite the fact that the BGR received about 650 mm precipitation per annum and the MNR about 200 mm. In both reserves the apparent sex ratios were skewed in favor of males (1:0.29 in the BGR and 1:0.1 in the MNR). The population was dispersed in a clumped manner which is ascribed to the formation of leks. Leks were positioned in open habitat while females invariably concealed themselves under cover.ConclusionsThe leks formed the pivot of the mating system of the Red-crested Korhaan and are identifiable social structures that field ecologists can use to monitor population stability.

Density dependence and habitat quality modulate the intensity of display territory defence in an exploded lekking species

We evaluated the effect of conspecific abundance and habitat quality of leks on the territorial behaviour of males in an exploded lekking species, the Little Bustard (Tetrax tetrax). The hypothesis that males more intensely defend territories with higher conspecific abundance and better habitat quality was evaluated experimentally analysing the agonistic response of experimental males to male decoys placed on their displaying areas. Decoy experiments showed that the intensity of display territory defence by little bustard males is density dependent. The time experimental males took to return to their display sites after decoy placement decreased with abundance of both males and females. The strength of their final response was positively associated to local male and female abundance in the vicinity of their display sites. Habitat quality also influenced males’ display territory defence since the intensity of male response increased with the degree of natural vegetation cover. Habitat quality was particularly relevant in explaining variation of experimental males’ snort call rate, which decreased with the degree in plough cover and increased with the number of fields in the lekking area. Snort call rate decreased with the level of male aggregation and was lowest in males exhibiting the strongest aggressive response to decoys. These results add new evidence for the density dependence of species’ breeding territorial behaviour, supporting density-dependent models of lek formation and reinforcing the role of resources defence in exploded lek mating systems.

Female Preference and Predation Risk Models Can Explain the Maintenance of a Fallow Deer (Dama dama) Lek and Its ‘Handy’ Location

We tested the predictions of three models (female preference; hotspot; predator avoidance) on lek formation in the fallow deer population of San Rossore, Tuscany. We collected behavioural observations in two leks and radiotracking data on 67 deer over 7 years. Two deer sub-populations were present in the northern and southern sides of the area, respectively, the two sectors being delimited by a river and including one lek each. Predictions were tested for one lek (SG), located in the south-side where we set up our 7-year radiotracking program. Data from a second lek (FO, north-side) were used to test those predictions which imply the occurrence of multiple leks in the same population. We showed that the majority of females made one single visit to one lek, only during the rut. The lek was located outside areas of higher female traffic and home range overlap, and females increased home range sizes during the rut to reach it. Twilight routes of females never crossed the lek; instead, females walked atypical routes and at a faster pace to reach the lek and mate. The distance between the two leks was higher than the average diameter of female home ranges, and only one lek was present within female home ranges. Males reached the lek one month before the arrival of females, corroborating that lekking is a female-initiated process (females moving towards large clumped male aggregations) rather than a male-initiated process (males moving towards female hotspots). Our results supported the female preference model, and rejected the predictions of the hotspot model. Also, leks were located far from areas with higher predation risk, supporting the predator avoidance model. The position of lek SG resulted ‘handy’ at the sub-population level because of the optimal trade-off between travel costs for females to reach it and avoidance of human predators.

Modelling female mating success during mass trapping and natural competitive attraction of searching males or females

AbstractSimulation models of insects encountering sex pheromone with or without mass trapping in which the searching sex is either male (moths and many insect species) or female (some true bugs, beetles, and flies) were developed. The searching sex moved as a correlated random walk, while the opposite sex remained stationary (calling) and released an attractive sex pheromone. The searching sex was caught when encountering a pheromone‐baited trap, and females mated when encountering a male. An encounter with pheromone was defined by the searcher's interception of a circle termed the effective attraction radius (EARc). Parameters of movement (speed and duration), initial numbers of calling sex and searching sex, number of traps, area, and EARc of traps and calling sex were varied individually to evaluate effects on the percentage of females mating. In the natural condition without traps, female mating success in both models was identical. Increasing the EARc of the calling sex caused diminishing increases in female mating success, suggesting that evolution of larger pheromone release and EARc is limited by increasing costs (production/sensitivity) relative to diminishing increases and benefits of mating encounters. With mass trapping, increasing the EARc of traps or density of traps caused similar declines in female mating in both models, but the female‐searching model predicted slightly lower mating success than the male‐searching model. Increasing the EARc of calling insects or the initial density of insects caused similar increases in female mating in both models, but again the female‐searching model had slightly lower mating success than the male‐searching model. The models have implications for mating lek formation and for understanding the variables affecting the success of mass trapping programs for insect pests with either male or female sex pheromones.

Spatial distribution and male mating success of Anopheles gambiaeswarms

BackgroundAnopheles gambiae mates in flight at particular mating sites over specific landmarks known as swarm markers. The swarms are composed of males; females typically approach a swarm, and leave in copula. This mating aggregation looks like a lek, but appears to lack the component of female choice. To investigate the possible mechanisms promoting the evolution of swarming in this mosquito species, we looked at the variation in mating success between swarms and discussed the factors that structure it in light of the three major lekking models, known as the female preference model, the hotspot model, and the hotshot model.ResultsWe found substantial variation in swarm size and in mating success between swarms. A strong correlation between swarm size and mating success was observed, and consistent with the hotspot model of lek formation, the per capita mating success of individual males did not increase with swarm size. For the spatial distribution of swarms, our results revealed that some display sites were more attractive to both males and females and that females were more attracted to large swarms. While the swarm markers we recognize help us in localizing swarms, they did not account for the variation in swarm size or in the swarm mating success, suggesting that mosquitoes probably are attracted to these markers, but also perceive and respond to other aspects of the swarming site.ConclusionsCharacterizing the mating system of a species helps understand how this species has evolved and how selective pressures operate on male and female traits. The current study looked at male mating success of An. gambiae and discussed possible factors that account for its variation. We found that swarms of An. gambiae conform to the hotspot model of lek formation. But because swarms may lack the female choice component, we propose that the An. gambiae mating system is a lek-like system that incorporates characteristics pertaining to other mating systems such as scramble mating competition.

Bat predation and the evolution of leks in acoustic moths

Theories of lek evolution generally invoke enhanced mating success experienced by males signalling in aggregations. Reduced predation has also been acknowledged as a potential factor driving lek formation, but its role is more ambiguous. Although lekking is a complex behaviour, few empirical studies have investigated the role of both claims. We studied the potential pressures imposed by mating success and predation in an acoustic moth, Achroia grisella, in which males gather in leks and broadcast a calling song attractive to females. We exploited the ability to manipulate the distribution of singing males in laboratory arenas to create different-sized leks and tested female preferences for these aggregations. Because A. grisella are vulnerable to predation by bats while in flight and on the substrate, we also tested the responses of a potential predator, Rhinolophus ferrumequinum, a bat species that feeds on moths, to the experimental leks. We found that the per capita attractiveness of A. grisella males to females rose with increasing lek size. R. ferrumequinum also oriented toward experimental A. grisella leks, but this attraction did not increase at larger leks. Thus, a male’s per capita exposure to predation risk declined as more moths joined the lek. A. grisella males appear to benefit from advertising in larger leks in terms of both increased mate attraction and reduced predation risk. Our results support the idea that multiple factors operating simultaneously may maintain lekking behaviour.

Aggression in closely related Malawi cichlids varies inversely with habitat complexity

Within the past 2 million years, the cichlids of Lake Malawi have diversified into well over 500 species resulting in one of the worlds largest lacustrine fish radiations. As a result, many of the habitats within the lake support a high diversity of species. In these highly species rich communities, male cichlids must acquire and defend a territory to successfully reproduce. Within the rock-dwelling cichlids of Lake Malawi (mbuna), this has resulted in the formation of polyspecific leks on the heterogeneous rocky benthos. Aggression is fairly common in these leks and has been tied not only to individual reproduction but to the larger phenomenon of community assembly and the maintenance of biological diversity. In this study, I examined the patterns of aggressive acts of four species within the mbuna genus Maylandia at two locations in the southern Lake Malawi. The number of aggressive acts of two sympatric species was examined at each location. At each site, one species defends territories over bedrock and the other over cobble. The number of aggressive acts across the four species was compared. The influence of habitat type on male aggression was examined and the targets of male aggression were identified to evaluate several hypotheses concerning the evolution of male aggression. The results show that aggression quantitatively varied among species, was largely directed towards heterospecifics, and was strongly influenced by habitat type. The aggressive behavior of one sympatric species pair, Maylandia benetos and Maylandia zebra, was observed under controlled laboratory conditions. Laboratory results support field observations: the bedrock associated species performed more aggressive acts and aggressive acts were directed equally at con- and heterospecifics. The results of this study suggest that habitat complexity plays a larger role in shaping aggressive behavior than other suggested factors such as competition for resources.

Male territoriality and mating system in the European beewolf Philanthus triangulum F. (Hymenoptera: Crabronidae): evidence for a “hotspot” lek polygyny

The evolution of the mating system of a species is strongly influenced by the spatial and temporal distribution of females and/or resources. Here, we describe aspects of the territorial behavior of males of a solitary digger wasp, the European beewolf (Philanthus triangulum) and characterize the mating system. We show that beewolf males establish small territories that do not contain any resources essential to females. These territories are intensively scent-marked with a pheromone from a cephalic gland and are defended against intruders in combat flights. We provide evidence that scent-marking constitutes a chemical display and that the pheromone serves to attract receptive females and, thus, represents a sex pheromone. Using spatial statistics, we show that beewolf territories are clumped in space both with respect to other male territories and, more importantly, with respect to female nesting sites. Additionally, the proportion of days a territory is occupied by a male is correlated with the number of female nests in the vicinity. Taking into account that beewolf males do not defend or provide resources essential to females, but merely display chemically to attract females for mating in an aggregation of territories close to female nesting sites, we conclude that the European beewolf exhibits a hotspot lek polygyny with female nesting sites constituting “hotspots” for lek formation.

Hotshots, hot spots, and female preference: exploring lek formation models with a bower-building cichlid fish

In many animals, males congregate in leks that females visit for the sole purpose of mating. We observed male and behavior on 3 different-sized leks of the bower-building cichlid fish Nyassachromis cf. microcephalus to test predictions of 3 prominent lek models: the hotshot, spot, and female models. In this system, we were able to refine these predictions by distinguishing between indirect mate choice, by which females restrict their set of potential mates in the absence of individual male assessment, and direct mate choice, by which females assess males and their territories through dyadic behavioral interactions. On no lek were males holding central territories favored by indirect or direct mate choice, contrary to the prediction of the hotshot model that leks form because inferior males establish territories surrounding hotshot males preferred by females. Average encounter rate of males increased with lek size, a pattern typically interpreted as evidence that leks form through preference for lekking males, rather than because males congregate in hot spots of high density. Female propensity to engage in premating behavior once courted did not increase with lek size, suggesting preference for males on larger leks operated through indirect choice rather than direct choice based on individual assessment. The frequency of male-male competitive interactions increased with lek size, whereas their foraging rate decreased, implying a cost to males maintaining territories on larger leks. Together these data most strongly support the preference model, where females may benefit through indirect mate choice for males able to meet the competitive cost of occupying larger leks.

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.

Capercaillie Tetrao urogallus Lek Formation in Young Forest

Abstract Capercaillie Tetrao urogallus leks have repeatedly been reported to be located in old forest. However, two conditions may have biased this widely held view. First, leks are known to be continuously used over several decades, and therefore might have been established when forest stands were younger. Second, stand-replacement logging (clearcutting) was not widely applied until the 1950s, leaving even-aged regenerating stands too young for leks to have been established in the latter part of the 20th century. Here we report eight cases of lek formation in young plantations from south-central Norway. Stand age ranged within 26-46 years when display activity started. At six of the sites, we confirmed that females were regularly feeding on pine trees in winter prior to lek establishment, and at four of these sites displaying males were observed courting the females in late winter. These findings support the hotspot model of lek formation put forward by Gjerde et al. (2000), and it offers promising optio...

LEKKING BEHAVIOR OF ANASTREPHA FRATERCULUS (DIPTERA: TEPHRITIDAE)

Anastrepha fraterculus (Wiedemann) (Diptera: Tephritidae) displays a lek mating system. Males form groups in which they simultaneously display signals (acoustical, visual, or chemical) to attract females with the purpose of mating. Females visit the lek and choose among signaling and courting males to mate. Scarce information is available in A. fraterculus about the main factors involved in female choice and the behavior of displaying males. This information could be important within the context of pest control programs with a sterile insect technique (SIT) component, because departures from normal sexual behavior caused by artificial rearing could affect males’ performance in the field. In this study we assessed A. fraterculus male behavior within the leks and analyzed the importance of behavioral and morphological traits on their copulatory success. The existence of preferred places for lek formation was evaluated in field cages with trees inside and analyzed by dividing the trees in sectors according to a 3-dimensional system. Males were individually weighed, marked, and observed every 15 min. Morphometric and behavioral characteristics of successful and unsuccessful males were compared. Most successful males grouped in a region of the tree characterized by the highest light intensity in the first 2 h of the morning. Results showed that pheromone calling activity is positively associated with copulatory success. Copulations were more frequent for males calling inside the lek, indicating that pheromone calling activity and presence in the lek are key factors for copulatory success. A positive association between copulatory success and eye length was found; some characteristics of the face were also associated with copula duration and latency.

Is more choice always desirable? Evidence and arguments from leks, food selection, and environmental enrichment

Recent studies on humans show that too much choice can make subjects less likely to choose any item. I consider general adaptive and non-adaptive explanations of why such choice aversion, or its converse, might occur in animals. There are three questions: is more choice always preferred, does it ever lead to less consumption (or a lower probability of consumption), and may it result in worse items being selected ? A preference for choice is one of the main explanations for lek formation and I draw attention to previously unrecognised parallels with models of human shopping behaviour. There is indeed evidence of female preference for larger leks, although much of the observational data are open to other interpretations. Unfortunately nobody has looked for choice aversion where it is most to be expected, in leks larger than normally occur. Evidence that too much choice of males confuses females is strongest in acoustically advertising frogs, but the widespread decrease of mating skew in larger leks might also have this explanation. A model reanalyses data on skew in black grouse Tetrao tetrix and suggests that considering only a random subset of a large lek may increase the chances of selecting the better males: larger leks are more likely to include better males, but these are less likely to be selected. These opposing effects may lead to an optimum lek size, but only with a sufficient decline in choice accuracy with size. With food choice, very few studies have avoided confounding choice with food quality, by manipulating only flavour. The widespread phenomena of stimulus-specific satiety and novelty seeking imply that monotonous diets are aversive, but no studies test whether animals choose sites where they know food diversity to be greater. Operant experiments that demonstrate mild preferences for free choice concern choice about the means to get food rather than the food itself. In some insect species even moderate choice of diet can be deleterious, and studies on search images and the confusion effect may be evidence of this in vertebrates. Environmental enrichment of captive animals often relies on increasing the options available, but it need not be the choice itself that is beneficial. I consider briefly further areas in biology where choice preference or aversion are potentially important.

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.