Physiological costs enforce the honesty of lek display in the black grouse (Tetrao tetrix)

Females may use condition-dependent sexual traits as reliable cues of male “quality” if the costs of the expression of such traits vary with male “quality”, and if there is positive genetic correlation between male traits and condition. However, there are multiple ways of measuring the changes in body condition which reflect physiological costs meaning that the multifaceted nature of the physiological costs associated with the expression of sexual traits has rarely been thoroughly examined. In the lekking black grouse (Tetrao tetrix), mating success is highly skewed towards males defending central territories and having high survival rates to the following year, but the mechanisms underpinning such superior performance remain unclear. In this study, we quantified the changes in five measures of body condition before and after the mating season and related these changes to male lek performance (fighting rate, territory centrality and mating success) to understand the physiological costs of male reproductive effort. Between the two capture sessions, male body mass decreased significantly, blood parasite counts and plasma carotenoid concentration increased substantially while the total immunoglobulin concentration tended to increase. There was no overall impairment of individual body condition as the changes in the five measures of body condition were unrelated. Male fighting rate was unrelated to changes in the condition measures but males losing more body mass defended central territories and had high mating success. Therefore, females preferring central, dominant males may select males better able to afford the energetic costs of lek performance thereby effectively enforcing the honesty of male display.     

No peace for estrous topi cows on leks

Male coercion, such as harassment, may be considered the thirdmain component in sexual selection alongside male competitionand female choice. In this study on lek-breeding topi antelopes(Damaliscus lunatus), I investigate whether female mating preferenceshave consequences for male investment in harassment and whetherharassing males are more likely to succeed in mating. I thenaddress the question of whether lek evolution in topi can beexplained by harassment avoidance. Judging from mating rate,I found that female topi antelopes in estrus preferred lek malesto resource defenders. In contrast to lek males, resource defendersdemonstrated significantly higher harassment rates before theysucceeded in mating than when they did not, and the precopulatoryharassment rate was significantly higher on resource territoriesthan on lek territories. After mating on resource territories,harassment dropped to low levels. Thus, resource defenders,but not lek males, seem to employ harassment as a strategy tocoerce females to mate against their preference. However, byusing various measures of harassment intensity, overall estrousfemales were found to experience higher harassment levels onlek, and chases by intruders were relatively rare on all territorytypes. These findings suggest that harassment avoidance is unlikelyas an explanation for lek evolution.     

Queuing in space and time reduces the lek paradox on an antelope lek

Lek systems, where females often use centrality to assess male quality, highlight a general paradox in evolutionary biology: how can female preferences for males providing good genes persist when consequential strong directional selection is predicted to deplete additive genetic variance in male quality and thereby obliterate benefits of choosiness? An explanation contributing to the resolution of this lek paradox may be that genetic variance is retained when an indirect mate choice cue, such as centrality on a lek, is an imperfect indicator of male genetic quality. Here I investigate whether the presence of alternative male mating tactics limits the reliability of centrality as a quality indicator in lek-breeding topi antelopes. Whereas males establishing territories directly on the central lek were relatively large, smaller peripheral males regularly shifted their territories centripetally and in this way also occasionally obtained central territories. By such opportunistic queuing, small males could increase their mating success drastically; however, their territorial tenure in the lek centre was relatively short, consistent with moderate competitive ability. These results suggest that male topi antelopes can obtain central lek territories through alternative mating tactics, providing scope for variance in male quality on the central lek. In a separate finding, the mating success of central males was found to increase during territorial tenure, independent of estimated age. The demonstration of queuing in both space and time on a mammalian lek highlights the importance of considering male tactical dynamics over time in order to avoid an inflated appearance of the lek paradox.     

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     

Resource defense polygyny by Hetaerina rosea Selys (Odonata: Calopterygidae): influence of age and wing pigmentation

Current evidence suggests that in Hetaerina damselflies males exhibit lek mating system. In this study, in order to answer if the same occurs in Hetaerina rosea Selys, we manipulated vegetation substrates used as territories and quantified the number of visiting females, males defending territories and fight intensity. We also examined whether body size and wing pigmentation are selectable traits in male-male competition, and if age affects male territorial behavior. Our results showed that males with larger pigmented areas won more contests, independently of body size. Old males changed from territoriality to sneaking strategy. Contrary to other Hetaerina species, males of H. rosea do not display lek behavior, but defend resources according to the resource defense polygyny strategy.     

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.     

Male dimorphism, territoriality and mating success in the tropical damselfly, Paraphlebia zoe Selys (Odonata: Megapodagrionidae)

The tropical damselfly Paraphlebia zoe has two male morphs: a black-winged (BW) male which is associated with territorial defense of oviposition sites; and a hyaline-winged (HW) male similar in appearance to females, and, compared to the black morph, less frequently found defending territories. In a wild population of this species, we first assessed the relationship between phenotypic traits [male morph, size and territorial status (being territorial or non-territorial)], their role on mating success, and the degree to which a particular territory may contribute to male mating success. Second, to relate a physiological basis of being territorial we compared both morphs in terms of muscular fat reserves and thoracic muscle, two key traits related to territory defense ability. Males of both morphs defended territories although the BW males were more commonly found doing this. BW males were larger than HW males and size predicted being territorial but only within HW males (territorial males were larger) but not in BW males. Male mating success was related to territorial status (territorial males achieved a higher mating success), but not to morph or size. Furthermore, territory identity also explained mating success with some territories producing more matings than others. The BW morph stored more fat reserves which may explain why this morph was more likely to secure and defend a place than the HW morph. However, the HW morph showed higher relative muscle mass which we have interpreted as a flexible strategy to enable males to defend a territory. These results are distant to what has been found in another male dimorphic damselfly, Mnais pruinosa, where the advantage of the non-territorial morph relies on its longevity to compensate in mating benefits compared to the territorial morph.     

Resource defence on exploded leks: do male little bustards, T. tetrax, control resources for females?

In lekking species, female choice of particular males for mating is based on male characters that are not related to immediate gains such as access to resources. This is, however, unclear in exploded-lekking species, as male territories may contain resources for females; it is thus the degree to which males regulate female access to these resources to obtain matings that should distinguish lekking from nonlekking species. We investigated the relationship between resources for females, male distribution and male attractiveness in little bustards. One parameter determining resource quality for females was invertebrate availability: females laid eggs in grassland fields (grass, alfalfa and ryegrass) with significantly higher invertebrate biomass than others of similar cover but nests were in areas that did not have higher invertebrate biomass, at a scale of 1 km². Males set up territories and females laid eggs in areas that had a high proportion of alfalfa, but not high mean invertebrate biomass. Thus males set up territories at resources potentially used by females, but that appeared not to be critical for breeding. In addition, females did not use the amount of resources defended as a criterion for mate choice, as male attractiveness was not related to various territorial resources. Locating the lek in areas attractive to females might be a way of increasing the probability of encounter between the sexes. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

When should males lek? Insights from a dynamic state variable model

A central focus in the study of lek evolution is to understandthe clustering of male mating territories. Lekking males typicallydefend small clumped territories and experience intense competitionassociated with dense aggregations. We used dynamic state variablemodeling to evaluate three alternative selective pressures proposedto explain the evolution of lekking. These are female matingbias for large clusters, reduction in predation risk in largeclusters, and male harassment of estrous females. We modeledmale mating decisions during a single breeding season usinga lekking ungulate as a model system. Males could choose fromeight alternative tactics that included a nonreproductive tactic,territorial tactics ranging from low to high clustering, andthe option to join a mixed-sex herd. The model predicted a state-and time-dependent strategy that maximizes mating success overthe course of the season. We then simulated a population of100 males that used the optimal strategy and calculated theproportion of the population that adopted each tactic. Our modelgenerated unique predictions for the three selective pressureswe considered. Female mating bias, when nonlinearly relatedto cluster size, had the greatest potential to generate largeclusters of territorial males, whereas predation risk and harassmentof females typically did not promote male clustering. More generally,our model highlights the conditions that will favor lekking.Lek-like clustering was consistently produced when the benefitsin clustering increased in specific nonlinear ways. Our modelthus emphasizes clarifying the shapes of relationships betweenpotential selective factors and the size of territory clusters.     

Non-random mating in classical lekking grouse species: seasonal and diurnal trends

This paper is the first to integrate both field and theoretical approaches to demonstrate that fertility benefits can be a direct benefit to females mating on the classical lek. Field data collected for male sharp-tailed grouse (Tympanuchus phasianellus), a classical lekking species, revealed potential fertility benefits for selective females. Adult males and individuals occupying centrally located territories on the lek were found to have significantly larger testes than juveniles and peripheral individuals. Further, using empirical data from previously published studies of classical lekking grouse species, time-series analysis was employed to illustrate that female mating patterns, seasonal and daily, were non-random. We are the first to show that these patterns coincide with times when male fertility is at its peak. Received: 26 February 1999 / Revised: 13 December 1999 / Accepted: 15 March 2000     

Indirect mate choice, direct mate choice and species recognition in a bower-building cichlid fish lek

Sexual selection arising through female mate choice typically favours males with larger, brighter and louder signals. A critical challenge in sexual selection research is to determine the degree to which this pattern results from direct mate choice, where females select individual males based on variation in signalling traits, or indirect mate choice, where male competition governs access to reproductively active females. We investigated female mate choice in a lekking Lake Malawi cichlid fish, Hemitilapia oxyrhynchus, in which males build and aggressively defend sand 'bowers'. Similar to previous studies, we found that male reproductive success was positively associated with bower height and centrality on the lek. However, this pattern resulted from males holding these territories encountering more females, and thus their greater success was due to indirect mate choice. Following initial male courtship, an increase in the relative mating success of some males was observed, but this relative increase was unrelated to bower size or position. Crucially, experimentally manipulating bowers to resemble those of a co-occurring species had no appreciable effect on direct choice by females or male spawning success. Together, these results suggest indirect mate choice is the dominant force determining male-mating success in this species, and that bowers are not signals used in direct mate choice by females. We propose that, in this species, bowers have a primary function in intraspecific male competition, with the most competitive males maintaining larger and more central bowers that are favoured by sexual selection due to higher female encounter rates.     

Are Centrally Displaying Males Always the Centre of Female Attention? Acoustic Display Position and Female Choice in a Lek Mating Subterranean Insect

Several conceptual models seek to explain patterns of male display and factors that influence female mate choice in lek mating systems. The central advantage model predicts that males displaying at or near the lek centre should be more attractive to females than are males positioned along the lek periphery. Females may exhibit biases toward these centrally displaying males based on either spatial or display‐related cues. We tested the prediction of the central advantage model in investigating the importance of male display position in the subterranean and lek mating prairie mole cricket (Gryllotalpa major). Gryllotalpa major males form mating aggregations in the early spring and produce an acoustic advertisement signal from a constructed calling chamber at the soil surface. Pair formation occurs in the calling chamber, and males typically maintain these structures for the duration of the reproductive season. To assess whether G. major females exhibit a preference for males calling from centrally located acoustic burrows, we documented the spatial position and number of female attractions for all advertising males across the focal lek. Six spatial attributes related to display position were reduced using principal component analysis and examined for an association with male attractiveness. We found that in general, female attractions were distributed randomly across the lek; male attractiveness was not related to proximity to the lek centre nor to any factor associated with display position. The most highly attractive males, however, were located further from the lek centre and from nearest calling neighbours than other attractive males. Advertising males that segregate themselves within the aggregation and locate nearer the lek margin may gain a geometric advantage resulting in the increased probability of attracting a searching female.     

Queuing for territory positions in the lekking black grouse (Tetrao tetrix)

The importance of a central territory position as a determinantof male mating success in lekking species has been debated.The process by which a male can obtain a central territory hasbeen especially lacking a quantitative analysis. We presenta stochastic queuing model describing territory succession towardthe lek center and apply it to a 8-year sequence of territorycentrality measures on a black grouse Tetrao tetrix lek. Thelek shows a value of intermediate queue discipline, which deviatessignificantly both from strict orderly queuing and from randomranking of males. Thus, high-ranking males are partly able tomaintain their superior position over years, but queue-jumpingis not excluded; especially because highly succesful males donot attempt to change their territory position toward the lekcenter. As a result of stochastic queuing, a central territoryindicates an older than average male, as well as a male witha history of high fighting rates. These results are consistentwith the hypotheses that territory position is an honest signalof male quality and that the long-term fighting effort and survivalrequired to acquire a central territory may increase the reliabilityof the signal over that of short-term display effort. The impartialqueue discipline, however, also leaves room for other male characteristicsto play a role in determining individual mating success.     

Non-defendable resources affect peafowl lek organization: a male removal experiment

A lekking mating system is typically thought to be non-resource based with male providing nothing to females but genes. However, males are thought to clump their display sites on areas where they are more likely to encounter females, which may depend on non-defendable resource location. We tested this hypothesis on a feral population of peacocks. In agreement, we found that, within the lek, display site proximity to food resources had an effect on female visitation rate and male mating success. The attractiveness of display sites to male intruders was explained by the distance to the feeding place and by the female visitation rate. We randomly removed 29 territorial males from their display sites. Display sites that were more attractive to male intruders before removal remained highly attractive after removal and display sites closer to the feeding area attracted the attention of intruders significantly more after removal. Similarly, display sites that were more visited by females before removal remained more visited after removal, suggesting again that the likelihood of encountering females is determined by the display site location. Overall, these results are in agreement with non-defendable resources affecting lek spatial organization in the peafowl.     

The social organization of feral peafowl

Peafowl are usually reported to have a mating system based on harem defence by adult males. In a small feral population near Oxford, males defended small (<1 ha) territories while females remained in one flock that ignored male territory boundaries. After mating, females become solitary. At no time did a female associate selectively with one male or remain within his territory, nor did males attempt to follow or guard female groups. Two out of four males were seen to mate. These differed from the other two in being neither very old nor very young; they held territories smaller than that of the young male and were no larger or longer-tailed. However, they spent more time displaying. We suggest that peafowl have a mating system similar to a lek: males defending small, clumped territories visited by females for mating.     

Black holes, mate retention, and the evolution of ungulate leks

In some ungulates living in unstable herds, females in estrusleave their usual groups and join males defending mating territories.During the 12-24 h before mating, females commonly move severaltimes between males. These movements are commonly caused byharems being disrupted by young males or by overenthusiasticcourtship by the territorial male. When females leave a territorythey typically move to its nearest neighbor. Under these circumstances,clusters of territories can retain estrous females until theymate and leave the lek. This paper develops a model to investigatethe benefits of defending clustered versus dispersed territoriesto males and the consequences of variation in the rate of femalemovement between territories (P_m) and the tendency for femalesto move from one territory to a neighboring territory (a). Wherefemales move between territories at least once every 24 h (P_m< 0.04) and usually move to neighboring territories (a <0.5), the mating success of males is inversely related to thedistance from their territory to its nearest neighbor, and malesdefending clustered territories have higher mating rates thanthose defending dispersed territories. This process may be importantin the initial evolution of ungulate leks, which may resembleblack holes, attracting and retaining estrous females untilthey mate and their estrus ceases. It provides one possibleexplanation of the evolution of ungulate leks that does notrely on female preferences for mating with particular phenotypiccategories of males.     

South American sea lions in Peru have a lek‐like mating system

Five years of behavioral observations revealed significant effects of high air temperatures and breeding site topography on the mating system of South American sea lions in Peru. Unlike most polygynous mammals that defend females or fixed territories, male sea lions in Peru maintained positions along the shoreline where females passed each day to thermoregulate, and where most copulations occurred. Sex ratios (1 male per 17 females) and male mating success were extremely skewed (14% of males achieved 50% of the copulations, and 25% of them did not copulate at all). The mass daily movements of females toward the water and cool substrate of the shoreline, along with a highly skewed sex ratio, accentuated the difficulty for males to monopolize and restrict female movements. Females moved freely and chose their mates, unlike in temperate regions of their range where male South American sea lions control groups of females or access to tide pools. Our observations indicate that the South American sea lion in Peru has a lek‐like breeding system. This is a rare alternative to the common male strategies of defending females and resources, and is likely an evolutionary product of their highly skewed sex ratio, protracted breeding season, and the extreme subtropical climate where they breed.     

Evolution of sexual size dimorphism in grouse and allies (Aves: Phasianidae) in relation to mating competition,fecundity demands and resource division

Sexual size dimorphism (SSD) is often assumed to be driven by three major selective processes: (1) sexual selection influencing male size and thus mating success, (2) fecundity selection acting on females and (3) inter‐sexual resource division favouring different size in males and females to reduce competition for resources. Sexual selection should be particularly strong in species that exhibit lek polygyny, since male mating success is highly skewed in such species. We investigated whether these three selective processes are related to SSD evolution in grouse and allies (Phasianidae). Male‐biased SSD increased with body size (Rensch’s rule) and lekking species exhibited more male‐biased SSD than nonlekking ones. Directional phylogenetic analyses indicated that lekking evolved before SSD, but conclusions were highly dependent on the body size traits and chosen model values. There was no relationship between SSD and male display agility, nor did resource division influence SSD. Although clutch mass increased with female body size it was not related to the degree of SSD. Taken together, the results are most consistent with the hypothesis that lekking behaviour led to the evolution of male‐biased SSD in Phasianidae.     

Absence of male reproductive skew,along with high frequency of polyandry and conspecific brood parasitism in the lekking Houbara bustard Chlamydotis undulata undulata

Behavioural studies have led to the perception that lekking species experience a high male reproductive skew as a consequence of females’ selective mate choice. In addition, observations suggest that females copulate only once and therefore polyandry seems unlikely as females are supposed to choose the best male available. In order to analyse the mating strategy of the Houbara bustard, an endangered lekking species under reinforcement in eastern Morocco, we used microsatellite data to perform paternity analyses. None of our observations followed common expectations under a lek mating system: we found no male reproductive skew suggesting no apparent selective female mate choice and no apparent male benefit from lekking. In contrast, a high level of polyandry (60 % of the nests) was recorded suggesting that sperm competition may operate. In addition, we present another case of conspecific brood parasitism in a lekking species and this was an unexpected alternative strategy for a species presenting high parental cost and low fecundity. The increasing number of studies contradicting common assumptions on lekking species suggests that alternative breeding strategies such as males pursuing an off‐lek mating strategy, female polyandry and even conspecific brood parasitism might be more widespread in lekking species than previously thought.     

Methoprene modulates the effect of diet on male melon fly,Bactrocera cucurbitae,performance at mating aggregations

The effect of access to dietary protein (P) (hydrolyzed yeast) and/or treatment with a juvenile hormone analogue, methoprene (M), (in addition to sugar and water) on male aggregation (lekking) behaviour and mating success was studied in a laboratory strain of the melon fly, Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae). Six‐day‐old males were treated with (1) protein and methoprene (M+P+), (2) only protein (M?P+), or (3) only methoprene (M+P?), and compared with 14‐day‐old sexually mature untreated males (M?P?). The lekking behaviour of the four groups of males when competing for virgin sexually mature females (14 –16 days old) was observed in field cages. The following parameters were measured at male aggregations: lek initiation, lek participation, males calling, male–male interaction, female acceptance index, and mating success. For all these parameters, the M+P+ males significantly outperformed the other males. Moreover, for all parameters, there was a similar trend with M+P+ > M?P+ > M?P? > M+P?. More M+P+ males called and initiated and participated in lek activities than all other types of male, which resulted in higher mating success. They had also fewer unsuccessful copulation attempts than their counterparts. Whereas treatment with methoprene alone had a negative effect in young males with only access to sugar, access to dietary protein alone significantly improved young male sexual performance; moreover, the provision of methoprene together with protein had a synergistic effect, improving further male performance at leks. The results are of great relevance for enhancing the application of the sterile insect technique (SIT) against this pest species. The fact that access to dietary protein and treatment of sterile males with methoprene improves mating success means that SIT cost‐effectiveness is increased, as more released males survive to sexual maturity.