Uganda kob prefer high-visibility leks and territories

In lekking species, where males provide estrous females withlittle more than sperm, it has been widely supposed that theonly possible benefits to females of mate choice are genetic.We studied female choice of leks and territories in a reduncineantelope, the Uganda kob (Kobus kob thomasi), and found thatfemales consistently preferred high-visibility mating sites.Leks were elevated and had shorter grass and fewer thicketsthan the surrounding areas. Changes in the number of male andfemale kob on 10 leks were correlated with changes in surroundinggrass height, and both females and males preferred leks withexperimentally reduced grass height over neighboring controls.Within a lek, territory popularity was the primary determinantof male daily mating success, and females preferred territoriesrelatively far from thickets, but removal of thickets did notaffect female territory preferences. Because lion hunting successon kob increases with grass height and thicket density, femalesmay benefit directly from these preferences by reducing therisk of predation.     

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.     

Lek size variation and its consequences in the ochre-bellied flycatcher, Mionectes oleagineus

Several hypotheses suggest that the costs and benefits of displayin aggregations of different sizes play a major role in boththe evolution of leks and in the distribution of males acrossleks of different sizes. We examined the consequences of variationin lek size for both males and females in a study of the ochre-belliedflycatcher, Mionectes oleagineus. We observed 41 solitary displaysites and leks, ranging in size from 1 to 5 mates, over 3 breedingseasons. Although mean visitation rate by females was positivelycorrelated with lek size, female visitation rate per male remainedconstant across lek sizes. The rate at which females visitedthe male who had the highest female visitation rate at eachlek was positively correlated with lek size as predicted bythe hotshot hypothesis. Neither mean nor per capita intrusionrates were correlated with lek size. For the top-ranked male,however, there was a significant correlation between intrusionrates and the size of the lek at which he displayed. Intrusionat leks may be costly, as 28% of female visits were interruptedby intruders. Solitary mates suffered no such interference.Females show no preferences for larger leks, visiting and matingat solitary sites as well as at leks. However, females preferentiallyvisit males with high singing rates, and this male trait maydetermine visitation patterns. Our data argue that preferencesfor larger leks are not important in the evolution of lekkingin this species, nor do they affect lek size. Instead, the dataare in accordance with the predictions of both the hotshot andhot-spot models. These processes may be operating simultaneouslyin this species     

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.     

Female mate choice across spatial scales: influence of lek and male attributes on mating success of blue-crowned manakins

Lekking males compete for females within and among leks, yet female choice is expected to work differently at each of these spatial scales. We used paternity analyses to examine how lek versus male attributes influence mate choice in the blue-crowned manakin Lepidothrix coronata. We tested the hypotheses that females prefer (i) to mate at larger leks where a larger number of potential mates can be assessed, (ii) to mate with unrelated or highly heterozygous males expected to produce high-quality offspring, (iii) to mate with males that display at higher rates, and that (iv) display honestly reflects male genetic quality. Our results show that (i) males at larger leks are not more likely to sire young, although females nesting close to small leks travel further to reach larger leks, (ii) siring males are not less related to females or more heterozygous than expected, (iii) within a lek, high-display males are more likely to sire young, and (iv) both male heterozygosity and display rate increased with lek size, and as a result display does not reliably reflect male genetic quality across leks. We suggest that female mate choice in this species is probably driven by a Fisherian process rather than adaptive genetic benefits.     

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.     

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.     

Great bustard population structure in central Spain: concordant results from genetic analysis and dispersal study.

We found significant sex differences in the mtDNA genetic structure and dispersal patterns of great bustards in a population of 11 breeding groups, "leks", in central Spain. The analysis of genetic distances showed that the female population was divided into three groups of leks separated by ca. 50 km, whereas male haplotypes were randomly distributed among leks. Genetic distances among pairs of leks were positively correlated with geographical distances in females but not in males. While female haplotype distributions were homogeneous among leks at close distances, differences in male genetic structure were highly variable even between two close leks. These results from genetic analyses were concordant with those from a radiotracking study on natal dispersal. Natal dispersal distances were higher in males than in females. Also, the frequency of movement of a female between two leks was positively correlated with their genetic affinity and geographical proximity. In males, the frequency of movement was correlated with geographical proximity but not with genetic affinity. Males dispersed among genetically unrelated leks, contributing to keep nuclear genetic diversity in the population, whereas females tended to be philopatric. These results suggest that isolation-by-distance influences the distribution of maternal lineages at a regional level.     

Mate copying versus preference for actively courting males by female Japanese medaka (Oryzias latipes)

We determined whether female Japanese medaka (Oryzias laiipes)copied the choice of other females or independently preferredactively courting males. We allowed focal females to observeand subsequently choose between a male that courted and potentiallyspawned with a receptive female and a male that courted andwas rejected by an unreceptive female. When the receptive femaledid not spawn, the focal female preferred the male with thehigher courtship rate; whereas when the receptive female spawned,the focal female preferred the spawning male. Our results suggestthat female medaka prefer actively courting males, unless theyhave the opportunity to copy the mate choice of another female.     

Do males choose their mates in the lekking moth Achroia grisella? Influence of female body mass and male reproductive status on male mate choice

Lekking males aggregate to attract females and contribute solely to egg fertilization, without any further parental care. Evolutionary theory therefore predicts them to be nonchoosy toward their mates, because any lost mating opportunities would outweigh the benefits associated with such preferences. Nevertheless, due to time costs, the production of energetically costly sexual displays, and potential sperm limitation, the mating effort of lekking males is often considerable. These factors, combined with the fact that many females of varying quality are likely to visit leks, could favor the evolution of male mate preferences. Here, we show that males of the lekking lesser wax moth, Achroia grisella, were indeed more likely to mate with heavier females in choice experiments, even at their virgin mating (i.e., when their reproductive resources have not yet been depleted by previous matings). This differential female mating success could not be attributed to female behavior as heavy and light females showed similar motivation to mate (i.e., latency to approach the males) and time to copulate. Males seem to benefit from mating with heavier females, as fecundity positively correlated with female mass. This new empirical evidence shows that male mate choice may have been underestimated in lekking species.     

Male cognitive performance declines in the absence of sexual selection

Sexual selection is responsible for the evolution of male ornaments and armaments, but its role in the evolution of cognition—the ability to process, retain and use information—is largely unexplored. Because successful courtship is likely to involve processing information in complex, competitive sexual environments, we hypothesized that sexual selection contributes to the evolution and maintenance of cognitive abilities in males. To test this, we removed mate choice and mate competition from experimental populations of Drosophila melanogaster by enforcing monogamy for over 100 generations. Males evolved under monogamy became less proficient than polygamous control males at relatively complex cognitive tasks. When faced with one receptive and several unreceptive females, polygamous males quickly focused on receptive females, whereas monogamous males continued to direct substantial courtship effort towards unreceptive females. As a result, monogamous males were less successful in this complex setting, despite being as quick to mate as their polygamous counterparts with only one receptive female. This diminished ability to use past information was not limited to the courtship context: monogamous males (but not females) also showed reduced aversive olfactory learning ability. Our results provide direct experimental evidence that the intensity of sexual selection is an important factor in the evolution of male cognitive ability.     

Experimental tests of copying and mate choice in fallow deer (Dama dama)

In fallow deer (Dama dama), as well as in other lek-breedingungulates, receptive females arriving at leks commonly joinmales that are defending large harems. This tendency enhancesdifferences in harem size and mating success between males.It could occur because females independendy move to the samemales, because females are attracted to males with females,or because females are attracted to each other. Using controlledexperiments with estrous female fallow deer, we show that, althoughfemales are more attracted to males with harems than to thosewithout, they are as frequently attracted to groups of femaleswithout a male as to female groups with males. We conclude thatfemale fallow deer joining leks are attracted to each otherand copy each other's movements. As yet, there is no firm evidencein fallow deer or in other lek-breeding ungulates that femalescopy each other's choice of mating partners. Key words: Damadama, fallow deer, lek breeding, mate choice, copying behavior.[Behav Ecol 4: 191–193 (1993)]     

Juggling priorities: female mating tactics in Phayre's leaf monkeys

Extended sexual receptivity in primates is thought to facilitate paternity confusion, thus decreasing the risk of infanticide. However, females might also provide some indication of ovulation to attract preferred males during fertile periods. We examined female mate preferences across defined receptive periods (N = 59) in a group of wild Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) at Phu Khieo Wildlife Sanctuary (February-September 2006; 2,603 contact hours). The group contained seven cycling adult females and three reproductively active males (one adult and two adolescents). We predicted that females would prefer the adult male during periovulatory (POP) receptive periods, but the adolescent males during nonperiovulatory (NPOP) and postconceptive (PC) periods. We collected focal and ad libitum data on sexual and agonistic behaviors to determine female preferences and male awareness of female fertility. We also determined the degree of mating overlap to assess if males were capable of monopolizing females. Our results indicate that females were more frequently proceptive and receptive toward the adult male during POP. By contrast, females were more proceptive and receptive toward one of the adolescent males during PC periods, but rarely interacted with the other adolescent. Patterns of attractivity and agonism across receptive periods suggested that the adult male could detect fertility, while the preferred adolescent could not. Finally, we found a high degree of overlap in total receptive period days, but a low degree of overlap in POP receptive days, suggesting that the adult male might have monopolized females, especially since he seemed to be aware of female fertility. Although these results suggest that females provide some information on ovulation, they also suggest that females attempt to confuse paternity, perhaps capitalizing on male differences in the ability to detect fertility.     

动物求偶场交配制度及其发生机制

求偶场是动物在繁殖期的集群求偶交配的场所, 是动物求偶与繁殖行为的表达空间。为探索求偶场的发生及其进化机制, 研究者们在短短20 年中提出了近10 种关于求偶场发生与进化机制的假说与模型。在鸟类、哺乳类、两栖爬行类和鱼类中已广泛地展开有关求偶场的研究, 但目前国内却还鲜见相关研究报道。本文介绍了求偶场的特征、类型及其进化假说与模型。结合保护生物学理论, 简单阐述了求偶场研究的保护生物学及行为生态学意义。     

Lek formation by female choice: a simulation study

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

Lekking without a paradox in the buff-breasted sandpiper

Females in lek-breeding species appear to copulate with a small subset of the available males. Such strong directional selection is predicted to decrease additive genetic variance in the preferred male traits, yet females continue to mate selectively, thus generating the lek paradox. In a study of buff-breasted sandpipers (Tryngites subruficollis), we combine detailed behavioral observations with paternity analyses using single-locus minisatellite DNA probes to provide the first evidence from a lek-breeding species that the variance in male reproductive success is much lower than expected. In 17 and 30 broods sampled in two consecutive years, a minimum of 20 and 39 males, respectively, sired offspring. This low variance in male reproductive success resulted from effective use of alternative reproductive tactics by males, females mating with solitary males off leks, and multiple mating by females. Thus, the results of this study suggests that sexual selection through female choice is weak in buff-breasted sandpipers. The behavior of other lek-breeding birds is sufficiently similar to that of buff-breasted sandpipers that paternity studies of those species should be conducted to determine whether leks generally are less paradoxical than they appear.     

Male southern water skinks (Eulamprus heatwolei) use both visual and chemical cues to detect female sexual receptivity

A wide range of organisms use chemical and visual cues in mate attraction and courtship; however, chemical discrimination relevant to reproduction and the interplay between these two types of communication are poorly understood in reptiles. We experimentally tested the ability of male Eulamprus heatwolei, a scincoid lizard, to discriminate between sexually receptive and non-receptive females in two ways. First, we conducted 155 staged encounters between males and females over 29 days to determine the start and the duration of the female receptive period based on the date of copulations. These data suggest that the receptive period lasted for approximately 7 days in late October under controlled laboratory conditions. We also recorded 6,330 individual male and female behaviours during these trials to evaluate the frequency of female courtship and rejection behaviours and the intensity of male courtship behaviour. Female courtship increased sharply during the receptive period and then diminished. The disparity between female courtship behaviours and female rejection behaviours was greatest during the receptive period. Male courtship intensity increased sharply prior to the receptive period, peaked during the receptive period and thereafter declined rapidly. To determine if males were using visual cues, chemical cues or both from females, we conducted an experiment during and after the receptive period wherein male lizards were presented with a choice of two retreat sites treated either with the odour of large sexually receptive females, odour of small sexually non-receptive females or no odour (control). Males preferred the scent of females over the odourless control, and analysis using a special form of a generalized linear model, the Bradley–Terry model, showed a clear order of retreat site preferences, with large sexually receptive females favoured over small non-receptive females over the odourless control. We speculate that males use vision to find females and then use their chemosensory ability to chemically evaluate female sexual receptivity once the pair are in close proximity.     

The resident's dilemma: a female choice model for the evolution of alternative mating strategies in lekking male ruffs (Philomachus pugnax)

Previous models for the evolution of alternative male matingbehavior have virtually ignored the role of female choice. Wepresent a model in which female choice favors the evolutionand maintenance of alternative mating strategies in male ruffs,Philomachus pugnax. Resident male ruffe establish and defendcourts on leks against other residents, while non-territorialsatellite males move between leks and among courts on a lek.Residents appear to actively recruit satellites to their courts,even though satellites may mate with females once there. Residentbehavior toward satellites and data on female behavior suggestthat residents benefit from a satellite's presence due to somefemale preference for mating on co-occupied courts. However,if all residents accept satellites, none gains any relativeadvantage, yet all pay the costs of having satellites on theircourt. We present a game theoretical model that shows that therelative nature of female choice places residents in an evolutionarydilemma with respect to satellite acceptance. Although all residentswould benefit if satellites could be cooperatively excludedfrom leks, the only evolutionarily stable strategy for individualresidents is to defect and accept satellites. The model alsodemonstrates that this "resident's dilemma" likely exists onlyin a local sense, since the failure of residents to cooperativelyexclude satellites from leks need not result in globally lowerpayoffs, due to frequency-dependent selection on the proportionof satellites in the population. Our analysis suggests thatthe resident-satellite relationship in ruffs, despite its obviouscompetitive elements, is fundamentally a cooperative associationfavored by female choice. Female choice has also been proposedas the primary mechanism selecting for male association to formleks in ruffe. In this context, resident-satellite associationsmay be thought of as transitory "leks within a lek     

Competition for landmark territories among male Polistes canadensis (L.) (Hymenoptera: Vespidae): large-size advantage and alternative mate-acquisition tactics

Alternative male mating tactics of insects at landmarks (leks)have only rarely been investigated. Some males of the paperwasp, Polistes canadensis (L.), were territorial at small treesalong the crests of dry ridges in Santa Rosa National Park,Costa Rica. Territories did not contain nests or resources forwhich females foraged. Contrary to other "hilltopping" species,male P. canadensis competed most intensely for territories insaddles along these ridges rather than at the highest points.Nonterritorial males patrolled small areas of the ridge line,following a path that took them to a number of territories.Many males switched between territoriality and patrolling, suggestingthat both size-related tactics belong to one conditional strategy.Males that were territorial on 2 or more days were larger thanthose that were territorial on only 1 day, and these in turnwere larger than permanent patrollers. Moreover, the mean sizeof territorial males was positively correlated with two measuresof territory attractiveness, suggesting that larger males monopolizepreferred sites. Mean age of territorial males was also relatedto territory attractiveness, but males of intermediate age claimedthe most attractive territories.     

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.