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Kyle A. Young, Martin J. Genner, Domino A. Joyce, Marcel P. In many animals, males congregate in leks that females visit for the sole purpose of mating. We observed male and female behavior on 3 different-sized leks of the bower-building cichlid fish Nyassachromis cf. 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 Saint-Marcel ga sexy girl 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 Saint-Marcel ga sexy girl hotshot males preferred by females. Average female encounter rate of males increased with lek size, a pattern typically interpreted as evidence that leks form through female preference for lekking males, rather than because males congregate in hot spots of high female density.

Female propensity to engage in premating behavior once courted did not increase with lek size, suggesting female 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 female preference model, where females may benefit through indirect mate choice for males able to meet the competitive cost of occupying larger leks. Research on lek systems has focused on 2 related issues: reconciling the relationship between female mate choice, male mating skew, and the maintenance of genetic variance, that is, the lek paradox Reynolds and Gross ; Kirkpatrick and Ryan ; Mackenzie et al.

One successful approach for assessing such models is to test their mechanistic predictions by comparing male and female behavior within and between leks of varying size Shelly; Alatalo et al. The female preference model was formalized by Bradbury and posits that males aggregate on leks because females prefer to mate with clustered males. Males form leks when the costs associated with clustering e. The hotshot model Beehler and Foster invokes female preference for particular males rather than clustered males in general.

The hot-spot model differs in that it does not invoke female preference for clustered males in general or dominant males in particular but instead proposes that males form leks in hot spots with high concentrations of females Bradbury and Gibson Defining mate choice as any behavior that reduces a female's pool of potential mates, direct mate choice, the focus of most sexual selection research, involves direct assessment and discrimination among males based on phenotypic traits such as feather length, call volume, or color.

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By contrast, indirect female choice from behaviors that reduce the set of potential mates without discrimination among individual males by the female. Such mate choice can occur if females prefer to mate in a certain location or at a certain time, and the outcome of indirect female choice is typically governed by male—male competition. The distinguishing predictions of the hot-spot model arise because it does not invoke female preference, through indirect or direct mate choice, for males in different-sized leks or for different males within leks. If males have accurate information about the distribution of females and the cost of occupying different-sized leks is constant, the distribution of males should match that of females Bradbury and Gibson As a result, the average female encounter rate of males should not vary with lek size.

Furthermore, because direct female choice should not vary across leks, the willingness of females to mate once encountered should not vary with lek size. The hot-spot model does not make explicit predictions about variation in male mating success within leks. If female preference for larger leks from indirect choice, the model predicts that males in larger leks should enjoy a reproductive advantage because females prefer to visit larger leks.

If females prefer males on larger leks through direct choice, the model predicts that females, once they have visited a lek, will be more likely to mate with males on larger leks. Average male mating success can increase with lek size through either mechanism, and the relative importance of each may vary between and within species. Like the hot-spot model, the female preference model makes no explicit predictions about variation in male mating success within leks.

The hotshot model explicitly predicts how male mating success should vary within leks. Females may prefer to visit such males through indirect choice if, for example, more central males happen to encounter a greater of receptive females. Alternatively, females may prefer to mate with such males through direct choice after assessment of a range of males across the lek. The second prediction, that the most successful males should occupy the largest leks, can be met through either form of mate choice; the hotter the shot the more females and males it should attract and the larger lek it should support Beehler Saint-Marcel ga sexy girl Foster Here we test these predictions using male and female behavior from 3 different-sized leks of Nyassachromis cf.

Three features of the system make it well suited for a comparative study of lek formation models. Second, unlike many systems, males do not actively search for females but instead reliably court those swimming past their territory. As a result, within and between lek variation in male courtship rate is primarily a consequence of indirect female choice.

Finally, after male courtship initiation, a predictable series of discrete interactions takes place. During this series, females decide whether to proceed to the next step of the series, and each of these decisions involves direct female choice based on assessment of individual Saint-Marcel ga sexy girl and their territories.

We studied 3 N. The species is a small maximum total length 10—13 cm planktivore that breeds in near-shore sand habitats between 3 and 20 m depth Konings Males defend territories centered on volcano-shaped bowers from which they court passing females. Prior research on lekking cichlids suggests female encounter rate and mating success increases with territory quality bower size and position centrality McKaye et al. Only recently, however, have there been attempts to distinguish between the roles of indirect and direct mate choice in bower-building cichlids Stauffer et al.

Characteristics of the 3 study leks. s in parentheses are one standard deviation of the mean. The study site: a Lake Malawi, b the 3 study leks in Lake Malawi National Park, and c the distribution of bowers within each lek. Closed circles represent locations of bowers from which observational data were collected. See Table 1 for a description of the leks.

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Prior to collecting behavioral data, we conducted preliminary surveys of the size and shape of each lek and used rope to establish 2 orthogonal axes for orientation and subsequent mapping Figure 1. We collected behavioral data during the daily period of peak breeding activity, between and Prior to formal observations, observers calibrated behavioral classifications by watching the same individual focal males.

During dives, each observer selected males haphazardly while moving inward or outward along an axis to ensure that we observed males from throughout the lek. Each male was observed once for a single 10 min period Kellogg et al. During each observation period, we recorded the of times the focal male engaged in the following behaviors: initiation of courtship with a passing female, aggression toward conspecific male, aggression toward a heterospecific male, bower-building activity picking up and moving sand by mouthand foraging in the Saint-Marcel ga sexy girl column.

Courtship follows a predictable sequence that females can terminate at any stage. After courtship initiation by a male, females follow the male to the bower, visit the bower platform, engage in circling bouts with male on the bower platform, and finally lay one or more eggs that are collected in the mouth of the female as they are fertilized by the male. We make 2 assumptions in our analyses and interpretation of the behavioral data. The first is that the of male courtship initiation events accurately reflects female visitation rate.

This requires that males reliably court females passing their territory. We confirmed this by following 21 females as they entered and traveled through one of the leks Thumbi West.

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The females passed males, of which courted. Because egg laying is rarely observed during focal male observations we observed only 4 spawning events in This assumption has been convincingly confirmed in studies of Lake Malawi bower-building cichlids Kellogg et al. After behavioral observations, we recorded the coordinates of every bower. Variation in bower size and shape within each lek 3 independent analyses was summarized using principal components analysis on the covariance matrix of the 12 measurements Supplementary material.

For each lek, we constructed a Spearman rank correlation matrix of the relationships between bower position, bower size, the 5 male behaviors, and female follows, enters and circles Supplementary material. Because testing the predictions of the models requires considering the absolute rather than relative of behaviors, we analyzed variation in the frequency of each male behavior per minute independently. To distinguish between the main predictions of the hot-spot and female preference models, we first tested whether female encounter rate increased with lek size using single factor analysis of variance ANOVA.

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To determine if an increase in female encounter rate with lek size came at a cost, we used the same model to test whether the cost of territory defense and maintenance increased, while feeding rate decreased, with lek size. The first ANOVA model above distinguishes between the principal prediction of the hot-spot model and the first prediction of the female preference model that Saint-Marcel ga sexy girl capita male mating success increases through indirect choice because females prefer to visit larger leks.

To investigate the second prediction, that courted females prefer through direct choice to mate with males on larger leks, we tested for lek effects using analyses of covariance ANCOVAs with follows, enters, and circles as response variables and courtship as a covariate.

For these models, we use only males that courted at least one female. If courted females prefer to mate with males on larger leks through direct choice, we expect ificant lek effects after controlling for variation in courtship rate among males. At each stage of the behavioral sequence, a proportion of females abandon the male. As a result, uninformative zeros accumulate in the response variables of these models e. We thus repeated the ANCOVAs on enters and circles using the preceding behavior as the covariate and including only those males that engaged in at least one such behavior.

These models have smaller sample sizes but provide more specific tests of whether direct female mate choice during different stages of the mating sequence varies with lek size. Before testing the specific predictions of the hotshot model, we first tested the possibility that females prefer to visit males Saint-Marcel ga sexy girl on aggressiveness, territory location, and quality, for each lek individually using multiple regression on courtship rate with conspecific attack rate, bower location, and bower size as predictor variables.

To test the prediction that courted females prefer hotshot males through direct choice, we repeated the analyses using only males that courted at least once with follows, enters, and circles as response variables and courtship as an additional predictor variable. As above, we repeated these analyses with the behavior as the covariate using only those males with at least one such behavior this sequence of models loses power rapidly for the Domwe lek; Table 1. We tested the second prediction of the hotshot model, that the most successful males should be on the largest leks, by selecting from each lek the 5 males with the most follows, enters, and circles these were generally, but not always, the same males.

For each of these increasingly accurate predictors of mating success, we used Kruskal—Wallis K-W non-parametric ANOVA to test whether estimates of mating success increased with lek size. We begin by describing the accumulation of male mating skew from courtship through circling on the 3 leks Figure 2. Ethogram summarizing accumulation of mating skew through the courtship sequence on the 3 study leks.

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For each behavior, we present the total of behaviors observed and in parentheses the percentage of males ing for all such behaviors. Bold arrows between behaviors indicate stages of direct female choice, with the percentage of positive responses shown. The bottom line shows the cumulative effect of post-courtship direct female choice on the accumulation of mating skew in each lek. Though females prefer males on larger leks through indirect choice, contrary to the second prediction of the female preference model, courted females showed no tendency to prefer males on larger leks through direct mate choice.

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Sequential ANCOVAs using the preceding behavior instead of courtship rate as the controlling covariate confirmed these .

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