Size and stability of vertebrate leks

Lek size varies greatly among lekking species. At present there is no explicit theoretical explanation for this diversity. We extend an existing model of optimal lek size that incorporates female mating preferences and male-male contest competition. The model shows that variation in lek size is predicted by the interaction between lek size, overall copulation rate and the proportion of copulations accruing to males of different rank. In species where females prefer to mate on the largest leks and high-ranking males are able to monopolize females irrespective of the size of the lek, the maximum lek size will be large. Conversely, in species where females show weak preference for mating on large leks or increasing lek size quickly results in scramble competition, the maximum lek size will be smaller. Thus, differences between species in lek size may be due largely to differences in the extent to which high-ranking males can monopolize mating opportunities. Leks become unstable and break down when high-ranking males can no longer get their 'expected' copulation success. Therefore, the mechanism that generates male clustering, that is, sexual parasitism of high-ranking males by subordinates, also sets a limit to the largest stable lek size. Copyright 1999 The Association for the Study of Animal Behaviour.     

How predation risk affects the temporal dynamics of avian leks: greater sage grouse versus golden eagles

Leks often attract predators as well as mates, yet most evolutionary models have assumed that sexual selection, not predation, drives lekking behavior. We explored the influence of predation on lek dynamics using a stochastic dynamic game model based on the lek-breeding greater sage grouse (Centrocercus urophasianus) and its principal avian predator, the golden eagle (Aquila chrysaetos). The model predicts time-dependent male lek attendance as a function of factors affecting both mating success (female arrival rate, male numbers, and social status) and predation risk (eagle arrival rate and group size). Dominant males are predicted to arrive sooner and leave later than subordinates, especially if mating skew is high, predation risk is low, or the relationship between lek size and female arrival rate is weak. Both high mean levels of predation risk and small lek size should reduce lek attendance, but the relative tendency of predators to attack large versus small leks has little influence on predicted lekking behavior. Field observations confirmed the predicted effects of female arrival rate, lek size, male dominance, and weather-dependent predator arrival rates on lek departure times. Predicted effects of female arrival rates and male dominance on seasonal lek attendance were also supported. Our model provides an empirically supported adaptive explanation for short-term lek dynamics. It also suggests alternative interpretations for phenomena previously invoked to support the hotshot and skew models of lek formation.     

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     

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.     

Non-linear ascension in a reproductive hierarchy of the Blue Manakin (Chiroxiphia caudata)

Blue Manakin Chiroxiphia caudata (Pipridae) has a lek mating system in which males cooperate in teams of two or more individuals for courtship displays to females. Males ascend through a dominance hierarchy to assume the rank of alpha male. Only alpha males copulate with females. In this study, we describe the ascension of a male to the alpha rank following the disappearance of the previous alpha male. The study was conducted in the seasonal Atlantic Forest of Southern Brazil. We observed nine banded males at two adjacent leks. The gamma male at the more successful lek also acted as an alpha male at a less successful lek 314 m away. Upon the disappearance of the alpha male at the more successful lek, the gamma gave up his alpha rank at the less successful lek and ascended to the alpha rank at the more successful lek, bypassing the beta male. Although a previous study suggests a strictly linear dominance hierarchy in C. caudata, with beta male ascending to alpha status, our results suggest that the rising to become the alpha male may be a more flexible and dynamic process than previously reported.     

The anatomy of population change in a black grouse population 1992–2008

We interrogate an 18-year-long dataset containing counts of displaying male black grouse Tetrao tetrix and incidental counts of females within an 800-km² region of Perthshire, Scotland. We examine the trends in the population and investigate how different components of the population might act as signposts of different stages of overall population change. We found statistical evidence for a decline in black grouse numbers between 1992 and 2000, and then a recovery from 2002 to 2008, but little evidence for a link between population change and weather during the decline phase. There was some evidence for a positive relationship between male and female counts. The two main components of male population size, lek size and lek frequency followed the overall population trend while it was increasing, but during the earlier decline, the two became uncoupled, to expose a complex structure within the data. During the decline, when black grouse numbers were approaching their minimum, mean lek size was actually increasing. Small leks lost proportionally more birds than did large leks, and lek longevity was positively correlated with lek size, indicating that maintenance of large leks is crucial in buffering the population against serious declines. During the decline, the spatial arrangement of leks changed, with remnant leks showing tight clustering at larger spatial scales, before expanding out to fill the large areas of unoccupied landscape during the population increase. We discuss these findings in terms of species monitoring and suggest that counts of young males may add much useful demographic information with little extra effort.     

Lek Site Selection by Male Mediterranean Fruit Flies

The mating system of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), is based on leks. In field cage choice experiments we examined lek site selection as influenced by the presence of fruit and the presence of conspecific flies. Males were allowed to choose between artificial trees containing fruit and trees without fruit. Males showed a significant preference for the tree baited with fruit. To determine which fruit-related stimuli were important, males were allowed to choose between visual and/or olfactory fruit-derived stimuli. Males preferred to lek on trees presenting both stimuli. To determine the influence of conspecifics on lek site selection, males were allowed to choose between trees containing male and/or female conspecific flies. The presence of pheromone-emitting males was preferred. In addition, both in the field and on artificial trees, we examined the effect of leaf size on lek site selection. In the field, males preferred to lek on leaves of a particular size. Moreover, leaf integrity was important, as males tended to select undamaged leaves as lek sites. In field cage experiments, males consistently chose to lek on trees containing the largest leaves. We conclude that the choice of lek site is influenced by the presence of fruit and of other lekking males, while leaf size and integrity determine male location within a lek.     

Reproductive Competition Among Males in Multimale Groups of Primates: Modeling the Costs and Effectiveness of Conflict

Multimale groups of primates are characterized by strong reproductive competition among males, generally resulting in an uneven division of male reproductive success (reproductive skew). The observed patterns of conflict and reproductive skew have often been attributed to the so-called tug-of-war model. We show, however, that two important assumptions of this model are not met in male primates. First, the tug-of-war model assumes that reproductive conflict reduces overall group productivity, but in male primates (and most other vertebrates) conflict likely involves mortality rather than fecundity costs. Second, the tug-of-war model does not account for the possibility that male primates can achieve some reproductive success without engagement in open conflict, such as when a single male cannot guard several receptive females at the same time. We therefore develop a dynamic version of the tug-of-war model, in which reproductive competition causes mortality costs, and in which individuals can gain uncontested shares of reproduction dependent on the degree of female receptive overlap. This model differs substantially from the original tug-of-war model, and derives a new and rich set of comparative predictions. For instance, it predicts that the level of conflict among males declines as the queuing success of subordinate males increases (as survival increases), and also, as their uncontested share of reproduction increases, e.g., as female receptive overlap increases. Our model shows how male–male conflict and female receptive overlap collectively determine the level of reproductive skew among male primates, and illustrates that this relationship is more complex than previously thought.     

Sexual signalling in female crested macaques and the evolution of primate fertility signals

ABSTRACT: BACKGROUND: Female signals of fertility have evolved in diverse taxa. Among the most interesting study systems are those of multimale multifemale group-living primates, where females signal fertility to males through multiple signals, and in which there is substantial inter-specific variation in the composition and reliability of such signals. Among the macaques, some species display reliable behavioural and/or anogenital signals while others do not. One cause of this variation may be differences in male competitive regimes: some species show marked sexual dimorphism and reproductive skew, with males fighting for dominance, while others show low dimorphism and skew, with males queuing for dominance. As such, there is variation in the extent to which rank is a reliable proxy for male competitiveness, which may affect the extent to which it is in females' interest to signal ovulation reliably. However, data on ovulatory signals are absent from species at one end of the macaque continuum, where selection has led to high sexual dimorphism and male reproductive skew. Here we present data from 31 cycles of 19 wild female crested macaques, a highly sexually dimorphic species with strong mating skew. We collected measures of ovarian hormone data from faeces, sexual swelling size from digital images, and male and female behaviour. RESULTS: We show that both sexual swelling size and female proceptivity are graded-signals, but relatively reliable indicators of ovulation, with swelling size largest and female proceptive behaviours most frequent around ovulation. Sexual swelling size was also larger in conceptive cycles. Male mating behaviour was well timed to female ovulation, suggesting that males had accurate information about this. CONCLUSION: Though probabilistic, crested macaque ovulatory signals are relatively reliable. We argue that in species where males fight over dominance, male dominance rank is surrogate for competitiveness. Under these circumstances it is in the interest of females to increase paternity concentration and assurance in dominants beyond levels seen in species where such competition is less marked. As such, we suggest that it may be variation in male competitive regimes that leads to the evolution of fertility signalling systems of different reliability.     

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.     

Size and Partner Choice in Dysdercus maurus Distant (Hemiptera: Pyrrhocoridae)

This study investigates whether females of Dysdercus maurus (Hemiptera) are considering male’s sizes in order to make their reproductive partner choice. Morphometric and biomass measurements were obtained from single males and copulating pairs. Positive correlations were identified for both sizes and weights between males and females in copula. The average ratios established between the couples sizes/weight were around 1.66. Female choice may operate in such a way that each female has her own optimal male size for mating, resulting from a trade-off between the negative influence of large partner size in female fecundity and the advantages of large male size for offspring fitness. Thus, the best choice for a D. maurus female may be a male of average size in relation with her own size.     

Male–female evolutionary game on mate-locating behaviour and evolution of mating systems in insects

We present a model of a male–female evolutionary game on mate-locating behaviour. Two major mating systems are considered: "lek polygyny" (in which males aggregate to wait for females searching for males) and "searching polygyny" (in which males search for females emerging or waiting for males). The model predicts that lek polygyny is favoured (i) when male survivorship during lekking is sufficiently higher than that during mate searching, (ii) when female survivorship while visiting a lek is sufficiently higher than at the emergence site, or (iii) when searching efficiency is higher at a lek than at an emergence site. Furthermore, the model shows that a reduction in the reproductive value of females later in the day, which prevents males from performing riskier mate-locating behaviour, can result in a change of mating system. In addition, mixed mating systems can be realized as transient states during this shift.     

Weather conditions and date influence male Sage Grouse attendance rates at leks

For lek‐breeding birds, lek attendance can be correlated with mating success. Variability in lek attendance could confound interpretation of male reproductive effort and complicate the use of lek counts as an index to monitor abundance. We assessed the daily probability of male Sage Grouse Centrocercus urophasianus lek attendance and explored implications of attendance on lek counts. We fitted 145 males with global positioning system (GPS) transmitters over 4 years in Carbon County, Wyoming. We evaluated influences of lek size and topography, date, weather, and bird characteristics such as age on daily morning lek attendance. The daily probability of attendance ranged considerably each year, from 0.120 (, 95% CI 0.051–0.259) in 2012 to 0.917 (95% CI 0.844–0.957) in 2013 with peak attendance dates ranging from 8 April (2012) to 11 May (2011). Attendance decreased with increasing precipitation on the observation day. Only 44–79% of lek counts occurred on days without precipitation and with high attendance (i.e. within 0.1 probability of peak predicted attendance). Although lek counts and population abundance, predicted using attendance rates, followed a similar trend, the relationship was not significantly correlated. We provide empirical evidence supporting current lek‐count protocols: managers should avoid counting leks on days with precipitation because attendance is reduced. Although managers sometimes only complete one to two lek counts per year on active leks, completing at least three lek counts as recommended in protocols increases chances for higher male counts and improves the relationship between counts and abundance. Attendance varies annually, making it challenging to use lek counts to assess regional population trends over short time periods unless attendance is accounted for.     

Combs and sexual selection in black grouse (Tetrao tetrix)

We studied supra-orbital combs in lekking black grouse (Tetrao tetrix) in relation to sexual selection at five leks in Finland1991-1998 and four leks in Sweden 1992-1995. Comb size wasestimated in two ways: by observing its natural size in thefield at different behaviors ("observed comb size"), and bymeasuring the comb size from captured birds ("measured combsize"). The size of combs is highly variable, and individualscan change it within seconds. Males express their larger combsduring display, as compared to other behaviors. Observed mean comb sizes were larger on leks with a higher number of malesand a higher number of copulations. Measured and observed combsizes and copulatory success did not significantly correlatewhen all males where analyzed, but a positive and significantrelationship between observed comb size and copulatory success was found within males that achieved copulations. Measured comblength correlated positively with the amount of testosterone.While females were present on the lek, displaying and successfulmales showed the largest observed comb size. When we comparedobserved comb size during fighting between successful and unsuccessfulmales and correlated comb size of pairs of fighting males withtheir fighting activity, no significant differences in combsize were found. The result that comb size correlated significantlywith an increase in testosterone level and that larger combsize, within successful males, predicted higher copulatorysuccess suggests that combs may be a cue for females to assessmale quality. The lack of a significant relationship betweenobserved comb size and fighting behavior suggests that combsize either has minor importance in male-male signaling onthe lek or that males may express similar-sized combs duringfighting to avoid serious fights and thus risk of comb injuries.     

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.     

Hormonal and behavioural correlates of male dominance and reproductive status in captive colonies of the naked mole-rat,Heterocephalus glaber.

Naked mole-rat colonies are societies with a high reproductive skew, breeding being restricted to one dominant female (the ''queen'') and 1-3 males. Other colony members of both sexes are reproductively suppressed. Experimental removal of breeding males allowed us to investigate the relationship between urinary testosterone and cortisol, dominance rank, and male reproductive status. Dominance rank was strongly correlated with body weight, age, and urinary testosterone titres in males. No relationship between urinary cortisol levels and male reproductive status or dominance was found. Breeding males were among the highest-ranking, heaviest and oldest males in their respective colonies, and were succeeded by other high-ranking, large, old colony males. In contrast to females, no evidence of competition over breeding status was observed among males. Male-male agonism was low both before and after removal of breeders and mate guarding was not observed. The lower reproductive skew for males compared with female skew or queen control over male reproduction may explain why males compete less strongly than females over breeding status after removal of same-sexed breeders.     

BODY SIZE AND SEXUAL SIZE DIMORPHISM IN MARINE IGUANAS FLUCTUATE AS A RESULT OF OPPOSING NATURAL AND SEXUAL SELECTION: AN ISLAND COMPARISON

Body size is often assumed to represent the outcome of conflicting selection pressures of natural and sexual selection. Marine iguana (Amblyrhynchus cristatus) populations in the Galápagos exhibit 10-fold differences in body mass between island populations. There is also strong sexual size dimorphism, with males being about twice as heavy as females. To understand the evolutionary processes shaping body size in marine iguanas, we analyzed the selection differentials on body size in two island populations (max. male mass 900 g in Genovesa, 3500 g in Santa Fé). Factors that usually confound any evolutionary analysis of body sizes—predation, interspecific food competition, reproductive role division—are ruled out for marine iguanas. We show that, above hatchlings, mortality rates increased with body size in both sexes to the same extent. This effect was independent of individual age. The largest animals (males) of each island were the first to die once environmental conditions deteriorated (e.g., during El Niños). This sex-biased mortality was the result of sexual size dimorphism, but at the same time caused sexual size dimorphism to fluctuate. Mortality differed between seasons (selection differentials as low as –1.4) and acted on different absolute body sizes between islands. Both males and females did not cease growth when an optimal body size for survival was reached, as demonstrated by the fact that individual adult body size phenotypically increased in each population under favorable environmental conditions beyond naturally selected limits. But why did marine iguanas grow “too large” for survival? Due to lek mating, sexual selection constantly favored large body size in males (selection differentials up to +0.77). Females only need to reach a body size sufficient to produce surviving offspring. Thereafter, large body size of females was less favored by fertility selection than large size in males. Resulting from these different selection pressures on male and female size, sexual size dimorphism was mechanistically caused by the fact that females matured at an earlier age and size than males, whereafter they constantly allocated resources into eggs, which slowed growth. The observed allometric increase in sexual size dimorphism is explained by the fact that the difference between these selective processes becomes larger as energy abundance in the environment increases. Because body size is generally highly heritable, these selective processes are expected to lead to genetic differences in body size between islands. We propose a common-garden experiment to determine the influence of genetic factors and phenotypic reaction norms of final body size.     

Microsatellite DNA analysis shows that greater sage grouse leks are not kin groups

The spectacular social courtship displays of lekking birds are thought to evolve via sexual selection, but this view does not easily explain the participation of many males that apparently fail to mate. One of several proposed solutions to this 'lek skew paradox' is that kin selection favours low-ranking males joining leks to increase the fitness of closely related breeders. We investigated the potential for kin selection to operate in leks of the greater sage grouse, Centrocercus urophasianus, by estimating relatedness between lekking males using microsatellite DNA markers. We also calibrated these estimates using data from known families. Mean relatedness within leks was statistically indistinguishable from zero. We also found no evidence for local clustering of kin during lek display, although males tended to range closer to kin when off the lek. These results make kin selection an unlikely solution to the lek skew paradox in sage grouse. Together with other recent studies, they also raise the question of why kin selection apparently promotes social courtship in some lekking species, but not in others.     

Age,condition and dominance‐related sexual ornament size before and during the breeding season in the black grouse Lyrurus tetrix

Male ornaments function as honest cues of male quality in many species and are subject to intra‐ and intersexual selection. These ornaments are generally studied during peak expression, however their size outside the breeding season may determine ultimate ornament size and costliness, and as such reproductive success. We investigated whether male black grouse Lyrurus tetrix eye comb size was related to age, condition and measures of male dominance before and during the breeding season. Total combined eye comb size began to increase ~70 d before the start of the breeding season. Adult males (aged ≥ 2 yr old) had consistently larger eye combs than younger males (1 yr old) both before and during the breeding season. Heavier and more dominant adult males (attending the lek more frequently and successfully reproducing) had larger eye combs. For younger males, those that were heavier had larger eye combs. Additionally, males that spent more time on the lek showed increased eye comb size as the breeding season approached. Overall we find that ornament size is positively related to dominance and condition before and during the breeding season. Since dominance is accrued through year‐round interactions in many species, the ability to maintain larger signals over prolonged periods, including outside of the breeding season, is likely to be beneficial for adults. For younger males, it is likely that they cannot sustain or are constrained from producing larger eye combs over long periods of time. They therefore prioritise growth of their ornaments later, and according to the amount of time they spend on the lek.