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Lek mating

Lek mating is a reproductive strategy in which males of certain species aggregate in a communal area known as a lek to perform courtship displays, while females visit these sites to select mates based on the quality of the displays, receiving no material resources or parental care from the chosen males.^[1] This system emphasizes indirect genetic benefits through female choice and intense male-male competition for display positions.^[1] Leks can vary in size and structure, from tightly clustered territories to more dispersed arenas, but they are typically not associated with essential resources like food or nesting sites.^[2] Lek mating occurs across diverse animal taxa, including birds, mammals, amphibians (anurans), fish, and insects, though it is most extensively studied in birds and insects.^[1] Prominent avian examples include manakins, where males perform acrobatic snaps and dances; sage-grouse, featuring inflated air sacs and strutting; Andean cocks-of-the-rock, with bowing and wing-flapping in noisy gatherings; and birds-of-paradise, known for elaborate plumage shakes and calls.^[3] In mammals, the blackbuck antelope forms leks where males display through leaping and posturing.^[4] Insect instances involve fruit flies such as the Mediterranean fruit fly (Ceratitis capitata), which uses pheromone emissions and wing-fanning in leks.^[1] A defining feature of lek mating is the strong sexual selection imposed by female mate choice, often favoring males in central or "hotspot" positions within the lek that receive disproportionate mating success.^[5] Males invest heavily in display traits—visual, acoustic, or chemical signals—while females benefit from reduced search costs by evaluating multiple suitors in one location.^[1] This dynamic raises the lek paradox, the puzzle of how genetic variation in attractive male traits persists despite directional selection that should erode it over time.^[6] Proposed resolutions include condition-dependent expression of traits, where displays honestly signal overall genetic quality, and fluctuating selection pressures.^[6]

Overview

Etymology and history

The term "lek" originates from the Swedish word lek, meaning "play" or "game," and was adopted into English ornithology in the late 19th century to describe communal display areas used by males for courtship. It was first employed in this context by Llewelyn Lloyd in his 1867 book The Game Birds and Wild Fowl of Sweden and Norway, where he documented the aggregated mating displays of species like the western capercaillie (Tetrao urogallus) and European black grouse (Tetrao tetrix) on traditional grounds in Scandinavia.^[7]^[8] Early observations of lekking date to the 19th century, when naturalists across Europe noted the dramatic communal gatherings of male European black grouse at display sites, often in open meadows or forest clearings, with records from locations including the United Kingdom's New Forest and southern England. These anecdotal reports highlighted the intense male-male competitions and vocalizations during spring breeding seasons but lacked systematic analysis. By the early 20th century, more detailed ethological studies emerged, such as those by Edmund Selous, who in 1909 described the territorial dynamics and aggressive interactions on black grouse leks in England, establishing lekking as a notable form of avian social behavior.^[9]^[10] In the 1930s to 1950s, pioneering ethologists including Niko Tinbergen advanced the scientific study of lek mating through observational and experimental approaches to courtship displays and innate behaviors, framing leks within broader principles of animal communication and sexual selection. Subsequent European research in the mid-20th century solidified lekking as a distinct mating system, distinct from resource-based polygyny. By the 1970s, the phenomenon was integrated into modern sexual selection theory, with seminal theoretical models by John W. Bradbury exploring the evolutionary origins of male clustering and female choice on leks.^[11]^[12]

Taxonomic range

Lek mating systems are predominantly observed in birds and mammals, with emerging evidence in select insect, fish, and amphibian species.^[13] Among vertebrates, these systems are characterized by males aggregating to display without providing resources such as food or nesting sites to females.^[2] In birds, lek mating is widespread, documented in approximately 150 species across various families, including the Paradisaeidae (birds of paradise) and Phasianidae (grouse and allies).^[14] Notable examples include the greater sage-grouse (Centrocercus urophasianus), which forms classical leks on open leksites where males perform elaborate strut displays during breeding seasons.^[15] Manakins in the family Pipridae, such as the wire-tailed manakin (Pipra filicauda), also exhibit coordinated lek displays involving cooperative courtship behaviors.^[16] Mammalian lek systems are rarer, occurring in about 15-20 species, primarily ungulates, which contrasts with the prevalence in birds.^[17] Key examples include antelopes like the Uganda kob (Kobus kob), where males defend small territories in leks on floodplains, and Jackson's hartebeest (Alcelaphus buselaphus johnstoni), which aggregates in similar resource-neutral arenas.^[18] These systems are less common in mammals overall, representing under 2% of species.^[19] Evidence for lek mating extends to other taxa, including insects such as certain butterflies and fruit flies (Drosophila), where males cluster to release pheromones and perform displays.^[15] In fish, lek-like aggregations occur in some cichlids, particularly bower-building species in Lake Malawi, such as Copadichromis and Ithophaga, where males construct sand bowers in communal courts.^[20] In amphibians, lek mating is confirmed in various anuran species, such as the neotropical frog Ololygon rubra, where males form choruses for communal displays.^[21] Recent studies have confirmed lek behavior in bats, including the greater mouse-eared bat (Myotis myotis), with 2025 research documenting male aggregations in church attics visited by females for mating.^[22] No confirmed cases exist in reptiles under strict lek definitions.^[13] The distribution of lek mating is influenced by environmental factors, particularly in resource-poor habitats where males cannot provide material benefits, favoring female choice based solely on genetic quality indicators.^[2] This pattern underscores leks' evolution in contexts of low resource defensibility and high female mobility.^[23]

Lekking behavior

Types of leks

Leks are classified primarily into classical and exploded types based on the spatial arrangement and size of male display territories. Classical leks feature small, tightly clustered territories where males defend areas typically less than 100 meters in diameter, allowing females to assess multiple males in close proximity during brief visits.^[5] In these systems, males aggregate in a central arena, often on bare ground or low vegetation, to perform coordinated displays. Representative examples include the white-bearded manakin (Manacus trinitatis), where males form leks of 10–30 individuals in understory forests, facilitating visual and auditory courtship signals.^[24] Exploded leks, in contrast, involve larger and more dispersed aggregations of male territories, with males separated by distances ranging from hundreds of meters to several kilometers, yet maintaining a loose overall clustering that influences female visitation patterns.^[25] This configuration allows males to defend broader areas while still benefiting from proximity to competitors. In mammals such as the topi antelope (Damaliscus lunatus), exploded leks span up to 2–3 km, where males use olfactory and vocal cues to attract females over greater distances.^[26] Similarly, in birds like the great bustard (Otis tarda), males display in dispersed sites connected by female movement corridors, emphasizing endurance-based displays rather than tight synchronization.^[27] Other variants include linear leks, where male territories align along ridges or linear features such as forest edges, often due to topographic constraints that concentrate display sites. In black grouse (Tetrao tetrix), these elongated leks form along lekking arenas up to several hundred meters long, with males spaced 5–10 meters apart to optimize acoustic signaling in open habitats.^[28] Arena leks, observed in insects, resemble classical leks but occur in temporary swarms or choruses without fixed territories; for instance, in tephritid fruit flies (Anastrepha spp.), males aggregate on fruiting trees to produce pheromonal and wing-fanning displays, with arenas forming dynamically based on resource hotspots.^[29] Distinctions among these types hinge on territory size, inter-male spacing, and environmental factors, with classical and arena forms typically under 1 km² and exploded and linear forms extending farther.^[5] Functionally, classical and arena leks prioritize visual and auditory displays for rapid female evaluation, as seen in avian and insect species where proximity enhances comparative assessment.^[30] Exploded leks, prevalent in mammals, rely more on olfactory cues and prolonged signaling, allowing females to traverse larger areas while detecting males via scent marks or calls.^[31] Recent studies frame these variations as emergent spatial patterns arising from collective behaviors, where simple rules of male attraction to high-density sites and female search efficiency drive clustering without centralized coordination.^[32]

Lek stability and dynamics

Lek formation begins with the seasonal aggregation of males at traditional sites, where they arrive weeks or months prior to the peak breeding period to establish and defend territories. This process is driven by social and environmental cues, leading to the rapid organization of display arenas in species such as ground-displaying birds, where males often reuse fixed locations across multiple years.^[33] In the wire-tailed manakin (Pipra filicauda), for instance, territorial males settle into leks during the pre-breeding phase, with recruitment of younger males increasing with age and plumage maturity to bolster the aggregation.^[34] Stability of leks is maintained through high site fidelity among males, particularly in birds like greater sage-grouse (Centrocercus urophasianus), where annual return rates to the same lek exceed 97% for adults.^[35] This fidelity is influenced by habitat quality, with persistent leks forming in areas offering suitable vegetation and low disturbance, and by predation risk, which can modulate male retention if sites provide visual advantages or escape cover.^[36] In sage-grouse, environmental stressors like severe winters can disrupt this stability, causing temporary shifts in lek locations by 200–400 meters and reduced male retention.^[36] Similarly, in Amazonian leks, annual survival probabilities of 0.74–0.79 for males correlate with lek persistence, with El Niño conditions enhancing survival by up to 10% through improved fruit availability and body condition.^[34] Lek dynamics involve pronounced fluctuations in male numbers, with daily variations tied to weather and female presence, and seasonal peaks during active breeding followed by post-breeding dissolution as males disperse. Attendance rates peak at 0.68 for adult sage-grouse males in March–April, declining sharply afterward, while subadults join later at lower rates of 0.42.^[37] Annual male turnover typically ranges from 20–50%, driven by mortality rates of 40–42% in sage-grouse and recruitment of new individuals, which can lead to reorganization if dominant males are lost.^[38] High-resolution GPS tracking from 2017 studies on great bustards (Otis tarda) has illuminated these dynamics, revealing fine-scale spatial shifts within leks—such as males adjusting territories by tens of meters daily—and overall movements that stabilize the arena during peak activity but allow dissolution outside breeding. Human-induced disruptions, particularly habitat fragmentation, significantly undermine lek stability by isolating sites and reducing male recruitment, as seen in lesser prairie-chickens (Tympanuchus pallidicinctus) where lek persistence declines with increasing land-use changes like agriculture and energy development.^[39] In fragmented landscapes, connectivity between leks decreases, leading to lower attendance and higher abandonment rates over years. Poaching exacerbates these issues in lekking ungulates, contributing to population declines that destabilize aggregations, though specific metrics for antelope leks remain limited. Metrics for assessing stability include peak attendance counts (e.g., 17–28 males per active sage-grouse lek) and turnover estimates derived from mark-recapture, providing indicators of persistence against anthropogenic pressures.^[40]^[38]

Female mating preferences

In lek mating systems, females select mates primarily based on male displays rather than resources or parental care, focusing on traits that signal genetic quality. For instance, in golden-collared manakins (Manacus vitellinus), females prefer males exhibiting vigorous cooperative courtship dances involving rapid flights and snaps, which correlate with higher mating success.^[41] Similarly, in birds of paradise such as Lawes' parotia (Parotia lawesii), females assess elaborate visual displays like tail fanning and head shaking, choosing males with the most synchronized and intense performances.^[42] Female preferences operate through multiple sensory modalities, including visual, auditory, and olfactory cues. Visually, plumage brightness and symmetry attract females; in blue-crowned manakins (Lepidothrix coronata), females favor males with high display rates over those with symmetric plumage alone.^[43] Auditory signals, such as complex calls, also play a key role; playback experiments in sage grouse (Centrocercus urophasianus) demonstrate that females approach and linger near recordings of vigorous male strutting and wing-whirring sounds, indicating a preference for dynamic vocal displays.^[44] In mammalian leks, like those of topi antelopes (Damaliscus lunatus), olfactory cues from soil in male territories influence choice, with females more likely to visit and mate in areas scented by prior female activity, amplifying stochastic mating variance.^[45] Females typically assess males during brief lek visits lasting 1-5 minutes, allowing comparison of multiple suitors before copulation.^[46] In lance-tailed manakins (Chiroxiphia lanceolata), females often mate with 1-3 males per visit, leading to multiple paternity in about 14% of nests, particularly when selecting inexperienced sires, followed by post-copulatory cryptic female choice that biases fertilization toward preferred sperm.^[47] Experimental evidence from playback studies further supports preferences for complex auditory traits, as females in black grouse (Tetrao tetrix) respond more to multifaceted calls indicating low parasite loads.^[48] Preferences exhibit context-dependent variation, with stronger selection for symmetry in parasite-rich environments. In black grouse leks, females indirectly favor symmetric males by avoiding those with high fluctuating asymmetry in tarsi—a marker of developmental instability linked to parasites—resulting in central, low-asymmetry males achieving more copulations.^[49] Recent 2025 research on the woodwasp Sirex noctilio, a lekking insect, shows that age is a key trait, with older males achieving higher mating success due to increased sexual maturity.^[50]

Costs and benefits

For males

In lek mating systems, males experience substantial benefits in reproductive potential due to the high degree of mating skew observed across species. Typically, a small proportion of males—often the top 10%—secure the majority of copulations, with at least 90% of matings in sage grouse leks (Centrocercus urophasianus) achieved by no more than 10% of the males present.^[10] This skew allows dominant males to achieve disproportionate reproductive success without providing resources or parental care to females or offspring, enabling them to allocate energy primarily toward elaborate displays and territory defense.^[51] However, these benefits come at significant costs, particularly in terms of energetic expenditure. In sage grouse, vigorously displaying males incur daily energy costs up to four times their basal metabolic rate, far exceeding that of non-displaying individuals.^[52] Similarly, in great snipe (Gallinago media), displaying males expend approximately four times their basal metabolic rate on average during lek activity.^[53] Aggregation on leks also elevates predation risk, as clustered males become more conspicuous to predators, potentially attracting them to the site despite possible dilution effects in larger groups.^[54] Additionally, intense territorial competition often results in physical injuries; in white-eared kob antelopes (Kobus kob leucotis), aggressive encounters lead to damaging wounds and disproportionately high mortality among breeding-age males.^[55] Trade-offs further complicate male strategies in lek systems. Older males often gain an advantage through accumulated experience, achieving higher mating success compared to younger counterparts, as seen in black grouse (Lyrurus tetrix) where yearlings in good condition can secure territories but rarely match the copulation rates of established adults.^[56] However, advancing age correlates with increased parasite loads, exacerbated by prolonged exposure and testosterone-driven behaviors that heighten susceptibility to infections in species like red grouse (Lagopus lagopus scotica).^[57] Recent genomic analyses reinforce these costs: a 2025 study on black grouse identified that loads of predicted deleterious mutations (measured via GERP++ scores) reduce lek attendance—a key behavioral trait for mating—by impairing overall genetic quality, with homozygous effects showing stronger negative impacts on lifetime mating success (β = -0.21).^[58] Ultimately, these dynamics result in high variance in male fitness outcomes, with most individuals securing zero copulations while a few dominant males monopolize reproduction.^[59] In antelopes such as the white-eared kob, lek participation contributes to reduced lifespan for breeding males through elevated injury and mortality risks, shortening their effective reproductive tenure compared to non-territorial individuals.^[60]

For females

In lek mating systems, females benefit from evaluating and mating with multiple high-quality males aggregated at display sites without the need for resource exchanges such as territories or parental care, allowing selection based primarily on genetic quality indicators like elaborate displays.^[61] This aggregation facilitates efficient comparison of potential mates, potentially yielding indirect genetic benefits through "good genes" that enhance offspring quality.^[61] Additionally, brief lek visits minimize exposure to predation risks compared to prolonged interactions in other mating systems, as females can quickly assess and depart after mating.^[62] Despite these advantages, lek mating imposes costs on females, including time and energy expended on travel to and from leks. In sage grouse, for example, extra travel associated with lek visits increases daily energetic expenditure by approximately 1% of the field metabolic rate, with total seasonal costs remaining low at around 12.4 kJ for typical movement patterns.^[62] Harassment risks exist but are generally low, as females retain control over mating decisions and can reject advances or leave territories at will.^[62] Lek visits may also increase exposure to parasites or pathogens due to male aggregation, though empirical evidence in birds shows variable transmission rates.^[63] Lek systems enhance the efficiency of female mate choice relative to resource-based mating, where females must invest time in assessing variable male-held resources; here, centralized male displays allow rapid evaluation of multiple suitors without such commitments.^[61] Females in lekking species often assess 5–20 males per breeding season by visiting one or more leks and sampling displays sequentially.^[43] Recent studies further indicate that environmental factors, such as fruit resource availability, can modulate these choice benefits; in white-bearded manakins, higher ripe fruit biomass near display courts boosts male display rates, indirectly increasing female visitation and mating success through enhanced assessment opportunities.^[64] Outcomes for females include elevated offspring viability from selecting males with superior display traits. Although no paternal care is provided, the potential for multiple paternity—observed in up to 44% of nests in some lekking birds—promotes genetic diversity within broods, potentially buffering against environmental variability.^[61]

Paradoxes and genetic considerations

The lek paradox

The lek paradox arises in lek mating systems where females impose intense directional sexual selection on male display traits, such as elaborate plumage, dances, or calls, which serve as indicators of "good genes" for offspring viability, including symmetry and overall vigor. Despite this strong selection favoring preferred traits, additive genetic variation in these traits does not erode as expected, persisting across generations and challenging core predictions of sexual selection theory that preferred alleles should rapidly fix in the population.^[65] This paradox stems from theoretical models of sexual selection, including Fisherian runaway processes—where female preferences and male traits coevolve through positive feedback—and the handicap principle, which posits that costly signals reliably indicate genetic quality. Both frameworks predict that sustained female choice would deplete heritable variation by driving advantageous alleles to fixation, yet empirical observations reveal sustained polymorphism in preferred traits. Empirical studies in lekking birds, such as manakins and grouse, demonstrate high heritability in display traits, typically ranging from 0.2 to 0.5, with no observed decline over multiple generations despite ongoing female preference. For instance, analyses of tail ornaments and vocalizations show elevated additive genetic variance compared to non-sexual traits, supporting the paradox's persistence in natural populations.^[66] The lek paradox has profound implications for indicator models of sexual selection, questioning their long-term efficacy in transmitting genetic benefits and highlighting potential roles for fluctuating environmental pressures or other factors in maintaining variation. Although resolutions involving genetic mechanisms, such as mutation-selection balance, have been proposed, the core conundrum underscores limits in our understanding of how sexual selection interacts with genetic diversity.^[67] Historically, the paradox was first articulated by R. A. Fisher in his 1930 treatise on natural selection, which laid the groundwork for understanding runaway processes, but it was formalized specifically for lek systems by Pomiankowski and Møller in the 1990s through comparative analyses of trait variance.^[65]

Genetic mechanisms

Quantitative genetics studies have demonstrated that sexually selected display traits in lekking species exhibit moderate heritability, typically ranging from 0.3 to 0.5, indicating a polygenic basis influenced by multiple loci of small effect.^[66] For instance, in songbirds, including some with lek-like mating, song complexity shows heritabilities in this range, reflecting additive genetic variance that supports variation in male attractiveness despite directional female preferences.^[66] This polygenic architecture allows for the maintenance of genetic diversity in traits like vocalizations and physical displays, as evidenced by biometric analyses in lek systems where mating success correlates with heritable components of performance.^[68] Recent research has highlighted the role of deleterious mutations in shaping lekking fitness, particularly through viability effects that reduce male participation without directly impairing ornamental traits. In the black grouse (Lyrurus tetrix), a classic lekking bird, whole-genome sequencing of 190 males revealed that loads of predicted deleterious mutations (identified via conservation scores and functional impact) negatively affect lifetime mating success by lowering lek attendance.^[69] Specifically, higher mutation loads decreased annual lek attendance (β = -0.13), indirectly reducing mating opportunities, while having minimal impact on display ornaments like lyre tail size or eye comb coloration.^[69] These findings from 2025 suggest that such mutations preserve genetic variation by imposing viability costs that prevent fixation of low-fitness alleles, thus sustaining polymorphism essential for female choice.^[69] Gene expression underlying lek traits is modulated by hormones such as testosterone, which activates androgen-responsive loci to enhance display performance. In golden-collared manakins (Manacus vitellinus), testosterone treatment upregulates hundreds of genes in wing muscles critical for acrobatic courtship snaps, including those involved in lipid metabolism and energy provision (e.g., APOB and MTTP), with over half containing androgen response elements for direct regulation.^[70] Genome-wide association studies (GWAS) and selection analyses in manakins have identified loci linked to plumage coloration, such as SLC45A2 under positive selection, influencing melanin-based ornaments that signal male quality during leks.^[71] Advances from 2024–2025 have elucidated the genetic architecture of lek species, emphasizing condition-dependent expression that preserves polymorphism. In black grouse, deleterious mutations in promoter regions exert strong fitness effects, indicating regulatory variants contribute to variable trait expression based on individual condition.^[69] These studies reveal a polygenic foundation where condition sensitivity allows environmental factors to reveal underlying genetic variance, resolving persistent questions about trait diversity in leks.^[69] While primarily DNA-based, non-genetic factors like epigenetics interact with these mechanisms; for example, a 2025 study on black grouse found DNA methylation changes before and after breeding associated with lek performance and reproductive investment in 50 males, modulating gene activity in response to environmental cues without altering the genome.^[72]^[73]

Evolution of lek mating

Hotshot hypothesis

The hotshot hypothesis, proposed by Beehler and Foster in 1988, explains lek formation as a male-driven process centered on dominant, highly attractive individuals known as "hotshots" who initiate aggregation by drawing in subordinate males seeking to parasitize their mating success. This model posits that hotshots establish territories that attract females due to their superior displays or dominance, prompting less competitive peripheral males to join nearby in hopes of intercepting females who arrive but do not mate with the central male. Unlike female-centered explanations, the hypothesis emphasizes intrasexual competition and male decision-making as key to lek development and maintenance.^[74] The core mechanism involves a skewed distribution of mating success, with hotshots securing the majority of copulations while satellites benefit from occasional spillover matings or errors in female choice. This leads to predicted spatial clustering, where subordinates position themselves peripherally around the hotshot to minimize aggression but remain close enough for interception opportunities. In classical leks, this dynamic fosters tight aggregation, as the presence of a successful hotshot signals high female traffic to joining males. Beehler and Foster drew from observations in avian systems to argue that such male-male conflict sustains lek structure over time.^[74] Empirical support derives from studies in birds of paradise, where dominant males occupy core territories and monopolize matings; for example, in the lesser bird-of-paradise (Paradisaea minor), a single hotshot in a group of eight males achieved 24 of 25 observed copulations. Experimental evidence further bolsters the model: in little bustards (Tetrax tetrax), attractive male decoys induced wild males to aggregate around them, mimicking hotshot attraction, whereas neutral or female decoys failed to elicit similar clustering. These findings confirm that males respond to cues of success in conspecifics, promoting lek formation around perceived dominants.^[74] The hypothesis predicts that lek size scales with hotshot quality, as more attractive central males recruit greater numbers of subordinates, enhancing overall stability in classical leks. However, it applies less effectively to exploded leks, where dispersed territories reduce the feasibility of satellite interception and dominance enforcement. In contrast to the hotspot model, the hotshot hypothesis underscores male-initiated dynamics over environmental cues for aggregation.^[27]

Hotspot model

The hotspot model posits that leks evolve as males aggregate at specific locations, or "hotspots," where females are predicted to search for mates, thereby maximizing male encounter rates with minimal search effort. Developed by Bradbury in 1981, this hypothesis suggests that such hotspots often occur at predictable sites like the edges of female foraging areas or migration routes, where female traffic is concentrated due to resource distribution.^[75] The mechanism underlying the hotspot model is game-theoretic: individual males, acting in their own interest, converge on locations that optimize the probability of female encounters, accounting for competition from other males and the costs of searching across the landscape. Computer simulations of male settlement patterns demonstrate that this leads to stable clustering at hotspots, as males adjust positions to balance personal mating opportunities against shared female access, predicting fixed lek sites rather than dispersed territories.^[75] Empirical evidence supports the model through field observations in lekking antelopes, such as topi (Damaliscus lunatus), where leks form near female foraging concentrations, with radiotelemetry data showing higher female visitation to these sites compared to random locations.^[76] Modeling studies further corroborate this, including dynamic simulations of seasonal hotspots that confirm lek persistence in areas of fluctuating female density, such as resource-rich patches during breeding seasons.^[75] The model predicts that lek locations are primarily determined by habitat features influencing female movement, with evolutionary pressures favoring aggregation only where it enhances female mate-search efficiency by concentrating options in accessible spots. However, it has limitations, as it focuses on female distribution without accounting for mechanisms like male-male attraction that might reinforce clustering.^[75]^[77]

Blackhole model

The black hole model of lek evolution, proposed by Stillman, Clutton-Brock, and Sutherland, describes leks as deceptive sinks or evolutionary traps that attract and retain males despite minimal expected mating benefits for most participants. In this framework, leks initially form when mobile estrous females searching for mates are more likely to be retained by clusters of male territories than by solitary males, as females tend to mate with the first male encountered and clusters reduce the chance of females escaping to competitors. This retention dynamic creates a feedback loop: the presence of females draws additional males to the cluster, amplifying the "trap" effect, where the system persists through rare mating successes by a few dominant males, even though the average male experiences zero net gain. The model emphasizes passive aggregation driven by female mobility rather than active female preferences for clustered males. Simulations extending the model to larger habitats and realistic female mobility limits illustrate its persistence mechanisms, showing that lek sizes increase with male density and that leks space evenly at intervals approximating one female home range diameter due to the retention advantage of clusters. Empirical patterns, such as in sage grouse (Centrocercus urophasianus), align with the model's implications of extreme skew in success, where roughly 90% of males at leks secure no copulations, yet aggregation endures because the visibility of females signals potential opportunities, sustaining influx despite high failure rates. These simulations highlight how variance in male success—concentrated among a small subset—upholds the structure without requiring female choice.^[78]^[10] The model predicts high male turnover, with unsuccessful individuals frequently replaced by newcomers attracted to established sites, a pattern observed in species like the Uganda kob (Kobus kob), where rapid shifts in male composition occur throughout breeding seasons. It particularly applies to taxa with male-biased dispersal, such as many ungulates, where young or dispersing males preferentially join existing leks to exploit perceived mating hotspots. However, the black hole model has been critiqued for limited empirical validation outside ungulates, with studies in avian lekking species suggesting it underperforms relative to alternatives like hotspot or hotshot hypotheses, as female movement patterns often fail to match the predicted retention dynamics in non-mammalian systems.^[33]

Kin selection hypothesis

The kin selection hypothesis applies W. D. Hamilton's framework of inclusive fitness to explain lek formation, positing that related males aggregate on leks to gain indirect reproductive benefits from the mating success of their kin, thereby increasing overall genetic representation despite limited direct benefits for subordinates. This model suggests that the extreme reproductive skew in leks—where few males monopolize matings—can be evolutionarily stable if non-breeding males are kin to successful breeders, as the indirect fitness gains satisfy Hamilton's rule (rB > C, where r is relatedness, B the benefit to the recipient, and C the cost to the actor). Under this hypothesis, the mechanism centers on nepotism in spatial arrangement, with males preferentially establishing display territories near relatives to minimize aggressive costs among kin and collectively enhance lek attractiveness to females, who may assess group displays or size. Limited male philopatry or kin recognition facilitates such clustering, allowing subordinates to contribute to lek persistence without direct mating opportunities. Genetic evidence supports kin structuring in several avian lek systems. In black grouse (Tetrao tetrix), microsatellite loci analysis demonstrated that average pairwise relatedness within leks was 0.09, significantly exceeding the 0.03 expected under random population sampling, indicating non-random kin aggregation. Comparable patterns occur in Indian peafowl (Pavo cristatus), where DNA fingerprinting revealed higher band-sharing coefficients (indicating relatedness) among males on the same lek compared to those on separate leks, persisting even without prior social or environmental familiarity. In the white-bearded manakin (Manacus manacus), leks comprise multiple discrete kin clusters of closely related males (often full siblings or father-son pairs), with elevated local relatedness promoting cooperative display elements. These studies highlight higher relatedness in stable avian systems, typically 10-20% above random expectations in clustered subgroups, though averages remain modest due to multi-family leks.^[79] The hypothesis predicts that kin-based clustering resolves the lek paradox—why males endure high competition with low direct success—by accruing indirect benefits that favor lek evolution in philopatric populations, potentially amplifying good-gene transmission through familial success.^[79] Limitations arise in systems lacking kin structure, such as dispersed lekking mammals, where genetic analyses show negligible relatedness within leks, suggesting kin selection plays a minor role compared to other drivers like female preference for aggregation.

Predation protection hypothesis

The predation protection hypothesis proposes that lek mating systems evolve primarily as an anti-predator adaptation, where males aggregate to dilute individual risk during conspicuous courtship displays that leave them vulnerable to attack. According to this view, leks function as "selfish herds," a concept introduced by Hamilton, in which each male reduces its own predation probability by clustering with others, thereby shifting the risk onto companions rather than isolating displays that would heighten personal exposure. This aggregation-based protection is particularly relevant during the brief, intense display periods when males are distracted and less vigilant.^[80] The underlying mechanisms include risk dilution across the group, where the per capita chance of being targeted decreases with larger lek sizes, and the confusion effect, which hampers visual predators' ability to isolate a single individual amid coordinated movements and displays. Central positions within the lek offer enhanced safety, as peripheral males bear disproportionately higher risk, though this benefit is offset by intensified intrasexual competition for prime spots and mates. Trade-offs arise because while grouping provides collective defense, it may amplify overall detectability to predators scanning for activity hotspots. Empirical evidence supports this hypothesis through observations of elevated survival rates in lek centers compared to edges; for instance, in ground-displaying birds like grouse, central territories are positioned farther from human disturbances and potential predator approaches, correlating with lower estimated predation rates in central positions in avian studies. In Neotropical manakins, lek participation aligns with varying threat levels, as larger groups form under higher eagle predation pressure, and overall predation events on leks remain notably low despite conspicuous displays. Recent analyses of collective behavior in lekking species further demonstrate emergent protective dynamics, where synchronized displays confuse approaching predators and enhance group-level vigilance. The hypothesis predicts that leks should predominate in habitats with elevated predation pressure, such as open savannas or forests with aerial hunters, where solitary displays would be suicidal; indeed, species like Uganda kob exhibit lekking in predator-rich environments, with aggregation reducing attack success rates. However, limitations include the potential for leks to increase overall conspicuousness, drawing more predators to the site and negating benefits in low-predation areas where dispersed mating suffices without aggregation costs. This hypothesis thus explains lek evolution in high-risk ecological contexts but is less applicable where other factors, such as resource distribution, dominate.

References

[1]
Sexual Selection on Leks: A Fruit Fly Primer - PMC - PubMed Central
Lek mating systems are relatively rare but occur in a diverse taxonomic array of animals, including birds, mammals, anurans, and insects.
[2]
Lekking - an overview | ScienceDirect Topics
A lek is an aggregation of males, each seeking to attract a mate, that is not associated with a key resource. Of Special Interest: Bachelor Herds. What do ...
[3]
Look at that Lek! 9 Birds That Put on A Show - Cool Green Science
Apr 11, 2022 · A lek is a gathering of male birds that strut, dance, or sing to show off for females, who then select the most fit male.
[4]
Lekking as collective behaviour - Journals
Feb 20, 2023 · Figure 1. Examples of lek-mating systems. (a) Blackbuck (Antilope cervicapra) lek at Velavadar National Park (picture credit: Shruti Hegde). (b ...
[5]
Lek Territory Size and the Evolution of Leks: A Model and a Test ...
Thus, to understand the maintenance of the lek mating system, we need to evaluate both evolutionary processes favoring key male and female mating decisions ...<|control11|><|separator|>
[6]
The lek paradox and the capture of genetic variance by condition ...
Our explanation offers a resolution to the lek paradox and rests on only two assumptions; condition dependence of sexually selected traits and high genetic ...
[7]
Lek - Etymology, Origin & Meaning
From Swedish leka "to play," the term means "to engage in courtship displays" among certain animals, highlighting playful mating behavior.
[8]
Leks: on JSTOR
The terms “lek” and “lekking behavior” were first used for the mating arenas of birds (L. Lloyd 1867), but more or less similar mating aggregations occur in a ...Missing: seminal | Show results with:seminal
[9]
Black grouse through history - Game and Wildlife Conservation Trust
Black grouse was first mentioned in literature in Medieval times, then referred to by their folk names blackcock and greyhen.
[10]
The Lek Mating System of the Sage Grouse - jstor
My own observations show that fe males arriving at a lek tend to follow one another and to stop as a group even when they are still some distance from the ...
[11]
Nikolaas Tinbergen – Facts - NobelPrize.org
Nikolaas Tinbergen used dummies in his experiments. One of his discoveries at the end of the 1930s was that birds preferred to brood eggs with exaggerated ...Missing: lek | Show results with:lek
[12]
Lek Mating Behavior in the Hammer‐headed Bat - Bradbury - 1977
Calling assemblies of Hypsignathus monstrosus were studied for 17 months in the field to determine if they were “leks” or mating arenas.
[13]
https://press.princeton.edu/books/hardcover/9780691634920/leks
[14]
Lek mating systems: a case study in the Little Bustard Tetrax tetrax
Leks are male display aggregations where females seek mates, with no male parental care, and females finding only male genes as resources.
[15]
9.2 Leks and arena mating systems - Animal Behavior
The lek mating system is characterized by intense male competition for mating opportunities and female mate choice based on male displays and traits · Leks ...
[16]
Social networks in the lek-mating wire-tailed manakin (Pipra filicauda)
Lek-mating manakins (Pipridae) are neotropical birds well known for their exaggerated courtship displays that involve complex movements by, and sometimes ...
[17]
Is lek breeding rare in bats? - Toth - 2013 - ZSL Publications - Wiley
Aug 15, 2013 · Bats (Mammalia: Chiroptera) are among the most successful mammals and likely display the widest range of mating systems within the Class.
[18]
What the Heck Is a Lek? The Quirkiest Mating Party on Earth.
Mar 15, 2017 · It's a gathering of males to show off their genes without any material benefits to a female. You might notice that birds aren't part of that definition.
[19]
Sexual selection in lekking antelope
Lekking—in which males gather in a single area to court visiting females—is a spectacular and rare example of such a mating system. With aerial videography, ...
[20]
Mate choice and sexual selection in a lekking cichlid fish ...
Sexual selection has two components: mate choice and intrasexual competition, both of which can result in differences in mating success.
[21]
Mating system and copulatory behavior of the greater mouse-eared ...
Jun 18, 2025 · Our findings suggest a lek mating system, where males aggregate and are visited by receptive females. Mating involves multiple copulations and distinct body ...
[22]
Evolution of ungulate mating systems: Integrating social and ...
Apr 15, 2020 · The quality and defensibility of resources affect mating systems, as does the defensibility of females. Population density of females also may ...Missing: poor | Show results with:poor
[23]
[PDF] PIPRIDAE - Prum Lab
rived from more concentrated or “classical” leks, and therefore literally “exploded” in size. ... Lekking in birds and mammals: behavioral and evolutionary issues ...
[24]
Exploded leks: What bustards can teach us - ResearchGate
Aug 6, 2025 · Exploded leks: what bustards can teach us. In lek mating systems, males defend small, clus-tered courts visited by females to mate (Höglund ...
[25]
Lek-breeding in topi antelopes (Damaliscus lunatus) - ResearchGate
These results suggest that male topi antelopes can obtain central lek territories through alternative mating tactics, providing scope for variance in male ...
[26]
[PDF] EXPLODED LEKS: WHAT BUSTARDS CAN TEACH US - Ardeola
a) Lek paradox scenario where benefits of search in a resource-based mating system are higher than in a lek mating system, while costs remain equal. b) A ...
[27]
Effects of density on lek-site selection by Black Grouse Tetrao tetrix ...
The Black Grouse is a primarily lekking species, but low population density and lack of suitable habitat can lead to the establishment of non-lekking ...
[28]
Sexual Selection on Leks: A Fruit Fly Primer | Journal of Insect Science
May 29, 2018 · The objective of this paper is to provide an overview of empirical findings on lek-forming tephritid and drosophilid flies.
[29]
Size and Plumage Dimorphism in Lek-Breeding Birds
This definition includes both classical leks (Bradbury 1981), where males display within sight of each other, and exploded leks (Gilliard 1969), where males ...
[30]
The evolution of leks through female choice: differential clustering ...
Bradbury JW (1977) Lek mating behavior in the hammer-headed bat. Z Tierpsychol 45:225–255. Google Scholar. Bradbury JW (1981) The evolution of leks. In ...
[31]
Lekking as collective behaviour - PMC - PubMed Central - NIH
Feb 20, 2023 · Lekking is a spectacular mating system in which males maintain tightly organized clustering of territories during the mating season, and females visit these ...
[32]
Leks in ground-displaying birds: hotspots or safe places?
Abstract. We studied the effects of female distribution, topography, and human infrastructures on lek-site selection in a ground-displaying bird, the great.Missing: insects | Show results with:insects
[33]
Climate, demography and lek stability in an Amazonian bird - PMC
Multiple ecological factors and their interactions have been hypothesized to influence lek dynamics, including high survival of territorial males, delayed ...Missing: poor | Show results with:poor
[34]
Fine-scale genetic structure among greater sage-grouse leks in ...
Very strong breeding-site fidelity has been documented for these specific leks across multiple years of monitoring, where 330 of 345 (98.5 %) among-year ...
[35]
[PDF] Lek Organization in Sage Grouse: Variations on a Territorial Theme
contributing to lek stability across years. The lek did alter over this period, however: indi- vidual 50% MAP areas and overlaps increased from 1984 to 1985 ...
[36]
Assessing lek attendance of male greater sage‐grouse using fine ...
Feb 7, 2019 · Overall, adult males attended leks at higher rates (0.683 at peak) and earlier in the season (19 March) than subadults (0.421 at peak on April ...Missing: turnover replacement
[37]
[PDF] Characteristics and Dynamics of Greater Sage-Grouse Populations
Mar 1, 2011 · Annual survival of male sage-grouse was estimated to be 59% in Wyoming (June 1963),. 58–60% in Idaho (Connelly et al. 1994, Wik 2002), and 30 ...
[38]
Patterns in lek persistence and attendance by lesser prairie‐chicken ...
Apr 11, 2025 · To model lek stability, we used Bayesian binomial regression to model the number of years each lek was active relative to the number of years ...
[39]
Sage grouse lek attendance remains steady
Sep 6, 2022 · This spring, officials counted an average of 17.9 male sage grouse per active lek.
[40]
Behavioural flexibility in the courtship dance of golden-collared ...
In manakins and other lekking species such as bowerbirds, females visit displaying males several times before choosing their mating partner; therefore, a ...
[41]
SEXUAL SELECTION THROUGH FEMALE CHOICE IN LAWES ...
We studied sexual selection in Lawes' Parotia, a lek-mating bird of paradise, during 1981-1983 in Papua New Guinea.
[42]
Female mate choice across spatial scales: influence of lek and male ...
Feb 25, 2009 · We used paternity analyses to examine how lek versus male attributes influence mate choice in the blue-crowned manakin Lepidothrix coronata.
[43]
Field playback of male display attracts females in lek breeding sage ...
The “after-response” following a playback suggests that some females present during a playback remembered its location and approached on a subsequent lek visit.
[44]
Olfactory cues influence female choice in two lek-breeding antelopes
Apr 16, 1992 · Females were probably attracted to the soil by smells that had accumulated from heavy use by other females. Because of this attraction, ...Missing: mammals | Show results with:mammals
[45]
A Morning on a Greater Sage-Grouse Lek - National Audubon Society
May 30, 2024 · One researcher observed a male mate 37 times in one morning with 37 different females. Twenty-three of those were in a 23-minute period!
[46]
Multiple paternity in a lek mating system: Females mate multiply when they choose inexperienced sires
### Summary of Multiple Paternity and Female Multiple Mating in Lance-tailed Manakins
[47]
The effects of parasites on male ornaments and female choice in the ...
We describe the results of two studies of parasitic infection in the black grouse (Tetrao tetrix). The first deals with our own observations of lekking black ...
[48]
Fluctuating asymmetry and copulation success in lekking black grouse
Höglund et al. The effects of parasites on male ornaments and female choice in the lek-breeding black grouse (Tetrao tetrix). Behav. Ecol. Sociobiol. (1992).
[49]
Determinants of mating success in a lek‐mating species - Queffelec
May 26, 2025 · Determinants of mating success in a lek-mating species. Joséphine Queffelec,. Corresponding Author. Joséphine Queffelec. queffelec.josephine ...Introduction · Methods · Discussion<|control11|><|separator|>
[50]
Sexual Selection on Leks: A Fruit Fly Primer - PubMed
May 1, 2018 · The objective of this paper is to provide an overview of empirical findings on lek-forming tephritid and drosophilid flies. The essay is ...
[51]
The energetic cost of display in male sage grouse - ADS
Daily energy expenditure for the most vigorously displaying males was two times higher than for a non-displaying male and four times higher than basal metabolic ...
[52]
The costs of secondary sexual characters in the lekking great snipe ...
The daily energy expenditure of displaying males was estimated to be on average 4 times the basal metabolic rate, which is close to the supposed maximum ...
[53]
Predation risk and lek-breeding in Uganda kob - ScienceDirect
While on a lek, the aggregation of potential prey may attract predators (Balmford and Turyaho, 1992). Because males spend more time on a lek and make more trips ...
[54]
Lek Breeding and Territorial Aggression in White‐eared Kob - 1987
Territorial aggression led to damaging injuries in several instances, and mortality of breeding age males was disproportionately high.
[55]
Determinants of yearling male lekking effort and mating success in ...
Jul 25, 2012 · Our results show that yearling male black grouse in good condition can establish territories and have some limited mating success, especially during increasing ...
[56]
testosterone increases parasite infection in red grouse - Journals
Sep 21, 2005 · We show that males with increased testosterone levels had greater parasite intensities than controls after one year. We discuss possible ...
[57]
Predicted deleterious mutations reveal the genetic architecture of ...
Aug 11, 2025 · For male black grouse, high lek attendance is critical for achieving reproductive success, as those males with the highest attendance ...Missing: replacement | Show results with:replacement
[58]
It takes two to tango: reproductive skew and social correlates of male ...
Mar 25, 2009 · Our data show that four out of six leks studied had significant reproductive skew, with success apportioned to very few individuals in each lek.<|separator|>
[59]
Lek breeding and territorial aggression in white-eared Kob.
Such fights often induced females to leave their original partners. TA led to damaging injuries, and mortality of breeding age males was disproportionately high ...
[60]
Costs and Benefits of Female Mate Choice: Is There a Lek Paradox?
In this paper, we emphasize natural selection acting directly on females and their offspring. We argue that, although females are expected to pay lower costs in ...
[61]
https://www.journals.uchicago.edu/doi/10.1086/285093
[62]
https://doi.org/10.1093/beheco/3.4.300
[63]
Offspring viability benefits but no apparent costs of mating with high ...
Females exposed to males with high courtship rates mated more often and produced offspring with greater egg–adult viability.
[64]
A resolution of the lek paradox - Journals
This hypothesis predicts that strong sexual selection, due to female choice, leads to fixation of most genetic variation in male sexual characters.
[65]
[PDF] The lek paradox and the capture of genetic variance by condition ...
Our explanation offers a resolution to the lek paradox and rests on only two assumptions; condition dependence of sexually selected traits and high genetic ...<|control11|><|separator|>
[66]
On the resolution of the lek paradox - ScienceDirect.com
The only resolution of the lek paradox is that the directional selection imposed by female choice is not enough to deplete the genetic variance in fitness.Missing: definition | Show results with:definition
[67]
Heritability and genetic correlation between the sexes in a songbird ...
Nov 17, 2010 · The threshold heritability of trait expression in females was lower, around 0.3, supporting autosomal-based trait expression in females.
[68]
Molecular evidence supports a genic capture resolution of the lek ...
Mar 25, 2019 · Together, these results support the main premise of the lek paradox that sexual selection erodes genetic variation and suggest a role for ...<|control11|><|separator|>
[69]
Predicted deleterious mutations reveal the genetic architecture of ...
Aug 11, 2025 · Specifically, the total GERP load reduces lek attendance, a crucial behavioural trait for mating success and an indicator of genetic quality in ...
[70]
Effect of Androgens on the Avian Muscular Transcriptome - PMC
Here we use RNA sequencing to explore how testosterone (T) modulates the muscular transcriptome to support male manakin courtship displays. In addition, we ...Missing: lek | Show results with:lek
[71]
Draft genome assemblies of four manakins | Scientific Data - Nature
Sep 13, 2022 · Selection analysis of plumage color related genes. Manakins are characterized by a variety of plumage colors57,58,59.
[72]
An epigenetic resolution of the lek paradox - PubMed
Female choice for traits signaling male genetic quality is expected to erode heritable variation in fitness, undermining the benefits of choice.
[73]
Hotshots, Hotspots, and Female Preference in the Organization of ...
The structure of leks is created by a complex of malemale interactions, with conflict between hotshots (who attempt to control lek mating) and subordinates, ...
[74]
Hotspots and the dispersion of leks - ScienceDirect
Males in the simulations were expected to settle at sites with high levels of female traffic (hotspots) and to correct for the sharing of females between ...
[75]
Dynamic Disturbance Processes Create Dynamic Lek Site Selection ...
Sep 22, 2015 · We specifically addressed stability of lek locations in the context of the fire-grazing interaction, and the environmental factors influencing ...Methods · Fig 2. Lek Trends · Fig 3. Frequency And...
[76]
Fruit resources shape sexual selection processes in a lek mating ...
Sep 25, 2024 · We demonstrate a link between fine-scale fruit resource availability and indices of male mating success in the white-bearded manakin (Manacus manacus).
[77]
https://royalsocietypublishing.org/doi/10.1098/rsbl.2024.0284
[78]
[PDF] Lek-kin in birds — provoking theory and surprising new results
I review studies relating to the question of whether kin selection is prevalent on leks and may help to explain their evolution. This is not a comprehensive ...
[79]
Kin selection does not explain male aggregation at leks of 4 ...
Abstract. In lek-mating systems, males aggregate at display arenas and females visit solely for the purpose of mating. This breeding system is characterize.