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Lek mating

March 03, 2023

For other uses, see Lek (disambiguation).

A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate.[1] A lek can also indicate an available plot of space able to be utilized by displaying males to defend their own share of territory for the breeding season. A lekking species is characterised by male displays, strong female mate choice, and the conferring of indirect benefits to males and reduced costs to females. Although most prevalent among birds such as black grouse, lekking is also found in a wide range of vertebrates including some bony fish, amphibians, reptiles, and mammals, and arthropods including crustaceans and insects.

Greater sage-grouse at a lek, with multiple males displaying for the less conspicuous females

A classical lek consists of male territories in visual and auditory range of each other. An exploded lek, as seen in the kakapo (the owl parrot), has more widely separated territories, but still in auditory range. Lekking is associated with an apparent paradox: strong sexual selection by females for specific male traits ought to erode genetic diversity by Fisherian runaway, but diversity is maintained and runaway does not occur. Many attempts have been made to explain it away,[2][3][4][5] but the paradox remains.

Etymology

The term derives from the Swedish lek, a noun which typically denotes pleasurable and less rule-bound games and activities ("play", as by children). English use of lek dates to the 1860s. Llewelyn Lloyd's The Game birds and wild fowl of Sweden and Norway (1867) introduces it (capitalised and in single quotes, as 'Lek') explicitly as a Swedish term.[6]

Taxonomic range

Lekking was originally described in the Tetraonidae (grouse, boldface in cladogram), in particular the black grouse (Swedish: "orrlek") and capercaillie (Swedish: "tjäderlek"), but it is widely distributed phylogenetically among other birds, and in many other animal groups within the vertebrates and the arthropods, as shown in the cladogram.[1][7][8]

The presence of a group name means that some species in that group lek; groups with no lekking members are not shown.

Lekking behaviour

 
Sage grouse lek mating arena, in which each male, alpha-male (highest ranking), beta-male, gamma-male, etc., guards a territory of a few meters in size. The dominant males may each attract eight or more females.[33] Higher-ranking individuals have larger personal space bubbles.[34] Bird leks may have 10-200 individuals. A strict hierarchy accords the top-ranking males the most prestigious central territories. Females come to choose mates when the males' hierarchy is established, and preferentially mate with the dominants in the centre.

Types

There are two types of lekking arrangement: classical and exploded. In the classic lekking system, male territories are in visual and auditory range of their neighbours. In an exploded lek, males are further away from one another than they would be in a classic lek. Males in an exploded lek are outside visual range of one another, but they stay within earshot.[35] Exploded lek territories are much larger than classic systems and more variable in size.[36] A well-known example of exploded leks is the "booming" call of the kakapo, the males of which position themselves many kilometres apart from one another to signal to potential mates.[37]

Stability

Lek territories of different taxa are stable and do not vary in terms of size and location.[38] Males often return to the same mating sites because of female fidelity.[39] Avian females such as the black grouse and great snipe are faithful to males and not to mating sites.[40] Successful males congregate in the same area as the previous breeding season because it is familiar to them, while females return to reunite with their males. Females do not return to a mating site if their male partner is not present.[40] Another possible explanation for lek stability is from male hierarchies within a lek. In some species of manakin, subordinate betas may inherit an alpha's display site, increasing the chances of female visitation.[40] Rank may also contribute to the stability of lek size, as lower ranking males may congregate to achieve a perceived optimal size to attract females.[41]

Female mating preferences

A meta analysis of 27 species found that qualities such as lekking size, male display rate, and the rate of male aggression exhibit positive correlation with male success rates.[1] A positive correlation was also found between attendance, magnitude of exaggerated traits, age, frequency of fights, and mating success.[1] This female preference leads to mating skew, with some males being more successful at copulating with females. The variation in mating success is quite large in lek mating systems with 70–80% of matings being attributed to only 10–20% of the males present.[42]

Costs and benefits

Further information: Sexual selection
Lekking behaviour in the Clusiid fly Paraclusia tigrina

The main benefit for both sexes is mating success. For males, the costs stem from females' preferences. The traits that are selected for may be energetically costly to maintain and may cause increased predation. For example, increased vocalization rate caused a decrease in the mass of male great snipes.[43] Other costs can derive from male combat. For example, male great snipes regularly fight to display dominance or defend their territory, with females preferring victorious males.[43] Aggressive male black grouse are preferred over non-aggressive males and when the males fight they tear feathers from each other's tails.[44] Lekking is associated with sexual dimorphism across a range of bird taxa.[7]

At first glance, it may seem that females receive no direct benefits from lekking, since the males are only contributing genes to the offspring in the absence of parental care or other benefits.[45] However, lekking reduces the cost of female searching because the congregating of males makes mate selection easier.[46] Females do not have to travel as far, since they are able to evaluate and compare multiple males within the same vicinity. Further, having the males in one place may reduce the amount of time a female is vulnerable to predators. When under predatory pressure, female marbled reed frogs consistently choose leks near their release sites; high male calling rates were observed to reduce female search time.[47]

The lek paradox

 
A group of three male great-tailed grackles trying to attract the attention of a receptive female
Main article: Lek paradox

Since sexual selection by females for specific male trait values should erode genetic diversity, the maintenance of genetic variation in lekking species constitutes a paradox in evolutionary biology. Many attempts have been made to explain it away, but the paradox remains.[48] There are two conditions in which the lek paradox arises. The first is that males contribute only genes and the second is that female preference does not affect fecundity.[49] Female choice should lead to directional runaway selection, resulting in a greater prevalence for the selected traits. Stronger selection should lead to impaired survival, as it decreases genetic variance and ensures that more offspring have similar traits.[50] However, lekking species do not exhibit runaway selection. In a lekking reproductive system, what male sexual characteristics can signal to females is limited, as the males provide no resources to females or parental care to their offspring.[51] This implies that a female gains indirect benefits from her choice in the form of "good genes" for her offspring.[52]

Amotz Zahavi argued that male sexual characteristics only convey useful information to the females if these traits confer a handicap on the male.[53][54] Zahavi's handicap principle may offer a resolution to the lek paradox, for if females select for the condition of male ornaments, then their offspring have better fitness. Another potential resolution to the lek paradox is Rowe and Houle's theory that sexually selected traits depend on physical condition, which might in turn, summarize many genetic loci.[52] This is the genic capture hypothesis, which describes how a significant amount of the genome is involved in shaping the traits that are sexually selected.[51] There are two assumptions in the genic capture hypothesis: the first is that sexually selected traits are dependent upon condition, and the second is that general condition is attributable to high genetic variance.[52] In addition, W. D. Hamilton and Marlene Zuk proposed that sexually selected traits might signal resistance to parasites.[55] One resolution to the lek paradox involves female preferences and how preference alone does not cause a drastic enough directional selection to diminish the genetic variance in fitness.[56] Another conclusion is that the preferred trait is not naturally selected for or against and the trait is maintained because it implies increased attractiveness to the male.[49]

 
In the little bustard, the presence of a hotshot male seems to attract males and females to the lek.

Evolution

Several possible mechanisms have been proposed as to why males cluster into leks, including the hotshot, hotspot, black hole, kin selection, and predation protection hypotheses, as described below.

Hotshot hypothesis

The hotshot hypothesis is the only model that attributes males as the driving force behind aggregation. The hotshot model hypothesizes that attractive males, known as hotshots, garner both female and male attention.[2] Females go to the hotshots because they are attracted to these males. Other males form leks around these hotshots as a way to lure females away from the hotshot. A manipulative experiment using the little bustard, Tetrax tetrax, was done to test the various lek evolution models.[4] The experiment involved varying the size and sex ratio of leks using decoys. To test whether or not the presence of a hotshot determined lek formation, a hotshot little bustard decoy was placed within a lek. After the fake hotshot was added to the lek, both male and female visitation to the lek increased, tending to confirm the hypothesis.[4]

Hotspot model

 
In manakins, males aggregate near hotspots with plentiful fruit, where females tend to go.

The hotspot model considers the female density to be the catalyst for the clustering of males. This model predicts that leks will form where females tend to reside as a way to increase female interaction.[3] Female manakin traffic has been observed to be concentrated around leks, bathing sites, and fruiting areas, with males aggregated near the most visited fruiting resources.[3] The hotspot model also predicts that lek size is dependent upon the number of females inhabiting a patch of land.[2] To test if the number of females affects lek formation, a group of female little bustard decoys were added to a lek. The presence of these female decoys did not have an effect on lek size, tending to refute the hypothesis.[4]

Blackhole model

The blackhole model proposes that females have a preference for neither size nor type of male, but rather that females tend to be mobile and mate wherever leks may be located.[4] This model predicts that female mobility is a response to male harassment.[57] This prediction is difficult to test, but there was a negative correlation found between male aggressiveness and female visitation in the little bustard population, suggesting that the model might be correct.[4] Evidence supporting the black hole model is mainly found in ungulates.[39]

Kin selection

 
In black grouse, leks are composed of brothers and half-brothers, suggesting a kin selection mechanism.
Further information: Kin selection

An alternative hypothesis for lekking is kin selection, which assumes that males within a lek are related to one another. As females rarely mate outside of leks, it is advantageous for males to form leks.[5] Although not all males within a lek mate with a female, the unmated males still receive fitness benefits. Kin selection explains that related males congregate to form leks, as a way to attract females and increase inclusive fitness.[40] In some species, the males at the leks show a high degree of relatedness, but this does not apply as a rule to lek-forming species in general.[58][59][60] In a few species such as peacocks and black grouse, leks are composed of brothers and half-brothers. The lower-ranking males gain some fitness benefit by passing their genes on through attracting mates for their brothers, since larger leks attract more females. Peacocks recognize and lek with their brothers, even if they have never met before.[61]

Predation protection

Another hypothesis is predation protection, or the idea that there is a reduction in individual predation risk in a larger group.[4] This could work both for the males within the group and any female who visits the lek.[62] Protection also explains the presence of mixed leks, when a male of one species joins a lek of another species for protection from a common set of predators. This occurs with manakins,[63] as well as other birds such as grouse species.[64]

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  64. ^ Gibson, Robert M.; Aspbury, Andrea S.; McDaniel, Leonard L. (2002). "Active formation of mixed–species grouse leks: a role for predation in lek evolution?". Proceedings of the Royal Society of London B: Biological Sciences. 269 (1509): 2503–2507. doi:10.1098/rspb.2002.2187. ISSN 0962-8452. PMC 1691199. PMID 12573063.

March 03, 2023
mating, other, uses, disambiguation, aggregation, male, animals, gathered, engage, competitive, displays, courtship, rituals, known, lekking, entice, visiting, females, which, surveying, prospective, partners, with, which, mate, also, indicate, available, plot. For other uses see Lek disambiguation A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals known as lekking to entice visiting females which are surveying prospective partners with which to mate 1 A lek can also indicate an available plot of space able to be utilized by displaying males to defend their own share of territory for the breeding season A lekking species is characterised by male displays strong female mate choice and the conferring of indirect benefits to males and reduced costs to females Although most prevalent among birds such as black grouse lekking is also found in a wide range of vertebrates including some bony fish amphibians reptiles and mammals and arthropods including crustaceans and insects Greater sage grouse at a lek with multiple males displaying for the less conspicuous females A classical lek consists of male territories in visual and auditory range of each other An exploded lek as seen in the kakapo the owl parrot has more widely separated territories but still in auditory range Lekking is associated with an apparent paradox strong sexual selection by females for specific male traits ought to erode genetic diversity by Fisherian runaway but diversity is maintained and runaway does not occur Many attempts have been made to explain it away 2 3 4 5 but the paradox remains Contents 1 Etymology 2 Taxonomic range 3 Lekking behaviour 3 1 Types 3 2 Stability 3 3 Female mating preferences 4 Costs and benefits 5 The lek paradox 6 Evolution 6 1 Hotshot hypothesis 6 2 Hotspot model 6 3 Blackhole model 6 4 Kin selection 6 5 Predation protection 7 ReferencesEtymology EditThe term derives from the Swedish lek a noun which typically denotes pleasurable and less rule bound games and activities play as by children English use of lek dates to the 1860s Llewelyn Lloyd s The Game birds and wild fowl of Sweden and Norway 1867 introduces it capitalised and in single quotes as Lek explicitly as a Swedish term 6 Taxonomic range EditLekking was originally described in the Tetraonidae grouse boldface in cladogram in particular the black grouse Swedish orrlek and capercaillie Swedish tjaderlek but it is widely distributed phylogenetically among other birds and in many other animal groups within the vertebrates and the arthropods as shown in the cladogram 1 7 8 The presence of a group name means that some species in that group lek groups with no lekking members are not shown Nephrozoa Deuterostomia Vertebrates Teleosts Atlantic cod 9 Desert pupfish 10 Cichlids 11 Tetrapoda Amphibians some frogs 12 13 Amniota Reptilia Marine iguanas 14 Birds Tetraonidae 1 grouse Neoaves Otididae 7 bustards Scolopacidae 7 sandpipers Psittacidae 7 parrots Passeriformes Cotingidae 7 cotingas Pipridae 7 manakins Pycnonotidae 7 bulbuls Ploceidae 7 weavers Paradisaeidae 7 birds of paradise Mammals Bats 15 16 Bovidae several species 17 18 19 20 Pinnipeds 21 22 Protostomia Decapoda fiddler crab 23 Insects Hymenoptera Vespidae paper wasps 24 Pompilidae tarantula hawk wasps 25 Formicidae some ants 26 Apoidea some bees 27 Lepidoptera some butterflies 28 and moths 29 30 Diptera fruit flies 31 druid flies 32 Lekking behaviour Edit A posing western capercaillie Sage grouse lek mating arena in which each male alpha male highest ranking beta male gamma male etc guards a territory of a few meters in size The dominant males may each attract eight or more females 33 Higher ranking individuals have larger personal space bubbles 34 Bird leks may have 10 200 individuals A strict hierarchy accords the top ranking males the most prestigious central territories Females come to choose mates when the males hierarchy is established and preferentially mate with the dominants in the centre Types Edit There are two types of lekking arrangement classical and exploded In the classic lekking system male territories are in visual and auditory range of their neighbours In an exploded lek males are further away from one another than they would be in a classic lek Males in an exploded lek are outside visual range of one another but they stay within earshot 35 Exploded lek territories are much larger than classic systems and more variable in size 36 A well known example of exploded leks is the booming call of the kakapo the males of which position themselves many kilometres apart from one another to signal to potential mates 37 Stability Edit Lek territories of different taxa are stable and do not vary in terms of size and location 38 Males often return to the same mating sites because of female fidelity 39 Avian females such as the black grouse and great snipe are faithful to males and not to mating sites 40 Successful males congregate in the same area as the previous breeding season because it is familiar to them while females return to reunite with their males Females do not return to a mating site if their male partner is not present 40 Another possible explanation for lek stability is from male hierarchies within a lek In some species of manakin subordinate betas may inherit an alpha s display site increasing the chances of female visitation 40 Rank may also contribute to the stability of lek size as lower ranking males may congregate to achieve a perceived optimal size to attract females 41 Female mating preferences Edit A meta analysis of 27 species found that qualities such as lekking size male display rate and the rate of male aggression exhibit positive correlation with male success rates 1 A positive correlation was also found between attendance magnitude of exaggerated traits age frequency of fights and mating success 1 This female preference leads to mating skew with some males being more successful at copulating with females The variation in mating success is quite large in lek mating systems with 70 80 of matings being attributed to only 10 20 of the males present 42 Costs and benefits EditFurther information Sexual selection source source source source source source source source source source Lekking behaviour in the Clusiid fly Paraclusia tigrina The main benefit for both sexes is mating success For males the costs stem from females preferences The traits that are selected for may be energetically costly to maintain and may cause increased predation For example increased vocalization rate caused a decrease in the mass of male great snipes 43 Other costs can derive from male combat For example male great snipes regularly fight to display dominance or defend their territory with females preferring victorious males 43 Aggressive male black grouse are preferred over non aggressive males and when the males fight they tear feathers from each other s tails 44 Lekking is associated with sexual dimorphism across a range of bird taxa 7 At first glance it may seem that females receive no direct benefits from lekking since the males are only contributing genes to the offspring in the absence of parental care or other benefits 45 However lekking reduces the cost of female searching because the congregating of males makes mate selection easier 46 Females do not have to travel as far since they are able to evaluate and compare multiple males within the same vicinity Further having the males in one place may reduce the amount of time a female is vulnerable to predators When under predatory pressure female marbled reed frogs consistently choose leks near their release sites high male calling rates were observed to reduce female search time 47 The lek paradox Edit A group of three male great tailed grackles trying to attract the attention of a receptive female Main article Lek paradox Since sexual selection by females for specific male trait values should erode genetic diversity the maintenance of genetic variation in lekking species constitutes a paradox in evolutionary biology Many attempts have been made to explain it away but the paradox remains 48 There are two conditions in which the lek paradox arises The first is that males contribute only genes and the second is that female preference does not affect fecundity 49 Female choice should lead to directional runaway selection resulting in a greater prevalence for the selected traits Stronger selection should lead to impaired survival as it decreases genetic variance and ensures that more offspring have similar traits 50 However lekking species do not exhibit runaway selection In a lekking reproductive system what male sexual characteristics can signal to females is limited as the males provide no resources to females or parental care to their offspring 51 This implies that a female gains indirect benefits from her choice in the form of good genes for her offspring 52 Amotz Zahavi argued that male sexual characteristics only convey useful information to the females if these traits confer a handicap on the male 53 54 Zahavi s handicap principle may offer a resolution to the lek paradox for if females select for the condition of male ornaments then their offspring have better fitness Another potential resolution to the lek paradox is Rowe and Houle s theory that sexually selected traits depend on physical condition which might in turn summarize many genetic loci 52 This is the genic capture hypothesis which describes how a significant amount of the genome is involved in shaping the traits that are sexually selected 51 There are two assumptions in the genic capture hypothesis the first is that sexually selected traits are dependent upon condition and the second is that general condition is attributable to high genetic variance 52 In addition W D Hamilton and Marlene Zuk proposed that sexually selected traits might signal resistance to parasites 55 One resolution to the lek paradox involves female preferences and how preference alone does not cause a drastic enough directional selection to diminish the genetic variance in fitness 56 Another conclusion is that the preferred trait is not naturally selected for or against and the trait is maintained because it implies increased attractiveness to the male 49 In the little bustard the presence of a hotshot male seems to attract males and females to the lek Evolution EditSeveral possible mechanisms have been proposed as to why males cluster into leks including the hotshot hotspot black hole kin selection and predation protection hypotheses as described below Hotshot hypothesis Edit The hotshot hypothesis is the only model that attributes males as the driving force behind aggregation The hotshot model hypothesizes that attractive males known as hotshots garner both female and male attention 2 Females go to the hotshots because they are attracted to these males Other males form leks around these hotshots as a way to lure females away from the hotshot A manipulative experiment using the little bustard Tetrax tetrax was done to test the various lek evolution models 4 The experiment involved varying the size and sex ratio of leks using decoys To test whether or not the presence of a hotshot determined lek formation a hotshot little bustard decoy was placed within a lek After the fake hotshot was added to the lek both male and female visitation to the lek increased tending to confirm the hypothesis 4 Hotspot model Edit In manakins males aggregate near hotspots with plentiful fruit where females tend to go The hotspot model considers the female density to be the catalyst for the clustering of males This model predicts that leks will form where females tend to reside as a way to increase female interaction 3 Female manakin traffic has been observed to be concentrated around leks bathing sites and fruiting areas with males aggregated near the most visited fruiting resources 3 The hotspot model also predicts that lek size is dependent upon the number of females inhabiting a patch of land 2 To test if the number of females affects lek formation a group of female little bustard decoys were added to a lek The presence of these female decoys did not have an effect on lek size tending to refute the hypothesis 4 Blackhole model Edit The blackhole model proposes that females have a preference for neither size nor type of male but rather that females tend to be mobile and mate wherever leks may be located 4 This model predicts that female mobility is a response to male harassment 57 This prediction is difficult to test but there was a negative correlation found between male aggressiveness and female visitation in the little bustard population suggesting that the model might be correct 4 Evidence supporting the black hole model is mainly found in ungulates 39 Kin selection Edit In black grouse leks are composed of brothers and half brothers suggesting a kin selection mechanism Further information Kin selection An alternative hypothesis for lekking is kin selection which assumes that males within a lek are related to one another As females rarely mate outside of leks it is advantageous for males to form leks 5 Although not all males within a lek mate with a female the unmated males still receive fitness benefits Kin selection explains that related males congregate to form leks as a way to attract females and increase inclusive fitness 40 In some species the males at the leks show a high degree of relatedness but this does not apply as a rule to lek forming species in general 58 59 60 In a few species such as peacocks and black grouse leks are composed of brothers and half brothers The lower ranking males gain some fitness benefit by passing their genes on through attracting mates for their brothers since larger leks attract more females Peacocks recognize and lek with their brothers even if they have never met before 61 Predation protection Edit Another hypothesis is predation protection or the idea that there is a reduction in individual predation risk in a larger group 4 This could work both for the males within the group and any female who visits the lek 62 Protection also explains the presence of mixed leks when a male of one species joins a lek of another species for protection from a common set of predators This occurs with manakins 63 as well as other birds such as grouse species 64 References Edit a b c d e Fiske P Rintamaki P T Karvonen E 1998 Mating success in lekking males a meta analysis Behavioral Ecology 9 4 328 338 doi 10 1093 beheco 9 4 328 a b c Foster M S Beehler B M 1998 Hotshots Hotspots and Female Preferences in the Organization of Lek Mating Systems The American Naturalist 131 2 203 219 doi 10 1086 284786 S2CID 85295271 a b c Thery M 1992 The evolution of leks through female choice Differential clustering and space utilization in six sympatric manakins Behavioral Ecology and Sociobiology 30 3 4 227 237 doi 10 1007 bf00166707 S2CID 1886240 a b c d e f g Jiguet F Bretagnolle V 2006 Manipulating lek size and composition using decoys An experimental investigation of lek evolution models The American Naturalist 168 6 758 768 doi 10 1086 508808 PMID 17109318 S2CID 2527129 a b Duraes R Loiselle B A Blake J G 2008 Spatial and temporal dynamics at manakin leks Reconciling lek traditionality with male turnover Behavioral Ecology and Sociobiology 62 12 1947 1957 doi 10 1007 s00265 008 0626 0 S2CID 36424100 Lloyd Llewelyn 1867 The Game Birds and Wild Fowl of Sweden and Norway London Frederick Warne amp Co pp 219ff Lloyd also loans Lek stalle Swedish lekstalle play place for pairing ground a b c d e f g h i j Oakes E J 1992 Lekking and the Evolution of Sexual Dimorphism in Birds Comparative Approaches The American Naturalist 140 4 665 684 doi 10 1086 285434 ISSN 0003 0147 PMID 19426038 S2CID 26713188 Bourke Andrew F G 2014 Hamilton s rule and the causes of social evolution Philosophical Transactions of the Royal Society B Biological Sciences The Royal Society 369 1642 20130362 doi 10 1098 rstb 2013 0362 ISSN 0962 8436 PMC 3982664 PMID 24686934 Ponomarenko I Ja 1965 Comparative characteristics of some biological indices of the bottom stages of 0 group cod belonging to the 1956 1958 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evolution and trait introgression across an avian hybrid zone Molecular Ecology 21 6 1477 1486 doi 10 1111 j 1365 294X 2012 05474 x ISSN 1365 294X PMID 22320709 S2CID 10795904 Brumfield Robb T Liu Liang Lum David E Edwards Scott V 1 October 2008 Comparison of Species Tree Methods for Reconstructing the Phylogeny of Bearded Manakins Aves Pipridae Manacus from Multilocus Sequence Data Systematic Biology 57 5 719 731 doi 10 1080 10635150802422290 ISSN 1063 5157 PMID 18853359 Gibson Robert M Aspbury Andrea S McDaniel Leonard L 2002 Active formation of mixed species grouse leks a role for predation in lek evolution Proceedings of the Royal Society of London B Biological Sciences 269 1509 2503 2507 doi 10 1098 rspb 2002 2187 ISSN 0962 8452 PMC 1691199 PMID 12573063 Retrieved from https en wikipedia org w index php title Lek mating amp oldid 1138159247, wikipedia, wiki, book, books, library,

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