Lek mating

The term derives from the Swedish lek (), a noun which typically denotes pleasurable and less rule-bound games and activities ("play", as by children), or from the Swedish verb "leka", to play. Written English use of lek dates from the late 1860s and early 1870s. 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. ==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. The presence of a group name means that some species in that group lek; groups with no lekking members are not shown. }} There is also evidence from scrape marks left by dinosaurs that suggest they also indulged in leks. ==Lekking behaviour==

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. Higher-ranking individuals have larger personal space bubbles. 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 classical lek. Males in an exploded lek are outside visual range of one another, but they stay within earshot. Exploded lek territories are much larger than classic systems and more variable in size. A well-known example of exploded leks is the "booming" call of the kākāpō, the males of which position themselves many kilometres apart from one another to signal to potential mates. Stability Lek territories of different taxa are stable and do not vary in terms of size and location. Males often return to the same mating sites because of female fidelity. Avian females such as the black grouse and great snipe are faithful to males and not to mating sites. 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. 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. ==Costs and benefits==

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. 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. Lekking is associated with sexual dimorphism across a range of bird taxa. However, lekking reduces the cost of female searching because the congregating of males makes mate selection easier. 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. ==The lek paradox==

s trying to attract the attention of a receptive female 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. 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. 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. 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. This implies that a female gains indirect benefits from her choice in the form of "good genes" for her offspring. Amotz Zahavi argued that male sexual characteristics only convey useful information to the females if these traits confer a handicap on the male. 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. 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. 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. ==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 presence of a hotshot male seems to attract males and females to the lek. 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. 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. Hotspot model s, 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. 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. 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. 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. 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. Predation protection Another hypothesis is predation protection, or the idea that there is a reduction in individual predation risk in a larger group. 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, as well as other birds such as grouse species. ==References==