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four graphs showing the relatedness coefficient of organisms leks of different species of manakin.
Displaying Relatedness Among Members of Manakin Leks
This graph shows the relatedness coefficients of organisms in leks of different species of manakin. The average relatedness is 0, which led Loiselle and team to conclude that relatedness did not have an effect on lek formation and persistence (Loiselle 2006).

McDonald concludes that while some studies have shown kin selection to play a role in lekking, the results of this study demonstrate that kin selection is unlikely to be the driving evolutionary force in lek formation (McDonald 2007). To completely rule out kin selection as a mechanism for lek formation in manakins, however, the relationship between male status and reproductive success within lek must be analyzed. Additionally, future experiments must be done to determine whether or not males can recognize unfamiliar kin (Loiselle 2006).


The study of lekking animals has led to important hypotheses and explanations for their paradoxical behavior, but a conclusive explanation has yet to be drawn. The preference, hotshot, hotspot, and hypotheses are all attempts to explain the peculiar behavior of this classification of animals, but no theory accurately accounts for all of the observations observed across all lekking species. The diversity of animals that mate in leks further complicates the research and explanation process, as behaviors that can be explained in one species may persist for different reasons in organisms of another species, making it difficult for researchers to draw definite conclusions. The phenotypic gambit enables researchers to assume a genetic basis for lek behavior, but the exact mechanism is yet unknown.

Discussion questions

  1. Do you think lekking is selected for by similar or different selective forces in different species?
  2. Why does lekking appear paradoxical from an evolutionary standpoint? How is this paradox overcome?


  • Bower - a shelter, dwelling, or nest
  • Conspecific - referring to the same species as the organism in question
  • Coverts - in birds, the small feathers that conceal the base of the animal’s larger tail and wing feathers
  • Dimorphism - often the result of sexual selection or disruptive natural selection, dimorphism refers to the existence of two distinct forms of the same species that differ in one or more characteristics such as size, color, pattern, etc.
  • Epigamic - any mal characteristic, such as coloring, behavior, or patterning, that serves to attract the other sex during courtship
  • Fecundity - having abundant offspring
  • Frugivorous : having a diet consisting mostly of fruit
  • Gene - the fundamental unit of inheritance and evolution; a hereditary unit that results in the expression of a phenotype through protein manufacture
  • Genotype - the genetic makeup of an organism
  • Heterozygosity - the state of being heterozygous, i.e. having more than one allele at a locus
  • Hierarchy - a system of ranking determined by escalations between individuals of a species. Often, the hierarchy is determined by strength, fitness, and/or age.
  • Home range - the limits to the area in which an animal lives the majority of the time
  • Hotspot - an area characterized by high encounter rates with females of a given species. One explanation for lekking behavior is that male settlement is determined by these hotspots and, thus, leks are formed by the aggregation of individual males settling where they are most likely to be encountered by a female.
  • Inclusive fitness - the fitness of an organism determined by taking into account not only its direct offspring but those of its kin as well. Because an organism shares many of its genes with its kin, certain cost-benefit tradeoffs increase its fitness by helping its kin reproduce rather than reproducing itself.
  • Kin selection - an evolutionary theory that posits some behaviors, though they may not directly improve an individual’s fitness, improve the fitness of relatives that share their genes and thus are selected for. Kin selection can be explained by Hamilton’s Rule.
  • Lek - an area in which males of a certain species gather together in competitive sexual courtship and mating displays. Leks themselves are small and are composed of the individual territories of many males. Leks are often characterized by a lack of resources and by a clear hierarchy among males.
  • Monogamy - the practice of having only one sexual partner throughout a period of time; males mate with the same female for multiple attempts during a period of time.
  • Obligate monogamy - monogamy that occurs across an organisms life span; a life-pair
  • Multilocus fingerprinting - a form of DNA analysis designed to identify polymorphisms at multiple loci
  • Natural selection - the process, defined by Charles Darwin, by which animals best suited for their environment are the most successful in reproduction and thus pass on their adaptive genes in a greater percentage to the next generation
  • Networking - male-male social interactions, as seen in coordinated displays
  • Phenotype - observable, physical expression of a trait
  • Phenotypic gambit - the assumption that, because genes control the expression of phenotypic characteristics and behaviors, evolution of characteristics/behaviors can be assumed to have a genetic basis, even if the genes are unknown. This enables researchers to study the persistence of traits without having to determine the exact chemical mechanism by which they persist.
  • Polyandry - the state of having more than one male mate at a time
  • Polygyny - the state of having more than one female mate at a time
  • Sexual selection - behaviors or characteristics that are not necessarily of adaptive advantage but increase the likelihood of successful mating of an organism
  • Sympatric - of or relating to the same geographic region (without interbreeding)
  • Variability - the description of deviation between individuals in genetic characteristics in a population ; the potential of a genotype to diverge when subjected to environmental pressures


  • Alatalo, Rauno V., Jacob Höglund, Arne Lundberg, and William J. Sutherland. 1992."Evolution of Black Grouse Leks: Female Preferences Benefit Males in Larger Leks." Evolution of Black Grouse Leks: Female Preferences Benefit Males in Larger Leks. 3(1): 53-59.
  • Bro-Jørgensen, Jakob. 2002. "Overt Female Mate Competition and Preference for Central Males in a Lekking Antelope." Proceedings of the National Academy of Sciences of the United States of America. 99(14): 9290-293.
  • Brown, Jerram L. 1997. “A Theory of Mate Choice Based on Heterozygosity.” Behavioral Ecology . 8(1): 60-65.
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  • Dastagir, Shekib, Kim Di Minni', Jeffrey Pritsky, and Hamid Saadati. "Evolution of Leks." Thesis. New York University, 1997. Print.
  • Duncan, Brittany A., Patrick D. Ulam, and Ronald C. Arkin. "Lek Behavior as a Model for Multi-Robot Systems." Scientific Commons . Institute for Media and Communications Management, 2009. Web.
  • Durães, Renata. 2009. "Lek Structure and Male Display Repertoire of Blue-Crowned Manakins in Eastern Ecuador." The Condor. 111: 453-61.
  • Durães, Renata, Bette A. Loiselle, Patricia G. Parker, and John G. Blake. 2009. "Female Mate Choice Across Spatial Scales: Influence of Lek and Male Attributes on Mating Success of Blue-Crowned Manakins." Proceedings of the Royal Society. 276(1663): 1875-881.

    This study concludes that while females will travel to reach a larger lek, males at larger leks are not individually more likely to have offspring. Additionally, males that display more are more likely to have offspring. Also concluded was that as the size of the lek increases, display frequency and heterozygosity both increase as well.

  • Emlen, Stephen T. 1976. “Lek Organization and Mating Strategies in the Bullfrog.” Behavioral Ecology and Sociobiology . 1(3): 283-313.
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  • Hamilton, Ian M., Marcel P. Haesler, and Michael Taborsky. 2006. "Predators, Reproductive Parasites, and the Persistence of Poor Males on Leks." Behavioral Ecology. 17(1): 97-107.
  • Hernandez, Marcel L., Alasdair I. Houston, and John M. McNamara. 1999. "Male Rank and Optimal Lek Size." Behavioral Ecology. 10(1): 73-79.
  • Isvaran, Kavita, and Colette M. St. Mary. 2003. "When Should Males Lek? Insights From a Dynamic State Variable Model." Behavioral Ecology 14(6): 876-86.
  • Knopp, T., M. Heimovirta, H. Kokko, and J. MerilÄ. 2008. "Do Male Moor Frogs (Rana arvalis) Lek with Kin?" Molecular Ecology . 17(10): 2522-530.
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  • Loiselle, Bette A., Thomas B. Ryder, Renata Durães, Wendy Tori, John G. Blake, and Patricia G. Parker. 2007. "Kin Selection Does Not Explain Male Aggregation at Leks of 4 Manakin Species." Behavioral Ecology. 18(2): 287-91.

    This article describes how a common explanation for lekking behavior is the theory of inclusive fitness, because some hypothesize that males gain indirect benefits from genetically related reproducing males. This, however, is unlikely to be an explanation for the behavior of manakins, because the birds studied were no more significantly related to one another than would be expected from random distribution.

  • McDonald, David B. 2009. "Young-Boy Networks Without Kin Clusters in a Lek-Mating Manakin." Behavioral Ecology and Sociobiology. 63(7): 1029.

    The author’s previous work showed that the networking of young males predicts their future reproductive success. In this article, the author attempted to determine whether kinship has an effect on the networking of the young males. He found that males that were not as closely related tended to be found in leks that were further apart and relatedness and distance between individuals were inversely related. However, the mean in relatedness was negative, so overall, the author concluded that kin selection does not play a role in the cooperation of leks.

  • Miller, Christine W., and Allen J. Moore. 2007. "A Potential Resolution to the Lek Paradox Through Indirect Genetic Effects." Proceedings of the Royal Society . 274(1615): 1279-286.
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  • Ryder, T. B., Blake, J. G.,&Loiselle, B. R. A. 2006. “A Test of the Environmental Hotspot Hypothesis for Lek Placement in Three Species of Manakins (Pipridae) in Ecuador.” The Auk, 123 (1): 247-258.
  • Ryder, Thomas B., David B. McDonald, John G. Blake, Patricia G. Parker, and Bette A. Loiselle. 2008. "Social Networks in the Lek-Mating Wire-Tailed Manakin." Proceedings of the Royal Society. 275(1641): 1367-374.
  • Ryder, Thomas B., Patricia G. Parker, John G. Blake, and Bette A. Loiselle. 2009. "It Takes Two to Tango: Reproductive Skew and Social Correlates of Male Mating Success in a Lek-Breeding Bird." Proceedings of the Royal Society. 276(1666): 2377-384.
  • Starr, Cecie; Taggart, Ralph (1992). Biology – the Unity and Diversity of Life, 6th Ed. . Wadsworth Publishing Company.
  • Théry, Marc. 1990. "The Evolution of Leks Through Female Choice: Differential Clustering and Space Utilization in Six Sympatric Manakins." Behavioral Ecology and Sociobiology. 30: 227-37.
  • Westcott, David A. 1994. "Lek of Leks: A Role for Hotspots in Lek Evolution?" Proceedings: Biological Sciences . 258(1353): 281-86.
  • Westcott, D. A. 1997. “Lek locations and patterns of female movement and distribution in a Neotropical frugivorous bird..” Animal Behaviour. 53 : 235-247.
  • Young, Kyle A., Martin J. Genner, Domino A. Joyce, and Marcel P. Haesler. 2009. "Hotshots, Hot Spots, and Female Preference: Exploring Lek Formation Models with a Bower-Building Cichlid Fish." Behavioral Ecology. 20(3): 609-15.

    This study focuses on the three hypotheses for lek formation, the hotshot hypothesis, the hotspot hypothesis, and female choice. The results favor the female preference hypothesis, as female encounters increase with the size of the lek but behavior did not increase with size once courted. Competitive interactions between males also increased with the lek size but their hunting rate decreased, which also supports the female choice model because it indicates that males experience a high cost of upholding their territories in the larger leks. The experiments in the study were performed on a species of cichlid fish.

Author’s biography

a picture of the author

Kirby Kempe is a first-year Economics major and first-time city dweller at Rice University in Houston, Texas. Originally from Florida, Kempe grew up on the beach and quickly fell in love with the outdoors, taking the phrase “tree hugger” to the extreme. In those rare moments of tranquility hiding in the life of a college student, Kempe enjoys reading, laughing, photographing, Houston highways, loose-leaf tea, and StumbleUpon. She has dreams of becoming a neonatologist, and is most frequently seen on shift for EMS on the ambulance or in the Emergency Rooms of Houston’s local hospitals.

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Source:  OpenStax, Mockingbird tales: readings in animal behavior. OpenStax CNX. Jan 12, 2011 Download for free at http://cnx.org/content/col11211/1.5
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